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The Mode of Inheritance of Factors for Resistance

About 40 years ago a standard set of differential varieties was selected from among several hundred varieties of different species groups of

B. The Host 1. Sources of Resistance to Pathogens

5. The Mode of Inheritance of Factors for Resistance

It is difficult to generalize about the mode of inheritance of factors for resistance except to say that determinations must be made in each specific case. In his pioneer studies, Biffen (1907) demonstrated that resistance to yellow rust of wheat was inherited as a simple recessive.

It was at first thought that this mode of inheritance might be universal, but it is not. The immunity of Kanred wheat from a considerable number of races of wheat stem rust is due to a single dominant gene. In other cases of resistance to stem rust two or more factor pairs are involved;

in still others there are multiple factors, some major, some minor. The mode of inheritance, then, within one kind of crop plant to one kind of pathogen may differ greatly in different crosses.

Almost every type of genie relationship has been reported in connec-tion with the genetics of bunt resistance in wheat (Fischer and Holton, 1957). There are many physiologic races of the pathogen and many wheat varieties have been crossed in attempts to checkmate them. In the many crosses between resistant and susceptible varieties, many factors apparently are involved. In some cases a single factor was involved, with or without modifiers; in others there are two, three, or more factors for resistance. Multiple factors have been recorded in many cases. Factors may be strong or weak, linked or independent, effective against a single race or against several. There may be inhibitors of factors for resistance to certain smut races but not to others. In resistant X resistant crosses, there may, of course be no segregation; in some, there is transgressive segregation.

Generalization regarding mode of inheritance cannot be applied to specific cases. Each must be considered separately. It can only be said, therefore, that the number of genes for resistance varies greatly with the crop plant and with the pathogen, and the way in which they are inherited varies greatly also.

Naturally there are many factors for virulence in species of pathogens that comprise many physiologic races. The final outcome of breeding for disease resistance will be determined by the ability of breeders to find and combine genes for resistance that counteract the genes for virulence in pathogens. As different genes for resistance are often dis-persed among many varieties of crop plants and those for virulence often are widely dispersed among many physiologic races of pathogens, it is

not an easy task to prognosticate the outcome of many breeding pro­

grams. Information is continually being accumulated about the genes for resistance, and some has been accumulated about the genes for virulence, but much more is needed.

Much more needs to be known about the genetics of pathogens. The writers prepared a brief summary in 1953 (Stakman and Christensen, 1953). Flor (1954, 1956) has made an especially comprehensive study of virulence in flax rust, Melampsora lini, and of resistance in flax. Flor suggested the presence in flax of a gene for susceptibility or resistance to match each gene for virulence or avirulence in the pathogen. He obtained 30 lines of flax, each apparently homozygous for an individual gene that governs reaction to rust. By using 18 of the lines as differential varieties, it was possible to distinguish 239 races of rust. In crosses between races of rust it appeared that avirulence was dominant over virulence and that from one to three factor pairs were involved. In crosses between resistant and susceptible varieties of flax, resistance was dominant and one to three factor pairs were involved. According to Flor, varieties of flax that carry " L L " genes are resistant to all North American races of rust. More work of this kind is needed.

V. F U T U R E PROSPECTS

Progress in the future should be more rapid and more nearly permanent than in the past. Many breeding programs must be inter­

national in scope to be successful, simply because different genes for resistance and for virulence may exist in different parts of the world.

Varieties with different genes for resistance must be assembled and then tested extensively in different geographical areas and intensively under optimum conditions for disease development. Breadth of experimentation must be combined with intensity of research. Facilities for both types of effort are being improved, but they must be expanded. There must be facilities for more travel in order that all possible information may be obtained from the international disease nurseries that are now grow­

ing in number. Large phytotrons are needed at strategic locations so that the most promising plant materials may be tested under necessary combinations of environmental conditions. Appropriate elimination plots are needed where plant materials can be assembled and exposed to infection by a world-wide collection of pathogens. There must be much more intensive research on the genetics, ecology, and physiology of hosts and pathogens and on the nature and variability of disease resistance.

These things are prerequisite to most rapid and assured progress. In the meantime progress is being made.

