Mapping quantitative trait loci and predicting candidate genes for Striga resistance in maize using resistance donor line derived from Zea diploperennis.

The parasitic weed, Striga is a major biological constraint to cereal production in sub-Saharan Africa (SSA) and threatens food and nutrition security. Two hundred and twenty-three (223) F2:3 mapping population involving individuals derived from TZdEI 352 x TZEI 916 were phenotyped for four Striga-adaptive traits and genotyped using the Diversity Arrays Technology (DArT) to determine the genomic regions responsible for Striga resistance in maize. After removing distorted SNP markers, a genetic linkage map was constructed using 1,918 DArTseq markers which covered 2092.1 cM. Using the inclusive composite interval mapping method in IciMapping, twenty-three QTLs influencing Striga resistance traits were identified across four Striga-infested environments with five stable QTLs (qGY4, qSC2.1, qSC2.2, qSC5, and qSC6) detected in more than one environment. The variations explained by the QTLs ranged from 4.1% (qSD2.3) to 14.4% (qSC7.1). Six QTLs each with significant additive × environment interactions were also identified for grain yield and Striga damage. Gene annotation revealed candidate genes underlying the QTLs, including the gene models GRMZM2G077002 and GRMZM2G404973 which encode the GATA transcription factors, GRMZM2G178998 and GRMZM2G134073 encoding the NAC transcription factors, GRMZM2G053868 and GRMZM2G157068 which encode the nitrate transporter protein and GRMZM2G371033 encoding the SBP-transcription factor. These candidate genes play crucial roles in plant growth and developmental processes and defense functions. This study provides further insights into the genetic mechanisms of resistance to Striga parasitism in maize. The QTL detected in more than one environment would be useful for further fine-mapping and marker-assisted selection for the development of Striga resistant and high-yielding maize cultivars.

Genetics of extra-early-maturing yellow and orange quality protein maize inbreds and derived hybrids under low soil nitrogen and Striga infestation.

The development and commercialization of extra-early quality protein maize (QPM)-provitamin A (PVA) hybrids that are tolerant of low soil N (LN) and Striga resistant are essential for addressing the food insecurity and undernourishment challenges currently faced by sub-Saharan Africa (SSA). This study was designed (a) to determine the genetic effects regulating grain yield (GY) and important secondary traits of extra-early yellow and orange QPM-PVA inbred lines under LN, Striga-infested, and high-N (HN) conditions, (b) to investigate whether maternal genes influenced the inheritance of GY and other secondary traits, (c) to assess the GY and stability of the hybrids across the three management conditions, and (d) to examine the relationship between single nucleotide polymorphism (SNP) marker-based genetic distances and GY. Twenty-four inbred lines were used to produce ninety-six single cross hybrids using the North Carolina Design II. The performance of the hybrids plus four checks was assessed across LN, Striga-infested, and HN management conditions in Ghana and Nigeria in 2018. Additive genetic variances were preponderant over nonadditive genetic variances for GY and most secondary traits in each and across environments. TZEEQI 358 exhibited significant and positive male and female GCA effects for GY under LN, Striga infestation, HN, and across management conditions indicating that favorable alleles for GY could be donated by TZEEQI 358. Maternal effects regulated the inheritance of plant height under the Striga-infested conditions. Genetic distances were associated with GY under LN, Striga infestation, and HN conditions. TZEEIORQ 58 × TZEEQI 397 demonstrated high GY and stability of performance; therefore, it should be further tested under multiple environments for commercialization.

Combining ability of extra-early biofortified maize inbreds under Striga infestation and low soil nitrogen.

Striga hermonthica (Del.) Benth parasitism, low soil N, and nutritional deficiencies of normal-endosperm maize (Zea mays L.) threaten maize yield and exacerbate nutritional problems in sub-Sahara Africa (SSA). This study was conducted (a) to evaluate genetic variation among extra-early maturing maize hybrids with provitamin A and quality protein characteristics, (b) to investigate gene action governing the inheritance of Striga resistance, grain yield, low N tolerance, and other measured traits under low-N, high-N, and Striga-infested environments, and (c) to identify hybrids with high yield and stability across environments. One hundred and fifty hybrids developed using North Carolina Design II were evaluated with six checks under low-N, high-N, and Striga-infested environments in Nigeria. Mean squares for hybrids were highly significant (P < .01) for grain yield and other traits across environments. Only general combining ability (GCA) for female and/or male mean squares were significant for measured traits under low N. In addition to significant GCA effects for most traits, specific combining ability was significant (P < .05) for Striga emergence count under Striga infestation, and ear height and ears per plant under high N, indicating that additive and nonadditive genetic effects controlled the inheritance of few traits under Striga and high N, whereas additive genetic effect governed the inheritance of the traits under low N. Hybrids TZEEIORQ 55 × TZEEIORQ 26, TZEEIORQ 49 × TZEEIORQ 75, and TZEEIORQ 52 × TZEEIORQ 43 were high yielding and stable across environments and have potential for improving nutrition and maize yields in SSA.

