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.     

Combining ability of maize inbred lines containing genes from Zea diploperennis for resistance to Striga hermonthica (Del.) Benth

Maize ( Zea mays ) is a staple food crop in sub-Saharan Africa, but its production is threatened by Striga hermonthica (Del.) Benth. Transfer of resistance genes from wild relatives may increase resistance to S. hermonthica in tropical maize. The objective of this study was to determine the combining ability of resistance to S. hermonthica among lines containing Zea diploperennis and tropical germplasm. Forty-five diallel crosses of 10 inbred lines were evaluated in an alpha-lattice design with and without artificial Striga infestation at two locations each in the Republic of Benin and Nigeria for 3 years. Results of analyses showed that only general combining ability (GCA) mean square was significant (P = 0.01) for number of emerged Striga plants (NESPP), while both GCA and specific combining ability (SCA) mean squares were significant for host damage score (HDS) and grain yield under Striga infestation. The ratio of GCA to SCA mean squares for the three traits varied from 3.5 to 57.5. Although GCA × environment interaction was significant for the three traits, two inbred lines containing Z. diploperennis (ZD 551) and tropical (TZL TC 87) germplasm had negative and significant GCA effects for NESPP and HDS and positive GCA effects for grain yield under Striga infestation in the two countries. Correlation between NESPP and HDS was strong and significant ( r  = 0.87, P = 0.01). Our results highlight the importance of harnessing useful genes from wild relatives to improve resistance to S. hermonthica in adapted maize germplasm.     

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.     

Development of synthetic maize populations for resistance to Striga hermonthica

The parasitic witchweed, Striga hermonthica (Del.) Benth, is the greatest biological constraint for cereal crop production by resource-poor farmers in sub-Saharan Africa. Maize, Zea mays L., is a widely grown cereal crop in this region (22 × 10⁶ ha). Striga-resistant maize populations were produced and tested as half-sib families in West and Central Africa. Three populations with white (W), yellow (Y), or mixed (Y/W) grain colour were formed by: (1) intercrossing Striga resistant maize inbred lines followed by two generations of random mating; (2) testing far under artificially induced S. hermonthica infestations in Nigeria, Cameroon, and Ivory Coast and selection of resistant families; (3) two generations of random mating; and (4) two years of testing for resistance. Striga-resistant synthetic W, Y and Y/W populations were produced by compositing resistant half-sib families. Outstanding performance in grain yields and host plant resistance was observed. Maize damage ratings and number of harvested ears were highly correlated with grain yield. High variation was observed for Striga emergence counts. The populations have combined resistance to Striga, maize streak virus (MSV), and other major biotic constraints for maize cultivation in Africa, thus providing the opportunity for improved sustainable maize production under stress environments. Breeder's seed of these synthetic varieties are being multiplied for distribution to national programmes.     

Genetic markers linked to Striga gesnerioides resistance for the improvement of Ghanaian cowpea (Vigna unguiculata) cultivars

The parasitic plant, Striga gesnerioides (Willd.) Vatke, is one of the most important constraints of cowpea production and food security in West Africa. Currently, few Striga resistant cowpea varieties have been developed that are well‐adapted to the dry savannah regions of Ghana. While genes conferring resistance to Striga races SG1, SG3 and SG5 have been mapped, the genetic locus of resistance to the race of Striga found in Ghana (SG‐GH) has not been characterized. Here, we report identification of genetic markers linked to SG‐GH resistance and define the relationship of this locus to SG3 resistance in recombinant inbred line populations generated from crosses between Striga resistant IT97K‐499‐35 and the Striga susceptible varieties Apagbaala and SARC‐LO2. The populations were genotyped with five genetic markers associated with SG3 and SG5 Striga resistance genes and a genetic map was developed. Genes conferring resistance to SG‐GH and SG3 mapped 4.2 cM from each other on chromosome Vu11. The identification of genetic markers linked to SG‐GH resistance will facilitate the marker‐assisted development of high‐quality Striga resistant cowpea varieties in Ghana.     

