UNIT

The Diversification of Life

29 Fungi
Chaiwatphotos/Shutterstock

Netted stinkhorn In this chapter you will learn about

(Phallus indusiatus)
Fungi, the master traders and recyclers in
is an edible fungus
terrestrial ecosystems
related to mushrooms. Fungi as
This species forms by asking by examining mutualists
a greenish, slimy, by
Why and how do Major themes in
foul-smelling cap focusing on Fungi as
biologists study fungi? their diversification
that produces spores. decomposers
Insects attracted by the
29.1, 29.2 29.3
foul smell help disperse
then, by looking Variation in
the spores. more closely at reproductive
Key lineages structures and
of fungi life cycles
29.4

F ungi are eukaryotes that grow as single cells or as large, branching networks of multicellular fila-
ments. Familiar fungi include the mushrooms you’ve encountered in the woods or a grocery store,
the moulds and mildews that grow in homes, the organism that causes athlete’s foot, and the yeasts
used in baking and brewing.
Along with the land plants and the animals, the fungi are one of three major lineages of large, multi-
cellular eukaryotes that occupy terrestrial environments. When it comes to making a living, the species
in these three groups use radically different strategies. Land plants make their own food through photo-
synthesis. Animals and fungi both feed on plants, protists, or each other by releasing digestive enzymes
and absorbing small molecules. But while many animals carry out digestion in a dedicated digestive tract,
This chapter is part of the fungi release their digestive enzymes to the external environment.
Big Picture. See how on Fungi that absorb nutrients from dead organisms are the world’s most important decomposers.
pages 734–735. Although a few types of organisms are capable of digesting the cellulose in plant cell walls, certain fungi

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 and a handful of bacterial species are the only organisms capable
of digesting both the lignin and cellulose that make up wood. 29.1 Why Do Biologists Study Fungi?
Without fungi, Earth’s surface would be piled high with dead tree
Given their importance to life on land and their intricate rela-
trunks and branches. tionships to other organisms, it’s no surprise that fungi are fas-
Other fungi specialize in absorbing nutrients from living cinating to biologists. But there are important practical reasons
organisms. When fungi absorb these nutrients without benefit- as well for humans to study fungi. They can cause debilitating
ing their host, they lower the fitness of their host organism and diseases in humans and crop plants. They nourish the plants
act as parasites. If you’ve ever had athlete’s foot or a vaginal that nourish us. They affect global climate change, because they
yeast infection, you’ve hosted a parasitic fungus. are critical to the carbon cycle on land. Let’s take a closer look at
Most of the fungi that live in association with other organ- some of the ways that fungi affect land-dwelling organisms.
isms benefit their hosts, however. In these cases, fungi are not
parasites but mutualists. Fungi Have Important Economic
• The roots of most land plants are colonized by an array of and Ecological Impacts
mutualistic fungi that provide water and key nutrients such In humans, parasitic fungi cause athlete’s foot, vaginitis, diaper
as nitrogen and phosphorus to the host plant. The soil around rash, ringworm, pneumonia, and thrush, among other miser-
you is alive with an enormous network of fungi that are fertil- ies. But even though these maladies can be serious, in reality no
izing the plants you see aboveground. more than approximately 300 species of fungi—out of the hun-
dreds of thousands of predicted species—regularly cause illness
• Fungi living inside the shoots of certain plants help ward off
in humans. Compared with the frequency of diseases caused by
herbivores by making toxic compounds.
bacteria, viruses, and protists, the incidence of fungal infections
• Many insects harbour single-celled fungi in their guts that aid in humans is low.
their digestion of plant material. It would be easy to argue, in fact, that fungi have done more
• Some insects grow gardens of fungi that they feed with pieces to promote human health than degrade it. The first antibiotic
of leaves. The insects then feed off the fungi, which they main- that was widely used, penicillin, was isolated from a fungus,
tain and continue to cultivate. and soil-dwelling fungi continue to be the source of many
of the most important antibiotics prescribed against bacterial
In short, fungi are the master traders and recyclers in terres-
infections.
trial ecosystems. Some fungi release nutrients from dead plants
The major destructive impact that fungi have on people is
and animals into the soil; others obtain nutrients and then
through the food supply. Fungi known as rusts, smuts, mildews,
transfer them directly to living plants and animals. Because wilts, and blights cause annual crop losses estimated in the
they recycle key elements such as carbon, nitrogen, and phos- billions of dollars. These fungi are particularly troublesome
phorus and because they transfer key nutrients to plants and in wheat, corn, barley, and other grain crops (Figure 29.1a).
animals, fungi profoundly influence ecosystem productivity Saprophytic fungi are also responsible for enormous losses
and biodiversity. In terms of nutrient cycling, fungi make the due to spoilage—particularly for fruit and vegetable growers
world go around. (Figure 29.1b).

(a) Parasitic fungi infect corn and other crops. (b) Saprophytic fungi rot fruits and vegetables.

Figure 29.1 Fungi Cause Problems with Crop Production and Storage. (a) A wide variety of grain crops are
parasitized by fungi. Corn smut is a serious disease in sweet corn, although in Mexico the smut fungus is eaten as a
delicacy. (b) Fungi decompose fruits and vegetables as well as leaves and tree trunks.
L to R: (a) Inga Spence/Photo Researchers, Inc./Science Source; (b) EGON/Alamy Stock Photo

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 On the other hand, fungi also play many important positive and Canada. Before these fungal epidemics occurred, chest-
roles in the human food supply: nuts and elms dominated these habitats, providing abundant
• Mushrooms are consumed in many cultures; in the industrial- food and shelter for many animals, including humans. In
ized nations they are used in sauces, salads, pizza, and many southern Ontario, less than 1 percent of the original popu-
other diverse types of foods. lation of 2 million chestnut trees survive. However, a small
• The yeast Saccharomyces cerevisiae was domesticated thou- population of chestnut trees remains healthy. These southern
sands of years ago; today it and other fungi are essential to the Ontario trees, which seem to be resistant to the epidemic, may
manufacture of bread, soy sauce, cheese, beer, wine, whiskey, be key to the recovery of the species.
and other products. In most cases, domesticated fungi are Alberta and British Columbia are still free of Dutch elm dis-
used in conditions where the cells grow via fermentation, cre- ease. In fact, Edmonton has one of the largest populations of
ating ethanol and by-products like the CO2 that causes bread healthy American elm trees in the world. A single diseased tree
to rise and beer and champagne to fizz. was discovered in Wainwright, Alberta, in 1998, but the tree was
quickly destroyed and the disease did not spread. With the loss of
• Chocolate is made from the seeds of Theobroma cacao, but it
so many trees from disease came new opportunities for other tree
is edible only after the seeds are fermented by several species
species to dominate the landscape.
of fungi.
Despite the widespread harmful impact fungi can have on an
• Enzymes derived from different types of fungi are used to ecosystem, the role of fungi in the healthy functioning of terres-
improve the characteristics of foods ranging from fruit juice trial ecosystems is critical. Let’s look at two important benefits
and candy to meat. fungi provide.
In terms of ecosystems, the impact of fungi can be wide-
spread. Epidemics caused by fungi have killed 4 billion American
chestnut trees and tens of millions of American elm trees. The
Mycorrhizal Fungi Provide Nutrients
fungal species responsible for these epidemics were accidentally for Land Plants
imported on species of chestnut and elm native to other regions Fungi that live in close association with plant roots are said
of the world. When the fungi arrived in North America and began to be mycorrhizal (literally, “fungal root”; see Figure 29.2a).
growing in native chestnuts and elms, the results were cata- The fungi along with the roots they are associated with are
strophic. The local chestnut and elm populations had virtually referred to as mycorrhizae. When biologists first discovered
no genetic resistance to the pathogens and quickly succumbed. how extensive these fungi–plant associations are, they asked
Living chestnut root systems continue to sprout shoots, but these an obvious question: Does plant growth suffer if mycorrhizal
trees rarely live long enough to reproduce. fungi are absent?
The chestnut and elm epidemics radically altered the composi- Figure 29.2b shows a result typical of many experiments. In
tion of upland and floodplain forests in the eastern United States this case, seedlings were grown in the presence and absence of the

(a) Mycorrhizal fungi form extensive networks in soil. (b) Mycorrhizal fungi increase plant growth.

With mycorrhizal
fungi

Without mycorrhizal
fungi

Seedling root
David Read/University of Sheffield

Mycorrhizal network Mycorrhizal Applications, Inc

Figure 29.2 Plants Grow Better in the Presence of Mycorrhizal Fungi. (a) Root system of a larch tree seedling;
the white threads of a mycorrhizal fungus are visible. (b) Typical experimental results when plants are grown with
and without their normal mycorrhizal fungi.

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 mycorrhizal fungi normally found on their roots. The photograph do many microorganisms. In addition, when fungi die or are
documents that this species grows three to four times faster in the eaten, the molecules are passed along to a wide array of other
presence of its normal fungal associates than it does without them. organisms.
For farmers, foresters, and ranchers, the presence of normal Fungi play an important role in cycling carbon atoms through
mycorrhizal fungi can mean the difference between profit and terrestrial ecosystems. Note that there are two basic components
loss. Fungi are critical to the productivity of forests, croplands, of the carbon cycle on land:
and rangelands. Many commercially available products improve 1. The fixation of carbon by land plants—meaning that car-
crop yield by supplying plants with mycorrhizal fungi. bon in atmospheric CO2 is reduced to form sugar, which is
then used to synthesize cellulose, lignin, and other complex
Saprophytic Fungi Accelerate organic compounds in the bodies of plants. (See Chapter 10
the Carbon Cycle on Land to review carbon fixation.)
Fungi that make their living by digesting dead plant material are 2. The release of CO2 from nearly all organisms as the result of
called saprophytes. To understand how saprophytic fungi play a cellular respiration—meaning the oxidation of glucose and
key role in today’s terrestrial environments, recall that cells in the production of the ATP that sustains life. (See Chapter 9 to
vascular tissues of land plants have secondary cell walls contain- review cellular respiration.)
ing both lignin and cellulose (see Chapter 28). Wood forms when The fundamental point is that, for many carbon atoms, sapro-
stems grow in girth by adding layers of lignin-rich vascular tissue. phytic fungi connect the two parts of the carbon cycle.
When trees die, certain fungi are among the organisms that If fungi had not evolved the ability to digest lignin and cel-
break down wood into sugars and other small organic com- lulose soon after land plants evolved the ability to make these
pounds (Figure 29.3). Fungi use these molecules as food, as compounds, carbon atoms would have been sequestered in wood
for millennia instead of being rapidly recycled into glucose mol-
ecules and CO2. Terrestrial environments would be radically dif-
ferent than they are today, and probably much less productive.
Atmospheric On land, fungi make the carbon cycle turn much more rapidly
carbon (CO2) than it would without fungi.
To summarize, biologists study fungi because they affect a
wide range of species in nature, including humans. What tools
CARBON CYCLE are helping researchers understand the diversity of fungi?
LIVE PLANTS

Carbon atoms released Carbon atoms If you understand the carbon cycle, you should be able to
during cellular respiration fixed by plants explain how fungi help connect two major parts of the cycle:
by plants and other during photo-
organisms synthesis carbon fixation and carbon release.

