Combining Ability and Heterosis for Grain Yield and Other Agronomic Traits in Provitamin A Enriched Maize Inbred Lines

Combining Ability and Heterosis for Grain Yield and Other Agronomic Traits in Provitamin A Enriched Maize Inbred Lines | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Combining Ability and Heterosis for Grain Yield and Other Agronomic Traits in Provitamin A Enriched Maize Inbred Lines Belay Garoma, Kassahun Bante, Girum Azimach, Abebe Menkir This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4839825/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Combining ability of enriched Pro-vitamin A yellow maize inbred lines can provide information for breeders to develop biofortified maize hybrids with high yield and desirable traits. Sixty-eight F1 hybrids along with two checks were arranged in alpha lattice design, replicated twice and evaluated across three locations at mid-altitude agro-ecology of Ethiopia. The combined analysis of variance revealed a significant difference (P ≤ 0.01) in mean square due to PVA hybrids, environment and hybrid interaction with environments for grain yield and most of the agronomic traits and it indicates that genotypes performed differently across environments. Seven PVA hybrids were selected based on average yield performance ranging from 7.11 to 7.95 t/ha and 18.69–29.04% yield advantage over the best check and these hybrids should be further evaluated across environments. The mean squares analysis of variance for GCA and SCA showed a significant difference (p ≤ 0.001) for all traits studied, indicating that both additive and non-additive gene effects were important for controlling these traits. Significant positive SCA effect on grain yield was recorded for crosses of L7/L12 (3.16 t/ha), L7/L9 (2.43 t/ha) and L10 x L12 (2.05 t/ha). Only one cross (L9 x L12) displayed significant (p < 0.05) positive standard heterosis (34.52%) over BH 549, whereas non-significant positive heterosis was detected over the BHQPY 545 check. Inbred lines L12, L8, L6, L10, and L2 had the highest positive and significant GCA effects on grain yield and other traits. Out of these lines, L8 and L12 were good combiners for most traits and will be used as alternative testers as they had the highest GCA effect for yield as well as other traits, and also their cross combination were among the top performing hybrids for yield. Our study highlighted that those inbred lines that had a high GCA effect within the heterotic group, could be used as a new source of PVA inbred line development. Furthermore, L8 and L12 can be used as alternative testers for future PVA maize breeding programs. Biological sciences/Genetics Biological sciences/Plant sciences Combing ability yield heterosis tester inbred lines maize 1. Introduction Maize ( Zea mays , L.) is one of the most important staple food crops in developing countries and provides over 20% of the calories in the human diet 1 . However, most of the maize varieties cultivated in these countries have low micronutrient as well as vitamin A content 2 which leads to micronutrient deficiency including vitamin A deficiency. Diverse and nutrient-rich foods such as fruits, vegetables and animal-based products are often inaccessible for many rural communities in low and middle-income countries. Thus, biofortification of staple foods through conventional breeding and application of biotechnological tools is a sustainable and affordable approach to increase vitamin and mineral content in staple food crops 3 . Maize kernels accumulate diverse carotenoids with provitamin A activity and biofortification breeding has been initiated; subsequently, efforts have been made to increase PVA concentration in maize inbred lines 4 . The usefulness of inbred lines for maize breeding program is based on their combining ability and high heterosis expression in hybrid performance. Diallel mating scheme was developed by Griffing (1956) and this scheme is used to study combing ability and heterosis phenomena among inbred lines 5 , 6 . Combing ability studies also provide useful information about the nature of gene action for particular traits 7 and to select superior parents for new sources of population and hybrid developments 8 . Heterosis refers to the superior performance of F 1 hybrids relative to their parents in terms of vigor or other desirable agronomic traits 9 . Cross hybrids expressed significantly higher than mid-parent heterosis for grain yield and provitamin A content 10 . Likewise, hybrids exhibited the highest specific combining ability and heterosis for grain yield compared to commercial hybrids 11 . Moreover, 12 demonstrated that the combining ability among yellow maize inbred lines showed significant general combining ability (GCA) and specific combining ability (SCA) effects for grain yield, provitamin A content and other traits. Combing ability analysis also helps to identify testers that are important in maize breeding to employ early screening progeny, grouping inbreeds, to select superior parents and top hybrids in trials performance. The choice of desirable testers can depend on the genetic difference in their progeny for tested traits 13 , high GCA effect on yield 14 , gene frequency of favorable alleles 8 and the discrimination of the differences among tested parents 15 . Many studies have reported on the selection of effective testers in partial diallel crosses. For example, 16 identified testers that had the highest GCA effects for provitamin A and other traits. 14 ; 17 selected testers that showed strong positive significant and high GCA effects for grain yield and ears per plant. Thus, continuous assessment of early and new provitamin A-enriched yellow maize inbred lines is important for their combing ability to select ideal testers and superior parents for hybrid development. In the earlier study, we identified medium to high PVA content of yellow/orange maize inbred lines from different genetic background sources, pedigrees recorded and heterotic groups. However, no information is available on combining abilities of newly developed PVA-enriched inbred lines and heterosis for grain yield, other agronomic traits and suitable testers. Therefore, the objectives of the study were: i) To evaluate the combing ability of PVA-enriched inbred lines for grain yield and important agronomic traits. ii) to identify provitamin A testers, the best-performing hybrids and estimate heterosis of F1s over check and mid-parents for yield and other agronomic traits. 2. Materials and Methods 2.1. Source of plant materials and crossing techniques The twelve PVA enriched yellow/orange maize inbred lines included in this study were developed at the Maize Improvement Program of IITA, CIMMYT and Bako National Mazie Research Centre of the Ethiopian Institute of Agricultural Research (BNMRC/ EIAR) as following. The two PVA enriched inbred lines CML486/(CML297-B KUICarotenoidsyn-FS17-3-2-B/KUI3 B77))-B-11-1-B-B-B-B-B-B-B-#-B-B-B (L10) and (CML537/[BETASYN]BC1-10-2-1-#/CML-305-B)-B-11-1-B-B-#-B-B-B (L12) were introduced from CIMMYT evaluated over years and selected for adaptation to local condition at Bako, Ethiopia. The three PVA enriched inbred lines TZMI1003-#-# (L8), TZMI1003-#-# (L9) and CLHP00306 (L11) introduced from International Institute of Tropical Agriculture (IITA) were also selected for adaption to locational condition at Bako, Ethiopia. Other inbred lines introduced from CIMMYT and used as donor parents for beta carotene include KUI carotenoid syn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-3(MAS:L4H1); KUI carotenoid syn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-1(MAS:L4H1; KUI carotenoid syn-FS3-3-2-B-B-B(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1; proA-HydB1)-#bal]FS and Florida A plus syn-FS2-2-1-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1) were crossed with locally adapted elite white maize inbred lines of BNMC/ EIAR includes; DE78-Z-126-3-2-2-1-1(g), Gibe1-91-1-1-1-1, IL00'E-9-1-1-1-1-1, Obatanpa and CML 161 as recipients. The main donors of high b-carotene for the IITA, CIMMYT and yellow/orange Ethiopian lines included in this study were originally temperate inbred line and source populations including DE3, [BETASYN]BC1-10-2-1-#, KUI carotenoids syn-FS17-3-2-B, KUI Carotenoids syn-FS11-1-1-B-B-B, KUI carotenoid syn-FS3-3-2-B-B-B, and (ProA-HydB1)-#ba1] FS. Yellow inbred lines were then derived from biparental crosses through re-repeated generation of inbreeding with selection for desirable kernel colour and agronomic attributes using the pedigree breeding approach. The advanced yellow/orange maize inbred lines were then evaluated across locations for their adaptation and agronomic performance and reaction to disease under natural condition. The advanced yellow/orange maize inbred lines along with selected IITA and CIMMYT lines were planted in trials evaluated in 2020 and 2021 at Bako, Ethiopia. Seed samples drawn from these trials were shipped to CIMMYT-Mexico for carotenoid analyses 18 . Based on the results of carotenoids analysis, twelve yellow/orange maize inbred lines with diverse genetic backgrounds and PVA content exceeding 6 µg/g were selected from the different heterotic groups based on available of seed in stock (Table 1 ). The twelve PVA enriched yellow/orange maize inbred lines were crossed in a 12 x 12 half-diallel mating design to generate 66 F 1 crosses at National maize research centre during the 2021 dry season at Bako, Ethiopia. Table 1 List of Provitamin A enriched maize inbred lines used to generate half diallel crosses Inbred lines code Pedigree Source of yellow maize inbred lines PVA content (µg/g) Seed source in stock Stock ID Heterotic group L1 (DE78-Z-126-3-2-2-1-1(g)/(KUI carotenoid syn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-3(MAS:L4H1)-1-B-B-B)-B-1-1-1-B-B BNMRC 9.96 2019K-MQ19N7-45-1 QL53-33 A L2 (DE78-Z-126-3-2-2-1-1(g)/(KUI carotenoid syn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-3(MAS:L4H1)-1-B-B-B)-B-13-1-2-B -B BNMRC 6.99 2019K-MQ19N7-47-1 QL53-36 A L3 (Gibe1-91-1-1-1-1/(KUI carotenoid syn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-1(MAS:L4H1)-5-B-B-B)-B-7-1-1-B-B BNMRC 6.77 2019K-MQ19N7-51-1 QL53-38 A L4 (IL00'E-9-1-1-1-1-1/(KUI carotenoid syn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-1(MAS:L4H1)-5-B-B-B)-B-13-1-4-B-B BNMRC 9.97 2019K-MQ19N7-14-1 QL53-7 B L5 (IL00'E-9-1-1-1-1-1/(KUI carotenoid syn-FS3-3-2-B-B-B(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1)-2-B-B-B)-B-2-2-1-B-B BNMRC 7.16 2019K-MQ19N7-17-1 QL53-9 B L6 [(Obatanpa(proA-HydB1)-#bal]FS-93-3-1-1-2-B-B BNMRC 9.78 2019K-BK18MN33-25 QL53-33 A L7 CML161/(Florida A plus syn-FS2-2-1-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1)-2-B-B-B//CML161-6-1-2-B-B BNMRC 6.61 2019K-MQ19N7-64-1 QL53-49 A L8 TZMI1003-#-# IITA 11.36 2019K-MQ19N30-34 QL75-33 A L9 TZMI1065-#-# IITA 9.79 2019K-MQ19N30-34 QL75-34 B L10 (CML486/(CML297-B KUICarotenoidsyn-FS17-3-2-B/KUI3 B77))-B-11-1-B-B-B-B-B-B-B-#-B-B-B CIMMYT 10.2 2019K-MQ19N30-132 A L11 CLHP00306 IITA 10.70 2019K-MQ19N30-31 QL75-31 B L12 (CML537/[BETASYN]BC1-10-2-1-#/CML-305-B)-B-11-1 -B-B-#-B-B-B CIMMYT 7.10 2019K-MQ19N30-134 QL75-30 B Notice: BNMRC/EIAR = Bako National Maize Research Centre under Ethiopian Institute of Agricultural Research, Bako, Ethiopia; IITA = International Institute of Tropical Agriculture, Ibadan, Nigeria. CIMMYT = International Mazie and wheat Improvement Centre, Stock ID and seed source indicates the availability of seed in seed store or in cold room at Bako National maize research centre during inbred lines planted. 