LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION

Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson

Chapter 31

Fungi

Lectures by Erin Barley Kathleen Fitzpatrick

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Overview: Mighty Mushrooms

Fungi are diverse and widespread They are essential for the well-being of most terrestrial ecosystems because they break down organic material and recycle vital nutrients About 100,000 species of fungi have been described It is estimated there are actually 1.5 million species of fungi

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Figure 31.1

Concept 31.1: Fungi are heterotrophs that feed by absorption

Despite their diversity, fungi share key traits, most importantly the way in which they derive nutrition

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Nutrition and Ecology

Fungi are heterotrophs and absorb nutrients from outside of their body Fungi use enzymes to break down a large variety of complex molecules into smaller organic compounds The versatility of these enzymes contributes to fungis ecological success

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 Fungi exhibit diverse lifestyles

Decomposers Parasites Mutualists

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Body Structure
The most common body structures are multicellular filaments and single cells (yeasts) Some species grow as either filaments or yeasts; others grow as both

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Animation: Fungal Reproduction and Nutrition

Right-click slide / select Play
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 The morphology of multicellular fungi enhances their ability to absorb nutrients Fungi consist of mycelia, networks of branched hyphae adapted for absorption A myceliums structure maximizes its surface areato-volume ratio Fungal cell walls contain chitin

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Figure 31.2

Reproductive structure Hyphae

Spore-producing structures

60 m

Mycelium

Figure 31.2a

Reproductive structures of penny bun fungi

Figure 31.2b

Hyphae

60 m

Figure 31.2c

Mycelium

 Most fungi have hyphae divided into cells by septa, with pores allowing cell-to-cell movement of organelles Coenocytic fungi lack septa and have a continuous cytoplasmic mass with hundreds or thousands of nuclei

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Figure 31.3

Nuclei

Cell wall Pore Septum

Cell wall

Nuclei (b) Coenocytic hypha

(a) Septate hypha

Specialized Hyphae in Mycorrhizal Fungi

Some unique fungi have specialized hyphae called haustoria that allow them to penetrate the tissues of their host

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Figure 31.4

Nematode

Hyphae

25 m

(a) Hyphae adapted for trapping and killing prey

Fungal hypha Plant cell wall

Plant cell Plant cell plasma membrane

(b) Haustoria

Haustorium

Figure 31.4a

Nematode

Hyphae

25 m

(a) Hyphae adapted for trapping and killing prey

Figure 31.4b

Fungal hypha

Plant cell wall

Plant cell Plant cell plasma membrane

(b) Haustoria

Haustorium

 Mycorrhizae are mutually beneficial relationships between fungi and plant roots Ectomycorrhizal fungi form sheaths of hyphae over a root and also grow into the extracellular spaces of the root cortex Arbuscular mycorrhizal fungi extend hyphae through the cell walls of root cells and into tubes formed by invagination of the root cell membrane

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 Mycorrhizal fungi deliver phosphate ions and minerals to plants Most vascular plants have mycorrhizae

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Concept 31.2: Fungi produce spores through sexual or asexual life cycles
Fungi propagate themselves by producing vast numbers of spores, either sexually or asexually Fungi can produce spores from different types of life cycles

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Figure 31.5-1

Key Haploid (n) Heterokaryotic Diploid (2n) Spore-producing structures Spores Mycelium ASEXUAL REPRODUCTION

GERMINATION

Figure 31.5-2

Key Haploid (n) Heterokaryotic Diploid (2n) Spore-producing structures Spores Mycelium ASEXUAL REPRODUCTION SEXUAL REPRODUCTION PLASMOGAMY Heterokaryotic stage

KARYOGAMY

Zygote

GERMINATION

Figure 31.5-3

Key Haploid (n) Heterokaryotic Diploid (2n) Spore-producing structures Spores Mycelium ASEXUAL REPRODUCTION SEXUAL REPRODUCTION PLASMOGAMY Heterokaryotic stage