Carbon atoms
stored in plant bodies
(living and dead)
29.2 How Do Biologists Study Fungi?
About 110 000 species of fungi have been described and named
Carbon-containing to date, and several hundreds more are discovered each year. But
nutrients returned
to the carbon cycle the fungi are so poorly studied that the known species are widely
regarded as a tiny fraction of the actual total.
Consider what American biologist Elizabeth Arnold and her
DEAD PLANTS

colleagues found when they analyzed fungi growing on Barro

B. G. Thomson/Photo Researchers, Inc./Science Source

Colorado Island, Panama: Living on the healthy leaves of just two

tropical tree species were a total of 418 distinct morphospecies
of fungi. (Recall from Chapter 24 that morphospecies are distin-
guished from each other by some aspect of morphology.) Because
over 310 species of trees and shrubs grow on Barro Colorado, the
data suggest that tens of thousands of fungi may be native to this
Fungi digest lignin island alone. If further work on fungal diversity in the tropics
and cellulose supports these conclusions, there may turn out to be many mil-
to obtain carbon-
containing nutrients
lions of fungal species. Indeed, a similar and more recent study
on soil fungi suggests that as many as 6 million species of fungi
may be found worldwide.
This viewpoint of fungal diversity was reinforced by an analysis
of the fungi living in the guts of beetles. In this study, American
Figure 29.3 Fungi Speed Up the Carbon Cycle as They Break biologist Meredith Blackwell and her colleagues isolated fungi
Down Dead Trees in Terrestrial Ecosystems. from 27 species of beetles and characterized them by observing

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 various morphological and physiological traits. They also used The fossil record for land plants and animals has been
the direct sequencing approach to analyze the gene that codes for extremely useful to understanding their early evolution (see
the RNA molecule in the small subunit of fungal ribosomes (see Chapters 22 and 28). In contrast, fossils of fungi are very rare, so
Chapter 17). The data showed that in the beetle guts there were biologists rely on morphological and molecular data to explore
over 650 fungal species, 200 of which had never been described. their past. Because most fungi form mycelia and because this
Biologists are only beginning to realize the extent of species body type is so fundamental to the absorptive mode of life, most
diversity in fungi. studies of fungal morphology have focused on mycelia.
Let’s consider how biologists are working to make sense of all
this diversity, beginning with an overview of fungal morphology. The Nature of the Fungal Mycelium If food sources are plenti-
ful, mycelia can be long lived and grow to be extremely large. In
Analyzing Morphological Traits 2000, U.S. Forest Service researcher Catherine Parks and her
colleagues in Oregon discovered a mycelium of the fungus Armil-
Compared with animals and land plants, fungi have simple
laria growing beneath 9 km2 of soil—an area substantially larger
bodies. Only two growth forms occur among them: (1) single-
than most college campuses. The biologists estimated the indi-
celled forms called yeasts (Figure 29.4a), and (2) multicellular,
vidual’s weight at hundreds of tonnes and its age at thousands of
filamentous structures called mycelia (singular: mycelium;
years, making it one of the largest and oldest organisms known.
Figure 29.4b). Many species of fungus grow either as yeasts or
Although most mycelia are much smaller and shorter lived
as a mycelium, but some regularly adopt both growth forms.
than the individual in Oregon, all mycelia are dynamic. Mycelia
constantly grow in the direction of food sources and die back in
(a) Single-celled fungi are called yeasts.
areas where food is running out. The body shape of a fungus can
change almost continuously throughout its life.

Eye of Science/Photo Researchers, Inc./Science Source

The Nature of Hyphae The filaments making up a mycelium are
called hyphae (singular: hypha). As Figure 29.5a shows, hyphae
are long and narrow.
In most terrestrial fungi, each filament is divided into cells by
cross-walls called septa (singular: septum; Figure 29.5b). Septa
do not close off the cells along hyphae completely. Instead, gaps
called pores enable a wide variety of materials, even nuclei and
other organelles, to flow from one cell-like compartment to the
next.
5 om Some fungal lineages have hyphae that are coenocytic
(“common celled”; pronounced see-no-SIT-ick)—meaning that
they are not divided into separate cells; they lack septa entirely
(b) Multicellular fungi have web-like bodies called mycelia. (Figure 29.5c). Coenocytic fungi have hundreds or thousands of
nuclei scattered throughout the mycelium.
Fungi lack complex, long-distance transport systems like
those found in plants and animals. One advantage of having
pores in septa (or no septa) is that nutrients can move rapidly
from regions of uptake to regions of mycelial growth. Because
nutrients and some organelles can flow through the entire myce-
lium—at least to a degree—the fungal mycelium is intermediate
between a multicellular land plant or animal and an enormous
Hecker/Sauer/AGE Fotostock

single-celled organism.
Some fungi produce hyphae with unique adaptations for prey
capture and nutrient uptake. For example, some are able to cap-
ture microscopic nematodes (roundworms) by means of sticky
substances on their cell walls, or in snares consisting of looped
20 mm hyphae (Figure 29.6). Once a worm is captured, hyphae invade
its body, digest it, and absorb the nutrients that are released.
Figure 29.4 Fungi Have Just Two Growth Forms. Fungi grow
(a) as single-celled yeasts and/or (b) as multicellular mycelia made Mycelia Have a Large Surface Area It’s important to appreciate
up of long, thin, highly branched filaments. The scanning electron just how thin hyphae are. Plant root tips are typically about 1 mm
micrograph in (a) is colourized. in diameter, but fungal hyphae are typically less than 10 μm in
CAUTION People sometimes mistakenly refer to yeasts as diameter, or 1/100th the width of a root tip. Fungal mycelia can
bacteria. What is a fundamental difference between yeasts (which penetrate tiny fissures in soil and absorb nutrients that are inac-
are eukaryotic) and bacteria (which are prokaryotic)? cessible to plant roots.

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 (a) Both the reproductive structure and mycelium are
composed of hyphae. Hyphae
Nematode

Susumu Nishinaga/Photo Researchers, Inc./Science Source

Dense hyphae

Reproductive
structure

N. Allen and G. Barron

50 om
Mycelium

Figure 29.6 Some Fungi Are Predators. This nematode-trapping

Spaced hyphae 100 om fungus (Arthrobotrys anchonia) forms specialized hyphae with
lasso-like traps.

(b) Most hyphae are divided into compartments by septa. Perhaps the most important aspect of mycelia and hyphae,
however, is their overall shape. Because mycelia are composed
of complex, branching networks of extremely thin hyphae, fungi
Septa
have the highest surface-area-to-volume ratios observed in mul-
Photo Researchers, Inc./Science Source ticellular organisms and are therefore the best at absorption.
To drive this point home, consider that the hyphae found
in any fist-sized ball of rich soil typically have a surface area
Pore equivalent to half a page of this book. Because of their large sur-
face area, fungi are extremely efficient at absorbing nutrients.
The extraordinarily high surface area of a mycelium has a
downside, however. The amount of water that evaporates from
George L. Barron

an organism is a function of its surface area—meaning that fungi

are prone to drying out. As a result, fungi are most abundant in
25 om 1 om moist habitats.
When conditions dry, the fungal mycelium may die back par-
tially or completely. Reproductive cells called spores that are
(c) Coenocytic hyphae consist of multinucleate cells. produced by sexual or asexual reproduction are resistant to dry-
ing, however. As a result, spores can endure dry periods and then
germinate to form a new mycelium when conditions improve.
Nuclei Mycelial growth is dynamic, changing with moisture availability
and food supply.
Patrick Hickey/NIPHT LIMITED

The oldest known fungi are 440 million year old microfossils
found in Sweden, Scotland, and New York. Although these tiny
fossilized fungi are fragments shorter than the width of a human
hair, they display the branching that is characteristic of mycelial
networks.

25 om
Reproductive Structures Mycelia are an adaptation that sup-
ports external digestion and the absorptive lifestyle of fungi.
Figure 29.5 Multicellular Fungi Have Unusual Bodies. Many fungi also produce dense, fleshy, multicellular struc-
(a) The feeding portion of a fungus is a mycelium, which is made tures—such as mushrooms and puffballs—that do not absorb
up of hyphae. In some species, hyphae are densely clustered, food. Instead, they function in reproduction. Typically they are
forming multicellular, fleshy structures such as mushrooms, the only part of a fungus that is exposed to air, where drying is
brackets, or morels that emerge from the ground. (b) Hyphae a potential problem. The mass of hyphal filaments on the inside
of some terrestrial fungi are divided into cell-like compartments
of mushrooms are protected from drying by the densely packed
by partitions called septa that contain pores. As a result, the
hyphae forming the surface (see the micrograph in Figure 29.5a).
cytoplasm of different compartments is continuous. (c) Hyphae of
some coenocytic fungi, which lack septa entirely, are composed In many fungi, including some entire lineages, sexual repro-
of giant, multinucleate cells. duction has never been observed. Among those lineages of fungi
that do reproduce sexually, important morphological differences

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 (a) Swimming gametes and (b) Zygosporangia: spore- (c) Basidia: club-shaped (d) Asci: sac-like cells where
spores producing structures formed cells where meiosis occurs, meiosis and one round of mitosis
when hyphae are yoked forming 4 spores occur, forming 8 spores

Gametes
Zygosporangium
Spore Spores

Ascus
Hypha

Flagella Hypha Basidium Hypha

Spores

Biophoto Associates/Photo Researchers,

Ed Reschke/Photolibrary/Getty Images

Inc./Science Source

Nino Santamaria
Melvin Fuller

1 om 25 om 20 om 50 om

Figure 29.7 Four Types of Sexual Reproductive Structures Are Observed in Fungi. The light and dark dots in
the illustrations represent nuclei.

are seen. Most fungal species that undergo sexual reproduction conidia by the trillions (Figure 29.8). Conidia can be dispersed
produce one of four types of distinctive reproductive structures: by water or wind currents and grow into new hyphae when
1. Swimming gametes and spores In certain species that conditions are right. Mould, for example, spreads rapidly across
live primarily in water or wet soils, the gametes produced
during sexual reproduction have flagella, as do the spores
produced during asexual reproduction (Figure 29.7a). Penicillium roquefortii
These are the only motile cells known in fungi. Species
with swimming gametes are traditionally known as chytrids
(pronounced KI-trids).
Conidia
2. Zygosporangia In some species, haploid hyphae from two
individuals meet and become joined, like oxen with a yoke,
as shown in Figure 29.7b. Cells from yoked hyphae fuse to

E. Gueho/Photo Researchers, Inc./Science Source

form a distinctive spore-producing structure called a zygo-
sporangium (plural: zygosporangia; the Greek root zygo
means “to be yoked”[see BioSkills 13]). Species with a zygospo-
rangium are traditionally known as zygomycetes.
3. Basidia Mushrooms, brackets, and puffballs form special-
ized club-shaped cells at the ends of hyphae called basidia
(singular: basidium; “little club”). Each basidium produces
four spores by meiosis (Figure 29.7c). Species with basidia
are traditionally called basidiomycetes or “club fungi.” WebCat/Shutterstock

4. Asci Cups, morels, and some other types of fungi form spe- 20 om
cialized sac-like cells called asci (singular: ascus) at the tips
of hyphae. Within each ascus, meiosis and one round of mito- Figure 29.8 Asexual Spores. Penicillium roquefortii is the dark-
sis produces eight spores (Figure 29.7d). Species with asci are coloured fungus used in the production of blue cheese. A close-up
traditionally known as ascomycetes or “sac fungi.” of the fungus reveals asexual spores (conidia).
Many fungi are quite good at asexual reproduction as well Are conidia produced from a single fungus genetically identical
as sexual reproduction. Some produce asexual spores called or different? Explain.

