basidiomycota mushrooms, bracket fungi, puffballs, rusts and smuts
TRANSCRIPT
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Basidiomycota
Mushrooms, bracket fungi, puffballs, rusts and smuts
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Basidiomycota
• Large phylum – 32,000 spp of higher fungi
• All produce a basidium – club shaped structure (site of karyogamy & meiosis) that produces haploid basidiospores
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Basidia• Basidia divided into two types depending on
whether they are septate• Septate basidia – phragmobasidia
– Septa can be transverse or longitudinal
• Aseptate basidia - holobasidia
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Hyphae• Hyphae are regularly
septate • Specialized type of pore
can occur – dolipore septum with parenthosome
• Allows cytoplasmic movement but prevents nuclear migration from one compartment to the next
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Basidiomycota• Three subphyla• Agaricomycotina – produce a basidioma,
(multicellular fruiting body), includes the mushrooms, brackets, puffballs, jelly fungi, stinkhorns (21400 spp)
• Pucciniomycotina – no basidioma, teliospore produces basidia, obligate plant parasites, the rusts (8400 spp)
• Ustilaginomycotina – no basidioma, teliospore produces basidia, facultative plant parasites, the smuts (1200 spp)
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Subphylum - Agaricomycotina
• Produce macroscopic basidiomata
• Most produce holobasidia, phragmobasidia produced by some jelly fungi
• Large and diverse class
• In life cycle, dikaryotic mycelium becomes the major vegetative mycelium – dikaryotic phase is extended over the Ascomycota
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Life cycle• Haploid basidiospores germinate to produce
haploid mycelium = primary mycelium
• In some species, the primary mycelium contains one nucleus in each compartment, in others, there may be multiple nuclei/compartment – also called monokaryon since it contains nuclei of one genotype
• Primary mycelium exhibits limited growth
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Life cycle
• Plasmogamy occurs shortly after basidiospore germination
• Most are heterothallic• Plasmogamy occurs between two compatible
hyphae (somatogamy) – no specialized sex cells
• Plasmogamy initiates dikaryotic phase or the secondary mycelium (dikaryon)
• After plasmogamy, nuclei migrate into monokaryotic cells to establish dikaryotic condition
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Dikaryon
• As mycelium grows, nuclei divide conjugately to maintain dikaryotic condition in cells
• Dolipore septum prevents nuclear migration so that each compartment contains two nuclei, pore is 0.1-0.2 μm in diameter
• Under certain conditions (e.g. after plasmogamy), dolipore septum breaks down to allow nuclear migration
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Dikaryon
• Secondary mycelium is the major vegetative phase of Basidiomycota
• Many also form clamp connections (formation is reminiscent of crozier formation in Ascomycota)
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Clamp connections
• Branch that grows back
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Clamp connections• Thought to be a mechanism to maintain
dikaryotic condition• Only found in dikaryotic hyphae, but not all
dikaryotic hyphae form them
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Dikaryon• Dikaryon grows – may reproduce asexually,
but asexual reproduction is not widespread in Agaricomycotina
• Favorable environmental conditions stimulate formation of basidiomata
• Multiple basidiomata can be produced from mycelium - one plasmogamy
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Basidioma• Composed entirely of dikaryotic hyphae (unlike
ascoma) • May be considerable tissue differentiation• Typically macroscopic, forms button then
expands
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Basidioma• Within the basidioma, cells in fertile regions – the
hymenium - will develop into basidia and produce basidiospores
• As the basidium develops, karyogamy and meiosis take place
karyogamy
meiosis
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Basidium• Sterigma (pl. –mata) –
projections from basidium to basidiospores
• Basidia may or may not produce sterigmata
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Basidioma• Basidia are produced in hymenia – fertile
regions in the basidioma (will discuss position in different basidiomata)
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Basidia• Basidia with
sterigmata can actively discharge basidiospores
• Basidiospores are positioned at a 45 angle
• 5-10 seconds before discharge a droplet forms
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Basidia• Thought that force to discharge spore comes
from droplet flowing around the spore from a hydrophobic region to a wettable region
• Change in center of mass of spore rapidly is thought to provide the force to discharge it
• Drop of fluid accompanies the spore
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Basidia• Spore is discharged less than a mm, then drops
straight down• Hymenia must be vertical with respect of gravity
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Basidioma
• Many basidiomata exhibit geotropism – Hymenium exhibits
positive geotropism– Basidioma exhibits
negative geotropism
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Basidioma• Basidiospores fall out of basidioma, into
turbulent air current and are carried away
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Basidioma• Composed entirely of dikaryotic hyphae• Basidioma is generally divided into 3
zones– Hymenium – layer of basidia with
basidiospores– Subhymenium – distinct area that gives rise to
hymenium– Trama – comprises the bulk of the basidioma,
separates hymenia– May also have cuticle – skin like layer on
surface of some
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Overall life cycle
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Representative Agaricomycotina• Jelly Fungi - Basidiomycetes that can be
grouped into the “jelly fungi” belong to 6 orders• Basidioma of these fungi is typically gelatinous –
during development hyphae in some tissues disintegrate and leave a gelatinous mass
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Jelly fungi• Most are saprotrophic on wood, some may be
parasitic• Most of little economic value• Some used as food – wood ear• Shape of basidiomata varies – cushion shaped,
ear shaped• May dry out to a hard, horny texture and then
become rewetted to become gelatinous again• Most produce phragmobasidia, those that
produce holobasidia have a unique morphology
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Auricularia– ear fungus, wood ear• Hymenium on
surface of basidoma
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Tremella• cushion shaped
basidioma that can vary in color (white, yellow, brown, black)
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Dacrymyces
