ISSN 00954527, Cytology and Genetics, 2015, Vol. 49, No. 1, pp. 13–19. © Allerton Press, Inc., 2015. Original Ukrainian Text © A.V. Galaev, Yu.M. Sivolap, 2015, published in Tsitologiya i Genetika, 2013, Vol. 49, No. 1, pp. 18–25.
Description of the Soft Wheat Varieties of Ukrainian and Russian Breeding by Alleles of Locus csLV34 Closely Connected with Multipathogen Resistance Gene Lr34/Yr18/Pm38 A. V. Galaev and Yu. M. Sivolap† Plant Breeding and Genetics Institute, National Center of Seed and Cultivar Investigations, National Academy of Agricultural Sciences, ul. Ovidiopolska 3, Odessa, 65036 Ukraine email:
[email protected] Received November 2, 2012
Abstract—Codominant moleculargenetic marker csLV3 was used to examine a collection of soft wheat vari eties created in the main plant breeding centers of Ukraine and Russia from 1912 to 2011. Two allelic variants csLV34a and csLV34b associated, respectively, with the absence and presence of gene Lr34/Yr18/Pm38 were revealed. Allele csLV34b, commonly found among varieties of Ukrainian selection, is especially common in varieties of the Plant Breeding and Genetics Institute (Odessa). The results can be used in breeding programs to select wheat varieties with gene Lr34/Yr18/Pm38 for use in crossing with varieties carrying the race non specific genes and develop new varieties with better resistance to fungal diseases. Keywords: soft wheat, genes resistance to fungal diseases, moleculargenetic marker, PCR DOI: 10.3103/S0095452715010041
INTRODUCTION
ated with tolerance to barley yellow draft virus Bdv1 [8, 9] and flag leaf tip necrosis Ltn1 [9].
Wheat damages with pathogenic fungi have become critical factors of harvest losses and reducing of its quality due to mycotoxins. Selection of plant pathogen resistant varieties of wheat is an effective method of control over fungal diseases. At present, sci entists have identified almost 67 brown leaf rust resis tant genes, 49 stripe rust resistant genes, and 45 mil dew resistant genes [1] that are used in plant breeding programs all over the world. Most of these genes are genus specific and they become less efficient if new virulent genera appear. Genus specific resistance ensures high efficiency protection, but resistance that is not based on specific recognition between host and pathogen is often more stable and prolonged. Genus specific resistance is not characterized by supersensi tivity but with partial sensitivity connected with long latent period of disease development [2, 3]. We con sider that mildew resistant loci Lr34/Yr18/Pm38 (located on chromosome 7DS) and Lr46/Y29/Pm39 (located on chromosome 1BL) are the most impor tant, since they allow one to gain simultaneously genus specific and prolonged resistance to three of the most important biotrophic wheat pathogens that are always inherited together as one Mendelian character [4, 5]. Locus Lr/Y18/Pm38, combined with other genus spe cific genera, ensures a high resistance level for many decades [6, 7]. The locus mentioned above is associ
As genes of multipathogenic resistance locus 1 Lr34/Yr18/Pm38 are expressed mostly at the stage of adult plant and can be concealed by other genes, there is a considerable interest in development of efficient detection methods. Singh et al. [9] used phenotype display of gene Ltn1 for detection of gene Lr34, but multigene influence on flag leaf tip necrosis and unequal Ltn1 expression in different conditions can cause controversial results. Xgwm295 and Xgwm1220 were the first moleculargenetic markers associated with Lr34/Yr18/Ltn1, but they were not widely used because of their low marking ability in different wheat varieties. Two new microsatellite markers Xcsm10 [10] and csLVMS1 [11] showed high diagnostic ability for multipathogenic resistance locus Lr34/Yr18/Pm38 in 1 wheat varieties of different origin. While using these markers, it is difficult to differentiate stable haplotype from a sensitive one due to the subtle difference between alleles 206 and 208 bp long by marker Xcs10 and 224 and 226 bp long by marker csLVMS1, corre spondingly. Moleculargenetic marker csLV34 is widely used for detection of multipathogenic resistance locus 1 Lr34/Yr18/Pm38 [18]. Codominant marker csLV34 bound to locus Lr34/Yr18/Pm38 is intensively used for detection of gene Lr34 in wheat varieties from Austra lia [13], the United States, Great Britain [14], India [15], Canada [16], and the world collection [14]. But
† Deceased.