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 decaying food and other items by producing lots and lots of and humans shared a common ancestor relatively recently. As a
asexual spores. result, their enzymes and cell components are similar in struc-
In sum, morphological studies allowed biologists to describe ture and function. Drugs that disrupt fungal enzymes and cells
and interpret the mycelial growth habit as an adaptation that are also likely to damage humans.
makes external digestion and nutrient absorption extremely effi- In addition to DNA sequence data, three key morphological
cient. Careful analyses of morphological features also allowed traits link animals and fungi:
researchers to identify four major types of sexual reproductive 1. Many animals and most fungi synthesize the tough structural
structures. material called chitin (see Chapter 5). Chitin is a prominent
Now the question is, within the Fungi, do species that produce component of the cell walls of fungi.
either swimming gametes and spores, zygosporangia, basidia,
or asci each form monophyletic groups—meaning that these dis- 2. The flagella that develop in chytrid spores and gametes are
tinctive reproductive structures evolved just once? Also, which similar to those observed in animals: As in animals, the fla-
eukaryotes are most closely related to fungi? These questions gella in chytrids are single, are located at the back of repro-
and more are explored in the next section. ductive cells, and move in a whip-like manner.
3. Both animals and fungi store food by synthesizing the poly-
Evaluating Molecular Phylogenies saccharide glycogen. (Green plants, in contrast, synthesize
starch as their storage product.)
Researchers have sequenced and analyzed an array of genes
and genomes to establish where fungi fit on the tree of life. In
What Are the Relationships among the Major Fungal Groups? To
fact, the first complete genome sequence of a eukaryotic species
understand the relationships among species with swimming
was that of a fungus, the model organism baker’s yeast, Sac-
gametes and spores, zygosporangia, basidia, or asci, biologists
charomyces cerevisiae (see BioSkills 9). The phylogenetic position
have sequenced a series of genes from an array of fungal spe-
of the Fungi is now well established; the position of lineages
cies and used the data to estimate the phylogeny of the group.
within the Fungi is still the subject of intense research.
The results, shown in Figure 29.10 on page 623, support several
important conclusions:
Fungi Are Closely Related to Animals Figure 29.9 shows that
fungi are much more closely related to animals than they are • The single-celled, parasitic eukaryotes called microsporidians
to land plants. The close evolutionary relationship between are fungi.
fungi and animals explains why fungal infections in humans Interpretation: They are not a distantly related sister group to
are much more difficult to treat than bacterial infections. Fungi fungi, as initially thought. This point is important. Research-
ers are now testing the hypothesis that fungicides—­
substances that can kill fungi or slow their growth—can cure
Bacteria microsporidian infections in bee colonies, silkworm colonies,
and AIDS patients.
Archaea
• The chytrids and zygomycetes are poorly resolved. The order
of branching events among lineages with these reproductive
Amoebozoa
structures is still not known (in Figure 29.10, they are col-
lapsed into a polytomy; see Chapter 25).
Fungi
Interpretation: Swimming gametes and the zygosporangium
Choanoflagellates evolved more than once. Or, both structures were present in a
Fungi and common ancestor but then were lost in certain lineages.
Animals
animals • An important group called the Glomeromycota is monophyletic.
share
a common Excavata Interpretation: The adaptations that allow glomeromycete
ancestor at species to live in association with plant roots as mycorrhizae
this point (discussed in Section 29.3) evolved once.
Glaucophyte algae
• The basidiomycetes are monophyletic—they form a lineage
Red algae
called Basidiomycota, or club fungi.
Green algae Green Interpretation: The basidium evolved once.
plants • The ascomycetes are monophyletic—they form a lineage
Land plants
called Ascomycota, or sac fungi.
Other protists Interpretation: The ascus evolved once.

Figure 29.9 Fungi Are More Closely Related to Animals than • Together, the Basidiomycota and Ascomycota form a mono-
to Land Plants. This phylogenetic tree shows the evolutionary phyletic group.
relationship of the Fungi to other major lineages. (Recall that Interpretation: Because basidiomycetes and ascomycetes both
choanoflagellates are solitary or colonial protists found in freshwater; form septate hyphae and large “fruiting” structures, this
see Chapter 27.) growth habit evolved once.

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 Choanoflagellates Although recent advances have been made in understanding
Animals the evolutionary history of fungi, the phylogenetic tree in Figure
29.10 is still a work in progress. For example, it is not yet clear
MICROSPORIDIA where microsporidians should be placed relative to several lin-
eages of chytrids and zygomycetes. (Microsporidians lack both
CHYTRIDS and ZYGOMYCETES swimming gametes and zygosporangia.) Future work should
clarify exactly how fungi diversified.

Polytomy Check Your Understanding

If you understand that …
Make chytrid-like motile • The bodies of fungi are either single-celled yeasts or multicellular
gametes and spores mycelia.
or • During sexual reproduction, different groups of fungi produce
Make zygosporangium distinct reproductive structures.
with tough outer coat • Some fungi produce vast numbers of asexual spores called
conidia.

You should be able to …

FUNGI 1. Explain why mycelia are interpreted as an adaptation to an
absorptive lifestyle.
2. Identify the four types of sexual reproductive structures ob-
served in fungi.
GLOMEROMYCOTA 3. Explain how mould on a single piece of fruit can easily spread
Monophyletic
Make spore with to nearby fruit.
tough outer coat
Answers are available in Appendix A.
BASIDIOMYCOTA

29.3 What Themes Occur in the

Monophyletic
Make club-like
basidium
Diversification of Fungi?
Why are there so many different species of fungi? This question
is particularly puzzling given that fungi share a common attri-
Monophyletic ASCOMYCOTA bute: They all make their living by absorbing food directly from
their surroundings. In contrast to the diversity of food-getting
strategies observed in bacteria, archaea, and protists, all fungi
make their living in the same basic way. In this respect, fungi
Make sac-like ascus
are like plants—virtually all of which make their own food via
photosynthesis.
The diversification of land plants was driven not by novel
ways of obtaining food, but by adaptations that allowed plants
to grow and reproduce in a diverse array of terrestrial habitats
(see Chapter 28). Recall that associations with animal pollina-
Figure 29.10 Phylogeny of the Fungi. A phylogenetic
tree based on analyses of DNA sequence data. The icons tors were especially important in the diversification of flowering
represent the types of sexual reproductive structures observed plants. What drove the diversification of fungi? The answer is
in each major lineage. the evolution of novel methods for absorbing nutrients from a
SOURCE: Based on James T. Y., F. Kauff, C. L. Schoch, et. al. 2006. Reconstructing the diverse array of food sources.
early evolution of fungi using a six-gene phylogeny. Nature 443: 818–822, Fig. 1.
This section introduces a few of the ways that fungi go about
absorbing nutrients from different food sources, as well as how
they produce offspring. Let’s explore the diversity of ways that
• The sister group to fungi comprises animals plus protists
fungi do what they do.
called choanoflagellates.
Interpretation: Because choanoflagellates and the most
ancient groups of animals are aquatic, and because chytrids Fungi Often Participate in Symbioses
are aquatic, it is reasonable to hypothesize that the earli- The first land plants in the fossil record are closely associated
est fungi were aquatic and that the switch to terrestrial life with fungal fossils; the ability to absorb nutrients from fungi
occurred early in the evolution of the Fungi. may have been crucial in the early evolution of land plants.

