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Сельскохозяйственная биология, 2012, № 3, с. 68-74.

doi: 10.15389/agrobiology.2012.3.68eng

УДК 633.11.«324»:575.224.232.3

RYE TRANSLOCATIONS IN THE VARIETIES OF WINTER COMMON WHEAT

N.A. Kozub^{1, 5}, I.A. Sozinov¹, T.A. Sobko^{1, 5}, O.S. Dedkova², E.D. Badaeva³, V.P. Netsvetaev⁴

Twenty winter common wheat varieties from the competitive testing of Belgorod State Research Institute of Agriculture were studied with respect to the storage protein loci. Special attention was given to two varieties, Synthetic and Bogdanka, with rye chromatin material, which were also studied by cytogenetic analysis. Among the studied varieties, rye material marked by the gliadin loci was identified in Synthetic, Kryzhinka, and Bogdanka varieties. Synthetic and Kryzhinka varieties have 1BL/1RS rye translocation, and Bogdanka variety carries 1AL/1RS translocation. These data were confirmed cytologically. Alleles at the Gli-1 (= Gld) loci, which control synthesis of gliadins, and alleles at the Glu-1 loci, which control synthesis of HMW glutenin subunits, were identified in 20 varieties. Genealogy of Bogdanka, the first winter common wheat variety of Russian breeding, with 1AL/1RS translocation was clarified.

Keywords: common wheat, translocations, gliadin, glutelin, chromosomes.

 

Analysis of allelic forms of wheat genes for storage proteins is widely used in genetic studies, breeding work and seed production (1). Gliadin is encoded by gene clusters of six major loci – Gli-А1, Gli-B1, Gli-D1, Gli-A2, Gli-B2 and Gli-D2 located distally on short arms of chromosomes of the 1st and 6th homoeologous groups (1). Loci controlling the synthesis of high-molecular weight (HMW) subunits of glutenin (Glu-A1, Glu-B1, Glu-D1) are located on long arms of chromosomes of the 1st homoeologous group (2). Genes for storage proteins have multiple alleles (3-5). Allelic forms in the loci of storage proteins are directly related to bread-making properties (1, 2, 6).
Wheat-rye translocations 1BL/1RS and 1AL/1RS show the most common type of introgressions in commercial wheat varieties (7). A source of 1BL/1RS translocation in modern varieties of soft wheat is mainly the line Riebesel 47-51 (author G. Riebesel) carrying a translocation from rye cv Petkus (2x) (7). 1AL/1RS translocation in most cases originates from cv Amigo created in the USA in 1976. A fragment of rye chromosome 1R in cv Amigo was obtained from Argentine rye cv Insave (Secale cereale L.) (8). There are several methods proposed for identification of rye translocations 1BL/1RS and 1AL/1RS (biochemical, cytological, DNA-labeling). The most common method is electrophoresis of alcohol-soluble storage protein of grains in an acidic medium (1).
The translocation 1BL/1RS in the genome of soft wheat is known to affect its bread-making properties  (1). This fact can be partly compensated by presence of alleles positively affecting grain quality in respect to other loci, in particular, HMW loci for glutenin subunits. Effects of 1AL/1RS translocation were investigated, and it was found to cause a smaller reduce in grain quality than 1VL/1RS (9, 10).
In this work the authors studied winter wheat varieties to identify genetic material of rye introduced into storage protein loci with a special attention to recently zoned varieties Sintetik and Bogdanka aimed at obtaining more data on their genealogy.
Technique. A set of 20 cultivars (competitive testing in Belgorod Research and Development Institute of Agriculture) included varieties Sintetik and Bogdanka zoned for Central Chernozem Zone of RF and registered in the State register of breeding achievements permitted for use (respectively, 2008 and 2009). Cv Sintetik was proposed for zonal growing in Kursk region, cv Bogdanka - in Belgorod region (11). The first variety, as a rule, produces gluten of II quality group (12, 13), the second – a little better one.
Endosperm storage proteins were analyzed in individual kernels. Electrophoresis of prolamins was performed in acidic medium in polyacrylamide gel (14). Notations of prolamins correspond to ones of E.V. Metakovsky (15, 16) and F.A. Poperele, T.A. Sobko (17, 18). Electrophoresis of glutenin HMW subunits was performed using the method of U.K. Laemmli in 10% separating gel (19). Alleles of glutenin HMW subunits were identified from the catalog of P. Payne et al. (3).
Cytological analysis of chromosomes was carried out by examination of roots of germinating seed. C-differential staining of  chromosomes was performed according to E.D. Badaeva (20, 21).
Genealogy of cv Sintetic: {[(Odesskaya 130 x Ol’via) Odesskaya 51] [Odesskaya 51 (Mironovskaya 808 x Avrora)] [(Odesskaya 51 x Iniya 66) (Odesskaya 51 x W.S. 1877 “modified”) (Ol’via x Odesskaya 130 )]}.
Genealogy of cv Bogdanka: {[BC1 (Odesskaya semi-dwarf x Aegilops cylindrica, Odessa population) Amigo] x Wheat grass, spontaneous hybridization} x Volzhskaya 16.
Results. Genotypes of winter wheat varieties (both passing state testing and cultivated in the 5th region of Russia - Central Chernozem Zone) in respect to gliadin and glutenin revealed by electrophoresis of storage proteins in grains are shown in Table 1. A sample Selyanka odesskaya was obtained from originators as a tall version of this variety, so variants of its formula described in this work may not coincide with those of other researchers. The maximum number of alleles (six) were identified in the locus Gli-A1, four alleles – in loci Gli-B1 and Gli-D1. In the studied group of varieties gliadin alleles were dominant: Gli-A1b, Gli-B1b, Gli-D1b and Glu-D1g (Table 2). For the loci of glutenin HMW subunits there were detected alleles determining high bread-making quality (Glu-A1a, Glu-A1b, Glu-B1b, Glu-B1c, Glu-D1d) (22). There should be noted coincidences in the set of identified dominant alleles with those of the well-known cv Bezostaya 1 (Gli-A1b, Gli-B1b, Gli-D1b, Glu-A1b, Glu-B1c, Glu-D1d), which probably indicates their adaptive role.
It is known that wheat-rye translocation 1BL/1RS or 1B substitution of wheat chromosome to 1R rye chromosome in wheat genome results in a significant reduce of grain quality. Chromatin of the short arm of 1R chromosome can be easily identified by analysis of gliadin composition (1, 17, 23, 24). A marker of 1BL/1RS translocation is secalin block synthesized under the control of allele Gli-B1l (15) or GLD 1B3 (1), for 1AL/1RS translocation - secalin block GLD 1A17 with a specific  spectrum of components controlled by allele Gli-B1l. It was proposed to be designated as Gli-A1w (25).

