The host-pathogen interaction between barley and casual agent of spot blotch (Bipolaris sorokiniana) disease: a review

Document Type : Research Paper


Seed and Plant Improvement Institute, Agricultural Research, Education and Extension Organization, Karaj, Iran


Understanding of the host-pathogen interaction is key to uncovering the defence mechanisms for being used in breeding programs and integrated disease management. Spot blotch is one of the most common foliar diseasesof barley worldwide. Bipolaris sorokiniana (Sacc.) Shoemakeris the causal agent of spot blotch and can cause other destructive diseases of barley such as common root rot, seedling blight and black point or smudge. Adequate knowledge about epidemiology and pathogenicity of the pathogen can provide great assistance to inhibit the outbreak of spot blotch. In this review, first the distribution and host range of the pathogen has beenreviewed. Then, disease symptoms and yield loss caused by spot blotch in barley are reviewed. Subsequently, virulence diversity and other pathogenic aspects ofB. Sorokinianasuch as epidemiology, primary and secondary infections, survival and effect of environmental conditions on epidemic of the disease arediscribed in detail. Later, different aspects of interaction between barley and B. Sorokinianasuch as host response and genetics of resistance has beendiscussed.The importance of molecular markers for studying population structure of the pathogen and genetics of resistance in barley genotypes are also covered. Finally, different disease control measures have beenpresented and discussed.


Adlakha, K.L., Wilcoxson, R.D., and Ray-chauduri, S.P. 1984. Resistance of wheat tospot blotch caused by Bipolaris sorokiniana. Plant Dis. 68:320-321.
Agrios, G.N. 1997. Plant Pathology. 4th ed. Academic Press. San Diego, CA. 633 pp.
Al-Sadi, A.M. 2016. Variation in resistance to spot blotch and the aggressiveness of Bipolaris sorokiniana on barley and wheat cultivars. J. Plant Pathol. 98(1): 97-103.
Al-Daoude, A., Al-Shehadah, E., Shoaib, A., Jawhar, M., and Arabi, M.I.E. 2019. Transcriptional changes of Salicylic Acid dependent signaling pathwaysin barley-Cochliobolus sativus interaction. J. Plant Biochem. Physiol. 7(1). DOI: 10.4172/2329-9029.1000228.
Anderson, W.H., and Banttari, E.E. 1976. The effect of Bipolaris sorokiniana on yield,kernel weight and kernel discoloration in six-row spring barley. Plant Dis. Reporter.60:754-758.
Anonymous. 1986. Cochliobolus sativus. Distribution Maps of Plant Diseases. No. 322(4thedition). CAB International, Wallingford, UK.
Arabi, M.I.E., and Jawhar, M. 2003. Germinability of Cochliobolus sativus conidiaexposed to solar radiation. J. Phytopathol. 151:620-624.
Arabi, M.I.E., and Jawhar, M. 2004. Identification of Cochliobolus sativus (spot blotch)isolates expressing differential virulence on barley genotypes in Syria. J.Phytopathol. 152:461-464.
Bailey, K.L., Johnston, A.M., Kutcher, H.R., Gossen, B.D., and Morrall, R.A.A. 2000.Managing crop losses from foliar diseases with fungicides, rotation, and tillage inthe Saskatchewan Parkland. Can. J. Plant Sci. 80:169-175.
Bailey, K.L., Gossen, B.D., Lafond, G.P., Watson, P.R., and Derksen, D.A. 2001. Effectof tillage and crop rotation on root and foliar diseases of wheat and pea inSaskatchewan from 1991 to 1998: Univariate and multivariate analyses. Can. J.Plant Sci. 81:789-803.
Bailey, K.L., Gossen, B.D., Gugel, R.K., and Morrall, R.A.A. 2003. Diseases of fieldcrops in Canada. 3rd ed. The Canadian Phytopathological Society. Ontario,Canada. 304 pp.