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 Close associations between land plants and fungi continue today. of plant–mycorrhizal interactions are particularly common,
Researchers estimate that over 90 percent of land plants live in involving ectomycorrhizal fungi (EMF) and arbuscular
physical contact with fungi. Stated another way, fungi and land mycorrhizal fungi (AMF). The two major types of mycor-
plants often have a symbiotic (“together living”) relationship. rhizae have distinctive morphologies, as shown in Figure 29.12
Although some species of fungi live in association with an on page 626, as well as distinctive functions and geographic
array of different land plant species, some fungi–plant associa- distributions.
tions are very specific. It is not uncommon for one fungal species
to live in only a particular type of tissue, in one plant species. Ectomycorrhizal Fungi (EMF) EMF like the one shown in Figure
Scientists categorize these symbiotic relationships as mutu- 29.12a are found on many tree species in the temperate regions
alistic if they benefit both species, parasitic if one species ben- of the world, where warm summers alternate with cold winters
efits at the other’s expense, or commensal if one species benefits (see Chapter 49). In this type of association, hyphae form a dense
while the other is unaffected. However, it is sometimes difficult network that covers a plant’s root tips. Individual hyphae pen-
to know if a species really benefits from the relationship. etrate between cells in the outer layer of the root, but hyphae
To understand the nature of the association, biologists have do not enter the root cells. The Greek root ecto, which means
turned to experimental approaches. In these experiments, treat- “outer,” describes this association accurately: The fungi form
ments lacking fungal symbionts are created by sterilizing soils an outer sheath on root tips that is often 0.1 mm thick. Hyphae
with heat or by treating soils and seeds with fungicides. Pres- also extend out from the sheath-like portion of the mycelium
ence–absence experiments have generally shown that plants into the soil.
grow much larger with their normal symbiotic fungi than they How do these trees and fungi interact, once they start liv-
do without (see Figure 29.2b). Similarly, fungi that are typically ing together? In the habitats where EMF are abundant, nitro-
symbiotic are usually unable to grow and reproduce if their regu- gen atoms—for example, in the form of amino acids—tend
lar host plant is absent. to remain in dead tissues instead of dispersing into the soil.
To explore the nature of fungi–plant symbioses in detail, The hyphae of EMF penetrate decaying material and release
researchers have used isotopes as tracers for specific elements. enzymes called peptidases that cleave the peptide bonds
For example, to test the hypothesis that fungi obtain food in the between amino acids in dead tissues. The amino acids released
form of carbon-containing compounds from their plant associ- by this reaction are absorbed by the hyphae and transported
ates, biologists introduced radioactively labelled carbon dioxide to spaces between the root cells of trees, where they can be
into the air surrounding plants that did or did not contain sym- absorbed by the plant.
biotic fungi (see Figure 29.11). They incorporated labelled CO2 EMF also can acquire phosphate ions that are bound to soil
molecules into the sugars produced during photosynthesis and particles and transfer the ions to host plants. The fungi receive
then followed the location of the radioactive atoms over time sugars and other complex carbon compounds from the tree.
by using a device that detects radioactivity. If plants feed their Researchers have found that when birch tree seedlings are
fungal symbionts, then labelled carbon compounds should be grown with and without their normal EMF in pots filled with
transferred from the plant to the fungi. forest soil, only the seedlings with EMF are able to acquire
To further test the hypothesis that plants are receiving nutri- significant quantities of nitrogen and phosphorus. Inspired by
ents from their symbiotic fungi, American biologist Heike Bück- such data, one biologist has referred to ectomycorrhizae as the
ing and her colleagues added radioactive phosphorous atoms “dominant nutrient-gathering organs in most temperate forest
or the heavy isotope of nitrogen (15N) to potted plants that did ecosystems.”
or did not contain symbiotic fungi (see Figure 29.11). If fungi The hyphae of EMF are like an army of miners that discover,
facilitate the transfer of nutrients from soil to plants, then plants excavate, and deliver precious nuggets of nitrogen and phospho-
grown in the presence of their symbiotic fungi should receive rus to trees. The productivity of the world’s most important com-
much more of the radioactive phosphorus or heavy nitrogen than mercial forests depends on EMF.
plants grown in the absence of fungi. Ectomycorrhizal fungi are important mutualists in Canada’s
Other experiments with isotopes used as tracers have shown vast boreal forests and play a vital role in regeneration of forest
that sugars and other carbon-containing compounds produced stands following clearcutting. University of British Columbia
by plants via photosynthesis are transferred to their fungal sym- Okanagan botanist Melanie Jones found that the diversity of
bionts. In some cases, as much as 20 percent of the sugars pro- ectomycorrhizal fungi on a clearcut site is important. When the
duced by a plant end up in their symbiotic fungi. In addition, the roots of conifer seedlings are associated with several different
symbiotic fungi facilitate the transfer of phosphorus or nitro- types of EMF, tree seedlings are more likely to be able to obtain a
gen—or both—from soil to the plant. variety of nutrients as soil conditions change.
Phosphorus and nitrogen are in short supply in many envi- Conifers are linked by a complex mycorrhizal network of fun-
ronments. Thus, the role of symbiotic fungi in transferring ions gal connections. Each tree is associated with many individual
and other nutrients that contain these atoms has likely been fungal hyphae from hundreds of species, while a fungal myce-
critical to the evolutionary success of plants. lium from a generalist fungal species may be associated with
Not long after associations between fungi and the roots of the roots of several trees. Mycorrhizal fungi obtain sugars from
land plants were discovered, researchers found that two types healthy, growing host plants, as shown in Figure 29.11, but what

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 RESEARCH

QUESTION: Are mycorrhizal fungi mutualistic?

HYPOTHESIS: Host plants provide mycorrhizal fungi with sugars and other photosynthetic products. Mycorrhizal fungi provide host plants with
phosphorus and/or nitrogen from the soil.

NULL HYPOTHESIS: No exchange of food or nutrients occurs between plants and mycorrhizal fungi. The relationship is not mutualistic.

EXPERIMENTAL SETUP:

Labelled carbon treatment: Labelled carbon control: Labelled P or N treatment: Labelled P or N control:
Labelled CO2 Labelled CO2
added to air around plant added to air around plant

Mycorrhizal No fungi No
fungi present present Fungi fungi

Labelled P or N added to soil Labelled P or N added to soil

PREDICTION FOR LABELLED CARBON: A large percentage of the PREDICTION FOR LABELLED P OR N: A large percentage of the labelled
labelled carbon taken up by the plant will be transferred to mycorrhizal P or N taken up by the fungi will be transferred to the plant. In the
fungi. In the control, little labelled carbon will be present in the soil control, little or no labelled P or N will be taken up by the plant.
surrounding the roots.
PREDICTION OF NULL HYPOTHESIS, LABELLED P OR N: There will
PREDICTION OF NULL HYPOTHESIS, LABELLED CARBON: There will be no difference between amounts of labelled P or N found in the plant
be no difference in the localization of carbon in the two treatments. in the presence or absence of fungi.

RESULTS:

Large amounts of labelled P or N Little labelled P or N

are found in host plant is found in host plant

No
Fungi fungi No
Fungi fungi
Up to 20% of labelled CO2 Little to no labelled carbon
taken up by plant is transferred is found in soil
to mycorrhizal fungi

CONCLUSION: The relationship between plants and mycorrhizal fungi is mutualistic. Plants provide mycorrhizal fungi with carbohydrates.
Mycorrhizal fungi supply host plants with nutrients.

Figure 29.11 Experimental Evidence that Mycorrhizal Fungi and Plants Are Mutualistic. Sugars flow from
plants to mycorrhizal fungi; key nutrients flow from mycorrhizal fungi to plants.
SOURCES: Based on Finlay, R. D., and B. Söderström. 1992. Mycorrhiza and carbon flow to the soil. In Mycorrhizal Functioning. An Integrative Plant-Fungal Process. Ed.
M. J. Allen. Chapman and Hall, New York, 134–160. Bücking, H., and W. Heyser. 2001. Microautoradiographic localization of phosphate and carbohydrates in mycorrhizal
roots of Populus tremula: Populus alba and the implications for transfer processes in ectomycorrhizal associations. Tree Physiology 21: 101–107.

PROCESS OF SCIENCE What do these “labelled nutrient” experiments tell you that you didn’t already know from
experiments like that shown in Figure 29.2, where plants are grown with and without mycorrhizae?

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 (a) Ectomycorrhizal fungi (EMF) form sheaths (b) Arbuscular mycorrhizal fungi (AMF)
around roots and penetrate between root cells. contact plasma membranes of root cells.

20 om 10 om

Brundrett, Mark
EMF AMF

Hyphae extend Hyphae extend

outward outward
into the soil into the soil

Hyphae extend inward, Hyphae extend inward,

between cells penetrate cell wall, and
contact plasma membrane

Hyphae form a dense,

continuous sheath
around root

Figure 29.12 Mutualistic Fungi Interact with the Roots of Plants in Two Distinct Ways. (a) Ectomycorrhizal fungi (EMF) form a dense
network around the roots of plants. Their hyphae penetrate the intercellular spaces of the root but do not enter the root cells. (b) The hyphae of
arbuscular mycorrhizal fungi (AMF) penetrate the walls of root cells, where they branch into bushy structures or balloon-like vesicles that are in
close contact with the root cell’s plasma membrane.

happens when host plants are stressed by drought or attacked as do EMF. Instead, they form a pipeline extending from inside
by defoliating insects? As Figure 29.13 in Canadian Research plant roots into the soil well beyond the root.
29.1 shows, University of British Columbia botanist Suzanne AMF are found in a whopping 80 percent of all land plant spe-
Simard and her colleagues investigated this question, which is cies. They are particularly common in grasslands and in tropical
vitally important to Canadian forests stressed by the warm, dry forests. They are also widespread in temperate climates.
weather that comes with climate change and the associated out- What do AMF do? Plant tissues decompose quickly in the grass-
breaks of forest pests. lands and tropical forests where AMF flourish, because the grow-
ing season is long and warm. As a result, nitrogen is often readily
Arbuscular Mycorrhizal Fungi (AMF) In contrast to the hyphae available to plants. Phosphorus is usually in short supply, though,
of EMF, which grow around and between root cells, the hyphae because it tends to leach out of soils that experience high rainfall.
of arbuscular mycorrhizal fungi (AMF) grow into the cells of root
tissue without penetrating cell membranes. The name arbus- Endophytes Mycorrhizae aren’t the only type of symbiotic fungi
cular (“little tree”) was inspired by the bushy, highly branched found in plants. Researchers have also become interested in
hyphae, shown in Figure 29.12b, that form between the cell fungi that are endophytes (“inside plants”)—organisms that live
walls and the plasma membrane of root cells. AMF are also called between and within plant cells. Endophytic fungi live in close
endomycorrhizal fungi, because they grow inside root cell association with the roots or the aboveground tissues (leaves and
walls. stems) of land plants.
The key point is that the hyphae of AMF penetrate the cell Although endophytic fungi were unknown before the 1940s,
wall and make direct contact with the plasma membrane of root they are turning out to be both extremely common and highly
cells. The highly branched hyphae inside the plant cell wall of diverse:
AMF are thought to be an adaptation that increases the surface • Biologists in Brazil are examining tree leaves for the presence
area available for exchange of molecules between the fungus and of fungi; each time they do, they are discovering several new
its host. However, AMF do not form a tight sheath around roots, species of endophytes.

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 Canadian Research 29.1 Photosynthetic Carbon Is Transferred from Stressed to Healthy Plants through
Ectomycorrhizal Networks

RESEARCH

QUESTION: Do ectomycorrhizal fungi transfer photosynthetic carbon from defoliated Douglas-fir seedlings to
healthy ponderosa pine seedlings?

HYPOTHESIS: Manual defoliation of Douglas-fir seedlings will result in transfer of labelled photosynthetic carbon to neighbouring ponderosa pine
seedlings via networks of ectomycorrhizal fungi.
NULL HYPOTHESIS: Manual defoliation of Douglas-fir seedlings will not result in transfer of labelled photosynthetic carbon to neighbouring
ponderosa pine seedlings via networks of ectomycorrhizal fungi.

EXPERIMENTAL SETUP: Forty pairs of Douglas-fir and ponderosa pine seedlings were grown in pots with 800 grams of soil and watered once a
week. Seedlings were colonized by Wilcoxina rehmii, a generalist fungus that forms ectomycorrhizae with both species of host trees.
Ponderosa pine seedlings were grown in two treatments: (1) in a 35 µm mesh bag that allowed ectomycorrhizal fungal hyphae from the
Douglas-fir seedling to come in contact with ponderosa plant roots but prevented contact between the roots of the two seedlings and (2) in a
0.5 µm mesh bag that allowed movement of water and solutes but prevented formation of an ectomycorrhizal fungal network shared by the two
seedlings. Douglas-fir seedlings were either (1) left intact or (2) manually defoliated with removal of all needles.
Douglas-fir seedlings were exposed to radioactively labelled CO2. Six days later, all seedlings were harvested and ponderosa pine seedlings
were assessed to see if they had taken up labelled carbon.

