ISSN 10227954, Russian Journal of Genetics, 2011, Vol. 47, No. 1, pp. 13–18. © Pleiades Publishing, Inc., 2011. Original Russian Text © N.V. Trubacheeva, L.P. Rosseeva, I.A. Belan, T.S. Osadchaya, L.A. Kravtsova, Yu.V. Kolmakov, N.P. Blokhina, L.A. Pershina, 2011, published in Genetika, 2011, Vol. 47, No. 1, pp. 18–24.
PLANT GENETICS
Characteristics of Common Wheat Cultivars of West Siberia Carrying the Wheat–Rye 1RS.1BL Translocation N. V. Trubacheevaa, L. P. Rosseevab, I. A. Belanb, T. S. Osadchayaa, L. A. Kravtsovaa, Yu. V. Kolmakovb, N. P. Blokhinab, and L. A. Pershinaa,c a Institute
of Cytology and Genetics, Russian Academy of Sciences, Novosibirsk, 630090 Russia; email:
[email protected] b Siberian Research Institute of Agriculture, Russian Academy of Agricultural Sciences, Omsk, 644012 Russia; email:
[email protected] c Novosibirsk State University, Department of Cytology and Genetics, Novosibirsk, 630090 Russia Received January 10, 2010
Abstract—Using genomic in situ hybidization, among the common wheat cultivars produced in West Siberia (Siberian Research Institute of Agriculture, Omsk) with the involvement of the winter wheat cultivar Kavkaz carrying the wheat–rye 1RS.1BL translocation we identified three cultivars with this translocation: Omskaya 29, Omskaya 37, and Omskaya 38. The protein and crude gluten contents in the grain of these cultivars are equal to or exceed the levels observed in cultivars without the wheat–rye translocation. The common wheat culti vars carrying the wheat–rye translocation were evaluated in terms of resistance of plants reaching wax ripe ness to leaf rust and powdery mildew in the natural field conditions. The cultivars Omskaya 37 and Omskaya 38 displayed a high field resistance to leaf rust and were resistant to a variable extent to powdery mildew. The cultivar Omskaya 29 was susceptible to leaf rust and powdery mildew pathogens. Importance of the selection direction and the role of the genetic background in developing common wheat cultivars carrying the wheat– rye translocation is discussed. DOI: 10.1134/S1022795411010157
INTRODUCTION
reason, the rye chromosome 1R and its short arm 1RS are able to completely compensate the absence of homeologous wheat chromosomes in wheat–rye sub stitution and translocation forms and provide cytoge netic stability and fertility of plants [13].
Improvement of the common wheat characteristics has been attempted for a long time through introgres sive hybridization between wheat and rye (Secale cere ale L.). Rye chromosomes are able to substitute homeologous wheat chromosomes with the formation of wheat–rye substitution and translocation lines, which takes place as a result of backcrossing of wheat– rye hybrids and triticale with common wheat [1, 2]. Common wheat cultivars of different origins carry ing rye chromosomes contain mainly the rye chromo some 1R in which the short arm is represented in the wheat–rye translocations 1RS.1BL or 1RS.1AL, and the whole chromosome in the wheat–rye substitution 1R(1B) [3–10]. According to R. Schlegel, there are known over 650 common wheat cultivars with the 1RS.1BL translocation or 1R(1B) substitution (the substitution being present only in approximately 20 cultivars) [11]. The predominant participation of the rye chromo some 1R in the formation of substitution and translo cation common wheat forms seems to be determined by the fact that this chromosome, in contrast to other rye chromosomes, was not involved in the evolution in multiple interchromosomal rearrangements and has preserved complete homeology with respect to homeologous chromosomes of wheat [12]. For this