Genetic studies of extra-early provitamin-A maize inbred lines and their hybrids in multiple environments.

Vitamin A deficiency, drought, low soil nitrogen (low-N), and Striga hermonthica parasitism of maize (Zea mays L.) cause malnutrition and food insecurity in sub-Saharan Africa. The objectives of this study were to determine combining abilities of extra-early provitamin A (PVA) lines, classify them into heterotic groups (HGs), identify testers, and determine yield stability of hybrids under contrasting environments in two trials. In Trial 1, 190 F1 hybrids plus six checks were tested under Striga-infested, drought, and stress-free environments in Nigeria from 2015-2017. In Trial 2, 35 extra-early yellow hybrids were evaluated under low-N, Striga-infested, and stress-free environments in 2018. TZEEIOR 202 and TZEEIOR 205 had PVA concentrations of 23.98 and 22.56 µg g-1. TZEEIOR 197 × TZEEIOR 205 (20.1 µg g-1) and TZEEIOR 202 × TZEEIOR 205 (22.7 µg g-1) contained about double the PVA level of the commercial check, TZEEI 58 × TZEE-Y Pop STR C5 (11.4 µg g-1). Both general (GCA) and specific (SCA) combining ability variances were significant for most agronomic traits, although GCA was larger than SCA effects, indicating GCA effects primarily controlled the inheritance of those traits. TZEEIOR 97 and TZEEIOR 197 were identified as inbred testers. TZEEIOR 197 × TZEEIOR 205 was identified as a single-cross tester and the most stable and highest-yielding hybrid across environments. TZEEIOR 202 and TZEEIOR 205 should be invaluable resources for breeding for high PVA. Provitamin A level was independent of hybrid yield potential, indicating that selection of superior hybrids with elevated PVA levels should be feasible.

Yield gains and associated changes in an early yellow bi-parental maize population following genomic selection for Striga resistance and drought tolerance.

BACKGROUND: Maize yield potential is rarely maximized in sub-Saharan Africa (SSA) due to the devastating effects of drought stress and Striga hermonthica parasitism. This study was conducted to determine the gains in grain yield and associated changes in an early-maturing yellow bi-parental maize population (TZEI 17 x TZEI 11) F3 following genomic selection (GS) for improved grain yield, Striga resistance and drought tolerance. Fifty S1 lines were extracted from each of cycles C0, C1, C2 and C3 of the population and crossed to a tester TZEI 23 to generate 200 testcrosses. The testcrosses were evaluated under drought, artificial Striga-infested and optimal (free from Striga infestation and without limitation of water and nitrogen) environments in Nigeria, 2014-2017. RESULTS: Gains in grain yield of 498 kg ha- 1 cycle- 1 (16.9% cycle- 1) and 522 kg ha- 1 cycle- 1 (12.6% cycle- 1) were obtained under Striga-infested and optimal environments, respectively. The yield gain under Striga-infested environments was associated with increased plant and ear heights as well as improvement in root lodging resistance, husk cover, ear aspect and Striga tolerance. Under optimal environments, yield gain was accompanied by increase in plant and ear heights along with improvement of husk cover and ear rot resistance. In contrast, genomic selection did not improve grain yield under drought but resulted in delayed flowering, poor pollen-silk synchrony during flowering and increased ear height. Genetic variances and heritabilities for most measured traits were not significant for the selection cycles under the research environments. Ear aspect was a major contributor to grain yield under all research environments and could serve as an indirect selection criterion for simultaneous improvement of grain yield under drought, Striga and optimal environments. CONCLUSION: This study demonstrated that genomic selection was effective for yield improvement in the bi-parental maize population under Striga-infested environments and resulted in concomitant yield gains under optimal environments. However, due to low genetic variability of most traits in the population, progress from further genomic selection could only be guaranteed if new sources of genes for Striga resistance and drought tolerance are introgressed into the population.

Genetic Variances and Heritabilities of Traits of an Early Yellow Maize Population after Cycles of Improvement for Striga Resistance and Drought Tolerance.