13
14
GALAEV, SIVOLAP csLV34F
csLV34R a
5'
3' b Fig. 1. Schematic image of gene of sulfatetransporter of wheat with the section inside intron 4 limited with pair of primers csLV34F and csLV34R [12]. Alleles csLV34a and csLV34b differ in nucleotide deletion, which is 79 bp in size. Exons are marked with black rectangles.
this marker is not diagnostic for some wheat genotypes from Canada and the United States that require Cana dian line RL4137 [16, 17]. Krattinger et al. [18] identified gene ABCtrans porter as the only candidate for all three resistant genes using chemical and physical mutagenesis. Initial anal ysis of ABCtransporter sequence of locus Lr34/Yr18/Pm38 discovered two main haplotypes. Resistant variety Chinese Spring (Lr34+) and sensi tive variety Renan (Lr34+) differ in three mutations inside ABCtransporter sequence. We identified two onenucleotide substitutions in intron 4 (A/T) and exon 12 (C/T) and one threenucleotide deletion in exon 11 (Del/TTC) [18]. Stable haplotype A/Del/C was marked as Lr34/Yr18a and sensitive haplotype T/TTC/T was marked as Lr34/Yr18b. Then we identi fied five more sensitive haplotypes that are characterized by different mutations in DNA sequence of ABCtrans porter gene: onenucleotide substitutions in exon 22 (G/T) (haplotype Lr34/Yr18d) [17, 19], exon 12 (C/T) (Lr34/Yr18g)[20], introns 4 and 6 (Lr34/Yr18e) [19] and onenucleotide deletions in intron 4 (A/–) (Lr34/Yr18c) [17] and exon 10 (A/–) (Lr34/Yr18f) [20]. Karelov et al. [21] detected allele state of locus Lr34 in 81 varieties of Ukrainian selection on the basis of findings of Evans et al. [22]. This concerned the usage of two pairs of allelespecific primers according to the analysis of deletion in exon 11. The authors showed that stable allele Lr34+ was found in 44% of all analyzed varieties. The aim of our research was to analyze the allele state of locus csLV34 in 260 winter soft wheat varieties of different geographical origin in order to detect gene Lr34/Yr18/Pm38 and also to determine the origin of this gene in existing wheat varieties. MATERIALS AND METHODS 253 soft wheat varieties from working collection of genetics and genomics departments of the Plant Breeding and Genetics Institute of the National Cen ter of Seed and Cultivar Investigations were used in our study. There were 214 varieties of different plant breeding centers of Ukraine and 39 varieties from Rus sia. We used varieties Chinese Spring and Farandol – bearers of gene Lr34 for positive control, and variety Renan and varieties with genes Lr9 (Transfer), Lr19
(Agatha), Lr24 (Agent), and Lr37 (VPM1) for nega tive control. DNA was isolated from 5day sprouts using CTAB buffer [23]. We studied DNA of five individual plants of each variety. To detect gene Lr34/Yr18/Pm38 among 260 soft wheat varieties, we used codominant marker csLV34 with sulfatetransporter gene bound with ABCtransporter. The latter ensures multipatho 1 genic resistance. This marker pointed to two alleles: csLV34a 229 bp long and csLV34b 150 bp long. They are associated with the presence or absence of gene Lr34/Yr18/Pm38, correspondingly [12, 14]. These alleles differ in nucleotide deletion of 79 bp in size in the fourth intron of sulfatetransporter gene (Fig. 1). PCR with directed primes to locus csLV34 was carried out on the Tertsik thermocycler (DNAtechnology, Russian Federation). Reaction mixture with a volume of 