PREDICTION: Labelled photosynthetic carbon will be transferred from manually defoliated Douglas-fir seedlings to neighbouring ponderosa pine
seedlings only when the conifer seedlings share an ectomycorrhizal fungal network.
PREDICTION OF NULL HYPOTHESIS: Labelled photosynthetic carbon will not be transferred from manually defoliated Douglas-fir seedlings to
neighbouring ponderosa pine seedlings, even when the conifer seedlings share an ectomycorrhizal fungal network.

RESULTS:
0.024
Ponderosa pine needles
uptake by ponderosa pine

0.02
Ponderosa pine roots
(in mg equivalents of 12C)

0.016

0.012

0.008

0.004
13C

-0.004 35 om 0.5 om 35 om 0.5 om

mesh mesh mesh mesh
Undefoliated Manually defoliated
Douglas-fir Douglas-fir
Treatments
Figure 29.13 Ectomycorrhizal fungi transfer carbon from stressed Douglas-fir trees to their neighbours.

CONCLUSION: Following manual defoliation, Douglas-fir seedlings export carbon-containing compounds to their roots. These compounds are
then transported to neighbouring ponderosa pine seedlings through the mycorrhizal network shared by both seedlings. Mycorrhizal transfer of
carbon-containing compounds from stressed to healthy plants may play a key role in recovery of forests from damage due to abiotic factors
such as drought or high temperature.

SOURCE: Based on Song, Y. Y., S. W. Simard, A. Carroll, W. W. Mohn, and R. S. Zeng. 2015. Defoliation of interior Douglas-fir elicits carbon transfer and stress signaling
to ponderosa pine neighbors through ectomycorrhizal networks. Scientific Reports 5: 8495–8504.

Think About It: How do mycorrhizal fungi benefit from transferring organic compounds from stressed
Douglas-fir seedlings to neighbouring ponderosa pine seedlings?

• Recall from Section 29.2 that a study in Panama found hun- Recent research has shown that some endophytes increase the
dreds of fungal species living in the leaves of just two tree spe- drought tolerance of their host plants. Other endophytes, found in
cies. These fungi are all endophytes. some grasses and morning glories, also produce compounds that

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 (a) Lichen on a tree branch (b) Cross section through a lichen
Ralf Wagner

Dense hyphae

Green algae
Marko König/imagebroker/AGE Fotostock

Loose hyphae

Dense hyphae

25 om

Figure 29.14 Lichens Are Symbiotic Associations between a Fungus and a Cyanobacterium or Green Alga.
(a) Lichens often colonize surfaces, such as tree bark or bare rock, where other organisms are rare. (b) In a lichen,
cyanobacteria or green algae live within a dense network of fungal hyphae.

benefit plants by deterring or even killing herbivores. Endophytes the “zombie ant” (Figure 29.15). From the high perch atop the
have been shown to benefit from these symbioses, too, by absorb- plant, the fungal spores are readily dispersed.
ing sugars from the plants. Fungal endophytes also help protect
forest trees. An endophyte living on white spruce trees makes the
spruce needles less palatable to spruce budworm, a major forest
What Adaptations Make Fungi
pest in Canada’s eastern forests. Another endophyte species pro- Such Effective Decomposers?
tects white pine against blister rust disease, a fungal disease. The saprophytic fungi are master recyclers. Although bacteria
Based on these results, biologists have concluded that the and archaea are also important decomposers in terrestrial envi-
relationship between endophytes and some plants is mutualistic. ronments, some fungi and a few bacterial species are the only
In other types of plants, however, researchers have not been able organisms that can digest wood completely. Given enough time,
to document benefits for the plant host. The current consensus is fungi can turn even the hardest, most massive trees into soft soils.
that at least some endophytic fungi may be commensals—mean- How do fungi do it? You’ve already been introduced to two key
ing the fungi and the plants simply coexist with no observable adaptations:
effect, either bad or good, on the host plant.
• The large surface area of a mycelium makes nutrient absorp-
The take-home message about mycorrhizae and other endo-
tion exceptionally efficient.
phytes is that most plants are covered with fungi, inside and out,
from the ends of their branches to the tips of their roots. Through- • Saprophytic fungi can grow toward the dead tissues that sup-
out their lives, many or even most plants are involved in several dis- ply their food.
tinct types of commensal or mutualistic relationships with fungi. What other adaptations help fungi decompose plant tissues?

Other Symbioses Fungi also take up residence with species

other than land plants. The many unique properties of fungi
have made them interesting partners in a wide array of symbiotic
relationships: Ophiocordyceps

Thailand Wildlife/Alamy Stock Photo

fungus reproductive
• Many plant-eating insects harbour diverse arrays of yeast spe- structures
cies in their guts, which may aid digestion or detoxification of
plant compounds.

• Lichens are a mutualistic partnership usually between a

species of ascomycete and either a cyanobacterium or an alga
(Figure 29.14).
• Some fungi that parasitize insects dramatically change their
hosts’ behaviour. For example, the fungus Ophiocordyceps
causes ants to follow a zombie-like sequence of motions: An Figure 29.15 “Zombie Ant” Fungus. After being infected by
infected ant climbs to the top of a plant, bites down hard on the Ophiocordyceps fungus, an ant will climb a plant, bite down
the stem or leaf, and eventually dies. The fungus then pro- hard, and soon die. The fungus sprouts from the ant’s head and
duces a reproductive structure that sprouts from the head of releases spores.

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 Extracellular Digestion Large molecules such as starch, lig- to metabolize, and as a result they accumulate in soil. If wood-
nin, cellulose, proteins, and RNA cannot diffuse across plasma rotting fungi don’t use lignin as food, why do they produce
membranes. Only sugars, amino acids, and other small mol- enzymes to digest it? The answer is simple. In wood, lignin
ecules can enter the cytoplasm of cells through membrane forms a dense matrix around long strands of cellulose. Degrad-
carrier proteins or transporters (see Chapter 6). As a result, ing the lignin matrix gives hyphae access to huge supplies of
fungi, like most animals, have to digest their food before they energy-rich cellulose. Similar to the mycorrhizal fungi dis-
can absorb it. cussed earlier, saprophytic fungi are like miners. But instead
Fungi perform extracellular digestion—digestion that takes of seeking out rare, gem-like nitrogen or phosphorous atoms,
place outside the organism. The simple compounds that result the saprophytes use lignin peroxidase to blast away enormous
from enzymatic action are then absorbed by the hyphae. Most lignin molecules, exposing rich veins of cellulose that can fuel
animals also perform extracellular digestion, but it takes place in growth and reproduction.
a digestive tract (see Chapter 41).
The two most abundant organic molecules on Earth—lignin Cellulose Digestion Once lignin peroxidase has softened
and cellulose—are digested by fungi: wood by stripping away its lignin matrix, the long strands
• Lignin is an extremely strong, complex polymer built from of cellulose that remain can be attacked by enzymes called
monomers that contain six-carbon rings. Recall that most cellulases.
lignin is found in the secondary cell walls of plant vascular Like lignin peroxidase, cellulases are secreted into extracel-
tissues, where it provides structural support (see Chapter 28). lular spaces by fungi. But unlike the uncontrolled chain reaction
catalyzed by lignin peroxidase, degradation of cellulose by cel-
• Cellulose is a polymer of glucose and is found in the primary
lulases is extremely predictable.
and secondary cell walls of all plant cells.
Biologists have purified seven different cellulases from the
Some basidiomycetes can degrade lignin and digest cellulose fungus Trichoderma reesei. Two of these enzymes catalyze a criti-
completely—to CO2 and H2O. Let’s look at how they do it. cal early step in digestion—they cleave long strands of cellulose
into a disaccharide called cellobiose. The other cellulases are
Lignin Degradation Biologists have been keenly interested in equally specific and also catalyze hydrolysis reactions. In com-
understanding how basidiomycetes digest lignin. Paper manu- bination, the suite of seven enzymes in T. reesei transforms long
facturers are also interested in this process because they need strands of cellulose into a simple monomer—glucose—that the
safe, efficient ways to degrade lignin in order to make soft, absor- fungus can absorb and use as a source of food.
bent paper products.
To find out how lignin-digesting fungi do it, biologists began
analyzing the proteins that these species secrete into extracel-
Variation in Reproduction
lular spaces. After purifying these molecules, the investigators Recall from Section 29.2 that fungi may produce swimming
tested each protein for the ability to degrade lignin. Using this gametes and spores, yoked hyphae in which nuclei from dif-
approach, investigators from two labs independently discovered ferent individuals fuse to form a zygosporangium inside a pro-
an enzyme called lignin peroxidase. tective structure, or specialized spore-producing cells called
Lignin peroxidase catalyzes the removal of a single electron basidia and asci. As fungi diversified, in addition to adapting
from an atom in the ring structures of lignin. This oxidation step to a variety of food sources, they evolved an array of ways to
creates a free radical—an atom with an unpaired electron (see reproduce.
Chapter 2). This is an extremely unstable electron configuration,
and it leads to a series of uncontrolled and unpredictable reac- Spores as Key Reproductive Cells The spore is the most funda-
tions that split the polymer into smaller units. mental reproductive cell in fungi. Spores are the dispersal stage
Biologists have referred to this mechanism of lignin degrada- in the fungal life cycle and are produced during both asexual and
tion as enzymatic combustion. The phrase is apt: The uncon- sexual reproduction. (Recall from Chapter 13 that asexual repro-
trolled oxidation reactions triggered by lignin peroxidase are duction is based on mitosis, while sexual reproduction is based
analogous to the uncontrolled oxidation reactions that occur on meiosis.) Fungi produce spores in such prodigious quanti-
when gasoline burns in a car engine. ties that it is not unusual for fungal spores to outnumber pollen
The unpredictable nature of the reaction is remarkable, grains in air samples.
because virtually all of the other reactions catalyzed by enzymes If a spore falls on a food source and is able to germinate, a
are extremely predictable. The lack of predictability here makes mycelium begins to form. As the fungus expands, hyphae grow in
sense, however. Unlike proteins, nucleic acids, and most other the direction in which food is most abundant. But if food begins
polymers with a regular and predictable structure, lignin is to run out, mycelia respond by making spores, which are dis-
extremely heterogeneous. Over 10 types of covalent linkages are persed by wind or animals.
routinely found between the monomers that make up lignin. But Why would mycelia reproduce when food is low? The lead-
once lignin peroxidase has created a free radical in the ring struc- ing hypothesis is that spore production allows starving mycelia
ture, any of these linkages can be broken. to disperse offspring to new habitats where more food might be
However, fungi cannot grow with lignin as their sole source available. Thus, spore production is favoured by natural selec-
of food. The six-carbon rings in lignin are extremely difficult tion when individuals are under nutritional stress.