Bakonyi, J., Aponyi, I., and Fischl, G. 1997. Diseases caused by Bipolaris sorokinianaand Drechslera tritici repentis in Hungary. Pp. 80-85. In: E. Duveiller, H. J. Dubin, J. Reeves, and A. McNab (eds.)Helminthosporiumblights of wheat: spot blotch and tan spot.  CIMMYT, Mexico D.F., Mexico.
Biles, C.L. and Hill, J.P. 1988. Effect of Trichoderma harzianum on sporulation ofCochliobolus sativus on excised wheat seedling leaves. Phytopathol. 78:656-659.
Bilgic, H., Steffenson, B., and Hayes, P.M. 2005. Comprehensive genetic analyses revealdifferential expression of seedling and adult plant resistance to spot blotch inpopulations of barley. Theor. Appl. Genet. 111:1238-1250.
Bilgic, H., Steffenson, B.J., and Hayes, P.M. 2006. Molecular mapping of loci conferringresistance to different pathotypes of the spot blotch pathogen in barley.Phytopathol. 96:699-708.
Bonman, J.M., Bockelman, H.E., Jackson, L.F., and Steffenson, B.J. 2005. Disease andinsect resistance in cultivated barley accessions from the USDA National SmallGrains Collection. Crop Sci. 45:1271-1280.
Boosalis, M.G. 1962. Precocious sporulation and longevity of conidia ofHelminthosporium sativum in soil. Phytopathol. 52:1172-1177.
Bulat, S.A., and Mironenko, N.V. 1993. Genetic differentiation of phytopathogenicfungus Cochliobolus sativus (Ito and Kurib.) Drechsl. ex Dastur (Bipolarissorokiniana (Sacc.: Sorok.) Shoem.) revealed by a universally primed polymerasechain reaction UP-PCR technique: Correlation with host-specificity. Genetika. 29:1295-1301.(in Russian).
Chester, K.S. 1950. Plant disease losses: their appraisal and interpretation. Plant Dis.Reporter Suppl. 193:189-362.
Chinn, S.H.F. 1965. Changes in the spore population of Cochliobolus sativus inSaskatchewan wheat fields. Can. J. Plant Sci. 45:288-291.
Chinn, S.H.F. 1976. Cochliobolus sativus conidia populations in soils following variouscereal crops. Phytopathol. 66:1082-1084.
Chinn, S.H.F. 1977. Influence of fungicide sprays on sporulation of Cochliobolus sativuson cypress wheat and on conidial populations in soil. Phytopathology 67:133-138.
Christensen, J.J. 1922. Studies on the parasitism of Helminthosporium sativum. University of Minnesota Agricultural ExperimentalStation. Tech. Bull. No. 11:1-42.
Clark, R.V. 1971. Influence of Vitavax seed treatment and loose smut infection on yields
of barley and wheat. Can. Plant Dis. Surv. 51:131-133.
Clark, R.V. 1979. Yield losses in barley cultivars caused by spot blotch. Can. J. PlantPathol. 2:113-117.
Clark, R.V., and Dickson, J.G. 1958. The influence of temperature on diseasedevelopment in barley infected by Helminthosporium sativum. Phytopathol.48:305-310.
Conner, R.L., Duczek, L.J., Kozub, G.C., and Kuzyk, A.D. 1996. Influence of croprotation on common root rot of wheat and barley. Can. J. Plant Pathol. 18:247-254.
Cook, R.J., and Timian, R.G. 1962. Lesion type as a means of detecting resistance inbarley leaves to the spot blotch Helminthosporium. Phytopathol. 52:1086-1089.
Couture, L., and Sutton, J.C. 1978a. Control of spot blotch in barley by fungicideapplications timed according to weather factors. Phytoprotect. 59:65-75.
Couture, L., and Sutton, J.C. 1978b. Efficacies of fungicides in controlling spot blotch ofbarley. Can. J. Plant Sci. 58:311-317.
Couture, L., Sutton, J. C. 1978c. Relation of weather variables and host factors toincidence of airborne spores of Bipolaris sorokiniana. Can. J. Bot. 56:2162-2170.
Couture, L., and Sutton, J.C. 1980. Effect of dry heat treatments on survival of seed borneBipolaris sorokiniana and germination of barley seeds. Can. Plant Dis. Surv.60:59-61.
