2015s module6 print - university of san...

FUNGI

•  Key points: •  Heterotrophs that obtain nutrition by

absorption. •  Shared common ancestor of animal-like

protists with Animalia. •  Vital symbionts of plants. •  Classification based on reproductive

modes.

I. Mycology: Mushrooms, molds, and yeasts. General Characteristics

•  Eukaryotes. •  Predominant stage

haploid.


•  Both multicellular (hyphae made up of filaments) and unicellular (yeasts).

Hyphae made up of filaments are organized into mushroom body (mycelium and fruiting body)

Fruiting body

Mycelium


•  Not photosynthetic, all heterotrophic by absorption.

I. Mycology: Mushrooms, molds, and yeasts. Evolutionary History

•  Share common ancestor with animals and some Protists (to the exclusion of other Protista and Plants).

•  Ancient, fossils associated with earliest land plants, ca. 400mya.

–  Molecular data suggests the split between the lineage leading to fungi and the lineage leading to animals occurred ca. 1 bya.

•  Have diversified into five main lineages

Five ‘lineages’ of Fungi But Chytrids and

Zygomycetes both paraphyletic

‘Chytridiomycota’:euchytrids

‘Zygomycota’

microsporidia

1087 Coprinopsis cinerea480 Lycoperdon pyriforme626 Coprinus comatus

563 Clavaria zollingeri673 Amanita brunnescens

625 Pluteus romellii285 Cortinarius iodes564 Pleurotus ostreatus449 Armillaria mellea558 Flammulina velutipes

556 Marasmius alliaceus542 Ampulloclitocybe clavipes

557 Collybia tuberosa468 Henningsomyces candidus

729 Hygrocybe aff. conica439 Calostoma cinnabarinum

713 Boletellus projectellus714 Hygrophoropsis aurantiaca717 Suillus pictus

576 Fibulorhizoctonia sp.455 Echinodontium tinctorium682 Lactarius deceptivus

452 Bondarzewia montana492 Stereum hirsutum

447 Coltricia perennis688 Fomitiporia mediterranea

484 Phlebia radiata767 Climacodon septentrionalis776 Phanerochaete chrysosporium562 Grifola sordulenta701 Grifola frondosa770 Fomitopsis pinicola

