Inheritance of resistance to broomrape (Orobanche cumana Wallr.) race F in a sunflower line derived from wild sunflower species

Genetic resistance to broomrape (Orobanche cumana Wallr.) race F in sunflower line J1, derived from the wild perennial species Helianthusgrosseserratus Martens and Helianthus divaricatus L., has been reported to be controlled by dominant alleles at a single locus, Or6. However, deviations from this monogenic inheritance have been observed. The objective of the present study was to gain insight into the inheritance of resistance to broomrape race F in the sunflower line J1. F₁, F₂, F₃ and BC generations from crosses between J1 and three susceptible lines, P21, NR5 and HA821 were evaluated. F₁ hybrids showed both resistant (R) and moderately resistant (MR) plants, the latter having a maximum of five broomrape stalks per plant compared with >10 in the susceptible parents. This indicated incomplete dominance of the Or6 alleles. F₂ plants were classified as R, MR or susceptible (more than five broomrape stalks per plant). Three different segregation ratios were observed: 3 : 1, 13 : 3 and 15 : 1 (R + MR : S), suggesting the presence of a second gene, Or7, whose expression was influenced by the environment. A digenic model was confirmed, based on the evaluation of F_2:3 families.     

Screening for resistance to broomrape (Orobanche cernua) in cultivated sunflower

Racial evolution of sunflower broomrape (Orobanche cernua) has been very rapid in Spain during recent years, in which resistance has been overcome several times and there has been an important increase in areas infested with this parasitic angiosperm. In order to find resistance to a highly virulent population of sunflower broomrape that could be used directly in breeding programmes, three different sets of cultivated plant material composed of 429 entries were tested by artificial inoculation. All evaluated inbred lines from Moden, Canada, were fully susceptible. Out of the 240 P.I. accessions tested, only 10 segregated for resistance to broomrape, the rest being susceptible. From the 160 USDA breeding lines evaluated, 5% were resistant and 19% segregated for resistance to O. cernua. These lines traced back mainly to crosses of RHA 274 and RHA 801 with Russian, Turkish and Romanian hybrids. The origin of P.I. accessions that segregated for resistance were primarily derived from the former USSR and from Romania.     

Evaluation of Differential Lines and a Collection of Sunflower Parental Lines for Resistance to Broomrape (Orobanche cernua) in Spain

Broomrape (Orobanche cernua Loelf.) is a sunflower parasite which has become a potential threat to this crop in Spam with an increasing incidence in recent last years. In order to investigate the racial situation of the parasite in Spam and to evaluate the resistance of differentials and inbred lines currently used in breeding programmes, two experiments were carried out. In the first, two sets of European sunflower differentials were inoculated with three broomrape populations from different geographical origin. The second experiment studied the reaction to the parasite of 38 inbred lines in comparison with resistant and susceptible checks. Reactions of differential lines resistant to known European broomrape races ranged from complete immunity of lines P-1380-2A and ‘Jdanovski 8281’ to complete susceptibility of ‘Record’ and the universal suscepts. These results confirmed the existence of several physiological races of broomrape in Spain and a different racial composition from that of Eastern Europe. Results from the second experiment showed good resistance of half the restorer lines tested and 36 % of the maintainers and cms lines. This material together with the resistant differential lines constitutes good starting material to be used in developing cultivars resistant to broomrape populations in Spain.     

Reproductive behaviour and broomrape resistance in interspecific hybrids of sunflower

Interspecific hybrids and backcross generations between the wild perennial species Helianthus resinosus, Helianthus paucifiorus, Helianthus laevigatus, Helianthus nuttallii ssp. nuttallii T. & G. and Helianthus giganteus, resistant to broomrape (Orobanche cernua) and susceptible inbred lines were obtained to study crossability to cultivated sunflower and the transmission and expression of resistance to this parasitic weed. Conventional crosses with all the species tested were successful except for the crosses with diploid H. giganteus, for which embryo rescue techniques were needed to overcome hybrid incompatibility. Pollen viability and seed set were highest for F₁ hybrids with hexaploid species and lowest for those with the diploid H. giganteus. We evaluated F₁, BC₁F₁, some BC₂F₁ plants and the wild and cultivated parents. The wild species and interspecific hybrids were resistant to broomrape infection except for H. nuttallii, which showed segregation, indicating that the resistance is dominant. The crossability and resistance of F₁, and back-cross generations of species with different ploidy levels indicate that the transfer of broomrape resistance to cultivated sunflower is feasible.     