De Oliveira, A.M., Matsumura, A.T., Prestes, A.M., and Van Der Sand S.T. 2002.Intraspecific variability of Bipolaris sorokiniana isolates determined by randomamplifiedpolymorphic DNA (RAPD). Genet. Mol. Res. 1:350-358.
Dickson, J.G.1956. Diseases of field crops. 2nd ed. McGraw-Hill Book Co., New York.517 pp.
Dosdall, L. 1923. Factors influencing the pathogenicity of Helminthosporium sativum.University of Minnesota Agricultural ExperimentalStation. Tech. Bull. No. 17.
Dostaler, D., Couture, L., and Pelletier, G.J. 1987. Étude de la tolérance de cultivars d'orge à la tache helminthosporienne. Can. J. Plant Sci. 67:153–157.
Duczek, L.J. and White, G.P. 1986. Chalara heteroderae a fungal antagonist thatparasitizes hyphae of Bipolaris sorokiniana. Soil Biol. Biochem. 18:655-659.
Duczek, L.J., Sutherland, K.A., Reed, S.L., Bailey, K.L., and Lafond, G.P. 1999. Survivalof leaf spot pathogens on crop residues of wheat and barley in Saskatchewan.Can. J. Plant Pathol. 21:165-173.
Duveiller, E., and Garcia Altamirano, I. 2000. Pathogenicity of Bipolaris sorokinianaisolates from wheat roots, leaves and grains in Mexico. Plant Pathol. 49:235-242.
Fetch, T.G., and Steffenson, B.J. 1999. Rating scales for assessing infection responses ofbarley infected with Cochliobolus sativus. Plant Dis. 83: 213-217.
Fokkema, N.J., Den-Houter, J.G., Kosterman, Y.J.C., and Nelis, A.L.1979. Manipulationof yeasts on field grown wheat leaves and their antagonistic effect on Cochliobolus sativus and Septoria nodorum. Trans. Br. Mycol. Soc. 72:19-30.
Gamba, F., and Estramill, E. 2002. Variation in virulence within Uruguayan population of
Cochliobolus sativus. Pp. 59-62. In: Proc. of the Second International Workshop onBarley Leaf Blights. April 7-11, 2002, ICARDA, Aleppo, Syria.
Gayed, S.K. 1962. The pathogenicity of six strains of Helminthosporium sativum to threecereals with special reference to barley. Mycopathologia 18:271-279.
Ghazvini, H., and Tekauz, A. 2004. Yield loss in barley inoculated with high and lowvirulence isolates of Bipolaris sorokiniana. Pp. 774-780. In: Proc. of the 9th International Barley Genetic Symposium, June 20-26, 2004, Brno, Czech Republic.
Ghazvini, H., and Tekauz, A. 2007. Virulence Diversity in Population of Bipolaris sorokiniana. Plant Dis. 91:814-821.
Ghazvini, H., and Tekauz, A. 2008. Host × pathogen interactions among barley genotypes and Bipolaris sorokiniana isolates. Plant Dis. 92:225-233.
Ghazvini, H. 2012. Adult plant resistance and yield loss in barley cultivars inoculated with a newly-emerged pathotype of Bipolaris sorokiniana in Manitoba, Canada. Crop Breeding Journal. 2 (1): 9-15.
Ghazvini, H., and Tekauz, A. 2012. Molecular diversity in the barley pathogen Bipolaris sorokiniana (Cochliobolus sativus). Aust. Plant Pathol. 41:283–293.
Ghazvini , H. 2014. Identification, by selective genotyping, of quantitative trait loci conferring resistance to Cochliobolus sativus in barley line TR 251. Crop Breeding Journal 4(1) : 35-45.
Gilbert, J., Wood, S.M., and Tekauz, A. 1998. Relationship between environmentalvariables and the prevalence and isolation frequency of leaf-spotting pathogens inspring wheat. Can. J. Plant Pathol. 20:158-164.
