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PAT 601 TAXONOMY OF FUNGI (2+1) An Introduction to Mycology R.S. Mehrotra, K.R. Aneja Introductory Mycology- C. J.

Alexopoulous Fungi include eucaryotic spore bearing achlorophyllous organisms that generally reproduce asexually and sexually and usually filamentous-branched somatic structures are typically surrounded by cell walls containing Chitin or Cellulose or both of these substances together with many other organic molecules.

Classification: In the world today, about 4,00,000 species of plants have already been described. Algae 20,000 sp. Fungi 1,00,000 sp. Liverworts and Mosses 25,000 Ferns 10,500 Phanerogams 2,50,000 Saccardo (1884) classified fungi into six classes. Schizomycetes (Bacteria) Myxomycetes (slime moulds) Phycomycetes Ascomycetes Basidiomycetes Deuteromycetes At one time some mycologists placed slime moulds in animal kingdom (Bessey, 1950) but today it was accepted under Fungi. Ainsworth (1966) treated fungi bas a separate kingdom or as kingdom of plant kingdom with two divisions Myxomycota or plasmodial form and Eumycota for non-plasmodial form. Eumycota then divided into five subdivisions

History: In 400 B.C. itself Buddha noticed diseases of plants caused by fungi. There are some records are available on plant diseases in the Vedas (1200 B.C.) about mildews and blasts. Clusius (1526-1609) wrote a book Rariorium Plantarum Historia. This book has 28 pages. Gaspard- Plnax Theatri Botanici (1623) includes 100 species of fungi, Agaricaceae, Boletaceae, Clavariaceae, Auriculariaceae, Lycoperdaceae, Phallaceae, Pezziaceae. Robert Hooke- Micrographa (1687) include Phragmodium and Mucor. Joseph P. Tournefort (1694) wrote Element de Botanique. He studied Boletus, Agaricales, Clathrus, Morchella, Aurucularia, Polyporus, Lycoperdon, Tubercules. Butler (1874-1944) studied Sexuality in Fungi. Tillet (1755) - Seed treatment 1766- Grain Smut. History of Mycology in India: Buddha 400 B.C. noticed diseases 1200 B.C. in the Vedas period, rust, smuts, Agaricales. About 10,000 species of fungi have been reported from India. Butler and Prisby (1931), Vasudeva (1960), Mundkar (1938), Vasudeva (1962), Subramanium and Ramakrishnan (1956), Tandan and Sudir Chandra (1963), Tilak and Rao (1970), mukerji and Juneja (1974), Bilgrami et al (1979) listed the Fungi and reported. Landmark in the study of Mycology: Linnaeus book Species Plantarum. Carl Linnaeus (1707-1778), Sweedish botanist brought description of all known species of plants. He established Binomial System of Nomenclature, Generic and Species. Modern classification was done by C.H. Persson. Christian Hendrix Persoon (17611836), South American. 1. Observations Mycologicae (1796-1799). 2. Tentamen diespositionis Methodicae Fungorum (1797) 3. Icones description Fungorum (1798) 4. Synopsis Methodica Fungorum (1801) 5. Mycologia Europaea (1822-1828)

Elias Magnus Fries (1794-1878) called Father of Systematic Mycology wrote Systema Mycologicum- Three Volumes. Saccardos Sylloge Fungorum (1882-1931)- 26 volumes. Listed out all genera and species of Fungi. De Bary (1866)- Father of Modern Mycology. He discovered Heteroecism in rust, Puccinia graminis tritici (1830-1840). Late blight-1846-Epidemic-Ireland- Famine. De Bary only proved the damage due to Phytophthora infestans. Woronin (1838-1903)- Club root. Bessey(1950), Sparrow (1958), Martin(1961), Alexopoulos and Mims (1974), Hawksworth et al. (1983). At one time some botanist placed slime moulds in animal kingdom and called Mycetozoa. But today placed in fungi (Myxomycetes-multinucleate, free living, naked plasmodium, zoospore, heterokont, whiplash type of flagella). Ainsworth placed myxomycetes as a separate division (plasmodial forms). The classification of fungi is still in a state of flux. Stable for ideal system is yet to be proposed by taxonomists. In the 7th edition of the Dictionary of Fungi by Hawksworth:; Sutton and Ainsworth (1983) brought out some important change in the classification of Fungi. Separate class uredinomycetes and ustilaginomycetes have been formed instead of teliomycetes. Britishers like Cunningham and Barclay (1871) Mucorales, Ustilaginales and Uredinales were listed by Cunningham. Barclay (1886) investigated wheat rust in India and Simla. Sir, Edwin John Butler (1874-1943)- Father of Indian Mycology initiated and organized Mycological and Phytopathological research in India. He worked in Imperial (IARI) at PUSA. Born in Ireland, 1874. He graduated in Medicine. He worked first on Pythium and published in India and discovered the genus Allomyces. He wrote a book Fungi and Diseases in plants(1918). Butler and Prisby The Fungi of India(1931). K.C. Mehta-(1892-1940)- Rust- Berberis and Mahonia Mundkur (1896-1952)- Fungi and Plant Diseases and Ustilaginales of India.

McRae, Subramanium, C.V., Sundaraman, T.S. Ramakrishnan, K.Ramakrishnan, M.J.Thirumalachar.

According to Saccardo in his Sylloge Fungorum 1. Acrasiomycetes- Assimilative phase free living amoeba which unite as psedoplasmodium before reproduction. 2. Hydromyxomycetes Assimilative phase is plasmodium (net plasmodium) 3. Myxomycetes Plasmodium saprobic, free living 4. Plasmodiophoromycetes- plasmodium parasitic within the host plant.

Thallus- Vegetative body is known as plasmodium. Plasmodium is characterized by presence of naked, multinucleate, motile mass of protoplasm. Most of the life cycle spends in the root cells of host when host dies, the roots disintegrate and resting spores are liberated into the soil. Resting Spores Minute, round uninucleate haploid enveloped in a chitinous smooth wall. They live in soil for more than 8 years without host under favourable condition in the presence of host germinate gives rise to uninucleate bean shaped biflagellate zoospores (swarm cell). In the absence of host, these spores perish. Before infection, they lose flagella and become amoeboid.

MYXOMYCETES- No Parasites Plasmodiophoromycetes They are endoparasitic slime moulds consists of holocarpic plasmodial thallus with plasmodial moment and feeding. They are obligate parasites. They cause excess enlargement of cell and excess multiplication of cell hypertrophy and hyperplasia. There are two types of plasmodium in the life cycle. 1. Sporangiogenous and 2. Cystogenous Sporangiogenous plasmodium form from sporangia aggregated in loose masses or sporangiosori consist of few or numerous small or large sporangia which produces single or many secondary zoospores. The zoospores are biflagellates with two anteriorly inserted whiplash flagella of unequal size (anisokont). These zoospores possibly unite

(but remain binucleate) before infecting the host to form cystogenous plasmodium which gives rise to thick walled cyst (resting spores). Then karyogamy and meiosis will take place. Hypertrophy of host cells is apparently brought out by blocking of cell division and is accompanied by enhanced DNA synthesis. The life cycle is initiated when cysts germinate, each giving rise to zoospores capable of infecting host cell or plant. The cyst zoospores attaches to the wall of root hair flagella become inactive and zoospores encyst. Within the encysted zoospores, a long tubular cavity termed the Rohr then appears. The end of the Rohr oriented towards the host wall is filled with a light-staining plug. Within the Rohr lies a sharp dark staining rod called Stachel. Following an evagination of the Rohr to form a bulbolous structure that adheres the host cell wall. The satchel punctures the host wall and the protoplasts of zoospores enter the (cell) root hair. According to Aist and William (1971), the protoplasts require only about one second to enter the (host) root hair. After penetration of a host by the cyst zoospores of small sporangiogenous plasmodia appears within the host cell. After the plasmodium reaches a certain apparently determined to a great extent by the size of the host cell; it cleaves into segments that develops into zoosporangia. It may occur singly or they may be loosely aggregated. Then the zoospores are released from zoosporangium (Plasmogamy, Karyogamy). There are gametes and undergo plasmogamy prior to penetration and infection. Due to infection, cytogenous plasmodia develop. Karyogamy appears to be delayed until shortly before cyst formation. Meiosis then takes place in the form of two successive divisions before formation of cyst. Cytogenous plasmodia gives rise to thick walled called cysts.

Plasmodiophoromycetes Phorales Phoraceae Myxomycetes slime moulds and endoparasitic

Mastigomycotina: (Alexopoulous) More like a true fungi. Spongospora subterranea Powdery scab of potatoes. It also transmit potato mop top virus. The viruses are carried by zoospores and persist for longer period in the spore balls in the soil. Mastigomycotina -----------------------------------------------------------------------------------------Uniflagellate Zoospore one flagellate

Flagella

Biflagellate -------------

----------------------------------------------------Posterior whiplash (pertshen)

Anterior tinsel (Flimmer)

Flagella equal posterior whiplash and anterior tinsel cell wall (cellulose) Oomycetes Chitridiomycetes True mycelium lacking. Rhizoids are called Rhizomycelium True mycelium present. Sexual reproduction

Plasmodiophoromycetes Zoospore unequal size.

Sexual reproduction by fusion of planogametes

Unknown. Zoospore released

Sexual reproduction by fusion of motile motile female gametes (Monoblepharidiales)

by circum sessile rupture (Harpochytridiales). (Blastocladiales)

Zoospores with annular cup male and non

Mastigomycotina The mastigomycotina are zoosporic fungi which includes chytridiomycetes, hypochytridiomycetes, plasmodiophoromycetes and Oomycetes. Sparrow, 1973,

Alexopoulos and Mims (1979) have included in myxomycetes. Classification of the Mastigomycotina into different classes is based mainly on the flagellation of the zoospores. Ainsworth (1973), Hawksworth et al. (1983) included

plasmodiophoromycetes under myxomycetes because of presence of plasmodium.

Chitridiomycetes: The motile cells of the fungi have posterior whiplash flagellum. Sexual reproduction- by motile isogamous gametes (Olpidium, Synchytrium). Synchytrium endobioticum-black wart of potato. Urophlyctis alfalfae crown wart. Physoderma maydis Brown spot of maize. Olpidium brassicae- parantheon root of cabbage, lettuce.

Somatic structure: Unicelluar, thallus without specialized vegetative part later which becomes converted into reproductive organ (Holocarpic). Chytridiomycetes: 1. 2. Chytridiales: True mycelium lacking. Rhizoids are Rhizomycelium Harpochytridiales: True mycelium present. Sexual reproduction unknown,

zoospores released by circum sessile rupture. 3. Blastocladiales: Sexual reproduction by fusion of planogametes. Zoospores

with annular cup. Reinstat Sporangia are formed. 4. Monoblepharidales: Sexual reproduction by fusion of motile male and non-

motile female gametes. No Reinstat Sporangia are formed.

Chytridiales Life cycle: This order contains 93 genera and 460 species. Generally called chytrids. Synchytrium endobioticum (potato wart), (Olpidium brassicae), Pleotrachelus virulentus (Lettuce big vein). The simple chytridiales are endobiotic, live anteriorly within the host. Epibiotic: Live outside (upon the cell) or the surface of the host, but nutrient absorbing structure may be immersed in the host tissue. Physoderma: Both epibiotic and endobiotic Monocentric: Rhizoidal system bears a single sporangium-resting spore. Polycentric: Extensive rhizoidal system and bear more sporangium. The asexual reproductive organs are sporangia spherical, pear shaped and inside zoospores are produced. Inoperculate: Discharge tubes are presentor exit papillae. The tip become gelatinous and dissolves through which zoospores are produced where as inoperculate, the discharge tubes are provided with well defined cup.

Chytridiales The classificationis still uncertain. Nine families. Sparrow (1960-1973). Chytridiales: Inoperculate: 1. Olpidiaceae 2. Achlyogetonaceae 3. Synchytriaceae 4. Phlyctidiaceae 5. Rhizidiaceae 6. Cladochytridiaceae 7. Physodermataceae Later Lange and Olson (1980) classified under Blastocladiales

Olpidiaceae: 1. Holocarpic 2. Endobiotic thallus which converted into single zoosporangium.

Sexual reproduction by fusion of isoplanogametes or resting sporangia.

Olpidium, Rozella Thallus- spherical, cylindrical Endobiotic-inside Holocarpic-entire thallus Zoosporangium- cleaves Zoospores Isoplanogametes Discharge from the tube and open to the exterior Zoospores Few minute tadpole Swim 20 minutes Root hair infection empty cyst coat outside Thalli a membrane Zoosporangium release Thick walled zoosporangium without discharge tube Sexual fusion, Zygote-fusion takes place outside Resting sporangium sexual Division-germination-zoospores The Olpidium thallus is spherical or cylindrical and there may be one or more in host cell. The entire thallus converts into zoosporangium and cleaves into uniflagellate zoospores. The zoospores escape from the discharge tube, which penetrate the outer wall of the host and open to the exterior. The release of zoospores takes place within few minutes of washing the roots free from the soil. The zoospores are like tadpole like with whiplash f;lagellum. After swarming for about 20 min, the zoospore encyst on the root hair and lose their flagellum. The root hair get infected and entire content will be transferred to the inside leaving only empty cyst. The thallus secretes a membrane around itself and grows into zoosporangium within the host cell. The nuclei divide repeatedly and within a few days, zoospores are formed and discharged from tubes in the sporangia and it repeats in asexual lifecycle.

Sexual phase: In addition to smooth zoosporangium with discharge tube, stellate bodies with thickly folded walls that lack of discharge tubes are also formed in the roots. There are resting

sporangia, which are believed to be formed after sexual fusion. Sexual reproduction of Zoospores takes place only between those spores that have been produced from different sporangia. The Zoospores copulate outside the host cell and produce biflagellate zygote which infects like zoospores, but develops into thick walled reinstant sporangium that discharge zoospores after several months. The resting sporangia are initially binucleate but karyogamy takes place during the division of the fusion nucleus before zoospore formation. Resting sporangia germinate by producing numerous zoospores (uninucleate). These escape and reinfect the host plant.

Synchytriaceae: Holocarpic, endobiotic thallus which is converted into a sorus(heap) of sporangia or prosorus surrounded by a common membrane or resting spore at the time of reproduction. The sporangia are inoperculate as in the family Olpidiaceae. Synchytrium is the largest genus with about 200 species, which are parasitic on flowering plants. S. lagenariae S. trichosanthidis

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S. sesamicola Sesamum S. endobioticum Black wart only in Darjeeling S. aureum 198 species of 123 genera and 34 families of flowering plants S. macroporum 1465 species in 918 genera of 176 families

S. endobioticum causes galls on the aerial shoots and tubers. During the infection on the aerial shoot, they become green with convoluted leaf like masses of tissue. The tuber infection induces the cells to divide and form into warts, which are dark brown warty excrexences. The size of the wart increases and in final rotting will be noticed and putrid mass excluding a dark brown liquid. It causes considerable yield loss. The pathogen starts with the release of the zoospores from the dark warts on the infected tubers, which contain resting sporangia. The resting sporangia are spherical cells with dark brown walls that have folded, plate like extensions and are released by the decay of warts. The germination of resting sporangia takes place by the formation of vesicles within which a single sporangium differentiates. Thus the resting sporangium

function as prosporangium on germination. After the release of the zoospores, move about 2 hrs and contact with tubers or any other parts (eye) and withdraw flagella and infect by means of infection pegs (outer empty cyst membrane end). The uninucleate contents passed onto the host cell. The infected cell induces galls and after sometime the cell dies. The thallus of the pathogen passes to the bottom of the host cell, enlarges, becomes spherical and fills the cell. As it matures, it becomes a prosorus or summer spores which has a thick orange exospore and a thin hyaline endospore. An extrusion, the endospore with a cytoplasm and the nucleus forms a vesicle; which fills upper part of the host cell. During its passage to the vesicle, the nucleus divides until 32 nuclei are formed. The content of the vesicle cleave into number of multinucleate blocks the sporangia each of which is finally surroundedby a thin wall and forms a sorus. Further, nuclear divisions and rounding off of the mass of cytoplasm around each nucleus produces large number of zoospores from each sporangium. At maturity, the sporangium absorbs water and bursting of host cell. The release of zoospores takes place in the sporangium through a slit. The zoospores of 500 to 600 in sporangium resembles like that of resting sporangium. After swimming for sometime, they infect healthy potato. The fusion of gametes takes place in pairs to form hyphae. After swimming for sometime, the zygote encysts on the surface of the host epidermis and penetrates the host cell. But before infection, the zygote penetrates the host, nuclear fusion occurs. As a result, hypertrophy and hyperplasia occurs whereas the zoospore does not cause the same. The resting sporangia are formed due to repeated cell division. The resting sporangia are shed into the soil by decay of warts. Before germination, the diploid nucleus divides repeatedly and initiates asexual reproduction. Occasionally the zoospores from the resting spores, sporangia are formed.

OOMYCETES It contains about 74 genera with 580 species. 1. Production of biflagellate zoospores with two types of flagella. The shorter flagellum of tinsel and two types of zoospores, one reniform and the other is pear shaped. 2. The cell walls are composed of glucan, cellulose and lack chitin.

3. An advanced oogamous reproduction takes place by the passage of gamete nuclei. The male gametes are not however free swimming is always conveyed from a well-defined antheridium by tube. Sometimes a mere pore in the wall of well defined oogoniumcontaining oospheres (egg) thus regulating the formation of oospore. Both gametes could be called endothallic. 1. Meiosis is gametangial rather than zygotic and the vegetative thallus is diploid (as in plants). 2. The life cycle is of the haplobiotic type.

Oomycetes includes unicellular, holocarpic, endobiotic parasites of other water molds and fungi. The peronosporales are the most advanced oomycetes- biotrophic parasites like downy mildews, white rust that completes life cycle entirely on the living plants (Albugo and Peronospora) directly. Zoospores germinate and give rise to germ tube hence they are called conidia. This indicates how conidia might have originated from sporangia. The name oomycetes originated from the oospore. But mainly zoospore flagellation has been accepted as the basis of the division of mastigomycotina and oospores have been given secondary importance.

Sparrow (1960-1973) Oomycetes: Spore Description: 1. Monomorphic / Dimorphic: Spores formed within the sporangium (One type or two type of zoospores) rarely aplanetic. a. Holocarpic or Eucarpic: Hyphae when present, lack of constrictions Eg. Saprolegniales b. Eucarpic: Hyphae with constrictions Eg. Leptomitales 1. Monomorphic / Reniform: Spores are formed within the sporangium or evanescent sporangial vesicle a. b. Holocarpic: eg. Lagenidiales Eucarpic : eg. Perenosporales

Saproleginales: They are water molds. Saprolegnia is a pathogen of fish. Aphanomyces euteiches causes root rot and sugarbeet of peas.

Saproleginaceae: Pyriform primary with two flagella, one whiplash and one tinsel attached anteriorly and reniform (secondary) with two flagella with whiplash and tinsel type. Somatic structure: profuse branched coenocytic mycelium, easily visible as it forms a colony around some bit of decaying plant or animal tissue.

Order:- Peronosporales: The peronosporales are the most specialized of the oomycetes are either saprophyte in water or soil or highly specialized parasites of angiosperms. The Peronosporaceae and Albuginaceae are biotrophic (obligate parasites).

Late blight of potato- Irish Famine Grapes downy mildew- Plasmopara viticola Downy mildews, Sclerospora, Peranosclerospora Sclerophthora are very important. Mycelium: Coenocytic, free branching, delicate septa may be formed to delimit the reproductive organs or in older mycelium. The colourless (hyaline) hyphae have walls composed of mainly glucan-cellulose complex and protein containing hydroxyproline. The biotropic species, the mycelium is intercellular and posses haustoria that vary in form from a minute spherical or cylindrical form.

Asexual reproduction: By zoospores, biflagellate with one whiplash and other is tinsel type. Peronospora and Peronosclerospora: Sporangium function as conidium. Sexual reproduction:

By oogamous and meiosis occurs in the gametangia (oogonium and antheridium) resulting in the formation of haploid oospheres and antheridial nuclei. The oospore wall is thick, smooth or ornamented. The mature oospore generally lies free within the oogonial wall and may fill the oogonium. Apleurotic Fill Pleurotic may not fill The oospores germinate either by producing zoospores or germtubes. Classification: Peronosporales: a. Sporangiophores are usually undifferentiated from the mycelium branched, indeterminate growth Eg. Pythiaceae b. 1. Sporangiophores are easily differentiated from the mycelium. Sporangia are in chain (Basipetal) produced by a short club shaped, unbranched sporangiophore. Eg. Albuginaceae 2. Sporangia or conidia are produced in single at the tip of sporangiophore of determinate growth. Eg. Peronosporaceae

PYTHIACEAE The member of the Pythiaceae is either saprophytes or facultative parasites and grew readily on artificial medium. The sporangiophores cannot be differentiated from the vegetative hyphae and resume growth after they produce sporangium either below or within the previous empty sporangium. In Pythium however the sporangiophore has determinate growth. The sporangia may remain attached to the sporangiophore or may be caduceus and liberate zoospores or it may germinate and produce germ tube and all depends upon environment. Pythium ultimum var. ultimum germinate directly and these sporangia are called conidia.

Sexual Reproduction: By well developed oogonia and antheridia, which are produced by single zoospores (Homothallic). Pythium sylvaticum, P. heterothallicum, P. intermedium, P. catenulatum and P. sphendens are found to be heterothallic. The oogonia originate as terminal or intercalary spherical swellings of the hyphae, which gradually increase in size until they become spherical. The oogonium is then cut off from the adjacent mycelium by the formation of cross wall. The antheridia arise as club shaped swollen hyphal tips often as branches of the oogonial stalk (monoclinous or from another hyphae, diclinous). The antheridia are smaller than oogonium and number of antheridia come into contact varies from one as in P. ultimum to several as in P. debaryanum. The young oogonium is multinucleate. Though young antheridia are also multinucleate, all nuclei degenerate except one. Fertilization takes place through the fertilization tube and male gametes enter into the oogonium. But before fertilization, meiosis occurs in the oogonium. The nucleus in periplasm also dies. The matured antheridium contains only one nucleus and fertilization takes place. This undergoes meiotic division so that the older antheridium has four nuclei. Although the number of antheridia attaches to the oogonium varies from one to four. Fusion takes place between single nucleus of antheridia and egg nucleus resulting in zygote. The fertilized oosphere secretes a double wall and develops into smooth thick walled spore. The oospore germinates after a resting period of several weeks.