Most frequent in common wheat cultivars is the wheat–rye translocation 1RS.1BL of which the short arm was introgressed from the winter rye cultivar Pet kus [5, 6]. The short arm of the rye chromosome 1R contains genes controlling resistance of plants to pathogenic fungi: leaf rust (Lr26), stem rust (Sr31), yellow rust (Yr9), and powdery mildew (Pm8) [14, 15]. At the same time, in some common wheat cultivars with the 1RS.1BL translocation the expression of the Pm8 gene is suppressed [16, 17]. This is explained by the influence of a dominant suppressor gene localized on the common wheat chromosome 7D and distrib uted with unequal frequencies in different geographic zones [18–20]. Depending on the genetic background, wheat cul tivars carrying the 1RS.1BL translocation can be char acterized by drought resistance [21], increased grain yield, and an increase of general biomass [22]. It turned out that the poor breadmaking quality in dif ferent wheat cultivars with the 1RS.1BL translocation is also essentially determined by the influence of the genetic background [23, 24]. 13
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Table 1. Pedigrees of the common wheat cultivars produced in the Siberian Research Institute of Agriculture of the Russian Academy of Agricultural Sciences with the involvement of the winter wheat cultivar Kavkaz Cultivar, main biological characteristic Omskaya 20, midripening Omskaya 29, midripening Omskaya 30, midlate Omskaya 33, midripening Omskaya 35, midlate Omskaya 37, midlate Omskaya 38, midripening Kazanskaya Yubileinaya, midripening
Pedigrees Skala/Saratovskaya 36/3/Grekum 114//Kavkaz Druzhina//Grekum 114/Kavkaz/4/Skala/Saratovskaya 36/3/Grekum 114//Kavkaz Omskaya 20/3/Druzhina//Grekum 114/Kavkaz Omskaya 20/3/Druzhina//Grekum 114/Kavkaz/4/Omskaya 28 Omskaya 29/Omskaya 30 Kavkaz/Grekum 114//Venets/3/Burgas/4/Taifun/5/Omskaya 20/Omskaya 24 Kavkaz/Grekum 114//Venets/3/Burgas/4/Taifun/5/Omskaya 20/Omskaya 24 Omskaya 20/3/Druzhina//Grekum 114/Kavkaz/4/Lutescence 3/886
Note: /, first cross; //, second cross; the numbers of all subsequent crosses are given between single oblique strokes.
Among common wheat cultivars of domestic ori gin, the translocation 1RS.1BL is present in the winter wheat cultivar Kavkaz, which was obtained in 1972 by Academician P.P. Lukyanenko with the involvement of the substitution line 1R(1B) Neuzucht 14/14 [25]. The cultivar Kavkaz was widely used to produce new common wheat cultivars in many countries of the world and appears in the pedigree of more than 80 cul tivars with an introgressed 1RS.1BL translocation [11]. The cultivar Kavkaz is also present in the pedi gree of a number of common wheat cultivars produced in the Siberian Research Institute of Agriculture of the Russian Academy of Agricultural Sciences (Omsk) [26]. However, no studies have been carried out for finding among them cultivars with the wheat–rye translocation. The tasks of this work were as follows: (1) to reveal, using genomic in situ hybridization, the presence of the wheat–rye translocation among the common spring wheat cultivars produced by breeders from Omsk with the involvement of the wheat cultivar Kavkaz and (2) to study the manifestation of a number of characters whose expression in common wheat cul tivars can be influenced by the wheat–rye transloca tion 1RS.1BL. MATERIALS AND METHODS Material. The common wheat cultivars Omskaya 20, Omskaya 29, Omskaya 30, Omskaya 33, Omskaya 35, Omskaya 37, Omskaya 38, and Kazanskaya Yubilein aya were studied. Their pedigrees include the winter wheat cultivar Kavkaz (Table 1). Genomic in situ hybridization (GISH) was carried out according to the previously described technique [27]. The probe for genomic in situ hybridization was biotinlabeled total DNA isolated from S. cereale plants. No less than ten plants were analyzed for each cultivar. Analysis of the characteristics of the wheat cultivars. The wheat cultivars were grown on the fields of the
Siberian Research Institute of Agriculture (southern forest–steppe) in 2002–2008 on four 10 m2 plots according to recommendations [28]. The cultivars were analyzed for grain yield per 1 ha, 1000kernel mass, protein and gluten contents in grain, and loaf volume as described [29]. The data obtained were sta tistically analyzed [30]. The common wheat cultivars were evaluated in the natural field conditions for resis tance of plants at the stage of wax ripeness to leaf rust (Puccinia recondite f. sp.tritici) and powdery mildew (Blumeria graminis f. sp. tritici) in accordance to the methods adopted for estimating the resistance of cere als to rust diseases and powdery mildew [31]. The