25 µL contained buffer (67 mM of trisHCl pH 8.8; 16.6 mM of (NH4)2SO4; 1.5 mM of MgCl2; 0.03% Tween20); 0.2 mM of each dNTP; 0.25 µM of primer; 20 ng of DNK; 0.8 ea of Taqpolymerase. We put 30 µL of mineral oil over the reaction mixture. The conditions of reaction are the following: denaturation at 94°C during 30 s (initial denaturation of 2 min), annealing at 55°C for 30 s, elongation at 72°C for 1 min (final elongation of 4 min). Amplification prod ucts (10 µL of aliquot of PCRmixture) were frac tioned in 90% polyacrylamide gel in 1 × TBE. We con ducted electrophoresis in polyacrylamide gel at per manent voltage of 500 V in the device for vertical gel electrophoresis (Hoefer Scientific Instruments, United States). Visualization of electrophoresis distri bution products was conducted by gel impregnation using argentic nitrate. Video image and sizes of ampli fied fragments were obtained by means of ImageMas ter VDS video system (Amersham Pharmacia Biotech, United States) according to the manufacturer’s guide lines. Molecular weight of obtained amplicons was calibrated using the standard pUC19.MspI and 10 bp DNA Ladder. Statistical processing of data was carried out according to generally accepted techniques [24]. RESULTS AND DISCUSSIONS Origin of gene Lr34/Yr18/Pm38 in wheat germ plasma. According to literature data, allele csLV34b associated with multipathogenic resistance gene 1 Lr34/Yr18/Pm38 originates from old Italian popula tion variety Rieti. In 1907 Italian selectionist Naza reno Strampelli created wheat varieties Mentana, Ardito, Ballila, Villa Glori, etc. by means of inter breeding of resistant pure wheat strain taken from vari ety Rieti, highyield Dutch variety Wilhelmina, and precocious lodging resistant Japanese variety Akago mughi. Selectionists from many countries of the world used newly created varieties during interbreedings [25]. For example, using variety Mentana (bearer of allele csLV34b) new variety Frontana was obtained. Through Fontana allele csLV34b appeared in germ CYTOLOGY AND GENETICS
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DESCRIPTION OF THE SOFT WHEAT VARIETIES
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Table 1. Analysis of allele state of locus csLV34 in 26 old Ukrainian and Russian varieties of soft wheat (enumeration according to date of creation) Variety
Locus alleles csLV34 (a, b)
Genus
Variety
1912–1933 Years
Locus alleles csLV34 (a, b)
Genus
Ulyanivka
a
Selection from variety Bilokosa bezosta
Kooperatorka
a
Selection from Krymka
Lisostepka 75
a
Ukrainka × Lisostepka 74
Krymka mistseva
a
Old variety
Odeska 12
a
Zemka × Gostianum 237
Banatka
a
Old variety of Hungary Artemivka
a
Selection of variety Mistseva
Zemka
a
Selection of variety Mistseva
Ukrainka
a
Selection of variety Banatka
Odeska 16
a
Selection from variety Odeska 12
Ukrainka 0246
a
Selection of variety Banatka
Zenitka
a
Selection from variety Gostianum 237
Gostianum 237
a
Selection of variety Mistseva @
Veselopodolyan ska 499
a
Covail (United States) × Milturum 14
Giarka 0274
b
Selection of variety Mistseva Odeska
Bilotserkivska 198
a
Erytrospermum × Covail (United States)
Odeska 26
a
Odeska 3 × Lutestsens 17
a
Odeska 3 × Lutestsens 17
1934–1947 Years
1948–1959 Years
Odeska 3
a
Kooperatorka × Gos tianum 237
Odeska bezosta
Milturum pererod
a
Genus unknown
Stepova
Milturum 553
a
Milturum 321 × Kitch ener
Erytrospermum 15
a