2.2. Trial management Sixty-eight F1 hybrids plus two checks were arranged in a 17 x 4 alpha lattice design and planted with two replications. Each hybrid was planted on 4 m long rows with 16 plants per row, with 75 cm and 25 cm spacing between rows and plants, respectively. At planting, two seeds per hill were planted, and after two weeks of emergency, they were thinned out to maintain one plant per hill. Trials were planted and evaluated at Bako, Jima and Pawe agricultural research centres in the 2022 main cropping season. The parental lines of the diallel crosses were also planted separately adjustment to the diallel cross trial in a randomized complete block design with two replications in the 2022 main cropping season. Agronomic management, such as weeding and fertilizer application was applied as per research recommendations at the respective field-testing site. Description of testing locations are presented in Table 2 . Table 2 Description for the test locations used to evaluate Provitamin A maize hybrids Location Country EIAR Co-ordinates Altitude (m.a.s.l) Rainfall (mm) Minimum Temperature ( 0 C) Maximum Temperature ( 0 C) Soil type Bako Ethiopia EIAR/BNMRC 9 0 06’N and 37 0 09' E 1650 1244 12.3 29.8 Nitosol Jima Ethiopia EIAR/JARC 7 0 40’N and 36 0 49’E 1753 1572 11 26.3 Eutric Nitosol Pawe Ethiopia EIAR/PARC 11°19'N and 36°24'E 1120 1238 16.3 32.6 Nitisol Notice: EIAR/BNMRC = Ethiopian Institute of Agricultural Research/Bako National Maize Research Centre; JARC = Jima Agricultural Research Centre; PARC = Pawe Agricultural Research Centre; JARC and PARC are under EIAR; masl = meters above sea level. 2.3. Data collection Necessary parameters were collected per plot for each location. These data (parameters) include the number of days to flowering for male (DM) and female (DF), i.e., when 50% of the plants within a plot start shedding pollen and silks emerge (2–3 cm). Plant height (PH) and ear height (EH) were measured in cm as the distance from the base of the plant to the first tassel branch and the node bearing the upper ear, respectively. The general performance of hybrids was scored as a plant aspect on a 1–9 scale, where 1 = very good and 9 = very poor phenotypic performance in terms of uniformity, vigorousness, resistance to disease and overall phenotypic appearance. Foliar diseases such as Turcicum leaf blight, grey leaf spot, and rust were also rated using 1 to 9 scoring scale 19 , where. 1 = very resistant and 9 = susceptible to the diseases. The number of plants and ears per plot was counted. Ear aspect was scored on 1 to 9 scales, where 1 = well-filled, uniform kernel arrangement, free from ear disease, and large or medium ears, while 9 = ear disease, poorly filled, and shrivelled ears. The total number of ears per row was weighted (kg/plot area), and the moisture content of the representative sample grain was measured for each unshelled plot using a portable moisture tester. Grain yield (tone/ha) was calculated on the basis of ear weight per plot, adjusted to 12.5% moisture content and harvested plot size area. Adjusted grain yield (tone/ha) = \(\:\frac{FW\left(100-MC\right)*0.8*10}{87.5*PA}\) Where; FW = Field weight of total ear weighted in kg/plot area, MC = moisture content of grain sample at harvest, 0.8 = shelling percentage, 10 = to convert quintal into tone, 87.5 = 100 minus from 12.5 standard moisture, PA = plot size area 2.4. Statistical analysis Individual site and combined analysis across locations were carried out using PROC GLM procedure in SAS version 9.3 20 . Genotype was considered as fixed effect whereas replication (location), block (replication), location × genotype was considered as random effects. The significances of genotypes and interaction effects were tested and mean separation was done using least significant difference (LSD) at 0.05. General combing ability analysis Diallel data analysis procedure was carried out using analysis of genetic designs with R (AGD-R) 21 software. The general and specific combining abilities and variance components for crosses and important traits was esteemed using Griffing's method II and model I 22 . Model for multilocation diallel test Yijrl = µ + r(l) + l + gi + gj + sij + l*gi + l*gj + l*sij + eijk, Where µ is the grand mean, r is the replication effect, r(l) = replication nested under location (r = 1..2), l = location(1..2), gi and gj are GCA effect (1…12), sij is the SCA effect, l*gi and l*gj are location by GCA effect interactions, l*sij is location by SCA effect interaction and eijrl is the experimental pooled error effect Estimate Heterosis Heterosis, mid parents as well as standard heterosis and their significant was calculated according to formula indicated in 23 , 24 MPH= \(\:\frac{\left(F1-Mp\right)\text{*}100}{MP}\) ; MP= \(\:\frac{\left(P1+P2\right)}{2}\) , where, MPH= Mid Parent Heterosis, MP= is an average value of the corresponding parents for the hybrid, F1= hybrid value. Economic (standard heterosis, SH) or percent of heterosis, SH (%) = \(\:\frac{\left(F1-SC\right)\text{*}100}{SC}\) , where, SH= standard heterosis, SC= standard check, F1= mean performance of F1 (hybrid) Significance of each hybrid value with respect to the corresponding MP and SH values were tested using t test 24 . \(\:tmp=\frac{\text{F}1-\text{m}\text{p}}{\frac{\sqrt{3MSE}}{2r}}\) , \(\:SH\left(t\right)=\frac{\text{F}1-\text{S}\text{C}}{\frac{\sqrt{2\text{M}\text{S}\text{E}}}{\text{r}}}\) , where, MSE= error mean square 3. Result The ANOVA showed a highly significant difference (P ≤ 0.001) among PVA hybrids for yield and most traits at each location, suggesting the presence of genetic variation among provitamin A hybrids performance under field conditions and feasibility selection of hybrids for grain yield and other traits. Furthermore, combined analysis of variance revealed a significant difference (P ≤ 0.01) in mean square due to PVA hybrids interaction with locations for grain yield and other agronomic traits studied (Table 3 ). Mean performance of PVA hybrids The mean performance of PVA hybrids along with checks for grain yield, days to anthesis, days to silking and plant height is presented in Table 4 . The highest grain yield (7.95 t/ha) was recorded for cross L9/L12, whereas L4/L5 showed the lowest yield (2.45 t/ha) with a mean of 5.20 t/ha across locations. The mean values for traits across locations ranged from 69.66 to 84.33 for days to anthesis with a mean of 76.04, 71.50 to 87.16 for days to silking with a mean of 78.75 and plant height (cm) from 223.39 to 161.89 with a mean of 198.15 and 6.00 to 2.31 for GLS with a mean of 3.7. Seventeen PVA hybrids showed better performance for grain yield than checks; out of these, seven PVA hybrids were selected with a mean yield ranged from 7.11 t/ha to 7.95 t/ha and showed 18.69–32.72% yield advantage over the better check (Table 4 ). In addition, the selected PVA hybrids showed earlier flowering (male anthesis and silking days) than checks without yield penalty across locations. Table 3 Mean squares from ANOVA for yield and important agronomic traits of 68 PVA maize hybrids evaluated across three locations in 2022 Notice = SV = source of variation, DA = days to anthesis, DS = days to silking, Plht = plant height, Erht = ear height, GLS = grey leaf spot, TLB = turcicum leaf blight SV df Mean square of yield and agronomic traits Yield DA DS Plht Erht GLS TLB Loc 2 127.04 *** 3658.26 *** 3837.88 *** 1077881.64 ** 100508.32 ** 254.05 ** 281.75 Rep 1 21.28 8.53 35.29 10.05 21.04 6.83 6.75 Blk(rep) 6 0.44 5.21 11.86 320.31 82.91 2.31 2.19 Genotype 67 8.95 *** 38.06 *** 47.46 ** 934.27 ** 505.9 7*** 4.09 *** 1.96 ** Rep(loc) 2 13.31 20.63 44.21 2291.32 628.05 6.91 3.59 Loc*Genotype 134 4.53 *** 18.00 ** 22.47 ** 591.00 ** 207.73 ** 1.96 *** 1.30 Error 195 01.63 0.12 7.14 0.16 0.58 1.02 0.88 Table 4 Selected PVA hybrids for yield and other agronomic traits entry Hybrids Yield advantage over best check (%) Yield (t/ha) DA DS Plant height 63 L9 x L12 32.72 7.95 75.50 78.33 219.00 61 L9 x L10 29.54 7.76 75.16 77.61 220.50 65 L10 x L12 29.04 7.73 72.66 74.83 209.00 62 L9 x L11 27.37 7.63 77.83 80.00 198.62 44 L5 x L11 22.03 7.31 74.83 76.46 221.28 42 L5 x L9 19.69 7.17 75.16 78.43 207.72 60 L8 x L12 18.69 7.11 76.50 79.33 220.50 67 Check, BHQPY545 5.99 81.33 84.00 201.11 68 Check, BH 549 5.91 84.33 87.16 206.72 Mean 5.20 76.04 78.75 198.52 CV 24.55 3.75 4.21 9.22 LSD.05 1.45 3.09 3.78 20.82 Analysis of variance: GCA for inbred lines and SCA for cross effects across locations Mean squares due to location, crosses, general combining ability (GCA), specific combining ability (SCA), location x crosses, GCA × location, GCA x lines, SCA × location, and SCA x crosse are presented in Table 5 . Significant mean squares due to GCA for lines and SCA for crosses indicate that both additive and non-additive gene actions control the inheritance of most traits studied. Furthermore, the mean square of SCA x locations and crosses x locations showed a significant difference (P ≤ 0.001) in the combined ANOVA for all traits studied (Table 5 ), indicating that SCA and crosses were affected by environments. Except for GLS and TLB traits, the mean square due to GCA of lines x location was non-significant in the combined ANOVA, suggesting that the inheritance of PVA-enriched lines is controlled by additive gene action and/or is unlikely to be affected by environment interaction for studied traits. Table 5 Summary mean squares of combined ANOVA across environments for yield and important agronomic traits of 66 PVA crosses SV df Mean square of yield and agronomic traits Yield DA DS Plht Erht GLS TLB PA EA Location 1 73.70** 1138.00ns 144.05** 141869.82* 110454.00ns 106.75** 141.89ns 29.78*** 3.59ns Rep(loc) 2 1.35* 21.05ns 23.75ns 1208.96* 434.97ns 29.73ns 17.65** 8.14ns 8.62*** Crosses 77 11.35*** 29.45*** 39.18*** 2219.72*** 892.60*** 3.29*** 2.52** 3.49** 4.26* GCA 11 18.98* 84.76*** 125.89*** 4781.88*** 2196.67** 7.78* 9.51** 6.20ns 6.54ns SCA 66 10.28** 20.26** 24.73ns 1792.70* 675.26ns 2.55*** 1.37ns 3.04* 3.81ns Crosses x Location 77 5.50*** 11.11*** 17.38*** 981.58*** 433.19*** 1.44ns 1.42ns 2.04** 2.76*** GCA x Location 11 4.51ns 12.36ns 17.02ns 603.ns 124.70ns 2.55* 3.03** 3.37ns 3.91ns SCA x Location 66 6.00*** 10.90*** 17.44*** 1044.52*** 484.61*** 1.25ns 1.18ns 1.82* 2.57*** Error 150 1.24 5.32 8.09 340.33 157.99 1.21 1.40 1.12 1.28 GCA: SCA 0.14 0.37 0.50 0.21 0.28 0.35 0.50 0.19 0.16 General combining ability effect of lines for grain yield and other traits The general combining ability (GCA) effect showed significant differences ( P < 0.05 to 0.001) among PVA-enriched lines for yield and other traits (Table 6 ). This suggests that at least one inbred line differs from other lines in terms of additive gene and additive x additive gene effects for agronomic traits studied. The GCA effects of lines for yield varied from − 0.85 to 0.99 t ha - 1 . Among all lines, L12, L8, L6, L10, and L2 showed desirable positive GCA effects on grain yield. In contrast, L3 had the lowest GCA effect (-0.85) for grain yield. L12 had the highest GCA effect for yield as well as a good combiner for grain yield, meanwhile it had a negative GCA effect for DA, DS, GLS, and TLB. Similarly, L8 was a good combiner for grain yield and other desirable agronomic traits. Thus, L12 (Tester B) and L8 (Tester A) can be used as alternative testers for further PVA maize breeding programs. Specific Combining Ability effect of crosses for grain yield and other traits The SCA effects of crosses which indicates whether their performance is better or worse than what would be expected from the GCA effects of their respective parents (Table 7 and Supplementary Table S1 ). The highest positive SCA effect for grain yield was recorded for crosses of L7/L12 (3.16 t/ha), followed by L7/L9 (2.43 t/ha). At the same time, these crosses had negative significant effects for