Dynamics of genetic diversity in winter common wheat Triticum aestivum L. cultivars released in Russia from 1929 to 2005

Genealogical analysis was used to study the dynamics of genetic diversity in Russian cultivars of winter common wheat from 1929 to 2005. The Shannon diversity index of the total set of released cultivars remained almost unchanged, although the number of original ancestors (landraces and genetic lines) increased almost tenfold in the period under study. This was explained in terms of the dependence of the modified Shannon diversity index on two parameters, the number of original ancestors and the mean coefficient of parentage. Significant direct effects were revealed: a positive effect of the former parameter and a negative of the latter. As a result, the increase in the number of original ancestors was compensated by the increase in relatedness of cultivars. Genetic erosion of released diversity was observed, as a half of Russian landraces were lost. Although the mean coefficient of parentage did not reach its critical value $(\bar R = 0.25)$ , cultivars of some regions (Central and Volga-Vyatka) proved to be closely related. A favorable gradual decrease in the mean coefficient of parentage was observed in the past 15 years. A set of modern winter wheat cultivars, which were introduced in the Russian Official List from 2002 to 2005, displayed a cluster structure. The overwhelming majority of cultivars formed two clusters originating from Bezostaya 1 (67% of cultivars) and Mironovskaya 808 (31%).     

Genealogical analysis of resistance to fusarium head blight in Russian and Ukrainian cultivars of common wheat Triticum aestivum L.

The GRIS3.5 information analytical system of wheat genetic resources was used to trace the possible ways of the transmission of fusarium head blight resistance from ancestors to progenies in extended pedigrees of 149 Russian and Ukrainian cultivaris of winter common wheat. Analysis of variance was performed for the coefficient of parentage computed between the cultivars under study and the putative sources of resistance and revealed that groups of resistant and susceptible cultivars differed in the distribution of contributions of the sources. In the resistant group, significant results were obtained for the contributions of Odesskaya 16, Hostianum 237, and Frontana. Pedigree analysis showed that fusarium head blight resistance was most commonly transmitted from Hostianum 237 through Odesskaya 16 and its derivatives. The landrace Khar’kovskaya probably served as a source of resistance in the case of Hostianum 237. In addition, the set of resistance sources included Kooperatorka, Hope, SanPastore, Triticum timopheevii Zhuk., and Secale cereale. Some well-known sources of fusarium head blight resistance varying in genetic determinants—Sumai 3, Wangshuibai, Wuhan 1, Nyubay (China), Nobeokabozukomugi, Shinchunaga (Japan), Arina (Switzerland), Fundulea-201R (Romania), and Renan (France)—have so far not being employed in breeding in Russia and provide an important reserve for breeding for resistance.     

Genealogical and statistical analyses of the inheritance of gliadin-coding alleles in a model set of common wheat <Emphasis Type="Italic">Triticum aestivum</Emphasis> L. cultivars

Allelic diversity of the gliadin-coding loci Gli-1 and Gli-2 was compared with the genealogical profiles of common wheat cultivars developed in Saratov. Allele tracking through their pedigrees and hierarchic cluster analysis associated 31 Gli alleles with groups of original ancestors. The cultivars Poltavka (12 alleles of six loci) and Selivanovskii Rusak (six alleles of six loci) were identified as sources of the majority of alleles. The results of the cluster analysis fully coincided with the results of allele tracking for alleles occurring at high frequencies. For rare alleles, the resolution of the cluster analysis was somewhat lower and depended on the similarity/distance measure. Thus, it proved possible to indirectly identify the donors of gene alleles by multidimensional statistics even when data on alleles identified in ancestors are unavailable. This approach to the analysis of inheritance has two limitations: detailed pedigree data should be known, and relatively high frequencies (no less than 15–20%) should be observed for the alleles in a sample under study. Cluster analysis was used to study the association of gliadin alleles with commercial quality classes. The most important gliadin-coding alleles, which mark strong cultivars, were identified. In the Saratov cultivars, such alleles include Gli-A1f, GliB1e, Gli-D1a, Gli-A2q, Gli-B2s, and Gli-D2e, which were inherited from the landrace Poltavka, and Gli-A1i, Gli-A2s, and Gli-B2q, which were inherited from the landrace Selivanovskii Rusak.     

Resistance of bread wheat (Triticum aestivum L.) to preharvest sprouting: An association analysis

A statistical analysis of the data about 1422 bread wheat accessions with estimated preharvest sprouting was carried out. Close associations of preharvest sprouting resistance with the grain color and with resistance to Fusarium head blight were revealed, as well as weak, but statistically significant, associations with the habit, awnedness, and reduced height genes Rht-B1 and Rht-D1 (insensitive to gibberellin GA3). The pedigree analysis showed that the cluster structures of the gene pools of the North American red-grained and white-grained varieties are practically identical. In both groups, varieties that are resistant to preharvest sprouting differ from susceptible ones in the percentage of the contributions of the Crimean and Mediterranean landraces. Resistance is associated with a high contribution by the Crimean landrace and susceptibility is associated with a high contribution by the Mediterranean landrace.     

