We are fun-guys! Figure 31.1 Can you spot the largest organism in this forest?

There’s fungus among us! 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!

Fungi are heterotrophs that feed by absorption Despite their diversity, fungi share key traits, most importantly the way in which they derive nutrition

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

Fungi exhibit diverse lifestyles: Decomposers Parasites Mutualists

Animation: Fungal Reproduction and Nutrition 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 Nutrition

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

Reproductive structure Fig. 31-2 Reproductive structure Hyphae Spore-producing structures Figure 31.2 Structure of a multicellular fungus 20 µm Mycelium

Specialized Hyphae in Mycorrhizal Fungi Some unique fungi have specialized hyphae called haustoria that allow them to penetrate the tissues of their host

ascomycete Fig. 31-4 Hyphae Nematode 25 µm (a) Hyphae adapted for trapping and killing prey Plant cell wall Fungal hypha Figure 31.4 Specialized hyphae Plant cell Plant cell plasma membrane Haustorium (b) Haustoria

Lasso sequence at 3 minutes Fig. 31-4a Hyphae Nematode 25 µm Figure 31.4 Specialized hyphae (a) Hyphae adapted for trapping and killing prey Lasso sequence at 3 minutes

Morgellons Disease? Figure 31.4 Specialized hyphae

Plant cell Fungal hypha wall Plant cell Plant cell plasma membrane Fig. 31-4b Plant cell wall Fungal hypha Plant cell Figure 31.4 Specialized hyphae Plant cell plasma membrane Haustorium (b) Haustoria

Mycorrhizae are mutually beneficial relationships between fungi and plant roots Two types:

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

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

Haploid (n) Heterokaryotic (unfused nuclei from different parents) Fig. 31-5-1 Key Haploid (n) Heterokaryotic (unfused nuclei from different parents) Diploid (2n) Spore-producing structures Spores ASEXUAL REPRODUCTION Mycelium Figure 31.5 Generalized life cycle of fungi GERMINATION

Haploid (n) Heterokaryotic stage Heterokaryotic (unfused nuclei from Fig. 31-5-2 Key Haploid (n) Heterokaryotic stage Heterokaryotic (unfused nuclei from different parents) PLASMOGAMY (fusion of cytoplasm) Diploid (2n) KARYOGAMY (fusion of nuclei) Spore-producing structures Zygote SEXUAL REPRODUCTION Spores ASEXUAL REPRODUCTION Mycelium Figure 31.5 Generalized life cycle of fungi GERMINATION

Haploid (n) Heterokaryotic stage Heterokaryotic (unfused nuclei from Fig. 31-5-3 Key Haploid (n) Heterokaryotic stage Heterokaryotic (unfused nuclei from different parents) PLASMOGAMY (fusion of cytoplasm) Diploid (2n) KARYOGAMY (fusion of nuclei) Spore-producing structures Zygote SEXUAL REPRODUCTION Spores ASEXUAL REPRODUCTION Mycelium Figure 31.5 Generalized life cycle of fungi MEIOSIS GERMINATION GERMINATION 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 (“sexes”) Fungi use sexual signaling molecules called pheromones to communicate their mating type (like many animals including primates!)

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”.

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

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

Fig. 31-6 2.5 µm Figure 31.6 Penicillium, a mold commonly encountered as a decomposer of food

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

Sacchromyces cervisiae Fig. 31-7 10 µm Parent cell Figure 31.7 The yeast Saccharomyces cerevisiae in several stages of budding (SEM) Bud Sacchromyces cervisiae

Many molds and yeasts have no known sexual stage

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

The Origin of Fungi Fungi, animals, and their protistan relatives form the opisthokonts clade

UNICELLULAR, FLAGELLATED Nucleariids ANCESTOR Opisthokonts Chytrids Fig. 31-8 Animals (and their close protistan relatives) UNICELLULAR, FLAGELLATED ANCESTOR Nucleariids Opisthokonts Chytrids Figure 31.8 Fungi and their close relatives Fungi Other fungi

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

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

Figure 31.11 Fungal diversity Hyphae 25 µm Chytrids (1,000 species) Zygomycetes (1,000 species) Fungal hypha Glomeromycetes (160 species) Ascomycetes (65,000 species) Figure 31.11 Fungal diversity Basidiomycetes (30,000 species)

Chytrids (1,000 species) Hyphae 25 µm Fig. 31-11a Figure 31.11 Fungal diversity

They can be decomposers, parasites, or mutualists 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: Allomyces Zoospore Release Video: Phlyctochytrium Zoospore Release

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes Fig. 31-UN1 Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Fig. 31-12 Figure 31.12 Flagellated chytrid zoospore (TEM) Flagellum 4 µm

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

Zygomycetes (1,000 species) Fig. 31-11b Zygomycetes (1,000 species) Figure 31.11 Fungal diversity

