phylum basidiomycota

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Phylum Basidiomycota The Club Fungi Eric Swann and David S. Hibbet plus notes from Alexopoulos and Blackwell

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Phylum Basidiomycota The

Club FungiEric Swann and David S. Hibbet plus

notes from Alexopoulos and Blackwell

Fungi comprising the phylum Basidiomycota are commonly known as basidiomycetes

Basidiomycetes are a large and diverse lot

Includes forms commonly as mushrooms, boletes, puffballs, earthstars, stinkhorns, bird’s nest fungi, jelly fungi, bracket or shelf fungi, and rust and smut fungi

Introduction

huge impact on human affairs and ecosystem functioning. Many Basidiomycota obtain nutrition by decaying dead organic matter, including wood and leaf litter and thus play a significant role in the carbon cycle. Unfortunately, Basidiomycota frequently attack the wood in buildings and other structures, which has negative economic consequences for humans.

Well-developed symbiotic life styles (intimate associations with other living organisms): rusts, smuts, mycorrhizae, associations with insects,

Importance

Basidiomycetes that attack dead woody plants are the principal agents that decay cellulose and lignin and are esssential components of the ecosystem

Some are used as biological pulping and bleaching agents in pulp and paper production and in removal of toxic substances from the environment

Important mycorrhizal species in natural and forest ecosystems

Importance, cont.

Smut and rust fungi destroy crops annually◦Stinking smut and black stem rust of wheat

◦Many others attack food and ornamental plants

◦Cause diseases of forest and shade trees

◦Destroy a variety of wood products inc. lumber, railroad ties, landscape timbers, utility poles

Importance, cont.

 Mushrooms, both cultivated and wild, are eaten in many countries

 toxin phalloidin (from the mushroom Amanita phalloides) binds actin, which is a component of microfilaments. Fluorescent stains that incorporate phalloidin are used by cell biologists to visualize the cytoskeleton

hallucinogens, which are produced by members of the genus Psilocybe (and other groups) - used in Central American indigenous cultures as a spiritual tool, and are now cultivated for the illicit drug trade

 astaxanthin, a red pigment produced by the basidiomycetous yeast Phaffia (used to add color to farmed salmon)

certain enzymes from wood-decaying Basidiomycota that have potential applications in paper production and bioremediation (decontamination of polluted environments using biological agents).

Importance, cont.

 unicellular or multicellular, sexual or asexual, and terrestrial or aquatic. 

 Basidiomycota are so variable that it is impossible to identify any morphological characteristics that are both unique to the group and constant in the group

The most diagnostic feature is the production of basidia (sing. basidium), which are the cells on which sexual spores are produced, and from which the group takes its name

Characteristics

Produce their sexual spores termed basidiospores on the outside of a specialized, microscopic, spore-producing structure called the basidium

The basiodiospores are ballistospores and are discharged forcibly from their basidia by means of an elaborate discharge mechanism

Basidiospores occur in a variety of sizes, shapes, colors and may be thin/thick, smooth or ornamental

Characteristics

A long-lived dikaryon, in which each cell in the thallus contains two haploid nuclei resulting from a mating event, is another characteristic feature.

clamp connections are a kind of hyphal outgrowth that is unique to Basidiomycota, although they are not present in all Basidiomycota. 

Characteristics

http://tolweb.org/tree/ToLimages/hymenium7.gif

SEM of the surface of a mushroom gill showing several basidia, some with four basidiospores attached

One of the most fascinating characteristics of Basidiomycota is the production of forcibly discharged ballistospores (Fig. 2), which are propelled into the air from the sterigma. Ballistospores may be sexual or asexual, and may be produced by basidia, hyphae, yeast cells, or even other ballistospores.

Ballistospory is associated with the production of a liquid filled "hilar droplet" that forms at the base of the spore, just above its attachment to the sterigma 

 Spore discharge occurs when the hilar droplet fuses with a film of liquid on the surface of the spore. The rapid coalescence of the liquids causes a sudden shift in the center of mass of the spore and contributes to its release from the sterigma. This mechanism has been termed a "surface tension catapult" and it results in spores being discharged with a force of about 25,000 g 

Mating in Basidiomycota involves fusion of haploid cells, but fusion of the nuclei is usually delayed until the basidia are formed. Thus, the dominant phase of the life cycle in most Basidiomycota is a dikaryon, in which the two nuclei brought together in mating exist side-by-side in each cell (Fig. 3A). Sometimes a dikaryon can donate a nucleus to a uninucleate "monokaryon", resulting in a "di-mon" mating.

