Breeding cowpea for resistance to Striga gesnerioides in the Nigerian dry savannas using marker‐assisted selection

Historically, conventional breeding has been the primary strategy used to develop a number of Striga‐resistant varieties currently grown in the Sahel of Western Africa. In this study, we have successfully developed and applied a marker‐assisted selection strategy that employs a single backcross programme to introgress Striga resistance into farmer preferred varieties of cowpea for the Nigeria savannas. In this strategy, we have introduced the Striga resistance gene from the donor parent IT97K‐499‐35 into an elite farmer preferred cowpea cultivar ‘Borno Brown’. The selected 47 BC₁F₂ populations confirmed the recombinants with desirable progeny having Striga resistance gene(s). The 28 lines selected in the BC₁F_2:4 generation with large seed size, brown seed coat colour and carrying marker alleles were evaluated in the field for resistance to Striga resistance. This led to the selection of a number of desirable improved lines that were immune to Striga having local genetic background with higher yield than those of their parents and standard varieties.     

Genetic analysis of Fusarium wilt resistance in cowpea (Vigna unguiculata Walp.)

This study investigated the inheritance of resistance to Fusarium oxysporum f.sp. tracheiphilum (Fot) in cowpea lines. Resistant and susceptible cowpea lines were crossed to develop F₁, F₂ and backcross populations. Reaction to Fot was evaluated in 2015 and 2016 using seed soak and modified root‐dip inoculation methods. The expression of resistance reaction in the F₁ and segregation in F₂ generations indicated the role of dominant gene controlling Fot in cowpea. These results were further supported by the result of backcross (BC₁P₁F₁ and BC₁P₂F₁) progeny tests. The backcross of F₁ with the resistant parent produced progeny that were uniformly resistant, whereas backcross of F₁ with the susceptible parent produced progeny that segregated into 1:1 ratio. The F₂ segregation ratio in the reciprocal cross showed no evidence of maternal effect in the inheritance of the resistance. Allelism test suggests that the gene for resistance in TVu 134 was the same in TVu 410 and TVu 109‐1. We also identified an SSR marker, C13‐16, that cosegregated with the gene conferring resistance to Fot in cowpea.     

Efficiency of secondary traits in selecting for improved grain yield in extra‐early maize under Striga‐infested and Striga‐free environments

A base index involving Striga damage, number of emerged Striga plants and ears per plant is used for selecting for maize (Zea mays L.) grain yield under Striga infestation. There are contradictory reports on the reliability of number of emerged Striga plants for selecting for Striga resistance. The objective of this study was to confirm reliability of the secondary traits for selecting for improved grain yield under Striga infestation. Ten Striga‐resistant extra‐early cultivars were evaluated for 3 years under artificial Striga‐infested and Striga‐free environments in Nigeria. Analysis of variance combined across years and locations showed significant mean squares for genotype, year, location and their interactions for most traits. Sequential path analysis identified ear aspect as the only trait with significant direct effect on yield under artificial Striga infestation, while GGE biplot confirmed ear aspect, ears per plant and Striga damage as the most reliable traits. Ear aspect should be included in the base index for selecting for improved grain yield of extra‐early maize under Striga infestation, while the number of emerged Striga plants should be excluded.     

Inheritance of resistance to the peach root‐knot nematode (Meloidogyne floridensis) in interspecific crosses between peach (Prunus persica) and its wild relative (Prunus kansuensis)