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

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes Fig. 31-UN2 Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

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

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

Fig. 31-14 Figure 31.14 Pilobolus aiming its sporangia Fungus Cannon! 0.5 mm

Glomeromycetes (160 species) Fig. 31-11c Fungal hypha Glomeromycetes (160 species) Figure 31.11 Fungal diversity

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

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes Fig. 31-UN3 Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Branched hyphae inside root cell (cleared) Fig. 31-15 Figure 31.15 Arbuscular mycorrhizae Branched hyphae inside root cell (cleared) 2.5 µm

Ascomycetes (65,000 species) Fig. 31-11d Ascomycetes (65,000 species) Figure 31.11 Fungal diversity

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

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 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

Ascomycetes include plant pathogens, decomposers, and symbionts Ascomycetes reproduce asexually by enormous numbers of asexual spores called conidia

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

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes Fig. 31-UN4 Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Ascomycetes: The Good the Bad and the Ugly Part I: The Good Morchella esculenta, the tasty morel Tuber melanosporum, a truffle Figure 31.16 Ascomycetes (sac fungi)

Morchella esculenta, the tasty morel Fig. 31-16a Figure 31.16 Ascomycetes (sac fungi)

ca. $600/lb Tuber melanosporum, a truffle Fig. 31-16b Figure 31.16 Ascomycetes (sac fungi) 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).

Alexander Fleming’s “moldy plate” (1928) Fig. 31-26 Alexander Fleming’s “moldy plate” (1928) Staphylococcus Penicillium Zone of inhibited growth Figure 31.26 Fungal production of an antibiotic

Sacchromyces cervisiae Fig. 31-7 10 µm Parent cell Figure 31.7 The yeast Saccharomyces cerevisiae in several stages of budding (SEM) Bud Sacchromyces cervisiae

6,000 years old Starter = old dough Figure 31.16 Ascomycetes (sac fungi) 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.

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!

Ascomycota: Claviceps (“ergot”) Parasitic on grains of grasses: (rye, barley, wheat)

Ascomycota: Claviceps (“ergot”) Parasitic on grains of grasses: (rye, barley, wheat) infects flowers and replaces mature grain with sclerotia sclerotia

Ascomycota: Claviceps (“ergot”) Parasitic on grains of grasses: (rye, barley, wheat) infects flowers and replaces mature grain with sclerotia hard sclerotia allow the fungus to survive freezing and desiccation sclerotia

Ascomycota: Claviceps (“ergot”) Parasitic on grains of grasses: (rye, barley, wheat) infects flowers and replaces mature grain with sclerotia hard sclerotia allow the fungus to survive freezing and desiccation “argot” = spur sclerotia

Alkaloids produced by Claviceps purpurea: Ergotamine - vasoconstrictor

Alkaloids produced by Claviceps purpurea: Ergotamine - vasoconstrictor Ergonovine - spontaneous abortions

Alkaloids produced by Claviceps purpurea: Ergotamine - vasoconstrictor Ergonovine - spontaneous abortions Ergine - gastrointestinal toxin, seizures, formication

Alkaloids produced by Claviceps purpurea: Ergotamine - vasoconstrictor Ergonovine - spontaneous abortions Ergine - gastrointestinal toxin, seizures, formication Lysergic Acid Hydroxyethylamide - hallucinations

Ascomycota: Claviceps (“ergot”) “Holy Fire” (Ignis Sacer)

Ascomycota: Claviceps (“ergot”) “Holy Fire” (Ignis Sacer) 994 - 40,000 died in France

Ascomycota: Claviceps (“ergot”) “Holy Fire” (Ignis Sacer) 994 - 40,000 died in France 1093 - “St. Anthony’s Fire”

Ascomycota: Claviceps (“ergot”) “Holy Fire” (Ignis Sacer) 994 - 40,000 died in France 1093 - “St. Anthony’s Fire” 1692 - Salem Witch Trials

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

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)

“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

Cordyceps fly Ascomycetes Part II: The Ugly Bullet ants in trouble Figure 31.4 Specialized hyphae moth

Geomyces destructans White nose Syndrome Causes early arousal? Up to 97% mortality Now in Missouri

Conidiophores w/ conidia colony conidia Conidiophores w/ conidia

Basidiomycetes (30,000 species) Fig. 31-11e Basidiomycetes (30,000 species) Figure 31.11 Fungal diversity

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

Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes Fig. 31-UN5 Chytrids Zygomycetes Glomeromycetes Ascomycetes Basidiomycetes

Fig. 31-18 Maiden veil fungus (Dictyphora), a fungus with an odor like rotting meat Puffballs emitting spores Shelf fungi, important decomposers of wood Figure 31.18 Basidiomycetes (club fungi)

(eat slime pass spores) Fig. 31-18a Maiden veil fungus (Dictyphora), a fungus with an odor like rotting meat Figure 31.18 Basidiomycetes (club fungi) Fly dispersed (eat slime pass spores)

Puffballs emitting 70 trillion! spores Fig. 31-18b Figure 31.18 Basidiomycetes (club fungi) 70 trillion!