PHYTOPHTHORA Anton de Bary first coined the name Phytophthora (Gr.Phyton- plant, Pathoradestruction) the plant destroyer in1876 when he described late blight of potato. South America- Europe 16th century - late blight appeared in Europe USA during 1830,1840. In 1845-46, epidemics outbreak complete failure and caused Irish Famine. It is estimated that hundreds of thousand people died of starvation and resulted in reduction of population of Ireland by more than 1/4th. Carefoot and Sprott (1967) in their book entitled Famine on the wind blame the late blight epidemic of potato for the defeat of Germans by the British in World War I. During the growing seasons in 1916, the weather in Germany similar to that of Ireland in

1846. The Bordeaux mixture is only the recommendation for the management. But the military leaders could not allow the use of copper, which is needed to prepare it. Copper was diverted to shell castings electric wire and for brass buttons. The Nation went hungry in 1916-17. Many of the soldiers family starved to death and this resulted in lowering the morale of the soldiers and collapse of military might in 1918. The major role of P. infestans in this defeat could be summarized by the old saying for want of a nail, the shoe was lost and for want of a shoe, the horse was lost and for want of a horse, the battle was lost. It has been reported that consumption of blighted potatoes by pregnant women may result in serious birth defects. The pathogen believed to produce a powerful mycotoxin in infected tubers (Alexopoulos and Mims, 1979). In the most recent and useful publication brought out by American Phytopathological Society, 1983 (Erwin et al) on International Symposium on Phytophthora. Phytophthora, the hyphae is wider, lemoniform, obpyriform or ovoid sporangium. The zoospores are fully flagellated. The smooth, rough or warted oogonila wall, which may be thicker and brown. The antheridium are amphigynous(very few paragynous). Haustoria are always present. It also produces oospores and chlamydospores. The mycelium is non-septate which branches at right angle and have constrictions at the point of origin. Sporangia and Sporangiophores are indeterminate. Zoospores direct and indirect.

Sexual reproduction: P. infestans is a heterothallic fungus and requires two matting types for sexual reproduction. Clinton, 1911 first discovered the sexual stage of this fungus in pure culture. At present, two matting types were discovered A1 and A2, which were (Gaundo and Gallegly, 1960). It is only when isolates of the A1 type are paired with isolates of the A2 type that the oospores are formed. Brasier (1971-75) showed that volatile substances produced by Trichoderma species can induce strain A2 to fertilize themselves. Several workers have studied the fine structure of sporangia in Phytophthora spp. Cleavage vesicles and flagella develop prior to induction of zoosporogenesis in the

sporangia of P.infestans. Mature sporangia are ready for indirect germination once the required, temperature and moisture are obtained. Zoospore initially arises as a result of the cleavage of the cytoplasmic contents. Cleavage is brought about by cleavage vesicles, which arise from the Golgi apparatus. In terrestrial forms, the sporangia are caduceus and the detachment of the sporangium is facilitated by hygroscopic twisting of the dry sporangia, which are dispersed by the wind before germination. In aquatic forms, zoospore release commonly occurs while the sporangia are still attached. Production and germination of sporangia is largely governed by temperature, moisture and relative humidity. Sporangia are formed at a minimum relative humidity of 91 per cent and optimally at 100 per cent and a temperature range of 3-260C. For abundant sporangia formation, the optimum temperature is 210C. Sporangia rapidly lose their viability at temperatures above 220C in moist air and less rapidly in dry air. In the presence of water, the germination of sporangia may occur either directly by a germtube or indirectly by means of zoospores. Higher temperature induces sporangial germination by a germtube, which grows directly through the sporangial wall or through the sporangial papilla. In P. infestans for example, multinucleate germtubes arise above 200C, whilst below 150C, uninucleate zoospores are produced. As the aged sporangia lose their capacity to produce zoospores and germinate by means of a germtube. Since cleavage vesicles and flagella are already present. Sexual reproduxction is by antheridia and oogonium of the opposite matting type. The oogonia penetrate and grow through the antheridium and develop into a globose structure above the antheridium. The mature antheridium thus forms a funnel shaped collar around the base of the oogonium. P. cactorum paragynous- Erwin (1972) Both antheridium and oogonia are multinucleate. At the time of fertilization during oosphere maturation, only a single nucleus remains in the center. Oospore has a dormant period of several weeks and germinates. But before germination, nuclear fusion takes place and the fusion nucleus divides several times.

Sclerophthora In Sclerophthora, the unbranched or sympodial sporangiophores with their citriform or obpyriform sporangia are like Phytophthora. The sex organs would fit either genus Phytophthora and Sclerospora but mature oospore has a thick epispore which fuses with a oogonial wall to form a thick brown covering called pleurotic which is characteristic of Sclerospora (S.P. Ray Chaudri, 1971 and Thirumalachar, 1969). Sclerophthora macrospora is parasitic on 46 species belongs to Graminae and Cyperaceae. Sclerophthora rayssiae var.zea on maize.

Peronosporaceae Highly specialized family of the order peronosporales. Most of them are biotropic ie. Obligate parasites. 1. 2. 3. 4. 5. Peronospora parasitica Crucifers (Guttenberg and Schmoller, 1958) P. farinose Ing ram and Joachim (1971). Sclerospora graminicola (Tiwari and Arya, 1969) Plasmopara viticola (Morel, 1944; Ingram, 1975) Pseudoperonospora humuli (Hope) were grown in tissue culture.

Mycelium is intercellular and made up of colourless, coenocytic hyphae and provided with various types of haustoria. From intercellular mycelium arise well differentiated branched sporangiophores are emerged through stomata. The

sporangiophores reach maturity, stop growing and then produce a group of sporangia on sterigmata. The sporangia are colourless either round or oval or lemon shaped. They are always deciduous and disseminated by wind. On reaching suitable host, the sporangium germinates either by germtube (Peronospora, Bremia germtube. Zoospores Pseudoperonospora, Basidiospora and Bremiella). Zoospores and germtubes Sclerospora Zoospore or plasma (sporangia emitting the entire content in a mass and later zoospores are fashioned externally

Plasmopara: Sexual reproduction: Same as that of Pythiaceae, oogonia are larger, antheridium is broader compared to Pythiaceae. The oogonial wall is thin and smooth. The oospores are smaller than oogonium. Sclerospora Pleurotic Oospore germtube Oospore Zoospores Peronospora tabacina and Plasmopara viticola. Sclerospora graminicola a. white basal medium b. casein hydrolysate c. 2,4-D. d. kinetin

Plasmopara viticola One of the most notorious and important downy mildews. Discovery of Bordeaux mixture in Bordeaux University by Alexis Millardet Botany Professor in 1885. Symptom: Destruction of leaves, dwarfing and killing of the shoots, rotting and cracking of berries. It is a biotropic parasite intercellular and with knob like haustoria. Sporangiophortes emerge through through stomata and truncate branches at right angle to the axis. The sporangia are hyaline and lemon shaped. Germination either by emission of plasma from which zoospores is formed externally or internally or by escape of entire sporangial contents which later produce germtubes. Oospore Oogonium and antheridium are paragynous Oospore by germination gives germtube terminate to sporangium.

Sclerospora graminicola Sclerospora graminicola is the causative agent of an important downy mildew and green ear disease of pearl millet (bajra) and has also been reported on other millets from India and several other countries (Bhat, 1973). The losses in yield ranges from six per cent to 30 per cent. Mathur and Dalela (1971)

estimated the losses, in monetary terms to be worth Rs. 20 million (approx. US$ 2.5 million) every year in the state of Rajasthan alone. During Kharif of 1975, a serious outbreak of the disease in hybrid bajra was reported in Karnataka and Maharashtra covering several thousand hectares and resulting in almost 100 per cent loss (Govindu, 1982). The inflorescence is mainly affected and the solid spicate ear is converted in part or wholly into a loose head composed of small, twisted, leaf like structures which give it a green bearded appearance, justifying its name; green ear disease. Bristles, glumes, stamens and pistils become leafy and green. But anatomically the green ear leaves are very different from normal leaves. In addition to differences in orientation, in green ear leaves the vascular bundles are devoid of sclerified bundle sheath and marginal fibres. The linear arrangement of epidermal cells and the stomata is missing (Patel, 1978). The infected plants show excessive tillering and remain dwarf because the internodes shorten.

Albuginaceae: This family has only one genus and 30 species. The sporangia are produced in chains and they are basipetal.

Zygomycotina
Reproduce asexually by means of aplanospores. The spores are produced in sporangia, which may be dispersed actively, but wind, rain or animals more frequently disperse them passively. Reproduce through sexually by means of copulation of two equal gametangia resulting in the formation of a thick walled Zygosporangium that contain Zygospore. The mycelium is coenocytic and the cell wall contains chitin. This class contains 145 genera and 765 species that may be saprobes. Facultative or weak parasites of plants, specialized parasites of arthropods and obligate parasites.

Zygomycotina: Zygomycetes: Saprobes or if parasites or predaceous having mycelium immersed in host tissue. Trichomycetes: Associated with arthropods and attached to the cuticle or digestive tract by a holdfast and not immersed in the host tissue. Zygomycetes: Coenocytic mycelium. Absence of motile cells and sexual reproduction by gametangial copulation and results in the formation of zygospore. Asexual reproduction is by nonmotile sporangiospore. Saprobes or weak parasites. Human pathogens or Mycorrhizae.

Mucorales: Saprobes, weak parasites on plants or endoparasite or vertebrates. Asexual reproduction is by non motile sporangiospore produced on sporangiophores borne in sporangia, mesosporangia or sporangiola or in one sporangia or conidia. Entomophthorales: Chiefly parasitic on invertebrates, asexual reproduction is by modified sporangia functioning as conidia or by true conidia. Conidia are borne singly or in chain and passively discharged.

Zoophagales: Modified sporangia functioning as conidium or and forcibly discharged.

Mucorales: Spoilage of food, mostly saprobes, weak parasites on plants and animals. Some of them are obligate parasites on mushrooms. They are also called pin moulds because of their sporangia that appear black dots in cobweb like hyphae. Rhizopus and Mucor causes soft rot of sweet potatoes, strawberries, raspberries, peach and other fruits.

Breadmould Absidia, Mucor, Rhizopus, Cunninghamella causes mucoromycosis in man and animals. The genus, Saksenaea is a virulent human pathogen produces lactic acid. 1. 2. 3. 4. R. sinensis Lactic acid R. stolonifer Lactic acid R. oryzae Fumaric acid and alcohol R. nodosus - Fumaric acid and alcohol

Under appropriate fermentation condition, numbers of mucorales are capable of rearranging steroid molecules. 1. 4. Epoxidation Actinomucor elegans are utilized for making Chinese cheese from soybean. Hydrogenation 2. Dehydrogenation 3. Hydroxylation

Mucorales: Martin (1961) 9 families Ainsworth (1971) - 11 Hawksworth et al. (1983) - 11 Alexopoulos and Mims (1979) - 11 Hesseltine and Ellis (1973) 14

Mucoraceae: 21 genera with 119 species. Mucor do not produce rhizoids and stolon. Acaulospora: Azygospores bud laterally form funnel shaped stalk of large inflated hyphal terminus. By spore maturity, the funnel stalked cell collapses. Gigaspora: Azygospores bud from the bulbous, suspensor like tip of hyphae, As spores mature, 1-sexual septa form below the suspensor like bulb, produce oneseveral narrow out going hyphae. No fusion of hyphae with spore. Glomus: Chlamydospores form at hyphal tip, usually one per tip. By maturity, spore contents are separated from attached hyphae by a septum.Two or more hyphae attached to the spores. Eg. Glomus mosseae. Sclerocystis: Chlamydospores form is sporocarp as a single crowded layer of excess spores that surrounds the sides and top of the spore free central mass of tightly interoven hypha. Endogyne sexual reproduction R. sexualis Homothallus

Endogonaceae For many years, Endogonaceae was considered to be ascomycetous but Bcuholtz (1912) described the formation of zygospore.Some members of Endogonaceae form spores that are found free in the soil and some produce sporocarps (fruiting body). The sporocarp varies

from 1mm in diameter and contains zygospores or chlamydospores. Many of the members mainly hypogeous forming endomycorrhizae. Mossae (1973) and Sanders and coworkers

(1976) 400 million years ago, the earliest known vascular plants. The Rhyme fossil of Devonian age has mycorrhizal root infections similar to modern infections caused by

Endogonaceae. Net sieving and decanting of soil samples. VAM (Vesicular Arbuscular Mycorrhizae) helps in the uptake of phosphorus and other nutrients.

Endomophthorales Very small group of fungi, parasitic on insects and parasitic on fern prothalli. Basidiobolus haplosporus Conidiobolus coronatus - causes Zygomycosis in humans. Asexual Fragmentation of hyphae ie. Hyphal bodies contain chitin and are forcibly discharged from sporangiophores. Sexual Gametangial copulation takes place by fusion of equal or unequal gametangia and forms zygospores. Zygospores also formed without the fusion. Some will not reproduce. Chlamydospores are thick walled resting spores. desmanthus or

Entomophthoraceae: Endomophthora with 82 species. Mycelium breaks up into multinucleate hyphal bodies, each one is capable of forming sporogenous cells (also called conidiophores) which is like a basidium and bears one or more conidium which are discharged forcibly. Entomophthora muscae Fly fungus (viable for 3-5 days). If then conidia gets the host or infect by producing appressorium. Otherwise it germinates and gives rise to another (secondary conidium). Like that tertiary conidia and germinate till a suitable substrates is reached or protoplasm is exhausted. Multinucleate spherical resting bodies are formed partheno- genetically within the body of the host. There are azygosporangia. So E. muscae do not reproduce sexually. Their azygosporangia act as resting spore. It germinates only after 2 to 3 years and germinates by inducement of chitin decomposing bacteria. E. sepulchralis reproduce through copulation of hyphal bodies and forms zygospore.

. HETEROTHALLISM AND PARASEXUAL CYCLE A.F.Blakeslee, an American Geneticist in 1904 made an important observation with Mucor, which resuited in the discovery of Heterthallism. Blakeslee observed that while some isolates of Mucor formed sporangia as well as zygospores (e.g. M.tenuis), some others failed to form the zygospores and reproduced only by sporangiospores. When he grew these non-sexually - reproducing isolate with other similar isolates, zygospres appeared in the region where the hyphae of the different isolates came in contact with each other. Blakeslee coined the terms homothallism and heterothallism to explain this phenomenon. The homothallic species were those that produced zygospores independently while heterothallic species required the presence of the opposite mating type. M.hiemalis,M.mucedo, Rhizopus nigricans are examples are examples of Heterothallic species. Since the two mating types were morphologically indistinguishable, Blackslee designated them as ther (+) and (-) mating types or strains (not male or female)Blackslee assumed that the (+) strain represented the female and the (-) strain

represented the male sex. We now don't speak of (+) and (-) sexes but only as (+) and (-) strains or mating types.

Pattern of Distribution of Sex Organ in Fungi On the basis of the distribution of sex organs fungi can be put in the following categories A. Hermaphoridite in which both male and female sex organs occur on the same thallus. B. Diccious (sexually dimorphic). The two sex organs are present on different thalli C. Sexually differentiated The male and female sex organs are morphologically similar and therefore indistinguishable. A hermophroditic fungus having both the sex organs may be homothallic or heterothallic. When the two sex organs, present on the same mycelium are unable to mate this is because of self-sterility and is called physiological heterthallism (Whitehouse, 1949). Such fungi need genetically different nuclei, which does not occur when the same thallus forms both the sex organs.

The dioecious fungi in which the male and the female sex organs are borne on different thalli are by necessity heterothallic. This is called Morphological heterothallism (Whitehouse, 1949).

In this case heterothallism is made obligatory because the opposite and morphologically distinct sex organs are formed only on different thalli.

The sexually undifferentiated fungi e.g. Mucor, Rhizopus and several members of Asco-and Basidiomycotina, do not have morphologically distinguishable sex organs. These can also be homo-or heterothallic. The heterothallic forms provide another example of physiological heterothalism. The requirement for the other thallus does not lie in morphologically distinct sex organs but in genetically different nuclei, which are not available in the same mycelium.

So, heterothallism according to Whitehouse (1949) can be caused by the absence of the morphological sex organs of opposite type (morphological heterothallism) or by the absence of genetically different nuclei (Physiological heterothallism).

Whatever be the reason of heterothallism the fact remains that different thalli are needed for the sexual reproduction. A heterothallic species may not be only two mating types. There can be four types of thalli and one thallus can mate only with only one of the rest thre. This is called tetrapolar heterothallism. Bipolar Heterothallism Fungi in this category have two mating types each containing genetically different nuclei. The sexual compatibility is controlled by a pair of genetic factors A and a located at the same locus on different chromosomes. This is therefore also called as ' two allele heterothallism'During meiosis the two chromosomes containing the alleles A and a are separated in the haploid spores (germ spores, ascospores or basidiospores). The spores give rise to two types of thalli which must come together the two nuclei carrying the compatibility factors A and a The two mating type are designated as (+) and (-) strain The 'two allele' or 'bipolar' heterothallism is found in

Mucorales (Mucor, Rhizopus, Phycomyces). Ascomycotina (Neurospora, Ascobolous). Basidiomycotina (Puccinia graminis and the smut fungus Ustilago levis). Tetrapolar Heterothallism Fungi in this group form thalli of four mating types. This type of heterothallism is goverened by two pairs of caompatibility factors Aa and Bh, located at different chromosomes, which segregate independently during meiosis. If crossing over occurs between the mating type loci,

four types of segregations (4B,Ab,aB,ab) are possible depending on the chromosomal arrangement.

Thus four types of spores (AB, Ab, aB and ab)are formed which give rise to four types of thalli. Only those thalli that have nuclei carrying oposite genes for both the factors, can mate. The resulting zygote must have the genotype Aa, Bb. Majority (630) of the heterothallic Basidiomycotina are tetrapolar forming four types of basidiospores. However if crossing over does not take place only two types of spores (AB and ab or Ab and aB) are formed and only two types of thalli are produced. since it is goverened by two factors it is called tetrapolar. Hormonal or physiological basis for heterothallism In case of zygomycetes, Burgeff at the year of 1924 reported that diffusable substances are responsible for initiation of sexual reproduction. + And - strains are separated by the collodion membrane makes it still attracks by opposite strains.

Plempel et al., 1957 reported that there are some reactions is being taken place while the two compatible strains approaches towards each other. The reactions are as follows,

I. Teleomorphic reactions: In this reaction aerial zygophores are formed which is in club shaped and yellow in colour. (carotene is responsible for the yellow color) II. Zygotrophic reaction Zygophores of + and - strains are grow towards each other. III. Thigomototrophic reaction

In this reaction events are taking place after gametangial fusion and septation.

Gooday (1968) isolated the substances responsible for the sexual reproduction at the point of contact of + and - strains. The isolated substances are generally called as Trisporic acid A, Tripsoric acid B, Tripsoric acid C. In + strains this acid is chemically termed as Methyl dihydrotrisporates whereas in - strains it termed as Triporols.

Heterothalism in Oomycetes In case of Oomycetes if two thalli were contacted together, then it was found to produce four kinds of different thalli. They are as follows,

I. Pure male thalli II. Predominantly male thalli and latent stages it produces OOgonia III. IV. Pure female thalli Predominantly female thalli and latent stages it produces antheridia.

Heterothallism in phytopthora sp Phytophthora infestans :Galindo and Galligly reported divided into A1 and A2 compatibility type . They paired and produce abundant oospores in lima bean medium.

In one combination act predominantly as a male when paired with stronger male then act as the female. Gametangial fusion occur between A1 and A2 type.

(ii) Phytothora arecae pethyb reported * * N-90 and N-91 isolates when paired does not produce sexual spores. When both isolates paired with A1 type of P.infestance; they produce abundance

oospores but not with A2 type. hence which is designated as A2 type.

(iii) Phytopthora cambivora * * N-98 and N99 paired with 7 isolates of P .sp An abundance of oospores formed when paired with A1 type of P. cinnamomi

(iv) Phytopthora capsici : 5 isolates N14 N58,N59,N313,N100.

* N14,N59 and N313,were designated as type A1 because which forms oospores with 60A isolates of P.infestans. * N58, and N100 were A2 type because paired with 60B type.

Phytopthora cinnamomi * Isolates -N2,N33,N66,N67 and N310, N101 to N106, N160, N202, N210, N311, N312. * Hasis and Nelson observed abundance of Oospores in hempseed medium.

(vi) Phytophthora palmivora: 32 isolates were received parings of the first 18 isolates with A1and A2 types of P.infestans on half strength lima bean Agar medium revealed the presence of A1and A2 compatibility types in P.palmivora

Based on oospore production with N241 isolates : * A1 isolates were N136, N137, N146, N240, N242, N243, N248 and N251. * A2 isolates oospores production with N137: N7, N8, N119- N123, N125, N130, N138, N136 141, N143, N144, N244, N245, N247. Phytopthora parasitica var Nicotianae:(Tucker) : 18 isolates were received.

Numerous oospores are formed when isolates N235, N236 and N237 were paired interspecifically with type A1 of P.parasitica,where as small number were produced when N238 was paired with A2 of P.parastitica A1 types : N4, N6, N15- N18 and N238 A2 types : N19- N26, N235, N237.

Heterothallism in Ascomycetes Many of the fungi coming under this sub-division are homothallic and some others are heterothallic. Eg for homothallism. Sordaria macrospora Sordaria fimicola

Eg for heterothallism: Neurospora crassa Neurospora sitophila In the process of ascosporogensis, the ascus produced 8 ascospore in which two mating types A, a were found to produced. Eg for Secondary Homothallism in ascomycestes: Neurospora tetrasperma Podospora anserine Gelasinospora tetrasperma Heterothallism in basidiomycotina: 10% of the fungus coming under this sub-division are homothallic. Eg) Coprinus sterquilinus Sexual compatibility is being controlled by pair of genetic factors then it can be termed as tetra polar heterothallism. Eg) Majority of basidiomycotina fungus are tetrapolar.

PARASEXUALITY Until 1944 the sexual cycle was the only means of exchange of genetic material it is to the credit of microbial geneticists that aseries of novel methods of genetic recombination are now known inbacteria which do not involve karyogamy and meiosis. these aretransformation conjugation transduction, Iysogeny and sexduction which differ from the standard sexual cycle.

A similar alternative to sexual reproduction was discoveredin the imperfect fungus Aspergillus nidulants in 1952 by Pontecorvo and Roper of Glasgow. This they called Parasexual cycle. In this genetic recombination occurs in vegetative cells by the mechanism of mitotic crossing over which brings the same result as is achieved by the meiotic crossing over.

The parasexual cycle involves the following steps 1. Formation of heterokaryotic mycelium

2. Nuclear fusions and multiplication of the diploid nuclei. 3. Mitotic crossing over during divisions of the diploid cells. 4. Sorting out of the diploid strains. 5. Haploidization

Formation of heterokaryotic Mycelium -Heterokaryosis Heterokaryopsis is the main source of variation in the imperfect fungi (deuteromycotina) which lack of sexual reproduction. The term Heterokaryosis was proposed by Hansen and Smith in 1932, who reported it for the first time in Botrytis cineraca.

The presence of genetically different nuclei in an individual is called heterokaryosis and the organism heterokaryon. Essentiallly a heterkaryon possess two sets of chromosomes just like a diploid organism but instead of being contained chromosomes just like a diploid organism but instead of being contained in a single nucleus the two sets of chromosomes lie in separate nuclei sharing the same cytoplasm.heterokaryons show dominance and thus resemble diploids in many respects. Heterokayons show dominance and thus resemble diploids in many respects. Heterokaryosis is a major factor in natural variability and sexuality. The heterokayotic condition can arise in a fungus by three methods viz., (i) Mutation , (ii) Anastomosis ie., fusion between genetically different hyphae and (iii) Diplodization - fusion between haploid nuclei to form diploid nuclei.

