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Page 1: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

We are fun-guys!

Page 2: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

• Fungi are diverse and widespread

• 100,000 described species

• Really as many as 1.5 MILLION?

• Honey mushroom - 2000 years old, 1,800 football fields (underground) or a yeast cell

• Essential in ecosystem nutrient recycling

• Mycology

There’s fungus among us!

Page 3: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

There’s fungus among us!

QuickTime™ and aYUV420 codec decompressor

are needed to see this picture.

Page 4: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fungi are heterotrophs that feed by absorption

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

Page 5: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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 fungi’s ecological success

Page 6: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

• Fungi exhibit diverse lifestyles:

– Decomposers

– Parasites

– Mutualists

Page 7: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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

Animation: Fungal Reproduction and NutritionAnimation: Fungal Reproduction and Nutrition

Page 8: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fungal Morphology

• The morphology of multicellular fungi enhances their ability to absorb nutrients

• Fungi consist of mycelia, networks of branched hyphae adapted for absorption

• Most fungi have cell walls made of chitin

Page 9: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-2

Reproductive structure

Spore-producingstructures

Hyphae

Mycelium

20 µm

Page 10: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Specialized Hyphae in Mycorrhizal Fungi

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

Page 11: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-4

(b) Haustoria

(a) Hyphae adapted for trapping and killing prey

Nematode

Plantcellwall

Haustorium

Plant cellplasmamembrane

Plant cell

Fungal hypha

Hyphae25 µm

ascomycete

Page 12: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-4a

(a) Hyphae adapted for trapping and killing prey

NematodeHyphae

25 µm

Lasso sequence at 3 minutes

Page 13: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

MorgellonsDisease?

Page 14: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-4b

(b) Haustoria

Plantcellwall

Haustorium

Plant cellplasmamembrane

Plant cell

Fungal hypha

Page 15: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

• Mycorrhizae are mutually beneficial relationships between fungi and plant roots

• Two types:

Page 16: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

• 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

Page 17: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fungi produce spores through sexual or asexual life cycles

• Fungi propagate themselves by producing vast numbers of spores, either sexually or asexually

• Fungi have different types of life cycles

Page 18: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-5-1

Spores

Spore-producingstructures

GERMINATION

ASEXUALREPRODUCTION

Mycelium

Key

Heterokaryotic(unfused nuclei fromdifferent parents)

Haploid (n)

Diploid (2n)

Page 19: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-5-2

Spores

Spore-producingstructures

GERMINATION

ASEXUALREPRODUCTION

Mycelium

Key

Heterokaryotic(unfused nuclei fromdifferent parents)

Haploid (n)

Diploid (2n)

SEXUALREPRODUCTION

KARYOGAMY(fusion of nuclei)

PLASMOGAMY(fusion of cytoplasm)

Heterokaryoticstage

Zygote

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Fig. 31-5-3

Spores

Spore-producingstructures

GERMINATION

ASEXUALREPRODUCTION

Mycelium

Key

Heterokaryotic(unfused nuclei fromdifferent parents)

Haploid (n)

Diploid (2n)

SEXUALREPRODUCTION

KARYOGAMY(fusion of nuclei)

PLASMOGAMY(fusion of cytoplasm)

Heterokaryoticstage

Zygote

Spores

GERMINATION

MEIOSIS

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Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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 (“sexes”)

• Fungi use sexual signaling molecules called pheromones to communicate their mating type (like many animals including primates!)

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• Plasmogamy is the union of 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

• The different nuclei may even swap DNA similar to “crossing over”.

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• Hours, days, or even centuries may pass before the occurrence of karyogamy, nuclear fusion (kary = nucleus)

• During karyogamy, the haploid nuclei fuse, producing diploid cells

• The diploid phase is short and undergoes meiosis (halving!), producing haploid spores

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Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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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Fig. 31-6

2.5 µm

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• Other fungi that can reproduce asexually are yeasts, which inhabit moist environments

• Instead of producing spores, yeasts reproduce asexually by simple cell division and the pinching of “bud cells” from a parent cell

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Fig. 31-7

10 µm

Parentcell

Bud

Sacchromyces cervisiae

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• Many molds and yeasts have no known sexual stage

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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

Page 30: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

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The Origin of Fungi

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

Page 31: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

Fig. 31-8

Animals (and their closeprotistan relatives)