KARYOGAMY

Zygote

GERMINATION

GERMINATION

MEIOSIS

Spores

Sexual Reproduction
Fungal nuclei are normally haploid, with the exception of transient diploid stages formed during the sexual life cycles Sexual reproduction requires the fusion of hyphae from different mating types Fungi use sexual signaling molecules called pheromones to communicate their mating type

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 Plasmogamy is the union of cytoplasm from two parent mycelia In most fungi, the haploid nuclei from each parent do not fuse right away; they coexist in the mycelium, called a heterokaryon In some fungi, the haploid nuclei pair off two to a cell; such a mycelium is said to be dikaryotic

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 Hours, days, or even centuries may pass before the occurrence of karyogamy, nuclear fusion During karyogamy, the haploid nuclei fuse, producing diploid cells The diploid phase is short-lived and undergoes meiosis, producing haploid spores The paired processes of karyogamy and meiosis produce genetic variation

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Asexual Reproduction
In addition to sexual reproduction, many fungi can reproduce asexually Molds produce haploid spores by mitosis and form visible mycelia

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Figure 31.6

1.5 m

Figure 31.6a

Figure 31.6b

1.5 m

 Other fungi that can reproduce asexually are yeasts, which are single cells Instead of producing spores, yeasts reproduce asexually by simple cell division and the pinching of bud cells from a parent cell Some fungi can grow as yeasts and as filamentous mycelia

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Figure 31.7

10 m

Parent cell Bud

 Many molds and yeasts have no known sexual stage Mycologists have traditionally called these deuteromycetes, or imperfect fungi This is not a sound taxonomic group; fungi are reclassified once their sexual stage is discovered

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Concept 31.3: The ancestor of fungi was an aquatic, single-celled, flagellated protist
Fungi and animals are more closely related to each other than they are to plants or other eukaryotes

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The Origin of Fungi

Fungi, animals, and their protistan relatives form the opisthokonts clade

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Figure 31.8

Animals (and their close protistan relatives)

Opisthokonts

UNICELLULAR, FLAGELLATED ANCESTOR

Nucleariids Chytrids Other fungi

Fungi

 DNA evidence suggests that

Fungi are most closely related to unicellular nucleariids Animals are most closely related to unicellular choanoflagellates

This suggests that multicellularity arose separately in animals and fungi The oldest undisputed fossils of fungi are only about 460 million years old

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Figure 31.9

50 m

Are Microsporidia Fungi?

Microsporidia are unicellular parasites of animals and protists They have tiny organelles derived from mitochondria but not conventional mitochondria Molecular comparisons indicate they are fungi or are closely related to fungi

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Figure 31.10

Host cell nucleus Developing microsporidian Spore

10 m

The Move to Land

Fungi were among the earliest colonizers of land and probably formed mutualistic relationships with early land plants

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Concept 31.4: Fungi have radiated into a diverse set of lineages

Molecular analyses have helped clarify evolutionary relationships among fungal groups, although areas of uncertainty remain

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Figure 31.11
Chytrids (1,000 species)

Hyphae

25 m

Zygomycetes (1,000 species)

Fungal hypha

25 m

Glomeromycetes (160 species)

Ascomycetes (65,000 species)

Basidiomycetes (30,000 species)

Figure 31.11a

Hyphae

25 m

Chytrids (1,000 species)

Figure 31.11b

Zygomycetes (1,000 species)

Figure 31.11c

Fungal hypha

25 m

Glomeromycetes (160 species)

Figure 31.11d

Ascomycetes (65,000 species)

Figure 31.11e

Basidiomycetes (30,000 species)

Chytrids
Chytrids (phylum Chytridiomycota) are found in freshwater and terrestrial habitats They can be decomposers, parasites, or mutualists Molecular evidence supports the hypothesis that chytrids diverged early in fungal evolution Chytrids are unique among fungi in having flagellated spores, called zoospores

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Figure 31.UN01

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Figure 31.12

Flagellum 4 m

Zygomycetes
The zygomycetes (phylum Zygomycota) exhibit great diversity of life histories They include fast-growing molds, parasites, and commensal symbionts The life cycle of black bread mold (Rhizopus stolonifer) is fairly typical of the phylum Its hyphae are coenocytic Asexual sporangia produce haploid spores

