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BackgroundOrange maize genotypes are sources of provitamin A (PVA) carotenoids, which are precursors of vitamin A. PVA deficiency and drought constitute major challenges causing increasing food and nutritional insecurity in sub-Saharan Africa (SSA). Breeding of drought-tolerant provitamin A hybrid maize can mitigate these challenges. This study was undertaken to determine the combining ability of newly developed extra-early orange inbreds for grain yield and related traits under managed drought stress and rain-fed conditions, determine the mode of gene action conditioning the inheritance of the traits, and classify the inbreds into heterotic groups.MethodologyOne hundred and ninety-six extra-early orange hybrids comprising 180 testcrosses, 10 single crosses, and 6 commercial checks were evaluated under managed drought and rain-fed conditions at Ikenne. In addition, 41 inbreds comprising 36 orange lines and 5 PVA testers involved in the hybrid development were assessed under drought and rain-fed conditions.ResultsThe means square for general combining ability (GCA) and specific combining ability (SCA) were significant for grain yield and most other traits under both growing conditions. The contributions of GCA to performance were larger than SCA in each growing condition. Broad-sense and narrow-sense heritability estimates for grain yield were 66% and 37% under managed drought and 88% and 32% under rain-fed conditions, respectively. Mid-parent heterosis and better-parent heterosis for grain yield were 338% and 247% under managed drought, while 173% and 137% under rain-fed conditions. Significant positive correlations existed among grain yield of hybrids, heterosis, and specific combining ability under managed drought. The 41 inbred lines were classified into three heterotic groups under both growing conditions. Sixteen testcross hybrids out-yielded the best commercial check under managed drought.ConclusionThe testcross hybrids have great potential for commercialization to address the problem of drought and PVA deficiency in SSA. Inbred TZEEIOR 510 showed desirable GCA effects for grain yield and 04 other traits under drought.

Abstract Background Maize production in lowland agro-ecologies in West and Central Africa is constrained by the fungus Exserohilum turcicum, causal agent of Northern Corn Leaf Blight (NCLB). Breeding for resistance to NCLB is considered the most effective management strategy. The strategy would be even more effective if there is adequate knowledge of the characteristics of E. turcicum in a target region. Maize leaves showing NCLB symptoms were collected during field surveys in three major maize growing areas in Nigeria: Ikenne, Ile-Ife, and Zaria during 2018/2019 and 2019/2020 growing seasons to characterize E. turcicum populations interacting with maize using morphological and molecular criteria. Results A total of 217 E. turcicum isolates were recovered. Most of the isolates (47%) were recovered from the Ikenne samples while the least were obtained from Zaria. All isolates were morphologically characterized. A subset of 124 isolates was analyzed for virulence effector profiles using three primers: SIX13-like, SIX5-like, and Ecp6. Inter- and intra-location variations among isolates was found in sporulation, growth patterns, and presence of the effectors. Candidate effector genes that condition pathogenicity and virulence in E. turcicum were found but not all isolates expressed the three effectors. Conclusion Morphological and genetic variation among E. turcicum isolates was found within and across locations. The variability observed suggests that breeding for resistance to NCLB in Nigeria requires selection for quantitative resistance to sustain the breeding efforts.

In West and Central Africa (WCA), drought and low soil nitrogen (low N) impede increased maize (Zea mays L.) productivity and production. Due to climate change, the two stresses usually occur together, leading to food, nutritional, and economic insecurity in the sub-region. There is, therefore, the need for the development and availability of high-yielding extra-early maturing white Quality Protein Maize (QPM) synthetics and hybrids with resilience to the prevailing stresses through the identification of superior climate smart (extra-early maturing) QPM inbreds under stress (drought and low soil N) conditions. The study was conducted to identify stress-resilient QPM inbred lines for hybrid production and assess the association between grain yield and other studied characters. During the 2012 minor and major rainy seasons, 96 extra-early white QPM inbreds and four (4) normal endosperm maize inbred checks were assessed in multi-location trials under stress and optimal conditions in Nigeria. The experiments were laid out in a 10 × 10 simple lattice design with two replications. Data were recorded on grain yield and other agronomic traits. Significant variations (p < 0.01) were detected among the inbred lines for measured characters, indicating adequate genetic variability among the inbreds to allow for selection and improvement of grain yield and other measured traits. Grain yield was interrelated with all the traits used in the selection index. Moderate to high estimates of heritability were observed for most of the measured traits under stress conditions, indicating that the traits could be easily transmitted to the progenies. Fifty-seven out of the 96 QPM inbreds evaluated exhibited varying degrees of resilience to drought and low N. The QPM inbreds with desired traits may be used as genetic resources for the incorporation of tolerance genes into QPM populations in the tropics, as well as for the development of drought and low N resilient synthetics and hybrids in WCA.

Frequent occurrence of drought, heat, low soil fertility and Striga infestation are the main stress factors reducing maize yield in the Sahel. Adoption of stable multiple stress tolerant maize cultivars in the region is crucial for achieving food security. However, selection of a stable high yielding cultivar is complicated by genotype × environment interaction (GEI) due to differential responses to growing conditions. Eleven extra-early maturing multiple-stress tolerant maize hybrids and two checks arranged in a randomized complete block design was evaluated across nine locations for two years in Mali and Niger. The objectives of this study were to identify (i) stable and high-yielding maize hybrids, and (ii) suitable test locations for selecting promising extra-early maize hybrids. GGE biplot was used for graphical analysis. Significant genotype, location and GEI effects were detected for grain yield and number of ears per plant. EEWQH-13 produced the highest grain yield (3860 kg ha−1) while EEYQH-1 had the poorest yield (2663 kg ha−1) with trial mean of 3395 kg ha−1 for all hybrids. GGE biplot explained 69.6 % of the total variation in grain yield among the hybrids. The polygon view identified EEWQH-13 as the best hybrid across six of the nine test locations. EEPVAH-58 was identified as the most stable high yielding hybrid across the nine test locations followed by EEWQH-16 and EEWQH-13. The nine locations were clustered under two mega-environments (ME1, ME2). Among the nine test locations, Tara and Aderaoua clustered in ME1 were the most suitable ones for selecting promising extra-early maize hybrids for wider adaptation. The three hybrids, EEPVAH-58, EEWQH-16, and EEWQH-13, identified in this study could be recommended for on-farm evaluation to confirm the consistency of their yield performance for possible release and commercialization in Mali and Niger.