Clamp connections are hyphal outgrowths that form when cells in dikaryotic hyphae divide. One of the nuclei divides in the main axis of the hypha, while the other divides into the clamp (Fig. 3B). Septa are formed across each of the mitotic spindles. The apex of the backward-growing monokaryotic clamp cell fuses with the subapical cell, reestablishing the dikaryotic condition (Fig. 3C). All fungi that produce clamp connections are members of the Basidiomycota, but not all Basidiomycota produce clamp connections.

Agaricomycotina one of three major clades of Basidiomycota. The

Agaricomycotina contains roughly 20,000 described species, which is almost 70% of the (known) Basidiomycota. About 98% of the species of the Agaricomycotina are in a clade called the Agaricomycetes, which includes mushrooms, bracket fungi, puffballs, and others. The other major groups are the Tremellomycetes and Dacrymycetes. These latter groups include "jelly fungi" (Fig. 1), which have gelatinous, often translucent fruiting bodies (e.g., "witches butter" Tremella mesenterica), as well as many yeast-forming species (Hibbett 2007).

Filobasidiella (Cryptococcus) neoformans

Calocera viscosaGomphus

http://tolweb.org/tree/ToLimages/Calocera_viscosa.jpg

http://tolweb.org/tree/ToLimages/Filobasidiella_neoformans.jpg

http://tolweb.org/tree/ToLimages/Gomphus.jpg

The class Ustilaginomycetes comprises more than 1400 species of basidiomycetous plant parasites, which are distributed in approximately 70 genera. They occur throughout the world, although many species are restricted to tropical, temperate or arctic regions. Some species of Ustilago and Tilletia, e.g. the barley, wheat or maize smut fungi, are well known because they are of economic importance. For example, from 1983 to 1988 the barley smut fungi reduced annual yields by 0.7% to 1.6% in the prairie provinces in central Canada, causing average annual losses of about U.S. $8,000,000 (Thomas 1989).

Ustilaginomycotina

Tilletia contraversa is important in the international wheat trade (Trione 1982), and 2-5% of the plants in a corn field are generally infected by Ustilago maydis, while up to 80% of a field can be infected if conditions are good for the smut fungus. On the other hand, the galls of U. maydis are considered a delicacy in the Mesoamerican tradition. They are known in Mexico as "Huitlacoche" and in the U.S.A. as "maize mushroom", "Mexican truffles" or "caviar azteca" (Valverde et al. 1995).

Cont.

Galls on the roots of Juncus articulatus induced by Entorrhiza casparyana

Corn galls on Zea mays induced by Ustilago maydis

Sporulation of Exobasidium vaccinii on Vaccinium vitis-idaea

 remarkably diverse, possessing a wide range of structural and ecological characters.

these fungi have in common, however, is their rather miniscule size, making them challenging to locate and study.

contains approximately 7,400 species, distributed among approximately 215 genera, but will eventually be much larger as many cryptic species are recognized. More than 95% of the species and 75% of the genera in this group are placed in the Pucciniales (Pucciniomycetes), the plant parasitic rust fungi.

Pucciniomycotina

The rusts are undeniably the most economically important fungi in this clade, as obligate parasites on a wide range of crop plants including cereal grains, legumes, and trees such as coffee, apple, and pine, where they can cause extensive reduction in yield and even host death. Other phytopathogenic Pucciniomycotina can be of economic importance, but on a much smaller scale than rust fungi.

Cont.

Species of Helicobasidium (Pucciniomycetes: Helicobasidiales) cause root rot diseases of many economically important plants such as asparagus, beet, mulberry, and pear. Microbotryum violaceum   (Microbotryomycetes: Microbotryales) causes anther smut of plants in the Caryophyllaceae, while smut on buckwheat is caused by Sphacelotheca fagopyri (Microbotryomycetes: Microbotryales). These smuts are not closely related to the smuts in the Ustilaginomycotina with which they were confused until molecular sequence data became available.

Cont.

Eocronartium muscicola

fruitbodies on moss host

Hemileia vastatrix, Coffee Rust on coffee leaf

Helicobasidium purpureum on Picea abies (L.) Karst.

Order 1. TREMELLALES (Jelly fungi)◦Basidiocarp has gelatinous consistency

◦Basidia of various shapes: deeply divided like a tuning fork, longitudinally septate or transversely septate

Philippine representative species

◦Examples:Dacryopinax – a very common jelly fungus. The fruiting body is shaped like a spoon, flattened on one end, and yellowish to orange in color. It grows on recently cut logs and is often seen on newly-built fences.