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 Multiple Mating Types In some fungal lineages, hyphae come in In many fungi, the process of sexual reproduction begins when
many different mating types. Instead of having morphologically hyphae from two different mating types grow near each other and
distinct males and females that produce sperm and eggs, hyphae fuse to form a hybrid cell. When the cytoplasm of two individu-
of different mating types appear identical. als fuses in this way, plasmogamy (pronounced plaz-MAH-ga-
Hyphae of the same mating type will not combine during mee) is said to occur (Figure 29.16). Often, plasmogamy does not
sexual reproduction. When zygomycete hyphae grow close to immediately lead to nuclear fusion. Rather, the genetically dis-
each other, for example, they will not fuse unless the individuals tinct, haploid nuclei persist in the same cell or mycelium. When
have different alleles of one or more genes involved in mating. If genetically distinct nuclei from the two different mating types
chemical messengers released by two hyphae indicate that they exist within a single mycelium, it is considered heterokaryotic
are of different mating types, then fusion and zygosporangium (“different kernel”). Most hyphae in heterokaryotic mycelia are
formation follows. dikaryotic (“two kernel”)—their cells are divided by septa, and
In this way, mating types function as sexes. Instead of having each cell contains two nuclei, one from each mating type.
just two sexes, a single fungal species may have tens of thousands The distinct nuclei in dikaryotic and heterokaryotic mycelia
of sexes. The basidiomycete Schizophyllum commune, for exam- function independently, even though gene expression must be
ple, is estimated to have 28 000 mating types. coordinated for growth and development to occur. For example,
Why so many? The leading hypothesis to explain the exis- in septate dikaryotic fungi the two types of nuclei divide as the
tence of mating types is that it helps generate genetic diversity hyphae expand, so each compartment that is divided by a septum
in offspring by increasing the probability of outcrossing. Genetic contains one of each of the two types of nuclei. Biologists are
diversity, in turn, is known to be advantageous in fighting off currently investigating how the activities of the two nuclei are
infections and responding to changes in the environment (see coordinated.
Chapter 13). In a dikaryotic or heterokaryotic mycelium, pairs of unlike
nuclei may eventually fuse to form diploid zygotes. The fusion
How Does Fertilization Occur? Compared with fertilization of nuclei is called karyogamy (pronounced ka-ree-AH-ga-mee).
in land plants, protists, and animals, fertilization in fungi has The diploid nuclei that are produced by karyogamy then divide
important unique features: by meiosis to form haploid spores. As a result of meiosis, the
spores are genetically distinct and different mating types are
• Among all the lineages of fungi, only some chytrids produce
produced.
gametes, which are not considered eggs and sperm. Both male
If you understand the relationship between plasmog-
and female gametes are motile, and female gametes are only
amy and karyogamy, you should be able to compare and con-
slightly larger than male gametes.
trast these events with the life cycles of other eukaryotes.
• In all other lineages of fungi, fertilization occurs in two dis- For example, which human cells undergo plasmogamy (cell
tinct steps: (1) fusion of hyphae and (2) fusion of nuclei from fusion) and karyogamy (nuclei fusion)? Are any human cells
the fused hyphae. These two steps can be separated by long similar to a heterokaryotic (possessing two or more geneti-
time spans and even long distances. cally distinct nuclei) fungal cell? Explain.

Heterokaryotic
mycelium Haploid (n)
(n + n) Heterokaryotic (n + n)
Nuclei
Spore-producing MITOSIS MY Diploid (2n)
structure GA
MO KA
(n) AS RY
PL OG
of AM
ion (fu
(fus plasment nu sio
cle n
o
Y

r
cyt diffe ls) d o
ind iffe i fro f
S

m a
fro dividu ivi ren m
MITOSI

in SEXUAL du t Zygote
ASEXUAL REPRODUCTION als
(2n)
)
REPRODUCTION

Spore-producing
structure
(n) S
SI
IO
Spores M IT ME
OSI
(n) MITOSIS S
Mycelium Spores
(n) (n)
Figure 29.16 Fungi Have Unusual Life Cycles. A generalized fungal life cycle, showing both asexual and sexual
reproduction.
Fungi spend most of their lives feeding. Which is the longest-lived component of this life cycle?

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 Asexual Reproduction As the left side of Figure 29.16 indi- Zygomycete mycelia frequently also reproduce asexually by
cates, most fungal species can reproduce asexually as well as making sporangia whose haploid spores (conidia) are produced
sexually. In fact, large numbers of ascomycetes have never by mitosis and dispersed by wind.
been observed to reproduce sexually. Researchers don’t yet
know if these fungi are unable to reproduce sexually, or if it Basidiomycete Life Cycle Mushrooms, bracket fungi, boletes,
simply has not happened while they have been growing under stinkhorns, and puffballs are sexual reproductive structures pro-
observation. duced by members of the Basidiomycota (Figure 29.17c). Even
During asexual reproduction, spore-forming structures are though their size, shape, and colour vary enormously from spe-
produced by a haploid mycelium, and spores are generated by cies to species, all basidiomycete reproductive structures origi-
mitosis. As a result, offspring are clones—meaning that they are nate from the dikaryotic hyphae of mated individuals.
genetically identical to their parent. Inside a mushroom or bracket fungus or puffball, the club-
Producing microscopic spores that are carried by wind was shaped, spore-producing cells called basidia form at the ends of
an adaptation to life on land that allowed terrestrial fungi to dikaryotic hyphae. Karyogamy occurs within the basidia. The
disperse widely and to colonize new habitats. The production diploid nucleus that results undergoes meiosis, yielding four
of large numbers of asexual spores has proven to be an effective haploid spores.
strategy for many fungi. Spores are eventually ejected from the end of the basidia and
are dispersed by the wind. It is not unusual for a single puffball or
mushroom to produce a billion spores. If they land in a suitable
Four Major Types of Life Cycles location, the spores germinate and grow by mitosis to form new
Among the sexually reproducing species of fungi, the presence hyphae.
of a heterokaryotic stage and the morphology of the spore-
producing structure vary. Morphologies of reproductive struc-
Ascomycete Life Cycle As in basidiomycetes, the mature sexual
tures were introduced in Section 29.2. Let’s now take a closer
reproductive structure in ascomycetes is also produced by a
look at the sexual reproduction portion of each of the four major
dikaryotic hypha.
types of life cycles that have been observed in fungi. Each life
As Figure 29.17d illustrates, the process usually begins when
cycle is summarized in Figure 29.17 on pages 632–633.
hyphae or specialized structures from different mating types
fuse, forming a heterokaryotic cell containing many independent
Chytrid Life Cycle The chytrids are the only types of fungi with nuclei. A short dikaryotic hypha with cells containing two nuclei,
species that exhibit alternation of generations. Recall that in one from each parent, emerges and eventually grows into a
organisms with alternation of generations, there are multicel- complex reproductive structure whose hyphae have the sac-like,
lular haploid and diploid forms (see Chapter 27). Figure 29.17a spore-producing structures called asci at their tips.
shows how alternation of generations occurs in the well-studied After karyogamy occurs inside each ascus, meiosis and one
genus Allomyces. Here are the key stages in its life cycle: round of mitosis takes place, resulting in the production of eight
1. Haploid adults form gametangia, in which male and female haploid spores. When the ascus matures, the spores inside are
swimming gametes are produced by mitosis. forcibly ejected.
2. Gametes from the same individual or different individuals
fuse to form a diploid zygote.
3. The zygote grows by mitosis into a diploid sporophytic Check Your Understanding
mycelium. If you understand that …
The life cycle continues when meiosis occurs in the sporophytic • Most fungi live in close association with land plants as
mycelium, inside a structure called a sporangium. The haploid mycorrhizae, endophytes, or saprophytes.
spores produced by meiosis disperse by swimming; then each • When plants are alive, mycorrhizae and some endophytes
associate with their roots; many other endophytes grow in
grows by mitosis into a haploid gametophytic mycelium.
aboveground plant tissues.
• When plants die, saprophytes degrade their tissues and release
Zygomycete Life Cycle In zygomycetes, sexual reproduction nutrients.
starts when hyphae from different mating types fuse, as shown • Instead of being based on the fusion of gametes, sexual
in Figure 29.17b. Plasmogamy forms a spore-forming structure reproduction in most fungi begins with plasmogamy, or
called a zygosporangium that develops a tough, resistant coat. the fusion of the cytoplasm of haploid hyphae, followed by
Inside the zygosporangium, nuclei from the mating partners karyogamy, or nuclear fusion, and meiosis. The resulting spores
fuse—meaning that karyogamy occurs. grow by mitosis into new hyphae.
The zygosporangium can persist if conditions become too You should be able to …
cold or dry to support growth. When temperature and moisture
1. Describe evidence that mutualism occurs in EMF.
conditions are again favourable, however, meiosis occurs and 2. Compare and contrast what happens inside a zygosporan-
the zygosporangium germinates and forms a sporangium. The gium, basidium, and ascus.
meiotic products (haploid spores) within the sporangium are
Answers are available in Appendix A.
released and germinate, growing into new mycelia.

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 (a) Chytrids include the only fungi in which alternation of generations occurs. There is no heterokaryotic stage.
(Species: Allomyces macrogynus)

Zygote
FUSION (2n)

S IS MIT
OS
TO Plasmogamy IS
MI and karyogamy
Gametophytic Sporophytic
mycelium Gametes mycelium
(n) Gametes form (n) Spores form (2n)
in gametangia in sporangia
Gametes and spores (n)
1 mm disperse by water 1 mm

MI
TO
SIS

2n
Spore MEIOSIS
(n)
n

Haploid (n)
Heterokaryotic (n + n)
Diploid (2n)

(b) Zygomycetes form yoked hyphae that produce sporangia (spore-forming structures).
(Species: Rhizopus stolonifer, black bread mould)

0.1 mm 1 mm

Asexual PLASMO Spores form in

GAMY zygosporangia
sporangium
(produces
conidia) Hyphae of n+n
different Hyphae
mating types fuse

Sexual
Spores sporangium
(n) (releases
spores)
Zygote n+n

MITO MY
SIS YOGA
2n KAR
MEIOSIS
n

Figure 29.17 Variation in Sexual Reproduction in Fungi. The sexual part of the life cycle in four major groups of fungi: (a) chytrids,
(b) zygomycetes, (c) basidiomycetes, and (d) ascomycetes.

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 (c) Basidiomycota have reproductive structures with many spore-producing basidia.
(Species: Amanita muscaria, fly agaric)
Spores (n)
dispersed
by wind
KARYOGAMY
MEIO
SIS
Basidium 2n
Mature spore-
producing body
(n + n) 20 om Spores

10 om

SIS
1 cm Dikaryotic

TO
mycelium (n + n) I
n M
begins to grow
Spores (n)
germinate
PLASMOGAMY to form hyphae

n+n n

Haploid (n)
Dikaryotic (n + n)
Diploid (2n)

(d) Ascomycota have reproductive structures with many spore-producing asci.