It is possible that attitudes must be changed. It is entirely possible

that a succession of varieties may have to be produced to control some diseases, as there can be no assurance that new pathogens or new races of old ones may not appear periodically to disrupt the status quo. The important thing is to anticipate the changes and prepare to meet them.

Even if permanent immunity is not attainable, practical field resistance or tolerance may reduce the destructiveness of many diseases. Quality may sometimes have to be sacrificed in order to attain greater resistance and productivity, especially when the standard of quality is not based on real values.

If the efforts to produce resistant varieties are really intelligent, efficiently systematic, and world-wide in scope, there is great promise for the future. Above all, the lessons of the past must not be forgotten and many new ones must be learned in the future. Progress in scientific attitudes and attainments can guarantee progress in the production of resistant varieties.

R E F E R E N C E S

Anwar, A. A. 1949. Factors affecting the survival of Helminthosporium sativum and Fusarium lini in soil. Phytopathohgy 3 9 : 1005-1019.

Ausemus, E. R., E. C. Stakman, E. W. Hanson, W. F. Geddes, and P. P. Merritt.

1944. Newthatch wheat. Minn. Univ. Agr. Expt. Sta. Tech. Bull. 1 6 6 .

Bain, S. M., and S. H. Essary. 1906. Selection for disease resistant clover: a prelim­

inary report. Tenn. Univ. Agr. Expt. Sta. Bull. 7 5 .

Barker, H. D. 1923. A study of wilt resistance in flax. Minn. Univ. Agr. Expt. Sta.

Tech. Bull. 2 0 .

Biffen, R. H. 1907. Studies in the inheritance of disease resistance. /. Agr. Sci. 2 : 109-128.

Biffen, R. H. 1912. Studies in the inheritance of disease resistance II. /. Agr. Sci.

4 : 421-429.

Biffen, R. H. 1931. The cereal rusts and their control. Trans. Brit. Mycol. Soc. 1 6 : 19-37.

Bingefors, S. 1957. Studies on breeding red clover for resistance to stem nematodes.

Vaxtodling 8 : 1-123.

Bolley, H. L. 1901. Flax wilt and flax-sick soil. N. Dakota Agr. Expt. Sta. Bull. 5 0 . Borlaug, Ν. E. 1945. Variation and variability of Fusarium lini. Minn. Univ. Agr.

Expt. Sta. Tech. Bull. 1 6 8 .

Borlaug, Ν. E. 1957. The development and use of composite varieties based upon the mechanical mixing of phenotypically similar lines developed through back-crossing. Kept. 3rd Wheat Rust Conf. Mexico, D. F., March 18-24, 1956. pp.

12^18.

Borlaug, Ν. E. 1958. II. The impact of agricultural research on Mexican wheat production. Trans. Ν. Y. Acad. Sci. [2] 2 0 : 278-295.

Borlaug, Ν. E., J. A. Rupert, and J. G. Harrar. 1949. New wheats for Mexico. (Mex.) Ofic. Estud. Esp., Sec. Agr. y Ganaderia, Fol. Divulgacion 5 .

Briggs, F. N. 1938. The use of the backcross in crop improvement. Am. Naturalist 7 2 : 285-292.

Broadfoot, W. C. 1926. Studies on the parasitism of Fusarium lini Bolley.

Phyto-pathology 1 6 : 951-978.

Campbell, W. S. 1912. An historical sketch of William Farrer's work in connection with his improvements in wheats for Australian conditions. Australasian Assoc.

Advance. Sci. 1 3 : 525-^536.

Christensen, J. J., and T. W. Graham. 1934. Physiologic specialization and variation in Helminthosporium gramineum Rab. Minn. Univ. Agr. Expt. Sta. Tech. Bull.

9 5 .

Clark, J. A. 1936. Improvement in wheat. Yearbook Agr. U. S. Dept. Agr. 1 9 3 6 : 207-302.

Coons, G. H. 1953. Breeding for resistance to disease. Yearbook Agr. U. S. Dept.

Agr. 1 9 5 3 : 174-192.

DuCharme, E. P., and W. Birchfield. 1956. Physiologic races of the burrowing nematode. Phytopathology 4 6 : 615-616.