Maize production in sub-Saharan Africa (SSA) faces challenges due to the damage caused by the parasitic weed, Striga hermonthica (Del.) Benths and low soil nitrogen. To address these constraints and improve food security and nutrition, this study assessed the combining ability of 47 inbred lines and four testers, grouped them into heterotic groups, identified effective testers, and determined the stability of the lines in hybrid combinations under contrasting research conditions. The study was conducted at Mokwa and Abuja during the 2019 and 2020 growing seasons. One hundred and ninety-six hybrids comprising 188 testcrosses, 6 hybrids derived by intermating the four testers, and two commercial checks were evaluated using a 14 × 14 lattice design with two replicates. Results revealed that under Striga infestation, the best quality protein maize (QPM) hybrid, TZEEQI 468 × TZEEQI 321, outyielded the best check, TZEEQI 342 × TZEEQI 7, by 24%. Under low-N, QPM hybrid, TZEEQI 515 × TZEEQI 321 outyielded the best check, TZEEQI 507 × TZEEQI 7 by 11% while under optimal conditions the best QPM hybrid, TZEEQI 506 × TZEEQI 321 outyielded the best check, TZEEQI 342 × TZEEQI 7 by 2%. General combining ability (GCA) and specific combining ability (SCA) significantly influenced grain yield and other measured traits across the test environments. These indicated the importance of both additive and non-additive genetic variances in trait inheritance. GCA was more important than SCA for grain yield and most traits in contrasting environments. Four inbred lines had significant and positive GCA effects for grain yield under Striga-infested conditions, while three lines had similar GCA effects under low-nitrogen conditions. These lines demonstrated outstanding potential for developing Striga-resistant and low-nitrogen-tolerant hybrids. The study identified four heterotic groups using the heterotic grouping method based on the general combining ability of multiple traits (HGCMAT). Inbred lines TZEEQI 490 and TZEEQI 460 were identified as testers. The QPM hybrid TZEEQI 515 × TZEEQI 321 exhibited outstanding yield and stability across contrasting environments, highlighting the need for extensive on-farm trials to confirm its superiority and potential for commercialization in SSA.

The consistently low yield turnout of maize on farmers’ fields owing to drought and the nutritional challenges attributable to the consumption of white endosperm maize pose a major threat to food and nutritional security in Sub-Saharan Africa (SSA). The objectives of this study were to assess the performance of newly developed extra-early maturing orange hybrids under managed drought and well-watered conditions, compare the outcomes of multiple-trait base index and multi-trait genotype–ideotype distance index selection procedures, and identify drought-tolerant hybrids with stable performance across contrasting environments for commercialization in SSA. One hundred and ninety orange hybrids and six checks were evaluated under managed drought and well-watered conditions at Ikenne for two seasons between 2021 and 2023. A 14 × 14-lattice design was used for the field evaluations under both research conditions. Drought stress was achieved by the complete withdrawal of irrigation water 25 days after planting. Results revealed significant differences among the hybrids under drought and well-watered conditions. Grain yield, ears per plant, and plant aspect under managed drought were correlated to the same traits under well-watered conditions, suggesting that the expression of these traits is governed by common genetic factors. Twenty-nine hybrids were identified as top-performing drought-tolerant hybrids by the multiple-trait base index and the multi-trait genotype–ideotype distance index. Of the selected outstanding 29 hybrids, 34% were derived from crosses involving the tester TZEEIOR 197, demonstrating the outstanding genetic potential of this inbred line. Further analysis of the 29 selected hybrids revealed TZEEIOR 509 × TZEEIOR 197 as the hybrid that combined the most drought-tolerant adaptive traits. However, the hybrids TZEEIOR 526 × TZEEIOR 97, TZEEIOR 384 × TZEEIOR 30, TZEEIOR 515 × TZEEIOR 249, TZEEIOR 510 × TZEEIOR 197, TZEEIOR 479 × TZEEIOR 197, and TZEEIOR 458 × TZEEIOR 197 were identified as the most stable hybrids across drought and well-watered conditions. These hybrids should be extensively tested in multi-location trials for deployment and commercialization in SSA.

Low soil nitrogen (low-N), Striga hermonthica infestation and vitamin A deficiency in normal endosperm maize are major challenges confronting maize production and nutrition of the people of sub-Saharan Africa (SSA). Development of pro-vitamin A (PVA) maize hybrids with combined resistance/tolerance to the two stress factors is crucial in mitigating the food insecurity and nutrition challenges resulting from low-N deficiency and Striga infestation. One hundred and fifty hybrids plus six hybrid checks were evaluated under low-N, Striga-infested and optimal conditions in Nigeria for two years. The study examined the combining ability of the PVA inbreds in contrasting environments, classified them into heterotic groups, examined the inter-relationships of grain yield and other agronomic traits of the inbreds in hybrid combinations and assessed the performance and stability of the PVA hybrids across environments. Additive gene action conditioned the inheritance of grain yield under Striga infestation and optimal conditions while non-additive gene action played the major role in the inheritance of grain yield under low-N. Hybrids TZEEIOR 217 × TZEEIOR 197 and TZEEIOR 245 × TZEEIOR 195 were the top-yielding under Striga and low-N conditions, respectively. Inbred lines were classified into three heterotic groups. Inbreds TZEEIOR 195 and TZEEIOR 221 were identified as testers while TZEEIOR 197 × TZEEIOR 82, TZEEIOR 132 × TZEEIOR 195 and TZEEIOR 205 × TZEEIOR 221 were identified as single-cross testers. Ear aspect had direct contribution to grain yield, justifying its inclusion in the multiple trait base index used for selection of Striga resistant and low-N tolerant maize genotypes in SSA. Hybrids TZEEIOR 195 × TZEEIOR 149, TZEEIOR 195 × TZEEIOR 150, TZEEIOR 245 × TZEEIOR 195, TZEEIOR 30 × TZEEIOR 150 and TZEEIOR 245 × TZEEIOR 30 were high-yielding and stable across test environments. These hybrids should be tested extensively in on-farm trials and commercialized to contribute to food security in SSA.