The peach root‐knot nematode, Meloidogyne floridensis (MF), infects majority of available nematode‐resistant peach rootstocks which are mostly derived from peach (Prunus persica) and Chinese wild peach (P. davidiana). Interspecific hybridization of peach with its wild relative, Kansu peach (P. kansuensis), offers potential for broadening the resistance spectrum in standard peach rootstocks. We investigated the inheritance of resistance to MF in segregating populations of peach (‘Okinawa’ or ‘Flordaguard’) × P. kansuensis. A total of 379 individuals from 13 F₂ and BC₁F₁ families were challenged with a pathogenic MF isolate “MFGnv14” and were classified as resistant (R) or susceptible (S) based on root galling intensity. Segregation analyses in F₂ progeny revealed the involvement of a major locus with a dominant or recessive allele determining resistance in progeny segregating 3R:1S and 1R:3S, respectively. Testcrosses with a homozygous‐susceptible peach genotype (‘Flordaguard’ or ‘UFSharp’) confirmed P. kansuensis as a source of new resistance and the heterozygous allelic status of P. kansuensis at the locus conferring resistance to MF. We propose a single‐locus dominant/recessive model for the inheritance of resistance.     

Genetic analysis of grain mould resistance in coloured sorghum genotypes

Grain moulds are a major constraint to sorghum production and to adoption of improved cultivars in many tropical areas. Information on the inheritance of grain mould reaction is required to facilitate breeding of resistant cultivars. The genetic control of grain mould reaction was studied in 7 crosses of 2 resistant sorghum genotypes. P₁, P₂, F₁, F₂, BC₁ and BC₂ families of each cross were evaluated under sprinkler irrigation for field grade and threshed grade scores and subjected to generation mean analysis. Frequency distributions for grain mould reaction were derived and F₂ and BC₁ segregation ratios were calculated. Grain mould reaction in crosses of coloured grain sorghum was generally controlled by two or three major genes. Resistance to grain moulds was dominant. Significant additive gene effects were also found in all cross/season combinations. Significant dominance effects of similar magnitude to additive effects were also observed in five out of ten cross/season combinations. Gene interactions varied according to the parents with both resistant and susceptible parents contributing major genes. Choice of parents with complementary resistance genes and mechanisms of resistance will be critical to the success of resistance breeding. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance pattern of mungbean yellow mosaic disease resistance and gene action for different traits in mungbean (Vigna radiata (L.) Wilczek) under protected and unprotected field conditions

Understanding of mungbean yellow mosaic disease resistance is crucial to develop resistant varieties to combat worldwide threat of the disease epidemics. This study was aimed to determine nature of disease resistance in terms of number of genes governing it and genetics of related traits. Experimental site was located on 31.43°N and 73.06°E with an elevation of 186 m, and evaluation trials were conducted during spring season due to high disease epidemics in this season. Two contrasting genotypes, that is, NM 6‐68‐2 (resistant) and NM 1‐32‐1 (highly susceptible), were crossed to raise six populations, that is, P₁, P₂, F₁, F₂, BC₁ and BC₂ for evaluation under protected and unprotected field conditions using chi‐square test and generation mean and variance analysis. It was discovered that disease resistance was governed by two major genes with additive effects. Disease resistance can easily be incorporated through backcrossing and direct selection following hybridization. Direct selection should be practised at earlier generation for plant height, chlorophyll contents and number of seeds per pod due to preponderance of additive effects whereas at later generations for seed yield per plant, 100 seed weight, harvest index, number of pods per plant and pod length due to involvement of duplicate epistasis.     

Genetic effects of the soft starch (h) and background loci on volume of starch granules in five inbreds of maize

Larger particle volume is beneficial for many aspects of maize starch processing, and may improve the performance of some starch attributes. This study focused on the soft starch (h) locus to identify its potentially influential role in starch particle volume distribution. The objectives were to study the genetic expression of starch particle volume of the h locus in different genetic backgrounds and the gene action conditioning starch particle volume of other loci in both normal‐starch and h‐starch backgrounds. Forty‐five populations (five intra‐inbred F_1s, 10 hybrid F_1s 10 F_2s, 10 BC₁F_1s to h/h parent, and 10 BC₁ to h:h conversion of normal parent) were planted in 1993 at two locations and in 1995 at one location. Selfed heterozygotes (±/h) in all generations provided intra‐ear comparisons of normal and h starch, and F₃ and BC₁F₂ generations provided inter‐ear comparisons. Significant differences were found between normal and h:h genotypes in all intra‐ear and inter‐ear comparisons. In all cases, general combining ability effects were highly significant, suggesting the presence of additive gene effects. Generation mean analysis of normal and h:h starch materials yielded similar results, indicating the predominance of additive and some dominance effects for other loci on starch particle volume. These results indicate the usefulness of the soft starch gene and additional genetic variation among inbreds in the improvement of starch particle volume for increased starch recovery in wet milling.     