Gonzalez Ceniceros, F. 1990. Assigning genes conferring resistance to spot and netblotch in barley to a specific chromosome. Ph.D. dissertation. North Dakota StateUniversity, Fargo.
Hampton, J.G. 1979. The presence and effects of seed-borne and soil-borne Drechslerasorokiniana on barley in New Zealand. III. Effect on grain yield. New Zeal. J. Exp.Agric. 7:391-393.
Herrman, T.J., Forster, R.L., and Martin, J.M. 1990. Imazalil seed treatment reducescommon root rot and increases yield of barley under commercial conditions. PlantDis. 74:246-247.
Hetzler, J., Eyal, Z., Mehta, Y.R., Campos, L.A., Fehrmann, H., Kushnir, U., Zekaria-Oren, J., and Cohen, L. 1991. Interactions between spot blotch (Cochliobolussativus) and wheat cultivars. Pp. 146-164. In: Saunders, D.A., and Hettel, G. (eds.)Wheat for the Non-traditional WarmAreas. CIMMYT, Mexico D.F., Mexico.
Ibeagha, A., Hückelhoven, R., and Kogel, K.H. 2005. Model wheat genotypes as tools touncover effective defense mechanisms against the hemibiotrophic fungusBipolaris sorokiniana. Phytopathology 95:528-532.
James, W.C. 1974. Assesment of plant diseases and losses. Annu. Rev. Phytopathol.12:27-48.
Jawhar, M., Till, B.J., Albaterni, A.,  Skiheita, A., Arabi, M.I.E., Bakri, Y., and MirAli N. 2017. Optimizing Eco-Tilling technique for tracking field Cochliobolus sativus population. Journal of Plant Pathol. 99: 339-345.
Kiesling, R.L. 1985. The diseases of barley. Pp. 269-312. In: Barley. Rasmusson, D.C. (ed.) Agronomy Monograph No. 26, ASA. Madison WI.
Knight, N.L., Platz, G.J., Lehmensiek, A., and Sutherland, M.W. 2010. An investigation of genetic variation among Australian isolates of Bipolaris sorokiniana from different cereal tissues and comparison of their abilities to cause spot blotch on barley. Aust. Plant Pathol. 39:207–216.
Krupinsky, J.M., Tanaka, D.L., Lares, M.T., and Merrill, S.D. 2004. Leaf spot diseases ofbarley and spring wheat as influenced by preceding crops. Agron. J. 96:259-266.
Kumar, J., Hückelhoven, R., Beckhove, U., Nagarajan, S., and Kogel, K.H. 2001. Acompromised Mlo pathway affects the response of barley to the necrotrophicfungus Bipolaris sorokiniana (teleomorph: Cochliobolus sativus). Phytopathology91:127-133.
Kumar, J., Schäfer, P., Hückelhoven, R., Langen, G., Baltruschat, H., Stein, E.,Nagarajan, S., and Kogel, K.H. 2002. Bipolaris sorokiniana, a cereal pathogen ofglobal concern: Cytological and molecular approaches towards better control.Mol. Plant Pathol. 3:185-195.
Kutcher, H.R., Bailey, K.L., Rossnagel, B.G., and Legge, W.G. 1996. Identification ofRAPD markers for common root rot and spot blotch (Cochliobolus sativus)resistance in barley. Genome 39:206-215.
Lee, C.J., Manthey, F.A., Ellefson, R.S., Horsley, R.D., and Schwarz, P.B. 1997. Effectsof the fungicide propiconazole on agronomic and quality traits of hulled andhulless barley. J. Prod. Agric. 10:320-322.
Leisova-Svobodova, L., Minarikova, V., Kucera, L., and Pereyra, S.A. 2012. Structure of the Cochliobolus sativus population variability. Plant Pathol. 61: 709-718.
Leng, Y., Wang, R., Ali, S., Zhao, M., and Zhong, S. 2016. Sources and genetics of spot blotch resistance to a new pathotype of Cochliobolus sativus in the USDA Small Grains Collection. Plant Dis. 100:1988-1993.