AbstractIdentification of genes associated with StrigaStrigaStrigaStrigaStrigaStriga

Abstract Background Assessment and effective utilization of genetic diversity in breeding programs is crucial for sustainable genetic improvement and rapid adaptation to changing breeding objectives. During the past two decades, the commercialization of the early and extra-early maturing cultivars has contributed to rapid expansion of maize into different agro-ecologies of sub-Saharan Africa (SSA) where maize has become an important component of the agricultural economy and played a vital role in food and nutritional security. The present study aimed at understanding the population structure and genetic variability among 439 early and extra-early maize inbred lines developed from three narrow-based and twenty-seven broad-based populations by the International Iinstitute of Tropical Agriculture Maize Improvement Program (IITA-MIP). These inbreds were genotyped using 9642 DArTseq-based single nucleotide polymorphism (SNP) markers distributed uniformly throughout the maize genome. Results About 40.8% SNP markers were found highly informative and exhibited polymorphic information content (PIC) greater than 0.25. The minor allele frequency and PIC ranged from 0.015 to 0.500 and 0.029 to 0.375, respectively. The STRUCTURE, neighbour-joining phylogenetic tree and principal coordinate analysis (PCoA) grouped the inbred lines into four major classes generally consistent with the selection history, ancestry and kernel colour of the inbreds but indicated a complex pattern of the genetic structure. The pattern of grouping of the lines based on the STRUCTURE analysis was in concordance with the results of the PCoA and suggested greater number of sub-populations (K = 10). Generally, the classification of the inbred lines into heterotic groups based on SNP markers was reasonably reliable and in agreement with defined heterotic groups of previously identified testers based on combining ability studies. Conclusions Complete understanding of potential heterotic groups would be difficult to portray by depending solely on molecular markers. Therefore, planned crosses involving representative testers from opposing heterotic groups would be required to refine the existing heterotic groups. It is anticipated that the present set of inbreds could contribute new beneficial alleles for population improvement, development of hybrids and lines with potential to strengthen future breeding programs. Results of this study would help breeders in formulating breeding strategies for genetic enhancement and sustainable maize production in SSA.

Maize (Zea mays L.) breeders in the West and Central Africa have developed and commercialized extra-early and early-maturing maize hybrids, which combine high yield potentials with tolerance/resistance to drought, low soil-N and Striga infestation. Hybrids of both maturity groups have not been investigated for tolerance to high plant density and N application and are new to the farmers; thus, the urgent need to recommend appropriate agronomic practices for these hybrids. We investigated the responses of four hybrids, belonging to the extra-early and early-maturity groups, to three plant densities and three N rates in five locations of different agroecologies. The early-maturing hybrids consistently out-yielded the extra-early maturing hybrids in all the five agroecologies. The hybrids showed no response to N-fertilizer application above 90 kg ha−1. All interactions involving N had no significant effect on grain yield and other measured agronomic traits except in few cases. The extra-early and early-maturing hybrids had similar response to plant density; their grain yield decreased as density increased. Contrarily, flowering was delayed and expression of some other agronomic traits such as plant and ear aspects were negatively impacted by increased density. Optimal yield for hybrids of both maturity groups was obtained at approximately 90 kg N ha−1 and 66,666 plants ha−1. Most of the measured traits indicated high repeatability estimates across the N levels, densities and environments. Evidently, the hybrids were intolerant of elevated density. It therefore, becomes necessary to improve maize germplasms for high plant density tolerance in the region.

Abstract Background Information on population structure and genetic diversity of germplasm in a breeding programme is useful because it enhances judicious utilisation of genetic resources to achieve breeding objectives. Seventy early maturing provitamin A (PVA) quality protein maize (QPM) inbreds developed by the IITA- maize improvement programme were genotyped using 8171 DArTseq markers. Furthermore, 96 hybrids derived from 24 selected inbreds plus four checks were evaluated under low-N and optimal environments in Nigeria during 2016 and 2017. Genotypic and phenotypic data of inbreds and hybrids respectively, were analysed to (i) assess the level of genetic dissimilarities and population structure of the inbreds, and (ii) investigate the grain yield performance of derived hybrids under low-N, optimal and across environments. Results Genetic diversity among the seventy inbreds was high varying from 0.042 to 0.500 with an average of 0.357. Sixty-six inbred lines with probabilities ≥0.70 were assigned to a single group. The population structure analysis, the UPGMA phylogeny, and the principal Coordinate Analysis (PCoA) of the DArTseq markers revealed a clear separation of five groups and each followed pedigree records. Clustered inbreds displayed common characteristics including high PVA levels, and drought and low-N tolerance. The top performing hybrid, TZEIORQ 40 × TZEIORQ 26 out-yielded the best hybrid control, TZEIOR 127 × TZEIOR 57 by 8, 3, and 9% under low-N, optimal, and across environments, respectively. High repeatability estimates were detected for grain yield under each and across environments. Similarly, high breeding efficiency of 71, 70 and 72% were computed under low-N, optimal, and across environments, respectively. Conclusions The UPGMA clustering, the structure analysis, and the PCoA consistently revealed five groups which largely followed pedigree information indicating the existence of genetically distinct groups in the inbred lines. High repeatability and breeding efficiency values estimated for grain yield of hybrids under low-N, optimal and across environments demonstrated that high productive hybrids could be developed using inbreds from the opposing clusters identified by the DArTseq markers. The 15 top performing hybrids identified, particularly TZEIORQ 40 × TZEIORQ 26 and TZEIORQ 29 × TZEIORQ 43 should be further evaluated for release and commercialization in SSA.