Other fungi

Nucleariids

Chytrids

UNICELLULAR,FLAGELLATEDANCESTOR

Fu

ng

i

Op

istho

kon

ts

Page 32: We are fun-guys!. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fungi are diverse and widespread 100,000 described

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• The oldest undisputed fossils of fungi are only about 460 million years old

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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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Fig. 31-11

Chytrids (1,000 species)

Zygomycetes (1,000 species)

Hyphae 25 µm

Glomeromycetes (160 species)

Fungal hypha

Ascomycetes (65,000 species)

Basidiomycetes (30,000 species)

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Fig. 31-11a

Chytrids (1,000 species)

Hyphae 25 µm

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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

Video: Video: AllomycesAllomyces Zoospore Release Zoospore Release Video: Video: PhlyctochytriumPhlyctochytrium Zoospore Release Zoospore Release

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Fig. 31-UN1

Chytrids

Basidiomycetes

Zygomycetes

Glomeromycetes

Ascomycetes

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Fig. 31-12

Flagellum

4 µm

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• Until recently, systematists thought that fungi lost flagella only once in their evolutionary history

• Molecular data indicate that some “chytrids” are actually more closely related to another fungal group, the zygomycetes; chytrids are a paraphyletic group

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Fig. 31-11b

Zygomycetes (1,000 species)

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Zygomycetes

• The zygomycetes (phylum Zygomycota) exhibit great diversity of life histories

• They include fast-growing molds, parasites, and commensal symbionts

• The zygomycetes are named for their sexually produced zygosporangia

• Zygosporangia, which are resistant to freezing and drying, can survive unfavorable conditions

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Fig. 31-UN2

Chytrids

Basidiomycetes

Zygomycetes

Glomeromycetes

Ascomycetes

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• The life cycle of black bread mold (Rhizopus stolonifer) is fairly typical of the phylum

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• Some zygomycetes, such as Pilobolus, can actually “aim” their sporangia toward conditions associated with good food sources

• Toward light = grass = cows = dispersal!

• 25m/sec = 1,million times its length/sec.

• Fastest aerial acceleration in nature

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Fig. 31-11c

Glomeromycetes (160 species)

Fungal hypha

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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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Fig. 31-UN3

Chytrids

Basidiomycetes

ZygomycetesGlomeromycetes

Ascomycetes

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Fig. 31-15

2.5 µmBranched hyphae inside root cell (cleared)

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Fig. 31-11d

Ascomycetes (65,000 species)

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Ascomycetes

• Ascomycetes (phylum Ascomycota) live in marine, freshwater, and terrestrial habitats

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Ascomycetes

• Ascomycetes (phylum Ascomycota) live in marine, freshwater, and terrestrial habitats

• The phylum is defined by production of sexual spores in saclike asci, usually contained in fruiting bodies called ascocarps

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Ascomycetes

• Ascomycetes (phylum Ascomycota) live in marine, freshwater, and terrestrial habitats

• The phylum is defined by production of sexual spores in saclike asci, usually 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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• Ascomycetes include plant pathogens, decomposers, and symbionts

• Ascomycetes reproduce asexually by enormous numbers of asexual spores called conidia

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• 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

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Fig. 31-UN4

Chytrids

Basidiomycetes

ZygomycetesGlomeromycetes

Ascomycetes

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Tuber melanosporum, a truffle

Morchella esculenta,the tasty morel

Ascomycetes: The Good the Bad and the Ugly Part I: The Good

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Fig. 31-16a

Morchella esculenta,the tasty morel

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Fig. 31-16b

Tuber melanosporum, a truffle ca. $600/lb

The Associate Press reported that a real estate investor and his wife from Hong Kong have paid €125,000 ($160,406 USD) for a gigantic

Italian White Alba truffle which is reportedly the world’s most expensive ever. The most expensive truffle weighs in 1.51 kilograms

(3.3 lbs).

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Fig. 31-26

Staphylococcus

Zone ofinhibitedgrowth

Penicillium

Alexander Fleming’s “moldy plate” (1928)

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Fig. 31-7

10 µm

Parentcell

Bud

Sacchromyces cervisiae

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A seal around 4,000 years old is a Sumerian "Hymn to Ninkasi", the goddess of brewing. This

"hymn" is also a recipe for making beer.