Shelf fungi, important decomposers of wood Fig. 31-18c Figure 31.18 Basidiomycetes (club fungi)

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

Dikaryotic mycelium Haploid mycelia Mating type (–) Mating type (+) Fig. 31-19-1 Dikaryotic mycelium Haploid mycelia PLASMOGAMY Mating type (–) Mating type (+) SEXUAL REPRODUCTION Figure 31.19 The life cycle of a mushroom-forming basidiomycete Key Haploid (n) Dikaryotic (n +n) Diploid (2n)

Dikaryotic mycelium Haploid mycelia Mating type (–) Mating type (+) Fig. 31-19-2 Dikaryotic mycelium Haploid mycelia PLASMOGAMY Mating type (–) Mating type (+) Gills lined with basidia SEXUAL REPRODUCTION Basidiocarp (n+n) Basidia (n+n) Figure 31.19 The life cycle of a mushroom-forming basidiomycete Key Haploid (n) Dikaryotic (n +n) Diploid (2n)

Dikaryotic mycelium Haploid mycelia Mating type (–) Mating type (+) Fig. 31-19-3 Dikaryotic mycelium Haploid mycelia PLASMOGAMY Mating type (–) Mating type (+) Gills lined with basidia SEXUAL REPRODUCTION Basidiocarp (n+n) Basidia (n+n) Figure 31.19 The life cycle of a mushroom-forming basidiomycete KARYOGAMY Key Haploid (n) Dikaryotic (n +n) Diploid nuclei Diploid (2n)

Dikaryotic mycelium Haploid mycelia Mating type (–) Mating type (+) Fig. 31-19-4 Dikaryotic mycelium Haploid mycelia PLASMOGAMY Mating type (–) Mating type (+) Gills lined with basidia Haploid mycelia SEXUAL REPRODUCTION Basidiocarp (n+n) Dispersal and germination Basidiospores (n) Basidium with four basidiospores Basidia (n+n) Basidium Figure 31.19 The life cycle of a mushroom-forming basidiomycete Basidium containing four haploid nuclei KARYOGAMY MEIOSIS Key Haploid (n) Dikaryotic (n +n) Diploid nuclei 1 µm Basidiospore Diploid (2n)

Fig. 31-20 Figure 31.20 A fairy ring

Concept 31.5: Fungi play key roles in nutrient cycling, ecological interactions, and human welfare Fungi interact with other organisms in many ways

Fungi as Decomposers Fungi are efficient decomposers They perform essential recycling of chemical elements between the living and nonliving world

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

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

30 15 20 10 10 5 E–P+ E+P+ E–P+ E+P+ RESULTS Fig. 31-21 RESULTS Endophyte not present; pathogen present (E–P+) Both endophyte and pathogen present (E+P+) 30 15 20 10 Leaf mortality (%) Leaf area damaged (%) Figure 31.21 Do endophytes benefit a woody plant? 10 5 E–P+ E+P+ E–P+ 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 and termites use the digestive power of fungi by raising them in “farms”

Fig. 31-22 Figure 31.22 Fungus-gardening insects For the Discovery Video Leafcutter Ants, go to Animation and Video Files.

Figure 31.22 Fungus-gardening insects For the Discovery Video Leafcutter Ants, go to Animation and Video Files.

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

Crustose (encrusting) lichens A foliose (leaflike) lichen Fig. 31-23  Crustose (encrusting) lichens A fruticose (shrublike) lichen  A foliose (leaflike) lichen Figure 31.23 Variation in lichen growth forms

A fruticose (shrublike) lichen Fig. 31-23a A fruticose (shrublike) lichen Figure 31.23 Variation in lichen growth forms

Crustose (encrusting) lichens Fig. 31-23b Figure 31.23 Variation in lichen growth forms

A foliose (leaflike) lichen Fig. 31-23c Figure 31.23 Variation in lichen growth forms

The fungal component of a lichen is most often an ascomycete Algae or cyanobacteria occupy an inner layer below the lichen surface

Ascocarp of fungus Soredia Fungal hyphae Algal layer Algal cell Fig. 31-24 Ascocarp of fungus Soredia Fungal hyphae Algal layer Figure 31.24 Anatomy of an ascomycete lichen (colorized SEM) Algal cell 20 µm Fungal hyphae

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

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

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

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

Fig. 31-UN6

Fig. 31-UN6a

Fig. 31-UN6b

Fig. 31-UN6c

Fig. 31-UN6d

Fig. 31-UN6e