Abstract Background Striga hermonthica (Benth.) parasitism militates against increased maize production and productivity in savannas of sub-Saharan Africa (SSA). Identification of Striga resistance genes is important in developing genotypes with durable resistance. So far, there is only one report on the existence of QTL for Striga resistance on chromosome 6 of maize. The objective of this study was to identify genomic regions significantly associated with grain yield and other agronomic traits under artificial Striga field infestation. A panel of 132 early-maturing maize inbreds were phenotyped for key agronomic traits under Striga-infested and Striga-free conditions. The inbred lines were also genotyped using 47,440 DArTseq markers from which 7224 markers were retained for population structure analysis and genome-wide association study (GWAS). Results The inbred lines were grouped into two major clusters based on structure analysis as well as the neighbor-joining hierarchical clustering. A total of 24 SNPs significantly associated with grain yield, Striga damage at 8 and 10 weeks after planting (WAP), ears per plant and ear aspect under Striga infestation were detected. Under Striga-free conditions, 11 SNPs significantly associated with grain yield, number of ears per plant and ear aspect were identified. Three markers physically located close to the putative genes GRMZM2G164743 (bin 10.05), GRMZM2G060216 (bin 3.06) and GRMZM2G103085 (bin 5.07) were detected, linked to grain yield, Striga damage at 8 and 10 WAP and number of ears per plant under Striga infestation, explaining 9 to 42% of the phenotypic variance. Furthermore, the S9_154,978,426 locus on chromosome 9 was found at 2.61 Mb close to the ZmCCD1 gene known to be associated with the reduction of strigolactone production in the maize roots. Conclusions Presented in this study is the first report of the identification of significant loci on chromosomes 9 and 10 of maize that are closely linked to ZmCCD1 and amt5 genes, respectively and may be related to plant defense mechanisms against Striga parasitism. After validation, the identified loci could be targets for breeders for marker-assisted selection (MAS) to accelerate genetic enhancement of maize for Striga resistance in the tropics, particularly in SSA, where the parasitic weed is endemic.

Abstract Availability of multiple-stress tolerant maize is critical for improvement in maize production in West and Central Africa (WCA). A study was carried out to (i) assess a set of inbred lines for combining ability under stressed and optimal conditions, (ii) determine the performance of the testcrosses under different conditions, and (iii) identify outstanding hybrids across the conditions. Two hundred and five testcrosses were planted with five hybrid checks under Striga-infested, low soil nitrogen, drought and optimal conditions between 2015 and 2016 in Nigeria. The grain yield inheritance under optimal condition was largely regulated by additive gene effect whereas non-additive gene effects largely regulated grain yield under the three stresses. Four of the inbreds had significant positive general combining ability effects each under low N and drought, and three under Striga infestation for grain yield. The inbreds could be vital sources of beneficial alleles for development and improvement of tropical yellow maize hybrids and populations. Hybrids TZEI 443 x ENT 13 and TZEI 462 x TZEI 10 were high yielding and stable; they out-performed the three early maturing released hybrids in WCA. The new hybrids should be extensively assessed and released in the sub-region to improve food security.

Information on the genetic diversity, population structure, and trait associations of germplasm resources is crucial for predicting hybrid performance. The objective of this study was to dissect the genetic diversity and population structure of extra-early yellow and orange quality protein maize (QPM) inbred lines and identify secondary traits for indirect selection for enhanced grain yield under low-soil nitrogen (LN). One hundred and ten inbred lines were assessed under LN (30 kg ha -1) and assayed for tryptophan content. The lines were genotyped using 2500 single nucleotide polymorphism (SNP) markers. Majority (85.4%) of the inbred lines exhibited wide pairwise genetic distances between 0.4801 and 0.600. Genetic distances were wider between yellow and orange endosperm lines and predicted high heterosis in crosses between parents of different endosperm colors. The unweighted pair group method with arithmetic mean (UPGMA) and the admixture model-based population structure method both grouped the lines into five clusters. The clustering was based on endosperm color, pedigree, and selection history but not on LN tolerance or tryptophan content. Genotype by trait biplot analysis revealed association of grain yield with plant height and ear height. TZEEQI 394 and TZEEIORQ 73A had high expressivity for these traits. Indirect selection for high grain yield among the inbred lines could be achieved using plant and ear heights as selection criteria. The wide genetic variability observed in this study suggested that the inbred lines could be important sources of beneficial alleles for LN breeding programs in SSA.

Information on combining ability and heterotic patterns of multiple stress-tolerant inbred lines are fundamental prerequisites for devising appropriate breeding strategies for the development of climate-resilient maize hybrids. In the present study, we evaluated 150 single cross hybrids derived from the North Carolina Design II (NCD II) along with six commercial checks under terminal drought stress (TDS), heat stress (HS), and combined drought and heat stress (CHDS)conditions. The objectives of the study were to: (i) determine the combining ability of the inbred lines and identify the best testers across the stresses; (ii) classify the inbred lines into heterotic groups (HGs) based on the general combining ability of multiple traits (HGCAMT) and sequencing-based diversity array technology (DArTseq) and (iii) assess the performance and stability of the lines in hybrid combinations. The inbred lines showed significantly (p < 0.01 and p < 0.05) positive and negative general combining ability (GCA) and specific combining ability (SCA) effects for grain yield (GY) and most other measured traits. The inbred line TZEI 135 displayed relatively larger positive GCA effects for GY when mated either as male or female and was identified as the best tester. TZEI 135 × TZEI 182 was identified as the best single-cross tester across environments. Results of the assessment of the relative importance of GCA and SCA effects revealed the predominance of additive gene action over the non-additive. Six HGs of inbreds were identified using the HGCAMT and three, based on the DArTseq marker genetic distance method, were the most efficient. The best hybrids in this study significantly out-yielded the best checks by 21, 46, and 70% under CHDS, HS, and TDS, respectively. These hybrids should be extensively tested in on-farm trials for possible commercialization in sub-Saharan Africa.

Maize (Zea mays L.) is an important staple, as well as cash crop, in sub-Saharan Africa (SSA). However, its production is severely constrained by low soil nitrogen (low N). Fifty-four early-maturing hybrids developed during three breeding periods, (2008–2010, 2011–2013 and 2014–2016) were evaluated under low (30 kg ha−1) and high (120 kg ha−1) soil nitrogen (N) in Ile-Ife and Mokwa, Nigeria, from 2017 to 2019. The study was designed to (i) determine the genetic gains in grain yield of the early-maturing maize hybrids developed during the three breeding periods, (ii) determine the relationship between grain yield and other agronomic traits and (iii) identify the highest-yielding and most stable hybrids under low- and high-N environments. The 54 hybrids were evaluated using a 6 × 9 lattice design with three replications. Mean squares for hybrids were significant for measured traits under low- and high-N environments, except the mean squares for stalk lodging and EPP under low N. Annual genetic gains in grain yield were 75 kg ha−1 year−1 (2.91%) and 55 kg ha−1 year−1 (1.33%) under low- and high-N environments, respectively, indicating that substantial gains were achieved in the genetic enhancement of the early-maturing hybrids. The hybrids TZdEI 314 × TZdEI 105, TZdEI 378 × TZdEI 173, ENT 12 × TZEI 48 and TZdEI 352 × TZdEI 315 were identified as the highest-yielding and most stable across test environments and should be tested extensively on farms and commercialized in SSA.