Analysis of Genetic Diversity of Spring Durum Wheat (<Emphasis Type="Italic">Triticum durum</Emphasis> Desf.) Cultivars Released in Russia in 1929–2004

Based on genealogical analysis, the genetic diversity of 78 spring durum wheat cultivars released in Russia in 1929–2004 have been examined. The temporal trends of change in diversity were studied using series of n × m matrices (where n is the number of the cultivars and m is the number of original ancestors) and calculating coefficients of parentage in sets of cultivars released in particular years. The pool of original ancestors of spring durum wheat cultivars includes 90 landraces and old varieties, more than a half (57%) of which originate from European countries, including Russia and Ukraine (45%). The original ancestors strongly differ in the frequency of presence in the cultivar pedigrees. Landraces Beloturka, Sivouska, Kubanka (T. durum Desf.), Transbaikalian emmer, Yaroslav emmer (T. dicoccum Schuebl.), Poltavka (T. aestivum L.), and the original ancestors of cultivars Kharkov 46, Narodnaya, and Melanopus 1932 enter in the pedigrees of more than half of cultivars created within the framework of various breeding programs. At that, their distribution by cultivars from different breeding centers strongly varies. Analysis of temporal dynamics of genetic diversity, based on genetic profiles and coefficients of parentage, has shown that the genetic diversity of Russian durum wheats increased during the period examined. Nevertheless, genetic erosion of the local material—a loss of approximately 20% of the pool of Russian original ancestors—has been found. The contribution of the original ancestors to the pedigrees of different cultivars, constructed in different breeding centers and recommended for cultivation in different regions, has been estimated. The variation of the released cultivars was highest in the Lower Volga region and lowest in the Ural region. In all, the lower threshold of genetic diversity in all regions does not reach the critical level, corresponding to the similarity of half-sibs. The set of modern cultivars included in the Russian Official List 2004 has a cluster structure.     

Analysis of diversity of russian and Ukrainian bread wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) cultivars for high-molecular-weight glutenin subunits

The allelic diversity of high-moleculat-weght glutenin subunits (HMWGS) in Russian and Ukrainian bread wheat cultivars was analyzed. The diversity of spring wheat cultivars for alleles of the Glu-1 loci is characterized by medium values of the polymorphism polymorphism information content (PIC), and in winter wheats it varies from high at the Glu-A1 locus to low at the Glu-D1 locus. The spring and winter cultivars differ significantly in the frequencies of alleles of the glutenin loci. The combination of the Glu-A1b, Glu-B1c, and Glu-D1a alleles prevails among the spring cultivars, and the combination of the Glu-A1a, Glu-B1c, and Glu-D1d alleles prevails among the winter cultivars. The distribution of the Glu-1 alleles significantly depends on the moisture and heat supply in the region of origin of the cultivars. Drought resistance is associated with the Glu-D1a allele in the spring wheat and with the Glu-B1b allele in the winter wheat. The sources of the Glu-1 alleles were identified in the spring and wheat cultivars. The analysis of independence of the distribution of the spring and winter cultivars by the market classes and by the alleles of the HMWGS loci showed a highly significant association of the alleles of three Glu-1 loci with the market classes in foreign cultivars and independence or a weak association in the Russian and Ukrainian cultivars. This seems to be due to the absence of a statistically substantiated system of classification of the domestic cultivars on the basis of their quality.     

Genealogical analysis of the use of two wheatgrass (<Emphasis Type="Italic">Agropyron</Emphasis>) species in common wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) breeding for disease resistance

During the last 80 years, in order to increase the genetic variability of wheat, translocations containing nine elongated wheatgrass (Agropyron elongatum) and eight intermediate wheatgrass (Agropyron intermedium) genes, which control resistance to pathogens, were transferred to this crop culture. Genealogical and statistical analysis of 1500 varieties developed using the wheatgrass gave evidence of the continuing increase in the proportion of such varieties in the total number of wheat varieties over the last half-century. Translocations from Ag. elongatum most commonly occur in the pedigrees of the varieties from the United States, less frequently they can be found in Australian and Chinese varieties, and they are extremely rare—in European and African ones. Ag. intermedium most frequently occurs in the pedigrees of the Eastern European varieties, mainly in those from Russia, as well as in the varieties from China. The observed uneven distribution of such varieties may be associated with either the effectiveness of the translocation in the development of resistance to the local populations of pathogens or with the effect of the translocation on the adaptive traits of plants. By computer tracking of pedigrees, we performed an inventory of the translocation donors from Ag. elongatum and Ag. intermedium used in the breeding programs in the United States, Russia, Australia, India, and China. The most widely occurring combinations of the gene complex Lr24/Sr24 of Ag. elongatum with other resistance genes were revealed. In Russia, there were developed varieties in which the 6D chromosome was substituted by the 6Ai chromosome of Ag. intermedium, which controls disease resistance and the adaptivity of plants. The identification and introgression of new translocations indicates that the possibilities of using wheatgrass species for broadening of genetic variability of wheat are far from being exhausted.     