518 Hyphoderma praetermissum700 Cotylidia sp.

466 Gautieria otthii724 Ramaria rubella471 Hydnum albomagnum

438 Calocera cornea454 Dacryopinax spathularia

867 Cintractia sorghi vulgaris870 Tilletiopsis sp.

865 Tilletiaria anomala675 Agaricostilbum hyphaenes

674 Rhodotorula hordea456 Endocronartium harknessii

1459 Puccinia graminis

138 Scutellospora heterogama139 Glomus mosseae

845 Glomus intraradices574 Geosiphon pyriformis

141 Mortierella verticillata144 Umbelopsis ramanniana

184 Phycomyces blakesleeanus1241 Rhizopus oryzae

136 Dimargaris bacillispora140 Coemansia reversa

1062 Orphella aff. haysii29 Smittium culisetae

185 Spiromyces aspiralis142 Rhopalomyces elegans145 Piptocephalis corymbifera

28 Entomophthora muscae

19 Physoderma maydis18 Coelomomyces stegomyiae

24 Polychytrium aggregatum27 Cladochytrium replicatum

21 Batrachochytrium dendrobatidis689 Rhizophydium macroporosum

43 Rhizophlyctis rosea182 Spizellomyces punctatus

635 Synchytrium macrosporum25 Monoblepharella sp.26 Hyaloraphidium curvatum

1068 Encephalitozoon cuniculi1089 Antonospora locustae

Ciona intestinalisHomo sapiens

Populus trichocarpaOryza sativa

Chlamydomonas reinhardtii

0.05 substitutions per site

to Ascomycota

709 Colacogloea peniophorae

‘Zygomycota’:Mucormycotina

Dikarya

Fungi

Y

Y

Y

Y

Y

Cyanidioschyzon merolaeArabidopsis thaliana

Phytophthora sojaeThalassiosira pseudonana

Monosiga brevicollis

Cryptosporidium parvumToxoplasma gondii

Dictyostelium discoideum

Drosophila melanogaster

Caenorhabditis elegans297 Rozella allomycis Y

300 Allomyces arbusculus20 Rhizoclosmatium sp.

638 Neocallimastix sp.

137 Conidiobolus coronatus‘Zygomycota’:

Entomophthorales

301 Basidiobolus ranarum633 Olpidium brassicae

844 Paraglomus occultum

539 Endogone pisiformis

710 Platygloea disciformis

1088 Cryptococcus neoformans505 Ustilago maydis

1078 Neurospora crassa216 Sordaria fimicola

1085 Podospora anserina217 Chaetomium globosum

1081 Magnaporthe grisea935 Diaporthe eres

952 Gnomonia gnomon51 Xylaria hypoxylon63 Xylaria acuta1082 Fusarium graminearum

161 Fusarium aff. solani186 Hydropisphaera erubescens

52 Hypocrea citrina914 Microascus trigonosporus

413 Lindra thalassiae424 Lulworthia grandispora

1 Leotia lubrica147 Coccomyces dentatus

744 Potebniamyces pyri151 Chlorociboria aeruginosa76 Mollisia cinerea

279 Monilinia fructicola59 Botryotinia fuckeliana

941 Dermea acerina166 Cudoniella clavus

49 Lachnum virgineum56 Geoglossum nigritum64 Trichoglossum hirsutum1004 Pleopsidium chlorophanum1005 Acarospora schleicheri1007 Acarospora laqueata

106 Echinoplaca strigulacea958 Diploschistes ocellatus

78 Acarosporina microspora398 Stictis radiata

296 Orceolina kerguelensis962 Trapelia placodioides224 Pertusaria dactylina

358 Dibaeis baeomyces645 Umbilicaria mammulata

687 Hypocenomyce scalaris134 Peltigera degenii

196 Mycoblastus sanguinarius639 Lecanora hybocarpa

6 Canoparmelia caroliniana3 Cladonia caroliniana

642 Bacidia schweinitzii84 Physcia aipolia1079 Aspergillus fumigatus

1080 Aspergillus nidulans426 Monascus purpureus

1083 Histoplasma capsulatum1084 Coccidioides immitis

430 Spiromastix warcupii657 Capronia pilosella668 Exophiala dermatitidis659 Ramichloridium anceps

669 Exophiala pisciphila684 Agonimia sp.91 Dermatocarpon miniatum661 Endocarpon pallidulum

697 Staurothele frustulenta342 Pyrgillus javanicus387 Pyrenula pseudobufonia

891 Peltula umbilicata892 Peltula auriculata896 Lichinella iodopulchra

101 Anisomeridium polypori1036 Trematosphaeria heterospora

1037 Westerdykella cylindrica283 Pyrenophora phaeocomes54 Cochliobolus heterostrophus940 Pleospora herbarum110 Trypethelium sp.

274 Dothidea sambuci921 Dothidea insculpta

939 Capnodium coffeae355 Dendrographa minor

126 Roccella fuciformis80 Simonyella variegata

148 Cheilymenia stercorea62 Scutellinia scutellata

65 Aleuria aurantia949 Pyronema domesticum50 Sarcoscypha coccinea

152 Caloscypha fulgens176 Gyromitra californica179 Disciotis sp.60 Morchella aff. esculenta66 Helvella compressa181 Ascobolus crenulatus507 Peziza vesiculosa71 Peziza proteana

905 Orbilia vinosa906 Orbilia auricolor

1069 Saccharomyces cerevisiae1070 Saccharomyces castellii1073 Candida glabrata

1071 Kluyveromyces waltii1072 Ashbya gossypii1075 Kluyveromyces lactis

1074 Candida albicans1269 Candida tropicalis

1270 Candida guilliermondii1077 Debaryomyces hansenii1268 Candida lusitaniae

1076 Yarrowia lipolytica1199 Schizosaccharomyces pombe

265 Taphrina wiesneri266 Protomyces inouyei

1192 Pneumocystis carinii

Pezizomycotina

Y

PhagotrophPhototrophLichenizedMycorrhizalPlant pathogenAnimal pathogenMycoparasiteInsect commensalSaprobeUncertain

GenomesequencedMotile cell stagepresent

SaprobeUncertain

PathogenMutualistPhototrophPhagotroph

Viridiplantae

Metazoa

Glomeromycota

Basidiomycota:Pucciniomycotina

Basidiomycota:Ustilaginomycotina

Basidiomycota:Agaricomycotina

100/100

100/92

100/73

100/71

--/85

‘Chytridiomycota’:Blastocladiales

...

...