Availability of maize (Zea mays L.) hybrids with elevated provitamin A (PVA) levels and tolerance to contrasting stresses would improve food self-sufficiency and combat malnutrition in sub-Saharan Africa (SSA). This study was conducted to (i) analyze selected PVA inbreds of extra-early maturity for carotenoid content, (ii) estimate the combining abilities of the inbred lines for grain yield and other agronomic traits, (iii) assign inbred lines to distinct heterotic groups (HGs), (iv) identify testers among the inbred lines, and (v) determine grain yield and stability of the PVA hybrids across contrasting environments. Thirty-three extra-early maturing inbred lines selected for high carotenoid content were crossed with four inbred testers to obtain 132 testcrosses. The testcrosses, six tester × tester crosses and two hybrid checks, were evaluated across three Striga-infested, four drought and five optimal growing environments in Nigeria, 2014–2016. Results of the chemical analysis revealed that inbred lines TZEEIOR 109, TZEEIOR 30, TZEEIOR 41, TZEEIOR 97, TZEEIOR 42, and TZEEIOR 140 had intermediate PVA levels. Both additive and nonadditive gene actions were important in the inheritance of grain yield and other measured traits under stress and optimal environments. However, additive gene action was preponderant over the nonadditive gene action. The inbred lines were classified into three HGs across environments. Inbreds TZEEIOR 249 and TZEEIOR 30 were identified as testers for HGs I and II, respectively. The hybrid TZEEI 79 × TZEEIOR 30 was the most outstanding in terms of grain yield and was stable across environments. This hybrid should be tested extensively in on-farm trials for consistency in performance and commercialized to combat malnutrition and food insecurity in SSA.

An increase in the average global temperature and drought is anticipated in sub-Saharan Africa (SSA) as a result of climate change. Therefore, early white quality protein maize (QPM) hybrids with tolerance to combined drought and heat stress (CDHS) as well as low soil nitrogen (low-nitrogen) have the potential to mitigate the adverse effects of climate change. Ninety-six early QPM hybrids and four checks were evaluated in Nigeria for two years under CDHS, low-nitrogen, and in optimal environments. The objectives of this study were to determine the gene action conditioning grain yield, assess the performance of the early QPM inbred lines and identify high yielding and stable QPM hybrids under CDHS, low-nitrogen and optimal environment conditions. There was preponderance of the non-additive gene action over the additive in the inheritance of grain yield under CDHS environment conditions, while additive gene action was more important for grain yield in a low-nitrogen environment. TZEQI 6 was confirmed as an inbred tester under low N while TZEQI 113 × TZEQI 6 was identified as a single-cross tester under low-nitrogen environments. Plant and ear aspects were the primary contributors to grain yield under CDHS and low-nitrogen environments. TZEQI 6 × TZEQI 228 and the check TZEQI 39 × TZEQI 44 were the highest yielding under each stress environment and across environments. Hybrid TZEQI 210 × TZEQI 188 was the most stable across environments and should be tested on-farm and commercialized in SSA.

Striga is an important biotic factor limiting maize production in sub-Saharan Africa and can cause yield losses as high as 100%. Marker-assisted selection (MAS) approaches hold a great potential for improving Striga resistance but requires identification and use of markers associated with Striga resistance for adequate genetic gains from selection. However, there is no report on the discovery of quantitative trait loci (QTL) for resistance to Striga in maize under artificial field infestation. In the present study, 198 BC1S1 families obtained from a cross involving TZEEI 29 (Striga resistant inbred line) and TZEEI 23 (Striga susceptible inbred line) plus the two parental lines were screened under artificial Striga-infested conditions at two Striga-endemic locations in Nigeria in 2018, to identify QTL associated with Striga resistance indicator traits, including grain yield, ears per plant, Striga damage and number of emerged Striga plants. Genetic map was constructed using 1,386 DArTseq markers distributed across the 10 maize chromosomes, covering 2076 cM of the total genome with a mean spacing of 0.11 cM between the markers. Using composite interval mapping (CIM), fourteen QTL were identified for key Striga resistance/tolerance indicator traits: 3 QTL for grain yield, 4 for ears per plant and 7 for Striga damage at 10 weeks after planting (WAP), across environments. Putative candidate genes which encode major transcription factor families WRKY, bHLH, AP2-EREBPs, MYB, and bZIP involved in plant defense signaling were detected for Striga resistance/tolerance indicator traits. The QTL detected in the present study would be useful for rapid transfer of Striga resistance/tolerance genes into Striga susceptible but high yielding maize genotypes using MAS approaches after validation. Further studies on validation of the QTL in different genetic backgrounds and in different environments would help verify their reproducibility and effective use in breeding for Striga resistance/tolerance.

Vitamin A deficiency is the leading cause of night blindness, total blindness, maternal and childhood mortality in developing countries. Drought, low soil nitrogen and Striga hermonthica parasitism are major constraints to maize production in sub-Saharan Africa (SSA). Thus, the development of multiple stress tolerant maize varieties with elevated levels of PVA is an economically feasible approach to simultaneously tackle malnutrition and food insecurity in SSA. The objectives of this study were to determine the gene action modulating the inheritance of grain yield and other traits, group the inbred lines, investigate inter-trait relationships among grain yield and other traits and assess the performance and stability of single-cross hybrids derived from a set of inbred lines under stress and non-stress environments. One hundred and ninety diallel crosses plus six hybrid checks were evaluated under managed drought at Ikenne during the 2016/17 and 2017/18 dry seasons, low soil N conditions at Mokwa and Ile-Ife, Striga infestation at Abuja and Mokwa, and optimal management conditions at Ikenne, Kadawa, Abuja, Bagauda and Mokwa during the 2016 and 2017 growing seasons. Both additive and non-additive gene actions were prominent in the inheritance of grain yield and other measured traits under stress and optimal management conditions. However, additive gene action was preponderant over the non-additive. The PVA inbreds were classified into three heterotic groups with TZEI 25 and TZEIOR 164 identified as inbred testers for heterotic groups 2 and 3, respectively. Plant and ear heights, ears per plant, plant and ear aspects were identified as reliable secondary traits for genetic enhancement of grain yield under both stress and non-stress conditions. Hybrids TZEIOR 4 × TZEIOR 158 and TZEIOR 119 × TZEIOR 158 were outstanding in performance and should be tested extensively for possible commercialization to combat malnutrition and food insecurity in SSA.

