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Cochliobolus carbonum
''Cochliobolus carbonum'' (anamorph: ''Helminthosporium carbonum'') is one of more than 40 species of filamentous ascomycetes belonging to the genus ''Cochliobolus'' (anamorph: ''Bipolaris/Curvularia''). This pathogen has a worldwide distribution, with reports from Australia, Brazil, Cambodia, Canada, China, Congo, Denmark, Egypt, India, Kenya, New Zealand, Nigeria, Solomon Islands, and the United States. ''Cochliobolus carbonum'' is one of the most aggressive members of this genus infecting sorghum (''Sorghum'' spp. ()), corn (''Zea mays'' ()) and apple (''Malus domestica'' ()).〔Manamgoda, D., Cai, L., Bahkali, A., Chukeatirote, E., and Hyde, K. (2011). Cochliobolus: an overview and current status of species. Fungal Diversity 51, 3-42.〕 As one of the most devastating pathogens of sweet corn, ''C. carbonum'' causes Northern leaf spot and ear rot disease 〔 while the asexual stage causes ''Helminthosporium'' corn leaf spot.〔Jones, M.J. and L.D. Dunkle, Analysis of Cochliobolus carbonum races by PCR amplification with arbitrary and gene specific primers. Phytopathology, 1993. 83(4): p. 366-370.〕 ''Cochliobolus carbonum'' is pathogenic to all organs of the corn plant including root, stalk, ear, kernel, and sheath.〔Sindhu, A., et al., A guardian of grasses: Specific origin and conservation of a unique disease-resistance gene in the grass lineage. Proceedings of the National Academy of Sciences, 2008. 105(5): p. 1762-1767.〕 However, symptoms of infection show distinct manifestations in different plant parts: whole plant - seedling blight affects the whole plant, leaf discoloration and mycelial growth, black fungal spores and lesions appear on inflorescences and glumes, and grain covered with very dark brown to black mycelium which gives a characteristic charcoal appearance due to the production of conidia.
==Morphology and keys to identification==
''Cochliobolus carbonum'' is divided into at least five different races based on pathogenicity, particularly lesion types on corn leaves.〔Lu, C., et al., Identification of races and mating types of Cochliobolus carbonum from corn in the yunnan province in China. Journal of Phytopathology, 2013: p. n/a-n/a.〕 ''Cochliobolus carbonum'' race 0 (CCR0) is essentially non-pathogenic.〔Welz, H.G. and K.J. Leonard, Phenotypic variation and parasitic fitness of races of Cochliobolus carbonum on corn in North Carolina. Phytopathology, 1993. 83(6): p. 593-601.〕 ''Cochliobolus carbonum'' race 1 (CCR1) produces host specific toxins and oval to circular spots on leaves.〔Tsukiboshi, T., T. Kimigafukuro, and T. Sato, Identification of races of Bipolaris zeicola, the casual fungus of Helminthosporium leaf spot on corn in Japan. Annals of the Phytopathological Society of Japan, 1987. 53(5): p. 647-649.〕 While infection by ''C. carbonum'' race 2 (CCR2) is rare, it is characterized by the production of oblong, chocolate-colored spots.〔Welz, G. and K.J. Leonard, Genetic variation in field population of race 0, 2 and 3 of Bipolaris zeicola in 1987. Phytopathology, 1988. 78: p. 1574.〕 ''C. carbonum'' race 3 (CCR3) produces linear and narrow lesions〔 while ''C. carbonum'' race 4 (CCR4) is weakly pathogenic, forming oval to concentric circular spots.〔Dodd, J.L. and A.L. Hooker, Previously undescribed pathotype of Bipolaris zeicola on corn. Plant Disease, 1990. 74(7): p. 530.〕
The genus ''Cochliobolus'' is distinguished by the presence of dark to black ascomata with a unilocular, globose pseudothecium and a short, cylindrical neck.〔 Ascomata also bear hyphae and conidiophores and asci are bitunicate, 2-8 spored, and cylindrical to obclavate.〔Sivanesan, A., Graminicolous species of Bipolaris, Curvularia, Drechslera, Exserohilum and their teleomorphs. Mycologia, 1987. 158: p. 1-261.〕 Ascomata are black, globose to ellipsoidal, and setae are present over the upper half of the wall mixed with conidiophores. Pseudoparaphyses are filiform, hyaline, septate, and branched.〔Navi, S.S., et al., A pictorial guide for the identification of mold fungi on sorghum grain. Information Bulletin, International Crops Research Institute for Semi Arid Tropics , Patancheru, Andhra Pradesh, India., 1999. 58.〕 The shapes of asci are cylindrical to clavate, short-stalked, straight to slightly curved having 1-8 spores, and vestigial bitunicate.〔 Conidia are straight to moderately curved, occasionally cylindrical but usually broad in the middle and tapering towards the rounded ends, distoseptate, and 36-100 X 12-18 µm.〔Nelson, R.R., The genetic control of conidial morphology and arrangement in Cochliobolus carbonum. Mycologia 1966. 58(2): p. 208-214.〕 The surface of the conidia is often granulose and the hilum is inconspicuous. Conidia are produced from the apex of an unbranched conidiophore.〔 Generally, the conidiophore arises singly or in small groups which are straight or flexuous, mid to dark brown, smooth, septate, cylindrical, and up to 250 μm long, 5-8 μm thick.〔Nelson, R.R., A major gene locus for compatibility in Cochliobolus heterostrophus Phytopathology 1957. 47: p. 742-743.〕 Variation in mycotoxin production by ''Cochliobolus'' species is used to distinguish some taxa and these mycotoxins are host-specific and non-host specific. HC toxin produced by ''C. carbonum'' race 1 and T toxin produced by ''C. heterostrophus'' are host-specific toxins while ophiobolins produced by ''C. miyabeanus'' and carbotoxin produced by ''C. carbonum'' are non-host-specific toxins.〔
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