Levitin, M.M., Petrova, A.N., and Afanasenko, O.S. 1985. Comparative analysis ofBipolaris sorokiniana population for virulence. Mikol. Fitopatol.19:154-158.(in Russian).
Luz, W.C., and Vieira, J.C. 1982. Seed treatment with systemic fungicides to controlCochliobolus sativus on barley. Plant Dis. 66:135-136.
Maraite, H., Di Zinno, T., Longrée, H., Daumerie, V., and Duveiller, E. 1998. Fungiassociated with foliar blight of wheat in warmer areas. Pp. 293-300. In: E. Duveiller, H.J., Dubin, J., Reeves, and A., McNab, (eds.)Proc. of theInternational Workshop on Helminthosporium Diseases of Wheat: Spot Blotchand Tan Spot. February9-14, 1997. CIMMYT, El Batán, Mexico.
Mathre D.E. 1997. Compendium of barley diseases, 2nd ed. American PhytopathologicalSociety Press, St. Paul, Minnesota. 90 pp.
McGee, D.C. 1995. Epidemiological approach to disease management through seedtechnology. Ann. Rev. Phytopathol. 33:445-466.
Mead, H.W. 1942. Host-parasite relationships in a seed-borne disease of barley caused byHelminthosporium sativum Pammel, King and Bakke. Can. J. Res., Sect. C.20:501-523.
Meldrum, S.I., Ogle, H.J., and Platz, G.J. 2004. Pathotypes of Cochliobolus sativus onbarley in Australia. Aust. Plant Pathol. 33:109-114.
Meronuck, R.A., and Pepper, E.H. 1968. Chlamydospore formation in conidia ofHelminthosporium sativum. Phytopathol. 58:866-867.
Mills, J.T., and Wallace, H.A.H. 1969. Effect of fungicides on Cochliobolus sativus andother fungi on barley seed in soil. Can. J. Plant Sci. 49:543-48.
Mironenko, N.V., and Bulat, S.A. 2001. Genetic structure of Cochliobolus sativus(Bipolaris sorokiniana) populations isolated from different hosts as revealed byUP-PCR (RAPD-like) technique. J. Russ. Phytopathol. Soc. 2:25-30.
Mumford, D.L. 1966. Factors associated with resistance in barley to spot blotch.Phytopathol. 56:79-82.
Nakada, M., Tanaka, C., Tsunewaki, K., and Tsuda, M. 1994. RFLP analysis for speciesseparation in the genera Bipolaris and Curvularia. Mycoscience 35: 271-278.
Nelson, R.R., and Kline, D.M. 1961. The pathogenicity of certain species ofhelminthosporium to species of the Gramineae. Plant Dis. Reporter. 45:644-648.
Nelson, R.R., and Kline, D.M. 1962. Intraspecific variation in pathogenecity in the genusHelminthosporium to Gramineous species. Phytopathol. 52:1045-1049.
Nelson, R.R., and Kline, D.M. 1963. Pathogenicity of isolates of Cochliobolus sativusfrom cultivated and wild Gramineous hosts from the western hemisphere tospecies of the Gramineae. Plant Dis. Reporter. 47:890-894.
Nutter, F.W. Jr., Pederson, V.D., and Foster, A.E. 1985. Effect of inoculation withCochliobolus sativus at specific growth stages on grain yield and quality ofmalting barley. Crop Sci. 25:933-938.
Pammel, L.H., King, C.M., and Bakke A.L. 1910. Two barley blights, with comparison ofspecies of Helminthosporium upon cereals. Iowa Agric. Exp. Sta. Bull. 116:178-190.
Patil, I.S., Kulkarni, S., and Hegde, R.K. 1987. Antagonistic action of species ofTrichoderma, Bacillus and Streptomyces on Drechslera sorokiniana (Sacc.)Subram and Jain. Pesticides 21:22.
Prasad, K.S.K., Kulkarni, S., and Siddaramaiah, A.L. 1978. Antagonistic action ofTrichoderma sp. and Streptomyces sp. on Drechslera sativum (Helminthosporiumsativum Pam, King and Bakke) Subram. and Jain. Curr. Res. 12:202-203.