Mistseva × Ukrainka
Lutestsens
a
Mistseva × Ukrainka
plasma of wheat varieties from the United States, Canada, and CIMMYT. Mentana was also introduced in other parts of the world, such as Turkey and China, where it was grown in the huge territories during pro longed period. Mostly the reason was their rust resis tance [26, 27]. Variety Ardito is a donor of allele csLV34b of European winter wheat varieties, including Bezosta 1 [14]. Thus, considering literature data on varieties origin, allele csLV34b was found among Ukrainian and Russian wheat varieties due to Bezosta 1. Since foreign authors used only varieties Bezosta 1 and Caucasus for detection of gene Lr34/Yr18/Pm38, the previous findings of Ukrainian and Russian variet 1 ies concerning multipathogenic resistance gene arouse interest. Table 1 shows that allele csLV34b of 150 bp long was found only in varieties Girka 0274, Stepova, Skorospelka 1, and Skorospelka 2. The presence of allele csLV34b associated with gene Lr34/Yr18/Pm38 in varieties Skorospelka 1 and Skorospelka 3b is obvi CYTOLOGY AND GENETICS
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Skorospelka
a, b
Bezosta × Odeska 16
b
CunderFulxter (United States) 266 287 × Clein (Argentina) 33
b
The same
ous, since these varieties were created by P. Luky anenko. It also concerns variety Skorospelka 2, which was created by means of hybridization and using variety Ardito. After interbreeding of varieties Lutestsens 17 and Skorospelka 2, we obtained new variety Bezosta 4, which was later used for creation of Bezosta 1. Popula tion variety Stepova contained two genotypes by locus csLV34aa and bb. At the same time, allele csLV34b was connected with variety Bezosta 4 (Table 1). We still know nothing about the origin of gene Lr34/Yr18/Pm38 in old Odessa slowmaturing spring wheat variety Girka 0274, which was created by A. Sapegin using the method of individual selection from local population varieties. Probably, this gene was present in the first population varieties that were brought to Ukraine by migrant farmers in the end of the 19th–the beginning of the 20th centuries. Data on rust resistance of variety Girka 0274 [28] confirms the presence of gene Lr34/Yr18/Pm38. Since variety
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GALAEV, SIVOLAP
Table 2. Distribution of alleles of locus csLV34 in varieties of Ukrainian selection Alleles
n
Genotype frequency p ± sp, %
Genus, strain North
a
19
52.7 ± 8.3
Kyianka, Kyivska ostysta, Zolotovolosa, Snigurka, Ekspromt, Kryzhynka, Favoritka, Bogdana, Myronivska 264, Myronivska 808, Myronivska yuvileina, Illichivka, Myronivska 33, Myrych, Myronivska 65, Murkhard, Myronivska ostysta, Smuglyanka, Ivanivska ostysta
b
13
36.2 ± 8.0
Veselka, Olesya, Perlyna lisostepu, Lybid, Yasochka, Tsyganka, Kyivska 8, Elegia, Vesnyanka, Volodarka, Khurtovyna, Podolyanka, Myronivska 27
4
11.1 ± 5.2
Bilotserkivska semidwarf, Yatran 60, Columbia, Myrleben
a+b
@ a
6
54.5 ± 15.0
Donetska 48, Bilosnizhka, Dar Luganshchyny, Napivkarlyk 1, Kharkivska 105, Astet luganskii
b
5
45.5 ± 15.0
Donetska semidwarf, Kharkivska 96, Kharus, Vasylyna, Apogei luganskii @
a
4
80.0 ± 17.9
Dykanka, Ukrainka poltavska, Kolomak 3, Kolomak 5
b
1
20.0 ± 17.9
Spivanka @
a
34
23.3 ± 3.5