DA and DS, suggesting the tendency of earliness. In contrast, L4 and L5 had the lowest cross-combinations for grain yield (-2.43 t/ha) with negative SCA effects. Furthermore, crosses of L1 and L10 exhibited a significant positive effect of PA and EA. Similarly, crosses of L10 x L12, L5 x L9, and L8 x L12 showed a positive effect for plant and ear height and a significant negative SCA effect for flowering as well as for TLB and GLS, suggesting that these parental combinations could be good sources of hybrid development for resistance to disease and earliness. Table 6 Estimates of General Combining Ability (GCA) effects for grain yield and important agronomic traits of twelve PVA enriched maize lines evaluated in 2022. Lines Grain yield and important agronomic traits GY DA DS Plht Erht GLS TLB PA EA 1 -0.40*** -1.92*** -2.29*** 2.24ns -0.89ns -0.26* 0.18ns 0.37*** -0.01ns 2 0.23* -0.67* -1.18*** 2.92ns 3.01* -0.39*** -0.07ns -0.45*** -0.47*** 3 -0.85*** 1.06** 1.45*** -6.18* -1.50ns 0.40*** 0.24ns 0.21* 0.05ns 4 -0.18ns 1.54** 1.93*** 7.42*** 0.73ns -0.17ns -0.34** -0.13ns -0.20ns 5 -0.17ns 0.17ns 0.20ns 1.52ns -1.82ns 0.07ns 0.26* -0.05ns 0.01ns 6 0.41*** 0.65** 0.54ns 3.80* 3.86** 0.23ns 0.22ns 0.06ns -0.01ns 7 -0.11ns 2.13** 2.66*** -7.35*** -3.65** -0.70*** -0.73*** 0.05ns 0.06ns 8 0.93*** 0.31ns 0.23ns 13.70*** 11.72 *** 0.03ns -0.07ns -0.65*** -0.60*** 9 -0.38*** -1.28*** -1.34*** -8.01*** -5.65 *** 0.69*** 0.80*** 0.33* 0.44*** 10 0.26* -1.44** -1.68*** 0.43ns -2.01ns 0.11ns 0.20ns 0.13ns 0.39*** 11 -0.71*** -0.44ns -0.48ns -20.32*** -12.07 *** 0.21ns -0.13ns 0.42** 0.58*** 12 0.99*** -0.10ns -0.04ns 9.83*** 8.26 *** -0.21ns -0.56*** -0.28* -0.24* GCA SE 0.10 0.21 0.25 1.66 1.13 0.99 0.10 0.09 0.10 Table 7 Specific combining ability (SCA) effect of twelve yellow maize inbred lines for grain yield L1 L2 L3 L4 L5 L6 L7 L8 L9 L10 L11 L12 L1 0.47 0.18 1.16* -1.13* -0.06 -0.29 0.99 0.34 -0.81 0.92 -0.16 L2 1.13* 2.41** 0.44 -0.55 -1.67*** 1.20* 0.39 0.53 -0.38 -0.46 L3 -0.83 0.29 -0.05 1.96*** 0.28 0.92 -2.54 0.67 0.14 L4 -2.43*** -0.08 0.02 2.08*** -0.28 0.69 2.05*** 0.17 L5 1.42*** -0.76 -0.36 2.00*** 1.67*** 0.29 1.07* L6 -1.42*** 1.11 0.51 1.40*** 0.85 2.26* L7 0.19 2.43*** 0.18 0.93 3.16** L8 -1.56** 1.97*** -0.26 1.75** L9 -0.24 -0.32 -0.83 L10 -0.85 2.05*** L11 -0.26 L12 Heterosis effect Mid-parent heterosis The values of mid-parent and standard heterosis were estimated for grain yield, DA, DS, and GLS across locations (supplementary Table S2 ). Significant (P < 0.05) to highly significant difference (P < 0.01) positive mid-parent heterosis (MPH) estimate for grain yield was recorded for 20 crosses (supplementary Table S2 ) which ranged from 99.30–685.59%, suggesting that the crosses performed better than the mean performance of the mid-parent. Regarding flowering, thirty-one and fourteen crosses exhibited significant (P < 0.05) negative mid-parent heterosis for DA and DS, respectively. L10 and L12 showed the highest mid-parent heterosis (685.59%), followed by L9 and L10 (509.59%) for grain yield. These crosses also showed significant negative MPH for flowering (DA and DS), indicating good yield performance with earliness. Standard Heterosis Only one cross (L9/L12) displayed significant (p < 0.05) positive standard heterosis (34.52%) over BH 549 (normal maize hybrid check for grain yield), while eleven crosses exhibited significant (p < 0.05) negative heterosis, ranging from − 58.84% to -34.01% for grain yield. On the other hand, none of the crosses showed significant positive standard heterosis, while nine crosses exhibited significant (p < 0.05) negative heterosis for grain yield compared to BHQPY 545, a yellow quality protein maize check (Supplementary Table S2 ), suggesting further improvement of PVA hybrids for grain yield. But it is to be noted that the cross L7/ L12 had the same yield (zero heterosis) compared to BHQPY 545 for grain yield. Regarding days to anthesis and silking, most of the crosses showed significant (p < 0.05) to highly significant (p < 0.01) negative heterosis for days to tasselling and silking as compared to BH 549 and BHQPY 545 checks, which indicates the tendency of these crosses for earliness. The estimated standard heterosis for GLS in four crosses (L2 / L6, L4 / L5, L4 / L10, and L5 / L6) were positive and significant, ranging from 76.67–100% over the standard checks BH549 and BHQPY545, indicating their likely susceptible to GLS when compared to the checks. On the other hand, some crosses showed negative standard heterosis for GLS, implying a tendency towards resistance to the disease. It was interesting that three crosses showed zero heterosis compared to checks for GLS recorded, suggesting that theses crosses and checks had similar GLS disease severity. 4. Discussion Performance of maize hybrids, Combining Ability Effects and Tester Identification Our result demonstrated that the significant mean square observed for provitamin A hybrids, locations, and locations x hybrids for grain yield and most other measured agronomic traits indicated the ranking of genotypes across environments at mid-altitude subhumid agro-ecology of Ethiopia. Maize hybrids with high grain yields were selected for (L9/LL12), (L10/LL12), (L9/LL10), (L9/LL11), L5/L11, L5/L9 and L8/L12 that showed 18.69–32.72% yield advantages over checks as well as early flowering without yield penalty. Moreover, L10/L12, L5/L9, and L8/L12 had high yield potential with positive SCA effect and could be changed into three-way crosses with the opposite testers. The better performance of those hybrids in terms of yield and other traits might be due to the contribution of favorable alleles 25 and other associated genes in their parents during cross-combination. Thus, the selected PVA hybrids should be tested further, including their enriched provitamin A content and stability across environments 26 The current study revealed that the mean square due to GCA for lines and SCA for crosses showed a significant difference for all traits measured, indicating that both additive and non-additive gene actions control the inheritance of most traits studied 14 , 27 , suggesting that improvement in these traits could be achieved through the selection of parents and their crosses for hybrid maize improvement. However, the ratio of SCA variance was higher than the GCA variance components for all traits studied, indicating that the predominance of non-additive gene action was more important in controlling the inheritance of traits, which differed from results of other studies 28 , 29 , but it was consistent with 30 , 31 who reported the non-additive gene effect for governing grain yield and other traits. Such contradictory result reported on the mode of inheritance of traits highlighted that might be due to the type of germplasm used and the environmental condition under which the study carried out 32 . Thus, further testing is required on Pro-vitamin A enriched with germplasm having diverse genetic background across multiple environments. The GCA effect of lines is important for the selection of parent and hybrid development in the direction of yield improvements and other traits. In the current study, L2, L6, L8, L10, and L12 showed positive contributions to grain yield and other traits, suggesting the presence of additive and additive x additive gene action in the inheritance of studied traits 14 . Also, these inbred lines that are within heterotic groups can be used for further inbred line development and hybridization 33 . Among the lines, L8 and L12 showed the highest GCA effect for grain yield and seemed to have the best average performance for yield when crossed with other lines; had positive contribution to height and negative to GLS and TLB, suggesting the development of hybrid resistance to disease. Moreover, crosses of L8 and L12 exhibited one of the top average performances for hybrid yield and other traits. The choice of tester in maize hybrid breeding programs is determined by the capacity of the tester to discriminate among maize inbred lines, GCA effects for traits and average testcross performance for yield 34 ; 15 . This result was consistent with the findings of other studies 14 ; 17 . Thus, L8 (Tester A) and L12 (Tester B) can be used as alternative opposite testers to evaluate new inbred lines and/or progenies across optimum conditions. Twenty crosses showed yield performance ranging from 99.30–685.59% compared to mid-parent heterosis. This result is in line with that of 14 who reported that the top crosses for grain yield ranged from 488 to 539% of mid-parent heterosis. Similarly, 10 who reported cross hybrids that expressed significantly higher than mid-parent heterosis for grain yield and other traits. In this study, the top three crosses (L10 / L12 (685.59%), L9/LL10 (509.55%), and L9/LL11 (505.05%) supplementary Table S2 ) showed the highest mid-parent heterosis for grain yield. Moreover, their single-cross combination was also found to be the top hybrid performance for yield which might be due to difference in the origins of parents, genetic divers and dominant gene action. The aim of maize breeders is to develop and select stable superior maize hybrids that are better than checks in terms of grain yield and other desirable traits. Our result showed that only one cross (L9 / L12) had significant standard heterosis over BH 549, a normal maize hybrid check, whereas there was a non-significant positive heterosis with BHQPY 545, a yellow quality protein maize hybrid check for grain yield. On the other hand, cross L7 and L12 had zero heterosis with BHQPY 545 for grain yield, indicating that this cross had the same yield as BHQPY 545. Thus, further PVA breeding is required with comprehensive allelic variants and genetic divergence of parents for crosses combination in heterosis manifestation for yield and other agronomic traits. 5. Conclusion The current study showed that F1 hybrids (L9 / L12), (L10/LL12), (L9/LL10), (L9/LL11), L9/L11, L5/L11, L5/L9, and L8/L12 exhibited better yield performance over checks and it need further testing across locations. Inbred lines L2, L6, L8, L10, and L12 had the highest GCA effects for grain yield, suggesting the transfer of favorable alleles to their progenies; thus, these inbred lines within the heterotic group could be used as new sources of PVA inbred line development. Out of these, L8 and L12 showed the highest GCA effect for grain yield, better average performance for yield when crossed with other lines, and negative contributions for GLS and TLB diseases. Thus, L8 (Tester A) and L12 (Tester B) can be used as alternative opposite testers to evaluate new PVA inbred lines and/or progenies across optimum conditions. Declarations Ethics approval and consent to participate Plants materials involved in this research are used for scientific research. This article did not contain any studies with human participants or animals and did not involve any protected species. Competing interest The authors declare that no conflict of interest Author Contribution BG: Writing – original draft, Investigation, Data curation, Formal analysis, Software. GA: Writing – editing draft, Methodology, Visualization and Project administration; KB: Supervision, Writing – review & editing, AM: Writing – review and editing, Project administration, Funding acquisition. Acknowledgment The authors highly acknowledge the Harvest Plus Challenge Program for funding this work, particularly for carotenoid analyses. The author would like to thank EIAR/BNMRC for funding this activity, particularly for testing trials across locations. The authors also express appreciation to all BNMRC and EIAR collaboration staff who participated in planting and managing trials. The authors also gratefully acknowledge the help of anonymous editorials and reviewers for improving this article Data Availability all relevant collected data are included in this article. 