Presser, R.W. 1991. Studies on the injurious effect of Drechslera sorokiniana (Sacc.)Subram. et Jain in spring barley (Hordeum vulgare L.). Arch. Phytopathol. PFL.27:127-133.
Qasem, J.R., Al-Abed, A.S., and Abu-Blan, H.A. 1995. Antifungal activity of clammyinula (Inula viscosa) on Helminthosporium sativum and Fusarium oxysporum f.sp. Lycopersici. Phytopathol. Mediterr. 34:7-14.
Raemaekers, R.H., and Tinline, R.D. 1981. Epidemic of diseases caused by Cochliobolussativus on rainfed wheat in Zambia. Can. J. Plant Pathol. 3:211-214.
Reis, E.R. 1984. Efeito da incorporacao e do tipo de restos culturais de trigo namultiplicacao de Helminthosporium sativum, no solo. Fitopatol. Bras. 9:537-541.
Reis, E.R., and Wünsche, W.A. 1984. The sporulation of Cochliobolus sativus on residuesof winter crops and its relationship to the increase of inoculum density in soil.Plant Dis. 68:411-412.
Richardson, L. T. 1972. Effectiveness of systemic fungicide seed dressings as protectantsof barley seedlings against Cochliobolus sativus. Can. J. Plant Sci. 52:949-953.
Sarhan, A.R.T., Király, Z., Sziráki, I., and Smedegaard-Petersen, V. 1991. Increasedlevels of cytokinins in barley leaves having the systemic acquired resistance toBipolaris Sorokiniana (Sacc.) Shoemaker. J. Phytopathol. 131:101-108.
Schäfer, P., Hückelhoven, R., and Kogel, K.H. 2004. The white barley mutant albostriansshows a super-susceptible but symptomless interaction phenotype with the hemibiotrophicfungus Bipolaris sorokiniana. Mol. Plant-Microbe Interact. 17:366-373.
Shaner, G. 1981. Effects of environment on fungal leaf blights of small grain. Ann. Rev.Phytopathol. 19:273-296.
Sharma-Poudyal, D., Duveiller, E., and Sharma, R.C. 2005. Effects of seed treatment andfoliar fungicides on Helminthosporium leaf blight and on performance of wheat inwarmer growing conditions. J. Phytopathol. 153:401-408.
Sheehy, J., and Huguelet, J.E. 1967. Diurnal periodicity in Helminthosporium andAlternaria spores in the air. Phytopathology 57:830. (Abstract)
Singh, D., Pande, S.K.,Singh,S.P.,and Singh, D.K.2017. Evaluation of the barley genotypes against spot blotch disease caused by bipolaris sorokiniana.Plant Archives.17(1): 167-170.
Sivanesan, A. 1987. Graminicolous species of Bipolaris, Curvularia, Drechslera,Exserohilum and their teleomorphs. Mycological Papers 158, CAB International,Wallingford, UK. 261 pp.
Sprague, R. 1950. Diseases of cereals and grasses in North America. The Ronald PressCo., New York. 538 pp.
Spurr, H.W. Jr., and Kiesling, R.L. 1961. Field and host studies of parasitism byHelminthosporium sorokinianum. Plant Dis. Reporter. 45:941-943.
Stack, R.W., and McMullen, M.P. 1991. Effect of fungicidal seed treatments on commonroot rot of spring wheat and barley. North Dakota Farm Res. 49:13-16.
Steffenson, B.J., and Smith, K.P. 2004. Breeding barley for multiple disease resistance inupper Midwest region of the USA. Pp. 294-301. In: Proc. of the 9th InternationalBarley Genetic Symposium, June 20-26, 2004, Brno, Czech Republic.
Steffenson, B.J., Hayes, P.M., and Kleinhofs, A. 1996. Genetics of seedling and adultplant resistance to net blotch (Pyrenophora teres f. teres) and spot blotch(Cochliobolus sativus) in barley. Theor. Appl. Genet. 92: 552-558.