2. Frequency of alleles for storage proteins in the studied set of winter soft wheat varieties (competitive testing, Belgorod Research and Development Institute of Agriculture)
Locus, allele	Frequency of allele
Gli-A1:
b	0,475
c	0,050
f	0,150
o	0,200
x	0,075
w	0,050
Gli-B1:
b	0,700
d	0,150
e	0,050
l	0,100
Gli-D1:
b	0,425
f	0,025
g	0,400
j	0,150
Glu-A1:
a	0,475
b	0,525
Glu-B1:
b	0,475
c	0,525
Glu-D1:
a	0,025
d	0,975

In the studied group of wheats the genetic material of rye was detected in three varieties - Sintetik, Kryzhinka and Bogdanka (Table 1). Analysis of endosperm storage proteins showed in cv Sintetik the presence of proteins determined by allele Gli-B1l (GLD 1B3) (Fig. 1) peculiar to carriers of 1B/1R substitution or 1BL/1RS translocation. The type of such introduction in cv Sintetik was revealed by C-differential staining (Fig. 2): in the short arm of 1B chromosome occurred an intense staining of telomeric part typical to rye chromatin. The long arm didn’t differ from a typical wheat. Therefore, cv Sintetik is a carrier of 1BL/1RS translocation. This was also confirmed by the presence of wheat c allele in locus Glu-B1 on the long arm of 1B chromosome, which was revealed by electrophoretic analysis of glutelin. In general, cv Sintetik has the following alleles for endosperm storage proteins: Gli-A1b, Gli-B1l, Gli-D1b + g, Glu-A1b, Glu-B1c, Glu-D1a + d. Genealogy of this sample suggests that such modification of 1B chromosome most likely originates from cv Aurora. In variety Kryzhinka the presence of rye translocation 1BL/1RS was also proved by analysis of gliadin and glutenin.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Electrophoresis of gliadins in cv Bogdanka (Fig. 1) showed the presence of GLD 1A17 proteins in the endosperm and, respectively, allele Gli-A1w specific to cv Amigo and varieties carrying the translocation 1AL/1RS (18, 25). At the same time, such variant of gliadin (GLD 1A17) is known for samples of wheat with 1B/1R replacement originated from octaploid triticale AD825 (24). In this regard chromosomes of cv Bogdanka were investigated using C-differential staining (Fig. 2), which showed in it 1B chromosome typical to hexaploid wheat and a modified chromosome 1A. Its short arm with striations looks similar to the short arm of 1B chromosome of cv Sintetik. Consequently, cv Bogdanka has rye translocation on 1A chromosome  (1AL/1RS). This fact was also confirmed by the presence of wheat b allele in Glu-A1 locus on the long arm of 1A chromosome shown by electrophoresis of glutelin. Genealogy of cv Bogdanka reported by V.P. Netsvetaev and N.P. Domanov (11) has no data on a source of rye chromatin. However, it is known that during the breeding work on cv Bogdanka interspecific hybrids were sown nearby the cultivar Amigo. The latter can be assumed as a participant of spontaneous hybridization resulted in a source material for cv Bogdanka. One of the variants with participation of cv Amigo in cv Bogdanka was mentioned above (see Technique). Probably, it occurred at a late (penultimate) stage of creation of cv Bogdanka. This variety carries the following alleles determining synthesis of storage proteins: Gli-A1w, Gli-B1e, Gli-D1g, Glu-A1b, Glu-B1c, Glu-D1d. According to the catalog proposed by E.V. Metakovsky et al. (16), cv Amigo is a carrier of Gli-A1o allele located on 1A chromosome. This allele is peculiar to samples with typical wheat chromosome 1A, such as Belgorodskaya 12, Belgorodskaya 16, Povaga. At the same time, cv Amigo has rye translocation 1AL/RS (26), which coincides with data of analysis of its storage proteins (18). Therefore, the authors assume that Gli-A1o allele was erroneously identified in cv Amigo.