Assessing the suitability of stress tolerant early-maturing maize (Zea mays) inbred lines for hybrid development using combining ability effects and DArTseq markers

Identification of hybrids for commercialization is crucial for sustainable maize production in sub-Saharan Africa (SSA). One hundred and ninety test crosses, 10 tester × tester crosses + 10 hybrid checks were evaluated across 11 environments, 2017 to 2019. Inheritance of grain yield under Striga infestation, optimal and across environments was influenced by additive genetic action, but there was greater influence of nonadditive gene action under drought stress conditions. Nine, seven and two inbreds had significant and positive general combining ability (GCA) effects for grain yield under Striga-infested, optimal and drought stress environments, respectively, and would contribute high grain yield to their progenies. Heterotic grouping methods based on specific and GCA, GCA effects of multiple traits and DArTseq markers classified the inbreds into five, three and two heterotic groups, respectively, across research conditions. The DArTseq markers method that classified the inbred lines into two major heterotic groups and was one of the most efficient methods should be adopted for practical purposes in maize breeding programmes in SSA. Hybrids TZEI 7 × TZdEI 352, TZEI 1238 × TZEI 7 and TZEI 1252 × TZEI 7 had outstanding grain yield under contrasting environments and should be tested on-farm for commercialization in SSA.     

Genetic analysis of grain mold resistance in white seed sorghum genotypes

Grain molds in rainy season sorghums can cause poor grain quality resulting in economic losses. Grain molds are a major constraint to the sorghum production and for adoption of the improved cultivars. A complex of fungi causes grain mold. Information on genetics of grain mold resistance and mechanisms is required to facilitate the breeding of durable resistant cultivars. A genetic study was conducted using one white susceptible, three white resistant/tolerant sources, and one colored resistant source in the crossing programme to obtain four crosses. P₁, P₂, F₁, BC₁, and BC₂, and F₂ families of each cross were evaluated for the field grade and threshed grade scores, under sprinkler irrigation. Generation mean analyses and frequency distribution studies were carried out. The frequency distribution studies showed that grain mold resistance in the white-grained resistance sources was polygenic. The additive gene action and additive × additive gene interaction were significant in all the crosses. Simple recurrent selection or backcrossing should accumulate the genes for resistance. Epistasis gene interactions were observed in colored resistance × white resistance cross. Gene interaction was influenced by pronounced G × E. Pooled analysis showed that environment × additive gene interaction and environment × dominant gene interaction were significant. The complex genetics of mold resistance is due to the presence of different mechanisms of inheritance from various sources. Evaluation of segregating population for resistance and selection for stable derivatives in advanced generations in different environments will be effective.     

Generation means analysis of resistance to peanut bud necrosis caused by peanut bud necrosis tospovirus in peanut

This study was conducted to evaluate the types of gene action governing the inheritance of resistance to peanut bud necrosis disease (PBND) in populations derived from three crosses involving two resistant (ICGV 86388 and IC 10) and one susceptible (KK 60–1) peanut lines. Populations were composed of P₁ P₂, F₁ F₂, BC₁₁, BC₁₂, BC₁₁S and BC₁₂S. These populations were evaluated for PBND incidence in a farmer's field in Kalasin province in north‐east Thailand, where PBND is a recurring problem. Results showed variations between crosses in the relative contributions of different types of gene effect. The results indicate that multiple genes control the PBND resistance trait, and that the two resistant lines differ in some of these genes. As non‐additive gene effects are important in all three crosses, selection for low PBND incidence in these crosses would be more effective in later generations.     