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Agronomy . 10 , 1–14. 10.3390/agronomy10020184 (2020). Sandesh, G. M. et al. Heterosis and combining ability studies for yield and its component traits in Maize (Zea mays L). Electron. J. Plant. Breed. 9 (3), 1012–1023 (2018). Amegbor, I. K., Van Biljon, A., Shargie, N. G., Tarekegne, A. & Labuschagne, M. T. Combining ability estimates for quality and non-quality protein maize inbred lines for grain yield, agronomic, and quality traits. Front. Sustain. Food Syst. 7 , 1123224. 10.3389/fsufs.2023.112322 (2023). Udo, E., Abe, A., Meseka, S., Mengesha, W. & Menkir, A. Genetic Analysis of Zinc, Iron and Provitamin A Content in Tropical Maize (Zea mays L). Agronomy . 13 , 266. https://doi.org/10.3390/agronomy13010266 (2023). Makumbi, D., Betran, F., Beanziger, M. & Ribaut, J. M. Combining ability, heterosis and genetic diversity in tropical maize (Zea mays L.) under stress and non-stress conditions. Euphytica . 180 , 143–162 (2011). Guimarães, L. M. et al. L.V.D. Performance of testers with different genetic structure for evaluation of maize inbred lines. Ciência Rural . 42 , 770–7769 (2012). Additional Declarations No competing interests reported. Supplementary Files SupplemTableS2MidParentStandHeterosis.docx SupplementaryTableS1SCAeffect.docx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4839825","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":385305464,"identity":"66c67a41-2b4b-4926-bf79-f5f8f7f6730c","order_by":0,"name":"Belay Garoma","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA7klEQVRIiWNgGAWjYBACAwbmBhib8QGQ4OEjrIURroXZAKSFjRQtbBJgkpAWc/bGxs88v+zy+cUOP6v8mmMnw8bA/PDRDTxaLHsONkvz9iVbzpydZnZbdlsy0GFsxsY5+Bx2I7FBmreH2cDgdoLZbcltzEAtPGzSeLXcf9j8m7enHqgl/Vux5LZ6IrTcYGyT5vlxGKglx4zx47bDhLVY9iS2Wc5tOG4gOTunWJpx23EeNmYCfjFnP3z4xps/1Qb80ukbP/7cVm3Pz9788DE+LSDAxNsGYTDzgEkCykGA8ccfGIMI1aNgFIyCUTDyAACpUkcePHOhUwAAAABJRU5ErkJggg==","orcid":"","institution":"Ethiopian Institute of Agricultural Research","correspondingAuthor":true,"prefix":"","firstName":"Belay","middleName":"","lastName":"Garoma","suffix":""},{"id":385305465,"identity":"68b78746-a7c4-4dc2-821e-8f8f2484a9fd","order_by":1,"name":"Kassahun Bante","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Kassahun","middleName":"","lastName":"Bante","suffix":""},{"id":385305466,"identity":"7807deda-cfd2-469e-9272-a14145451a29","order_by":2,"name":"Girum Azimach","email":"","orcid":"","institution":"Ethiopian Institute of Agricultural Research","correspondingAuthor":false,"prefix":"","firstName":"Girum","middleName":"","lastName":"Azimach","suffix":""},{"id":385305467,"identity":"d306a7db-9920-4ec1-8bff-dfe3c1301a4c","order_by":3,"name":"Abebe Menkir","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Abebe","middleName":"","lastName":"Menkir","suffix":""}],"badges":[],"createdAt":"2024-08-01 06:44:20","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4839825/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4839825/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":87160029,"identity":"0b5f7654-e0fc-4606-9e8e-8beb536ecf5d","added_by":"auto","created_at":"2025-07-21 04:23:50","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1376412,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4839825/v1/abfac366-06eb-4f21-9427-a0474634ce23.pdf"},{"id":70528338,"identity":"9820f9dd-e908-44bb-ab16-5d79a150d3af","added_by":"auto","created_at":"2024-12-04 05:31:27","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":29893,"visible":true,"origin":"","legend":"","description":"","filename":"SupplemTableS2MidParentStandHeterosis.docx","url":"https://assets-eu.researchsquare.com/files/rs-4839825/v1/bfc908d1e7086adc06b784f2.docx"},{"id":70528337,"identity":"92a70788-9b37-48e5-9a95-a64dfe0594c5","added_by":"auto","created_at":"2024-12-04 05:31:27","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":29231,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryTableS1SCAeffect.docx","url":"https://assets-eu.researchsquare.com/files/rs-4839825/v1/e88895e23a02732bd63ff11a.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Combining Ability and Heterosis for Grain Yield and Other Agronomic Traits in Provitamin A Enriched Maize Inbred Lines","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eMaize (\u003cem\u003eZea mays\u003c/em\u003e, L.) is one of the most important staple food crops in developing countries and provides over 20% of the calories in the human diet\u003csup\u003e\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u003c/sup\u003e. However, most of the maize varieties cultivated in these countries have low micronutrient as well as vitamin A content\u003csup\u003e\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u003c/sup\u003e which leads to micronutrient deficiency including vitamin A deficiency. Diverse and nutrient-rich foods such as fruits, vegetables and animal-based products are often inaccessible for many rural communities in low and middle-income countries. Thus, biofortification of staple foods through conventional breeding and application of biotechnological tools is a sustainable and affordable approach to increase vitamin and mineral content in staple food crops\u003csup\u003e\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e\u003c/sup\u003e. Maize kernels accumulate diverse carotenoids with provitamin A activity and biofortification breeding has been initiated; subsequently, efforts have been made to increase PVA concentration in maize inbred lines\u003csup\u003e\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eThe usefulness of inbred lines for maize breeding program is based on their combining ability and high heterosis expression in hybrid performance. Diallel mating scheme was developed by Griffing (1956) and this scheme is used to study combing ability and heterosis phenomena among inbred lines\u003csup\u003e\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e,\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u003c/sup\u003e. Combing ability studies also provide useful information about the nature of gene action for particular traits\u003csup\u003e\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u003c/sup\u003e and to select superior parents for new sources of population and hybrid developments\u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e\u003c/sup\u003e. Heterosis refers to the superior performance of F\u003csub\u003e1\u003c/sub\u003e hybrids relative to their parents in terms of vigor or other desirable agronomic traits\u003csup\u003e\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. Cross hybrids expressed significantly higher than mid-parent heterosis for grain yield and provitamin A content\u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e\u003c/sup\u003e. Likewise, hybrids exhibited the highest specific combining ability and heterosis for grain yield compared to commercial hybrids\u003csup\u003e\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e. Moreover, \u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u003c/sup\u003edemonstrated that the combining ability among yellow maize inbred lines showed significant general combining ability (GCA) and specific combining ability (SCA) effects for grain yield, provitamin A content and other traits.\u003c/p\u003e \u003cp\u003eCombing ability analysis also helps to identify testers that are important in maize breeding to employ early screening progeny, grouping inbreeds, to select superior parents and top hybrids in trials performance. The choice of desirable testers can depend on the genetic difference in their progeny for tested traits\u003csup\u003e\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e\u003c/sup\u003e, high GCA effect on yield\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e, gene frequency of favorable alleles \u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e\u003c/sup\u003e and the discrimination of the differences among tested parents\u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u003c/sup\u003e. Many studies have reported on the selection of effective testers in partial diallel crosses. For example, \u003csup\u003e\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u003c/sup\u003eidentified testers that had the highest GCA effects for provitamin A and other traits. \u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e; \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u003c/sup\u003eselected testers that showed strong positive significant and high GCA effects for grain yield and ears per plant. Thus, continuous assessment of early and new provitamin A-enriched yellow maize inbred lines is important for their combing ability to select ideal testers and superior parents for hybrid development.\u003c/p\u003e \u003cp\u003eIn the earlier study, we identified medium to high PVA content of yellow/orange maize inbred lines from different genetic background sources, pedigrees recorded and heterotic groups. However, no information is available on combining abilities of newly developed PVA-enriched inbred lines and heterosis for grain yield, other agronomic traits and suitable testers. Therefore, the objectives of the study were: i) To evaluate the combing ability of PVA-enriched inbred lines for grain yield and important agronomic traits. ii) to identify provitamin A testers, the best-performing hybrids and estimate heterosis of F1s over check and mid-parents for yield and other agronomic traits.\u003c/p\u003e"},{"header":"2. Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Source of plant materials and crossing techniques\u003c/h2\u003e \u003cp\u003eThe twelve PVA enriched yellow/orange maize inbred lines included in this study were developed at the Maize Improvement Program of IITA, CIMMYT and Bako National Mazie Research Centre of the Ethiopian Institute of Agricultural Research (BNMRC/ EIAR) as following. The two PVA enriched inbred lines CML486/(CML297-B\u0026thinsp;\u0026lt;\u0026thinsp;d7\u0026thinsp;\u0026gt;\u0026thinsp;KUICarotenoidsyn-FS17-3-2-B/KUI3\u0026thinsp;\u0026lt;\u0026thinsp;d7\u0026thinsp;\u0026gt;\u0026thinsp;B77))-B-11-1-B-B-B-B-B-B-B-#-B-B-B (L10) and (CML537/[BETASYN]BC1-10-2-1-#/CML-305-B)-B-11-1-B-B-#-B-B-B (L12) were introduced from CIMMYT evaluated over years and selected for adaptation to local condition at Bako, Ethiopia. The three PVA enriched inbred lines TZMI1003-#-# (L8), TZMI1003-#-# (L9) and CLHP00306 (L11) introduced from International Institute of Tropical Agriculture (IITA) were also selected for adaption to locational condition at Bako, Ethiopia. Other inbred lines introduced from CIMMYT and used as donor parents for beta carotene include KUI carotenoid syn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-3(MAS:L4H1); KUI carotenoid syn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-1(MAS:L4H1; KUI carotenoid syn-FS3-3-2-B-B-B(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1; proA-HydB1)-#bal]FS and Florida A plus syn-FS2-2-1-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1) were crossed with locally adapted elite white maize inbred lines of BNMC/ EIAR includes; DE78-Z-126-3-2-2-1-1(g), Gibe1-91-1-1-1-1, IL00'E-9-1-1-1-1-1, Obatanpa and CML 161 as recipients. The main donors of high b-carotene for the IITA, CIMMYT and yellow/orange Ethiopian lines included in this study were originally temperate inbred line and source populations including DE3, [BETASYN]BC1-10-2-1-#, KUI carotenoids syn-FS17-3-2-B, KUI Carotenoids syn-FS11-1-1-B-B-B, KUI carotenoid syn-FS3-3-2-B-B-B, and (ProA-HydB1)-#ba1] FS. Yellow inbred lines were then derived from biparental crosses through re-repeated generation of inbreeding with selection for desirable kernel colour and agronomic attributes using the pedigree breeding approach. The advanced yellow/orange maize inbred lines were then evaluated across locations for their adaptation and agronomic performance and reaction to disease under natural condition. The advanced yellow/orange maize inbred lines along with selected IITA and CIMMYT lines were planted in trials evaluated in 2020 and 2021 at Bako, Ethiopia. Seed samples drawn from these trials were shipped to CIMMYT-Mexico for carotenoid analyses\u003csup\u003e\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e\u003c/sup\u003e. Based on the results of carotenoids analysis, twelve yellow/orange maize inbred lines with diverse genetic backgrounds and PVA content exceeding 6 \u0026micro;g/g were selected from the different heterotic groups based on available of seed in stock (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The twelve PVA enriched yellow/orange maize inbred lines were crossed in a 12 x 12 half-diallel mating design to generate 66 F\u003csub\u003e1\u003c/sub\u003e crosses at National maize research centre during the 2021 dry season at Bako, Ethiopia.