Expression of mature plant resistance to Striga hermonthica in maize

Twelve maize genotypes with different degrees of resistance were evaluated in plots inoculated with seeds of the phytoparasite Striga hermonthica (Del.) Benth. and in Striga-free (control) plots for three seasons between 1991 and 1993. Resistant genotypes, although showing similar levels of underground infection as susceptible 9 weeks after maize planting (WAP), had significantly fewer emerged parasitic plants and sustained lower damage. Host damage was not determined by the severity of infection. Percent height and dry matter reductions increased from 3 to 6 WAP for resistant and susceptible genotypes. While percent height and dry matter reductions for resistant genotypes declined at 9 and 12 WAP, susceptible genotypes either maintained or increased their levels of damage. Correlation between the severity of height and dry matter reductions at the vegetative (3 and 6 WAP) and reproductive (9 and 12 WAP) stages of maize growth were not significant, indicating that damage at early stages of growth cannot be used to predict mature plant response. Maize damage score (1–9) at 8 WAP, a non destructive and composite shoot indicator of host performance under Striga infestation, was significantly correlated (r = 0.88, r = 0.82; p < 0.01) to the level of shoot reduction at mature plant stages. Emerged Striga count at 8 WAP was significantly correlated (r = 0.98, p < 0.01) to the count at 10 WAP, the time when parasite emergence was highest. Under high and uniform infestation, mature plant resistance can be detected at 8 WAP, the onset of flowering in maize. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic diversity of Tainan-white maize inbred lines and prediction of single cross hybrid performance using RAPD markers

To evaluate the genetic diversity of 13 maize inbred lines, and to determine the correlation between genetic distance and single cross hybrid performance, we employed the randomly amplified polymorphic DNA(RAPD)- a PCR-based technique. Six of these lines came from the Taichung population, and others derived from seven different sites. Forty different primers were used to give a total of 646 reproducible amplification products, 547 (84.7%) of them being polymorphic. Genetic divergence was determined using Jaccard's similarity coefficient, and a final dendrogram was constructed by UPGMA (unweighted pair-group method with arithmetical averages) cluster analysis in the CLUSTER procedure of the SAS system. The RAPD analysis was a useful tool in determining the extent of genetic diversity among Tainan-white maize inbred lines in the present case. Cluster analysis showed that the 13 inbred lines could be classified into distinct heterotic groups. There was no significant linear regression of grain dry weight heterosis value and mean performance of hybrids on genetic distance. And their coefficients of determination(R²) are small, so that predictive value is limited. The present results showed that the Jaccard's similarity coefficients based on RAPD data cannot be used to precisely predict the F₁ hybrids yield performance and heterosis value. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic characterization of 218 elite CIMMYT maize inbred lines using RFLP markers

Characterization of genetic diversity among maize inbred lines can facilitate organization of germplasm and improve efficiency of breeding programs. A set of 218 phenotypically diverse inbred maize lines developed at CIMMYT for hybrid production was characterized using 32 RFLP markers to: (1) analyze the genetic diversity present; (2) define potential heterotic groups based on clusters formed with marker data; and (3) identify the most representative testers for each potential heterotic group. Lines were clustered using five different genetic distance measurements to find consensus non-hierarchical clusters. Dendrograms were produced to study hierarchical classification within smaller groups of lines. A very high average allelic diversity was seen in this germplasm. Lines did not cluster based on phenotype, environmental adaptation, grain color or type, maturity, or heterotic response (as determined based on hybrid performance with testers), but lines related by pedigree usually did cluster together. Previously defined testers from opposite heterotic groups were not genetically differentiated, and did not represent well their heterotic group. Discrete clusters were difficult to find; thus, potential heterotic groups will be difficult to suggest using RFLP markers alone. However, suggestions on how to use molecular markers and cross performance information to refine heterotic groups and select representative testers are presented.     

Genetic diversity of maize inbred lines in relation to downy mildew

A major emphasis in maize breeding in Asian countries has been the improvement for resistance to downy mildew, a serious disease that causes significant yield losses. A total of 102 inbred lines, including lines from Asian breeding programs, Mexico, USA and Germany, were analyzed with 76 SSR markers to measure diversity and investigate the effect of selection for downy mildew resistance. A mean polymorphism information content of 0.59, with a range of 0.14 to 0.83, was observed. Diversity at the gene level showed an average of 5.4 alleles per locus and a range of two to 16 alleles per locus, with a total of 409 alleles. About half of the alleles in the Asian lines had frequencies of 0.10 or less, and only 2% had frequencies > 0.80, indicating the presence of many alleles, and thus a high level of diversity. Some of the high-frequency alleles were in chromosomal regions associated with disease resistance. However, the frequencies of alleles in three SSR loci that are linked to a QTL for resistance to downy mildews in Asia were not significantly different in the subtropical/tropical Asian lines as compared to all the lines in the study. Lines from the US, Germany, and China, comprised three clusters of temperate maize(GS = 0.31), while those from India, Indonesia, Philippines, Thailand, Vietnam and CIMMYT comprised seven indistinct clusters of subtropical and subtropical maize (GS = 0.29). We conclude that maize breeding activity in Asia has not caused a decline in the overall amount of diversity in the region. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic diversity and population structure of the selected core set in Amaranthus using SSR markers

