Basidiomycota Mushrooms, bracket fungi, puffballs, rusts and smuts

Basidiomycota Large phylum – 32,000 spp of higher fungi All produce a basidium – club shaped structure (site of karyogamy & meiosis) that produces haploid basidiospores

Basidia Basidia divided into two types depending on whether they are septate Septate basidia – phragmobasidia –Septa can be transverse or longitudinal Aseptate basidia - holobasidia

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

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)

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

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

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

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 μm in diameter Under certain conditions (e.g. after plasmogamy), dolipore septum breaks down to allow nuclear migration

Dikaryon Secondary mycelium is the major vegetative phase of Basidiomycota Many also form clamp connections (formation is reminiscent of crozier formation in Ascomycota)

Clamp connections Branch that grows back

Clamp connections Thought to be a mechanism to maintain dikaryotic condition Only found in dikaryotic hyphae, but not all dikaryotic hyphae form them

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

Basidioma Composed entirely of dikaryotic hyphae (unlike ascoma) May be considerable tissue differentiation Typically macroscopic, forms button then expands

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

Basidium Sterigma (pl. –mata) – projections from basidium to basidiospores Basidia may or may not produce sterigmata

Basidioma Basidia are produced in hymenia – fertile regions in the basidioma (will discuss position in different basidiomata)

Basidia Basidia with sterigmata can actively discharge basidiospores Basidiospores are positioned at a 45 angle 5-10 seconds before discharge a droplet forms

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

Basidia Spore is discharged less than a mm, then drops straight down Hymenia must be vertical with respect of gravity

Basidioma Many basidiomata exhibit geotropism –Hymenium exhibits positive geotropism –Basidioma exhibits negative geotropism

Basidioma Basidiospores fall out of basidioma, into turbulent air current and are carried away

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

Overall life cycle

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

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

Auricularia– ear fungus, wood ear Hymenium on surface of basidoma

Tremella cushion shaped basidioma that can vary in color (white, yellow, brown, black)

Dacrymyces cushion shaped basidiomata that are frequently bright yellow

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

Wood decomposition White rots – lignin preferentially decomposed Brown rots – cellulose preferentially decomposed

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)

Clavaria – coral fungi Produce club shaped or branched basidiomata Hymenium covers surface of branches except at base Resemble coral, brightly colored, yellow, orange, violet

Coral fungi Other species

Toothed or hedgehog fungi Hymenium formed on surface of spines or teeth like projections that point down

Toothed fungi Others

Chanterelles Form a funnel shaped basidioma with hymenium on ridges on lower surface

Chanterelles

Polypores – bracket fungi Hymenium lines the surface of pores in woody basidiomata Some may be perennial

Polypores Bracket fungi

Polypores

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)

Agaricales Basidiomata arise from extensive perennial mycelia Overall structure of basidioma

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

Gills Increased surface area for greater number of basidia

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)

Basidioma development Pseudoangiocarpic development – button has only a partial veil (forms annulus and cortina)

Gymnocarpic development

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

Basidia Basidiospores not actively liberated from basidia – are formed symetrically on short or long sterigmata Symetrically placed basidiospores vs those at an angle

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

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

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

Puffballs Large range in size Edible when young

Puffballs Rain on peridium or kicking allows spores to be released

Earthstars Have a complex, multilayered peridium Thick layers split and bend back to expose thin papery peridium layer

Earth balls Pisolithus widely used as a mycorrhizal fungus to inoculate pines before they are planted Gleba is divided into locules within the basidioma

Stinkhorns At maturity, the gleba becomes slimy and foul smelling Young stage resembles an egg, gleba surrounded by peridium

Stinkhorns Spongy tissue below gleba expands rapidly and becomes the receptacle – stalk and breaks the peridium (becomes the volva) and lifts the gleba up

Phallales The gleba attracts flies that disperse the spores Other species may have different forms

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

Nidulariales Peridioles have a coiled cord – funiculus Basidiomata are small, up to a cm in diameter Saprotrophs on wood or dung

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

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

Sphaerobolus