results of this evaluation are presented in the given work only for the common wheat cultivars carrying the wheat–rye translocation. RESULTS AND DISCUSSION Results of GISH Analysis The presence of the rye chromosome 1R in wheat– rye substitution genotypes of commom wheat or of the short arm of this chromosome in the translocations 1RS.1AL, 1RS.1BL, and 1RS.1DL in wheat is detected, depending on the tasks of the study, with the use of biochemical, molecular, cytogenetic, and molecular cytogenetic technologies [4, 5, 9, 10, 32]. Knowing the origin of the 1RS.1BL translocation, which is present in one of the parental cultivars (winter wheat cultivar Kavkaz), it was sufficient in our work to use genomic in situ hybridization (GISH). The GISH analysis makes it possible to identify the short arm of the chromosome in the translocation and thus to reveal the presence or absence of this translocation in plants [10, 32]. The presence of the wheat–rye translocation was established in three of the studied cultivars: Omskaya 29, Omskaya 37, and Omskaya 38. Each of the analyzed plants of these cultivars was found to have the short arm of the rye (S. cereale) chromosome 1R in the pair of translocated chromosomes (figure). Since the
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CHARACTERISTICS OF COMMON WHEAT CULTIVARS
Kavkaz cultivar carries the wheat–rye translocation 1RS.1BL, the abovementioned cultivars obtained with its involvement also have this type of transloca tion. As for plants of the cultivars Omskaya 20, Omskaya 30, Omskaya 33, Omskaya 35, and Kazanskaya Yubilein aya analyzed by GISH, no presence of rye chromo somes or their segments was detected. All chromo somes (2n = 42) in these cultivars belong to common wheat. These data show that in not all cases the wheat–rye translocation was transmitted to the new wheat culti vars. For instance, the wheat–rye translocation was not introgressed from the Kavkaz cultivar into the cul tivars Omskaya 20, Omskaya 30, Omskaya 33, and Kazanskaya Yubileinaya and from Omskaya 29 into Omskaya 35. Analogous results were obtained by some other authors. For example, about 55% of the common wheat cultivars registered during the last twenty years in Hungary have the 1RS.1BL translocation [21]. Among six winter cultivars of common wheat obtained with the use of the Magdalena cultivar carrying 1RS.1B, this translocation passed only to one cultivar, Mv Walzer. In the other five cultivars, differential staining of chromosomes revealed no segments of the rye chromosome [8]. The loss of the translocated chromosome can be explained by the fact that in plant genotypes heterozy gous for the translocation the frequency of gametes with the translocation is reduced as is the fertilization capacity of male gametes carrying the wheat–rye translocation [23]. In addition, it is possible that inter varietal hybrid genotypes with a substitution of the translocated 1RS.1BL chromosome by a pair of 1B common wheat chromosomes had an advantage at certain stages of selection for adaptability. It should be noted that among the common wheat cultivars studied in our work two cultivars, midlate Omskaya 37 and midripening Omskaya 38, carrying the wheat–rye translocation are sister lines selected from one combination (Table 1). A specific feature of the origin of these cultivars is that the cultivar Kavkaz was used as the maternal form. The remaining culti vars studied in this work were obtained with the involvement of the cultivar Kavkaz in crosses as the paternal form. Analysis of Grain Quality and Yield Table 2 summarizes the data of 2002–2008 on the grain quality and yield in the common wheat cultivars produced with the use of the winter wheat Kavkas. These parameters in the wheat cultivars carrying the wheat–rye translocation and in the cultivars lacking this translocation were compared within two groups differing in the length of the vegetation period. RUSSIAN JOURNAL OF GENETICS
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An example of genomic in situ hybridization (GISH) for the common wheat cultivar Omskaya 29. The arrows indi cate the wheat–rye translocation.