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Figure 31.UN02

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Figure 31.13

PLASMOGAMY Mating type () Rhizopus growing on bread Gametangia with haploid nuclei 100 m Young zygosporangium (heterokaryotic) SEXUAL REPRODUCTION Dispersal and germination Zygosporangium KARYOGAMY

Mating type (+ )

Sporangia

Flagellum

ASEXUAL REPRODUCTION Dispersal and germination 50 m Mycelium

Sporangium MEIOSIS

Diploid nuclei

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

Figure 31.13a

PLASMOGAMY Mating type () Gametangia with haploid nuclei

Mating type (+ )

Young zygosporangium (heterokaryotic) SEXUAL REPRODUCTION Dispersal and germination

KARYOGAMY

Sporangium MEIOSIS

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

Figure 31.13b

ASEXUAL REPRODUCTION Dispersal and germination Mycelium

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

Figure 31.13c

Figure 31.13d

Figure 31.13e

100 m

Zygosporangium

Figure 31.13f

Sporangia

50 m

 The zygomycetes are named for their sexually produced zygosporangia Zygosporangia are the site of karyogamy and then meiosis Zygosporangia, which are resistant to freezing and drying, can survive unfavorable conditions Some zygomycetes, such as Pilobolus, can actually aim their sporangia toward conditions associated with good food sources

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Figure 31.14

0.5 mm

Glomeromycetes
The glomeromycetes (phylum Glomeromycota) were once considered zygomycetes They are now classified in a separate clade Glomeromycetes form arbuscular mycorrhizae

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Figure 31.UN03

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Figure 31.15

2.5 m

Ascomycetes
Ascomycetes (phylum Ascomycota) live in marine, freshwater, and terrestrial habitats Ascomycetes produce sexual spores in saclike asci contained in fruiting bodies called ascocarps Ascomycetes are commonly called sac fungi Ascomycetes vary in size and complexity from unicellular yeasts to elaborate cup fungi and morels

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Figure 31.UN04

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Figure 31.16

Morchella esculenta, the tasty morel

Tuber melanosporum, a truffle

Figure 31.16a

Morchella esculenta, the tasty morel

Figure 31.16b

Tuber melanosporum, a truffle

 Ascomycetes include plant pathogens, decomposers, and symbionts Ascomycetes reproduce asexually by enormous numbers of asexual spores called conidia Conidia are not formed inside sporangia; they are produced asexually at the tips of specialized hyphae called conidiophores Neurospora crassa, a bread mold, is a model organism with a well-studied genome

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Figure 31.17

Conidia; mating type ()

Key Dispersal Germination Mating type (+ ) PLASMOGAMY Haploid (n) Dikaryotic (n + n) Diploid (2n)

ASEXUAL REPRODUCTION Hypha

Conidiophore

Ascus (dikaryotic) Mycelia Mycelium Germination Dispersal Asci Ascocarp SEXUAL REPRODUCTION Diploid nucleus (zygote) KARYOGAMY Dikaryotic hyphae

Eight ascospores

Four haploid nuclei

MEIOSIS

Figure 31.17a

Dispersal

Germination

ASEXUAL REPRODUCTION Hypha

Conidiophore

Mycelium Germination Key Haploid (n) Dikaryotic (n + n) Diploid (2n)

Figure 31.17b

Conidia; mating type ()

Mating type (+ )

Key Haploid (n) Dikaryotic (n + n) Diploid (2n) PLASMOGAMY Ascus (dikaryotic)

Mycelia Germination Dispersal Ascocarp

Dikaryotic hyphae

SEXUAL REPRODUCTION KARYOGAMY Diploid nucleus (zygote)

Eight ascospores Four haploid nuclei MEIOSIS

Asci

Figure 31.17c

Eight ascospores

Basidiomycetes
Basidomycetes (phylum Basidiomycota) include mushrooms, puffballs, and shelf fungi, mycorrhizae, and plant parasites The phylum is defined by a clublike structure called a basidium, a transient diploid stage in the life cycle The basidiomycetes are also called club fungi Many basidiomycetes are decomposers of wood