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eList of Provitamin A enriched maize inbred lines used to generate half diallel crosses\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInbred lines code\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePedigree\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSource of yellow maize inbred lines\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePVA content (\u0026micro;g/g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSeed source in stock\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eStock ID\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eHeterotic group\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(DE78-Z-126-3-2-2-1-1(g)/(KUI carotenoid syn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-3(MAS:L4H1)-1-B-B-B)-B-1-1-1-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9.96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N7-45-1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(DE78-Z-126-3-2-2-1-1(g)/(KUI carotenoid syn-FS11-1-1-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-3(MAS:L4H1)-1-B-B-B)-B-13-1-2-B -B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N7-47-1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(Gibe1-91-1-1-1-1/(KUI carotenoid syn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-1(MAS:L4H1)-5-B-B-B)-B-7-1-1-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N7-51-1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(IL00'E-9-1-1-1-1-1/(KUI carotenoid syn-FS17-3-2-B-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-1(MAS:L4H1)-5-B-B-B)-B-13-1-4-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N7-14-1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eB\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(IL00'E-9-1-1-1-1-1/(KUI carotenoid syn-FS3-3-2-B-B-B(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1)-2-B-B-B)-B-2-2-1-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N7-17-1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eB\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e[(Obatanpa(proA-HydB1)-#bal]FS-93-3-1-1-2-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-BK18MN33-25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCML161/(Florida A plus syn-FS2-2-1-B-B/(KU1409/DE3/KU1409)S2-18-2-B)-B-4(MAS:L4H1)-2-B-B-B//CML161-6-1-2-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N7-64-1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL53-49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTZMI1003-#-#\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIITA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N30-34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL75-33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTZMI1065-#-#\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIITA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N30-34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL75-34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eB\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(CML486/(CML297-B\u0026thinsp;\u0026lt;\u0026thinsp;d7\u0026thinsp;\u0026gt;\u0026thinsp;KUICarotenoidsyn-FS17-3-2-B/KUI3\u0026thinsp;\u0026lt;\u0026thinsp;d7\u0026thinsp;\u0026gt;\u0026thinsp;B77))-B-11-1-B-B-B-B-B-B-B-#-B-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCIMMYT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N30-132\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eA\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCLHP00306\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIITA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N30-31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL75-31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eB\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(CML537/[BETASYN]BC1-10-2-1-#/CML-305-B)-B-11-1 -B-B-#-B-B-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCIMMYT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2019K-MQ19N30-134\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eQL75-30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eB\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eNotice: BNMRC/EIAR\u0026thinsp;=\u0026thinsp;Bako National Maize Research Centre under Ethiopian Institute of Agricultural Research, Bako, Ethiopia; IITA\u0026thinsp;=\u0026thinsp;International Institute of Tropical Agriculture, Ibadan, Nigeria. CIMMYT\u0026thinsp;=\u0026thinsp;International Mazie and wheat Improvement Centre, Stock ID and seed source indicates the availability of seed in seed store or in cold room at Bako National maize research centre during inbred lines planted.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Trial management\u003c/h2\u003e \u003cp\u003eSixty-eight F1 hybrids plus two checks were arranged in a 17 x 4 alpha lattice design and planted with two replications. Each hybrid was planted on 4 m long rows with 16 plants per row, with 75 cm and 25 cm spacing between rows and plants, respectively. At planting, two seeds per hill were planted, and after two weeks of emergency, they were thinned out to maintain one plant per hill. Trials were planted and evaluated at Bako, Jima and Pawe agricultural research centres in the 2022 main cropping season. The parental lines of the diallel crosses were also planted separately adjustment to the diallel cross trial in a randomized complete block design with two replications in the 2022 main cropping season. Agronomic management, such as weeding and fertilizer application was applied as per research recommendations at the respective field-testing site. Description of testing locations are presented in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eDescription for the test locations used to evaluate Provitamin A maize hybrids\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCountry\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEIAR\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCo-ordinates\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAltitude (m.a.s.l)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eRainfall (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMinimum Temperature (\u003csup\u003e0\u003c/sup\u003eC)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eMaximum Temperature (\u003csup\u003e0\u003c/sup\u003eC)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSoil type\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBako\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEthiopia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEIAR/BNMRC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9\u003csup\u003e0\u003c/sup\u003e06\u0026rsquo;N and 37\u003csup\u003e0\u003c/sup\u003e09' E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1650\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1244\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e12.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e29.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eNitosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eJima\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEthiopia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEIAR/JARC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7\u003csup\u003e0\u003c/sup\u003e40\u0026rsquo;N and 36\u003csup\u003e0\u003c/sup\u003e 49\u0026rsquo;E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1753\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1572\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e26.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eEutric Nitosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePawe\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEthiopia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEIAR/PARC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e11\u0026deg;19'N and 36\u0026deg;24'E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1120\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1238\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e16.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e32.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eNitisol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eNotice: EIAR/BNMRC\u0026thinsp;=\u0026thinsp;Ethiopian Institute of Agricultural Research/Bako National Maize Research Centre; JARC\u0026thinsp;=\u0026thinsp;Jima Agricultural Research Centre; PARC\u0026thinsp;=\u0026thinsp;Pawe Agricultural Research Centre; JARC and PARC are under EIAR; masl\u0026thinsp;=\u0026thinsp;meters above sea level.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Data collection\u003c/h2\u003e \u003cp\u003eNecessary parameters were collected per plot for each location. These data (parameters) include the number of days to flowering for male (DM) and female (DF), i.e., when 50% of the plants within a plot start shedding pollen and silks emerge (2\u0026ndash;3 cm). Plant height (PH) and ear height (EH) were measured in cm as the distance from the base of the plant to the first tassel branch and the node bearing the upper ear, respectively. The general performance of hybrids was scored as a plant aspect on a 1\u0026ndash;9 scale, where 1\u0026thinsp;=\u0026thinsp;very good and 9\u0026thinsp;=\u0026thinsp;very poor phenotypic performance in terms of uniformity, vigorousness, resistance to disease and overall phenotypic appearance. Foliar diseases such as Turcicum leaf blight, grey leaf spot, and rust were also rated using 1 to 9 scoring scale\u003csup\u003e\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e\u003c/sup\u003e, where. 1\u0026thinsp;=\u0026thinsp;very resistant and 9\u0026thinsp;=\u0026thinsp;susceptible to the diseases. The number of plants and ears per plot was counted. Ear aspect was scored on 1 to 9 scales, where 1\u0026thinsp;=\u0026thinsp;well-filled, uniform kernel arrangement, free from ear disease, and large or medium ears, while 9\u0026thinsp;=\u0026thinsp;ear disease, poorly filled, and shrivelled ears. The total number of ears per row was weighted (kg/plot area), and the moisture content of the representative sample grain was measured for each unshelled plot using a portable moisture tester. Grain yield (tone/ha) was calculated on the basis of ear weight per plot, adjusted to 12.5% moisture content and harvested plot size area.