6,000 years oldStarter = old dough

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Ascomycetes Part II: The Bad

• Mycoses - Ringworm, indoor molds (sick buildings), athelete’s foot, Candida (diaper rash and worse)Candida albicans

• Fusarium (grains; Jurgenson research)

• Cryptococcus (pigeon poop)

• Ergot!

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Ascomycota: Claviceps (“ergot”)• Parasitic on

grainsof grasses: (rye, barley, wheat)

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Ascomycota: Claviceps (“ergot”)

sclerotiasclerotia

• Parasitic on grainsof grasses: (rye, barley, wheat)

• infects flowers and replaces mature grain with sclerotia

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Ascomycota: Claviceps (“ergot”)

sclerotiasclerotia

• Parasitic on grainsof grasses: (rye, barley, wheat)

• infects flowers and replaces mature grain with sclerotia

• hard sclerotia allow the fungus to survive freezing and desiccation

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Ascomycota: Claviceps (“ergot”)

sclerotiasclerotia

• Parasitic on grainsof grasses: (rye, barley, wheat)

• infects flowers and replaces mature grain with sclerotia

• hard sclerotia allow the fungus to survive freezing and desiccation

• “argot” = spur

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Alkaloids produced by Claviceps purpurea:

• Ergotamine - vasoconstrictor

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Alkaloids produced by Claviceps purpurea:

• Ergotamine - vasoconstrictor

• Ergonovine - spontaneous abortions

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Alkaloids produced by Claviceps purpurea:

• Ergotamine - vasoconstrictor

• Ergonovine - spontaneous abortions

• Ergine - gastrointestinal toxin, seizures, formication

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Alkaloids produced by Claviceps purpurea:

• Ergotamine - vasoconstrictor

• Ergonovine - spontaneous abortions

• Ergine - gastrointestinal toxin, seizures, formication

• Lysergic Acid Hydroxyethylamide - hallucinations

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Ascomycota: Claviceps (“ergot”)

• “Holy Fire” (Ignis Sacer)

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Ascomycota: Claviceps (“ergot”)

• “Holy Fire” (Ignis Sacer)

• 994 - 40,000 died in France

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Ascomycota: Claviceps (“ergot”)

• “Holy Fire” (Ignis Sacer)

• 994 - 40,000 died in France

• 1093 - “St. Anthony’s Fire”

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Ascomycota: Claviceps (“ergot”)

• “Holy Fire” (Ignis Sacer)

• 994 - 40,000 died in France

• 1093 - “St. Anthony’s Fire”

• 1692 - Salem Witch Trials

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Ascomycota: Claviceps (“ergot”)

• “Holy Fire” (Ignis Sacer)

• 994 - 40,000 died in France

• 1093 - “St. Anthony’s Fire”

• 1692 - Salem Witch Trials

• 1777 - 8000 in Sologne, France

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Ascomycota: Claviceps (“ergot”)

• “Holy Fire” (Ignis Sacer)

• 994 - 40,000 died in France

• 1093 - “St. Anthony’s Fire”

• 1692 - Salem Witch Trials

• 1777 - 8000 in Sologne, France

• 1943 - Lysergic acid diethylamide (LSD-25: Albert Hofmann)

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• “Holy Fire” (Ignis Sacer)

• 994 - 40,000 died in France

• 1093 - “St. Anthony’s Fire”

• 1692 - Salem Witch Trials

• 1777 - 8000 in Sologne, France

• 1943 - Lysergic acid diethylamide (LSD-25: Albert Hofmann)

• 1951 - Pont-St. Esprit, France

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25 µm

Ascomycetes Part II: The Ugly

Cordyceps

Bullet ants in trouble

fly

moth

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Geomyces destructans

White nose Syndrome

Causes early arousal?Up to 97% mortality

Now in Missouri

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colony

conidia Conidiophores w/ conidia

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Fig. 31-11e

Basidiomycetes (30,000 species)

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Basidiomycetes

• Basidomycetes (phylum Basidiomycota) include mushrooms, puffballs, and shelf fungi, mutualists, 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

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Fig. 31-UN5

Chytrids

Basidiomycetes

ZygomycetesGlomeromycetes

Ascomycetes

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Fig. 31-18

Shelf fungi, importantdecomposers of wood

Maiden veil fungus(Dictyphora), afungus with anodor like rottingmeat

Puffballs emittingspores

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Fig. 31-18a

Maiden veil fungus(Dictyphora), afungus with an odor like rotting meat

Fly dispersed(eat slime pass spores)

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Fig. 31-18b

Puffballs emittingspores

70 trillion!