Mutations occur frequently in fungi and a homokaryotic mycelium is frequently covereted into heteokaryotic one. Anastomosis between spores and hypahae is a universal feature of higher fungi and certainly must be a potential source of heterokaryosis and thus of variability. Whether nuclei migrate from one thallus to another is a debated point but the hyphae having nuclei --- parents arise at the point of fusion. Heterokayosis is often accompanied by parasexual cycle. Nuclear Fusions and Multiplication of the Diploid Nuclei Nuclear fusion invegetaitive heterokaryotic hyphae was first noted by Roper (1952) in Aspergillus nidulans. Nuclear fusion may occur between genetically similar and

dissimilar nuclei, resulting in the formation of homozygous and heterozygous diploid nuclei , respectively. Diploid heterzygous nuclei are formed very rarely (at a are present ; 2 types of haploid nuclei their two types of homozygous diploids and the one type of heterozygous diploids.

Mitotic Crossing Over Crossing over is a phenomenon which occurs during meiosis and gives rise to new linkage of genes gene recombination. However initotic crossing over was discovered in 1936 by Stern in Drosophila. a similar mitotic crossing over occurs during the multiplication of the diploid heterozygous nuclei though at a low frequency of 10-2 per nuclear division. However, in some other fungi e.g., Penicillium chrysogenum and Aspergillus niger the frequency of mitotic crossing over is as high as during meiosis in sexual reproduction ; both lack sexual reproduction. Mitotic crossing over is the most important , or key event in parasexual cycle, as it is during this step that genetic recombination occurs.

Sorting out of Diploid strains the segregation of the diploid strains occurs when unimucleate diploid comdia are formed. The colonies that are formed by diploid conidia are recognized by various methods e.g. higher DNA content and bigger size of the conidia and certain phenotype characters of the colony.

Haplodization The diploid colonies show appearance of sectors on the Petriplate, which produce haploid conidia. Thisindicates that some diploid nuclei must have undergone haplodization forming haploid nuclei which later get sorted out in haploid conidi. some of these haploids are genetically different from the original haploid parental nuclei. This is because of the recombination that occurred during the mitotic crossing over. Haploidation occurs at a constant frequency of 10-3 per nuclear division. The haplodization occurs not by a reduction division (meiosis) but by aneuploidy a

phenomenon in which chromosomes are lost during mitotic divisions. it happens like this. During mitosisof the diploid nucleus, the chromatids fail to separate (non-disjunction) in the anaphase stage.One daughter nucleus gets one chromosome more (2n+1), while the other gets one chromosome less(2n-1) than the normal 2 sets of chromosomes (2n). Both the daughter nuclei are called aneeuploid. the deficient aneuploid nucleus (2n-1) may lose more chromosomes in the successive mitotic divisions and finally reduced to haploid state (n). Mitotic crossing over and haploidization occur also with the diploid hymozygous nuclei but since the two nuclei are similar, crossing - over products or the haploid nuclei formed by haploidation are genetically no different from the haploid parent nuclei. The parasexual cycle thus like the sexual cycle involvesplasmogamy, karyogamy and haplodization but not at a specified time or place. Every step differs drastically. A comparison betweeen the sexual and the parasexual cycle

Sexual cycle Nuclear structures. fusion occurs in

Parasexual cycle specialized Nuclear fusion occurs in vegetative cells

Zygote usually persists for one generation Zygote persists for many generations by only. mitotic divisions.

Recombination occurs by crossing over Recombination by mitotic crossing over during meiosis. The cropping over occurs which is rare event and occurs in a single in all chromosomes by a parts and in is chromosome arm. Haplodizations unlike the meiosis is independent of crossing over.

accompanied

reduction

chromosome number. Products of meiosis readily recognizable Recombinant nuclear lie in the vegetative which can be isolatedeasily. cells and can be recognized only by suitable genetic markers. References Text book of fungi, Bacteria and Viruses by - Dubae A Text book of plant pathology by D.P.Tripathi, H.P. Shukla Phytopathology - Volume - 58 (1968) page 1004-1020.

Ascomycotina
Largest group of fungi contains 2720 genera and 28,650 species. (Hawksworth et al. 1983) which include yeasts, black mould, blue mould, ringworms, powdery mildews, cup fungi, morels and truffles. Destructive diseases like peach leaf cur-Taphrina deformans, ergot - Claviceps purpurea and take all - Gaumannomyces graminis. LSD-Lysergic acid, diethyl amine hallucinogenic drugs from ergot contains alkaloids. DEUTEROMYCETES This is the second largest subdivision of the fungi and contains 1680 genera, 17000 species (Hawksworth et al., 1983). Fungi reproducing exclusively by asexual means usually by conidia are placed in subdivision - Deutecomycotina. Mycelium - Well developed, branched, septate, multinucleate hyphae. Hyphae - well developed, branched, septate The sexual stages belongs to Ascomycotina & Sometimes Basidiomycotina. (e.g) Imperfect (Anamorph), Perfect (Teliomorph) Ascomycotina -Fusarium moniliforme - Gibberella fujikuroi Basidiomycotina - Rhizoctonia solani - Thanatephorus cucumeris

Deutecomycetes: (imperfect / asexual fungi) Mycelium well developed, septate branched, sexual reproduction rare (or) unknown, Asexual spores (Conidia). From on conidiophores existing singly or grouped in specialized structures. Eg: Sporodochia, synnemata or produced in pycnidia and acervuli.

Asexual fruiting bodies 1. Sporodochium : Is is cushion like aggregation of hypha which breaks through host and bears conidia. eg. Fusarium, Epicoccum .

2. Acervulus : Formed by the aggregation of somatic hyphae below the epidermis of the host forming cushion like structure on which conidiophores and conidia formed. Eg:

Colletotrichum capsici 3. Pycnidia : When spore and sporophores are formed inside a specially designed fruit bodies which have protective wall made up of fungal tissue, they are termed as pycnidia. eg: Macrophomina phaseolina.

4. Synnemata : Loose aggregation of errect conidiophore similar to mycelial strands eg: Arthrobotryum.

Deuteromycotina

Blatomycetes Coelomycetes * Budding yeast / yeastlike

Hyphomycetes

* Mycelium well developed

* Mycelium well developed

* with/without pseudomyclium Budding cells

* Budding cells absent

absent * True mycelium lacking/not well developed * Mycelium sterile / bears * Spores in pycnidium Or acervuli

spores directly/on sporophores. * Pycnidia/acervuli absent

Hyphomycetes Order

Moniliales (1) Moniliaceae Verticillium, Aganomycetaceae Aspergillus, Penicilium, Sarocladium.

Agonomycetels

(2) Dematiaceae Helminthosporium,

Pyricularia, Culvularia, Alternaria, Cercospora.

(3) Tuberculariaceae Fusarium . (4) Stillbellaceae synnemata,

Coelomycetes Order

Melanconiales Frucitification acervuli

Sphaeropsidales Fructification pycnidia (or)stomata

Melanconiaceae Colletotrichum Sphaeropsidaceae Phoma, Phomopsis Botryodiploida,

Coelomycetes: There are 650 genera, 8000 species in this class (Hawksworth et al., 1983). Grove (1919) introduced the term coelomycetes to include those imperfect fungi which are formed inside a cavity formed in the substrate on which fungus grows. Melan coniales: General character: It is characterised by the production of acelvuli which may develop sub epidermally / sub cuticulary. Aurvulus - It is a saucer - shaped fructification in which the conidiogenous cells develops on the floor of the cavity from a pseudoparenchymatous stroma beneath an integument of host tissues which rupture at maturity (Hawksworth et al., 1983).

The conidia from acervuli are released in the form of characteristic droplets which may be variously colored from hyaline, to cream, pink orange or black.

eg:

Over 120 genera are included in this family, Antheacnose, Colletotrichum, Pestolotia, Pestalotiopsis, monochactia,

melanconium.

Sphaeropsidales : General character : The conidia and conidiogenous cells (or) conidiophores are produced in pynidia. Mycelium may be immersed in the substrate / superficial. Pycnidia are flask shaped to globose fructification with slightly tapered apex and single ostiole. Inside the pynidium hymenium is present which bears conidia. They may be produced superficially, subimmersed (or) immersed in the substratum. The wall of the pynidium is made up of pseudoparenchymatous tissue and the conidiophores arise from cell of the inner wall.

Hypomycetes: moniliales(order): Tuberculariaceae: This order contains many genera. It includes Fusarium, Tubercularia, Volutella, Epicoccum, Exosporium.Special feature of this order is the production of sporodochia in which the spore mass is supported by a superficial, cushion like (Pulvinate) mass of conidiogenous celles (or) short conidiophores.

Aganomycetales : The fungi included in this order are often referred to as mycelia sterilia, as they lack even imperfect stage. ie., in which no spores of any kind are produced and reproduce only by fragmentation of the mycelium.

The do form sclerotia / chlamydospores which perpetuate and disseminate the pathogen. 42 genera included in this order (kendrick and Carmichavel, 1973) including Aegerita, Arbuscula, Burgoa, Papulaspora, Dactuliosphora, Rhizoctonia and Sclerotium.

Genus - Colletotrichum (Deuteromycetes, coelmycetes, Melanconiales, Melanconiaceae, Amero-, Hyalo-, Phialospores). This genus is a facultative parasite causing anthrocnose diseases on leaves, young twigs and fruits of many plant species. Important species: Colletotrichum capsici- Dieback and ripe fruit rot of chillies, other hosts chickpea, cotton, brinjal, tomato, turmeric. Colletotrichum coccodes Hosts belong to solanaceae, leguminosae, and cucurbitaceae. The species causes black dot of potato, and tomato and anthracnose of fruits of tomato and chilli.

Colletotrichum truncatum Attacks groundnuts, pigeonpea, soybean, pea, cowpea, Phaseolus spp. Causing anthranose and leaf dpots. Colletotrichum lini - Anthracnose of linseed. Colletotrichum falcatum (Glomerella tucumanensis)- Red rot of sugarcane. Colletotrichum lindemuthianum(Glomerella lindemuthianum)Anthracnose of beans.

Genera - Helminthosporium, Drechslera and Biopolaris: These three genera belong to series porosporae of hypohomycetes according to classification proposed by Hughes (1953) as modified by Tubaki (1963) and Barron (1968) and to family Helminthosporiaceae of Hyphomycetes as proposed by Subbraminian (1962). In the traditional classification the genus Helminthosporium is placed under Monilales, family Dematiaceae (darkconidia), phragmosporae. Perfect states where known belong to ascomycetes.

In Helminthosporium conidiophores are determinate, ceasing growth with production of the apical conidia, relatively short and simple or, spariangly branched. The branching is not dichotomous. In Drechslera and Biopolaris the conidiophores are simple or sparingly branched, indeterminate continuing growth sympodially. Conidia lack apical prolongation, and are cylindrical or fusiform, sometimes curved but then concolourous. In Drechslera conidia are cylindric and eminate form all the cells, while in Bipolaris conidia are fusiform germinating from end cells only. Most of the important plant pathogenic genera, especially those attacking Graminaceous hosts, are now considered Drechslera. The type species of this genus was Bipolaris maydis which was transferred from H maydis However, later publicationsa of CMI placed this species as D maydis with perfect state as cochlibolus heterostrophous. Helminthosporium species transferred to Drechslera are given in table;

Helminthosporium sp. H.carbonum H. gramineum H. heveae H. incurvatum H. maydis H. nodulosum H. oryzae H. sacchari H. sativum H. stenospilum H. teres H. turcicum

Drechslera sp. D. zeicola D. graminea D. heveae D. incurvata D. maydis D. nodulosum D. oryzae D. Sachari Bipolaris sativus D. stenospila D. teres D. turcica

Ascigenous state Cochliobolus carbonum Pyrenophora graminea C. heterostrophous C. nodulosum C. miyabeanus C. sativum P. teres Trichometasphaeria turcica

Genus - Alternaria Hyphomycetes (Moniliales - Dematiacae) Spores - Dictyo, Phaeo, Muriform.

The genus generally grows as a saprophyte on the decaying plant parts and also in the soil. some species are parasitic on higher plants. The leaf spots produced by Alternaria species are usually characterized by concentric rings hence the name "target spot". The mycelium is hyaline at first but becomes brown at maturity. In parasitic species the hyphae are intercellular at first and become intracellular later on and the cells are usually multinucleate.

Important Plant Parasitic Species Species Alternata brassicae A. brassicicola A. carthami A. citri A. longipes A. porri A. solani A. triticina Genus - Cercospora (Hyphomycetes, Moniliales, Dematiaceae, phragmo, phaeo-, sympodulo-) The genus cercospora includes about 700 species which are parasites causing leaf spot or lef blotch. Distinct necrotic spots are produced on leaves, stems, fruits, etc. Diseases caused Leaf spot of crucifers Leaf and pod spots of crucifers Leaf spot of safflower Black rot of oranges Brown spot of tobacco Purple blotch of onion Early blight of potato Leaf blight of wheat

Important species Cercospora beticola Cercospora apii Cercospora cruenta - Leaf spot of beet - Early bilght of celery - Leaf spot of phaseolus mungo

Cercospora arachidicola - Early leaf spot of groundnut Perfect state - Mycosphaerella arachidis

Genus - Pyricularia : Pyricularia forms hyaline or pale grey bi or tri celled, pyriform conidia (with a rounded base and basal papilla) singly at the tip of the conidiophore and its successive growing points. The conidiophores are simple, erect, septate and hyaline or grey. P.oryzae causes the most destructive "blast" disease of rice plants. There is much controversy about the name of the pathogen, as two generic names have been used Pyricularia and Dactylaria. The specific name of the fungus on rice was first described as Pyricularia oryzae. Comparing the various characteristics of P. oryzae and P. grisea, it has been concluded that there are not enough differences between these two to maintain them as different species. The morrice isolates is similar to Magnaporthe grisea (ascomycetes) and therefore, M.grisea is retained for the teleomorph stage of rice blast fungus.

Rhizoctonia and Selerotium: The fungi Rhizoctonia and Sclerotium are soil inhabitants and cause diseases on many hosts by affecting the roots, stems, tubers, corns, and other plant parts. These two fungi were known as sterile fungi because they were thought to produce only sclerotia and to be incapable of producing either asexual or sexual spores. The two were distinguished from one another by their mycelial characteristics and by the fact that Rhizoctonia sclerotia have a uniform texture through out where as sclerotium sclerotia are internally differentiated into three areas. It is known now that atleast some species within these two general produce basidiospores as their sexual spores, and, therefore, they are Basidiomycetes. On the other hand, some fungi previously thought to belong to sclerotium, for example, S. bataticola, are known to produce conidia (Macrophomina) and some (e.g. S. oryzae) produce ascospores (Magnaporthe). Others, however In any case, the spores of Rhizoctonia and Sclerotium either are produced under specific conditions in the laboratory or are extremely rare in nature and therefore of little value in diagnosing the fungus. For these reasons, these purposes, they behave as such they continue to be referred by the names Rhizoctonia and Sclerotium.

References : Alexopoules, C.J., C.W. Mims, M. Blackwell (1996). Deuteromycotiva -

Intrductory mycology. John wiley and Sons. pp: 214-258. Mehrotra, R.S. and K.R. Aneja (1999). Deuteromycotina - An introduction to mycology. New International Publication. pp. 561-637. Hriday, S. Chaube, Ramiji Singh (2001). Introductory plant Pathology. International Book Distributing Co., pp: 201-215. Drugs: P. notatum, P. chrysogenum Neurospora genetic analysis Shorter life to study heredity- Shear and Dodge (1927). Aspergillus flavus - oryzae processing many foods and acids.

Characters This group of fungi is characterized by the production of destructive structure, the ascus or asci (Askos = goat skin, sac) within which sexual spores, ascospores are formed by fine cell formation after Karyogamy and meiosis. Eight ascospores are usually produced within an ascus but the number may vary from one to thousand. Except Hemiascomycetes wherein naked asci are produced. The members of ascomycotina form large conspicuous fruiting bodies (ascocarps or ascomata) that enclose the asci. 1. Septate mycelium 2. Absence of motile (cells) spores or gametes 3. Formation of conidia (non-sexual) 4. Dikaryophase in life cycle 5. The cell wall filamentous contains chitin, aminosugars, protein, mannose and glucose also present. Somatic structure Except for some yeasts, these fungi possess well developed septate, profusely branched mycelium with one or more nucleus. Mitochondria, ribosomes, endoplasmic reticulum, vacuoles, microtubules, lipid bodies, lysosomes are the typical cellular components. Three kinds of nuclei may exist in hyphae of ascomycotina.

a. A homokaryotic mycelium that results from uninucleate haploid ascospores. b. A homokaryotic multinucleate condition arising by mitotic duplication of this nuclei and subsequent migration of daughter nuclei. c. Heterokaryotic condition may arise as a result of the mutation of nuclei in an originally homokaryotic mycelium or by fusion of genetically distinct entities that results in dikaryon. d. sclerotia. In certain cases, rhizomorphs, stroma are also produced (compact mass) with or without tissue. Reproduction : Both asexual and sexual ascus or sexual stage often called as ascigenous or perfect stage. Conidial stage is asexual or imperfect stage. Hennebent and Weressub (1977) used Teleomorph sexual form and Anamorph asexual Asexual - Conidia are the chief which are produced in conidiogenous cell which are borne on simple or branched hypha, the conidiophore Budding and fissions are the characteristic of the yeasts. The spores produced by budding are called blastospores. Schizosaccharomyces only budding fragmentation of hyphae and Aggregation of vegetative hyphae may lead to the production of

chlamydospores also other asexual reproductive means conidia are variously shaped, septate or non septate, various colours, single or in chains. There may be produced either directly from somatic hyphae or from specialized conidiogenous cells or from specialized structures called conidioma or conidiomata. Synnemata, sporodochia, pycnidia and acervuli when the hyphae are aggregated to form parallel fascicles of closely appressed hyphae are called synnematous and fructification called synnemata.

Sporodochium It is pad or cushion of mycelia that may get compact to become stromatic although most of it remains loosely interwoven. Simple or branched conidiophores are produced on the surface, eg. Fusarium

Acervulus is composed of a group of closely associated hyphae from the upper layer of which conidia are produced. The entire structure may be on or below the surface of plant host. Eg. Colletotrichum. Pycnidium is flask shaped structure with ostiole Sexual reproduction: Except Hemiascomycetes, Ascomycotina produce some form of fruiting bodies (ascocarp - ascoma) during sexual reproduction. Taxonomy of Ascomycotina is now largely on the development and structure of the fruiting body. The important features of sexual reproduction is the separation of plasmogamy and karyogamy (except Hemi ascomycetes), the sexually compatible nuclei that come to lie together during plasmogamy do not fuse immediately but remain paired and form a dikaryon. The dikaryon gives rise to more pairs of such dikaryons by successive conjugate divisions until karyogamy takes place. The paired male and female gamete nuclei (dikaryon) migrate from the female gametangium known as ascogonia into a hyphal protrusion (called ascogenous hyphae) where they undergo successive conjugate divisions. Ultimately forming binucleate or dikaryotic hyphal segments of female and male nucleus. In most instances, one of these binucleate or dikaryotic cells bends over to form a hook like structure. Within the hook, the two nuclei divide to produce daughter nuclei that are separated by septa that subsequently divide to hook into three cells. Of these three, the tip and basal ones are uninucleate whereas the hook cell is binucleate, the hook cell is known as ascus mother cell is formed at the tip of the ascogenous hyphae and in two nuclei fuse to produce a single diploid nucleus. Thus a dikaryotic phase is thrown between haploid and diploid phases of life cycle. Meiosis follows karyogamy and results in the formation of four haploid nuclei, which undergo mitosis to form 8 haploid nuclei The ascus mother cell gets transformed into an ascus generally containing eight haploid ascospores. In hemiascomycetes, there is no dikaryotic phase and zygote develops directly into an ascus. Plasmogamy in Ascomycotina is brought about by one of the many methods developed by them during evolution.

1. Gametangial contact Morphologically differentiated sex organs are formed in some

ascomycotina. The male is antheridium, the female is ascogonium. The antheridium are multinucleate and more or less elevate while the ascogonium are usually globular or cylindrical and may be single or many celled and commonly coiled. The ascogonium may sometimes have a hair like appendages or outgrowth called trichogyne which act as the receptive organ. Some times in Aspergillus and Penicillum, the ascogonia lack trichogyne and are cylindrical. The gametes are reduced to undifferentiated protoplasts and the male nucleus passes from antheridium into ascogonium through trichogyne or through pore. In some species, though antheridia is present they are non-functional and absent in some cases. In such cases, nuclear pairing will take place either by ascogonial nuclei themselves or from trichogyne and ascogonial nuclei.

2. Gametangial copulation As in Zygomycotina, two similar gametangia touch at their tip or coil around each other and fuses and the fusion cell develops into ascus. In the ascomycetes, species that reproduce by this method lack dikaryophase. Since plasmogamy is followed immediately by karyogamy. In yeast, the somatic cells themselves act as gametangia, fuse and form unicellular zygote which is converted into ascus. This is called hologamy. 3. Spermatization In this mode of plasmogamy, union of spermatia occurs with a receptive structure. Spherical or rod shaped male cells which are produced from short erect structures known as spermatiophores. They are carried by agents like insect or air or water to the receptive organs, the trichogyne or in their absence to somatic hyphae and empty their content into them. In discomycetes and pyrenomycetes, microconidia and conidia serve as spermatia and bring spermatization. Spermatia differ from microconidia or conidia in their enability to germinate and gives rise to a new thallus. 4. Somotogamy Many members in Ascomycotina, there are no sex organs and fusion takes place between two undifferentiated somatic cells of the same or different hyphae and this

process in called somatogamy. The nucleus from one male cell migrate into other and forms dikaryon. The ascus wall may be unitunicate or tritunicate and an important character in the taxonomy of Ascomycotina. The unitunicate asci have two thin wall which is appear as single wall. The tritunicate asci have an outer rigid wall and an inner extensible wall. The inner wall on maturity comes out after the breakage of the outer wall. The extensible inner wall has an apical pore through which the spores are forcibly ejected. Classification: Six classes on the basis of presence or absence of ascocarp and in shape. 1) Hemiascomycetes : Asci naked, no ascogenous hyphae, no ascocarp. 2) Loculoascomycetes: Asci bitunicate ascocarps formed in locules or cavities in stroma formed by hyphal aggregation. 3) Plectomycetes: Unitunicate asci which are typically globose are borne (not in a hymenium) at various level mostly in a cleistothecium ie. dehisce when the ascospores are mating. 4) Laboulbeniomycetes: Specialized ectoparasites on insects. The thallus consists of a foot cell(anchored in the exoskeleton of the host) and a filament of cell which form conidia Asci are formed in perithecia and are deliquescent. 5) Pyrenomycetes: Asci unitunicate cylindrical, persistent arranged in a hymenial layer in cleistothecium or perithecium, asci inoperculate. 6) Discomycetes: Cylindrical asci are arranged in a hymenium and borne in apothecium, epigeous or hypogean.