The group of midripening cultivars includes the cultivars Omskaya 29 and Omskaya 38 carrying the wheat–rye translocation. These cultivars did not sig nificantly differ in such characteristics as grain yield, 1000kernel mass, and loaf volume from the midrip ening cultivars Omskaya 20, Omskaya 33, and Kazan skaya Yubileinaya in which the translocation is absent (Table 2). No differences in the protein and crude glu ten contents in grain were discovered between the common wheat cultivars with the translocation (Omskaya 29 and Omskaya 38) and the common wheat cultivars Omskaya 20 and Kazanskaya Yubilein aya. Yet, the levels of protein and crude gluten in grain are significantly higher in the cultivar Omskaya 38 than in Omskaya 33 (Table 2). A significantly higher content of gluten was revealed in Omskaya 29 as com pared to that in Omskaya 33. In the group of midlate cultivars, the common wheat cultivar Omskaya 37 carrying the wheat–rye translocation did not differ in all characteristics from the cultivar Omskaya 30 that does not have this trans location. However, in the common wheat cultivar Omskaya 37 the levels of protein and gluten in grain proved to be considerably higher than in another mid late common wheat cultivar, Omskaya 35, in which the wheat–rye translocation is absent (Table 2). Thus, it has been established that in the studied common wheat cultivars carrying the wheat–rye translocation such characteristics of grain quality as the contents of protein and crude gluten are similar to or exceed their levels in the cultivars without this translocation. The compared wheat cultivars displayed no differences in other characteristics of grain quality and in the yield. These results are in conformity with the data of other authors who reported that the wheat– rye translocation 1RS.1BL could favor an increase of the content of grain protein and did not essentially affect the grain yield and the mass of 1000 kernels [33]. 2011
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Table 2. Yield and quality of grain of the common wheat cultivars produced with the involvement of the winter wheat cultivar Kavkaz (data for 2002–2008) Average values of the characteristics and their variation Cultivars
1000kernel mass, g Grain protein content, % Grain crude gluten content, % Loaf volume, cm3 Grain yield, t/ha
Midripening cultivars Omskaya 29 Omskaya 38 Omskaya 20 Omskaya 33 Kazanskaya Yubileinaya Midlate cultivars Omskaya 37 Omskaya 30 Omskaya 35
42.1 ± 0.8 39.0–44.8 39.1 ± 1.5 32.5–45.8 37.2 ± 1.9 26.4–44.2 41.2 ± 1.2 37.6–46.2 39.6 ± 1.7 32.5–45.6
15.81 ± 0.26 14.59–16.62 16.19 ± 0.31* 14.93–17.36 16.05 ± 0.27 15.00–17.33 15.24 ± 0.17 14.68–15.96 15.77 ± 0.26 15.0–16.82
31.24 ± 0.58* 28.6–32.2 31.69 ± 0.75* 28.2–34.4 31.86 ± 0.49 30.40–34.42 29.67 ± 0.32 28.6–31.0 31.29 ± 0.65 29.3–33.7
1038 ± 23 930–1100 1002 ± 62 735–1170 993 ± 31 905–1105 954 ± 23 875–1020 952 ± 34 850–1100
3.73 ± 0.45 2.69–6.23 3.77 ± 0.40 2.60–5.97 3.51 ± 0.46 2.15–6.00 4.26 ± 0.43 3.01–6.51 4.02 ± 0.39 2.96–6.13
37.2 ± 1.6 30.6–44.6 35.9 ± 1.3 29.6–39.0 39.1 ± 2.0 31.0–45.8
17.18 ± 0.20*** 16.53–18.13 16.53 ± 0.30 15.62–17.84 16.00 ± 0.21 15.13–16.84
33.54 ± 0.19*** 32.9–34.4 33.19 ± 0.45 31.5–34.7 31.71 ± 0.34 30.2–33.2
1006 ± 44 775–1130 1011 ± 14 960–1060 977 ± 32 855–1115
3.80 ± 0.37 2.86–5.83 3.42 ± 0.42 2.08–5.25 4.13 ± 0.53 2.96–6.13
Note: The difference from the cultivar Omskaya 33 is significant at * P < 0.05. The difference from the cultivar Omkaya 35 is significant at *** P < 0.001.
