Assessing fitness parameters of hybrids between weed beets and transgenic sugar beets

Herbicide resistance is a desired trait for sugar beet (Beta vulgaris L.) production because it is a low‐competitor crop that requires careful weed management. However, gene flow to weed beet (also B. vulgaris) could jeopardize the weed control strategy by causing the emergence of resistant weed beets; it could also lead to further adaptation of weed beet as a troublesome weed for other crops by selecting more competitive plants. To evaluate the hazard that such a selection process represents, apart the herbicide resistance, we investigated the morphology and reproduction of progeny of weed beets having inherited a herbicide resistance gene. First‐generation resistant weed beet exhibited traits likely counter‐selected. But such crop traits rapidly disappeared with backcrossing to weedy relatives: no biologically relevant difference was noted between resistant and susceptible near‐isogenic siblings in the various experiments. In the absence of resistance selection, our data indicate low chances for weed beet to evolve more competitive forms than present weed beet populations. However, they also suggest that there is no fitness cost limiting transgenes spread.     

QTL mapping of a partial resistance to the corn delphacid‐transmitted viruses in Lepidopteran‐resistant maize line Mp705

A partial resistance to maize mosaic virus (MMV) and maize stripe virus (MStV) was mapped in a RILs population derived from a cross between lines MP705 (resistant) and B73 (susceptible). A genetic map constructed from 131 SSR markers spanned 1399 cM with an average distance of 9.6 cM. A total of 10 QTL were detected for resistance to MMV and MStV, using composite interval mapping. A major QTL explaining 34–41% of the phenotypic variance for early resistance to MMV was detected on chromosome 1. Another major QTL explaining up to 30% of the phenotypic variation for all traits of resistance to MStV was detected in the centromeric region of chromosome 3 (3.05 bin). After adding supplementary SSR markers, this region was found to correspond well to the one where a QTL of resistance to MStV already was located in a previous mapping study using an F₂ population derived from a cross between Rev81 and B73. These results suggested that these QTL of resistance to MStV detected on chromosome 3 could be allelic in maize genome.     

Dissection of a major QTL for seed colour and fibre content in Brassica napus reveals colocalization with candidate genes for phenylpropanoid biosynthesis and flavonoid deposition

A major quantitative trait locus (QTL) influencing seed fibre and colour in Brassica napus was dissected by marker saturation in a doubled haploid (DH) population from the black‐seeded oilseed rape line ‘Express 617’ crossed with a yellow‐seeded B. napus line, ‘1012–98’. The marker at the peak of a sub‐QTL with a strong effect on both seed colour and acid detergent lignin content lay only 4 kb away from a Brassica (H+)‐ATPase gene orthologous to the transparent testa gene AHA10. Near the peak of a second sub‐QTL, we mapped a copy of the key phenylpropanoid biosynthesis gene cinnamyl alcohol dehydrogenase, while another key phenylpropanoid biosynthesis gene, cinnamoyl co‐a reductase 1, was found nearby. In a cross between ‘Express 617’ and another dark‐seeded parent, ‘V8’, Bna.CCR1 was localized in silico near the peak of a corresponding seed fibre QTL, whereas in this case Bna.CAD2/CAD3 lay nearby. Re‐sequencing of the two phenylpropanoid genes via next‐generation amplicon sequencing revealed intragenic rearrangements and functionally relevant allelic variation in the three parents.     

Identification of QTL controlling high levels of partial resistance to Fusarium solani f. sp. pisi in pea

Fusarium root rot is a common biotic restraint on pea yields, and genetic resistance is the most feasible method for improving pea production. This study was conducted to discover quantitative trait loci (QTL) controlling genetic partial resistance to Fusarium root rot caused by Fusarium solani (Mart.) Sacc. f.sp. pisi (F.R. Jones) W.C. Snyder & H.N. Hans (Fsp). A RIL population was screened in a Fusarium root rot field disease nursery for 3 years. Composite interval mapping was employed for QTL detection using the means of disease severity from three growing seasons. Five QTL were identified, including one QTL identified in all three years. The multiyear QTL Fsp‐Ps2.1 contributed to a significant portion of the phenotypic variance (22.1–72.2%), while a second QTL, Fsp‐Ps6.1, contributed 17.3% of the phenotypic variance. The other single growing season QTL are of additional interest as they colocate with previously reported pea–Fusarium root rot resistance QTL. QTL Fsp‐Ps2.1, Fsp‐Ps3.1, Fsp‐4.1 and Fsp‐Ps7.1 are flanked by codominant SSRs and may be useful in marker‐assisted breeding of pea for high levels of partial resistance to Fsp.     