(Species: Cookeina speciosa, Venus wine cup fungus)

20 om
Mature
ascus
2n MEIOSIS MITOS Spores (n)
KARYOGAMY
IS dispersed
by wind
Young
ascus 10 om
Mature spore-
producing body Eight spores
(n + n) formed

1 cm Dikaryotic Structures containing n

mycelium (n + n) many nuclei form
begins to grow

n+n
MITO
SIS Spores (n)
PLASMOGAMY
germinate
n to form
Hyphae of different hyphae
mating types make
contact and fuse

Figure 29.17 Variation in Sexual Reproduction in Fungi (continued).

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 Microsporidia
29.4 Key Lineages of Fungi More than 1200 species of microsporidia have been identified,
The lineages described here and summarized in Table 29.1 are and all are single-celled and parasitic on animal cells. They are
just a small sampling of fungal diversity. As you study the table, distinguished by a unique structure called the polar tube, which
it’s important to keep in mind that the wide array of absorptive allows them to enter the cells they parasitize. The tube shoots out
lifestyles found in the various lineages (saprophytic, mutualis- from the microsporidian, penetrates the membrane of the host
tic, parasitic, etc.) evolved independently of each other—that is, cell, and acts as a conduit for the contents of the microsporidian
after the groups split from their common ancestor and formed cell to enter the host cell. Once inside, the microsporidian DNA
distinct and separate lineages. replicates and produces a generation of daughter cells, which go
Based on the current hypothesized phylogeny for the Fungi, on to infect other host cells.
it is clear that neither chytrids nor zygomycetes form mono- Species from nine different genera can infect humans. Univer-
phyletic groups and should not be named as single lineages (see sity of British Columbia’s Patrick Keeling describes one patho-
Figure 29.10). Researchers continue to collect and analyze addi- genic microsporidian that lacks all genes for carbon metabolism.
tional data—mostly DNA sequences—to identify monophyletic This fungus cannot even carry out glycolysis and is entirely
groups and explain the early evolution of fungi. dependent on its human host for ATP. In most cases, however,

DIVERSITY Table 29.1 Key Lineages of Fungi

Known
Species Absorptive Life Style Life Cycle Relevance

Microsporidia 1300 All are parasites of Some appear to Many species can
animal cells, especially reproduce only infect humans,
Dr. J. I. Ronny Larsson, Professor

in insects or fish asexually. Others but cause serious

Spore produce various types infections only in
case Enter host cell through
polar tube (which of sexual spores patients with a
microsporidian shoots Some infect several compromised immune
Polar tube
into host cell) hosts to complete life system
cycle Some are pests in
of Zoology

honeybee and silkworm

5 om colonies
◀ Nosema tractabile

Chytrids 750 Many decompose The only fungi that Some are parasites of
Gametangia Aquatic; common plants by digesting produce motile cells– mosquitoes and are
in freshwater cellulose both their spores and being studied as a
Mutualistic chytrids live gametes swim via biological control agent
in guts of cows, deer, flagella Batrachochytrium
and other mammals Most exhibit alternation dendrobatidis is
and help digest plant of generations largely responsible for
material declines in amphibian
Parasitic chytrids infect populations worldwide
many species of plants
Thomas J. Volk

and animals

◀ Allomyces macrogynus
250 om
showing male and female
gametangia

Zygomycetes 1050 Many are saprophytes Asexual reproduction is Several species often
Asexual
Food moulds and live on plant very common; asexual seen growing on bread
Researchers, Inc./Science Source

sporangia
debris sporangia produce and soft fruits
spores
Gregory G. Dimijian/Photo

Some parasitize other Some are used in the

fungi, insects, or Sexual reproduction production of steroids,
spiders involves fusion of pigments, alcohols,
hyphae and formation and fermented foods
of zygosporangium and
sexual sporangia
200 om
Mycelium ◀ Rhizopus nigricans
(a bread mould)

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 microsporidians cause serious infections only in AIDS patients decomposers of plant material in wet soils, ponds, and lakes.
and other individuals whose immune systems are not function- However, many of the freshwater species are parasitic; some of
ing well. Some microsporidians are serious pests in honeybee these species cause disease epidemics in algae or aquatic insects.
and silkworm colonies. However, other species that infect grass- Other species parasitize mosses, ferns, flowering plants, or ani-
hoppers are used as a biological control agent to reduce crop mals. Chytrids are the likely culprit behind the decline of amphib-
damage caused by these insects. ian populations worldwide, including in Canada.

Chytrids Zygomycetes
Chytrids are largely aquatic and are common in freshwater envi- The zygomycetes (“yoked fungi”) are primarily soil-dwellers.
ronments. Some species live in wet soils, and a few have been When hyphae of opposite mating types grow near each other,
found in desert soils that are wet only during a rainy season. they yoke together and fuse during sexual reproduction and then
Species found in dry soils have tough spores that endure harsh form a durable, thick-walled zygosporangium. Asexual repro-
conditions. Spores from chytrids have been shown to germinate duction is also extremely common.
after a resting period of as long as 31 years. The common black bread mould Rhizopus stolonifer is a fre-
Many species of chytrids have enzymes that allow them quent household pest—it’s probably the zygomycete that is
to digest cellulose. As a result, these species are important most familiar to you. Saprophytic and parasitic members of this

DIVERSITY Table 29.1 Key Lineages of Fungi (continued)

Known
Species Absorptive Life Style Life Cycle Relevance

Glomeromycota 200 Form mutualistic Most produce large Play important roles in
associations with plant spores the ecology of prairies
roots in the form of Difficult to grow and and tropical forests
arbuscular mycorrhizal study
fungi
Sexual reproduction
has not been
Dr. Jim Deacon

documented
◀ An arbuscular mycor-
50 om rhizal fungus penetrating
a root

Basidiomycota 32 000 Decompose wood Produce haploid spores Mushrooms are used
(“club fungi”) by producing lignin in club-shaped basidia as a source of food
Mushrooms, peroxidase Basidia of mushroom- Some mushrooms are
boletes, Some form forming species line toxic or hallucinogenic
Zoonar/Jürgen Vogt/AGE Fotostock

stinkhorns, ectomycorrhizal gills found under the Ectomycorrhizal

puffballs, associations with tree cap basidiomycetes
brackets roots enhance growth
Smut and rust fungi are of hardwoods and
plant parasites softwoods
Smuts and rusts cause
serious economic
1 cm damage to crop plants
◀ Lycoperdon perlatu
(a puffball) releasing spores

Ascomycota 64 000 Many form Produce haploid spores Lichens are often
(“sac fungi”) ectomycorrhizal in sac-like asci sensitive to air pollution
associations with tree and are used as
Adrian Hepworth/Photoshot

Morels, truffles, Asci of many species

yeasts, cup roots found on fleshy, cup- bioindicators
fungi, lichen- Some form mutualistic shaped structures
forming fungi associations with Asexual reproduction
License Limited

photosynthetic algae or in many

bacteria in lichens
1 cm ◀ Cookeina speciosa
(a cup fungus) Some are predatory on
protists or nematodes

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 lineage are also responsible for rotting strawberries and other named because most ascomycetes form mycelia and produce
fruits and vegetables and causing large losses in these food indus- spores in structures called asci. Ascomycete mycelia also com-
tries. On the positive side, other species are used beneficially in monly reproduce asexually. In addition, single-celled yeasts are
commercial production of various products. part of the Ascomycota lineage. Yeasts reproduce when daughter
cells pinch off from the parent cell after mitosis. This asexual
Glomeromycota reproductive process is called budding.
All members of this group (except for a single species) comprise About half of ascomycete species grow in mutualistic associa-
the arbuscular mycorrhizal fungi (AMF). Recall from Section tion with cyanobacteria and/or single-celled green algae, forming
29.3 that AMF absorb phosphorus-containing ions or molecules the structures called lichens (see Figure 29.14). Over 17 500 dif-
in the soil and transfer them, along with nitrogen and other ferent lichens have been described to date; in most, the fungus
nutrients and water, into the roots of most of the plants living in involved is an ascomycete (although a few basidiomycetes par-
grasslands and tropical forests. The host plant provides the sym- ticipate as well).
biotic fungi with sugars and other organic compounds. Many ascomycetes form mutualistic EMF associations with
tree roots. Ascomycetes are also the most common endophytic
Basidiomycota fungi on aboveground tissues. Many other ascomycetes are sap-
rophytic and are abundant on forest floors and in grassland soils;
Almost all basidiomycetes form mycelia and produce multicellu-
parasitic forms are also common. In addition, several species are
lar reproductive structures that include basidia, the club-shaped
predatory—primarily on amoebae and other unicellular protists,
cells where meiosis and spore formation occur. About 31 000
but a few on larger prey. For example, the nematode-trapping
species of basidiomycetes have already been described, and more
fungus shown in Figure 29.6 is an ascomycete.
are discovered each year.
Some saprophytic ascomycetes can grow on jet fuel or paint
Throughout the world, mushrooms are cultivated or collected
and are used to help clean up contaminated sites. Penicillium
from the wild as a source of food. The white button, crimini,
is an important source of antibiotics, and Aspergillus produces
and portabello mushrooms you may have seen in grocery stores
citric acid used to flavour soft drinks and candy. Truffles are
are all the same species, Agaricus bisporus, at different stages of
so highly prized that they can fetch over $750 per kilogram,
development. Some of the toxins found in poisonous mushrooms
and the multibillion-dollar brewing, baking, and wine-making
are used in biological research; others are hallucinogens.
industries would collapse without the yeast Saccharomyces cere-
Basidiomycetes are important saprophytes. Along with a few soil-
visiae. A few parasitic ascomycetes cause infections in humans
dwelling bacteria, some members of this lineage are the only organ-
and other animals. In land plants, parasites from this group
isms capable of synthesizing lignin peroxidase and are therefore
cause several plant diseases, including Dutch elm disease and
important in wood decomposition. Some basidiomycetes are ecto-
chestnut blight.
mycorrhizal fungi (EMF) that improve the health and productivity
A recently discovered cold-loving ascomycote fungus, Geomy-
of temperate forests via their mutualistic associations with tree
ces destructans, is responsible for a deadly disease in bats called
roots. Temperate forests provide most of the hardwoods and soft-
white-nose syndrome. The fungal infection leaves the dead bats
woods used in construction, furniture-making, and papermaking.
looking as though they have been sprinkled with flour. Since 2006,
Basidiomycetes can also be parasitic. One subgroup of the
over 5 million hibernating bats have died, and many bat colonies
lineage consists entirely of parasitic forms called rusts, including
have been completely wiped out. White-nose syndrome was first
species that cause serious infections in many crop plants. Plant
detected in Canada in 2010 and has now spread to Ontario, Quebec,
parasites called smut fungi specialize in infecting grasses; a few
New Brunswick, Prince Edward Island, and Nova Scotia. Little
parasitic basidiomycetes even infect other fungi.
brown bats, one of the hardest-hit species, have been functionally
extirpated in New Brunswick. Only 12 of the bats were counted in a
Ascomycota 2015 survey of caves in southern New Brunswick. The syndrome is
Ascomycetes are found in virtually every terrestrial habitat as moving west each year, but as of 2017, had not yet been detected in
well as some freshwater and marine environments. They are so Manitoba.