Edgerton, C. W., and C. C. Moreland. 1920. Tomato wilt. Louisiana Agr. Expt.

Sta. Tech. Bull. 1 7 4 .

Eriksson, J. 1894. Ueber die Specialisierung des Parasitismus bei den Getreiderost-pilzen. Ber. deut. botan. Ges. 1 2 : 331.

Fischer, G. W., and C. S. Holton. 1957. "Biology and Control of the Smut Fungi."

Ronald Press, New York.

Flor, Η. H. 1954. Identification of races of flax rust by lines with single rust-con­

ditioning genes. U. S. Dept. Agr. Tech. Bull. 1 0 8 7 .

Flor, Η. H. 1956. The complementary genie systems in flax and flax rust. Advances in Genet. 8 : 29^54.

Freeman, Ε. M., and E. C. Johnson. 1911. The rusts of grains in the United States.

U. S. Dept. Agr. Bur. Phnt Ind. Bull. 2 1 6 .

Giddings, N. J. 1954. Two recently isolated strains of curly-top virus. Phytopathology 4 4 : 123-124.

Gordon, W. L. 1933. A study of the relation of environment to the development of the uredial and telial stages of the physiologic forms of Puccinia graminis avenae Erikss. and Henn. Sci. Agr. 1 4 : 184-237.

Hart, H. 1949. Nature and variability of disease resistance in plants. Ann. Rev.

Microbiol. 9 : 289-316.

Hayes, Η. Κ., E. R. Ausemus, E. C. Stakman, C. H. Bailey, Η. K. Wilson, R. H.

Bamberg, M. C. Markley, R. F. Crim, and Μ. N. Levine. 1936. Thatcher wheat. Minn. Univ. Agr. Expt. Sta. Bull. 3 2 5 .

Hayes, Η. K., F. R. Immer, and D. C. Smith. 1955. "Methods of Plant Breeding,"

2nd ed., Chapt. 9. McGraw-Hill, New York.

Hiratsuka, N. 1896. Report of the investigation on flax wilt-disease. Hokkai no Shokusan 4 8 (in Japanese).

Hiratsuka, N. 1903. On the cause of flax wilt-disease and its prevention. Bull. Agr.

Soc. Hokkaido 2 (in Japanese).

Jones, F. G. W. 1957. Resistance-breaking biotypes of the potato root eelworm (Heterodera rostochiensis Woll.). Nematologica 2 : 185-192.

Jones, L. R., J. C. Walker, and W. B. Tisdale. 1920. Fusarium resistant cabbage.

Wisconsin Univ. Agr. Expt. Sta. Research Bull. 4 8 .

Kernkamp, M. F., D. J. de Zeeuw, S. M. Chen, B. C. Ortega, C. T. Tsiang, and A. M. Khan. 1952. Investigations on physiologic specialization and parasitism of Rhizoctonia sohni. Minn. Univ. Agr. Expt. Sta. Tech. Bull. 2 0 0 .

Konzak, C. F. 1954. Stem rust resistance in oats induced by nuclear radiation.

Agron. J. 4 6 : 538-540.

Leach, J. G., P. Decker, and H. Becker. 1939. Pathogenic races of Actinomyces scabies in relation to scab resistance. Phytopathology 29: 204-208.

McFadden, E. S. 1930. A successful transfer of emmer characters to vulgare wheat.

Agron. J. 22: 1020-1034.

Murphy, A. M., C. W. Bennett, and F. V. Owen. 1959. Varietal reaction of sugar beets to curly top virus strain 11 under field conditions. /. Am. Soc. Sugar Beet Technologists 10 : 281-282.

Newton, M., and T. Johnson. 1944. Physiologic specialization of oat stem rust in Canada. Can. J. Research 22: 201-216.

Orton, W. A. 1900. The wilt disease of cotton and its control. Bull. U. S. Dept.

Agr. Div. Vegetable Physiol, and Pathol. 27.

Orton, W. A. 1909. The development of farm crops resistant to disease. Yearbook Agr. U. S. Dept. Agr. 1908: 453-464.

Peterson, R. F. 1958. Twenty-five years' progress in breeding new varieties of wheat for Canada. Empire J. Exptl. Agr. 26: 104-122.