Analysis of Resistance of Common Wheat Cultivars Carrying the Wheat–Rye Translocation to Leaf Rust and Powdery Mildew A massive spreading of the leaf rust pathogen (Puc cinia recondite f. sp. tritici) was noted in 2004 and in 2007. The common wheat cultivars studied carrying the wheat–rye translocation displayed different resis tances to leaf rust. For example, in all years of investi gation, including the years of massive spreading of the Table 3. Leaf rust resistance of common wheat cultivars carrying the wheat–rye translocation (2002–2008) Resistance to Puccinia recondite f. sp. tritici, score Cultivar Omskaya 29 Omskaya 38 Omskaya 37 LSD05
2002 2003 2004* 2005 2006# 2007* 2008# 4 9 9 0.71
6 9 9 0.95
4 8 8 0.89
3 9 9 0.76
– – –
2 9 8 0.79
– – –
Note: * The years of massive spreading of the leaf rust pathogen. # The years when no massive spreading of the leaf rust pathogen was observed (data not shown). 9, immune (very high resistance); 8, highly resistant; 6, resistant; 4–3, sus ceptible; 2, highly susceptible.
pathogen, plants of the Omskaya 37 and Omskaya 38 cultivars showed either a high (score 8) or very high (score 9) field resistance to the leaf rust pathotypes (Table 3). Regarding the cultivar Omskaya 29 carrying the wheat–rye translocation, by the period of wax ripeness it was susceptible (2002, 2004, 2005) or highly susceptible (2007) to the leaf rust pathogen. One of the mechanisms underlying the resistance of the wheat cultivars Omskaya 37 and Omskaya 38 to leaf rust may be associated with the action of the Lr26 gene localized on the short arm 1RS of the rye chro mosome [14, 15], which is present in the wheat–rye translocation in these cultivars. At the same time, the action of this gene is not expressed in the cultivar Omskaya 29 carrying an analogous wheat–rye translo cation. This result may be due to the specific features of the genetic background of this wheat cultivar, which differs in origin from the cultivars Omskaya 37 and Omskaya 38 (Table 1). The estimation of the resistance to powdery mildew in the common wheat cultivars carrying the wheat–rye translocation revealed differences between the cultivar Omskaya 29 and the cultivars Omskaya 37 and Omskaya 38 (Table 4). A mass spreading of the pow dery mildew pathogen (Blumeria graminis f. sp. tritici) was observed in 2002, 2004, 2007, and 2008. In all
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CHARACTERISTICS OF COMMON WHEAT CULTIVARS Table 4. Powdery mildew resistance of common wheat cultivars carrying the wheat–rye translocation (2002–2008) Resistance to Blumeria graminis f. sp. tritici, score Cultivar 2002* 2003 2004* 2005 2006 2007* 2008* Omskaya 29 Omskaya 38 Omskaya 37 LSD05
2 6 6 1.1
4 6 6 0.80
3 8 8 0.64
4 6 6 0.80
5 8 8 1.2
3 6 7 0.98
3 7 5 1.35
Note: * The years of mass spreading of the powdery mildew patho gen. 8, resistant; 7–6, midresistant; 5, midsusceptible; 4– 3, susceptible; 2, highly susceptible.