Plectomycetes The asci are produced at different level (not in a definite hymenium) inside the ascocarp, which is mostly a cleistothecium. The asci are unitunicate and evanescent ie. dehisce at maturity so that the ascospores lie free inside the cleistothecium. When the wall of cleistothecium breaks, the ascospores are released. There are three orders.

Hemiascomycetes

Absence of an ascocarp. Asci are naked and lack of sterile cells. The thallus is made up of a poorly developed mycelium or is represented by separate cells. Cells in the later (yeast) may often remain attached to each other, for varying period of time, thus forming pseudomycelium or cluster of cells. The fungi included in this class are considered to be the most primitive among ascomycotina. Yeast : Saccharomyces cerevisiae is an important in the processing of bread and alcoholic beverages. Ephedrine an important drug for asthma. Hemiascomycetes:Ascospores formed in a compound spore sac, which is equivalent to many asci. Eg. Protomycetales. Asci are formed in a synasius sac. Asci are not formed in a synasius : Asci arise from binucleate,

ascogenous, cells as thin walled sacs Zygote or single cell transformed bursting out of the epispore or by

directly into an ascus or a diploid elongation of the ascogenous cell wall. ascophore. Eg. Endomycetales Protomycetales : Protomyces macrosporus stem gall of Coriander. Single family: Protomycetaceae parasitic on vascular plants and cause galls. There are no ascocarps. The intercellular mycelium is apparently diploid and produces thick walled chlamydospores ie. resting spores. On germination, these form a vesicle, which may be interpreted as an ascus or synascus. Loss due to this disease is about 8.9 % gall size varies from 1.5 cm long. The endophytic and intercellular mycelium forms septa and is converted into multinuclear compartments. Some of these compartments are converted into thick walled resting spore or chlamydospores. The resting bodies are released from infected crop and undergo dormancy. At the time of germination, exospore (outer wall) and forms an elongated more or less cylindrical sac which is called synascus or sporangium. Eg. Taphrinales

A large vacuole develops in thin vesicle and pushes multinucleate protoplast to the periphery. The nuclei undergo meiotic division and produce endospores per nucleus. The four spores formed following each reduction compared with a wallless ascus and the large number of asci in the spore sac makes it a synascus. The endospores are uninucleate and multiply by budding and artificially they can be cultivated (yeast state). These endospores conjugate in pairs, with one member of each pair continuing to bud and form diploid mycelium prior to infection. These fused spores germinate only on the host and are capable of causing infection while the unfused fail to infect. Endomycetales or Saccharomycetales:

Yeast produces ascospores either through copulation of two cells or partheno genetically develops directly into an ascus, somatic structure may be unicellular, thallus or pseudomycelium. Asexual reproduction - Fission, budding. Sexual reproduction - Fusion of two equal or unequal cells of two ascospores. The diploid cell enlarges and become single ascus and produces ascospores. Taphrinales: Taphrinales are parasitic on higher plants and cause malformation of tissues causes leaf curl in which puckering of the leaves, witches broom, close clustering of twigs, fruits are wrinkled, shrivelled and has a cavity in the centre. Sub-cuticular or sub epidermal layer of binucleate ascogenous cells from the mycelium. The asci and spore sacs that arise terminally hyaline, sub-globose ascospores that often bud to form yeast like colonies.

Taphrinaceae Taphrina deformans Peach and Almond leaf curl. T. pruni- Plum (pockets) T. maculans Zinger, turmeric leaf spot

95 species are recognized. Thickening of leaves due to hypertrophy and hyperplasia.

Increased cytokinin activity and Indole 3- acetic acid and tryptophan and hormonal imbalance. While the ascospores are still present in the ascus or soon after their release from asci, they start budding and produce small ovoid blastospores. On reaching the surface of the leaves uninucleate haploid blastospores produce germ tube and penetrate the host tissue and form intercellular mycelium. But before penetration the host, the single haploid nucleus of the blastospores divides to establish dikaryon, the binucleate (dikaryotic) condition of the hyphae in the host is perpetuated by the conjugate division of the nuclei. However, the hyphae of thin fungus are multinucleate and the nuclei are occurring in pairs. These aggregate below the leaf cuticle and breaks up into binucleate ascogenous cell. Karyogamy takes place in the elongated ascogenous cell (Chlamydospores) and immediately in mitosis. One daughter nucleus migrates into apical portion and the other remains at the base; a septum form in the mother cell contains one nucleus each. The ascus mother cell continues to elongate and ruptures the leaf cuticle. The base stalk cell containing nuclei will disintegrate. The diploid nucleus of the ascus divides by meiotic and mitotic division and form nuclei. Then each nucleus gets enclosed by double delimiting membrane and forms eight ascospores. The ascospores bud either in ascus mother cell or and numerous ascospores are formed called blastospores. The asci are formed over the epidermis and its presence gives the leaf a waxy bloom. With the bursting of epidermis, the ascospores / blastospores liberate into air and repeat the cycle. Pyrenomycetes The pyrenomycetes are defined as ascomycetes with ascocarps entirely surrounded by a periodical wall and containing unitunicate asci, which are typically globose, or flask shaped perithecia and are provided with an opening, the ostiole which is lined by hypha like periphyses. The structures lying inside the ascocarp constitute centrum. It includes asci and paraphyses. Claviceps : The modern classification of pyrenomycetes is based upon relationship between the ascus and other components of the ascocarp occupying the perithecial cavity or locule within which asci develops.

Orders: Erysiphales, Meliolales, Xylariales, Clavicipitales, Diaporthales, Hypocreales, Coronophorales and Coryneliales.

Erysiphales: Powdery mildews The member produces superficial white mycelium attached by haustoria. Ascocarps are globose, non-ostiolated, open irregularly are often provided with characteristic appendages. The number of asci varies from one to many per ascocarp. Asci are pyriform to clavate, unitunicate, Ascospores are one celled. Asexual stage: (Oidium) Acrosporium oidiopsis and Ovulariopsis. Erysiphales : 28 genera, 105 species. E. polygoni (Salmons) 357 hosts and 157 genera. Podosphaera leucotricha on apple. The family Erysiphaceae has been placed in the Plectomycetes by some and by other in Pyrenomycetes (Miller 1941, 1949) included in Plectomycetes because of the non-ostiolated ascocarp. But Luttrel (1951) transferred to Pyrenomycetes because the asci are produced in basal fascicles and discharged forcibly (presence of asci in definite layer, the hymenium. Cleistothecium- mycelium with hymenium. Phyllactinia, Leveillula - Internal (Haustoria). Asexual conidia and conidiophores basipetal.

Clavicipitales: Parasitic on grasses (Claviceps), on insects and on other fungi. Characteristic features-perithecia which are superficial or immersed in fleshy, bright coloured stroma made up of fungal tissue. Narrow and cylindrical asci with a typical apical cap. Filiform or fusiform multiseptate ascospores that often fragments into short segments, each segment behave as a spore. Claviceps oryzae sativae False smut; C. fusiformis ergot on bajra; C. purpurea Rye ergot. Cyperaceae, gramineae. The genus is characterized by the erect stroma with a long stalk and globose head arising from sclerotia.

The perithecia are produced towards the periphery of the stroma completely immersed. The perithecia are isolated. Ergomatine, Ergotamine and Ergotoxime. Epichloe-E. typhina Choke disease: Formation of effused stroma that surrounds the host shoot. The ascospores are eight numbered. Hyaline, filiform, multiseptate and separate within the ascus and are bitunicate. Meliolales: Black mildews- since they are black covering the surface of the host. Similar to erysiphales, epiphytes are having obligate mycelium. Thick, dark septate hyphae. The ascocarps are superficial on the mycelium and are produced either ostiolated or rudimentary ostiole. The asci are 2 to 4 rarely 8. The ascospores are dark brown, smooth, thick walled and contain 5 cells. Asexual reproduction is not known. M. jasminicola, M. argentina, M. palmicola.

Sphaeriales: Syn.- Xylariales. Ascocarps are dark leathery or carbonous, globose or pear shaped. Single and immersed in stromata, ostiolate or non-ostiolated. Asci are unitunicate, ellipsoid to cylindrical and intermixed with paraphyses forming persistant hymenial layer. Mainly saprophytes. Wood decaying on herbaceous plants, parasiticPhyllachora graminis - tar leaf spot of wheat. Rosellinia necatrix - root rot of grape vine. Nummularia discinicola- Nail head canker in apple. Glomerella- bitter rot in apple. R. arcuata- black root rot of tea R. bunodes Coffee, rubber, cacao, tea etc. R. necatrix- white root rot of grapes. No Rhizomorphs. Black sclerotia are formed.

Rosellina necatrix-imperfect stage, Dematophora necatrix synnemata. The conidiophores aggregate to form synnemata. The fungus produces globose, black, shortly pedicillate perithecia embedded in septate hyphae. The asci are cylindrical, long stalk, 8 spored. Eg. Xylaria.

Diaporthaceae: No pathogens. Phyllachoraceae: Black ostiolate, ascomata embedded in the host tissue. Phyllachora -very large and about 107 species are there. And about 1120 hosts and 105 families. Tar leaf spot -P. maydis on maize. P. musicola on banana P. sorghi sorghum P. graminis grass The fructification is simple or compound at maturity. Ascocarps generally crowded in the mesophyll. The ascocarp is provided with a pore. The hymenium is made up of asci and filiform paraphyses. The asci are cylindrical or broadly ellipsoid and are uniformly thickened with an apical pore for liberation of ascospores. The ascospores are unicellular, hyaline to black. Variously shaped conidia of uncertain phylogeny are borne similar to fructification present in the same species. Spermatia are short and filiform. In leguminaceae family, the bacilloid spermatia whereas in graminae, myrtaceae produces filiform spermatia (Linochora genus).

Polystigmataceae: Non-stromatic or under developed stroma whose ascal apexes do not stain blue with iodine and ascospores are without germpores. Ascospores produces appressoria by the germination of ascospores. Most of them are parasites. Glomerella, Physalospora and Polystigmata. Production of spherical or flask shaped and sometimes hairy periphysate perithecia embedded in stroma or host tissue. The asci are short stalked; 8 spored and are intermingled with paraphyses. Ascospores are unicellular, hyaline and curved.

Imperfect Stage Colletotrichum causes anthracnose. Glomerella cingulata Bitter rot of apple. Colletotrichum gloeosporioides citrus, anona, avacado, mango and banana G. lindemuthianum Bean G. tucumanensis Physalospora tucumanensis sugarcane red rot. G. graminicola maize. Conidia - Acervuli conidia cylindrical, pinkish mass.

Diaporthales: It is characterized by a Diaporthe type centrum. Initially a mass of plectenchyma into which asci and paraphyses push themselves from the base, thus forming the centrum. The perithecia are usually immersed. The asci are discharged inside at the time of maturity. Diaporthaceae: Presence of stroma with the perithecia are buried in it. The ascospores are two celled, hyaline and these are liberated within the ascus and easily liberated. Diaporthe: Largest genus with 75 species. D. batatas All species (Phomopsis batatas) storage rot in sweet potatoes D. citri (P. citri) Stem end rot in citrus D. vexans Fruit rot of Brinjal Conidia Pycnidial fungi Two types hyaline a, a spores. Oval and unicelluar while spores called stylopores are filiform and hooked at one end like walking stick. The stromatoid cushions later gives rise to immersed or superficial perithecia. Strains: When + and strains mated in Petri dish culture, perithecia are formed on the line of contact. Two uninucleate branches from the somatic hyphae develops into an outer and inner coil. The inner coil act as ascogonium while the outer coil develops into peridium covering the inner. This is called as Protoperithecium. Plasmogamy takes place between two conidia of opposite strain. The conidia germinate to produce germ tube and grows towards protoperithecium and enters onto ascogonial coil. The wall dissolves and gives rise to ascogenous hyphae and crozia formation takes place.

Sordariaceae: Neurospora Drosophila of the fungus. Endotheca: E. parasitica Cheshnut blight or canker Perithecia having long beak, black and deeply immersed in a yellow and fleshy stroma. The asci are club shaped and spore without paraphyses. Ascospores are two celled, hyaline and constricted at the septum. Pycnidial formation is earlier than perithecia.

Gnomoniales Endotheca E.parasitica- chestnut blight or canker Perithcia having long black and deeply immersied in a yellow and fleshy stroma .The asci club shaped and 8 spores without paraphyses.Ascospores two ccelled hyaline and constructed at the septum. Gnomoniales-Non-stromatic family Gaumanomyces 4 species,G.graminis earhead and take all of creales. Hypocreales- Nectria type ,Apical paraphyses arise from the perithecial apex below the periphyses from the roof of the centrum and grow downward and they donot reach the base of cavity Imperfect Acremonium, Fusarium Trichoderma , Gliocladium, Verticillium,Tubercularia,Cylindrocarpon. Hypocreaceae- Perithecia superficial or sunken in pseudoparenchymatous stroma or without stroma. Hypomycetaceae -Perithecia are immersed in a loose Prosenchyma tousubicalum Hypocreaceae;-the ascocarps are fleshy bright often white and produced either superficially or immersed in stroma Nectria,Hypocrea ,Gibberrella, N.haematococoaconidial F.solani , N.radicola rootrot in temperate crops .the imperfect states of nectria belongs not only to a single genus but to a number of genera. -White

Acremonium, cylendrocarpin, Fusarium Tubercularia, indicating no direct correladon between perpect and imperpect stalks to genera like Eurotium and Talaromyces where all produces Aspergillus and penicillium like improvement states. Acremonium Gibberella Fusarium The stroma in erumpent and pseduoparen chymatous,blue or violet coloured perithecia are globose produced superficially in immersed in host tissue.

G. fuzikuroi Kakanae partheno carpy Gibberellin A1, A2, A3

Elongation and taller than healthy induction of

Plectomycetes The asci are produced at different level (no in a definite hymonium inside the ascocap. Which is mostly claistthecium. The asci are unitanicated and evanescent ie dehisce at maturity. So that the ascospores he free inside the cleistothecium when the wall of the cleistothecium breaks.

Three orders

Eurohales

Onygenales Microascal Eurotiarcxeae Aspergillus

Penicillium Conidia are produced in chain on phialidess

Discomycetes

Fungi that produce fructification with an exposed hymenium called apothecia Most of these are cup shape or saucer shaped but in some groups. They are club shaped or mushroom shaped. The morels, which also belong to this class, are spongy like and those of false morels have bell like apothecia. Ascopores are liberated forsibly except in Truffles (Dispensed through animals such as pigs and Deer etc.).

Sclerotinia (Monilinia fructicola) S.luxa, S.frucigena causes brown rot of apple, peas and peach Morchells most eatable fungi Helvellas Helvellic acid, Very good decomposer. Hymenium, subhymenium (hypothecium) exciputum. In some discomycetes, pseudoepithecium and epithecium may be present. The hymenium is made up of cylindrical or club shaped asci and paraphyes in a palisade layer. When paraphyses are longer than asci, they may be glued together in matrix to form pseudoepithicum. Classification The presence or absence of an operculum at the ascal apex, the mechanism of dehiscence and the reaction of the ascal tip to iodine are important character for classifying these fungi. 1. Medeolariales 2. Cyttariales a broad pore in present 3. Tuberales asci donot rupture 4. Pezizales apical or sub apical operculam 5. Phacidiales inoperculate 6. Ostropales - inoperculate 7. Helotiales - inoperculate Helotiales: Inoperculate discomycetes contains 345 genera, 2000 species Sclerotinia fructigena (Monilinia fructigena)

S. fructicola Brown rot of apple, pear. S. sclerotiarum- carrot, cabbage, potato. Lettuce - Stromatinia gladioli-corm rot Sclerotiniaceae: Characterised by cup shaped, yellowish brown stalked apothecia that develops from stromata or sclerotia. The ascus pore stains with iodine. The ascospores are hyaline unicellular, oval, 28 genera with 245 species. facultative saprophytes. Segregate genera: Botryotinia, Gloeotinia, Monilinia or Septotinia on the basis of asexual state, Botrytis, Endoconidium, Monilia, Septotis. Some Sclerotinia dont have conidial stage and only spermatial stages are present. Conidia oval, lemon shaped, unicellular, break easily and dispersed. The mummified fruits are fall on the ground and buried. Apothesia are arise after one year or so and some of them produces spermatia. They are parasite and

Laboulbeniomycetes: Mostly insect parasites superficial thallus with perithecia bearing trichogynes and antheridia with non-motile spermatia and do not produce true mycelium.

Loculoascomycetes: Bitunicate asci and septate ascospores produced in unwalled locales (pseudothecia) in ascostromatic ascocarps. The asci develop in the fruit body, which is regarded as an ascostroma. The ascostroma is cushion or mattress shaped. The stroma may be unilocular or multilocular. It is very difficult to distinguish from true perithecium and unilocular stroma. Important-Mycosphaerella musicola. Venturia inaequalis There are three types of centra. Elsinoe asci arise singly in monoascus locules, which are separated by relatively unaltered stromatal tissue. Dothidea- centrum is characterised by the production of aparaphysate asci-in fascicles in locules. The centrum tissue is pseudoparenchymatous.

Pleospora: The locule is formed by growth of vertical sterile hyphae. The pseudoparaphyses, which arise from the roof to meet the floor of the stroma, where ascogonium located. Pseudosphaeria : Monascus locules are formed close to each other. The bitunicate ascus is equipped with apical structure called asse apicale.

Classification: Myringiales: Produces typically globose, thick walled asci that develop singly in uniascal locules immersed in a stromatic body which opens by rupture or dehiscence. Myringiaceae: Asci, spherical or broadly clavate and are borne in uniascal locule. The asco spores are multiseptate. E. fawcetti Citrus. E. phaseoli - Lima bean Elsinoe Sphaceloma - conidial stage Melanobasidium - Conidial stage (Melanconiales). E.veneta -conidial Sphaceloma venetum - anthracnose of raspberries. E.mangi ferae scab on mango. E.sacchari - white rash sugarcane.

Dothidiales: Bitunicate with locule containing several cylindrical asci without paraphyses. The locule may be ostiolated or non-ostiolated. Mycosphaerella, Guignardia, Pleospora

Dothidiaceae or Mycosphaerellaceae: Small, cylindrical or spherical ascocarp (pseudothecia), 8 spored, bitunicate asci without filamentous paraphyses and 2 celled hyaline ascospores. The ascocarp is 2 celled hyaline ascospores. The ascocarp developes within the host cell.

Mycospharella

Coelomycetes Hyphomycetes

Mycosphaerella arachidis early leaf spot. M.cercosporidium Late leaf spot. M.brassicicola black ring. Mycosphaerella Septoria Cercoseptoria Cercospora Pseudocercospora Cercosporella Heterosporium Ramularia Ovularia, Cladosporium Plasmogamy takes place through spermatization in many species. The

spermatia are formed in flask shaped structure. Spermagonia, which resemble like pycnidia and confused with conidia.

Capnodiaceae: Sooty molds: Bitunicate asci borne in ostiolate ascocarps with many septate ascospores. It is an epiphytic saprobes living on honey dew exudates of insects. Capnodium, Limacinia genera.

Pleosporales: Bitunicate with paraphysate asci. Pleospora centrum in which locule are spherical and large or medium sized ostiolated or non ostiole and asci are surrounded by pseudoparaphyses. Pseudoparaphyses are originated in the upper wall grow downward and meet the basal wall attached to both the roof and floor of the locule. Venturia inaequalis - apple scab.

Pleosporaceae Multiseptate ascospores that lack germ pore or slits produced in parallel asci intermingled with filamentous paraphyses. Asexual Pycnidia Conidia: Phoma Coniothyrium Stagenospora Hendrosonia Gamarosporium Conidia are many septate. Spermatia are also present. Alternaria Drechslera Curvularia Stemphylium

Sexual: Pleospora, Pyrenophora Cochliobolus, Leptosphaeria Magnaporthe Pleospora herbarum(Stemphylium botryosum) leaf spot of legumes, lettuce and storage rot of apples.

Cochliobolus Filiform and many celled, ascospores coiled in the ascus. Dark, many septate conidia. (Drechslera, Bipolaris, Exerohilum, C.miyabeanus, C.victoriae -host specific toxins (Victorin) (Oats). Leptosphaeria: Leptosphaeria maculans (Phoma lingam) L.sacchari - sugarcane ring spot L. elaeidis - Pestalotiopsis. Magnaporthe

Venturiaceae: Pseudothecia are immersed or erumpent and are produced below the cuticle or epidermis. One septate -ovoid greenish or brown ascospore with smooth walls. V.inaequalis - apple scab. Venturia pirina -pear scab. conidial hyphomycetes. Venturia spilocaea or V. fusicladium(F. destriticum) V.inaequalis - Spilocaea pomi(imperfect stage). The spores are unequal. The ascospores are unequally two celled that is why the name V.inaequalis.

DEUTEROMYCETES This is the second largest subdivision of the fungi and contains 1680 genera, 17000 species (Hawksworth et al., 1983). Fungi reproducing exclusively by asexual means usually by conidia are placed in subdivision - Deutecomycotina. Mycelium - Well developed, branched, septate, multinucleate hyphae. Hyphae - well developed, branched, septate The sexual stages belongs to Ascomycotina & Sometimes Basidiomycotina. (e.g) Imperfect (Anamorph), Perfect (Teliomorph) Ascomycotina -Fusarium moniliforme - Gibberella fujikuroi Basidiomycotina - Rhizoctonia solani - Thanatephorus cucumeris

Deuteromycetes: (imperfect / asexual fungi) Mycelium well developed, septate branched, sexual reproduction rare (or) unknown, Asexual spores (Conidia). From on conidiophores existing singly or grouped in specialized structures. Eg: Sporodochia, synnemata or produced in pycnidia and acervuli.

Asexual fruiting bodies 2. Sporodochium : Is is cushion like aggregation of hypha which breaks

through host and bears conidia. eg. Fusarium, Epicoccum .

2. Acervulus : Formed by the aggregation of somatic hyphae below the epidermis of the host forming cushion like structure on which conidiophores and conidia formed. Eg: Colletotrichum capsici 3. Pycnidia : When spore and sporophores are formed inside a specially designed fruit bodies which have protective wall made up of fungal tissue, they are termed as pycnidia. eg: Macrophomina phaseolina.

4. Synnemata : Loose aggregation of errect conidiophore similar to mycelial strands eg: Arthrobotryum.

Deuteromycotina

Blatomycetes Coelomycetes * Budding yeast / yeastlike Mycelium well

Hyphomycetes

* Mycelium well developed

developed * with/without pseudomyclium * Budding cells absent * True mycelium lacking/not pycnidium well developed acervuli * Pycnidia/acervuli absent spores directly/on sporophores. Or * Mycelium sterile / bears * Spores in * Budding cells absent

Hyphomycetes Order

Moniliales (1) Moniliaceae Verticillium, Aganomycetaceae Aspergillus, Penicilium, Sarocladium.

Agonomycetels

(2) Dematiaceae Helminthosporium, Pyricularia, Culvularia, Alternaria, Cercospora.

(3) Tuberculariaceae Fusarium .