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Figure 31.UN05

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Figure 31.18

Shelf fungi

Puffballs emitting spores

Maiden veil fungus (Dictyphora)

Figure 31.18a

Shelf fungi

Figure 31.18b

Puffballs emitting spores

Figure 31.18c

Maiden veil fungus (Dictyphora)

 The life cycle of a basidiomycete usually includes a long-lived dikaryotic mycelium In response to environmental stimuli, the mycelium reproduces sexually by producing elaborate fruiting bodies call basidiocarps Mushrooms are examples of basidiocarps The numerous basidia in a basidiocarp are sources of sexual spores called basidiospores

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Figure 31.19
Key Haploid (n) Dikaryotic (n + n) Diploid (2n) PLASMOGAMY

Dikaryotic mycelium

Mating type () Mating type (+ ) Haploid mycelia SEXUAL REPRODUCTION Dispersal and germination Basidiospores (n) Gills lined with basidia

Basidiocarp (n + n)

Basidium

Basidium with four basidiospores Basidium containing four haploid nuclei

Basidia (n + n)

KARYOGAMY MEIOSIS

1 m

Basidiospore

Diploid nuclei

Figure 31.19a

PLASMOGAMY Mating type () Mating type (+ ) Gills lined with basidia

Dikaryotic mycelium

Haploid mycelia

SEXUAL REPRODUCTION

Basidiocarp (n + n)

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

Figure 31.19b

Basidiocarp (n + n) SEXUAL REPRODUCTION Dispersal and germination Basidiospores (n) Basidium with four basidiospores Basidium containing four haploid nuclei MEIOSIS Key Haploid (n) Dikaryotic (n + n) Diploid (2n) Basidia (n + n) Gills lined with basidia

KARYOGAMY

Diploid nuclei

Figure 31.19c

Basidium

1 m

Basidiospore

 Basidiomycetes can produce mushrooms quickly Some species may produce fairy rings

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Figure 31.20

Concept 31.5: Fungi play key roles in nutrient cycling, ecological interactions, and human welfare
Fungi interact with other organisms as decomposers, mutualists, and pathogens

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Fungi as Decomposers
Fungi are efficient decomposers of organic material including cellulose and lignin They perform essential recycling of chemical elements between the living and nonliving world Fungi are also used in bioremediation projects

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Fungi as Mutualists
Fungi form mutualistic relationships with plants, algae, cyanobacteria, and animals All of these relationships have profound ecological effects

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Fungus-Plant Mutualisms
Mycorrhizae are enormously important in natural ecosystems and agriculture Plants harbor harmless symbiotic endophytes, fungi that live inside leaves or other plant parts Endophytes make toxins that deter herbivores and defend against pathogens Most endophytes are ascomycetes

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Figure 31.21

RESULTS Endophyte not present; pathogen present (EP+ ) Both endophyte and pathogen present (E+ P+ ) 30 20 10 0

Leaf area damaged (%)

Leaf mortality (%)

15 10 5 0

EP+

E+ P+

EP+

E+ P+

Fungus-Animal Symbioses
Some fungi share their digestive services with animals These fungi help break down plant material in the guts of cows and other grazing mammals Many species of ants use the digestive power of fungi by raising them in farms

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Figure 31.22

Lichens
A lichen is a symbiotic association between a photosynthetic microorganism and a fungus Millions of photosynthetic cells are held in a mass of fungal hyphae The photosynthetic component is green algae of cyanobacteria The fungal component is most often an ascomycete

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Figure 31.23

A foliose (leaflike) lichen Crustose (encrusting) lichens

A fruticose (shrublike) lichen

Figure 31.23a

Crustose (encrusting) lichens

Figure 31.23b

A foliose (leaflike) lichen

Figure 31.23c

A fruticose (shrublike) lichen

 The symbioses are so complete that lichens are given scientific names Algae or cyanobacteria occupy an inner layer below the lichen surface