Tekauz, A., Gilbert, J., Mueller, E., Stulzer, M., Beyene, M., Ghazvini, H., Morgan, K.,and Reverchon, F. 2003. Survey for foliar diseases of barley in Manitoba in 2002.Can. Plant Dis. Surv. 83:60-61.
Terekhova, V.A., and Rochev, M.V. 1989. Comparative characterization of mycelia isozymes of Bipolaris sorokiniana Shoem. isolates of different origin. Moscow-Univ-Biol-Sci-Bull. 44:77-81.
Tinline, R.D. 1960. Pathogenic and cultural variation in Cochliobolus sativus. Trans. Brit.Mycol. Soc. 43:696-697.
Tinline, R.D. 1988. Cochliobolus sativus, a pathogen of wide host range. Pp. 113-122. In:D. S. Ingram, and P. H. Williams (eds.)Advances in Plant Pathology, Vol. 6.Academic Press, London, England.
Valim-Labres, M.E., Porto, M.D., and Matsumura, A.T.S. 1997. Effects of host resistanceon the isozymatic patterns of Bipolaris sorokiniana (Dematiaceae, Moniliales).Braz. J. Genet. 20:541-545.
Valjavec-Gratian, M., and Steffenson, B.J. 1997a. Pathotypes of Cochliobolus sativus onbarley. Plant Dis. 81:1275-1278.
Valjavec-Gratian, M., and Steffenson, B.J. 1997b. Genetics of virulence in Cochliobolussativus and resistance in barley. Phytopathol. 87:1140–1143.
Verma, P.R., Chinn, S.H.F., Crowle, W.L., Spurr, D.T., and Tinline, R.D. 1981. Effect ofimazalil seed treatment on common root rot and grain yields of cereal cultivars.Can. J. Plant Pathol. 3:239-243.
Whittle, A.M., and Richardson, M.J. 1978.Yield loss caused by Cochliobolus sativus onClermont barley. Phytopathol. Z. 91:238-256.
Wilcoxson, R.D., Rasmusson, D.C., and Miles, M.R. 1990. Development of barleyresistant to spot blotch and genetics of resistance. Plant Dis. 74:207-210.
Winter, W., Baenziger, I., Krebs, H., Rueegger, A., Frei, P., and Gindrat, D. 1996. Warmand hot water treatment of barley seed. Agrarforschung 3:25-28.
Wood, L.S. 1962. Relation of variation in Helminthosporium sativum to seedling blightof small grains. Phytopathology 52:493-497.
Yashwant, C.K., Rao, G.M., and Singh, S.K. 2017. Hostsof Bipolaris sorokiniana, the major pathogen of spot blotch of wheat in Uttar Pradesh. Indian J. of Res. 6(3): 593-594.
Yun, S.J., Gyenis, L., Hayes, P.M., Matus, I., Smith, K.P., Steffenson, B.J., andMuehlbauer, G.J. 2005. Quantitative trait loci for multiple disease resistance inwild barley. Crop Sci. 45:2563-2572.
Yun, S.J., Gyenis, L., Bossolini, E., Hayes, P.M., Matus, I., Smith, K.P., Steffenson, B.J.,Tuberosa, R. ,and Muehlbauer, G.J. 2006. Validation of quantitative trait loci formultiple disease resistance in barley using advanced backcross lines developedwith a wild barley. Crop Sci. 46:1179-1186.
Zhong, S., and Steffenson, B.J. 2001. Virulence and molecular diversity in Cochliobolussativus. Phytopathology 91:469-476.
Zhong, S., and Steffenson, B.J. 2002. Identification and characterization of DNA markersassociated with a locus conferring virulence on barley in the plant pathogenicfungus Cochliobolus sativus. Theor. Appl. Genet. 104:1049-1054.
Zhong, S., Steffenson, B.J., Martinez, J.P., and Ciuffetti, L.M. 2002. A molecular geneticmap and electrophoretic karyotype of the plant pathogenic fungus Cochliobolussativus. Mol. Plant-Microbe Interact. 15:481-492.
Zillinsky, F.J. 1983. Common diseases of small grain cereals: a guide to identification.CIMMYT, Mexico D.F. Mexico. 141 pp.