\u003c/p\u003e \u003cp\u003eAdjusted grain yield (tone/ha) = \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{FW\\left(100-MC\\right)*0.8*10}{87.5*PA}\\)\u003c/span\u003e\u003c/span\u003e\u003c/p\u003e \u003cp\u003eWhere; FW\u0026thinsp;=\u0026thinsp;Field weight of total ear weighted in kg/plot area, MC\u0026thinsp;=\u0026thinsp;moisture content of grain sample at harvest, 0.8\u0026thinsp;=\u0026thinsp;shelling percentage, 10\u0026thinsp;=\u0026thinsp;to convert quintal into tone, 87.5\u0026thinsp;=\u0026thinsp;100 minus from 12.5 standard moisture, PA\u0026thinsp;=\u0026thinsp;plot size area\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4. Statistical analysis\u003c/h2\u003e \u003cp\u003eIndividual site and combined analysis across locations were carried out using PROC GLM procedure in SAS version 9.3\u003csup\u003e20\u003c/sup\u003e. Genotype was considered as fixed effect whereas replication (location), block (replication), location \u0026times; genotype was considered as random effects. The significances of genotypes and interaction effects were tested and mean separation was done using least significant difference (LSD) at 0.05.\u003c/p\u003e \u003cp\u003e \u003cb\u003eGeneral combing ability analysis\u003c/b\u003e \u003c/p\u003e \u003cp\u003eDiallel data analysis procedure was carried out using analysis of genetic designs with R (AGD-R)\u003csup\u003e\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e\u003c/sup\u003e software. The general and specific combining abilities and variance components for crosses and important traits was esteemed using Griffing's method II and model I \u003csup\u003e\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eModel for multilocation diallel test\u003c/p\u003e \u003cp\u003eYijrl\u0026thinsp;=\u0026thinsp;\u0026micro;\u0026thinsp;+\u0026thinsp;r(l)\u0026thinsp;+\u0026thinsp;l\u0026thinsp;+\u0026thinsp;gi\u0026thinsp;+\u0026thinsp;gj\u0026thinsp;+\u0026thinsp;sij\u0026thinsp;+\u0026thinsp;l*gi\u0026thinsp;+\u0026thinsp;l*gj\u0026thinsp;+\u0026thinsp;l*sij\u0026thinsp;+\u0026thinsp;eijk, Where \u0026micro; is the grand mean, r is the replication effect, r(l)\u0026thinsp;=\u0026thinsp;replication nested under location (r\u0026thinsp;=\u0026thinsp;1..2), l\u0026thinsp;=\u0026thinsp;location(1..2), gi and gj are GCA effect (1\u0026hellip;12), sij is the SCA effect, l*gi and l*gj are location by GCA effect interactions, l*sij is location by SCA effect interaction and eijrl is the experimental pooled error effect\u003c/p\u003e \u003cp\u003e \u003cb\u003eEstimate Heterosis\u003c/b\u003e \u003c/p\u003e \u003cp\u003eHeterosis, mid parents as well as standard heterosis and their significant was calculated according to formula indicated in\u003csup\u003e\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e,\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e\u003c/sup\u003e\u003c/p\u003e \u003cp\u003eMPH=\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{\\left(F1-Mp\\right)\\text{*}100}{MP}\\)\u003c/span\u003e\u003c/span\u003e; MP= \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{\\left(P1+P2\\right)}{2}\\)\u003c/span\u003e\u003c/span\u003e, where, MPH= Mid Parent Heterosis, MP= is an average value of the corresponding parents for the hybrid, F1= hybrid value.\u003c/p\u003e \u003cp\u003eEconomic (standard heterosis, SH) or percent of heterosis, SH (%) = \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\frac{\\left(F1-SC\\right)\\text{*}100}{SC}\\)\u003c/span\u003e\u003c/span\u003e, where, SH= standard heterosis, SC= standard check, F1= mean performance of F1 (hybrid)\u003c/p\u003e \u003cp\u003eSignificance of each hybrid value with respect to the corresponding MP and SH values\u003c/p\u003e \u003cp\u003ewere tested using t test \u003csup\u003e\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003cspan class=\"InlineEquation\"\u003e \u003cspan class=\"mathinline\"\u003e\\(\\:tmp=\\frac{\\text{F}1-\\text{m}\\text{p}}{\\frac{\\sqrt{3MSE}}{2r}}\\)\u003c/span\u003e \u003c/span\u003e, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:SH\\left(t\\right)=\\frac{\\text{F}1-\\text{S}\\text{C}}{\\frac{\\sqrt{2\\text{M}\\text{S}\\text{E}}}{\\text{r}}}\\)\u003c/span\u003e\u003c/span\u003e, where, MSE= error mean square\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Result","content":"\u003cp\u003eThe ANOVA showed a highly significant difference (P\u0026thinsp;\u0026le;\u0026thinsp;0.001) among PVA hybrids for yield and most traits at each location, suggesting the presence of genetic variation among provitamin A hybrids performance under field conditions and feasibility selection of hybrids for grain yield and other traits. Furthermore, combined analysis of variance revealed a significant difference (P\u0026thinsp;\u0026le;\u0026thinsp;0.01) in mean square due to PVA hybrids interaction with locations for grain yield and other agronomic traits studied (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eMean performance of PVA hybrids\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe mean performance of PVA hybrids along with checks for grain yield, days to anthesis, days to silking and plant height is presented in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. The highest grain yield (7.95 t/ha) was recorded for cross L9/L12, whereas L4/L5 showed the lowest yield (2.45 t/ha) with a mean of 5.20 t/ha across locations. The mean values for traits across locations ranged from 69.66 to 84.33 for days to anthesis with a mean of 76.04, 71.50 to 87.16 for days to silking with a mean of 78.75 and plant height (cm) from 223.39 to 161.89 with a mean of 198.15 and 6.00 to 2.31 for GLS with a mean of 3.7. Seventeen PVA hybrids showed better performance for grain yield than checks; out of these, seven PVA hybrids were selected with a mean yield ranged from 7.11 t/ha to 7.95 t/ha and showed 18.69\u0026ndash;32.72% yield advantage over the better check (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). In addition, the selected PVA hybrids showed earlier flowering (male anthesis and silking days) than checks without yield penalty across locations.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eMean squares from ANOVA for yield and important agronomic traits of 68 PVA maize hybrids evaluated across three locations in 2022 Notice\u0026thinsp;=\u0026thinsp;SV\u0026thinsp;=\u0026thinsp;source of variation, DA\u0026thinsp;=\u0026thinsp;days to anthesis, DS\u0026thinsp;=\u0026thinsp;days to silking, Plht\u0026thinsp;=\u0026thinsp;plant height, Erht\u0026thinsp;=\u0026thinsp;ear height, GLS\u0026thinsp;=\u0026thinsp;grey leaf spot, TLB\u0026thinsp;=\u0026thinsp;turcicum leaf blight\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSV\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"7\" nameend=\"c9\" namest=\"c3\"\u003e \u003cp\u003eMean square of yield and agronomic traits\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYield\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePlht\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eErht\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eGLS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTLB\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLoc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e127.04\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3658.26\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3837.88\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1077881.64\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e100508.32\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e254.05\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e281.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRep\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e21.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e8.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e35.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e10.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e21.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e6.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.75\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBlk(rep)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e11.86\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e320.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e82.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2.19\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGenotype\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8.95\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e38.06\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e47.46\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e934.27\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e505.9\u003csup\u003e7***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e4.09\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e1.96\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRep(loc)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e13.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e20.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e44.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2291.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e628.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e6.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e3.59\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLoc*Genotype\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e134\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4.53\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e18.00\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e22.47\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e591.00\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e207.73\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e1.96\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e1.30\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eError\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e195\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e01.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e1.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.88\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSelected PVA hybrids for yield and other agronomic traits\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eentry\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHybrids\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYield advantage over best check (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eYield (t/ha)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eDS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePlant height\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL9 x L12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e32.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e75.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e78.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e219.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL9 x L10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e29.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e75.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e77.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e220.50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL10 x L12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e29.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e72.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e74.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e209.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL9 x L11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e27.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e77.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e80.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e198.62\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL5 x L11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e22.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e74.