Quantitative trait locus analysis of seed germination,seedling vigour and seedling‐regulated hormones in Brassica napus

Good germination and seedling vigour are major breeding targets in winter oilseed rape (Brassica napus), because seedling vigour and prewinter crop establishment are closely associated with postwinter growth and yield. Here, we identified quantitative trait loci (QTL) related to germination, seedling vigour and seedling‐regulated hormones in a doubled haploid (DH) mapping population from a cross between winter oilseed rape parents with high vigour (Express 617) and low vigour (1012‐98). By phenotyping in a climate‐controlled glasshouse, we identified a total of 13 QTL on nine chromosomes for germination and seedling‐related traits at 7 and 14 days after sowing (DAS), explaining up to 11.2% of the phenotypic variation for seedling vigour. Forty‐seven metabolic QTL on 15 chromosomes were identified for auxin, abscisic acid (ABA) and dihydrophaseic acid (DPA) at 5 and 12 DAS, explaining up to 49.4% of phenotypic variation in seedling hormone composition. Multitrait QTL hot spots contribute to our understanding of the genetics and metabolomics of germination and seeding vigour in B. napus, and represent potential targets to breed high‐vigour cultivars.     

Use of non‐adapted quantitative trait loci for increasing Fusarium head blight resistance for breeding semi‐dwarf wheat

Semi‐dwarf wheat is an important prerequisite for releasing a successful commercial cultivar in high‐yielding environments. In Northern Europe, this aim is achieved by using one of the dwarfing genes Rht‐B1 (formerly known as Rht‐1) or Rht‐D1 (Rht‐2). Both genes, however, result in a higher susceptibility to Fusarium head blight (FHB). We analysed the possibility to use the two non‐adapted FHB resistance quantitative trait loci Fhb1 and Fhb5 (syn. QFhs.ifa‐5A) to counterbalance the negative effect of the dwarfing allele Rht‐D1b in a winter wheat population of 585 doubled‐haploid (DH) lines segregating for the three loci. All lines were inoculated with Fusarium culmorum at four locations and analysed for FHB severity, plant height, and heading date. The DH population showed a significant (p < 0.001) genotypic variation for FHB severity ranging from 3.6% to 65.9% with a very high entry‐mean heritability of 0.95. The dwarfing allele Rht‐D1b reduced plant height by 24 cm, but nearly doubled the FHB susceptibility (24.74% vs. 12.74%). The resistance alleles of Fhb1 and Fhb5 reduced FHB susceptibility by 6.5 and 11.3 percentage points, respectively. Taken all three loci together, Fhb5 alone was already able to reduce FHB susceptibility to the same extent as Rht‐D1b increased it. This opens new avenues for selecting semi‐dwarf wheat by marker‐assisted introgression of Fhb5 without the enhancement of FHB susceptibility.     

Pyramiding three rust‐resistance genes confers a high level of resistance in soybean (Glycine max)

Pyramiding Asian soybean rust (ASR) resistance (Rpp) genes in a single genotype has been shown to increase ASR resistance in soybean. However, it remains unclear which combinations of Rpp genes are superior. Therefore, here, we developed six new Rpp‐pyramided lines carrying different combinations of Rpp genes and evaluated their resistance against 13 Bangladeshi rust (Phakopsora pachyrhizi) isolates (BdRPs) alongside seven previously developed Rpp‐pyramided lines. We found that lines carrying one, two and three Rpp genes had high ASR resistance without sporulation in 13.8%, 35.2% and 73.1% of the assays, respectively. Among the new lines that were developed, those with Rpp3 + Rpp4 and Rpp3 + Rpp4 + Rpp5 had high levels of ASR resistance, while the line containing Rpp2 + Rpp4 + Rpp5 showed immunity phenotype at two weeks after inoculation by the BdRP‐22 infection. Thus, pyramiding larger numbers of Rpp genes confers soybean with a higher level of resistance to ASR pathogens and can produce an immunity phenotype at two weeks after inoculation.     