Drought and Striga are principal constraints to maize (Zea mays L.) production in sub-Saharan Africa. An early yellow maize population, TZE-Y Pop DT STR, which had undergone five cycles of selection for resistance to Striga, followed by three cycles of improvement for drought tolerance, was investigated for yield gains, changes in genetic variance, and interrelationships among traits under drought stress and optimum environments. Two hundred and forty S1 lines comprising 60 each from the base population and subsequent populations from three selection cycles improved for grain yield and drought tolerance were assessed under drought and optimal environments in Nigeria from 2010 to 2012. Genetic improvements in grain yield of 423 and 518 kg ha-1 cycle-1 were achieved under drought stress and optimal environments. Predicted improvements in selection for yield were 348 and 377 kg ha-1 cycle-1 under drought stress and optimum environments, respectively. The highest yield observed in C3 was accompanied by reduced days to silking and anthesis-silking interval, improved plant aspect and ear aspect, and increased plant height and ears per plant across research environments, as well as improved stay-green characteristic under drought. The level of genetic variability for yield and a few other traits were maintained under drought and optimal environments in the population. The presence of residual genetic variability for yield and other assayed traits in C3 indicated that progress could be made from future selection in the population depending on the ability of breeders to identify outstanding genotypes and the precision level of experimentation. Substantial improvement has been made in yield and drought tolerance in C3 of the population.

Gains in Grain Yield of Extra-Early Maize during Three Breeding Periods under Drought and Rainfed Conditions.

Drought is a key maize (Zea mays L.) production constraint in sub-Saharan Africa. Fourteen, fifteen, and twenty-five extra-early maturing maize cultivars, with varying Striga resistance and drought and low soil N tolerance, were developed from 1995 to 2000 (Period 1), 2001 to 2006 (Period 2), and 2007 to 2012 (Period 3), respectively. The objectives of this study were to examine yield gains in the cultivars and to investigate inter-trait relationships and yield stability under six drought and 17 rainfed conditions in West Africa from 2013 to 2016. Annual rate of yield increase across cultivars was 0.034 (3.28%) and 0.068 Mg ha-1 (2.25%), whereas yield gains per period were 0.17 and 0.38 Mg ha-1 under drought and rainfed environments, respectively. Yield gains under drought and rainfed environments were related to prolonged flowering period, increased plant and ear heights, improved stalk lodging, and ear and plant aspects, whereas delayed leaf senescence and increased number of ears per plant accompanied yield improvement under drought only. Ear aspect and number of ears per plant were primary contributors to yield and could be used as selection criteria for yield enhancement under drought and rainfed conditions. High-yielding and stable cultivars across all environments based on additive main effects and multiplicative interaction (AMMI) biplot included '2004 TZEE-Y Pop STR C4' and 'TZEE-W Pop STR BC2 C0' of Period 2 and '2009 TZEE-W STR', 'TZEE-Y STR 106', 'TZEE-W STR 107', and 'TZEE-W DT C0 STR C5' of Period 3. These cultivars could be commercialized to improve food self-sufficiency in sub-Saharan Africa.

Combining ability and performance of extra-early maturing yellow maize inbreds in hybrid combinations under drought and rain-fed conditions.

Maize (Zea mays L.) is a major staple food and cash crop in sub-Saharan Africa (SSA). However, its production and productivity are severely constrained by drought. A total of 120 single-cross hybrids and an open-pollinated control variety were evaluated for 2 years at two locations under managed drought and rain-fed conditions in Nigeria. The objective of the present study was to assess their performance, classify them into distinct heterotic groups and identify promising hybrids for commercialization in the West and Central Africa sub-region. General combining ability and specific combining ability mean squares were highly significant for grain yield and other traits under the research environments. However, there was a preponderance of additive gene action over non-additive. Only six out of 39 inbreds were classified into distinct heterotic groups by the testers. The highest-yielding drought-tolerant hybrid, TZEEI 102 × TZEEI 95, out-yielded the open-pollinated control variety by 43·70%. The average yield reduction under drought was 54·90% of the yield under rain-fed conditions. The hybrids TZEEI 81 × TZEE1 79, TZEEI 100 × TZEEI 63 and TZEEI 64 × TZEEI 79 were the highest-yielding and most stable across environments. These outstanding drought-tolerant hybrids, which are also resistant to Striga, have the potential to contribute to food security and increased incomes in SSA and should be tested extensively on-farm and commercialized.

Relative changes in genetic variability and correlations in an early-maturing maize population during recurrent selection.