Tremella – found on dead wood in secondary growth forests

PRS, cont.

◦Examples, cont. Auricularia (taingang daga) Basidiocarps are ear-shaped or flattened and the basidia are found on the smooth side. Used as an ingredient in vegetable dishes, noodles and soups

The flattened type is tough, unpalatable and is not eaten

Septobasidium Basidiocarp is not gelatinous and grows in association with aphids

PRS, cont.

Auricularia sp.

http://www.kaimaibush.co.nz/Fungi/Auricularia4b.jpg

Microstructures of S. meredithiae. A. Cross section showing subiculum with pillars supporting the upper layer. B. Empty probasidial cells with basidia and spores. C. Irregularly coiled haustoria. Bars: A = 50 μm, B = 10 μm, C = 10 μm.

Microstructure of Septobasidium sp. A. Cross section of thallus showing dense context slightly looser at the surface. B. Punctate probasidia like structures C. Spindle-shape haustoria. Bars: A = 50 μm, B = 10 μm, C = 10 μm.

Order 2. UREDINALES (rusts)◦This group includes parasites on higher plants

◦So-called because of the conspicuous reddish color of the spores when they appear on plant parts in large masses

◦Complicated life history◦Some species involve 5 different kinds of spores and their production may involve 2 completely unrelated plant hosts

PRS, cont.

Hemileia Forms large patches of reddish to golden yellow spore on the upper surface of coffee leaves

Order 3. USTILAGINALES (Smuts)◦So-named because of the color of their spores (black like soot or smut)

◦The spores form a large, black ball in the ear of the corn

PRS, rusts, cont.

under surface of affected leaf, x 1⁄2; 2, section through same showing mycelium, haustoria, and a spore-cluster; 3, a spore-cluster seen from below; 4, a uredospore; 5, germinating uredospore; 6, appressorial swellings at tips of germ-tubes; 7, infection through stoma of leaf; 8, teleutospores; 9, teleutospore germinating with promycelium and sporidia; 10, sporidia and their germination (2 after Zimmermann, 3 after Delacroix, 4–10 after Ward)

http://www.cluesheet.com/images/image125.jpg

http://www.extension.umn.edu/yardandgarden/YGLNews/images2/Aug107/corn_smut_darker.jpg

For years, scientists have assumed that huitlacoche (WEET-LA-KO-CHEE) — a gnarly, gray-black corn fungus long-savored in Mexico — had nutritional values similar to those of the corn on which it grew. But test results just published in the journal Food Chemistry reveal that an infection that U.S. farmers and crop scientists have spent millions trying to eradicate, is packed with unique proteins, minerals and other nutritional goodies.

Order 4. THELEPORALES ◦Basidiocarps have a smooth, fertile layerCorticium (painted stick)Looks like paint (white or pink) poured over the substrate

Layer of fungal tissue lying flat on the surface of twigs or rotting logs

PRS, cont.

http://www.mycokey.com/asco/2005/asco2005/album/slides/JHP-05.030-CorticiumRose02.jpg

http://t1.gstatic.com/images?q=tbn:ANd9GcSHuHtKI5c7GtAXKcszW1sVcTnt1xMlo-AlRp5MKTDG51XB-3p94w

Corticium sp.

Stereum Similar to Coticium but basidiocarp is

semi-upright or resupinate Common on decaying wood or twigs

Cymatoderma Basidiocarps are erect and stalked Funnel-shaped and always found on

wood Upper side is brownish and the white,

vein-like configuration on the underside shows through

Hymenium is found in the lower side of the fruiting body

PRS, cont.

http://t0.gstatic.com/images?q=tbn:ANd9GcRDBP7zPdbbIfPb8pO-nPKDIDxCOj_8eCa-DPv5gwqfMFiwd0_C

http://www.biopix.com/photos/jcs-stereum-rugosum-50904.jpg

Stereum sp.

Cymatoderma sp.

http://4.bp.blogspot.com/_zKDjcLXZBTI/TEz7g50K4QI/AAAAAAAADUg/kW424WzAcD0/s1600/Cymatoderma_elegans_Funnel_fungi_leathery_02.jpg

http://2.bp.blogspot.com/_zKDjcLXZBTI/TEz7rRtaIZI/AAAAAAAADUo/Y7PtKVU0Fr0/s400/Cymatoderma_elegans_Funnel_fungi_leathery_01.jpg

Order 5. CLAVARIALES (Coral Fungi)◦Look like sea corals◦Fingerlike projections are simple or branched, and small to medium in size

◦Some species are colorless, others dark or light brown, red, blue and yellow

◦The hymenium covers the outer surface of the branches

PRS, cont.