RhodophytaStramenopiles

ApicomplexaMycetozoa

Choanoflagellida

‘Chytridiomycota’

‘Chytridiomycota’

100/80

100/100

100/81

100/94

100/93

100/98

100/94

Ascomycota:Lichinomycetes

Ascomycota:

Eurotiomycetes

Ascomycota:Lecanoromycetes

Ascomycota:Leotiomycetes

Ascomycota: Sordariom

ycetes

HOLMPA

IS?

SS

S

S

SS

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S

S

S

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SS

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PS A

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Ascomycota:Taphrinomycotina

Ascomycota:Saccharomycotina

Ascomycota: Orbiliomycetes

Ascomycota:Pezizomycetes

Ascomycota:Arthoniomycetes

Ascomycota:Dothideomycetes

Figure 1 | Phylogeny of the kingdom Fungi using bayesian analysis of thecombined, six-gene data set. Each fungal species begins with a unique‘Assembling the Fungal Tree of Life’ identifier, followed by genus andspecies. Indicated for each terminal taxon are: nutritional mode, whetherthey produce flagellated cells and if there is a genome sequence for the taxoncompleted or underway. Thickened branches indicate those that aresupported both by heterogeneous bayesian analysis (BPP $95%) and byMLBS ($70%). Almost every branch was supported by BPP and thus valuesare not shown. Where indicated, support values (percentage of trees in

agreement out of 58,611 trees) indicate BPP followed by MLBS. Branches areshaded according to reconstruction of nutritional mode. Microsporidiabranches have been shortened three times (double black break) to increasereadability. Red vertical ticks on branches indicate alternative placements ofmicrosporidia that might be significantly rejected (P , 0.05) and green ticksindicate placements that cannot be rejected. Quotation marks indicate non-monophyly of the taxon. The name ‘Mucormycotina’ will be validated in amanuscript that is in preparation.

ARTICLES NATURE | Vol 443 | 19 October 2006

820Nature Publishing Group ©2006

I. Mycology: Mushrooms, molds, and yeasts. The Absorptive Mode of Nutrition

•  Enzymes decompose organic materials.

•  Saprobes decompose and absorb nutrients from non-living organic matter.

•  Parasitic fungi absorb nutrients from cells of living hosts, some are pathogenic.

•  Mutualistic fungi also absorb from host organism, but reciprocate with beneficial functions, e.g. uptake of nutrients, minerals. –  Most plants depend on

mutualist relationship with fungi.

II. Structure: The mycelium •  Vegetative (non-reproductive)

body. •  Basic units are hyphae

–  Filaments of cells with a wall of chitin.

–  Septate cells (most common)

–  Aseptate & coenocytic cells •  Hyphae form interwoven mat

called the mycelium –  10 cm3 of soil can contain 1

km of hyphae.

•  Haustoria are specialized hyphae of parasitic fungi that penetrate plant cells.

Mycelium

II.  Structure: Reproductive body

•  Varies with taxonomic group

•  All reproduce by haploid spores in specialized structures peculiar to each group.

•  Evolution of fungi toward larger, more specialized spore-bearing structures.

Basidiomycete

Ascomycete

Zygomycete

II. Structure: Other Life Styles

•  Molds: Rapidly growing asexual stage. –  May develop sexual

stage as fruiting body distinctive of one of 5 phyla.

–  Some without known sexual stage, called imperfect fungi or deuteromycetes.

–  Important molds, e.g. Penicillium (penicillin, cheese)


•  Yeasts: Unicellular fungi in liquids or moisture including sap and animal tissue. –  Asexual cell division, or

budding; but some sexual ascomycetes and basidiomycetes are yeasts.

–  Saccharomyces: baker’s yeast, brewer’s yeast; active metabolically, release CO2 causes dough to rise, also ferments sugars to alcohol.

–  Candida: pathogenic yeast (vaginal, oral).


•  Mycorrhizae: “fungus roots” –  Important mutualism. –  Increases surface area

for absorption and exchange of nutrients.

–  Important in global Phosphorus cycle.

–  Over 95% of plants have mycorrhizae.

Mutualistic Symbioses


•  Mycorrhizae: “fungus roots” –  Ectomycorrhizae:

Hyphal sheath covers root and hartig net surrounds individual plant root cells.

•  Species of Zygomycetes, Ascomycetes, Basidiomycetes.