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Figure 31.24

Ascocarp of fungus Fungal hyphae Algal layer

Soredia

50 m

Fungal hyphae Algal cell

Figure 31.24a

50 m

Fungal hyphae Algal cell

 The algae provide carbon compounds, cyanobacteria also provide organic nitrogen, and fungi provide the environment for growth The fungi of lichens can reproduce sexually and asexually Asexual reproduction is by fragmentation or the formation of soredia, small clusters of hyphae with embedded algae

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 Lichens are important pioneers on new rock and soil surfaces Lichens may have helped the colonization of land by plants 550600 million years ago Lichens are sensitive to pollution, and their death can be a warning that air quality is deteriorating

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Fungi as Pathogens
About 30% of known fungal species are parasites or pathogens, mostly on or in plants Each year, 10% to 50% of the worlds fruit harvest is lost due to fungi Some fungi that attack food crops are toxic to humans

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Figure 31.25

(b) Tar spot fungus on maple leaves

(a) Corn smut on corn (c) Ergots on rye

Figure 31.25a

(a) Corn smut on corn

Figure 31.25b

(b) Tar spot fungus on maple leaves

Figure 31.25c

(c) Ergots on rye

 Ergot of rye is caused by an ascomycete, and produces toxins More than 40,000 people died from an epidemic of ergotism during the middle ages Ergotism is characterized by gangrene, nervous spasms, burning sensations, hallucinations, and temporary insanity Ergots contain lysergic acid, the raw material for LSD

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 Animals are much less susceptible to parasitic fungi than are plants The chytrid Batrachochytrium dendrobatidis might be the cause of the recent decline in amphibians worldwide

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Figure 31.26

California

Sixty Lake Basin

2004

2005

Yellow-legged frogs killed by B. dendrobatidis infection Key

2006

2007

Boundary of chytrid spread Lake status in 2009: Frog population extinct Treatment lake: frogs treated with fungicides 08 and released 20

Figure 31.26a

California

Sixty Lake Basin

2004

2005

Key Boundary of chytrid spread Lake status in 2009: Frog population extinct Treatment lake: frogs treated with fungicides 8 and released 00 2

2006

2007

Figure 31.26b

Yellow-legged frogs killed by B. dendrobatidis infection

 The general term for a fungal infection in animals is mycosis Ringworm and athletes foot are examples a human mycoses Systemic mycoses spread through the body
For example, coccidioidomycosis produces tuberculosis-like symptoms

Some mycoses are opportunistic

For example, Candida albicans, which causes yeast infections

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Practical Uses of Fungi

Humans eat many fungi and use others to make cheeses, alcoholic beverages, and bread Some fungi are used to produce antibiotics for the treatment of bacterial infections
For example, the ascomycete Penicillium

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Figure 31.27

Staphylococcus

Penicillium Zone of inhibited growth

 Genetic research on fungi is leading to applications in biotechnology

For example, scientists are using Saccharomyces to study homologs of the genes involved in Parkinsons and Huntingtons diseases For example, insulin-like growth factor can be produced in the fungus Saccharomyces cerevisiae

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Figure 31.UN06

Fungal Phylum Chytridiomycota (chytrids) Zygomycota (zygote fungi) Glomeromycota (arbuscular mycorrhizal fungi) Ascomycota (ascomycetes, or sac fungi)

Distinguishing Features of Morphology and Life Cycles Flagellated spores

Resistant zygosporangium as sexual stage Arbuscular mycorrhizae formed with plants

Sexual spores (ascospores) borne internally in sacs called asci; vast numbers of asexual spores (conidia) produced Elaborate fruiting body (basidiocarp) containing many basidia that produce sexual spores (basidiospores)

Basidiomycota (basidiomycetes, or club fungi)

Figure 31.UN07

Soil Temp. 30C 35C 40C 45C

Curvularia Presence E E+ E E+ E E+ E E+

Plant Mass (g) 16.2 22.8 21.7 28.4 8.8 22.2 0 15.1

Number of New Shoots 32 60 43 60 10 37 0 24

Source: R. S. Redman et al., Thermotolerance generated by plant/fungal symbiosis, Science 298:1581 (2002).

Figure 31.UN08

Figure 31.UN09

Figure 31.UN10