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e76.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e221.28\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL5 x L9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e19.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e75.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e78.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e207.72\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL8 x L12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e76.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e79.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e220.50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCheck, BHQPY545\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e81.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e84.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e201.11\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCheck, BH 549\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e84.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e87.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e206.72\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMean\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e76.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e78.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e198.52\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e24.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9.22\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLSD.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e20.82\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eAnalysis of variance: GCA for inbred lines and SCA for cross effects across locations\u003c/b\u003e \u003c/p\u003e \u003cp\u003eMean squares due to location, crosses, general combining ability (GCA), specific combining ability (SCA), location x crosses, GCA \u0026times; location, GCA x lines, SCA \u0026times; location, and SCA x crosse are presented in Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e. Significant mean squares due to GCA for lines and SCA for crosses indicate that both additive and non-additive gene actions control the inheritance of most traits studied. Furthermore, the mean square of SCA x locations and crosses x locations showed a significant difference (P\u0026thinsp;\u0026le;\u0026thinsp;0.001) in the combined ANOVA for all traits studied (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e), indicating that SCA and crosses were affected by environments. Except for GLS and TLB traits, the mean square due to GCA of lines x location was non-significant in the combined ANOVA, suggesting that the inheritance of PVA-enriched lines is controlled by additive gene action and/or is unlikely to be affected by environment interaction for studied traits.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSummary mean squares of combined ANOVA across environments for yield and important agronomic traits of 66 PVA crosses\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSV\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"9\" nameend=\"c11\" namest=\"c3\"\u003e \u003cp\u003eMean square of yield and agronomic traits\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYield\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePlht\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eErht\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eGLS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTLB\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eEA\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e73.70**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1138.00ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e144.05**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e141869.82*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e110454.00ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e106.75**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e141.89ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e29.78***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e3.59ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRep(loc)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.35*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21.05ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e23.75ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1208.96*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e434.97ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e29.73ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e17.65**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e8.14ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.62***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCrosses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.35***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e29.45***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e39.18***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2219.72***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e892.60***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e3.29***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.52**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.49**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e4.26*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGCA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18.98*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e84.76***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e125.89***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4781.88***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2196.67**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e7.78*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e9.51**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e6.20ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e6.54ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSCA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.28**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e20.26**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24.73ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1792.70*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e675.26ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2.55***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.37ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.04*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e3.81ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCrosses x Location\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.50***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e11.11***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e17.38***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e981.58***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e433.19***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.44ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.42ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e2.04**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e2.76***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGCA x Location\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.51ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e12.36ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e17.02ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e603.ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e124.70ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2.55*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e3.03**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.37ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e3.91ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSCA x Location\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.00***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10.90***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e17.44***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1044.52***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e484.61***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.25ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.18ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e1.82*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e2.57***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eError\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e150\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e8.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e340.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e157.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e1.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e1.28\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGCA: SCA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eGeneral combining ability effect of lines for grain yield and other traits\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe general combining ability (GCA) effect showed significant differences (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05 to 0.001) among PVA-enriched lines for yield and other traits (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). This suggests that at least one inbred line differs from other lines in terms of additive gene and additive x additive gene effects for agronomic traits studied.\u003c/p\u003e \u003cp\u003eThe GCA effects of lines for yield varied from \u0026minus;\u0026thinsp;0.85 to 0.99 t ha\u003csup\u003e-\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u003c/sup\u003e. Among all lines, L12, L8, L6, L10, and L2 showed desirable positive GCA effects on grain yield. In contrast, L3 had the lowest GCA effect (-0.85) for grain yield. L12 had the highest GCA effect for yield as well as a good combiner for grain yield, meanwhile it had a negative GCA effect for DA, DS, GLS, and TLB. Similarly, L8 was a good combiner for grain yield and other desirable agronomic traits. Thus, L12 (Tester B) and L8 (Tester A) can be used as alternative testers for further PVA maize breeding programs.\u003c/p\u003e \u003cp\u003e \u003cb\u003eSpecific Combining Ability effect of crosses for grain yield and other traits\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe SCA effects of crosses which indicates whether their performance is better or worse than what would be expected from the GCA effects of their respective parents (Table\u0026nbsp;\u003cspan refid=\"Tab7\" class=\"InternalRef\"\u003e7\u003c/span\u003e and Supplementary Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). The highest positive SCA effect for grain yield was recorded for crosses of L7/L12 (3.16 t/ha), followed by L7/L9 (2.43 t/ha). At the same time, these crosses had negative significant effects for DA and DS, suggesting the tendency of earliness. In contrast, L4 and L5 had the lowest cross-combinations for grain yield (-2.43 t/ha) with negative SCA effects.\u003c/p\u003e \u003cp\u003eFurthermore, crosses of L1 and L10 exhibited a significant positive effect of PA and EA. Similarly, crosses of L10 x L12, L5 x L9, and L8 x L12 showed a positive effect for plant and ear height and a significant negative SCA effect for flowering as well as for TLB and GLS, suggesting that these parental combinations could be good sources of hybrid development for resistance to disease and earliness.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEstimates of General Combining Ability (GCA) effects for grain yield and important agronomic traits of twelve PVA enriched maize lines evaluated in 2022.