Development, testing and selection of superior inbred lines is crucial for the success of a hybrid program targeting Striga-infested and low soil nitrogen (low-N) environments. The practical value of inbred lines is determined by multiple traits, most of which are inter-dependent. The main objective of this study was to identify early maturing inbred lines based on multiple traits under optimal, low-N and Striga-infested environments for hybrid development and population improvement. One hundred early maturing inbred lines were evaluated under artificial Striga-infestation, low-N and optimal growing environments for two years at Kwadaso and Nyankpala in Ghana. The inbred lines exhibited high levels of genetic variability for grain yield and other agronomic traits desirable for Striga resistance and low-N tolerance. Under optimal growing conditions, days to silking (DS), ears per plot (EHARV) and days to anthesis (DA) had high direct effects on grain yield (GYLD). Days to silking and ears per plant (EPP) had the highest positive direct effects on GYLD, while DA had the highest negative direct effect on grain yield in low-N environments. Under Striga-infestation, the highest negative direct effect on GYLD was obtained with EASP. All the measured traits previously identified to have direct influence on grain yield were associated with it and could be used for indirect selection for improved grain yield under the contrasting environments. Forty-eight of the 100 inbred lines studied were identified as low-N tolerant and forty-nine as Striga resistant.

A hemiparasitic plant, Striga hermonthica (Del.) Benth and soil nitrogen stress are the key constraints to maize (Zea mays L.) productivity in sub-Saharan Africa, where commonly cultivated maize is the normal endosperm type that is deficient in provitamin A, tryptophan and lysine (PVATL). Seventy-six extra-early maize inbreds with provitamin A, tryptophan, and lysine qualities (TZEEIORQ) were developed to address these constraints, and four checks were assessed under Striga, low and high nitrogen conditions at three locations in Nigeria. The inbreds were further genotyped with two beta-carotene hydroxylase 1 (crtRB1) markers, and their seeds were quantified for provitamin A content. Significant (p < 0.01) genetic variations were observed for grain yield and other agronomic attributes of the inbreds under varying environmental conditions. Levels of PVATL for the inbreds ranged from 2.21–10.95 µg g−1, 0.04–0.08%, and 0.19–0.39%, respectively. Beta-carotene marker, crtRB1-3′TE, was polymorphic and grouped the inbreds into two. The marker was effective in identifying inbreds with moderate provitamin A content. Inbreds TZEEIORQ 5, TZEEIORQ 52, and TZEEIORQ 55 exhibited resistance to Striga, tolerance to nitrogen stress with moderate levels of PVATL and could be invaluable sources of favorable alleles for breeding nutritionally improved maize varieties with resistance/tolerance to Striga and soil nitrogen stress.

Genetic gain studies in a breeding program are very important for measuring the efficiency of the program over a specific period. It also provides breeders with an understanding of the outcomes of the huge investments committed to the breeding programs. This study sought to (i) estimate the gains in grain yield under drought and rainfed conditions and (ii) identify high-yielding and stable hybrids developed in the last two decades under drought and rainfed conditions. Sixty extra-early maturing hybrids developed by the International Institute of Tropical Agriculture (IITA), Ibadan, during three breeding periods: (2008–2010, 2011–2013, and 2014–2016) were assessed under managed drought and rainfed conditions across 14 environments in Nigeria to estimate the genetic gains in grain yield using linear regression analysis. Considerably high genetic gains (4.1%) per year for grain yield was obtained for the extra-early maturing hybrids under drought environments. Six of the top ten hybrids identified were from period 3, an indication of the high genetic gains from selection in period 3. Additionally, two of the top six period 3 hybrids (TZEEI 29 × TZEEI 13) × TZEEI 64 and TZdEEI 64 × TZEEI 54 were found to be the best yielding and most stable across the 14 test environments and were highly recommended for extensive evaluations to warrant their commercialization in sub-Saharan Africa (SSA).

Recurrent drought and Striga hermonthica (Del.) Benth parasitism constrains maize production in sub-Saharan Africa (SSA). Transfer of resistance genes from wild relatives can improve resistance to drought and Striga in tropical maize. The objectives of this study were to (i) determine the combining ability of 12 extra-early yellow maize inbreds derived from Zea diploperennis and tropical maize germplasm; (ii) classify the inbreds into heterotic groups using heterotic grouping based on the general combining ability (GCA) of multiple traits (HGCAMT) method; (iii) examine hybrid performance under contrasting environments; and (iv) examine the stability of hybrid combinations involving the inbreds. Sixty-six diallel crosses involving the inbreds plus four checks were evaluated for two years under drought, Striga-infested and rainfed environments in Nigeria. Significant differences (p < 0.05) were observed for the effects of genotype, environment, genotype × environment, GCA and specific combining ability (SCA) on grain yield and other measured traits. Inbred lines such as TZdEEI 7 × TZEEI 63 derived from Z. diploperennis and tropical germplasm exceeded the checks by a range of 28 to 41%. Across environments, the hybrid TZdEEI 1 × TZdEEI 7, which was derived from Z. diploperennis, was the highest-yielding with a grain yield of 4302 kg ha−1. The results revealed the predominance of GCA over SCA effects for most measured traits, suggesting that additive gene action governed the inheritance of Striga resistance and drought tolerance related traits in the inbreds. The 12 inbreds were classified into three heterotic groups, while TZEEI 79 and TZdEEI 7 were identified as inbred testers and TZdEEI 7 × TZEEI 12 as a single-cross tester across environments. Hybrid TZdEEI 9 × TZEEI 79 was the highest-yielding and most stable. Other promising hybrids were TZdEEI 7 × TZEEI 79, TZdEEI 1 × TZdEEI 7 and TZdEEI 12 × TZEEI 95. These hybrids should be extensively tested on-farm for potential commercialization in SSA. Overall, our results highlighted the importance of harnessing beneficial alleles from wild relatives of maize for improvement of resistance to Striga and tolerance to drought in adapted maize germplasm.