Analysis of the Distribution of <Emphasis Type="Italic">Triticum timopheevii</Emphasis> Zhuk. Genetic Material in Common Wheat Varieties (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

The database of the world gene pool of wheat was scanned by pedigree and the participation of genetic material from T. timopheevii in the creation of 3088 varieties of common wheat was established. The spatial and temporal dynamics of the propagation of these varieties was studied. Using the analysis of pedigrees, a diversity of T. timopheevii donors was studied. The specificity of donors of the genetic material T. timopheevii for the regions of wheat breeding was established. The main source of resistance genes for most varieties is accession D-357-1 from the Georgian variety-population of Zanduri. This significantly reduces the diversity of the genetic material of T. timopheevii used in wheat breeding. In 369 varieties and 184 lines, the genes for resistance to pathogens from T. timopheevii were identified. The genes of T. timopheevii are distributed mainly in winter varieties, as well as spring varieties sown in autumn. The value of donors as sources of T. timopheevii genes is ambiguous, despite the fact that most of them come from the same D-357-1 accession. The Sr36 gene is most commonly found in the United States, Western Europe, and Australia; it was transferred from the Wisconsin-245 line through Arthur or TP-114-1965a. The Pm6 gene is distributed in Western Europe; it was transferred from the pre-breeding line Wisconsin 245/5*Cappelle-Desprez//Hybrid- 46/Cappelle Desprez. The gene Lr18 is more common in the United States; it was transmitted by the Blueboy or Vogel 5 varieties from the Coker-55-9 line. The extremely limited set of genes for resistance to pathogens from T. timopheevii used in commercial varieties and the specificity of their geographical distribution are possibly associated with the uniqueness of the G subgenome and plasmon in this species, its low potential for plasticity, and tolerance to drought. In addition, the imperfection of the methods of pre-breeding and recombination breeding prevents the elimination in translocation of close linkage of target genes with undesirable ones.     

Comparative Genealogical Analysis of the Resistance of Winter Wheat to Common Bunt

Comparative genealogical analysis of North American (the United States and Canada) and Eastern European (Russia and Ukraine) winter wheat cultivars resistant and susceptible to common bunt has been performed. Analysis of variance applied to North American wheats has demonstrated that resistant and susceptible cultivars significantly differ from each other with respect to the contributions of common ancestors. The contributions of Oro (Bt4and Bt7), Rio (Bt6), White Odessa (Bt1), and Florence (Bt3) to the resistant cultivars are significantly higher than their contributions to the susceptible ones. This demonstrates that the use of these resistance donors in wheat breeding for several decades has been effective. The contribution of PI-178383 (Bt8, Bt9,and Bt10) is considerably higher in the group of resistant cultivars bred after 1965. The mean contributions of Federation (Bt7) and Nebred (Bt4) are significantly higher in the group of resistant cultivars obtained before 1965; however, the differences in the contributions of these donors between new resistant and susceptible cultivars became nonsignificant. Among the Russian and Ukrainian cultivars, there are differences between groups of resistant and susceptible cultivars from different regions determined by the differences between the regional populations of the pathogen in racial composition. In the northern region, the contributions of the wheat grass (Agropyron glaucum) and the rye cultivar Eliseevskaya are significantly higher in the resistant cultivars; in the southern region, a local cultivar of the Odessa oblast is the prevalent resistant cultivar. In addition, cultivar Yaroslav Emmer is likely to be effective in the northern region; and foreign sources (Oro, Florence, Federation, and Triticum timopheevii), in the southern region. Very few sources of vertical resistance to common bunt are used for winter wheat breeding in Russia and Ukraine. The decrease in genetic diversity in favor of a few identical genes may cause adequate changes in the pathogen population and subsequent proliferation of the pathogen on the genetically identical substrate. A new interpretation of the resistance of line Lutescens 6028 as a source of new genes, Bt12 and Bt13, is suggested. Both genealogical and segregation analyses have shown that the genes determining the resistance of this line may be identical to those described earlier (Bt1, Bt3, Bt4, Bt6, and Bt7); and the high resistance of this line is determined by a combination of these genes.     

New data on the distribution of hybrid necrosis genes in winter bread wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) cultivars

Hybrid necrosis genotypes have been identified in 125 Russian cultivars of winter bread wheat. More than half of them (56%) carry the Ne2 gene (genotype ne1ne1Ne2Ne2); others are free of necrosis genes (genotype ne1ne1ne2ne2). The possible causes of the increase in the Ne2 allele frequency and the loss of the Ne1Ne1ne2ne2 genotype in modern Russian cultivars of winter wheat are discussed. The principal component method has been used to compare the structures of the genetic diversity of cultivars differing in the hybrid necrosis genotype. It has been found that the Ne2 allele in winter wheat cultivars from northern Russia has originated from the cultivar Mironovskaya 808, whereas the cultivar Bezostaya 1 is not a source of this gene. In cultivars from southern Russia, the presence of the Ne2 allele is also mainly accounted for by the use of Mironovskaya 808 wheat in their breeding. The recessive genotype is explained by the presence of descendants of the cultivar Odesskaya 16 in the pedigrees of southern Russian winter wheats. The genetic relationship of cultivars with identical and different necrosis genotypes has been analyzed in nine regions of the Russian Federation.