• cushion shaped basidiomata that are frequently bright yellow
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Aphyllophorales• Large order - ~2000 spp.• Includes species that primarily produce dry,
woody, corky basidiomata (some may be fleshy)
• Produce holobasidia• Hymenium is on the surface of the
basidioma in some; surface may be smooth, ridged or spiny
• Hymenium may line tubes or gills – if the hymenium lines tubes or gills then the basidioma is always woody, never fleshy
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Wood decomposition
• White rots – lignin preferentially decomposed
• Brown rots – cellulose preferentially decomposed
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Corticium
• Simplest type of basidioma – thin, flattened, resupinate basidiomata that are on the surface of wood or bark
• Hymenium is flat and covers the surface of the basidioma (resembles paint)
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Clavaria – coral fungi
• Produce club shaped or branched basidiomata
• Hymenium covers surface of branches except at base
• Resemble coral, brightly colored, yellow, orange, violet
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Coral fungi
• Other species
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Toothed or hedgehog fungi
• Hymenium formed on surface of spines or teeth like projections that point down
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Toothed fungi
• Others
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Chanterelles • Form a funnel shaped basidioma with
hymenium on ridges on lower surface
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Chanterelles
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Polypores – bracket fungi
• Hymenium lines the surface of pores in woody basidiomata
• Some may be perennial
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Polypores
• Bracket fungi
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Polypores
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Agaricales
• Large order - ~5000 spp • Hymenium lines surfaces of tubes or gills
in fleshy basidiomata• Includes the mushrooms and boletes• Primarily saprotrophs in soil, litter or wood• Many are biotrophic – form mycorrhizal
associations with trees• Economically important as cultivated
mushrooms and mycorrhizae (used in forestry)
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Agaricales
• Basidiomata arise from extensive perennial mycelia
• Overall structure of basidioma
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Basidioma structure
• Basidia produced in hymenia on gills – flat plate like structures
• Cystidia extend out from hymenia
• In many, the gills are geotropic – parallel to force of gravity so that basidiospores can fall out once they leave the basidia
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Gills
• Increased surface area for greater number of basidia
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Basidioma development• Hemiangiocarpic development – button is
surrounded by a layer of tissue, the universal veil (forms the volva and scales) , also contains a partial veil (forms the annulus and cortina)
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Basidioma development
• Pseudoangiocarpic development – button has only a partial veil (forms annulus and cortina)
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Gymnocarpic development
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Gasteroid Basidiomycetes
• Many Basidiomycetes bear their basidiospores internally – gasteroid
• May or may not form a hymenium
• If a hymenium is formed, the basidioma remains closed until basidia form basidiospores
• All produce holobasidia
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Basidia
• Basidiospores not actively liberated from basidia – are formed symetrically on short or long sterigmata
• Symetrically placed basidiospores vs those at an angle
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Basidiomata• Are macroscopic• Enclosed by one or more layers of tissue –
peridium which may be papery, leathery, membranous or scaly
• Peridium surrounds the inner fertile tissue of the basidioma
– the gleba which
consists of trama
(sterile tissue) with
the basidia
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Gasteroid Basidiomycetes
• Produce macroscopic basidiomata
• Generally saprotrophic on litter, wood or dung
• Some are mycorrhizal
• Orders are separated on characteristics of the gelba – whether it is dry, fleshy, slimy or compartmentalized
• There are 8 orders, we will discuss 5
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Puffballs and earthstars• A hymenium is present but may be highly
convoluted to form a chamber like gleba
• At maturity, the glebal tissue disintegrates and becomes powdery and dry
• Certain hyphae (thick walled) persist and form a network – capillitium – interspersed with basidiospores
• Peridium is papery and has an pore - ostiole
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Puffballs
• Large range in size• Edible when young
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Puffballs
• Rain on peridium or kicking allows spores to be released
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Earthstars• Have a complex,
multilayered peridium• Thick layers split and
bend back to expose thin papery peridium layer
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Earth balls
• Pisolithus widely used as a mycorrhizal fungus to inoculate pines before they are planted
• Gleba is divided into locules within the basidioma
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Stinkhorns
• At maturity, the gleba becomes slimy and foul smelling
• Young stage resembles an egg, gleba surrounded by peridium
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Stinkhorns• Spongy tissue below gleba expands rapidly and
becomes the receptacle – stalk and breaks the peridium (becomes the volva) and lifts the gleba up
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Phallales• The gleba attracts flies that
disperse the spores• Other species may have
different forms
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• Gleba is divided into separate chambers, each of which is surrounded by its own peridium = peridioles
• Peridium is funnel shaped and leathery
Bird’s nest fungi
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Nidulariales
• Peridioles have a coiled cord – funiculus
• Basidiomata are small, up to a cm in diameter
• Saprotrophs on wood or dung
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Nidulariales
• Peridium acts as a splash cup
• Peridioles forced out of peridium when raindrop hits
• Funiculus attaches to objects
• Peridiole breaks open and spores dispersed
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Nidulariales
• Sphaerobolus – the fungal cannon
• Basidiomata are small – 2 mm diam
• Complex peridium – two cups
• Inner cup everts rapidly and catapults the gleba out – up to 6 m
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Sphaerobolus