years of the study, the cultivar Omskaya 29 was suscepti ble to powdery mildew: from high susceptibility in 2002 (score 2) to medium susceptibility in 2006 (score 5). Distinct results were obtained for the cultivars Omskaya 37 and Omskaya 38. In particular, Omskaya 37 displayed a medium susceptibility to powdery mildew (score 5) in 2008. In the other years, the cultivars Omskaya 37 and Omskaya 38 were resistant (scores 9–8) or midresistant (scores 7–6) to the pathogen of this fungus (Table 4). It can be assumed that in the common wheat culti var Omskaya 29, in contrast to the cultivars Omskaya 37 and 38, the Pm8 gene localized on the short arm 1RS of the rye chromosome is suppressed [14, 15], as it was described by other authors for a number of wheat cul tivars carrying the wheat–rye translocation 1RS.1BL [18–20]. It was noted above that the midlate cultivar Omskaya 37 and the midripening cultivar Omskaya 38 differing in the length of the vegetation period are of common origin (Table 1). As follows from the charac teristics of these cultivars presented above, the selec tion on grain quality and resistance to the leaf patho gens of midripening and midlate genotypes from their common form was of the same direction. The differences detected between the common wheat cultivar Omskaya 29 carrying the wheat–rye translocation and the cultivars Omskaya 37 and Omskaya 38 also suggest a powerful influence of the genetic background on the manifestation of characters whose expression depends on the presence of the wheat–rye translocation. Moreover, in the cultivars Omskaya 37 and Omskaya 38 the genetic background was favorable for realization of the genetic potential of the wheat–rye 1RS.1BL translocation acquired by these cultivars from the winter wheat cultivar Kavkaz. These data are in agreement with the results of the works [21–24, 33] demonstraing the importance of the genetic background in creating new common wheat cultivars carrying this translocation. RUSSIAN JOURNAL OF GENETICS
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ACKNOWLEDGMENTS The work was supported by the Russian Founda tion for Basic Research (project no. 080000598), Complex Integration Project of the Siberian Branch of the Russian Academy of Sciences, Program of the Presidium of the Russian Academy of Sciences “Biodiversity and Gene Pool Dynamics”, and the Ministry of Education and Science of the Russian Federation (s/c 02.740.11.0277). REFERENCES 1. Lukaszewski, A.J., Gustafson, J.P., and Apolinarska, B., Transmission of Chromosomes through the Eggs and Pollen of Triticale Wheat F1 Hybrids, Theor. Appl. Genet., 1982, vol. 63, pp. 49–55. 2. Shchapova, A.I. and Kravtsova, L.A., Tsitogenetika pshenichnorzhanykh gibridov (Cytogenetics of Wheat– Rye Hybrids), Novosibirsk: Nauka, 1990. 3. Lukaszewski, A., Frequency of 1RS.1AL and 1RS.1BL Translocations in Unated States Wheat, Crop Sci., 1990, vol. 30, pp. 1151–1153. 4. Gupta, R.B. and Shepherd, K.W., Identification of Rye Chromosome 1R Translocations and Substitutions in Hexaploid Wheats Using Storage Proteins as Genetic Markers, Plant Breed., 1992, vol. 109, pp. 130–140. 5. Schlegel, R. and Korzun, V., About the Origin of 1RS.1BL Wheat–Rye Chromosome Translocations from Germany, Plant Breed., 1997, vol. 116, pp. 537– 540. 6. Rabinovich, S.V., Importance of Wheat–Rye Translo cations for Breeding Modern Cultivars of Triticum aes tivum L., Euphytica, 1998, vol. 100, pp. 323–340. 7. Ko, J.M., Seo, B.B., Suh, D.Y., et al., Production of New Wheat Line Possessing the 1BL.1RS Wheat–Rye Translocation Derived from Korean Rye Cultivar Paldanghomil, Theor. Appl. Genet., 2002, vol. 104, pp. 171–176. 8. Szakacs, E., Linc, G., Lang, L., and MolnarLang, M., Detection of the 1A/1R and 1B/1R Wheat/Rye Trans location in New Martonvasar Wheat Cultivars and Advanced Lines Using in situ Hybridization, Növeny termeles, 2004, vol. 53, pp. 527–534. 9. Landjeva, S., Korzun, V., and Tsanev, V., Distribution of Wheat–Rye Translocation 1RS.1BL among Bread Wheat Cultivars of Bulgaria, Plant Breed., 2006, vol. 125, pp. 102–104. 10. Tang, Z.X., Fu, S.L., Ren, Z.L., et al., Production of New Wheat Cultivar with a Different 1B.1R Transloca tion with Resistance to Powdery Mildew and Stripe Rust, Cereal Res. Commun., 2008, vol. 36, pp. 451–460. 11. Schlegel, R., Current List of Wheats with Rye and Alien Introgression, V0508, pp. 1–14, http://www.des icca.de/Wheatrye introgression. 12. Devos, K.M., Atkinson, M.D., Chinoy, C.N., et al., Chromosomal Rearrangements in the Rye Genome Relative to That Wheat, Theor. Appl. Genet., 1993, vol. 85, pp. 673–680. 13. Friebe, B., Jiang, J., Raupp, W.J., et al., Characteriza tion of WheatAlien Translocations Conferring Resis 2011
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