Clavulina Simple and fingerlike; grows from the soil

Deflexula Small and occurs in tiny clumps on the

surface of the bark Lentaria

Rounded branches are much branched Occurs on twigs and trunks

Scitinopogon Has a highly-branched flattened fruiting

body which develops from the soil◦Some are edible; none known to be poisonous

but some may be bitter in taste

PRS, cont.

Clavulina sp.

http://www.mushroomexpert.com/images/kuo3/clavulina_rugosa_01.jpg

http://www.nature-diary.co.uk/nn-images/0908/090825-clavulina-rugosa.jpg

Deflexula sp.

http://www.aebletoften.dk/jhp/fungi/Fungal%20Shapes/album/slides/DeflexulaSp.jpg

Scitinopogon sp.

http://www.svims.ca/council/illust/Lentaria%20pinicola%205%20Michael%20Beug.jpg

Lentaria sp.

http://3.bp.blogspot.com/_tpzJtMaDjnc/TNvOG5u2mEI/AAAAAAAABV0/7lPUixyDdG4/s1600/Lentaria%2Bsp.%2Bcopy.jpg

Order 6. POLYPORALES (Shelf or bracket fungi)◦From “polypore” meaning many holes◦The fertile side (hymenium) of the basidiocarp of a polypore is covered with many little pores lined by the basidia

◦They are distinguished from each other by the characteristic shape of their pores

PRS, cont,

Certain polypores cause ‘heart rot’ – hollow centers caused by mycelia rotting the middle portion of large dipterocarp trees of rain forests

Among the most common fungi in the Philippines

Important as wood rotters

Polyporales, cont.

Poria – flattened on the surface of the substrate

Daedalea – fruiting body is tough and white Polyporus – the most common genus;

rounded pores on the underside of erect, usually stalked basidium

Hexagona – 5-sided pores Fomes – woody, perennial polypore with

new hymenial layer consisting of many long tubes added to the fruiting body each season, forming concentric rings making the fruiting body resemble a shelf or bracket

Polyporales, cont.

http://upload.wikimedia.org/wikipedia/en/7/7a/Tuckahoe.jpg

Wolfiporia extensa

sclerotium (called "Tuckahoe", or Indian bread)

Polyporus sp.

http://www.treasurekeepers.rs/Library/forests/banjsuma/polyporus_squamosus.jpg

Fomes sp.

http://us.123rf.com/400wm/400/400/lianem/lianem0804/lianem080400155/3106218-fomes-fomentarius-wood-fungus-on-dead-wood.jpg

Hexagona sp.

http://www.banjorah.com/images/Photo%20Gallery/Fungus/Fungus%2025.jpg

Order 7. HYDNALES (tooth fungi)◦Look much like theleporoids or polypores but at closer look, have flattened, tooth-like projections covered by the hymeniumHydnum – somewhat tough and corallike

PRS, cont.

Hydnum sp.

http://www.mykoweb.com/CAF/photos/Hydnum_repandum(bk-01).jpg

Order 8. AGARICALES (mushrooms)◦The most easily recognized group of fungi

◦With large, umbrella-like fruiting bodies

◦Found in forest floor, cultivated or abandoned fields, secondary forests, lawns

◦Grow well in soil, dead wood, and on other decaying plant parts

◦The basidiocarp of the Agaricales is called a sporosphore

PRS, CONT.

The sporosphore is first formed as a small , compact knot of cells on mycelium already existing on the soil or other substrates then expands to four or five times its original size

The gills bear numerous basidiospores

Agaricales, cont.

How do we distinguish an edible mushroom from poisonous ones?

The only sure answer: KNOW THE MUSHROOM BEFORE EATING IT.

Rule of thumb: Poisonous mushrooms are said to have a cup at the base of the stem, a ring on the stem, and spots on the cap.

Some delicious mushrooms have one or a combination of these

Some poisonous mushrooms do not have any of the three

Agaricales, cont.

Philippine mushrooms: Schizophyllum (split leaf) Commonly found throughout the Philippines

Grows on any type of dead wood Gills consist of two halves which are easily pulled apart

Upper side of white, fan-shaped fruiting body has meaty appearance

http://www.mykoweb.com/CAF/photos/Schizophyllum_commune(bab-2).jpg

http://www.elfram.com/fungi/fungi_s/schizophyllum_commune_1_ng7.jpg

Schizophyllum sp.