Chapter 29 Review For media, go to MasteringBiology

29.1 Why Do Biologists Study Fungi? the plant with nitrogen, phosphorus, and water. The fungi benefit
by receiving sugars from photosynthesis along with other plant
• Parasitic fungi are responsible for devastating blights in crops and
products.
other plants.
• Once plants die, saprophytic fungi degrade the lignin and cellulose
• Fungi are used widely in the food processing, cheese-making, bak- in wood and use nutrients from decaying plant material.
ing, and brewing industries. • Because they free up carbon atoms that would otherwise be
• Many fungi species live in close association with land plants. The locked up in wood, fungi speed up the carbon cycle in terrestrial
roots of most plants are infected by mutualistic fungi that provide ecosystems.

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 fusion of nuclei, or karyogamy, does not occur immediately, a het-
29.2 How Do Biologists Study Fungi? erokaryotic mycelium forms, which in some species is dikaryotic.
Heterokaryotic cells may eventually produce spore-forming struc-
• Morphological analyses of fungi have revealed adaptations that
tures where karyogamy and meiosis take place.
make fungi exceptionally effective at absorbing nutrients from the
environment.
• Researchers have determined that fungi exhibit two growth forms— 29.4 Key Lineages of Fungi
single-celled “yeasts” or mycelia composed of long, filamentous • Microsporidia are parasitic, single-celled fungi that use a unique
hyphae. Mycelia provide fungi with extremely high surface-area-to- structure called a polar tube to infect host cells.
volume ratios. Some fungal bodies are extremely large. • Chytrids are usually aquatic with motile, flagellated spores
• Septa divide many hyphae into cells, but pores in the septa facili- and gametes. Some are terrestrial decomposers and some are
tate transport of nutrients and organelles through the hyphae. parasites.
• Scientists have determined from morphological studies that fungi • Zygomycetes are largely saprophytic on dead plant tissue. During
possess four known, distinctive types of sexual reproductive sexual reproduction, they form zygosporangia after hyphae from
structures: (1) Chytrids produce motile gametes; (2) zygomycetes different mating types fuse. Asexual reproduction is also common.
make tough zygosporangia; (3) basidiomycetes have club-shaped, • The Glomeromycota is a monophyletic group of soil fungi that
spore-forming structures; and (4) ascomycetes have sac-like, form arbuscular mycorrhizae with most land plants.
spore-forming structures. • The Basidiomycota is a monophyletic group of terrestrial fungi
• Fungi can reproduce asexually, and many do so by forming that includes mushrooms, bracket fungi, stinkhorns, and puff-
conidia—asexual spores that are released and have the ability to balls. During sexual reproduction, they form airborne spores in
grow into a new fungus. structures called basidia. Some can degrade wood. Others form
ectomycorrhizae with certain trees.
• Recent analyses of DNA sequence data have revealed that fungi are
more closely related to animals than they are to plants, and that • Ascomycota is a monophyletic group of terrestrial fungi. Dur-
the Basidiomycota, Ascomycota, and Glomeromycota each form ing sexual reproduction, they form airborne spores in structures
monophyletic groups. Researchers are still trying to understand called asci. Some form lichens in association with green algae or
how the various groups of chytrids, zygomycetes, and microspo- cyanobacteria.
ridians are related to each other and to the other fungal lineages.
Answers are available in Appendix A

29.3 What Themes Occur in the Test Your Knowledge

Diversification of Fungi? 1. The mycelial growth habit leads to a body with a high surface-area-
• All fungi make their living by absorbing nutrients from living or to-volume ratio. Why is this important?
dead organisms. a. Mycelia have a large surface area for absorption.
• Fungi often associate with other organisms, especially plants, b. The hyphae that make up mycelia are long, thin tubes.
in symbiotic relationships that can be mutualistic, parasitic, or c. Most hyphae are broken up into compartments by walls called
commensal. septa, although some exist as single, gigantic cells.
d. Hyphae can infiltrate living or dead tissues.
• Ectomycorrhizal fungi (EMF) and arbuscular mycorrhizal fungi
(AMF) form mutualistic symbiotic relationships with roots of 2. What are asci?
most plants. a. club-shaped, spore-forming cells found in mushrooms
• Many fungi live in leaves, stems, and roots as endophytes. Some of b. asexual spores
these endophytic fungi secrete toxins that discourage herbivores; c. sac-like structures that produce eight spores
others live as commensals, providing no apparent benefit to the d. the spore-producing structure commonly found in zygomycetes
plant. 3. The hyphae of arbuscular mycorrhizal fungi (AMF) form bushy
• Fungi form symbiotic relationships with many animal species structures after making contact with the plasma membrane of a root
and, in the form of lichens, with various species of green algae and cell. What is the function of these structures?
cyanobacteria. a. They anchor the fungus inside the root, so the association is more
permanent.
• Extracellular digestion, in which enzymes are secreted into food
b. They increase the surface area available for the transfer of nutri-
sources, enables fungi to break down extremely large molecules
ents.
before absorbing them.
c. They produce toxins that protect the plant cells against herbi-
• Lignin decomposes through a series of uncontrolled oxidation vores.
reactions triggered by the enzyme lignin peroxidase. d. They break down cellulose and lignin in the plant cell wall.
• Cellulose digestion occurs in a carefully regulated series of steps,
4. What does it mean to say that a hypha is dikaryotic?
each catalyzed by a specific cellulase.
a. Two nuclei fuse during sexual reproduction to form a zygote.
• Many species of fungi have never been observed to reproduce b. Two independent nuclei, derived from different individuals, are
sexually. Most species can produce haploid spores either sexually present in each cell.
or asexually, however. c. The nucleus is diploid or polyploid—not haploid.
• Sexual reproduction usually starts when hyphae from genetically d. It is extremely highly branched, which increases its surface area
different mating types fuse—an event called plasmogamy. If the and thus absorptive capacity.

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 Test Your Understanding
Just how fast can fungal spores fly?
5. The Greek root ecto means “outer.” Why are ectomycorrhizal fungi, Some fungi have elaborate mechanisms for dispersing spores. For
or EMF, aptly named? example, the “squirt gun” fungus Pilobolus, which grows in cow
a. Their hyphae form tree-like branching structures inside plant cell dung, forcibly shoots off tiny black sporangia filled with spores.
walls. Any sporangia that are flung onto fresh grass are likely to be eaten
b. They are mutualistic. by a grazing cow, passed through its digestive system unharmed,
c. Their hyphae form dense mats that envelop roots but do not pen- and deposited in a new batch of dung. The spores carried within the
etrate the cell walls. sporangia are perfectly positioned to grow into a new mycelium.
d. They transfer nitrogen from outside their plant hosts to the inte-
rior. 11. Pilobolus is a zygomycete, and the sporangia just described
produce asexual spores. In contrast, which of the following best
6. Explain why fungi that degrade dead plant materials are important describes what happens during the sexual phase of its life cycle?
to the global carbon cycle. Do you accept the text’s statement that, a. It produces eight ascospores in each ascus.
without these fungi, “Terrestrial environments would be radically b. It produces basidiospores that grow into mushrooms.
different than they are today, and probably much less productive”? c. It produces motile sperm and eggs.
Why or why not? d. Hyphae from two compatible mating types fuse and form a
7. Lignin and cellulose provide rigidity to the cell walls of plants. But in zygosporangium.
most fungi, chitin performs this role. Why is it logical that most fungi 12. MODEL Both asexual sporangia (shown in the photo) and
don’t have lignin or cellulose in their cell walls? zygosporangia can be found in cow dung. Make a simplified
8. SOCIETY Lawns are sometimes fertilized with nitrate that can be drawing that illustrates the events of plasmogamy, karyogamy,
washed into neighbouring woodlots by rain. If the trees in that and meiosis that occur during sexual reproduction in Pilobolus.
woodlot are associated with EMF, what effect might the excess Be sure to indicate and label the zygosporangium.
nitrate have on the fungi and/or trees? 13. Would you describe the association between Pilobolus and cows
a. Fungal growth may be stimulated, causing the trees to transport as a symbiotic relationship? Explain your answer.
more nitrogen to the fungi.
b. The fungi may secrete more peptidases to break down the nitrate. 14. Explain the process of extracellular digestion that occurs during
c. The trees may take up the nitrate directly into their roots and rely the growth of Pilobolus mycelia through dung.
less on the fungi. 15. QUANTITATIVE Using high-speed video, researchers have measured
d. The excess nitrate may stimulate the fungi to transport more several variables associated with sporangium discharge in
sugar to the trees. various spore-shooting fungi. Based on the data in the table
below, how do the launch speed and acceleration of Pilobolus
Test Your Problem-Solving Skills compare with those of the other fungi? How does launch speed
for each species compare with the top speed of 44 km/h reached
9. PROCESS OF SCIENCE Table 29.1 mentions that chytrids are
by Jamaican athlete Usain Bolt during his 2009 world record-
responsible for massive die-offs currently occurring in amphibians.
setting race?
Review Koch’s postulates (see Chapter 26), then design a study
showing how you would use Koch’s postulates to test the hypothesis Ascobolus Podospora Pilobolus
that chytrid infections are responsible for the frog deaths. immersus anserina kleinii
Launch speed (m/s) 14 21 9
10. PROCESS OF SCIENCE Many mushrooms are extremely colourful.
2
One hypothesis is that the colours serve as a warning to prevent Acceleration (m/s ) 1 800 000 1 500 000 210 000
animals from eating mushrooms, much like the bright yellow and Range (m) 0.3 0.2 2.5
black stripes on wasps help to deter potential predators. Design an
Turgor pressure (MPa) 0.40 0.40 0.55
experiment capable of testing this hypothesis.
Source: Yafetto, L., et al. 2008. PLoS ONE, 3(9): e3237.

PUT IT ALL TOGETHER: Case Study 16. PROCESS OF SCIENCE Propose an experiment to test whether
Pilobolus shoots its sporangia preferentially toward light.
Sporangium Hypothesize why such a preference might be advantageous.

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1 mm
Pilobolus crystallinus

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638 Unit 5  The Diversification of Life

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