Four cycles of S(1) family recurrent selection to improve grain yield and resistance to Striga hermonthica have been completed in TZE-Y Pop STR C(0.) In order to determine whether or not to continue with the recurrent scheme, it was desirable to evaluate the amount of residual genetic variance and associated parameters in the population. The objective of this study was to characterize the relative changes in the levels of the genetic variances, heritability estimates and genetic correlation coefficients, and to predict future gains from selection for grain yield, Striga resistance and other agronomic traits. Fifty S(1) families, derived from each cycle, were evaluated under Striga-infested and Striga-free conditions at Mokwa, Ikenne and Abuja, Nigeria, in 2005 and 2007. Under Striga infestation, genetic variances for grain yield, days to anthesis, plant height and Striga damage generally increased in the advanced cycles of selection. In contrast, the genetic variances for days to silk, anthesis-silking interval, ears per plant, ear aspect and number of emerged Striga plants decreased with selection. The advanced cycles of selection significantly out-yielded the original cycle in both research environments. Heritabilities for grain yield, Striga damage and number of emerged Striga plants were significantly greater than zero. The realized gains from selection for grain yield under Striga infestation (52 kg ha(-1) cycle(-1)) and Striga-free conditions (130 kg ha(-1) cycle(-1)) were remarkably lower than the predicted gains (350 and 250 kg ha(-1 )cycle(-1), respectively). Adequate genetic variability exists in cycle 4 of the scheme to ensure future gains from selection.

Assessment of testcross performance and genetic diversity of yellow endosperm maize lines derived from adapted x exotic backcrosses.

Introduction of exotic maize (Zea mays L.) into adapted tropical germplasm may enhance genetic variability and lead to greater progress from selection. The first objective of this study was to determine if yellow endosperm lines derived from adapted x exotic backcrosses contain exotic alleles that are superior to the recurrent adapted parental line for yield and other agronomic traits in tropical environments. Thirteen exotic yellow maize inbred lines were crossed to an adapted orange line (KUSR) and the F1s were backcrossed to KUSR to generate the first backcrosses. Fifty BC1F4 lines derived from these backcrosses and the recurrent parent were crossed to a common inbred tester (L4001) to form testcrosses, which were evaluated at eight environments in Nigeria. Testcrosses of the BC-derived lines differed significantly for grain yield and other agronomic traits. Only two testcrosses yielded significantly less than L4001 x KUSR, with the best 15 testcrosses producing between 289 and 1,056 kg/ha more grain yield than L4001 x KUSR. The best testcrosses were similar to or better than L4001 x KUSR for other agronomic traits. The second objective of this study was to assess the extent of genetic diversity present among the BC-derived lines. We genotyped 46 BC-derived lines including KUSR and L4001 with 10 AFLP primer pairs and found 491 polymorphic fragments. The average allelic diversity of the lines was 0.30 +/- 0.01. The genetic distance of each BC-derived line from KUSR ranged between 0.49 and 0.91. The average genetic distance for all pairs of the BC-derived lines was 0.68 +/- 0.004, varying from 0.34 to 0.92. The increased grain yield and genetic diversity observed in these studies provide evidence that exotic germplasm can contribute new alleles to expand the genetic base of tropical maize and develop high-yielding hybrids.

Molecular marker-based genetic diversity assessment of Striga-resistant maize inbred lines.

Striga-resistant maize inbred lines are of interest to maize breeding programs in the savannas of Africa where the parasitic weed is endemic and causes severe yield losses in tropical maize. Assessment of the genetic diversity of such inbred lines is useful for their systematic and efficient use in a breeding program. Diversity analysis of 41 Striga-resistant maize inbred lines was conducted using amplified fragment length polymorphism (AFLP) and simple sequence repeat (SSR) markers to examine the genetic relationships among these lines and to determine the level of genetic diversity that exists within and between their source populations. The two marker systems generated 262 and 101 polymorphic fragments, respectively. Genetic similarity (GS) values among all possible pairs of inbred lines varied from 0.45 to 0.95, with a mean of 0.61+/-0.002 for AFLPs, and from 0.21 to 0.92, with a mean of 0.48+/-0.003, for SSRs. The inbred lines from each source population exhibited a broad range of GS values with the two types of markers. Both AFLPs and SSRs revealed similar levels of within population genetic variation for all source populations. Cluster and principal component analysis of GS estimates with the two markers revealed clear differentiation of the Striga-resistant inbred lines into groups according to their source populations. There was clear separation between early- and late-maturing Striga-resistant inbred lines. Considering the paucity of germplasm with good levels of resistance to Striga in maize, the broad genetic diversity detected within and among source populations demonstrates the genetic potential that exists to improve maize for resistance to Striga.