The genus Amaranthus has gained much attention, particularly for its high economic and nutritional value. It is a genus of taxonomic complexity with many interspecific hybrids. For effective conservation and management of the germplasm collected, the development of a core set of accessions is especially important. A core set of 63 accessions was successfully developed from an entire collection of 634 accessions using the powercore 1.0. Among the tested methods for developing the core set – the advanced M strategy using the modified heuristic method (HCC), randomly chosen collections and stratified random collections – HCC was found to be best, with 100% coverage of alleles and minimum redundancy. Allele frequency distribution of the core set developed with HCC was highly correlated with that of entire collections (r = 0.91). Model‐based structure analysis revealed the presence of three subpopulations and 11 admixtures in the selected core set, which is basically consistent with clustering based on the genetic distance. The results from this study will provide effective information for future germplasm conservation and improvement programmes in Amaranthus.     

黑龙江省部分审定玉米品种亲本自交系的遗传多样性分析

黑龙江省气候条件特殊,血缘关系有些混杂,系谱关系已不太清楚,种质资源匮乏问题尤为突出。利用SSR分子标记技术,分析黑龙江省近年来审定玉米品种亲本自交系的遗传多样性,从79对SSR核心引物目录中,选出43对引物对18份玉米自交系和5个标准测验种进行遗传多样性研究,共检测到174个等位基因位点,每对引物检测到2～8个等位基因,平均每个位点的等位基因变异数4.35个,平均多态性信息量为0.586。UPGMA聚类分析结果表明,23份自交系划分为4个类群。结果表明黑龙江省玉米生产上推广品种的亲本自交系仍然集中在兰卡斯特群和瑞德群两大杂种优势群。     

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 diversity in tropical maize inbred lines: heterotic group assignment and hybrid performance determined by RFLP markers

Tropical maize inbred lines, eight derived from a Thai synthetic population (BR‐105) and 10 from a Brazilian composite population (BR‐106), were assayed for restriction fragment length polymorphisms with 185 clone‐enzyme combinations. The aim of this study was to investigate genetic distances among tropical maize material and their relationship to heterotic group allocation and hybrid performance. Genetic distances (GDs) were on average greater for BR‐105×BR‐106 lines (0.77) than for BR‐106×BR‐106 (0.71) and for BR‐105×BR‐105 (0.69) lines. Cluster analysis resulted in a clear separation of BR‐105 and BR‐106 populations and was according to pedigree information. Correlations of parental GDs with single crosses and their heterosis for grain yield were high for line crosses from the same heterotic group and low for line combinations from different heterotic groups. Our results suggest that RFLP‐based GDs are efficient and reliable to assess and allocate genotypes from tropical maize populations into heterotic groups. However, RFLP‐based GDs are not suitable for predicting the performance of line crosses from genetically different heterotic groups.     

基于SNP和InDel标记的巴西木薯遗传多样性与群体遗传结构分析

为了对巴西木薯种质资源进行遗传多样性、亲缘关系和群体遗传结构分析,本研究利用了7946个SNPs和1997个InDels分子标记,通过ADMIXTURE软件进行群体结构分析、GCTA软件进行主成分分析。结果显示,巴西木薯被划分为9个亚群。这与利用PHYLIP进行的聚类分析结果大概一致,其中亚群1、亚群2、亚群4、亚群6和亚群8能较好地分别聚在一起,而其他亚群中的样品大致能聚在一起,且样品间有一定的交叉。巴西木薯种质资源遗传多样性指数(0.274)高于中国、尼日利亚等,其中巴西木薯亚群5具有相对较高的遗传多样性水平(0.29)。巴西木薯各亚群的群体遗传分化程度较低(群体分化指数在0.03~0.15之间),但高于中国木薯种质资源的群体分化指数。各木薯材料间的遗传距离变幅为0.084~0.297,平均遗传距离为0.228。本研究结果可为后续关联分析发掘优良等位基因及引种提供依据。     