(4) Stillbellaceae synnemata,

Coelomycetes Order

Melanconiales Sphaeropsidales Frucitification acervuli (or)stomata Fructification pycnidia

Melanconiaceae Colletotrichum Sphaeropsidaceae Phoma, Phomopsis Botryodiploida,

Coelomycetes: There are 650 genera, 8000 species in this class (Hawksworth et al., 1983). Grove (1919) introduced the term coelomycetes to include those imperfect fungi which are formed inside a cavity formed in the substrate on which fungus grows. Melanconiales: General character: It is characterised by the production of acelvuli which may develop sub

epidermally / sub cuticulary. Acervulus - It is a saucer - shaped fructification in which the conidiogenous cells develops on the floor of the cavity from a pseudoparenchymatous stroma beneath an integument of host tissues which rupture at maturity (Hawksworth et al., 1983). The conidia from acervuli are released in the form of characteristic

droplets which may be variously colored from hyaline, to cream, pink orange or black. Over 120 genera are included in this family,

eg: Antheacnose, Colletotrichum, Pestolotia, Pestalotiopsis, monochactia, melanconium.

Sphaeropsidales : General character : pynidia. Mycelium may be immersed in the substrate / superficial. Pycnidia are flask shaped to globose fructification with slightly tapered The conidia and conidiogenous cells (or) conidiophores are produced in

apex and single ostiole. Inside the pynidium hymenium is present which bears conidia. They may be produced superficially, subimmersed (or) immersed in the substratum. The wall of the pynidium is made up of pseudoparenchymatous tissue and

the conidiophores arise from cell of the inner wall.

Hypomycetes: moniliales(order): Tuberculariaceae: This order contains many genera. It includes Fusarium, Tubercularia, Volutella, Epicoccum, Exosporium.Special feature of this order is the production of sporodochia in which the spore mass is supported by a superficial, cushion like (Pulvinate) mass of conidiogenous celles (or) short conidiophores.

Aganomycetales : The fungi included in this order are often referred to as mycelia sterilia, as

they lack even imperfect stage. ie., in which no spores of any kind are produced and reproduce only by fragmentation of the mycelium. the pathogen. 42 genera included in this order (kendrick and Carmichavel, 1973) The do form sclerotia / chlamydospores which perpetuate and disseminate

including Aegerita, Arbuscula, Burgoa, Papulaspora, Dactuliosphora, Rhizoctonia and Sclerotium.

Genus - Colletotrichum (Deuteromycetes, coelmycetes, Melanconiales, Melanconiaceae, Amero-, Hyalo-, Phialospores). This genus is a facultative parasite causing anthrocnose diseases on leaves, young twigs and fruits of many plant species. Important species: Colletotrichum capsici- Dieback and ripe fruit rot of chillies, other hosts chickpea, cotton, brinjal, tomato, turmeric. Colletotrichum coccodes Hosts belong to solanaceae, leguminosae, and cucurbitaceae. The species causes black dot of potato, and tomato and anthracnose of fruits of tomato and chilli.

Colletotrichum truncatum Attacks groundnuts, pigeonpea, soybean, pea, cowpea, Phaseolus spp. Causing anthranose and leaf dpots. Colletotrichum lini - Anthracnose of linseed. Colletotrichum falcatum (Glomerella tucumanensis)- Red rot of sugarcane. Colletotrichum lindemuthianum(Glomerella lindemuthianum)Anthracnose of beans. Genera - Helminthosporium, Drechslera and Biopolaris: These three genera belong to series porosporae of hypohomycetes according to classification proposed by Hughes (1953) as modified by Tubaki (1963) and Barron (1968) and to family Helminthosporiaceae of Hyphomycetes as proposed by Subbraminian (1962). In the traditional classification the genus Helminthosporium is placed under Monilales, family Dematiaceae (darkconidia), phragmosporae. Perfect states where known belong to ascomycetes.

In Helminthosporium conidiophores are determinate, ceasing growth with production of the apical conidia, relatively short and simple or, spariangly branched. The branching is not dichotomous. In Drechslera and Biopolaris the conidiophores are simple or sparingly branched, indeterminate continuing growth sympodially. Conidia lack apical prolongation, and are cylindrical or fusiform, sometimes curved but then concolourous. In Drechslera conidia are cylindric and eminate form all the cells, while in Bipolaris conidia

are fusiform germinating from end cells only. Most of the important plant pathogenic genera, especially those attacking Graminaceous hosts, are now considered Drechslera. The type species of this genus was Bipolaris maydis which was transferred from H maydis However, later publicationsa of CMI placed this species as D maydis with perfect state as cochlibolus heterostrophous. Helminthosporium species transferred to Drechslera are given in table; Helminthosporium sp. H.carbonum H. gramineum H. heveae H. incurvatum H. maydis H. nodulosum H. oryzae H. sacchari H. sativum H. stenospilum H. teres H. turcicum Drechslera sp. D. zeicola D. graminea D. heveae D. incurvata D. maydis D. nodulosum D. oryzae D. Sachari Bipolaris sativus D. stenospila D. teres D. turcica P. teres Trichometasphaeria turcica Ascigenous state Cochliobolus carbonum Pyrenophora graminea C. heterostrophous C. nodulosum C. miyabeanus C. sativum -

Genus - Alternaria Hyphomycetes (Moniliales - Dematiacae) Spores - Dictyo, Phaeo, Muriform. The genus generally grows as a saprophyte on the decaying plant parts and also in the soil. some species are parasitic on higher plants. The leaf spots produced by Alternaria species are usually characterized by concentric rings hence the name "target spot". The mycelium is hyaline at first but becomes brown at maturity. In parasitic species the hyphae are intercellular at first and become intracellular later on and the cells are usually multinucleate.

Important Plant Parasitic Species Species Alternata brassicae A. brassicicola crucifers A. carthami A. citri A. longipes A. porri A. solani A. triticina Genus - Cercospora (Hyphomycetes, Moniliales, Dematiaceae, phragmo, phaeo-, sympodulo-) The genus cercospora includes about 700 species which are parasites causing leaf spot or lef blotch. Distinct necrotic spots are produced on leaves, stems, fruits, etc. Leaf spot of safflower Black rot of oranges Brown spot of tobacco Purple blotch of onion Early blight of potato Leaf blight of wheat Diseases caused Leaf spot of crucifers Leaf and pod spots of

Important species Cercospora beticola Cercospora apii Cercospora cruenta - Leaf spot of beet - Early bilght of celery - Leaf spot of phaseolus mungo

Cercospora arachidicola - Early leaf spot of groundnut Perfect state Genus - Pyricularia : Pyricularia forms hyaline or pale grey bi or tri celled, pyriform conidia (with a rounded base and basal papilla) singly at the tip of the conidiophore and its successive growing points. The conidiophores are simple, erect, septate and hyaline or grey. P.oryzae causes the most destructive "blast" disease of rice plants. There is - Mycosphaerella arachidis

much controversy about the name of the pathogen, as two generic names have been used Pyricularia and Dactylaria. The specific name of the fungus on rice was first described as Pyricularia oryzae. Comparing the various characteristics of P. oryzae and P. grisea, it has been concluded that there are not enough differences between these two to maintain them as different species. The morrice isolates is similar to Magnaporthe grisea (ascomycetes) and therefore, M.grisea is retained for the teleomorph stage of rice blast fungus.

Rhizoctonia and Selerotium: The fungi Rhizoctonia and Sclerotium are soil inhabitants and cause diseases on many hosts by affecting the roots, stems, tubers, corns, and other plant parts. These two fungi were known as sterile fungi because they were thought to produce only sclerotia and to be incapable of producing either asexual or sexual spores. The two were distinguished from one another by their mycelial characteristics and by the fact that Rhizoctonia sclerotia have a uniform texture through out where as sclerotium sclerotia are internally differentiated into three areas. It is known now that atleast some species within these two general produce basidiospores as their sexual spores, and, therefore, they are Basidiomycetes. On the other hand, some fungi previously thought to belong to sclerotium, for example, S. bataticola, are known to produce conidia (Macrophomina) and some (e.g. S. oryzae) produce ascospores (Magnaporthe). Others, however In any case, the spores of Rhizoctonia and Sclerotium either are produced under specific conditions in the laboratory or are extremely rare in nature and therefore of little value in diagnosing the fungus. For these reasons, these purposes, they behave as such they continue to be referred by the names Rhizoctonia and Sclerotium. References : Alexopoules, C.J., C.W. Mims, M. Blackwell (1996). Deuteromycotiva -

Intrductory mycology. John wiley and Sons. pp: 214-258. Mehrotra, R.S. and K.R. Aneja (1999). Deuteromycotina - An introduction

to mycology. New International Publication. pp. 561-637. Hriday, S. Chaube, Ramiji Singh (2001). Introductory plant Pathology.

International Book Distributing Co., pp: 201-215.

BASIDIOMYCOTINA Systematic Classification: Kingdom : Fungi Division : Eumycota Sub Division: Basidiomycotina Class : Teliomycetes Order : Uredinales Family : Puccinaceae Melampsoraceae Genus: Puccinia Gymnosporangium Uromyces Haemilia Cronartium Melampsora Uredinales: 150 genera, 6000 species Intercellular, Haustoria Lack of clamp connection, dolipore septum Teliospores produced terminally Sporidia discharged forcibly Pleiomorphic life cycle No basidiocarps Obligate parasites Puccinia graminis tritici P. helianthi Melampsora lini Autoecious Heteroecious

Major Epidemic: 1946-47 Madhya Pradesh 1956-57 - Bihar 1971-72 Punjab, Uttar Pradesh 1972-73 Haryana, Hills of Himalaya 1974 Rajasthan One third of total Crop in 20,000 ha. Life cycle pattern: Uredinales: 1. Macrocyclic: Five productive stages Puccinia helianthi(A) and P. pennisetti(H) 2. 3. Microcyclic: Teliospore, Binucleate spore P. malvacearum Demicyclic: Uredinia absent G. juniperi- Virginianae (H) Xenodocus carbonarius (A) Five Stages of Macrocyclic Rust: Stage 0 Spermagonia Spermatia, Receptive hyphae Stage 1 Aecia Aeciospores Stage 2 Uredinia Urediniospores Stage 3 Telia Teliospores Stage 4 Basidia Basidiospores

Stage 0 : Spermagonia (Pycnia) Spermatia: male sex organ Receptive hyphae: Female sex organ Basidiospore primary mycelium Monokaryotic gametes Flask shaped, ostiole Spermatia insects Spermatization Self-sterile

Stage 1 - Aecia Aeciospores Group of Dikaryotic hyphal cells chain of dikaryotic aeciospores Non-repeating vegetative spores Usually associated with spermagonia Matured spores released Four types of Aecia: Cupulate / Aecioid Aecia Eg. Puccinia Peridermioid Aecia Eg. Cronartium Roestelioid Aecia: Eg. Roes telia Caeomoid Aecia: Eg. Melampsora

Stage 2: Uredinia Urediniospores (Red Rust spores, Summer Spores) Repeating Vegetative stage Mycelium dikaryotic Uredospores are unicellular, pedicellate, deciduous, echinulate (2/3 germpores). No peridia Urdinopsis and Hyalospora Amphispores

Stage 3: Telia Teliospores (winter spores, black rust spores) Perfect stage, unicellular/2/ more celled, sessile or pedicellate, not thick walled, ornamented. Initially dikaryotic Karyogamy (diploid, uninucleate) resting Ger. Promycelium (Meiosis) Four nuclei (Four cell) Basidiospores. Sterigma period deciduous,

Stage 4: Basidia Basidiospores (Sporidia) Promycelium Metabasidium Short lived exogenously Forcibly discharged Directly germinate Forms sterigma sporidia.

Autoecium Single host group

Heteroecium Two groups of host 0 & 1 one Puccinia helianthi II & III Another group Eg. P. graminis Berberis vulgaris (0 &1) II & III Cereals P. pennisetti Brinjal Cumbu

Specialization of Parasitism: Ability to attack various host P. graminis (third name) depending upon the host P.g. tritici, P. g. hordei, P. g. avenae and P. g. poae Races Differences in Pathogenicity Biotypes Population of individuals Genetically same. Uredinales Puccinaceae Melampsoraceae Coleosporiaceae

Puccinaceae: stalked teliospores, free, united. Important Genera: Puccinia : Heteroecious / Autoecious Macrocyclic / Microcyclic Largest genus, Teliospores are two celled. Black stem rust P. g. tritici. Brown / Leaf rust P. recondita Yellow / Stripe rust P. striiformis Sorghum rust P. purpurea Cotton rust P. cacabata

Sugarcane rust P. erianthi Groundnut rust P. arachidis Guava rust- P. psidii. Pucciniaceae: Important Genera: 1. Puccinia 2. Uromyces 3. Phragmidium 4. Gymnosporangium 5. Ravenelia 6. Haemilia Epidemiology of wheat rust India South India Inspection earlier North Black rust Throughout summer north Not play a significant role spreading the disease. Temperature- 7-130C, Incubation period 40 to 50 days Feb. Spread of the inoculum from south (Joshi et al., 1982) Genus: Gymnosporangium G. bermudianum Autoecious rust. All are Heteroecious demicyclic Spermagonia and Aecia belong to Rosaceae Telia Gymnosperms G. juniperi virginiae Apple cedar rusts Genus: Uromyces Teliospres stalked, unicellular, apical papilla U. appendiculatus bean rust- auto, macrocyclic U. fabae peas rust U. ciceris- arietini Gram rust Heterocious (S & A) U. pisi Peas rust Heteroecious Euphorbia cyparissias. Haemilia: Urediospores sub stomatal, reniform urediospores, specialized type Teliospores unicellular, without germ pores H. vastatrix Coffee rust 1870 Ceylon Export 50 m. Kg. 1885 export about zero. Rust sori lower surface of the leaf. Autoecious rust but Basidiospores do not infect coffee. Uredio and Teliospores of H.vastatrix.

Family: Melampsoraceae Sessile teliospores Genera: Cronartium Melampsora Cronartium Macrocyclic, Heteroecious Spermagonia and aecia - Pinus Uredinia and telia Dicotyledons Teliospores fused finger like columns C. ribicola Blister rust white pine. Melampsora has single celled, sessile teliospores Mostly heteroecious M. lini Rust of linseed / flax Autoecious, Macrocyclic Spermagonia, Aecia, Uredinia and Telia Flax.

ORDER : Agaricales The order Agaricales commonly called gill fungi contains 270 genera and around 4,000 species. These includes the mushrooms, the toadstools and the boletus.

Systematic Position: Kingdom : Fungi Division : Eumycota Subdivision: Basidiomycotina Class: Hymenomycetes Subclass: Holobasidiomycetidae Order : Agaricales Occurrence and Importance: Terrestrial, mostly saprophyte on dead leaves, wood and dung. Some are mycorrhizal, very few pathogenic eg. Armillaria and Marasmius. Nutritious Rich in minerals and proteins medicinal value stimulate digestion, cure hypertension and improve blood circulation.

mushroom cultivation 17th Century Paris mushroom 27,000 32,000 kg dry protein/acre/year beef 32 kg/acre/year. Fishfarm 260 kg/acre/year. Commercial Edible Mushrooms: Agaricus campestris var. bisporus (button mushroom) Volvariella volvacea ( Paddystraw mushrrom) Lentinus edodes (Shitake mushroom) Pleurotus ostreatus (Oyster mushroom) Tricholoma matsutake Agaricus brunnescens is commercially grown in Europe, America and temperate climate in India. Volvariella volvacea is a tropical mushroom which is cultivated in South east Asian countries. There is no single test of knowing whether a mushroom is edible or poisonous. The best precaution is that we should not touch any mushroom that has a volva at the base.

Poisonous Mushrooms: Amanita phalloides Death cap. A. virosa destroying angel A. verna Fools cap. A. muscaria Fly agaric.

Mushroom poisoning: Mycetism poisoning by large fungi in man. Types of mushroom poisoning: A. Toxins Cellular destruction, liver, kidney damage and death. Onset of symptoms 10 hours. Group 1 Deadly cyclopeptida (amanitin) poisoning Genera Amanita, Galerina.

Group 2 Deadly monomethylhydrazine (gyrometrin) poisoning Genus: Gyromitra (Helvella). B. Toxins Autonomic nervous system. onset of symptoms 20 min. to 2 hours. Group 3 Coprine (Antabuse-like) poisoning Genera: Coprinus. Group 4 Muscarine (sweating) poisoning Genera : Clitocybe and Inocybe.

C. Toxins: Central nervous system. onset of symptoms 20 min. to 2 hours.

Group 5: Ibotenic acid muscimol (delirium) Genera: Amanita. Group 6: Psilocybin psilocin (hallucinogenic) Genera: Psilocybe, Panaeoius.

D. Toxins: Gastro-intestinal irritation onset of symptoms 30 min. to 3 hours. Group 7: Gastro-intestinal irritation. Genera: Boletus, Hypholoma.

Bioluminescence: emit light of bluish green, green in dark. Fox-fire. Enzyme Luciferase. Pleurotus spp. Clitocybe illudens

Armillaria mellea. Mycorrhizae: Increase the solubility of minerals. Increased uptake of N, P & K. protect roots from the pathogens. obtain carbohydrate from host. Ecto, Endo and Ectendomycorrhizal form. Boletus, Cussula, Cortinarius, Tricholoma, Armillaria mellea and Amanita. These are mostly Ectomycorrhizal form.

General Characters: Somatic structures: Mycelium Basidiomycetous. Primary mycelium from homokaryotic basidiospore, short duration. Secondary mycelium interaction of compatible, homokaryotic mycelium dikaryotic hyphae may or may not have clamp connections. responsible for basidiocarp production eg. Marasmium oreades survive for 400 years (Butler & Jones, 1949).

Tertiary mycelium Fruiting bodies Rhizomorph parallel unbranched septate hyphae Armillaria mellea. Fairy rings: Secondary mycelium: grow outwards in all directions. Sporulation Basidiocarp produced periphery Fairy ring. Superstition path of dancing fairies. Three types: no effect on vegetation Lepiota morganii. increased vegetative growth Lepsta personatum

vegetation damaged Agaricus Marasmius oreodus.

Asexual Reproduction: Oidia thin walled spores (mycelial fragments) Wet type: sticky masses on oidiophores Coprinus cinereus Dry type: chains of cylindrical arthroconidia C. micaceous C. truncicola. Function: on germination mycelium. behave as spermatia. chlamydospores : give rise to mycelium. Coprinus lagopus, V. volvacea.

Sexual Reproduction: Majority Heterothallism Compatible thalli by - Hyphal fusion - Oidia. Dikaryotic mycelium formation of basidiocarp. Karyogamy and Meiosis Basidium. Basidiospore haploid, uninucleate.

Basidiocarp: Fruiting body Sporocarp, basidiocarp or basidioma A stalk and a pileus. Pileus: Conical to Flattern and red, yellow, white Gills: closely or distantly placed. Annulus: skirt like ring on stipe. Volva: Cup like membrane at the base of stipe.

Basidiocarp development: Gymnocarpus development Pseudoangiocarpous development Hemiangiocarpous development. Position of hymenium and presence or absence of protective tissues.

Hemiangiocarpous: Early stage hymenium always enclosed by tissues of basidiocarp A membrane connect pileus and stipe Inner veil. Veil severed from pileus and attached to stipe when spore mature and discharge Annulus. Veil tear and hangdown from cap Cortina. Amanita entire primordium universal veil. Sporophore enlarge universal veil tears volva (cup shaped body). remaining universal veil on pileus Scales. Gymnocarpous: Hymenium naked and never covered.

Pseudoangiocarpous: Hymenium later enclosed by incurving margin of pileus and sometimes by an outgrowth of the stipe. No vestigial structures left out in Gymnocarpous and Pseudoangiocarpous development. Anatomy of Basidiocarp: Pseudoparenchymatous tissue. A.campestris stipe are widely inflated. A.cinereus stipe are narrow thread like cell. Lactarius Laticiferous hyphae latex. Hymenium: beneath the pileus, closely packed basidai, cystidia, basidioles and cystidioles, palisade like layer, line inside of tube like structure Boletus. Line gills hang below pileus.

Gills: thin strips of tissues radiate from pileus to stalk. Aequi- hymeniferous: Wedge shaped Basidia shed spores evenly over gills. Inaequi hymeniferous: Coprinus type gill. Sides are parallel. Basidia shed spores upward. Autodeliquescence zone destroy parts of gill and spores discharged. Some species gills free from stalk. Adnate attach directly to stalk. Decurrent attach to stalk and rundown for a distance.

Trama: Central longitudinal running hyphae in gills Lactarius and Russula swollen cells spaerocysts Spongy texture of flesh. Subhymenium: Distinct layer of short cells on the sides of trama. Basidiospores: Basidia bear 4 spores. Forcibly discharged and fall below pileus gravity. Deposited in mass spore print. Taxonomically important character. Colour, shape, size and surface features (ornamentation) taxonomically important.

Classification: Order: Agaricales 16 families (Smith, 1973). 1. 2. 3. 4. Boletaceae Hygrophoraceae Tricholomataceae Entolomataceae 9. Bolbitaceae 10. Strophariaceae 11. Coprinaceae 12. Cortinariaceae

5. 6. 7. 8.

Amanitaceae Pleutoceae Lepiotaceae Agaricaceae

13. Paxillaceae 14. Gomphidiaceae 15. Russulaceae 16. Cantharellaceae

1.

Boletaceae: (Pore, M.R.) Hymenium is poroid, possess vertically arranged tubes, no gills, tube layer easily

ndetachable and soft, fleshy separate from Polyporaceae. Most spp. mycorrhizal on conifers and woody plants. 13 genera spore colour, shape, presence or absence of cystidia. Flesh oxidation of anthraquinone pigment. Blue colour (boletoquinone) by laccase. Genus: Boletus Basidiocarp conspicuous, fleshy, separable tubes, some are edible. B. mirabilis B. edulis fungi sulli. B.parasiticus grow on sporophores of Scleroderma. Family: Agaricaceae Basidiospore black or brown colour, pink or rose coloured free gills on pileus. Annulus present on the stipe.

Genera: Agaricus, Cystoagaricus, Melanophyllum. Genus: Agaricus Well known table delicacy. Deep purplish brown free gills and annulus, no volva. Stalk readily separates from pileus. All are edible except A. placomyces and A. silvaticus. A.campestris field, wild type, edible mushroom, basidia 4 spored. A.brunnescens cultivated mushroom, basidia 2 spored.

Mushroom cultivation: Spawn production pure culture of mycelium Spawning on substrate spawn running environmental condition. Casing 10 days after spawning conversion of vegetative to reproductive. Harvesting 6 to 8 weeks after spawning.

Parasites: Mycogone perniciosa Verticillium lamellicola. Clitocybe dealbata. Life Cycle of Agaricus campestris Fruiting body (Tertiary mycelium) Basidia Basidiospores Somatogamy Secondary mycelium Fruiting body (Tertiary mycelium) & Secondary mycelium

Mushrooms, toad stools, puff balls stinknown rust fruits bracket and jelly funger a special structure called basidium are produced. Within the baridium sucleare fusion takens place and merosis occur but four basidio spores are produced exgenously on an exterion called staimat where as in asco mycotina the qsiospores are formed edogeously. The baridia bearisy structure vary from a mixture parcels like rent to large perennial flattened structure in Ganoderma lam connection, dolipne septa and double layered wall are also important.