Prybii, Odeska 66, Chaika, Pivdenna zorya, Peresvit, Odeska 133, Odeska 162, Odom, Vympel odeskii, Odeska 266, Odeska 267, Strumok, Khvylya, Luzanivka odeska, Lyubava odeska, Khyst, Oksana, Bezmezhna, Turunchuk, Krasen, Polyovyk, Epokha odeska, Vatazhok, Odeska 120, Prokofyevka, Chervona, Povaga, Dovira, Erytrospermum 2917, Odeska ostysta semiintensive, Shestopalivka, Rozmai, Khersonska ostysta, Knopa
b
86
58.9 ± 4.0
Rosynka, Erytrospermum 127, Brygantyna, @, Progres, Odeska redeared, Olvia, Odeska 132, Yunnat odeskii, Zolotava, Fedorivka, Odeska 161, Odeska 265, Symvol odeskii, Fantaziya odeska, Tira, Viktoria odeska, Zastava odeska, Lada odeska, Leleka, Lyutestsens 23397, Nikosia, Panna, Pryma odeska, Syrena odeska, Selyanka, Zustrich, Kuyalnyk, Poshana, Dalnytska, Vadala, Zemlyacha odeska, Pysanka, Suputnytsa, Gospodnya, Zmina, Skarbnytsya, Yednist, Zamozhnist, Zaporuka, Kosovytsya, Lytanivka, Blagodarka odeska, Bunchuk, Goduvalnytsya odeska, Misiya odeska, Sluzhnytsya odeska, Zhaivir, Zagrava odeska, Istyna, Vykhovanka odeska, Zadumka, Zvytyaga, Zysk, Princess Olha, Lad, Nasnaga, Nyva, Bryz, Burevisnyk odeskii, Lan, Fregat odeskii, Yakir odeskii, Zlagoda, Nagoroda odeska, Gurt, Zhuravka, Lebidka, Lira, Otaman, Driada 1, Svitanok 1, Albatross odeskii, Ukrainka odeska, Krasunya, Veteran, Vilen, Era, Lanovii, Melodiya, Sofiika, Chornobrova, Khersonska bezosta, Nakhodka 4, Khersonska 99, 5/5591
a+b
26
17.8 ± 3.1
Stepova, Odeska 51, Odeska semidwarf, Zirka, Prometei, Odeska 117, Yuvileina 75, Porada, Lelya, Znakhidka odeska, Kiriya, Liona, Antonivka, Dyuk, Podyaka, Golubka, Darunok odeskii, Bilyava, Borvii, Uzhynok, Odeska 130, Dobrochyn, Zorepad, Lastivka, Nebokrai, Pylypivka
Girka 0274 was not used in selection programs, it is not considered to be a protosource of gene Lr34/Yr18/Pm38 in varieties of Ukrainian and Rus sian selection. Allele state of locus csLV34 in varieties of Ukrainian and Russian selection. We analyzed the collection from
198 varieties of Ukrainian and 30 Russian selection which were created in main plant breeding centers from 1959 (year of creation of Bezosta 1) to 2011 (Tables 2, 3). According to the results of PCR, we found only two identical alleles along with a pair of primers to locus csLV34. These alleles are the follow ing csLV34a 229 bp long and csLV34b 150 bp in length CYTOLOGY AND GENETICS
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DESCRIPTION OF THE SOFT WHEAT VARIETIES
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Table 3. Distribution of alleles of locus csLV34 in varieties of Russian selection Alleles
n
Genotype frequency
Genus, strain Western Siberia and Volga region
a
11
64.7 ± 11.6
Zorya, Pivnichna zorya, Omska zima, Kazanska 285, Kazanska 237, Omska 2, Saratovska 25, Lutestsens 7, Albidum 114, Albidum 12, Bagrationivska
b
4
23.5 ± 10.3
Omska 3, Omska 4, Omska 5, Gostianum 274
a+b
2
11.8 ± 7.8
Gostianum 273, Sibirska niva Northern Caucasus
a
8
61.5 ± 13.5
Avrora, Skifyanka, Yuna, Batko, Pobeda 50, Krasnodarska 99, Donska, Donskii syurpriz
b
5
38.5 ± 13.5
Bezosta 1, Caucasus, Donsimb, Spartanka, Stanichna
(Fig. 2). Ambiguous allele csLV34c [14] was not dis covered.
odeska, Ukrainka odeska, and Selyanka. High fre quency of allele csLV34a associated with the absence of gene Lr34/Yr18/Pm38 in varieties from the north and center of Ukraine and western Siberia is con nected with active usage in selection programs of frost resistant varieties adjusted to these regions—bearers of the following alleles: Ulyanivka, Myronivska 808, and Bagrationivka.