Molecular characterization of genetic basis of Sugarcane Yellow Leaf Virus (SCYLV) resistance in Saccharum spp. hybrid

Sugarcane (Saccharum Spp.) produces 80% of the world's sugar along with other by‐products. The production of sugarcane is vulnerable to infestation of sugarcane yellow leaf virus (SCYLV) worldwide. A study was conducted using an F₁ segregating population derived from CP95‐1039 × CP88‐1762 to identify the genetic factors underlying SCYLV resistance. The disease infection data were measured using tissue blot immunoassay after 6 years of exposure to the virus under natural field conditions. Genetic maps were created using genotyping by sequencing‐based markers for each parent separately following a pseudo‐testcross approach. Two quantitative trait loci (QTL) were detected for SCYLV resistance accounting for 28% of the phenotypic variation. A major QTL qSCYLR79 located on linkage group 79 and linked with marker 3PAV3154 appears to be unique for SCYLV resistance in sugarcane. Progeny having a combination of two major alleles had 31% less SCYLV incidence than progeny with a combination of major and minor alleles in the genomic region of qSCYLR79. Thus, selection against the minor allele may decrease the SCYLV incidence in sugarcane.     

Identification and marker‐assisted introgression of QTL conferring resistance to bacterial leaf blight in cowpea (Vigna unguiculata (L.) Walp.)

Bacterial leaf blight (BLB), caused by Xanthomonas axonopodis pv. vignicola (Xav), is widespread in major cowpea [Vigna unguiculata (L.) Walp.] growing regions of the world. Considering the resource poor nature of cowpea farmers, development and introduction of cultivars resistant to the disease is the best option. Identification of DNA markers and marker‐assisted selection will increase precision of breeding for resistance to diseases like bacterial leaf blight. Hence, an attempt was made to detect QTL for resistance to BLB using 194 F_2 : 3 progeny derived from the cross ‘C‐152’ (susceptible parent) × ‘V‐16’ (resistant parent). These progeny were screened for resistance to bacterial blight by the leaf inoculation method. Platykurtic distribution of per cent disease index scores indicated quantitative inheritance of resistance to bacterial leaf blight. A genetic map with 96 markers (79 SSR and 17 CISP) constructed from the 194 F₂ individuals was used to perform QTL analysis. Out of three major QTL identified, one was on LG 8 (qtlblb‐1) and two on LG 11 (qtlblb‐2 and qtlblb‐3). The PCR product generated by the primer VuMt337 encoded for RIN2‐like mRNA that positively regulate RPM1‐ and RPS2‐dependent hypersensitive response. The QTL qtlblb‐1 explained 30.58% phenotypic variation followed by qtlblb‐2 and qtlblb‐3 with 10.77% and 10.63%, respectively. The major QTL region on LG 8 was introgressed from cultivar V‐16 into the bacterial leaf blight susceptible variety C‐152 through marker‐assisted backcrossing (MABC).     

Beet mild yellowing virus resistance derived from wild and cultivated Beta germplasm

Sugar beet (Beta vulgaris ssp. vulgaris L.) has a relatively narrow genetic base and several programmes have sought to evaluate the potential for introducing novel traits from wild and cultivated Beta germplasm into the crop. In particular, resistance to important sugar beet diseases has been identified within individual Beta accessions. We report here the successful transfer of resistance to Beet mild yellowing virus (BMYV) from garden beet, fodder beet and leaf beet accessions to progeny populations in initial crosses with sugar beet. Twelve plant populations derived from different Beta accessions were inoculated with viruliferous aphids carrying BMYV and the virus content of individual plants subsequently quantified by an ELISA test. Seven populations were significantly more resistant than a control sugar beet cultivar (P ≤ 0.05). BMYV resistance was successfully inherited in BC₁ and BC₂ generations, suggesting that resistance could potentially be introgressed from these sources into elite sugar beet lines.     