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.     

Assessment of reactions of diverse maize inbred lines to Striga hermonthica (Del.) Benth

Striga hermonthica is the most widespread and destructive obligate root parasite infecting maize and other cereals in Africa. Maize inbred lines supporting reduced S. hermonthica emergence can form an important basis for developing Striga‐resistant maize cultivars. Twenty new inbred lines selected for field resistance to S. hermonthica, and five inbred checks with known resitance, tolerance and susceptibility reactions to S. hermonthica were evaluated in pots, greenhouse and field experiments under artificial Striga infestation for 3 years. The experiments were conducted to determine the extent of variation in parasite attachment to the roots of these lines and its relationship with emerged Striga plants and other traits. Significant differences (P < 0.0001) were detected among the inbred lines for the numbers of attached and emerged Striga plants and the results were consistent across test environments. Also, the lines exhibited significant differences for Striga damage symptom ratings and other traits recorded in the field. Parasite attachment to the roots was significantly correlated with emerged Striga count in the screenhouse (r = 0.67–0.68, P < 0.001) and in the field (r = 0.82–0.84, P < 0.0001) and with levels of grain yield reduction due to Striga (r = 0.71, P < 0.0001). Regression analysis of the numbers of attached parasites on the first principal component axis scores that integrated several traits recorded in the field was significant (P < 0.0001) and accounted for 62% of the total variation in numbers of attached parasites. The new inbred lines and the resistant inbred check were the least affected by S. hermonthica and exhibited yield losses of 0–37% compared with the yields of the tolerant and the susceptible inbred checks, which were reduced by 40–85%. Sixteen new inbred lines supported significantly fewer attached parasites compared with the susceptible inbred check. Some of these lines also supported significantly fewer emerged parasites and sustained lower damage symptoms and percentage yield losses due to Striga compared with the susceptible inbred check. These inbred lines would be useful in breeding programmes for developing resistant maize cultivars.     

Inheritance and allelic relationship among gene(s) for blast resistance in pearl millet [Pennisetum glaucum (L.) R. Br.]

Six blast‐resistant pearl millet genotypes, ICMB 93333, ICMB 97222, ICMR 06444, ICMR 06222, ICMR 11003 and IP 21187‐P1, were crossed with two susceptible genotypes, ICMB 95444 and ICMB 89111 to generate F₁s, F₂s and backcrosses, BC₁P₁ (susceptible parent × F₁) and BC₁P₂ (resistant parent × F₁) for inheritance study. The resistant genotypes were crossed among themselves in half diallel to generate F₁s and F₂s for test of allelism. The F₁, F₂ and backcross generations, and their parents were screened in a glasshouse against Magnaporthe grisea isolates Pg 45 and Pg 53. The reaction of the F₁s, segregation pattern of F₂s and BC₁P₁ derived from crosses involving two susceptible parents and six resistant parents revealed the presence of single dominant gene governing resistance in the resistant genotypes. No segregation for blast reaction was observed in the F₂s derived from the crosses of resistant × resistant parents. The resistance reaction of these F₂s indicated that single dominant gene conferring resistance in the six genotypes is allelic, that is same gene imparts blast resistance in these genotypes to M. grisea isolates.     

Analysis of generation means and components of variance for parthenocarpy in cucumber (Cucumis sativus L.)

Parthenocarpy (seedless fruit) has potential for increasing yield in cucumber (Cucumis sativus var. sativus L.). To determine the inheritance of parthenocarpy in gynoecious cucumber, P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂ generations derived from crossing two non‐parthenocarpic gynoecious inbred lines [Gy8 (P₂; processing type) and ‘Marketmore 80’ (P₂; MM, fresh market type)] with a highly parthenocarpic inbred line [2A (P₁; processing type)] were evaluated for fruit number in a greenhouse at Arlington, Wisc. in 1999 (designated 2A × Gy8 1999) and in the open‐field at Hancock, Wisc. in 2000 (designated 2A × Gy8 2000 and 2A × MM 2000). There were significant location and location × generation interaction effects, and therefore generation means analyses were conducted separately for each location. The minimum numbers of effective factors controlling parthenocarpy were estimated to be at least one (2A × Gy8 1999), two (2A × Gy8 2000) and four (2A × MM 2000). Results suggest that selection for parthenocarpy for multiple hand harvest operations will likely be more effective than that for once‐over machine harvest operations. However, the selection efficiency will likely vary across different populations and environments.     