Identification of PCR markers linked to different Or genes in sunflower

Broomrape is a parasitic plant that significantly decreases yield of sunflower. Breeding for resistance has proved to be the most efficient method for suppressing broomrape infestation in the field; however, new races of parasite constantly emerge, and new resistance genes need to be discovered and introduced into cultivated sunflower lines. The aim of this work was to test SSR markers from linkage group 3 (LG3) to investigate whether they could be used for identification of a particular Or gene. Twenty sunflower inbred lines were used, and polymorphism between the lines with various resistance genes and genetic background was investigated. The used markers revealed DNA polymorphism between the investigated lines. Strong association of markers from LG3 with Or₆, as well as Or₄ and Or₂ genes, was found. Identified markers could be used for introduction of these resistance genes into commercial sunflower lines and for establishment and identification of differential lines.     

Inheritance of quantitative resistance to downy mildew (Plasmopara halstedii) in sunflower (Helianthus annuus L.)

Quantitative resistance to sunflower downy mildew was studied on inbred lines and hybrids not carrying efficient major gene resistance, in field trials in one to four sites over 3 years. Hybrids from factorial crosses showed that inheritance is under additive control and comparison with reactions of parental inbred lines gave narrow sense heritabilities of 27–57%. Analysis of a polymorphic recombinant inbred line population without efficient major gene resistance indicated that two highly significant Quantitative Trait Loci (QTL) explained 42% of variation in field reaction to downy mildew. These QTL were mapped on linkage groups 8 and 10, and do not appear related to any of the known major resistance gene clusters. Possible bases of this type of resistance and its use in breeding are discussed.     

Inheritance of resistance to race F of broomrape in sunflower lines of different origins

A new race F of broomrape overcomes all known resistance genes in cultivated sunflower, but recently, sources of resistance against race F have been developed. The objective of the present research was to study the inheritance of resistance to race F in crosses between 12 resistant sunflower breeding lines, derived from three different sources of resistance, and the susceptible male‐sterile line P‐21. Parental lines and F₁, F₂, F₃ and BC₁ generations were evaluated for broomrape resistance. Segregations in the F₂ and BC₁ to resistant parent approached resistant to susceptible ratios of 1: 15 and 1: 3, respectively, in most of the crosses, suggesting a double dominant epistasis. However, segregations of 3: 13 and 1: 1 for F₂ and BC₁, respectively, indicating a dominant‐recessive epistasis, were also found. The F₃ data confirmed these results. Owing to the recessive nature of this resistance, it must be incorporated into both parental lines for developing resistant hybrid cultivars.     

不同向日葵品种根系分泌物对向日葵列当萌发的刺激作用

以不同列当抗性向日葵根系分泌物作为萌发刺激物,研究其对向日葵列当种子萌发的影响,探究不同向日葵品种对列当抗性差异响应的机制,为向日葵列当抗性品种的选育和应用提供理论依据。研究表明：根系分泌物的稀释倍数对向日葵列当的萌发具有显著影响,其中TP-3313和SH363-M根系分泌物在稀释50倍时达到最大萌发率,而3638C根系分泌物在稀释75倍时达到最大萌发率,整体表现为低浓度促进、高浓度抑制的特点;不同列当抗性的向日葵根系分泌物诱导向日葵列当萌发的效果不同,向日葵列当萌发率表现为TP-3313相似文献   

Colocation of downy mildew (Plasmopara halstedii) resistance genes in sunflower (Helianthus annuus L.)

Summary The Pl6 locus in the inbred sunflower (Helianthus annuus L.) line HA335 giving resistance to French races of downy mildew (Plasmopara halstedii (Farl.) Berl. & de Toni. was localized by molecular techniques. A bulked segregant analysis was made on the F2 progeny from a cross between this line and H52, a downy mildew susceptible line. The resistance gene in HA335 was found to have the same linked RFLP marker loci as those determined for Pl1 (resistance to race 1 in the line RHA266) on linkage group 1 of the consensus RFLP map of the cultivated sunflower. Pl1 and Pl6 thus appear either to be allelic or closely linked. The implications for sunflower breeding are discussed.     