Effect of 3B, 5A and 3A QTL for Fusarium head blight resistance on agronomic and quality performance of Canadian winter wheat

The objectives of this study were to investigate (i) the correlations between Fusarium head blight (FHB) index, deoxynivalenol (DON) accumulation and percentage of Fusarium‐damaged kernels (FDK) with agronomic and quality traits and (ii) the effect associated with the presence of single QTLs for FHB resistance on agronomic and quality traits in winter wheat. The population was derived from the cross between ‘RCATL33' (FHB resistance derived from ‘Sumai 3’ and ‘Frontana’) and ‘RC Strategy’. Parental lines and recombinant inbred lines (RILs) were genotyped with SSR markers associated with the 3B, 5A and 3A QTLs. The population was planted in FHB‐inoculated nurseries and in agronomy trials. Lines in the 3B QTL class had the lowest FHB index, DON content and FDK level and did not have a significantly lower yield, thousand kernel weight or protein content compared with the lines grouped in other QTL classes (including no QTL class). Marker‐assisted selection of the 3B QTL for FHB resistance into high‐yielding FHB‐susceptible winter wheat is the recommended approach for the development of lines with increased FHB resistance without significant yield and quality penalties.     

Yield of bolting winter beet (Beta vulgaris L.) as affected by plant density,genotype and environment

Winter beet roots and shoots might be a favorable substrate for biogas production in Central Europe. However, detailed information about the attainable yield of this crop is lacking. Thus, the impact of plant density, genotype and environmental conditions on total dry matter yield of winter beet crops that bolt after winter was investigated. A significant increase of the dry matter yield (esp. shoot) was expected by harvesting the 1st shoot after flowering in June followed by a final harvest of the whole plant in July. In 2009/10, 2010/11 and 2011/12, three series of field trials with (i) 3 target plant densities (148, 246, 370 thousand plants ha⁻¹) and (ii) 3 different sugar beet genotypes were conducted at Göttingen (Lower Saxony, GER) and Kiel (Schleswig-Holstein, GER); (iii), additional field trials with 5 different sugar beet genotypes cultivated at 2 target plant densities (148, 246 thousand plants ha⁻¹) were conducted in 2011/12, to investigate the relation between maximum taproot diameter and the shoot and taproot yield of bolting winter beet. The total dry matter yield considerably varied between 4 and 23 t ha⁻¹. It was predominantly affected by the environment and to a substantially lower extent by plant density. Increasing plant densities increased the total dry matter yield, resulting in a significantly higher total dry matter yield at plant densities ≥300,000 plants ha⁻¹ compared with lower plant densities. Genotypic differences in total dry matter yield were negligibly small. Pruning in June substantially increased the total dry matter yield in July by ca. 8 t ha⁻¹ only in one out of three environments.Final yield in June (without pruning) and July (pruning in June) was positively related with cumulated temperature and global radiation, but also with taproot dry matter yield before winter. The taproot, shoot (1st, 2nd) and total plant yield were positively correlated with maximum taproot diameter.In conclusion, high dry matter yields close to yields of established energy crops grown over winter were obtained with winter beet roots and shoots only under very favorable conditions (climate, single plant size). High yields can be achieved after good pre-winter development. However, for sufficient frost tolerance the taproot size of plants must be rather small. Hence, the cultivation of bolting winter beet under Central European climate conditions has to face a severe conflict of goals concerning winter survival and yield formation.     

QTL analysis of Cercospora leaf spot resistance in sugar beet

The inheritance of Cercospora leaf spot resistance in sugar beet was investigated by means of quantitative trait loci (QTL) analysis of a segregating population of 193 individuals, using 110 AFLP and 35 restriction fragment length polymorphism (RFLP) markers. Five QTL were found through composite interval mapping on linkage groups 1, 2, 3, and 9, respectively, two of which were linked on linkage group 3. The significance of these QTL was tested by permutation analysis The QTL had mostly additive, but also certain negative dominance effects; all the resistance alleles came from the Cercospora-resistant parent. Each quantitative trait locus accounted for 7-18% of the phenotypic variation, leaving 37% of the variation unexplained. The results are discussed in relation to the potential use of marker-assisted breeding for Cercospora leaf spot resistance in sugar beet.     