No specialized sex organs one produced except in the rents and the sexual feuds hematology is performed by less specilized structures usually the Semitic hepatitis. Formation of clamp connection somatic structures: Typical hypture are mycelia but some are yeast. There are three boards of mycelia the primary secondary and tertiary always found in this sequence in those member producing based carp on gernibration of basides spores giver rise to primary mycelium which in camellia short duration and limited extent. Where well developed primary mycelium in branched septet and haploid in nature with uninucleate sometimes mulinucleate cells. It is usually menokaryotic became of the predominant occurrence of the Helesothallis species among Basideomycolina. Soon the primary mycelium under go dikaryotization either by spermatozoon on somatogamy regularly in the production of secondary or dekayotic mycelium that occurs commonly in nature and is often perennial. The secondary mycelium does not give rise to directly basideocays but under goes certain morphogenetic changes and give rise to territory mycelium which taken ponts iin theconstruction of basidiocarp. In the baridiam the presence of dekampotic mycelium and the formahon of clamp connections are typical characteristic of dasideo mycoltina. The baridia are formed on definate nymenial layer mixed with sterile structure called cystidia The barium originates originates is a terminal help of a brodunuloate mycelium and in separates from it by a septum. The two munchkin the young basidium fare and undergoes reduction division giving rise to four helped nuclei for the mean time. Four stemmata are pushed out at the tap and thai tip enlonga forming basidiospores. The four nucleic now sequate through strenmetal passage and bassidiospores develops as uninucleate haploid basided spores. Clamp connections Except results a specilized structure known as plunges connection is formed on the secondary mycelium It is a typical out growth which makes a connections between the resulting two cells by fusion with the lower at cell division. Not all basies mycetes produce clamp connection but almost all mycelia which posses clamp connection are basideomycelie the clamp connection appears to sieve as a by pass mechanism where by dikayotic conditions can be maintained in successive daughter cells

Banidis mycolina Basidio camp lacking and replaced by tell spores or chlamydospores grouped in sori or slather within the host time parasitic on vaxalac plants Basidiocays usually well developed Besides typically organized us a hymenium sapropic and rarely parasitic. Basidiocay typically gymnocarpoar or semi angiocarpous Besides phragmo basidia or holo baridia barideo spores ballistospares (Violetiacally Hymeno micelles cosigned) Metabasidium in divided by primary septa usually vertically. Basided camps typically angiocarpous Basideo spores not ballisto spores Gastero mycelei statismospores Positive.

Asexual : Ocean commonly but camper to lower fungi is in less frequent It occur by budding fragmentation of mycelium and by cenidra altrospores and Vida. The

underscores act as consider very often the hyphen get fragmented into enucleate such on called arto conidra or arthrospore without bounding up or forming thick walls. Ordia who produced on primary mycelium.

Sexual: Sex organ are generally not found in the Basidiomycotina except in rent. But plasmogermy, Kanyogamy and meisis does taken place. Mostly sematogamy copulation between vegetative mycelium. Gametangial contact, copulation are reduced and in replaced by spermatization of a receptive hypha (Rust,)

Hasidim: It in the organ in which karyogermy and meiosis take places Stargemat : Protobasidicum Karyogermy taken plant Metabasidium Development stage where methods takes Stergmata extermion of metabands.

Helopasidium The Metabasidium in pasidium in which the Metabasidium is not divided by primary septa.

Phragmopasidium: Metabasidium in decided by primary septa, usually vertical Tehobasideaum consist of generally thick walled retting spores and promycelium are germinated Basidiospores Ballisospore virology discharged spores except Gasteromycele and Utilizaginater

Passively discharged based spores of Gassteromycele are statismospores. Tehomycetes Rust and fruits 200 genera 7600 species production of thick walled resting spore, the teliospore (or chlamydospora) in which karyogamy takes place. The tekospora are pant of bestial apparatus and also called tehopasidium. Tehomycete

Uredinaler

Usilaginale

The tako spores in terminal And germinate to give Promycelium pearing Basideospores on sterigmata From which they are Discharged forcibly.

The teko spores are * 8ml ) are usually interracially are baridiospores are produced either terminal or lateral but are sessile and rot discharged forcibly.

Uredinales Rent 6000 species 150 genera

Both manicotti, divot, farms, conifers Chemicals; 1. Mycelium generally intercellular and from heurstinia but back of clamp connection and doliporesepta. 2. Teliospores are produced terminally. 3. Four typical baridiospore (Spridia) formed and discharged forcibly.

4. Highly pleomosphic and complied life cycle(Media telia acexial, paridial, spermatia) autogous, heleroecious(Other heleroecious fungi coelomomyces psorophorae

(Blastocladiales) 5. Rent obligate But (Actinic cultures)

Gymnosporangium funnier Virginian p.graminis tritici, p.helianthi, Melampsora lini uromyces dianthi cronartium fusiforme c.ribicola has been reported. Stage O spermatic monokauptic (pycnia) produce, spermogenia orpycnia. These are produced from haploid primary mycelium after germination of baridiospores. The stage O called because in (1927) the function of the spores produced in spermagonium was not known.

Aeciospores: Plasmogamospores aecidiospores are unicellular produced in chaint then walled and verrucose germinate to give dukaytic mycelium. Macro cylic : all the five spores. Undeiospores: Repeating ehkayotic may produce uredia or telia But in Hemileia vastatrix inedispores produces basidea on germination is called telabal wredinispores (Rajendran91967) In some genera uredionophis and Hyalospora and few puccisia produces resting unedenospores called amphispores (Thickea danker and having dormancy tebospores testing spore on germination promycelium in which depoid sucuas moves and divides meitically. Bandiospores (Sporidea) haploid, uncellular short livied produced exogenoursly after meoisis. Mucrocycli- All five stage long cycle p.q.t. P.hehanthi p.pernesethi. Demi cyclic: Rent which lack ordinal state (spermagenia may be present) cedar apple sent) G. juniperi viriginiance Heleroecious demycyclic Gymno speragium funfifears gynoconis peckiana orgage rust of autoecious demilgibe rent. Microcylitic Teho spores (with or without) spermagenia in the only binuleate spores produced p.malveceareem p.hetero spora.

GASTEROMYCETES Subdivision Class Order Podaxales Family : Nictulariaceae, Geastraceae, Lycoperdaceae, Nidulariaceae, : : : Basidiomycotina Gasteromycetes Nictulariales, Lycoperdales, Nidulariales, Sclerodermatales,

Scleroderma, Podaxaceae, Sphaerobolaceae, Sclerodermataceae Genus : Cyathus, Geastrum, Lycoperdon, Nidula, Nidularia, Pisolithus,

Podaxis, Scleroderma, Sphaerobolus, Trichaster Characters Basidiospores are not discharged violently from their basidia Basidiospores are usually symmetrically poised on their sterigmata

Dring (1973) has termed the b.spores of Gastero statismospores. In Lycoperdon, the fruit-body opens by a pore thro which the spores escape. In Phallus, the spores are exposed in a sticky mass attractive to insects. All members of the gp are saprophytic and grow on soil, roting wood, dung. Rhizopogon and scleroderma are aquatic gastero Totally there are 9 orders, of which only 4 are, saprophytic and grow on soil, roting wood, dung.

LYCOPERDALES Common eg. of this gp are puff-balls Lycoperdon, Calvatia, Bovista, Geastnum. Their FB may begin devt. beneath the surface of soil, but mature above ground. Lycoperdon

FB are pear-shaped, most spp. grow on the ground but L. pyriforme occurs on old stumps, rotting wood and sawdust heaps. Clamp connecting are absent (Dowding and Bulmer, 1964) The basidia bear 1-4 basidiospores symmetrically arranged on sterigmata. Young basidia are binucleate

Calvatia C. gigantean forms FB about the size of a rugby foot balls.

Geastrum Produce FB that begin devt beneath soil or litter. G. triplex grows amongst the leaves of beach, pine. Young FB is onion-shaped. Basidia are pear-shaped, with 4-6 spores borne on a knob-like extension of the pointed end.

NIDULARIALES FB are globose or funnel shaped Eg. : Cyathus, Crucibulum, sphaerobolus First 2 are termed as birds nest fungi Cyathus. Funnal shaped FB in C. olla. Basidia form 4-8 basidiospores Basidia disappear soon after the formation of spores Nuclear fusion and meiosis occur in the basidia Both Cyathus and Crucibulum show tetrapolar heterothallism sphaerobolus S. stellatus forms globose orange FB about 2mm is dm attached to rotten wood, old dung of cow and sheep.

Usually heterothallic Most basidiospores germinate to give mycelia with simple septa.

SCLERODERMATALES Scleroderma 2 common spp. of earth-ball are S. aurantium and S. vernicosum Found in autumn in acid woodland and heaths growing with such trees as Pinus, Betula, Quercus Phallales Common Eg: Phallus impudicus Stinkhorn Mutinus caninus dogs stinkhorn

Phallus Can be detected readily by their smell that are attractive to flies. The smell consists of H2S, phenyl CH3CHO, CH3CHO, CH3COOH These have good vegetative growth on a wide range of carbohydrates and require thiamine. Mutinus is similar to Phallus, but the FB are smaller. The upper part of the stipe is orange in colour and the smell is less obvious.

Classification of Fungi according to Hawkswoth et al. (1983), von Arx (1981), Moore (1980), Moore (1980), Alexopoulos and mins 91979), Ainsworth (1973), Kreisel (1969), Gaumann (1964), Alexopoulos (1962), Martin (1961) and bessey (1950)
Hawksworth et al. (1983) Kingdom Division Von Arx (1981) FUNGI Myxomycota Kingdom MYCOTA Myxomycota Moore (1980) Kingdom FUNGI FUNGI INFERIOR Alexopoulos and mims (1979) Kingdom MYCETAE Gymnomycota Ainsworth (1973) Kingdom FUNGI Myxomycota

Kreisel (

Kingdom PROTOBIO (Myxomycot excluded fro

Classes

Protosteliomycetes Ceratiomyxomycetes Dictyosteliomycetes Acrasiomycetes Myxomycetes Plasmodiophoromycetes Labyrinthulomycetes

Myxomycetes Acrasiomycetes Plasmodiophoromycet es Labyrinthulomycetes

Moore (1971) included Oomycota, Hyphochytridiomycoata , Chytridiomycota and Zygomycota in 'PHYCOMYCOTERA'

Division sub divn.

Eumycota Mastigomycotina

Class

Chytridiomycetes Hyphochytridiomycetes Oomycetes

Oomycota Oomycetes Hyphochytridiomycete s Chytridiomycota Chytridiomycetes

Acrasiogymnomycotina Acrasiomycetes Plasmodiogymnomycoti na Protosteliomycetes Myxomycetes Ceratiomyxomycetidae Myxogastromycetidae Stemonitomycetidae Mastigomycota Haplomastigomycotina

Acrasiomycetes Hbydromyxomycetes Myxomycetes Plasmodiophoromycetes

Sub divn.

Zygomycotina Sygomycetes

Eu-Mycota Zygomycetes Endomycetes

Chytridiomycetes Hyphochytridiomycetes Plasmodiophoromycetes Diplomastigomycotina Oomycetes Amastigomycoa Zygomycotina Zygomycetes

Eumycota Mastigomycotina Chytridiomycetes HyphochytridioChytridiomycetes Mycetes Oomycetes

Eumycota (Chytridiomy excluded fro

(Oomycetes as a class of Chrysophyta Zygomycete

Zygomycotina Zygomycetes Trichomycetes

Endomycete Ascomyceie Euascomyce Loculoascom

Class

Trichomycetes

Ustomycetes Ascomycetes Basidiomycetes Deuteromycetes

FUNGI SUPERIOR (Moore 1971) included Hemiascomycetes and Euascomycetes)

Trichomycetes Ascomycotina Ascomycetes Hemiascomycetidae Plectomycetidae Hymenoascomycetidae Pyrenomycetes Discomycetes Laboulbeniomycetidae Loculoascomycetidae Basidiomycotina Basidiomycetes Teliomycetidae Phragmobasidiomycetid ae Holobasidiomycetidae Hymenomycetes Hymenomycetes Gasteromycetes

Ascomycotina

Sub division

Ascomycotina (No classes recognised)

Hemiascomycetes Plectomycetes Pyrenomycetes Discomycetes Laboulbeniomycetes Loculoascomycetes Basidiomycotina Teliomycetes Hymenomycetes Phragomobasidiomycetidae Holobasidiomycetidae Gasteromycetes

Sub divn. Class

Basidiomycotina Urediniomycetes Ustilaginomycetes Hymenomycetes Gasteromycetes

Sub divn.

Deuteromycotina Hyphomycefes Coelomycetes

BASIDIOMYCOTERA Ustomycota Ustomycetes Sporidiomycetes Basidiomycota Homobasidiomycia Hymenomycetes Gasteromycetes Heterobasidiomycia Holobasidiomycetes Phragmobasi diomycetes Deuteromycotera Blastomycota Ascoblastomycetes Basidioblastomycetes Deuteromycota

Basidiomyco

Pragmobasid dae Hymenobasi dae Gasteromyce

Deuteromycotina Blastomycetidae Hyphomycetidae Coelomycetidae

Deuteromycotina Blastomycetidae Hyphomycetidae Coelomycetidae

Endomyces Imperfecti Ascomycete Imperfecti Basidiomyce Imperfecti

HETEROTHALLISM AND PARASEXUAL CYCLE A.F.Blakeslee, an American Geneticist in 1904 made an important observation with Mucor, which resuited in the discovery of Heterthallism. Blakeslee observed that while some isolates of Mucor formed sporangia as well as zygospores (e.g. M.tenuis), some others failed to form the zygospores and reproduced only by sporangiospores. When he grew these non-sexually - reproducing isolate with other similar isolates, zygospres appeared in the region where the hyphae of the different isolates came in contact with each other. Blakeslee coined the terms homothallism and heterothallism to explain this phenomenon. The homothallic species were those that produced zygospores independently while heterothallic species required the presence of the opposite mating type. M.hiemalis, M.mucedo, Rhizopus nigricans are examples are examples of Heterothallic species. Since the two mating types were morphologically

indistinguishable, Blackslee designated them as ther (+) and (-) mating types or strains (not male or female) Blackslee assumed that the (+) strain represented the female and the (-) strain represented the male sex. We now don't speak of (+) and (-) sexes but only as (+) and (-) strains or mating types.

Pattern of Distribution of Sex Organ in Fungi On the basis of the distribution of sex organs fungi can be put in the following categories A. Hermaphoridite in which both male and female sex organs occur on the same thallus. B. Diccious (sexually dimorphic). The two sex organs are present on different thalli C. Sexually differentiated The male and female sex organs are morphologically similar and therefore indistinguishable. A hermophroditic fungus having both the sex organs may be homothallic or heterothallic. When the two sex organs, present on the same mycelium are unable to mate this is because of self-sterility and is called physiological heterthallism (Whitehouse, 1949). Such fungi need genetically different nuclei, which does not occur when the same thallus forms both the sex organs.

The dioecious fungi in which the male and the female sex organs are borne on different thalli are by necessity heterothallic. This is called Morphological heterothallism (Whitehouse, 1949). In this case heterothallism is made obligatory because the opposite and morphologically distinct sex organs are formed only on different thalli.

The sexually undifferentiated fungi e.g. Mucor, Rhizopus and several members of Asco-and Basidiomycotina, do not have morphologically distinguishable sex organs. These can also be homo-or heterothallic. The heterothallic forms provide another example of physiological heterothalism. The requirement for the other thallus does not lie in morphologically distinct sex organs but in genetically different nuclei, which are not available in the same mycelium.

So, heterothallism according to Whitehouse (1949) can be caused by the absence of the morphological sex organs of opposite type (morphological heterothallism) or by the absence of genetically different nuclei (Physiological heterothallism).

Whatever be the reason of heterothallism the fact remains that different thalli are needed for the sexual reproduction. A heterothallic species may not be only two mating types. There can be four types of thalli and one thallus can mate only with only one of the rest thre. This is called tetrapolar heterothallism.

Bipolar Heterothallism Fungi in this category have two mating types each containing genetically different nuclei. The sexual compatibility is controlled by a pair of genetic factors A and a located at the same locus on different chromosomes. This is therefore also called as ' two allele heterothallism'During meiosis the two chromosomes containing the alleles A and a are separated in the haploid spores (germ spores, ascospores or basidiospores). The spores give rise to two types of thalli which must come together the two nuclei carrying the compatibility factors A and a The two mating type are designated as (+) and (-) strainThe 'two allele' or 'bipolar' heterothallism is found in Mucorales (Mucor, Rhizopus,

Phycomyces). Ascomycotina (Neurospora, Ascobolous). Basidiomycotina (Puccinia graminis and the smut fungus Ustilago levis).

Tetrapolar Heterothallism Fungi in this group form thalli of four mating types. This type of heterothallism is goverened by two pairs of caompatibility factors Aa and Bh, located at different chromosomes, which segregate independently during meiosis. If crossing over occurs between the mating type loci, four types of segregations (4B,Ab,aB,ab) are possible depending on the chromosomal arrangement.

Thus four types of spores (AB, Ab, aB and ab)are formed which give rise to four types of thalli. Only those thalli that have nuclei carrying oposite genes for both the factors, can mate. The resulting zygote must have the genotype Aa, Bb. Majority (630) of the heterothallic Basidiomycotina are tetrapolar forming four types of basidiospores. However if crossing over does not take place only two types of spores (AB and ab or Ab and aB) are formed and only two types of thalli are produced. since it is goverened by two factors it is called tetrapolar. Hormonal or physiological basis for heterothallism In case of zygomycetes, Burgeff at the year of 1924 reported that diffusable substances are responsible for initiation of sexual reproduction. + And - strains are separated by the collodion membrane makes it still attracks by opposite strains.

Plempel et al., 1957 reported that there are some reactions is being taken place while the two compatible strains approaches towards each other. The reactions are as follows,

I.

Teleomorphic reactions: In this reaction aerial zygophores are formed which is in club shaped and yellow in

colour. (-carotene is responsible for the yellow color) II. Zygotrophic reaction Zygophores of + and - strains are grow towards each other.

III. Thigomototrophic reaction In this reaction events are taking place after gametangial fusion and septation.

Gooday (1968) isolated the substances responsible for the sexual reproduction at the point of contact of + and - strains. The isolated substances are generally called as Trisporic acid A, Tripsoric acid B, Tripsoric acid C. In + strains this acid is chemically termed as Methyl dihydrotrisporates whereas in - strains it termed as Triporols.

Heterothalism in Oomycetes In case of Oomycetes if two thalli were contacted together, then it was found to produce four kinds of different thalli. They are as follows,

I.

Pure male thalli

II. Predominantly male thalli and latent stages it produces OOgonia III. Pure female thalli IV. Predominantly female thalli and latent stages it produces antheridia.

Heterothallism in phytopthora sp Phytopthora infestans :Galindo and Galligly reported divided into A1 and A2 compatibility type . They paired and produce abundant oospores in lima bean medium.

In one combination act predominantly as a male when paired with stronger male then act as the female. Gametangial fusion occur between A1 and A2 type.

(ii) Phytothora arecae pethyb reported * N-90 and N-91 isolates when paired does not produce sexual spores. * When both isolates paired with A1 type of P.infestance; they produce abundance oospores but not with A2 type. hence which is designated as A2 type.

(iii) Phytopthora cambivora * N-98 and N99 paired with 7 isolates of P .sp * An abundance of oospores formed when paired with A1 type of P. cinnamomi

(iv) Phytopthora capsici : 5 isolates N14 N58,N59,N313,N100. * N14,N59 and N313,were designated as type A1 because which forms oospores with 60A isolates of P.infestans. * N58, and N100 were A2 type because paired with 60B type.

Phytopthora cinnamomi * Isolates -N2,N33,N66,N67 and N310, N101 to N106, N160, N202, N210, N311, N312. * Hasis and Nelson observed abundance of Oospores in hempseed medium.

(vi) Phytophthora palmivora: 32 isolates were received parings of the first 18 isolates with A1and A2 types of P.infestans on half strength lima bean Agar medium revealed the presence of A1and A2 compatibility types in P.palmivora

Based on oospore production with N241 isolates : * A1 isolates were N136, N137, N146, N240, N242, N243, N248 and N251. * A2 isolates oospores production with N137: N7, N8, N119- N123, N125, N130, N138, N136 141, N143, N144, N244, N245, N247. Phytopthora parasitica var Nicotianae:(Tucker) : 18 isolates were received.

Numerous oospores are formed when isolates N235, N236 and N237 were paired interspecifically with type A1 of P.parasitica,where as small number were produced when N238 was paired with A2 of P.parastitica A1 types : N4, N6, N15- N18 and N238 A2 types : N19- N26, N235, N237.

Heterothallism in Ascomycetes Many of the fungi coming under this sub-division are homothallic and some others are heterothallic. Eg for homothallism. Sordaria macrospora Sordaria fimicola Eg for heterothallism: Neurospora crassa Neurospora sitophila In the process of ascosporogensis, the ascus produced 8 ascospore in which two mating types A, a were found to produced. Eg for Secondary Homothallism in ascomycestes: Neurospora tetrasperma Podospora anserine Gelasinospora tetrasperma Heterothallism in basidiomycotina: 10% of the fungus coming under this sub-division are homothallic. Eg) Coprinus sterquilinus Sexual compatibility is being controlled by pair of genetic factors then it can be termed as tetra polar heterothallism. Eg) Majority of basidiomycotina fungus are tetrapolar.

PARASEXUALITY

Until 1944 the sexual cycle was the only means of exchange of genetic material it is to the credit of microbial geneticists that aseries of novel methods of genetic recombination are now known inbacteria which do not involve karyogamy and meiosis. these aretransformation conjugation transduction, Iysogeny and sexduction which differ from the standard sexual cycle.

A similar alternative to sexual reproduction was discoveredin the imperfect fungus Aspergillus nidulants in 1952 by Pontecorvo and Roper of Glasgow. This they called Parasexual cycle. In this genetic recombination occurs in vegetative cells by the mechanism of mitotic crossing over which brings the same result as is achieved by the meiotic crossing over.

The parasexual cycle involves the following steps 1. Formation of heterokaryotic mycelium 2. Nuclear fusions and multiplication of the diploid nuclei. 3. Mitotic crossing over during divisions of the diploid cells. 4. Sorting out of the diploid strains. 5. Haploidization

Formation of heterokaryotic Mycelium -Heterokaryosis Heterokaryopsis is the main source of variation in the imperfect fungi (deuteromycotina) which lack of sexual reproduction. The term Heterokaryosis was proposed by Hansen and Smith in 1932, who reported it for the first time in Botrytis cineraca.

The presence of genetically different nuclei in an individual is called heterokaryosis and the organism heterokaryon. Essentiallly a heterkaryon possess two sets of chromosomes just like a diploid organism but instead of being contained chromosomes just like a diploid organism but instead of being contained in a single nucleus the two sets of chromosomes lie in separate nuclei sharing the same cytoplasm.heterokaryons show dominance and thus resemble diploids in many respects. Heterokayons show dominance and thus resemble diploids in many respects. Heterokaryosis is a major factor in natural variability and sexuality. The heterokayotic condition can arise in a fungus by three methods viz., (i) Mutation , (ii) Anastomosis ie., fusion between genetically different hyphae and (iii) Diplodization - fusion between haploid nuclei to form diploid nuclei.