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Fig. 31-18cShelf fungi, importantdecomposers of wood

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• 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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Fig. 31-19-1

SEXUALREPRODUCTION

Diploid (2n)

Haploid (n)Dikaryotic (n +n)

Key

PLASMOGAMY

Matingtype (+)

Haploid myceliaDikaryotic mycelium

Matingtype (–)

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Fig. 31-19-2

SEXUALREPRODUCTION

Diploid (2n)

Haploid (n)Dikaryotic (n +n)

Key

PLASMOGAMY

Matingtype (+)

Haploid myceliaDikaryotic mycelium

Matingtype (–)

Basidia(n+n)

Gills linedwith basidia

Basidiocarp(n+n)

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Fig. 31-19-3

SEXUALREPRODUCTION

Diploid (2n)

Haploid (n)Dikaryotic (n +n)

Key

PLASMOGAMY

Matingtype (+)

Haploid myceliaDikaryotic mycelium

Matingtype (–)

Basidia(n+n)

Gills linedwith basidia

Basidiocarp(n+n)

KARYOGAMY

Diploidnuclei

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Basidium

Fig. 31-19-4

SEXUALREPRODUCTION

Diploid (2n)

Haploid (n)Dikaryotic (n +n)

Key

PLASMOGAMY

Matingtype (+)

Haploid myceliaDikaryotic mycelium

Matingtype (–)

Basidia(n+n)

Gills linedwith basidia

Basidiocarp(n+n)

KARYOGAMY

Diploidnuclei

MEIOSIS

Basidium containingfour haploid nuclei

Dispersal andgermination

Basidiospores(n)

Basidium withfour basidiospores

Basidiospore1 µm

Haploid mycelia

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Fig. 31-20

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Concept 31.5: Fungi play key roles in nutrient cycling, ecological interactions, and human welfare

• Fungi interact with other organisms in many ways

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Fungi as Decomposers

• Fungi are efficient decomposers

• They perform essential recycling of chemical elements between the living and nonliving world

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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 that live inside leaves or other plant parts

• Endophytes make toxins that deter herbivores and defend against pathogens

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Fig. 31-21

Both endophyte and pathogen present (E+P+)Endophyte not present; pathogen present (E–P+)

Le

af

are

a d

am

ag

ed

(%

)

Le

af

mo

rta

lity

(%

) 30

1020

10

15

5

00E+P+E–P+ E–P+E+P+

RESULTS

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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 and termites use the digestive power of fungi by raising them in “farms”

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Fig. 31-22

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QuickTime™ and aYUV420 codec decompressor

are needed to see this picture.

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Lichens

• A lichen is a symbiotic association between a photosynthetic microorganism and a fungus in which millions of photosynthetic cells are held in a mass of fungal hyphae

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Fig. 31-23

 A foliose(leaflike)lichen

A fruticose (shrublike) lichen

 Crustose(encrusting)lichens

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Fig. 31-23a

A fruticose (shrublike) lichen

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Fig. 31-23b

Crustose(encrusting)lichens

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Fig. 31-23c

A foliose(leaflike)lichen

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• The fungal component of a lichen is most often an ascomycete

• Algae or cyanobacteria occupy an inner layer below the lichen surface

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Fig. 31-24

Algal cell

Ascocarp of fungusSoredia

Fungal hyphae

Fungalhyphae Algal

layer

20 µ

m

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• The algae provide carbon compounds, cyanobacteria 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 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

• Some fungi that attack food crops are toxic to humans

• Animals are much less susceptible to parasitic fungi than are plants

• The general term for a fungal infection in animals is mycosis

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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

• Genetic research on fungi is leading to applications in biotechnology

– For example, insulin-like growth factor can be produced in the fungus Saccharomyces cerevisiae

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Fig. 31-UN6

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Fig. 31-UN6a

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Fig. 31-UN6b

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Fig. 31-UN6c

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Fig. 31-UN6d

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Fig. 31-UN6e