Marasmius one of the most frequently encountered mushrooms in the forest

The sporophores are often very tiny, with long stems, and are sometimes called “horsehair fungi”

The fungus threads penetrate twigs and sticks lying on the forest floor

Agaricales, cont.

http://farm1.static.flickr.com/222/505981098_c427fe887d.jpg

http://www.lincstrust.org.uk/species/fungi/fairy_l.jpg

Marasmius sp.

Panus Somewhat tough and has white spots on its brownish cap

Has lateral stem, grows on wood and is edible

Agaricales, cont.

Panus sp.

Volvariella - (kabuteng saging) a most desirable mushroom; grows

throughout the Philippines Grows on cut banana stalks; can be

cultivated on rice straw, or water lilies Cap is brownish and silky because of

the many long hairs occuring in a layer on the outer surface. The hairs are also present on stem and on the cup found at its base.

Gills are white when young, becoming pink when mature

Agaricales, cont.

Volvariella sp.

Collybia (Kabuting punso) An edible mushroom growing in the Batangas-

Laguna area in the months of July and August. White with brown tinge to the cap Has a long, tapering stem which extends far

below the surface of the soil Often seen growing on anthills The white ants apparently become attracted to

the mycelium for its sweet, sticky fluid and cultivate the mycelium in a chamber deep within the ant nest. The young mushrooms form int hese chambers and expand as they push their way out to the surface of the ground

Agaricales, cont.

Collybia sp..

Tricholoma A very large mushroom Sporophores grow in clusters on the soil

Fruting body is white; has no ring or cup on stem

Caps are edible but stems are somewhat tough and usually filled with insect larvae

Agaricales, cont.

Tricholoma sp. (Matsutake mushroom)

http://www.bayareamushrooms.org/images/pictures/matsutake.jpg

Chlorophyllum (laughing mushroom) With green gills and spores Cap with brown scales and a ring around the stem

Grows in garbage piles The white-gilled “buttons” are sometimes mistaken for those of the edible ones

Persons who eat this mushroom by mistake later go into an uncontrollable fit of laughter that may last for a long time

Agaricales, cont.

Chlorophyllum sp.

Chlorophyllum sp. Fairy ring

Order 9. LYCOPERDALES (Puffballs or Devil’s Snuff box)◦ Basidiospores are produced inside the basidiocarp◦ Basidiospores are not exposed to the air in

structures such as a gill

PRS, cont.

http://botit.botany.wisc.edu/toms_fungi/images/quarter.jpg

http://blogs.scottarboretum.org/gardenseeds/wp-content/uploads/2010/11/Calvatia-gigantea-Giant-Puffball-JWC.jpg

Geastrum sp.

http://fireflyforest.net/images/firefly/2006/August/Geastrum.jpg

Tulostoma sp.

Scleroderma sp.

Simblum sp.

Dictyophora sp.http://www.anhs.com.au/images/Stinkhorn.jpg

http://fiddlehead.smugmug.com/Fungi/Mushroom-Photos/DSC01222-copy/661770_WVNz5-M-1.jpg

http://t1.gstatic.com/images?q=tbn:ANd9GcThyy1oGDeViInVf2GMHDHzvl0Q4kDe25Ec5-iurOo6Tz86FnRXDw

The "eggs" in the nest are actually analogous to tiny puffballs in that they contain the basidiospores inside of them. The nest acts as a splash cup. When raindrops fall into the cup, the eggs (technically called peridioles) are splashed out some distance away from the nest. If they land on a suitable substrate, they will germinate and form new mycelium that can grow into new fruiting bodies. When immature, the nest is covered by a brown membrane called an epiphragm, which breaks before the rain can splash out the eggs.

http://www.sciencephoto.com/image/14631/large/B2501781-_Splash_cups_bird_s_nest_fungus,_Cyathus_striatus_-SPL.jpg

http://www.sciencephoto.com/image/14144/large/B2501285-Fungus_on_wooden_log-SPL.jpg

A=peridioles (eggs), B=funiculus and hapteron (cord and anchor)

http://www.mycokey.com/MycoKeySolidState/pictures/basi/gast/Nidularia/defo15S.jpg

http://calphotos.berkeley.edu/imgs/128x192/0000_0000/1208/0736.jpeg

http://www.anbg.gov.au/fungi/images/0141.jpg

The EndThank you