Mutations occur frequently in fungi and a homokaryotic mycelium is frequently covereted into heteokaryotic one. Anastomosis between spores and hypahae is a universal feature of higher fungi and certainly must be a potential source of

heterokaryosis and thus of variability. Whether nuclei migrate from one thallus to another is a debated point but the hyphae having nuclei --- parents arise at the point of fusion. Heterokayosis is often accompanied by parasexual cycle.

Nuclear Fusions and Multiplication of the Diploid Nuclei Nuclear fusion invegetaitive heterokaryotic hyphae was first noted by Roper (1952) in Aspergillus nidulans. Nuclear fusion may occur between genetically similar and dissimilar nuclei, resulting in the formation of homozygous and heterozygous diploid nuclei , respectively. Diploid heterzygous nuclei are formed very rarely (at a are present ; 2 types of haploid nuclei their two types of homozygous diploids and the one type of heterozygous diploids.

Mitotic Crossing Over Crossing over is a phenomenon which occurs during meiosis and gives rise to new linkage of genes gene recombination. However initotic crossing over was discovered in 1936 by Stern in Drosophila. a similar mitotic crossing over occurs during the multiplication of the diploid heterozygous nuclei though at a low frequency of 10-2 per nuclear division. However, in some other fungi e.g., Penicillium chrysogenum and Aspergillus niger the frequency of mitotic crossing over is as high as during meiosis in sexual reproduction ; both lack sexual reproduction. Mitotic crossing over is the most important , or key event in parasexual cycle, as it is during this step that genetic recombination occurs.

Sorting out of Diploid strains the segregation of the diploid strains occurs when unimucleate diploid comdia are formed. The colonies that are formed by diploid conidia are recognized by various methods e.g. higher DNA content and bigger size of the conidia and certain phenotype characters of the colony.

Haplodization The diploid colonies show appearance of sectors on the Petriplate, which produce haploid conidia. Thisindicates that some diploid nuclei must have undergone haplodization forming haploid nuclei which later get sorted out in haploid conidi. some of these haploids are genetically different from the original haploid parental nuclei. This is because of the recombination that occurred during the mitotic crossing over. Haploidation occurs at a constant frequency of 10-3 per nuclear division. The haplodization occurs not by a reduction division (meiosis) but by aneuploidy a phenomenon in which chromosomes are lost during mitotic divisions. it happens like this. During mitosisof the diploid nucleus, the chromatids fail to separate (non-disjunction) in the anaphase stage.One daughter nucleus gets one chromosome more (2n+1), while the other gets one chromosome less(2n-1) than the normal 2 sets of chromosomes (2n). Both the daughter nuclei are called aneeuploid. the deficient aneuploid nucleus (2n-1) may lose more chromosomes in the successive mitotic divisions and finally reduced to haploid state (n). Mitotic crossing over and haploidization occur also with the diploid hymozygous nuclei but since the two nuclei are similar, crossing - over products or the haploid nuclei formed by haploidation are genetically no different from the haploid parent nuclei. The parasexual cycle thus like the sexual cycle involvesplasmogamy, karyogamy and haplodization but not at a specified time or place. Every step differs drastically.

A comparison betweeen the sexual and the parasexual cycle Sexual cycle Nuclear fusion occurs in Nuclear vegetative cells Zygote persists for many Parasexual cycle fusion occurs in

specialized structures. Zygote usually persists for one generation only. Recombination occurs by crossing

generations by mitotic divisions. Recombination by mitotic

over during meiosis. The cropping over crossing over which is rare event and occurs in all chromosomes parts and is occurs in a single chromosome arm. accompanied by a reduction in the Haplodizations unlike meiosis is

chromosome number.

independent of crossing over.

Products of meiosis readily recognizable, Recombinant nuclear lies in the vegetative which can be isolatedeasily. cells and can be recognized only by suitable genetic markers. References Text book of fungi, Bacteria and Viruses by - Dubae A Text book of plant pathology by D.P.Tripathi, H.P. Shukla Phytopathology - Volume - 58 (1968) page 1004-1020.

Smut Fungi and Allied Taxa


Class Ustilaginomycetesnow recognized with three subclasses (Entorrhizomycetidae, Ustilaginomycetidae and Exobasidiomycetidae) and nine orders. For our purposes, we will cover two different groups, the smuts (seven orders; we will discuss representatives of four orders) and Exobasidiales, which are plant parasitic fungi but do not share most of the morphological features of smut fungi: Orders Urocystales, Ustilaginales, Entylomatales and Tilletiales (smut fungi). Exobasidiales: Plant pathogenic fungi (biotrophs) united by a) 5S rDNA secondary structure b) sequence analysis of large subunit rDNA, c) host/parasite zone of interaction with primary interactive vesicles, d) septal pores lacking parenthesomes, but with membrane

caps, some with dolipore septa, and e) parasitic dikaryotic phase and saprotrophic haploid phase. Smut Fungi Smut fungi, from the black, dusty masses of teliospores produced by most members of the order Economically important species infecting cultivated crops include Ustilago maydis (corn smut), U. avenae (loose smut of oats), Tilletia controversa (dwarf bunt of wheat), T. tritici (common bunt of wheat), T. indica (Karnal bunt of wheat), Urocystis cepulae (onion smut) Includes 1200 species in 50 genera, with over 4000 species of plant hosts in 75 families of angiosperms. Both monocot- and dicot-infecting smuts are known, but monocot-infecting smuts on cereals are the most economically important. Microbotryon violaceum (anther smut of Caryophyllaceae) and Ustilago maydis are important experimental organisms; Microbotryum, traditionally considered a smut fungus, is now known to belong in the Uredinales. Characterisitics: Heterothallic-unifactorial (bipolor) or bifactorial(tetrapolar) Ustilago maydistwo unlinked mating type loci, a and b, control compatibility. The a locus has two alleles (a1 and a2), encodes pheromones and pheromone receptors, and controls fusion of haploids to form a dikaryon. The b locus, with multiple alleles (25 in U. maydis), regulates differentiation of the dikaryon, filamentous growth, and is considered to be the main pathogenicity determinant (Casselton and Olesnicky, 1998) Putative unifactorial (bipolar) species of Ustilago have been shown to have the a and b loci linked on one chromosome H.H. Flor (gene-for-gene theory in host-pathogen interactions) demonstrated heterothallism in smut fungi (1931-1932) while at WSU early in his career Teliospores (probasidium) that germinate to produce metabasidium (also called a promycelium) and basidiospores (also called sporidia) similar to the rust fungi. Teliospores are produced in sori (sing. = sorus) in the host tissue, often in host reproductive organs (usually ovaries), but also in leaves, stems and roots in some taxa. Life cycle in smut fungi (U. maydis) (from Casselton and Olesnicky, 1998):

Two general types of teliospore germination are distinguished:

Ustilago-type, with transversely septate (=primary septa) metabasidium with basidiospores (on left) and Tilletia-type, with aseptate (or adventitiously septate) metabasidium with terminal whorl of basidiospores (on right) The infectious dikaryon is formed by conjugation of haploid basidiospores or by conjugation of basidium cells:

In Tilletia species, the dikaryon is formed by conjugation of adjacent basidiospores with nuclei of compatible mating type to form a structure called an Hbody because of the shape (see also example of germinating teliospore on previous page:

H-bodies may germinate directly, or indirectly through the formation of secondary basidiospores (secondary sporidia). Two types of secondary basidiospores are formed in Tilletia: lunate basidiospores formed on sterigmata that are forcibly discharged (on left), or filiform basidiospores that are passively dispersed (on right). Both types of secondary basidiospores are also formed from the hyphae:

n the traditional classification of the smut fungi, these two types of germination have provided the basis for delimiting the families Ustilaginaceae and Tilletiaceae. In the modern classification based on ultrastructural and molecular data, the smut fungi are polyphyletic (include taxa from another lineage). Sori are composed of host and fungal tissues. In some genera (Anthracoidea, Neovossia, Sphacelotheca and Tilletia), sori are formed mostly in host ovaries. In other genera (Entyloma, Burrillia, Doassania and Tracya), the sori are formed mostly in the leaves. In Entorrhiza, the sori are formed in galls on host roots. In Ustilago and other genera, the sori are formed in various plant organs in different species, including flowers, stems and leaves. Characters of taxonomic importance relating to the sorus include presence or absence of: thread-like structures (fungal), columella (host), peridium (host or fungus). The consistency of the sorus and whether it is permanently embedded in host tissue rather than becoming exposed at maturity are also important taxonomic characters. Teliospores are formed singly or in spore balls. Approximately half the smut fungi produce spore balls, which may be composed of only teliospores, or teliospores and sterile cells. Sterile cells may also be produced singly and interspersed with teliospores in the sorus. Sterile cells do not germinate. Teliospores are mostly globose, pigmented, and often have sculptured walls. Size ranges from 3.5 to 60 m diameter. Teliospores in mass are usually dark. Teliospores are resistant structures and can survive for 10-25 years in some species under optimal conditions. Host Specialization: Some genera are restricted to a single host family, for example Anthracoidea, Cintractia and Dermatosorus occur only on members of the Cyperaceae, Sphacelotheca occurs on the Polygonaceae, and Sporisorium and Tilletia occur on Poaceae. Species of Doassansia, Entyloma, Thecaphora, Urocystis and Ustilago parasitize plants in a wide range of host families, in some cases both monocot and dicot. However, recent studies have shown in Ustilago that the dicot-infecting species belong to a different genus, Microbotryum, which is now known to be more similar to the rust fungi (Urediniomycetes) than to the smuts. Smut Diseases Categorized based on location of sorus in host tissue: Inflorescence smuts

Leaf smuts Stem smuts Root smuts Infection types: Seedling infectionsystemic infection initiated at young seedling stage, with mostly intercellular dikaryotic hyphae and sporulation (sorus formation) in a host ovary. The mycelium grows with the apical meristem during elongation of the seedling. Examples include common and dwarf bunts of wheat (Tilletia tritici, T. laevis and T. controversa), loose smut of oats (Ustilago avenae), flag smut of wheat (Urocystis agropyri) and sorghum smuts (Sporisorium spp.) Embryo-infectionsystemic infection initiated through the developing embryo; after infection the intercellular mycelium remains dormant in the seed. When the infected seed germinates, the fungus grows with the seedling to infect the initials of the inflorescence. The developing sori replace most of the inflorescence except for the rachis. Examples include loose smut of barley and wheat (Ustilago nuda and U. tritici, respectively). Shoot-infectionsystemic infection resulting from infection through the shoots or young buds. This type of infection may result in the lack of floral development or aborted inflorescences, with the sori formed in the leaves, stems, or part of the aborted inflorescence. Examples include stem or culm smut of grasses (Ustilago hypodytes), sugarcane smut (Ustilago scitaminea) and anther smut of Caryophyllaceae (Microbotryum violaceum). Local infectionmycelium and sporulation are restricted to the region of the plant where infection occurs. The fungus does not become systemic. Examples include corn smut (Ustilago maydis), Karnal bunt of wheat (Tilletia indica) and kernel smut of rice (Tilletia horrida).

Common types of smut diseases: Bunta name given to the ovary-infecting species of Tilletia that infect cereals. The origin of the term is uncertain, but it is believed to come from the burnt appearance of infected cereal inflorescences. Stinking bunt (or stinking smut)the name given to the diseases caused by species of Tilletia because of the fetid (or fishy) odor of the teliospores due to the production of trimethylamine.

Partial buntonly a portion of the seeds on one inflorescence are bunted, and only part of each seed is replaced by the fungal sorus. The infected seed may still germinate if the embryo is not damaged. Occurs in T. indica and T. horrida. Covered smutcharacterized by a well developed, persistent peridium surrounding the sorus. The peridium may be of host or fungal origin. Loose smutcharacterized by a thin, delicate peridium that ruptures easily to expose the dusty mass of teliospores. The peridium is usually of fungal origin. Representative Genera: TilletiaSori usually in reproductive organs of host; teliospores formed singly, usually pigmented with sculptured walls (often reticulate or verrucose); sterile cells present in sorus, germination of the Tilletia-type. EntylomaSori in vegetative organs of host; teliospores formed singly, permanently embedded in host tissue, pale with smooth walls. Germination of the Tilletia-type. UrocystisSori mostly in leaves in stems, forming streaks, swellings or galls. Spore balls comprising several smooth-walled, pigmented teliospores surrounded by hyaline, smooth-walled sterile cells. Germination of the Tilletia-type. UstilagoSori in reproductive organs or vegetative tissues of host; teliospores formed singly, usually pigmented with sculptured walls (often verrucose or echinulate); sterile cells absent, germination of the Ustilago-type. SporisoriumSori in host inflorescence, with central columella of host tissue, teliospores in loose spore balls, sterile cells present. Germination of Ustilago-type.

AnthracoideaSori in host ovaries, teliospores formed singly but agglutinated, dark brown, usually ornamented (spines or warts). Germination of the Ustilago-type but with a 2-celled metabasidium. ThecaphoraSori in various parts of the host, mostly reproductive organs. Spore balls comprising several to many, pigmented, wedge-shaped teliospores permanently united. Teliospores with sculptured outer walls but smooth lateral walls. Germination by formation of a septate metabasidium; each cell developing a lateral hyphae that fuse with each other or a cell of the metabasidium. Order Exobasidiales This order contains plant parasitic fungi that are dimorphic and form holobasidia. They do not form a teliospore. In most of the species, the basidiospores

become septate during germination. Hosts include monocots and dicots. The basidia form predominantly on leaves. Four families are recognized in this order, we will discuss two, the Exobasidiaceae and Graphiolaceae. The Exobasidiaceae produce holobasidia that mostly emerge through stomates. The holobasidia are unique in producing ballistosporic basidiospores oriented abaxially to the apex of the basidium, a feature that is also seen in at least one member of the Tilletiales. The genus Exobasidium is the best known member of this family. Exobasidium species primarily infect hosts in the Ericaceae. Infection often results in distortion of host leaves (hypertrophy and hyperplasia). Exobasidium oxycocci causes a disease of cultivated cranberry (Vaccinium macrocarpon) called rose bloom because the infected leaves become red and thickened, resembling the flower of a rose. The lower surface of the infected leaves is covered with the white hymenium. Basidiospores have 1-3 septa, and germinate by budding to produce conidia. The Graphiolaceae are parasites of palms. In Graphiola, the cylindrical fruiting bodies occur on the leaves of palms, and contain globose basidia formed in chains. The passively dispersed basidiospores are formed laterally on the basidia. In culture, this fungus grows in a yeast-like manner. References: Bauer, R., D. Begerow, F. Oberwinkler, M. Piepenbring and M. L. Berbee. 2001. Ustilaginomycetes. Chap. 3. In: The Mycota VIII Part B. Systematics and Evolution. McLaughlin, McLaughlin and Lemke, eds. Springer-Verlag, Berlin. Bauer, R., F. Oberwinkler and K. Vanky. 1997. Ultrastructural markers and systematics in smut fungi and allied taxa. Can. J. Bot. 75: 1273-1314. Begerow, D., R. Bauer and F. Oberwinkler. 1997. Phylogenetic studies on nuclear large subunit ribosomal DNA sequences of smut fungi and related taxa. Can. J. Bot. 75: 2045-2056. Casselton, L. A. and N. S. Olesnicky. 1998. Molecular genetics of mating recognition in basidiomycete fungi. Microbiology and Molecular Biology Reviews 62: 55-70. Duran, R. and G. W. Fischer. 1961. The Genus Tilletia. Washington State University. Fischer, G. W. 1953. Manual of the North American Smut Fungi. The Ronald Press, New York. Fischer, G. W. and C. S. Holton. 1957. Biology and Control of the Smut Fungi. The Ronald Press, New York.

Vanky, K. 1987. Illlustrated Genera of Smut Fungi. Gustav Fischer Verlag. New York. Vanky, K. 1994. European Smut Fungi. Gustav Fischer Verlag. New York.

The classification of dimorphic basidiomycetes J.P. Sampaio and R. Bauer

Introduction Class Ustilaginomycetes


Introduction The current tripartite classification concept of the Basidiomycota is mainly based on sequence analyses of 18S rRNA (Taylor, 1995; Swann and Taylor, 1993; 1995a; 1995b; 1995c). As a consequence of those studies, the classes Urediniomycetes, Ustilaginomycetes and Hymenomycetes have been proposed (Swann and Taylor 1995a). Analysis of the D1/D2 region of the 26S rDNA shows basically the same pattern (Begerow et al., 1997). The data derived fromthe analysis of cell wall carbohydrates correlate well with the molecular phylogenies (Prillinger et al., 1990; 1991a; 1991b; 1993) and the same holds for the structure of the septal pores and the shape of the spindle pole bodies (McLaughlin et al., 1995a; 1995b and references therein).

Class Ustilaginomycetes
The Ustilaginomycetes are predominantly plant-parasites and include ca. 1400 species. They can be distinguished from the other basidiomycetes because they have a distinctive cell wall carbohydrate composition with dominance of glucose and absence of xylose

(Prillinger et al.,1990; 1993). The type B secondary structure of the 5S rRNA is shared with the Hymenomycetes (Gottschalk and Blanz, 1985) and, like the Urediniomycetes, they lack parenthesomes at the septal pores (Bauer et al., 1997). Several recent publications have dealt with the classification of the Ustilaginomycetes (Bauer et al., 1997; Bauer et al., 2001; Begerow et al., 1997) and a comprehensive classification scheme based mainly on the characteristics of host-parasite interactions and septal pore apparatus has been proposed. In general, sequence data gives support to the classification proposals listed below, derived mainly from Bauer et al. (2001) and Begerow et al. (2002). The supra-generic classification of mitosporic taxa belonging to the Ustilaginomycetes is also being improved. The order Malasseziales Moore emend. Begerow, Bauer & Boekhout was redefined and presently accommodates the species of Malassezia Baillon, a genus of the Exobasidiomycetidae (Begerow et al., 1997). The genus Pseudozyma Bandoni emend. Boekhout seems to represent the asexual counterpart of Ustilago (Persoon) Roussel, whereas Tilletiopsis Derx ex Derx belongs to the Exobasidiomycetidae but is polyphyletic (Begerow et al., 2000). Sympodiomycopsis paphiopedili Sugiyama, Tokuoka & Komagata and three species of Rhodotorula, namely Rh. bacarum (Buhagiar) Rodrigues de Miranda & Weijman, Rh. hinnulea (Shivas & Rodrigues de Miranda) Rodrigues de Miranda & Weijman and Rh. phylloplana (Shivas & Rodrigues de Miranda) Rodrigues de Miranda & Weijman belong to the order Microstromatales of the Exobasidiomycetidae (Fell et al., 2001; Begerow et al., 2001). The classification system of the Ustilaginomycetes is presented below.

CLASS USTILAGINOMYCETES Bauer, Oberwinkler & Vnky SUB-CLASS ENTORRHIZOMYCETIDAE Bauer & Oberwinkler
Members of the Ustilaginomycetes having local interaction zones and pores without membranous bands or caps (Bauer et al., 1997).

Order Entorrhizales Bauer & Oberwinkler Family Entorrhizaceae Bauer & Oberwinkler Entorrhiza Weber

SUB-CLASS USTILAGINOMYCETIDAE Jlich emend. Bauer & Oberwinkler


Members of the Ustilaginomycetes having enlarged interaction zones. Septa have pores with membranous caps or they are poreless at maturity.

Order Urocystales Bauer & Oberwinkler Family Melanotaeniaceae Begerow, Bauer & Oberwinkler Exoteliospora Bauer, Oberwinkler & Vnky Melanotaenium de Bary Yelsemia Walker

Family Doassansiopsaceae Begerow, Bauer & Oberwinkler Doassansiopsis (Setchell) Dietel

Family Urocystaceae Begerow, Bauer & Oberwinkler

Mundkurella Thirumalachar Urocystis Rabenhorst ex Fuckel Ustacystis Zundel Vankya Ershad

Order Ustilaginales Clinton emend. Bauer & Oberwinkler Family Ustilaginaceae L. Tulasne & C. Tulasne emend. Bauer & Oberwinkler Anthracoidea Brefeld Cintractia Cornu Cintractiella K. B. Boedijn Clintamra Cordas & Durn Dermatosorus Sawada ex Ling Farysia Raciborski Farysporium Vnky Franzpetrakia Thirumalachar & Pavgi emend. Guo, Vnky & Mordue Geminago Vnky et Bauer Heterotolyposporium Vnky Kuntzeomyces Hennings ex Saccardo & Sydow Leucocintractia Piepenbring, Begerow & Oberwinkler

Macalpinomyces Langdon & Fullerton emend. Vnky Melanopsichium Beck Moesziomyces Vnky Moreaua Liou & Cheng Orphanomyces Saville Pericladium Passerini emend. Mundkur Planetella Saville Pseudozyma Bandoni emend. Boekhout
Mitosporic

"Rhodotorula" acheniorum (Buhagiar & Barnett) Rodrigues de Miranda


Mitosporic

Schizonella Schrter Sporisorium Ehrenberg. ex Link Stegocintractia Piepenbring, Begerow & Oberwinkler Testicularia Klotzsch Tolyposporium Woronin ex Schrter Tranzscheliella Lavrov Trichocintractia Piepenbring, Begerow & Oberwinkler Uleiella Schrter

Ustanciosporium Vnky emend. Piepenbring Ustilago (Persoon) Roussel Websdanea Vnky

Family Glomosporiaceae Cifferi emend. Begerow, Bauer & Oberwinkler Glomosporium Kochman Thecaphora Fingerhuth emend. Vnky Tothiella Vnky

Family Mycosyringaceae Bauer & Oberwinkler Mycosyrinx Beck

SUB-CLASS EXOBASIDIOMYCETIDAE Jlich emend. Bauer & Oberwinkler


Members of the Ustilaginomycetes having local interaction zones. Septa have pores with membranous bands or caps or they are poreless at maturity.