Grain-yield potential of maize (Zea mays L.) is limited by the harsh climatic conditions during dry seasons in the savanna of sub-Saharan Africa (SSA). One hundred and five hybrids derived from diallel crosses involving 15 early-maturing yellow maize inbreds were screened under combined drought and heat stress (CDHS) and well-watered (WW) conditions at four locations representative of the dry savannas of Niger, 2012–2014. The objective of the study was to examine the general combining ability (GCA) effects of the inbred lines and specific combining ability (SCA) of the crosses under CDHS and WW environments, assess yield and stability of the derived hybrids under contrasting environments and investigate inter-relationships among traits of the hybrids under contrasting environments. GCA and SCA effects for grain yield and other measured traits were significant, with preponderance of GCA effects over SCA effects under both the CDHS and WW environments. Inbred ENT 13 possessed positive and significant GCA effects for grain yield under both CDHS and WW environments indicating that the inbred could be used for developing outstanding hybrids with good levels of tolerance to CDHS. This also implied that the inbred line could serve as invaluable genetic resource for introgression of favorable alleles into tropical early-maturing maize populations for accelerated genetic gains from selection for improved grain yield. Hybrids ENT 13 × TZEI 167 and TZEI 157 × ENT 13 were the most stable and among the highest yielding hybrids across research environments. These outstanding hybrids could be tested extensively for commercialization in SSA to improve food security. Plant and ear aspects and days to 50% silking could be combined with grain yield in a selection index for improvement of yield under CDHS without yield penalties under WW environments.

Adequate knowledge and understanding of the genetic diversity and inter-trait relationships among elite maize inbred lines are crucial for determining breeding strategies and predicting hybrid performance. The objectives of this study were to investigate the genetic diversity of 162 early maturing white and yellow tropical maize inbred lines, and to determine the population structure, heterotic groups and inter-trait relationships among the lines. Using 9684 DArT single nucleotide polymorphism (SNP) markers, a gene diversity (GD) of 0.30 was recorded for the inbred lines with polymorphic information content (PIC) ranging from 0.08 to 0.38. The genetic relatedness among the inbred lines evaluated revealed six different groups based on the history of selection, colour of endosperm and pedigree. The genotype-by-trait (GT) biplot analysis identified inbred 1 (TZEI 935) as outstanding in terms of combined heat and drought (HD) tolerance with the base index analysis identifying 15 superior inbreds in the HD environment. A wide range of genetic variability was observed among the inbred lines, indicating that they are an invaluable resource for breeding for HD tolerance in maize breeding programmes, especially in West and Central Africa.

Accurate estimation of genetic variability present in tropical maize inbreds with varying reactions to Strigahermonthica infestation is essential for efficient and sustainable utilization to ensure increased genetic gain in a breeding program. Thirty-six early maturing maize inbred lines and 156 single cross hybrids were evaluated under Striga-infested and non-infested conditions in Nigeria during the 2014 and 2015 cropping seasons. Under Striga infestation, grain yield ranged from 1134 kg ha−1 for TZEI 26 × TZEI 5 to 5362 kg ha−1 for TZdEI 173 × TZdEI 280. The average yield reduction of the hybrids under Striga infestation was 44% relative to the performance under non-infested environments. Using 4440 high-quality DArT markers, clustering and population structure analyses separated the inbred lines into three distinct groups based on the genetic distance indicating high level of genetic variability among the lines. The base index of the International Institute of Tropical Agriculture (IITA) identified 50% of the inbred lines as Striga resistant. The genetic diversity study provided an opportunity for selecting divergent parents for tagging candidate genes and quantitative trait loci for marker-assisted introgression of Striga resistance genes into early maturing tropical maize breeding populations. The most reliable secondary trait for indirect selection for grain yield under Striga infestation was the ear aspect.

Striga hermonthica parasitism is a major constraint to maize production in sub-Saharan Africa with yield losses reaching 100% under severe infestation. The application of marker-assisted selection is highly promising for accelerating breeding for Striga resistance/tolerance in maize but requires the identification of quantitative trait loci (QTLs) linked to Striga resistance/tolerance traits. In the present study, 194 F2:3 families of TZEEI 79 × TZdEEI 11 were screened at two Striga-endemic locations in Nigeria, to identify QTLs associated with S. hermonthica resistance/tolerance and underlying putative candidate genes. A genetic map was constructed using 1139 filtered DArTseq markers distributed across the 10 maize chromosomes, covering 2016 cM, with mean genetic distance of 1.70 cM. Twelve minor and major QTLs were identified for four Striga resistance/tolerance adaptive traits, explaining 19.4%, 34.9%, 14.2% and 3.2% of observed phenotypic variation for grain yield, ears per plant, Striga damage and emerged Striga plants, respectively. The QTLs were found to be linked to candidate genes which may be associated with plant defense mechanisms in S. hermonthica infested environments. The results of this study provide insights into the genetic architecture of S. hermonthica resistance/tolerance indicator traits which could be employed for marker-assisted selection to accelerate efficient transfer host plant resistance genes to susceptible genotypes.

Striga hermonthica is a major maize production constraint in West and Central Africa (WCA). Fifty-four early maturing maize hybrids of three breeding periods: 2008–2011, 2012–2013, 2014–2015, were evaluated under Striga-infested and non-infested environments in WCA. The study aimed at assessing genetic improvement in grain yield of the hybrids, identifying traits associated with yield gain during the breeding periods, and grain yield and stability of the hybrids in Striga infested and non-infested environments. Annual increase in grain yield of 101 kg ha−1 (4.82 %) and 61 kg ha−1 (1.24%) were recorded in Striga-infested and non-infested environments, respectively. The gains in grain yield from period 1 to period 3 under Striga-infested environments were associated with reduced anthesis-silking interval, reduced Striga damage, number of emerged Striga plants, improved ear aspect, and increased ears per plant. Ear aspect, ears per plant, and Striga damage at 8 and 10 weeks after planting (WAP) were significantly correlated with yield in Striga-infested environments, whereas ears per plant and plant and ear aspects had significant correlations with yield in non-infested environments. Hybrids TZdEI 352 × TZEI 355, TZdEI 378 × TZdEI 173, and TZdEI 173 × TZdEI 352 were outstanding in grain yield and stability in Striga-infested environments, whereas TZEI 326 × TZdEI 352, TZEI 495 × ENT 13, and TZdEI 268 × TZdEI 131 were superior in non-stress environments. These hybrids should be further tested extensively and commercialized. Significant genetic gains have been made in breeding for resistance to Striga hermonthica in early maturing maize hybrids.