Development of molecular markers linked to the Leptosphaeria maculans resistance gene Rlm6 and inheritance of SCAR and CAPS markers in Brassica napus × Brassica juncea interspecific hybrids

Leptosphaeria maculans causes blackleg disease on Brassica napus, an economically important oilseed crop. Brassica juncea has high resistance to blackleg and is a source for the development of resistant B. napus varieties. To transfer the Rlm6 resistance gene from B. juncea into B. napus, an interspecific cross between B. napus “Topas DH16516” and B. juncea “Forge” was produced, followed by the development of F₂ and F₃ generations. Sequence characterized amplified region (SCAR) and cleaved amplified polymorphic sequence (CAPS) markers linked to the L. maculans resistance gene Rlm6 were developed. Segregation of SCAR and CAPS markers linked to Rlm6 were confirmed by genotyping of F₂ and F₃ progeny. Segregation of CAPS markers and phenotypes for blackleg disease severity in F₂ plants had a Mendelian ratio of 3:1 in resistant vs. susceptible plants, respectively, supporting the assumption that genetic control of resistance was by a single dominant gene. The molecular markers developed in this study, which show linkage with the L. maculans resistance gene Rlm6, would facilitate marker‐assisted backcross breeding in a variety development programme.     

Stripe rust resistance gene Yr18 and its suppressor gene in Chinese wheat landraces

The slow‐rusting and mildewing gene Yr18/Lr34/Pm38/Sr57 confers partial, durable resistance to multiple fungal pathogens and has its origins in China. A number of diagnostic markers were developed for this gene based on the gene sequence, but these markers do not always predict the presence of the resistant phenotype as some wheat varieties with the gene are susceptible to stripe rust in China. We hypothesized that these varieties have a suppressor of Yr18. This study was undertaken to determine the presence of Yr18, the suppressor and/or another resistance gene in 144 Chinese wheat landraces using molecular markers and stripe rust field data. Forty‐three landraces were predicted to have Yr18 based on the presence of the markers, but had final disease severities higher than 70%, indicating that this gene may be under the influence of a suppressor. Four of these landraces, ‘Sichuanyonggang 2’, ‘Baikemai’, ‘Youmai’ and ‘Zhangsihuang’, were chosen for genetic studies. Crosses were made between the lines and ‘Avocet S’, with further crosses of Sichuanyonggang 2 ×  ‘Huixianhong’ and Sichuanyonggang 2 ×  ‘Chinese Spring’. The F₁ plants of Sichuanyonggang 2/Chinese Spring was susceptible indicating the presence of a dominant suppressor gene. The results of genetic analyses of F_2:3 and BC₁F₂ families derived from these crosses indicated the presence of Yr18, a Yr18 suppressor and another additive resistance gene. The Yr18 region in Sichuanyonggang 2 was sequenced to ensure that it contained the functional allele. This is the first report of a suppressor of Yr18/Lr34/Pm38/Sr57 gene with respect to stripe rust response.     

Inheritance of insensitivity to culture filtrate of Pyrenophora tritici-repentis, race 2, in wheat (Triticum aestivum L.)