1RS транслокация в сорте пшеницы Amigo, полученная от ржи сорта Insave, в хромосоме 1A несет ряд генов устойчивости к болезням и вредителям: гены Gb2 и Cm3, из которых первый определяет устойчивость к тле Schizaphis graminum биотипов B и C, второй — к клещу Aceria tosicheilla (Keifer), гены Pm17 и Sr1AR, обусловливающие устойчивость соответственно к мучнистой росе и стеблевой ржавчине (5). Сорта с 1AL/RS транслокацией были ранее идентифицированы среди форм украинской селекции, созданных в последние 15 лет (18, 25). Богданка — первый российской сорт, несущий эту транслокацию.
Таким образом, ряд сортов озимой мягкой пшеницы охарактеризованы по глиадин- и глютенинкодирующим локусам. На основе этого анализа выделены сорта, имеющие ржаные глиадины, а с помощью цитогенетического исследования определены хромосомы и плечи с наличием ржаной транслокации 1RS, ответственной за синтез этих белков. Уточнена генеалогия нового сорта озимой мягкой пшеницы Богданка. Сорт Богданка — первый сорт российской селекции, который несет 1AL/RS транслокацию.
In wheat cv Amigo 1RS translocation originated from rye cv Insave and detected on 1A chromosome  includes a number of genes for resistance to diseases and pests: genes Gb2 and Cm3, the first of which determines the resistance to aphids Schizaphis graminum biotypes B and C, the second - to mites Aceria tosicheilla (Keifer), along with Pm17 and Sr1AR genes for resistance to, respectively, powdery mildew and stem rust (5). Varieties with 1AL/RS translocation were previously identified among the forms of Ukrainian selection developed in the last 15 years (18, 25). Bogdanka is the first variety of Russian selection carrying this translocation.
Thus, a group of winter wheat varieties were characterized in respect to their loci for gliadin and glutelin. This analysis revealed among them the carriers of rye gliadin, as well as chromosomes and their arms with the presence of rye translocation 1RS responsible for the synthesis of these proteins. It also allowed to obtain more data on genealogy of the new winter wheat cv Bogdanka, which was confirmed as the first variety of Russian selection carrying 1AL/RS translocation.

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1Institute of Plant Protection, UAAS, Kyiv 03022, Ukraine, e-mail: sia1@i.com.ua; 2N.I. Vavilov Institute of General Genetics, RAS, Moscow 119991, Russia, 119991 г. Москва, ул. Губкина, 3, e-mail: olgadedkova@gmail.com; 3V.A. Engelhardt Institute of Molecular Biology, RAS, Moscow 119991, Russia, e-mail: k_badaeva@mail.ru; 4Belgorod Research and Development Institute of Agriculture, RAAS, Belgorod 308001, Russia, e-mail: netsvetaev@bsu.edu.ru; 5Institute of Food Biotechnology and Genomics, NASU, Kyiv 04123, Ukraine

Received November 1, 2010

All-Russian Institute for Selection and Seed-Breeding of Vegetables LIK Cosmetics Institute of Agricultural Biotechnology Joint Stock Company "GosNIISyntezBelok"