Resistance to Sclerotinia sclerotiorum of 'high oleic' sunflower inbred lines

Cultivation of sunflower (Helianthus annuus L.) is strongly affected by Sclerotinia sclerotiorum. To identify new sources of genetic diversity for sunflower breeding 25 sunflower inbred lines were analysed using eight Amplified Fragment Length Polymorphism (AFLP) primer combinations and their genetic similarities (GS) were estimated. Data were used to develop a Unweighted Pair Group Method using Arithmetic Averages (UPGMA) dendrogram. GS values of 0.58‐0.98 were observed but with no separate groupings dependant on oil quality. The inbred lines were screened for their reaction to inoculation with Sclerotinia sclerotiorum (Lib.) de Bary. Sunflower heads were artificially inoculated with S. sclerotiorum in three environments. Head infection was monitored after 1 week (lesion length) and 2 weeks (head rot). The F5 generation of a cross between a resistant (SWS‐B‐04) and a susceptible inbred line (SWS‐B‐01) was also tested for sclerotinia reaction across three environments. Significant differences in sclerotinia resistance, moderate heritabilities and a high correlation between the two assessments were observed. Inbred lines with a high level of resistance could be identified. These lines can be used for further breeding to improve sunflower sclerotinia resistance and to develop superior new hybrids.     

The genetics of resistance in sunflower capitula to Sclerotinia sclerotiorum measured by mycelium infections combined with ascospore tests

To enhance efficiency of breeding programmes for Sclerotinia sclerotiorum resistance in sunflower capitula, two separate resistance tests have been recommended. However, the time necessary to develop genotypes with two types of resistance makes this impractical. A strategy to combine the two tests was consequently proposed to reduce the number of seasons per selection cycle, but genetic studies were necessary to determine if it could be applied in breeding programmes. This was the objective of this work. Data from two genetically different sunflower inbred lines and their F1, F2 and backcross generations were analysed in two years to determine the genetics of resistance to S. sclerotiorum in capitula measured by application on the same plants of a mycelium test combined with an ascospore test. Effects of maternal origin were detected when the reciprocal generations were evaluated for the mycelium test. This suggests the importance of choice of the inbred line used as female in population formation by hybridisation. Progeny subjected to the mycelium test did not show any change in relative reactions to two S. sclerotiorum isolates suggesting that ranking genotypes according to the results of combined S. sclerotiorum tests is repeatable across Sclerotinia isolates and experimental seasons. Moderate narrow sense heritability indicated that selection of the best F2 plants should be effective. Genetic gain from selection is possible because a reduction of lesion areas produced by the mycelium test can be expected. Additive gene effects contribute significantly to reduction in lesion area. Consequences of results in population improvement for S. sclerotiorum resistance in capitula are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Optimal allocation of resources in evaluating current sunflower inbred lines for resistance to Sclerotinia

Sclerotinia sclerotiorum (Lib.) de Bary is a major limiting factor for sunflower (Helianthus annuus) production. This study was conducted to (1) evaluate a large number of current sunflower lines for their reaction to Sclerotinia, and (2) determine the optimum allocation of resources for their selection. A set of 85 inbred lines was screened under artificial leaf infection in three environments. The test revealed ample variation for all resistance traits in the current germplasm. A line selected from NDBLOS, an oilseed sunflower germplasm poo, and its derivatives were most resistant to Sclerotinia. Lines derived from interspecific crosses with the wild species Helianthus tuberosus and Helianthus argophyllus also showed satisfactory Sclerotinia resistance. The evaluation of five plants per plot in two replications and four environments appears to be a reasonable allocation of resources in order to optimize the selection response.     

向日葵列当寄生机理研究进展

总结归纳了温度、渗透势、预培养时间、生长调节物质及光照等预培养条件对向日葵列当种子萌发的影响,并对刺激向日葵列当种子萌发的信号物质和其吸器形成的诱导物质进行了综述,提出了向日葵列当寄生机理研究中存在的问题,如种子萌发机制不明确、萌发信号物质的分离与鉴定手段有待改进以及向日葵列当与向日葵间的识别机制问题等,同时对研究前景进行了展望。     