In most Ukrainian and Russian wheat varieties, we found one of two alleles of locus csLV34: 87.6 and 93.3%, correspondingly. Heterogeneous to this gene varieties included genotypes aa and bb with different ratio of genotype frequencies. Allele csLV34b was mostly spread among varieties from the south of Ukraine (58.9 ± 4.0%). Allele csLV34a was widely spread among soft winter wheat varieties in the north and center of Ukraine (from 52.7 to 80.0%) (Table 2). In the east of Ukraine, frequencies of alleles csLV34a (54.5 ± 15.0%) and csLV34b (45.5 ± 15.0%) do not differ much. In Russian varieties of Siberia and Volga region, frequency of allele csLV34b is equal to 23.5 ± 10.3%, and that in the Northern Caucasus is 38.5 ± 13.5%. High frequency of allele csLV34b associated with gene Lr34/Yr18/Pm38 in varieties from the south of Ukraine is connected with active usage of varieties— bearers of the mentioned allele: Bezosta 1, Odeska 51, Odeska semidwarf, Zirka, Albatross odeskii, Viktoria
It is necessary to admit that gene Lr34/Yr18/Pm38 is not effective in Ukraine and Russia. Wheat plants bearing this gene are characterized by a high level of infectious ness with leaf rust in the range of 15–90% [7, 29, 30]. Expression of resistance gene Lr34/Yr18/Pm38 depends on temperature of environment and stage of plant onto genesis. The display of gene Lr34/Yr18/Pm38 was con ducted in field conditions in the range 0–20°C. When temperature increases, gene expression is inhibited [31]. Despite low efficiency of gene Lr34/Yr18/Pm38, a number of scientists found that its combination with other genus specific genes, such as Lr13, Lr22a, Lr26, Lr35, and Lr37, increases the level of field resistance
csLV34a allele
csLV34b allele
1 2
3
4
5
6
M
7
8
9
10
11
12
13
Fig. 2. Electrophorogram of products of amplification of DNA of soft wheat varieties by locus csLV34: (M) marker of molecular weight pUC19 DNA/MspI; (1) negative control, variety Renan; (2) positive control, variety Chinese Spring—bearer of gene Lr34; (3, 4) Perlyna lisostepu; (5, 6) Albatross odeskii; (7, 8) Myronivska 808; (9–11) Odeska semidwarf; (12, 13) Selyanka. CYTOLOGY AND GENETICS
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GALAEV, SIVOLAP
Table 4. Distribution of soft winter wheat varieties of the Plant Breeding and Genetics Institute of the National Center of Seed and Cultivar Investigations by selection stages [33] and a number of studied varieties by locus csLV34 Selection Years stage of cultivation II III IV V VI
1960–1967 1968–1975 1976–1990 1991–1996 1997–2004
VII
2005
Odeska bezosta, Stepova Prybii, Odeska 51 Odeska semidwarf, Obrii, Zirka, Odeska redeared, Yunat odeskii Albatross odeskii, Ukrainka odeska, Fedorivka Fantaziya odeska, Krasunya odeska, Viktoria odeska, Nikonia, Zastava odeska, Luzanivka odeska Odeska 267, Selynkal, Kuyalnyk, Znakhidka odeska, Sirena odeska, Pysanka, Vdala, Antonivka, Turunchuk
[6, 7]. We also know about the positive influence of Lr34/Yr18/Pm38 upon stem rust resistant genes [32]. When soft winter wheat varieties of the Plant Breeding and Genetics Institute of the National Center of Seed and Cultivar Investigations were divided on the basis of stages of selection according to Lytvynenko’s recom mendations [3] (Table 4), we noticed the increase of fre quency of allele csLV34b with every new stage (Fig. 3). This proves the preservation of efficiency of gene Lr34/Yr18/Pm in the south of Ukraine (if there are other genes of genus specific resistance), which is con nected with leaf rust, yellow rust and mildew resis tance, and barley yellow dwarf virus resistance, which made for selection of plant genotypes with this gene. 80
csLV34a allel
40 20
1
2
3
4
5
6
Fig. 3. Histogram of distribution of frequencies (down, %) of alleles of locus csLV34 in soft wheat varieties of the Plant Breeding and Genetics Institute of the National Center of Seed and Cultivar Investigations at different stages of selec tion (across: (1) 1960–1967; (2) 1968–1975; (3) 1976–1990; (4) 1991–1996; (5) 1996–2004; (from 2005 to present)).