Tan spot of wheat is caused by the fungus Pyrenophora tritici‐repentis. On susceptible hosts, P. tritici‐repentis induces two phenotypically distinct symptoms, tan necrosis and chlorosis. This fungus produces several toxins that induce tan necrosis and chlorosis symptoms in susceptible cultivars. The objectives of this study were to determine the inheritance of insensitivity to necrosis‐inducing culture filtrate of P. tritici‐repentis, race 2, and to establish the relationship between the host reaction to culture filtrate and spore inoculation with respect to the necrosis component. The F₁, F₂, and BC₁F₁ plants and F_2:8 lines of five crosses involving resistant wheat genotypes ‘Erik’, ‘Red Chief’, and line 86ISMN 2137 with susceptible cultivars ‘Glenlea’ and ‘Kenyon’ were studied. Plants were spore‐inoculated at the two‐leaf stage. Four days later, the newly emerged uninoculated third leaf was infiltrated with a culture filtrate of isolate Ptr 92–164 (race 2). Reactions to the spore inoculation and the culture filtrate were recorded 8 days after spore inoculation. The segregation observed in the F₂ and BC₁F₁ generations and the F_2:8 lines of all crosses indicated that a single recessive gene controlled insensitivity to necrosis caused by culture filtrate. This gene also controlled resistance to necrosis induced by spore inoculation.     

A temporal genetic analysis of disease resistance genes: resistance to melon vine decline derived from Cucumis melo var. agrestis

Melon vine decline causes severe economic losses to melon crops all over the world. Two fungi. Acremonium cucurbitacearum and Monosporascus cannonballus, have been reported as the main causal agents of root damage, responsible for the decrease in root water and nutrient uptake capacity. However, melon vine decline aetiology is not completely understood, so screening assays must be conducted under natural infestation conditions. In field assays, the erratic effect of uncontrolled environmental factors makes the study of the trait's genetics difficult. The percentage of asymptomatic plants scored at individual moments during the infection process in the field is an imprecise indicator of the resistance level of each genotype. The analysis of disease progress curves does allow for the minimization of the stochastic fluctuations caused by environmental factors at various moments in time. Using this analysis the mode of inheritance of the partial resistance to melon vine decline derived from the accession Pat 81 of Cucumis melo var. agrestis was further studied. Data from the six generations assayed (susceptible and resistant parents. F₁, F₂, and backcrosses BC₁ and BC₂ were fitted to an additive/dominance model without epistatic effects using a scaling test. Monogenic control was proposed as the simplest model to explain the data. Assuming this simple model, the method of analysis also allowed for the characterization of the incomplete penetrance of resistance, which varies as a log probit function of the time. Simple genetic control would simplify the use of Pat 81 in breeding programmes aimed at developing commercial melon varieties with resistance to vine decline.     

Development of a SCAR marker linked with a MYMV resistance gene in mungbean (Vigna radiata L. Wilczek)

Yellow mosaic disease (YMD) caused by mungbean yellow mosaic virus (MYMV) is the most important disease of mungbean, causing great yield loss. The present investigation was carried out to study the inheritance and identify molecular markers linked with MYMV resistance gene by using F₁, F₂ and 167 F_2 : 8 recombinant inbred lines (RILs) developed from the cross ‘TM‐99‐37’ (resistant) × Mulmarada (susceptible). The F₁ was susceptible, F₂ segregated in 3S:1R phenotypic ratio and RILs segregated in 1S:1R ratio in the field screening indicating that the MYMV resistance gene is governed by a single recessive gene. Of the 140 RAPD primers, 45 primers showing polymorphism in parents were screened using bulked segregant analysis. Three primers amplified specific polymorphic fragments viz. OPB‐07_600, OPC‐06₁₇₅₀ and OPB‐12₈₂₀. The marker OPB‐07₆₀₀ was more closely linked (6.8 cM) with a MYMV resistance gene as compared to OPC‐06₁₇₅₀ (22.8 cM) and OPB‐12₈₂₀ (25.2 cM). The resistance‐specific fragment OPB‐07₆₀₀ was cloned, sequenced and converted into a sequence‐characterized amplified region (SCAR) marker and validated in twenty genotypes with different genetic backgrounds.