Somaclonal variants of sunflower with modified coumarin expression under stress

Summary Two coumarins, scopoletin and ayapin, have been shown to accumulate in sunflower following insect damage or fungal infection (Tal & Robeson, 1986a, 1986b & Olson, 1989). The technique of shoot regeneration from embryo-derived callus was used to produce somaclonal variants of sunflower with enhanced levels of coumarin expression after an applied stress. Five regenerates (R1–R5) of inbred lines RHA 297 produced viable seeds following self-pollination. One of the S1 families from these plants had significantly higher concentration of scopoletin (12.75 g/g fresh weight) compared to the parental inbred line (5.66 g/g). The increased levels of scopoletin in S1 plants were partially inherited after a second self-fertilization (S2 plants). The ploidy of the regenerates and their S1 and S2 progeny was not examined. No consistent correlation between higher levels of scopoletin in stimulated S1 and S2 plants, and deterrence to feeding of Heliothis virescens (Fab) larvae on unstimulated plants was shown after in vivo assays. From inbred line HA 300, three regenerates (H1–H3) were derived which produced S1 progeny with higher or lower concentrations of ayapin or scopoletin than the parental inbred. The inheritance of increased ayapin in the HA 300-derived lines was not assessed in the S2 generation because of poor seed germination. Feeding deterrence of H. virescens larvae was not consistently correlated with ayapin concentrations in S1 progeny of HA 300 regenerates. However, a correlation (-0.61) between ayapin concentration and daily feeding score was observed in one planting of these unstimulated S1 progeny. Sunflower germplasm was developed with enhanced levels of stress-induced coumarin expression through somaclonal variation and this germplasm may also show increased resistance to some leaf-feeding insects.     

新疆阿勒泰地区和内蒙古乌拉特前旗抗列当向日葵品种(系)的鉴定

向日葵列当是向日葵生产中重要的寄生性杂草。新疆阿勒泰地区和内蒙古乌拉特前旗是国内向日葵列当发生的重灾区。为筛选并为抗列当育种工作提供优质抗性材料,从根本上防治向日葵列当,利用田间自然发生向日葵列当的田块,采用随机区组设计设置小区试验;通过寄生率、寄生程度2 个指标判定对列当的抗感性。于2012 年在新疆阿勒泰对56 个向日葵品种（系）,2013—2014 年在内蒙古乌拉特前旗分别对22 个品种（系）和20 个品种（系）,进行抗列当鉴定。结果表明：在新疆22 个油葵品（系）中‘陇葵杂1 号’、‘F08-1’、‘辽丰F53’、‘赤CY101’、‘S26’、‘MGS’、‘S31’等16 个品种,34 个食葵品种（系）中‘JK106’品种,表现为免疫;在内蒙古乌拉特前旗2013—2014 年24 个油葵品种（系）中,‘赤CY102’、‘F08-1’等2 个品种表现为免疫,‘MSG’、‘新葵杂5 号’、‘赤029×115R’、‘赤128A×116R’等4个品种（系）表现为高抗,而18 个食葵品种（系）中,没有免疫品种,只有‘JK103’和‘JK106’表现为抗列当,其余16 个品种（系）均表现为感病和高度感列当。其中油葵品种‘赤CY102’在新疆和内蒙古抗列当鉴定中均表现免疫。在2 个不同地区进行向日葵品种抗列当鉴定,新疆地区表现免疫和高抗的品种较内蒙古地区多,说明内蒙古乌拉特前旗列当生理小种级别高于新疆阿勒泰地区列当生理小种。另外,油葵品种对列当的抗性较食葵高。     

Development of drought‐tolerant breeding lines derived from Helianthus annuus × H. argophyllus interspecific crosses

Helianthus argophyllus is a wild species known as “silver sunflower”, which possesses several traits, including morphological traits that increase drought tolerance. Therefore, introgressions between chosen cultivated lines and two H. argophyllus accessions were made, and segregating generations were established. Important agronomic traits including single heading, high pollen fertility, silver canopy (indicating more cuticular wax), and content of cuticular waxes and oil were selected over six segregating generations. The resulting F₆ lines showed introgression of water saving traits, as they had lower excise leaf water loss, with comparable yield to standard checks. However, these F₆ lines were late maturing and showed poor flowering synchronization between the cultivated and introgressed lines. Introgressed line “D‐22” was particularly promising as a breeding line, with superior agronomic and drought resistance traits. This line had the potential to be used as an inbred parental line for introgression of drought resistance traits into elite sunflower germplasm. Combining ability analysis of the introgressed lines further showed their potential for heterosis breeding or to be used as parental lines in breeding programme.     