CONCLUSIONS 1
2 2 25 15 26 68
150 bp long associated with the presence of absence of gene Lr34/Yr18/Pm38, correspondingly. The oldest variety where we found allele csLV34b is not connected with Italian variety Ardito. This is Odessa variety of spring wheat Girka 0274. As variety Girka 0274 was not used in selection programs, variety Bezosta 1 is a promosource of gene Lr34/Yr18/Pm38 in varieties of Ukrainain and Russian selection. This testifies to the generally accepted hypothesis. Allele csLV34a is the most wide spread in the north and center of Ukraine similar to soft winter wheat varieties of Russian selec tion (from 52.7 to 80.0%). On the contrary, allele csLV34b was widely spread among varieties of the south of Ukraine. REFERENCES
csLV34b allel
60
0
Number of varieties studied
Varieties (main representatives)
While using codominant moleculargenetic marker csLV34 bound with multipathogenic resistance gene Lr34/Yr18/Pm38, we conducted the analysis of allele state of locus csLV34 among soft wheat varieties of Ukrainian and Russian selection. We found two alleles of this locus: csLV34a 229 bp long and csLV34b
1. McIntosh, R.A., Dubcovsky, J., Rogers, W.J., et al., Catalogue of Gene Symbols for Wheat: 2011 suppl. http://www.shigen.nig.ac.jp/wheat/komugi/genes/ macgene/supplement2011.pdf 2. Caldwell, R.M., Breeding for general and/or specific plant disease resistance, in Proc. 3rd Int. Wheat Genet. Symp., Finlay, K.W. and Shepherd, K.W., Eds., Can berra, 1968, pp. 263–272. 3. Rubiales, D. and Niks, R.E., Characterisation of Lr34, major gene conferring nonhypersensitive resistance to wheat leaf rust, Plant Dis., 1994, vol. 94, pp. 1208– 1212. 4. Spielmeyer, W., McIntosh, R.A., Kolmer, J., and Lagudah, E.S., Powdery mildew resistance and Lr34/Yr18 genes for durable resistance to leaf and stripe rust coseg regate at a locus on the short arm of chromosome 7D of wheat, Theor. Appl. Genet., 2005, vol. 111, pp. 731–735. 5. Lillemo, M., Asalf, B., Singh, R.P., et al., The adult plant rust resistance loci Lr34/Yr18 and Lr46/Yr29 are important determinants of partial resistance to powdery mildew in bread wheat line Saar, Theor. Appl. Genet., 2008, vol. 116, pp. 1155–1166. 6. Kolmer, J.A., Genetics of resistance to wheat leaf rust, Annu. Rev. Phytopathol., 1996, vol. 34, pp. 435–455. 7. Shtubei, T.Yu., Cytophysiological aspects of age resis tance of common wheat to the brown rust pathogen, CYTOLOGY AND GENETICS
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31. Marshll, D., Virulence of Puccinia recondite in Texas from 1988 to 1990, Plant Dis., 1992, vol. 76, pp. 296– 299. 32. Kerber, E.R. and Aung, T., Leaf resistance gene Lr34 associated with non suppression of stem rust resistance in the wheat cultivate Canthatch, Phyropathology, 1999, vol. 88, pp. 518–521. 33. Litvinenko, M.A., Department of breeding and seed production of wheat in the 100year history of the insti tute, Zb. Nauk. Pr. SGINTsN, 2012, vol. 20, no. 60, pp. 11–27.
Translated by N. Berestetska