The success of a hybrid breeding program is dependent on available heterotic patterns for exploitation of grain-yield heterosis. The efficiency of the assignment of germplasm lines into heterotic groups is a prerequisite for obtaining useful heterotic patterns among germplasm lines. A total of 256 maize hybrids, comprising 244 top crosses, six diallel cross hybrids, and six checks, were grown under Striga infestation, drought, and optimal conditions, from 2015 to 2017. The study determined the combining abilities of the parental inbreds, classified the inbreds into heterotic groups, and compared the efficiencies of the following four grouping methods for classifying the inbreds: specific combining ability (SCA) effect of grain yield; general combining ability (GCA) effects of multiple traits (HGCAMT); SCA and GCA (HSGCA) for yield; and single nucleotide polymorphism-based genetic distance (SNP-based genetic distance (GD)). Significant GCA and/or SCA mean squares were revealed for most measured traits in all test environments. Sums of squares (SS) due to GCA were higher than SCA SS for measured traits in all test environments. The HSGCA, SCA, and SNP-based GD methods identified four heterotic groups, whereas the HGCAMT identified three groups, in all environments. The additive gene effect was preponderant in the inheritance of most measured traits. The efficiencies of the grouping methods varied with the test environments. The HSGCA and SCA methods were the most efficient for grouping in all test conditions. For practical breeding purposes, the HGCAMT and HSGCA methods were recommended under Striga infestation and drought, respectively. The heterotic patterns, which were revealed in this study, were effective for planning hybridization schemes for developing high-yielding, Striga-tolerant/resistant, and drought-tolerant maize hybrids for stressful environments.

Knowledge of the genetic mechanisms conditioning drought tolerance in maize is crucial to the success of hybrid breeding programs aimed at developing high-yielding cultivars under drought. The objectives of this study were to determine the combining ability of extra-early inbreds, compute the heritability of measured traits, assess the performance of inbreds in hybrid combinations and investigate the associations among traits under drought and optimal conditions. A total of 252 hybrids generated by crossing 63 inbreds to four testers, along with four commercial hybrid checks, were evaluated for 2 years under drought and rainfed conditions. General combining ability (GCA) and specific combining ability (SCA) for the traits were significant. A total of 57.1% and 53.4% of the genotypic sum of squares were attributable to GCA effects for grain yield under managed drought and rainfed conditions, respectively. Hybrids TZdEEI 91 × TZEEI 21 and TZdEEI 55 × TZEEI 13 out-yielded the best checks under drought and optimal conditions by 49.13% and 39.05%, respectively. The most promising hybrids with consistently high grain yield under drought and rainfed conditions, were TZdEEI 54 × TZEEI 13, TZdEEI 91 × TZEEI 21 and TZdEEI 55 × TZEEI 21 and should be further evaluated for possible commercial production in sub-Saharan Africa.

Information on genetic diversity and population structure are very important in any breeding programme for the improvement of traits of interest and the development of outstanding products for commercialization. In the present study, we assessed the genetic diversity of 94 early-maturing white and yellow tropical maize inbred lines using single nucleotide polymorphism (SNP) markers. The larger number of SNP markers used in this study allowed a clearer inference of the population structure of the 94 inbred lines. Cluster analysis resolved the inbred lines into different clusters based on their pedigree, selection history and endosperm colour. However, three heterotic groups were revealed by population structure analysis, but additional field evaluation could be more informative to confirm the heterotic groups identified. Nevertheless, wide genetic variability existed among the inbred lines making them unique with the potential to contribute new beneficial alleles to maize breeding programmes in the tropics, especially in the West and Central Africa (WCA) sub-region.

Maize landrace accessions constitute an invaluable gene pool of unexplored alleles that can be harnessed to mitigate the challenges of the narrowing genetic base, declined genetic gains, and reduced resilience to abiotic stress in modern varieties developed from repeated recycling of few superior breeding lines. The objective of this study was to identify extra-early maize landraces that express tolerance to drought and/or heat stress and maintain high grain yield (GY) with other desirable agronomic/morpho-physiological traits. Field experiments were carried out over two years on 66 extra-early maturing maize landraces and six drought and/or heat-tolerant populations under drought stress (DS), heat stress (HS), combined both stresses (DSHS), and non-stress (NS) conditions as a control. Wide variations were observed across the accessions for measured traits under each stress, demonstrating the existence of substantial natural variation for tolerance to the abiotic stresses in the maize accessions. Performance under DS was predictive of yield potential under DSHS, but tolerance to HS was independent of tolerance to DS and DSHS. The accessions displayed greater tolerance to HS (23% yield loss) relative to DS (49% yield loss) and DSHS (yield loss = 58%). Accessions TZm-1162, TZm-1167, TZm-1472, and TZm-1508 showed particularly good adaptation to the three stresses. These landrace accessions should be further explored to identify the genes underlying their high tolerance and they could be exploited in maize breeding as a resource for broadening the genetic base and increasing the abiotic stress resilience of elite maize varieties.

Climate change is expected to aggravate the effects of drought, heat and combined drought and heat stresses. An important step in developing ‘climate smart’ maize varieties is to identify germplasm with good levels of tolerance to the abiotic stresses. The primary objective of this study was to identify landraces with combined high yield potential and desirable secondary traits under drought, heat and combined drought and heat stresses. Thirty-three landraces from Burkina Faso (6), Ghana (6) and Togo (21), and three drought-tolerant populations/varieties from the Maize Improvement Program at the International Institute of Tropical Agriculture were evaluated under three conditions, namely managed drought stress, heat stress and combined drought and heat stress, with optimal growing conditions as control, for two years. The phenotypic and genetic correlations between grain yield of the different treatments were very weak, suggesting the presence of independent genetic control of yield to these stresses. However, grain yield under heat and combined drought and heat stresses were highly and positively correlated, indicating that heat-tolerant genotypes would most likely tolerate combined drought and stress. Yield reduction averaged 46% under managed drought stress, 55% under heat stress, and 66% under combined drought and heat stress, which reflected hypo-additive effect of drought and heat stress on grain yield of the maize accessions. Accession GH-3505 was highly tolerant to drought, while GH-4859 and TZm-1353 were tolerant to the three stresses. These landrace accessions can be invaluable sources of genes/alleles for breeding for adaptation of maize to climate change.