Effect of Extending the Growing Period on Yield Formation of Sugar Beet

Autumn sown sugar beets (winter beets) are expected to yield markedly higher than spring sown beets. This requires a continuous growth during an extended growing period. So far, bolting‐resistant sugar beet varieties are not available to test winter beets under field conditions in Central Europe. The objective of this study was therefore to analyse yield formation and sugar storage of sugar beet plants during an extended growing period to estimate whether sugar beet has the potential to generate the theoretically expected yield increase. From 2008 to 2012, pot experiments were carried out in the glasshouse with 11 sowing dates spread over the years with sequential harvests. The oldest plants were grown for 859 days (14 242 °Cd). Root fresh matter yield continuously increased till the latest harvest. In contrast, the sugar concentration reached an optimum value between 3400 and 5000 °Cd and then decreased with time. Despite longer growing periods, the number of cambium rings, which are regarded as essential for sugar storage, did not change. This points to an early and genetically fixed determination of the formation of cambium rings. Additionally, the rate of photosynthesis decreased concomitantly with the sugar concentration. In conclusion, there is some evidence that the sugar concentration of the storage root is limited by the sink capacity, which in turn controls the source activity by a feedback regulation of photosynthesis and leaf formation. The dry matter composition of the storage root changed towards lower sugar concentration and concurrent higher concentration of cell wall compounds (marc). The sugar yield still increased beyond a thermal time at which winter beets will probably be harvested in practice. Hence, the theoretical yield increase in autumn sown sugar beets can be realized, provided that the plants show sufficient winter hardiness and bolting resistance.     

Identification and validation of an over‐dominant QTL controlling soybean seed weight using populations derived from Glycine max × Glycine soja

Heterosis, or hybrid vigour, has been used to improve seed yield in several important crops for decades and it has potential applications in soybean. The discovery of over‐dominant quantitative trait loci (QTL) underlying yield‐related traits, such as seed weight, will facilitate hybrid soybean breeding via marker‐assisted selection. In this study, F₂ and F_2 : 3 populations derived from the crosses of ‘Jidou 12’ (Glycine max) × ‘ZYD2738’ (Glycine soja) and ‘Jidou 9’ (G. max) × ‘ZYD2738’ were used to identify over‐dominant QTL associated with seed weight. A total of seven QTL were identified. Among them, qSWT_13_1, mapped on chromosome 13 and linked with Satt114, showed an over‐dominant effect in two populations for two successive generations. This over‐dominant effect was further examined by six subpopulations derived from ‘Jidou12’ × ‘ZYD2738’. The seed weight for heterozygous individuals was 1.1‐ to 1.6‐fold higher than that of homozygous individuals among the six validation populations examined in different locations and years. Therefore, qSWT_13_1 may be a useful locus to improve the yield of hybrid soybean and to understand the molecular mechanism of heterosis in soybean.     

The potato R10 resistance specificity to late blight is conferred by both a single dominant R gene and quantitative trait loci

The R10 late blight differential of potato, 3681ad1, exhibits good field resistance. Progeny from the cross between 3681ad1 and the susceptible cultivar ‘Katahdin’ were assessed for late blight resistance to three Phytophthora infestans isolates, using a detached leaf assay. Progeny differed in response to the three isolates. Resistance to isolates IPO‐0 and 99018 was controlled by quantitative trait loci (QTL), whereas resistance to isolate 89148‐9 was inherited as a dominant R gene, designated as R10 in this study. Statistical analysis revealed that one of the resistance QTLs to isolates IPO‐0 and 99018 is linked to the R10 gene, which maps to chromosome 11 in a region where a complex late blight resistance locus has been reported previously. A high‐resolution map of R10 was constructed using a large segregating population, and the gene was delimited to a genetic interval of 0.26 cM. The clustering of the qualitative gene R10 with resistance QTLs could explain the field resistance observed with 3681ad1.     

Overwintering of genetically modified sugar beet, Beta vulgaris L. subsp. vulgaris, as a source for dispersal of transgenic pollen

The potential impact of transgenic crops on community ecology will depend on the distribution and establishment of the new transgenic traits, on the sexual transfer of their new genes to the environment (Bartsch &; Pohl-Orf, 1996) and on the potential ecological impact of the transgenic trait. Flowering and pollen dispersal is important for outcrossing of the genetically engineered trait. For a biennial plant, like the cultivars of Beta vulgaris L., overwintering is normally necessary to become generative and to produce pollen and seeds (Abe et al., 1997), which usually does not happen with sugar beet as a field crop harvested in autumn (Longden 1989). The starting point for the project was a transgenic sugar beet, Beta vulgaris L. subsp. vulgaris (Lange et al., 1998), with rhizomania and herbicide ( Basta®, Liberty®) resistance. Cold tolerance is one of the most important factors for survival of sugar beet in Central- and North-Europe. Among other ways, spreading of transgenic traits into weed beet (Boudry et al., 1993) or wild beet can occur if genetically engineered – biennial – plants survive the winter, flower in spring and spread their pollen. Field experiments were performed with transgenic breeding lines and their hybrids, transgenic and non-transgenic hybrids with Swiss chard and three conventional beet cultivars to evaluate winter survival rates at seven different field sites. We could show that survival of sugar beet – transgenic as well as conventional ones – in Germany and at the Dutch border is possible. Survival rates were well correlated with temperature data and were unexpectedly high. Differences between sugar beet hybrids and breeding lines could be detected but not within different breeding lines or hybrids. There were no differences detectable between transgenic and non-transgenic plants. The data are crucial for the risk assessment of the release of transgenic sugar beet and are the basis for further experiments towards outcrossing and establishment.     