基于SNP和InDel标记的巴西木薯遗传多样性与群体遗传结构分析

As a typical tropical crop, cassava (Manihot esculenta Crantz) has the characteristics of drought resistance, barren resistance, high biomass and so on. In addition to being used for food and forage, it can also be used for production, processing and starch extraction. Due to highly heterozygous cassava genome, breeding is more difficult. Enriching the genetic diversity of cassava germplasm, comprehensively evaluating its genetic background and traits, and discovering superior alleles that control excellent traits are of great significance for cassava breeding in the future. In order to analyze the genetic diversity, genetic relationship and population structure of cassava germplasm in Brazil, 7946 SNPs and 1997 InDels molecular markers were used. Population structure analysis was performed by ADMIXTURE software, and principal component analysis was performed by GCTA software. Brazilian cassava was divided into nine subgroups, and was roughly consistent with the results of cluster analysis using PHYLIP. Among them, subgroup 1, subgroup 2, subgroup 4, subgroup 6, and subgroup 8 could be clustered together respectively, while the samples of other subgroups could be roughly clustered, and there was a certain cross between the samples. The genetic diversity of cassava germplasm in Brazil (0.274) was higher than the genetic diversity level of cassava germplasm in China and Nigeria. Subgroup 5 of Brazil cassava had a relatively high genetic diversity (0.29). The genetic differentiation of subgroups was low (the genetic differentiation vary from 0.03 to 0.15), but higher than domestic cassava germplasm. The genetic distance between cassava accessions varied from 0.084 to 0.297, with the average of 0.228. The results of this study can provide a basis for subsequent association analysis to identify great alleles and introduction.     

Genetic diversity of oilseed Brassica napus inbred lines based on sequence-related amplified polymorphism and its relation to hybrid performance

Significant heterosis for seed yield in oilseed rape has created interest in the development of hybrid cultivars. The DNA‐based marker protocol, sequence‐related amplified polymorphism (SRAP) was used to determine genetic diversity among oilseed rape maintainer and restorer lines. This measure was used in an attempt to establish an association between genetic distance and heterosis in hybrids for various agronomic traits. A total of 118 polymorphic loci were generated by 18 SRAP primer combinations. Based on the polymorphism generated by the markers, calculated similarity index values ranged from 0.46 to 0.97. Cluster analysis grouped 10 maintainer and 12 restorer lines into three groups, with the exception of two maintainer lines, PM5 and PM9, which fell outside these groups. The grouping of the lines was largely in agreement with the available pedigree data on their origin and agronomic performance. Analysis of variance among inbred lines and their resulting F₁ hybrids over two locations revealed significant differences for plant height, days to maturity and seed yield, but not for oil content. Substantial mid‐parent heterosis was observed only for seed yield, and ranged from 26% to 169%. All hybrids surpassed their respective inbred lines for this trait, except for a single cross combination of related lines. In general, crosses of lines located in different clusters yielded more than those from the same clusters. Regression analysis revealed a statistically significant relationship between the genetic distance of the parents and seed yield in their hybrid, and their derived mid‐parent and high‐parent heterosis. The correlation coefficient between genetic distance and yield (0.64) indicated a moderately strong relationship, so it is possible that some of the SRAP markers might be linked to quantitative trait loci for seed yield.     

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.     

Identification of new sources of resistance to Striga gesnerioides in cowpea germplasm

There have been reports of breakdown of striga resistance in previously resistant cowpea cultivars in Burkina Faso. This could be attributed to new striga races emerging or to an increase in the aggressiveness of current striga races. Therefore, cowpea genotypes were evaluated in fields infested with S. gesnerioides at three striga hot spots in Burkina Faso and in pots under artificial infestation with striga races SR 1, SR 5 and SR Kp to identify new, adapted and striga‐resistant sources. Cowpea genotypes showed differential reactions for striga resistance over sites and for striga races in pot experiments, indicating differences in the races involved, and SR Kp was reported as a new race. Resistant sources conferring site‐specific or multiple striga‐race resistance were identified. Genotypes 58‐57, Sanga 2, IT84S‐2049, IT98K‐205‐8, IT93K‐693‐2, KVx771‐10, KVx775‐33‐2, KVx61‐1, Gorom local, Mouride and Melakh conferred resistance to all three striga races. These genotypes are potential donor parents for breeding new, adapted and striga‐resistant genotypes. Cowpea landraces including Moussa local and Niaogo local with farmers' preferred traits were susceptible and need improvement for striga resistance.     

几种分子标记技术在玉米自交系类群划分中的应用

分子标记技术的开发利用推动了玉米育种的发展,概述了5种常用分子标记SSR、RFLP、RAPD、AFLP、SNP的原理及特点。综述了分子标记技术在玉米自交系类群划分中的应用。