Salmon, E. S. 1904. On Erysiphe graminis D. C. and its adaptive parasitism within the genus Bromus. Ann. Mycol Notitiam Sci. Mycol. Univ. 2 : 255-267, 307-343.

Schaal, L. A. 1944. Variation and physiologic specialization in the common scab fungus (Actinomyces scabies). J. Agr. Research 69: 169^-186.

Simons, M. D. 1954. A North American race of crown rust attacking the oat varieties Landhafer and Santa Fe. Phnt Disease Reptr. 38: 505-506.

Simons, M. D. 1956. Physiologic races of crown rust of oats identified in 1955.

Plant Disease Reptr. 40: 810-813.

Smith, G. S. 1943. Two new durum wheat varieties. N. Dakota Agr. Expt. Sta.

Bimo. Bull. 5 : 2-3.

Stakman, E. C. 1955. Progress and problems in plant pathology. Ann. Appl. Biol.

42: 22-33.

Stakman, E. C , and J. J. Christensen. 1953. Problems of variability in fungi.

Yearbook Agr. U. S. Dept. Agr. 1953: 35-62.

Stakman, E. C , and W. Q. Loegering. 1944. The potential importance of race 8 of Puccinia graminis avenap in the United States. Phytopathology 34: 421-425.

Stakman, E. C , and W. Q. Loegering. 1951. Physiologic races of Puccinia graminis in the United States in 1950. U. S. Dept. Agr., Agr. Research Admin., and Minnesota Agr. Expt. Sta. Physiologic Races—1950.

Stakman, E. C , and F. J. Piemeisel. 1917. Biologic forms of Puccinia graminis on cereals and grasses. /. Agr. Research 10: 429-495.

Stakman, E. C , and H. A. Rodenhiser. 1958. Race 15B of wheat stem rust—what it is and what it means. Advances in Agron. 10: 143-165.

Stakman, E. C , F. J. Piemeisel, and Μ. N. Levine. 1918a. Plasticity of biologic forms of Puccinia graminis. J. Agr. Research 15 : 221-249.

Stakman, E. C , J. H. Parker, and F. J. Piemeisel. 1918b. Can biologic forms of stem rust on wheat change rapidly enough to interfere with breeding for rust resistance? /. Agr. Research 14: 111-123.

Stewart, D. M., and R. U. Cotter. 1958. A new and virulent culture of oat stem rust.

Phytopathology 48 : 389-390.

Tisdale, W. H. 1917. Flax wilt; a study of the nature and inheritance of wilt resist­

ance. /. Agr. Research 11: 573-606.

Vaheeduddin, S. 1942. The pathogenicity and genetics of some sorghum smuts.

Minn. Univ. Agr, Expt. Sta. Tech. Bull. 154.

Varro, Marcus Terentius.—On Agriculture I. 1934. "De Re Rustica," English transl.

by W. D. Hooper. [Loeb Classical Library.] Harvard Univ. Press, Cambridge, Massachusetts.

Walker, J. C , and R. Smith. 1930. Effect of environmental factors upon the resistance of cabbage to yellows. /. Agr. Research 4 1 : 1015.

Ward, Η. M. 1903. Further observations on the brown rust of the bromes, Puccinia dispersa (Erikss.) and its adaptive parasitism. Ann. Mycol. 1 : 132-151.

Waterhouse, W. L. 1929. Australian rust studies I. Proc. Linnean Soc. N. S. Wales 5 4 : 615-680.

Wellman, F. L., and D. J. Blaisdell. 1940. Differences in growth characters and pathogenicity of Fusarium wilt isolates tested on three tomato varieties. U. S.

Dept. Agr. Tech. Bull. 7 0 5 .

Wellman, F. L., and D. J. Blaisdell. 1941. Pathogenic and cultural variation among single spore isolates from strains of the tomato-wilt Fusarium. Phytopathology 3 1 : 103-120.

Welsh, J. N., and T. Johnson. 1954. Inheritance of reaction to race 7A and other races of oat stem rust, Puccinia graminis avenae. Can. J. Botany 3 2 : 347-357.