Gene expression analysis of four WIR1-like genes in floret tissues of European winter wheat after challenge with <Emphasis Type="Italic">G. zeae</Emphasis>

Fusarium head blight (FHB) is a highly destructive disease of wheat and other cereals which causes serious mycotoxin contaminations of grain. A number of molecular mapping studies led to the detection of QTL with small to moderate effects on FHB resistance in European winter wheat. Genes involved in the defence reaction of these genotypes remain largely unknown. WIR1 (wheat induced resistance 1) genes have been shown to be upregulated in cereals during attack of various fungal pathogens; however, their role in resistance is ambiguous. In this study, the expression of three WIR1 genes and a gene with high sequence similarity to WIR1 was investigated in European winter wheat genotypes after inoculation with Giberella zeae. Floret tissues of four winter wheat genotypes (Dream, Lynx, G16-92, Hussar) were challenged with G. zeae conidia or water (control) and sampled six times during 0–96 h after inoculation. Quantitative real-time PCR showed that all four genes were highly upregulated in the resistant genotypes compared to the susceptible ones. WIR1b and a gene with sequence similarity to WIR1 genes mapped to chromosome 5DS in the G16-92/Hussar mapping population. Two genes annotated as WIR1a mapped in the interval of a FHB resistance QTL on chromosome 7BS in the Dream/Lynx mapping population. These could be considered possible candidate genes for quantitative FHB resistance.     

Oligogenic control of resistance to soil‐borne viruses SBCMV and WSSMV in rye (Secale cereale L.)

Rye production in European growing areas is constrained by the soilborne cereal mosaic virus (SBCMV) and the wheat spindle streak mosaic virus (WSSMV). To date, no European rye cultivars are known to exhibit resistance against these viruses. In this study, we pursued a quantitative trait locus (QTL) mapping strategy to identify genomic regions for resistance to SBCMV and WSSMV in rye. Three populations, each comprising 100 lines segregating for resistance to SBCMV and/or WSSMV, were evaluated for disease response at two years in three locations in Germany where soils are naturally infested with SBCMV and WSSMV. In the combined analysis across environments, one QTL for SBCMV resistance on chromosome 5R explained 31.9% of the phenotypic variation in one of the populations. For WSSMV resistance, one QTL explaining up to 64.0% of the phenotypic variation was detected on chromosome 7R in each of the three populations. On the Triticeae homoeologous group 5, we found evidence for synteny of the major QTL for SBCMV resistance between the wheat and rye genomes.     

Rph23: A new designated additive adult plant resistance gene to leaf rust in barley on chromosome 7H

We report on a new adult plant resistance (APR) gene Rph23 conferring resistance to leaf rust in barley. The gene was identified and characterized from a doubled haploid population derived from an intercross between the Australian barley varieties Yerong (Y) and Franklin (F). Genetic analysis of adult plant field leaf rust scores of the Y/F population collected over three successive years indicated involvement of two highly additive genes controlling APR, one of which was named Rph23. The gene was mapped to chromosome 7HS positioned at a genetic distance 36.6 cM. Rph23 is closely linked to marker Ebmac0603, which is flanked by markers bPb‐8660 and bPb‐9601 with linkage distances of 0.8 and 5.1 cM, respectively. A PCR‐based marker was optimized for marker‐assisted selection of Rph23, and on the basis of this marker, the gene was postulated as being common in Australian and global barley germplasm. Pedigree and molecular marker analyses indicated that the six‐rowed black Russian landrace ‘LV‐Taganrog’ is the likely origin of Rph23.