Order Malasseziales Moore emend. Begerow, Bauer & Boekhout Malassezia Baillon

Order Georgefischeriales Bauer, Begerow & Oberwinkler

Family Georgefischeriaceae Bauer, Begerow & Oberwinkler Georgefischeria Thirumalachar & Narasimhan emend Gandhe Jamesdicksonia Thirumalachar, Pavgi & Payak

Family Tilletiariaceae Moore Phragmotaenium Bauer, Begerow, Nagler & Oberwinkler Tilletiaria Bandoni & Johri Tilletiopsis flava (Tubaki) Boekhout
Mitosporic

Tilletiopsis fulvescens Gokhale


Mitosporic

Tolyposporella Atkinson

Family Eballistraceae (Bauer, Begerow, Nagler & Oberwinkler) Eballistra Bauer, Begerow, Nagler & Oberwinkler

Order Tilletiales Kreisel ex Bauer & Oberwinkler Family Tilletiaceae L & C Tullasne emend. Bauer et Oberwinker Conidiosporomyces Vnky

Erratomyces Piepenbring & Bauer Ingoldiomyces Vnky Neovossia Krnicke Oberwinkleria Vnky & Bauer Tilletia L. & C Tullasne

Order Microstromatales Bauer & Oberwinkler Family Microstromataceae Jlich "Cerinosterus" cyanescens (de Hoog & de Vries) Moore Microstroma Niessl "Rhodotorula" bacarum (Buhagiar) Rodrigues de Miranda & Weijman
Mitosporic

"Rhodotorula" phylloplana (Shivas & Rodrigues de Miranda) Rodrigues de Miranda & Weijman
Mitosporic

Sympodiomycopsis Sugiyama, Tokuoka & Komagata


Mitosporic

Family Volvocisporaceae Begerow, Bauer & Oberwinkler Volvocisporium Begerow, Bauer & Oberwinkler

Superorder Exobasidianae Bauer & Oberwinkler


Members of the Exobasididiomycetidae having simple pores and interaction apparatus

Order Entylomatales Bauer & Oberwinkler Family Entylomataceae Bauer & Oberwinkler Entyloma de Bary

Order Doassansiales Bauer & Oberwinkler Family Melaniellaceae Bauer, Vnky, Begerow & Oberwinkler Mellaniella Bauer, Vnky, Begerow & Oberwinkler

Family Doassansiaceae (Azb. & Karat.) Moore emend. Bauer & Oberwinkler Burillia Setchell Doassansia Cornu Doassinga Vnky, Bauer & Begerow Heterodoassansia Vnky Nannfeldtiomyces Vnky

Narasimhania Thirumalachar et Pavgi emend. Vnky Pseudodermatosorus Vnky Pseudodoassansia (Setchell) Vnky Pseudotracya Vnky Tracya H. & P. Sydow

Family Rhamphosporaceae Bauer & Oberwinkler Rhamphospora Cunningham

Order Exobasidiales Hennings emend. Bauer & Oberwinkler Family Brachybasidiaceae Gumann Brachybasidium Gumann Dicellomyces Olive Exobasidiellum Donk Kordyana Racib. Proliferobasidium Cunningham

Family Exobasidiaceae Hennings Exobasidium Woronin

Muribasidiospora Kamat e & Rajendren

Family Cryptobasidiaceae Malenon ex Donk Botryoconis H. & P. Sydow Clinoconidium Pat. Coniodictyum Hariot & Pat. Drepanoconis Schrter & P. Hennings Laurobasidium Jlich

Family Graphiolaceae Fischer Graphiola Poiteau Stylina H. Sydow

Taxa of the Exobasidiomycetidae not ascribed to any family Ceraceosorus bombacis (Bakshi) Bakshi

Tilletiopsis albescens Gokhale


Mitosporic

Tilletiopsis cremea Tubaki


Mitosporic

Tilletiopsis lilacina Tubaki


Mitosporic

Tilletiopsis minor Nyland


Mitosporic

Tilletiopsis pallescens Gokhale


Mitosporic

Tilletiopsis washingtonensis Nyland


Mitosporic

Class Hymenomycetes (Tremellomycetidae)


The class Hymenomycetes as defined by Swann and Taylor (1995a; 1995b; 1995c) includes the complex-septate basidiomycetes (i.e. with dolipore), including the gasteromycetes. Moreover, the members of the Hymenomycetes have glucose as the major cell wall carbohydrate component and, at variance with the Urediniomycetes and Ustilaginomycetes, xylose is present. Swann and Taylor (1995b) distinguished two subclasses, viz. Hymenomycetidae and Tremellomycetidae. A few traits other than DNA sequences differentiate the two subclasses. Sacculate membrane elements are normally observed at the vicinity of the dolipore of the Tremellomycetidae (in some cases they are lacking), whereas the members of the Hymenomycetidae have smooth membranous septal pore caps of various types, but not sacculate. In addition, whereas the vast majority of the Tremellomycetidae produces a yeast phase, such ontogenetic stage is not observed

in the Hymenomycetidae. Therefore, the present review will deal only with the Tremellomycetidae. The classification system presented below is based mainly on the revisions of Wells (1994) and Wells and Bandoni (2001), which are derived mostly from comparisons of morphological and ultrastructural data. New teleomorphic genera described in Kirschner et al. (2001) and Sampaio et al. (2002) are also included. Although most mitosporic species of the Tremellomycetidae have been investigated in comprehensive sequence analysis studies (Fell et al., 2000; 2001; Scorzetii et al., 2002), several sexual taxa have not yet been studied using the same approaches. It is anticipated that relevant systematic changes will be proposed when such species are investigated by molecular phylogenetic methods.

References
Bauer, R. and Oberwinkler, F. 1991a. The colacosomes: new structures at the host-parasite interface of a mycoparasitic basidiomycete. Botanica Acta 104: 53-57. Bauer, R. and Oberwinkler, F. 1991b. The symplechosome: a unique cell organelle of some basidiomycetes. Botanica Acta 104: 93-97. Bauer, R. and Oberwinkler, F. 1994. Meiosis, septal pore architecture and systematic position of the heterobasidiomycetous fern parasite Herpobasidium filicinum. Canadian Journal of Botany 72: 1229-1242. Bauer, R., Oberwinkler, F. and Vnky, K. 1997. Ultrastructural markers and systematics in smut fungi and allied taxa. Canadian Journal of Botany. 75: 1273-1314. Bauer, R., Begerow, D, Oberwinkler, F. Piepenbring, M. and Berbee, ML. 2001. Ustilaginomycetes In The Mycota vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 58-83. Bauer, R., Begerow, D., Oberwinkler, F. and Marvanov, L. 2003. Classicula: the teleomorph of Naiadella fluitans. Mycologia (in press). Begerow, D., Bauer, R. and Oberwinkler, F. 1997. Phylogenetic studies on nuclear large subunit ribosomal DNA sequences of smut fungi and related taxa. Canadian Journal Botany 75: 2045-2056.

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McLaughlin, D.J. 1990. A new cytoplasmic structure in the basidiomycete Helicogloea: the microscala. Experimental Mycology 14: 331-338. McLaughlin, D.J., Berres, M.E. and Szabo, L.J. 1995a. Molecules and morphology in basidiomycete phylogeny. Canadian Journal of Botany 73: 684-692. McLaughlin, D.J., Frieders, E.M. and L, H. 1995b. A microscopists view of heterobasidiomycete phylogeny. Studies in Mycology 38: 91-109. Oberwinkler, Fand Bauer, R. 1989. The systematics of gasteroid, auricularioid Heterobasidiomycetes. Sydowia 41: 224-256. Prillinger, H., Drfler, Ch., Laaser, G. e Hauska, G. 1990. Ein Beitrag zur Systematik und Entwicklungsbiologie Hherer Pilze. Hefe-Typen der Basidiomyceten. Teil III:Ustilago-Typ. Zeitschrift fr Mykologie 56: 251-278.Prillinger, H., Laaser, G., Drfler, Ch., e Ziegler, K. 1991a. Ein Beitrag zur Systematik und Entwicklungsbiologie Hherer Pilze. Hefe-Typen der Basidiomyceten. Teil IV: Dacrymyces-Typ, Tremella-Typ. Sydowia 53: 170-218. Prillinger, H., Deml, G., Drfler, Ch., Laaser, G. e Lockau, W. 1991b. Ein Beitrag zur Systematik und Entwicklungsbiologie Hherer Pilze. Hefe-Typen der Basidiomyceten. Teil II: Microbotryum-Typ. Botanica Acta 104: 5-17. Prillinger, H., Oberwinkler, F., Umile, C., Tlachac, K., Bauer, R., Drfler, C. e Taufratzhofer, E. 1993. Analysis of cell wall carbohydrates (neutral sugars) from ascomycetous and basidiomycetous yeasts with and without derivatization. Journal of General and Applied Microbiology 39: 1-34. Sampaio, J.P., Bauer, R., Begerow, D. andOberwinkler, F. 1999. Occultifur externus sp. nov., a new species of simple-pored auricularioid heterobasidiomycete from plant litter in Portugal. Mycologia 91: 1094-1101. Sampaio, J.P., Wei, M., Gadanho, M. and Bauer, R. 2002. New taxa in the Tremellales: Bulleribasidium oberjochense gen. et sp. nov., Papiliotrema bandonii gen. et sp. nov. and Fibulobasidium murrhardtense sp. nov. Mycologia 94: 873-887. Sampaio, J.P., Gadanho, M., Bauer, R. and Wei, M. 2003. Taxonomic studies in the Microbotryomycetidae: Leucosporidium golubevii sp. nov., Leucosporidiella gen. nov. and the new orders Leucosporidiales and Sporidiobolales. Mycological Progress (in press).

Scorzetti, G., Fell, J.W., Fonseca, A. and Statzell-Tallman, A. 2002. Systematics of basidiomycetous yeasts: a comparison of large subunit D1/D2 and internal transcribed spacer rDNA regions. FEMS Yeast Research 2: 495-517. Swann, E.C. and Taylor, J.W. 1993. Higher taxa of basidiomycetes: an 18S rRNA gene perspective. Mycologia 85: 923-936. Swann, E.C. and Taylor, J.W. 1995a. Phylogenetic perspectives on basidiomycete systematics: evidence from the 18S rRNA gene. Canadian Journal of Botany 73: 649-1433. Swann, E.C. and Taylor, J.W. 1995b. Phylogenetic diversity of yeast-producing basidiomycetes. Mycological Research 99: 1205-1210. Swann, E.C. and Taylor, J.W. 1995c. Toward a phylogenetic systematics of the Basidiomycota: integrating yeasts and filamentous basidiomycetes using 18S rRNA gene sequences. Studies in Mycology 38: 147-161. Swann, E.C. Frieders, E.M. and McLaughlin, D.J. 1999. Microbotryum, Kriegeria and the changing paradigm in basidiomycete classification. Mycologia 91: 51-66.Swann, E.C. Frieders, E.M. and McLaughlin, D.J. 2001. Urediniomycetes In The Mycota vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 37-56 Takashima, M., Hamamoto, M. and Nakase, T. 2000. Taxonomic significance of fucose in the class Urediniomycetes: distribution of fucose in cell wall and phylogeny of urediniomycetous yeasts. Systematic and Applied Microbiology 23: 63-70. Taylor, J.W. 1995. Molecular phylogenetic classification of fungi. Archives of Medical Research 26: 307314. Wells, K. 1994. Jelly fungi, then and now! Mycologia 86: 18-48. Wells, K. and Bandoni, R.J. 2001. Heterobasidiomycetes. In The Mycota vol VII, Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin and P.A. Lemke (eds.), Springer-Verlag, Berlin, pp 85-120.

HYMENOMYCETES

GENERAL CHARACTERS: Basidiocarp typically gymnocarpous or semiangiocarpous Basidia either Phragmobasidia or Holobasidia Basidiospores are ballistospores Largest group of basidiomycotina fungi Includes toadstools, bracket fungi, fairy clubs, jelly fungi etc., Basidia arranged in a palisade like fashion to form hymenium which is fully exposed at maturity

CLASSIFICATION: Based basidial structure it is classified into 2 types: 1. Holobasidiomycetidae Phragmobasidiomycetidae

2.

HOLOBASIDIOMYCETIDAE: It includes 6 orders (Mc Nabb & Talbot, 1973). 1. 2. 3. 4. 5. 6. Agaricales Aphyllophorales Dacrymycetales Exobasidiales Tulsanellales Brachybasidiales Out of 6 family 2 are considered to be most important 1. 2. Agaricales Aphyllophorales

DIFFERENCE BETWEEN THESE TWO ORDERS: CHARACTERS AGARICALES APHYLLOPHORALES

Fruiting body

Thin walled hyphae which inflate monomitic

Hymenophore Nuclear spindle Basidiocarp AGARICALES: 250 Genera 16 families

Surrounded by 1 or more veil Transverse (Chiastobasidial) Gymnocarpus

Thin walled generative hyphae along with thick walled unbranched skeletal or thick walled much branched binding hyphae or both-mono\di\trimitic Veil not present Longitudinal (Stichobasidial) Gymnocarpus or Angiocarpus

Few are parasitic, majority are saprophytic Includes edible and poisonous mushrooms Mycelium shows bioluminescence, phenomenon called Fox Fire Eg. Omphalotus olerius Eg.Armillaria mellea Some of them produce rhizomorphs eg. Armillaria mellea Few species produces Thin walled oidia (Coprinus lagopus) Chlamydospores (Volvariella volvacea) Conidiospores Asexual reproduction rarely occurs Sexual reproduction by means of hyphal fusion Few are homothallic and majority are heterothallic Hymenium protected by partial veil Thin in Cortinarius spp called cortina Firmer in Agaricus called annulus

FRUIT BODY DEVELOPMENT:


1.GYMNOCARPIC:

Hymenophore naked Never enclosed Pileus develop at the tip of stipe Eg. Boletus subtomentosus
2. ANGIOCARPIC:

enclosed by tissue 2 kinds Primary angiocarpy: pileus margin, the hymenophore and sometimes the pilues and stipe differentiate beneath the surface. Eg. Stropharia semiglobata Secondary angiocarpy: extension of hyphae from pileus margin towards stipe to enclose the previously differentiated hymenophore (gills). Eg. Lentinus tigrinus TWO TYPES OF GILL STRUCTURE: Aequi-hymeniiferous (aequihymenial) Inaequi-hymeniiferous (inaequihymenial) Differences: AEQUI-HYMENIIFEROUS Most agarics Gills are wedge shaped Hymenium develops in equal manner all over the gill surface INAEQUI-HYMENIIFEROUS Characteristics of Coprinus (inkcaps) Not so but are parallel sided Unequal manner with basidia ripening in zones

TERMINOLOGIES: 1. HYMENOPODIUM: In bilateral trama, the layer immediately outside the central

layer . eg.Amanita rubescens 2. SUBHYMENIUM: Ends of tramal hyphae turn outward to form a distinct layer of

shorter cells 3. CYSTIDIOLES: Thin walled, sterile elements of hymenium, protruding slightly

out of hymenial surface

4.

CYSTIDIA: Enlarged conical or cylindrical cells which may arise in the hymenium along with the basidia

5. 6. 7. 8.

PLEUROCYSTIDIA: Cystidia on gill face CHEILOCYSTIDIA: Those on gill margin PILEOCYSTIDIA: Similar structure found on pilues surface CAULOCYSTIDIA: Similar structure found on the stipe

Important Families of the Order Agaricales are as follows : 1. 2. 3. 4. 5. 6. 7. Agaricaceae Boletaceae Russsulaceae Amanitaceae Coprinaceae Hygrophoraceae Tricholomataceae

AGARICACEAE: Stipe and pileus are clearly separated Cuticle of pileus is fibrillose or squamulose Annulus present on stipe Spore deposits are blackish or dark chocholate brown An apical germ pore is absent in the spores Includes edible mushrooms.eg.Agaricus campestris Based on the spore colour it is divided into 5 groups black spores Coprinus purple spores Agaricus brown spores Flammula pink spores Volvaria white spores Pleurotus

BOLETACEAE: Distinguishing character is the presence of tubular hymenophore Fruiting body is fleshy, soft, decay easily and quickly Mycorrhizae pine, birch, larch etc., Most species are edible eg. Boletus edulis Few are posisonous Important genera are Boletus, Boletinus, Strobilomyces GENERIC LEVEL CLASSIFICATION: Based on the following characters they are classified: 1. 2. 3. 4. 5. Spore colour- yellow, pink, brown, purple black, olive Spore shape-elliptical or subfusiform Cystidia- presence or absence Texture of the cap- glutinous, dry or scaly Markings on the stem- punctate, veined-reticulate, scaly.

Important genus are Boletus satanus- beech or oak Boletus parasiticus-parasite on Scleroderma Leccinum versipelle- birch Suillus bovines- pinus Paxillus involutus- birch RUSSULACEAE: It includes 2 important genera Russula and Lactarius. Presence of sphaerocysts is identifiable character. Russula: They are brittle, having short thick stalk and caps are coloured brilliantly. Russula emetica is a poisonous fungi in this group. Lactarius: Identified by milky juice exudates. It includes more than 40 spp. The latex produced may be colored or colorless, creamy, yellow, blue or red. Species differentiation can be done by colour changes and milk exposure.

AMANITACEAE: Presence of white gills and spore Presence of annulus on stem Presence of a volva (as cup) Presence of bilateral hymenophore trama Edible mushrooms: Amanita rubescens Amanita vaginata Amanita fulva Poisonous toadstools: Amanita verna and Amanita virosa Poison principles: They are generally called as amanita toxins which includes , And - amanitin Phalloidin Hallucinogenic substances are also present in some species. Eg. Amanita muscaria (fly agaric) which contains the following: isoxazole derivatives ibotenic acid muscimol COPRINACEAE: Presence of parallel sided gills Presence of Y-shaped gills 2 imporatant genus are Coprinus and Panaellus Coprinus identified by the dimorphic basidia which may be long or short

HYGROPHORACEAE: Commonly seen in pastures Hygrophorus sp. Are bright, viscid and waxy textured.

TRICHOLOMATACEAE: Important genera are: Tricholoma Armillaria Laccaria

Armillaria mellea: Commonly known as honey agaric. Parasitic on woody host. Fruit bodies are edible. Produces rhizomorphs, which can grow upto 1 meter per year. It has antibacterial and antifungal antibiotics, which are effective against gram-positive bacteria and Trichoderma. The major competitors to this fungi are Coriolus versicolor and Phlebia mersimoides Armillaria mellea spread by 3 way: o o o Basidiospores Root contact Boot lace like rhizomorphs

APHYLLOPHORALES: Fruiting body gymnocarpus Presence of flattened hymenophore Club shaped Tooth like Basidiocarp are cottony or papery Sterigmata typically cureved Stichobasidial 22 families 1. 2. 3. 4. 5. 6. 7. 8. Cantharellaceae Coniophoraceae Corticiaceae* Gomphaceae Punctulariaceae Stereaceae* Thelephoraceae Schizophyllaceae 12.Sparassidaceae 13.Auriscalpiaceae 14.Bankeraceae 15.Echinedontiaceaea 16.Heriaceae 17.Hydnaceae 18.Bondarzeloiceae 19.Fistulinaceae

9. 10. 11.

Cyphellaceae Clavariaceae Clavulinaceae

20.Ganodermataceae* 21.Hymenochataceae 22.Polyporaceae*

POLYPORACEAE: Includes 40 genera and 1000 speciesMost of them are wood rotting fungi. They are heterothallic in nature. Fan shaped brackets are produce which are lacking stipes. They are pore-bearing fungi. Hymenium lining interior pores. Hymenium is tubular. Eg. Heterobasidion annosum Heart rot of conifers (perfect stage.Oedocephalum lineatum) Eg.Pitoporus botulinus - Heart rot of birch HYPHAL ANALYSIS: To study the structure of a polypore fruit body the hyphal analysis can be followed. There are 3 kinds of hyphae:

Generative hypha Thin walled Branched Septate With or without clamp

Skeletal hypha Thick walled Sparsely branched Aseptate or psuedoseptate Without clamp

Binding hypha Thick walled Much branched Aseptate Without clamp Weave themselves between other hyphae

Gives rise to basidia or 2 Gives rigid frame work other kinds of hyphae Cyanophilus Unstained

Unstained

Based on the presence of these hyphae: 1. 2. Monomitic: presence of generative hypha alone. Eg.Polyporus adustus Dimitic: Presence of generative and skeletal hypha. Eg. Heterobasidion annosum -Presence of generative and binding hypha. Eg. Polyporus sulphureous 3. Trimitic: presence of all the three kinds of hyphae. Eg. Coriolus versicolor

GANODERMATACEAE: Commonly known as lignicolous fungi since they decay standing timber. It is differentiated from the family polyporaceae by the presence double walled spore with dark inner layer bearing ornamentation, which pierces the hyaline outer one. The spore surface appears as spiny here. Basidiocarp stalked or sessile. Trimitic hyphal construction was observed. GANODERMA: It is one of the important genera Ganoderma applanatum Heart rot of beach (wound parasite) Ganoderma lucidum Tanjore wilt of coconut Sporophores are perennial, brown, woody. Fan shaped brackets are produced which 50 cm to 1m across. Hymenial tube 2 cm length and 1mm diameter Spore germinates by producing germ tube. CORTICIACEAE: It includes 80 genera where some are parasitic and saprophytic. Basidiocarp monomitic and dimitic (skeletal hyphae). Hymenophore are smooth and tuberculate. Basidiospores are non cyanophilous and smooth walled Important genera are Corticium and Peniophora. Corticium Thin flat fruit body lying appressed to substratum. It is made up of thin layer of basidia bearing hyphae. Hymenium fully exposed. Corticium solmonicolor causing pink disease of tea and rubber Corticium rolfsii is a soil inhabiting parasite attacks groundnut and

cotton.(imperfect stage Sclerotium rolfsii) Peniophora : Possess cystidia and gloeocystidia. Presence of flat and waxy basidiocarps

Sterum

Peniophora gigantean used of bio control for the management of Fomes annosus

STEREACEAE: It includes 18 genera and 150 species. Fruit body is flattened, appressed, sessile or stalked. Hymenium smooth Dimitic usually.

They are mostly saprophytic. Some of the important species are Sterum hirsutum- yellowish fan shaped leathery brackets S. gausapatum- fruit body bleed when bruised and it causes pipe rot of oak S.rugosum - broad leaved trees attacked by this fungi and it bleeds S.sanguinolentum- attacks conifers S.purpureum- silver leaf disease DACRYMYCETACEAE: It includes 9 genera and 2 are found to be most important. Viz., Dacrymyces and Calocera

PHRAGMOBASIDIOMYCETIDAE: Basidia in which metabasidium is divided by septa. Basidiospores by repeated germination produce secondary spores. Fruit body of the fungus are gelatinous or waxy (jelly fungi). It includes 3 orders (Mc Nabb, 1973). Tremellales Auriculariales Septobasidiales

TREMELLALES: It has longitudinally divided basidia. The fruiting body of the fungus is

gelatinous and bright when it is wet and it become cartilaginous texture when it is dry. Important genera of this group are Tremella, Exidia and Psuedohydnum.

Tremella : Tremella frondosa attacks the stump of the trees like oak and beech. The fruiting body of this fungus is fleshy, colourless to pale brown in colour. The

basidiospore germinates by repeated germination, which resembles budding in case of yeast. Tremella mesenterica attacks oak, willow and beech trees. It produces yellow to orange lobed fructification. This fungus is heterothallic in nature. Exidia Exidia glandulosa commonly known as witches buffer attacks lime and oak. It produces black rubbery fructification. The hymenium is small black and has warty outgrowth. Exidia nucleata has fine structure. AURICULARIALES: It includes 18 genera (Mc Nabb, 1973). Metabasidium is divided by transverse septa in this order. They are parasitic or saprotrophic on plants or fungi. They produce gelatinous fruiting body. Eg. Helicobasidum brebisonii causes violet root rot. Eg.Auricularia auricula. Auricularia auricula: Since it produces rubbery ear shaped fruiting body it is commonly known as jews ear fungus. Fruit body dry to brittle mass, while wetting it absorbs moisture and discharg spores. Basidia are cylindrical with 3 septate and 4 celled. Each cell has epibasidium terminate in conical sterigmata bearing basidiospore. projected out as ballistospores. This fungus is heterothallic in nature. The spores are