•  Mycorrhizae: “fungus roots” –  Arbuscular

mycorrhizae: Hyphae enter plant cells via invagination of plant cell membrane (do not enter protoplast--the interior of the cell).

•  Only and all species of Glomeromycota.


•  Endophytes live within plant tissues.

–  Provide fungi with protection, nutrients, water.

–  May receive from fungi chemical protection from insects, protists, bacteria, other fungi.

–  May receive ability to tolerate stressful environmental conditions.

–  Found in every plant studied to date (do not know role of all).

–  Primarily Ascomyceta. –  Taxol, effective anti-cancer

drug derived from endophyte of the Pacific Yew.



•  Lichens are mutualisms between fungi and photosynthetic organisms.

1.  Green Algae 2.  Cyanobacteria 3.  Yellow-Green Algae† 4.  Brown Alga (1 known case)†

•  Rarely three-fold: Fungus + photosynthetic green alga + nitrogen-fixing cyanobacteria.

•  Fungi provide mineral nutrients and water, algae provide carbohydrates via photosynthesis.

•  Ascomyceta (mostly), Basidiomyceta (a few), Glomeromyceta (1)



•  Insect cultivation: Fungi are cultivated by insects, provided with protection, fungi break down cellulose in plant tissue.

•  Internal cultivation: Termites




•  External cultivation: –  Leaf-cutter ants:

cultivate underground fungus gardens. Fungi break down cellulose in leaf tissue, ants eat fungus.



•  External cultivation: –  Bark and ambrosia

beetles: colonize new trees and inoculate carved galleries with ascomycete fungus. Fungi break down cellulose in wood, beetles consume fungus.

III. Growth & Reproduction •  Growth not in bulk, but by

proliferation of hyphae growing into resource (e.g. giant fairy ring)

•  Reproduction mostly asexual by spores or simple cloning, only chytrids with flagellar stage.

•  Spores dispersed by wind, water, animals.

III. Growth & Reproduction: Generalized life cycle

1.  Plasmogamy: fusion of cytoplasm of haploid hyphae.

2.  Dikaryon: cells with two haploid unfused nuclei.

3.  Karyogamy: fusion of nuclei, diploid stage; followed immediately by:

4.  Meiosis in haploid spore producing structures.

5.  Spores are released as haploid and germinate into filaments.

IV. Diversity

A.  Chytridiomycota (chytrids) B.  Zygomycota (zygote fungi) C.  Glomeromycota (arbuscular

mycorrhizae) D.  Ascomycota (sac or cup fungi) E.  Basidiomycota (club fungi)

A. Chytridiomycota (chytrids) •  ~1,000 species,

paraphyletic. •  Only fungi with flagellated

(spore) stage. •  A link between ancestral

protists and true fungi? •  Once excluded from Fungi,

but share biochemical characters, cell walls of chitin, absorptive mode of nutrition.

•  Mainly aquatic saprobes, some parasitic on plants and animals.

B. Zygomycota (zygote fungi) •  ~1,000 species •  Terrestrial, soil, decaying

plant/animal tissue. •  Mycorrhizae: mutualistic

association with plant roots.

•  Hyphae coenocytic. •  Zygosporangia are the

reproductive structures that give the name.

•  E.g. Rhizopus stolonifer, black bread mold.

C. Glomeromycota (arbuscular mycorrhizae)

•  160 species, once considered Ascomycota, but now recognized as distinct.

•  Nearly all form arbuscular mycorrhizae with plants--ecologically important. –  Ca. 90% of all plant

species have them as mutualists!

D. Ascomycota (sac or cup fungi)

•  Include such tasties as truffles and morels.

•  Unicellular to complex multicellularity.

•  Some extreme plant pathogens; other important saprobes.

•  Half symbiotic with Chlorophyta: lichens.

D. Ascomycota (sac or cup fungi)

•  Karyogamy in an ascocarp; sexual spore in asci.

•  Asexual: spores (conidia) come from specialized structures (conidiophores), wind dispersed.

E. Basidiomyceta (club fungi)

•  Include the commonly encountered mushrooms, shelf fungi, puffballs.

•  Basidium (“little pedestal”) transient diploid stage.

•  Most important plant (wood) decomposers.

E. Basidiomyceta (club fungi)

•  Karyogamy in basidiocarp (sexual); puff balls release spores explosively.

•  Most complex of fungi: long-lived dikaryotic mycelium.

•  Fairy rings; giant ring in Michigan ~40 acres (!); genetically uniform mycelium.

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