Field evaluation of somaclonal variation in sunflower (Helianthus annuus L) and its application for crop improvement

Immature zygotic embryos from the American fertility restorer line RHA-857 were used as donor material for induction of direct organogenesis in sunflower (Helianthusannuus L.). The range of spontaneous somaclonal variation among the progenies of regenerants was studied. The genetic modifications observed in regenerants included agronomic traits such as oil content in seed, 1000 seed weight, plant height, leaf width, leaf length, petiole length, internode length, head diameter, number of branches, length of branches, number of ray florets, seed width, seed length, and seed thickness. RAPD molecular analysis carried out on sunflower materials in the R-11 generation showed the absence of a specific 358 bp band in somaclonal line 11/2/51 R. This line showed a modified architecture, full resistance to Phomopsis helianthiand higher oil content in seed in comparison to the standard RHA-857. Line31/3/53 R was with modified architecture and higher 1000 seed weight. Hybrid No. 144 produced with the participation of somaclonal line 20/5/52 R demonstrated high production capacity, shorter vegetation period and reduced height. The combination of these favourable changes is desirable in breeding work on sunflower. Somaclonal variation through director ganogenesis has facilitated the creation of genetically heritable variation in sunflower, which can be used with great success for hybrid seed production of highly productive hybrids. This revised version was published online in July 2006 with corrections to the Cover Date.     

Interspecific hybrids of sunflower as a source of Sclerotinia resistance

Sclerotinia sclerotiorum is considered the most devastating pathogen of sunflower grown in humid environments. In this study, progenies of partial hybrids between Helianthus maximiliani, a wild species that has been shown to be resistant to S. sclerotiorum, and H. annuus were characterized by amplified fragment length polymorphism (AFLP) analysis to identify whether there are introgressions from H. maximiliani into the cultivated sunflower at the molecular level. Wild species‐specific fragments as well as fragments not found in either parent were detected. Progenies tended to cluster together according to the original partial hybrids in the dendrogram by the use of bootstrap procedures. The progenies were studied for their reaction to S. sclerotiorum using artificial inoculation of sunflower heads. Some of the progenies showed a higher level of resistance compared with resistant inbred lines. It was possible to identify two AFLP‐fragments which seem to be linked to Sclerotinia resistance.     

Interspecific hybridization between sunflower and wild perennial Helianthus species via embryo rescue

Interspecific crosses were made between the cultivated sunflower inbred line HA89 and accessions of five wild perennial Helianthus species (H. giganteus L., H. laevigatus T. & G., H. resinosus Small, H. pauciflorus Nutt. and H. decapetalus L.) resistant to broomrape ( Orobanche cernua Loefl.). Using the genetic male-sterile isogenic version of that line as female, successful reciprocal crosses were also obtained with all these wild species except with H. decapetalus. Five-day-old hybrid embryos were excised and cultured in vitro. In all cases, few mature plants were obtained from embryos in early developmental stages (early heart and globular) but up to 28% mature plants were obtained from embryos in later stages of development. Very immature embryos were difficult to excise without damage. Hybrid embryos and mature plants were obtained from all five wild species. Interspecific hybrid embryos from different wild species showed distinct developmental potentials, the proportion of hybrid embryos in different developmental stages varying among species. Differences in the proportion of hybrid embryos at the several developmental stages were also obtained for the reciprocal crosses (cultivated × wild), which showed higher proportion of fully developed embryos. Hybrids involving H. giganteus and cultivated sunflower were difficult to obtain without the use of embryo culture. Embryo culture proved to be an useful tool to overcome post-zygotic hybrid incompatibility in different Helianthus spp., and facilitated interspecific transfer of resistance to O. cernua. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of Striga hermonthica adaptive traits in an early‐maturing white maize inbred line containing resistance genes from Zea diploperennis

Striga hermonthica can cause as high as 100% yield loss in maize depending on soil fertility level, type of genotype, severity of infestation and climatic conditions. Understanding the mode of inheritance of Striga resistance in maize is crucial for introgression of resistance genes into tropical germplasm and deployment of resistant varieties. This study examined the mode of inheritance of resistance to Striga in early‐maturing inbred line, TZdEI 352 containing resistance genes from Zea diploperennis. Six generations, P₁, P₂, F₁, F₂, BC₁P₁ and BC₁P₂ derived from a cross between resistant line, TZdEI 352 and susceptible line, TZdEI 425 were screened under artificial Striga infestation at Mokwa and Abuja, Nigeria, 2015. Additive‐dominance model was adequate in describing observed variations in the number of emerged Striga plants among the population; hence, digenic epistatic model was adopted for Striga damage. Dominance effects were higher than the additive effects for the number of emerged Striga plants at both locations signifying that non‐additive gene action conditioned inheritance of Striga resistance. Inbred TZdEI 352 could serve as invaluable parent for hybrid development in Striga endemic agro‐ecologies of sub‐Saharan Africa.