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eLines\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"9\" nameend=\"c10\" namest=\"c2\"\u003e \u003cp\u003eGrain yield and important agronomic traits\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGY\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePlht\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eErht\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eGLS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eTLB\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eEA\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.40***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-1.92***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-2.29***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.24ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.89ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.26*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.18ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.37***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.01ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.23*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.67*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-1.18***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.92ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.01*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.39***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.07ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.45***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.47***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.85***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.06**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.45***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-6.18*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-1.50ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.40***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.24ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.21*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.05ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e 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colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.41***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.65**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.54ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.80*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.86**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.23ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.22ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.06ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.01ns\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e 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colname=\"c1\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.38***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-1.28***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-1.34***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-8.01***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-5.65\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.69***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.80***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.33*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.44***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e 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align=\"left\" colname=\"c1\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.71***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.44ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.48ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-20.32***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-12.07\u003csup\u003e***\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.21ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.13ns\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.42**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.58***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e 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colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eL2\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eL3\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eL4\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eL5\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eL6\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eL7\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eL8\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eL9\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eL10\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e \u003cp\u003eL11\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c13\"\u003e \u003cp\u003eL12\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.16*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-1.13*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e-0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-0.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e0.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-0.16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.13*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.41**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-0.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e-1.67***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e1.20*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e0.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-0.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-0.46\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e1.96***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-2.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e0.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e0.14\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-2.43***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2.08***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e0.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e2.05***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e0.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e1.42***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e-0.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-0.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e2.00***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e1.67***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e1.07*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e-1.42***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e1.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e1.40***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e2.26*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e2.43***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e0.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e0.93\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e3.16**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-1.56**\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e1.97***\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-0.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e1.75**\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e 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colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e2.05***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-0.26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eHeterosis effect\u003c/b\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eMid-parent heterosis\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe values of mid-parent and standard heterosis were estimated for grain yield, DA, DS, and GLS across locations (supplementary Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e). Significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) to highly significant difference (P\u0026thinsp;\u0026lt;\u0026thinsp;0.01) positive mid-parent heterosis (MPH) estimate for grain yield was recorded for 20 crosses (supplementary Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e) which ranged from 99.30\u0026ndash;685.59%, suggesting that the crosses performed better than the mean performance of the mid-parent. Regarding flowering, thirty-one and fourteen crosses exhibited significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) negative mid-parent heterosis for DA and DS, respectively. L10 and L12 showed the highest mid-parent heterosis (685.59%), followed by L9 and L10 (509.59%) for grain yield. These crosses also showed significant negative MPH for flowering (DA and DS), indicating good yield performance with earliness.\u003c/p\u003e \u003cp\u003e \u003cb\u003eStandard Heterosis\u003c/b\u003e \u003c/p\u003e \u003cp\u003eOnly one cross (L9/L12) displayed significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) positive standard heterosis (34.52%) over BH 549 (normal maize hybrid check for grain yield), while eleven crosses exhibited significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) negative heterosis, ranging from \u0026minus;\u0026thinsp;58.84% to -34.01% for grain yield. On the other hand, none of the crosses showed significant positive standard heterosis, while nine crosses exhibited significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) negative heterosis for grain yield compared to BHQPY 545, a yellow quality protein maize check (Supplementary Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e), suggesting further improvement of PVA hybrids for grain yield. But it is to be noted that the cross L7/ L12 had the same yield (zero heterosis) compared to BHQPY 545 for grain yield.\u003c/p\u003e \u003cp\u003eRegarding days to anthesis and silking, most of the crosses showed significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) to highly significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.01) negative heterosis for days to tasselling and silking as compared to BH 549 and BHQPY 545 checks, which indicates the tendency of these crosses for earliness.\u003c/p\u003e \u003cp\u003eThe estimated standard heterosis for GLS in four crosses (L2 / L6, L4 / L5, L4 / L10, and L5 / L6) were positive and significant, ranging from 76.67\u0026ndash;100% over the standard checks BH549 and BHQPY545, indicating their likely susceptible to GLS when compared to the checks. On the other hand, some crosses showed negative standard heterosis for GLS, implying a tendency towards resistance to the disease. It was interesting that three crosses showed zero heterosis compared to checks for GLS recorded, suggesting that theses crosses and checks had similar GLS disease severity.\u003c/p\u003e"},{"header":"4. Discussion","content":"\u003cp\u003e \u003cb\u003ePerformance of maize hybrids, Combining Ability Effects and Tester Identification\u003c/b\u003e \u003c/p\u003e \u003cp\u003eOur result demonstrated that the significant mean square observed for provitamin A hybrids, locations, and locations x hybrids for grain yield and most other measured agronomic traits indicated the ranking of genotypes across environments at mid-altitude subhumid agro-ecology of Ethiopia. Maize hybrids with high grain yields were selected for (L9/LL12), (L10/LL12), (L9/LL10), (L9/LL11), L5/L11, L5/L9 and L8/L12 that showed 18.69\u0026ndash;32.72% yield advantages over checks as well as early flowering without yield penalty. Moreover, L10/L12, L5/L9, and L8/L12 had high yield potential with positive SCA effect and could be changed into three-way crosses with the opposite testers. The better performance of those hybrids in terms of yield and other traits might be due to the contribution of favorable alleles\u003csup\u003e\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u003c/sup\u003e and other associated genes in their parents during cross-combination. Thus, the selected PVA hybrids should be tested further, including their enriched provitamin A content and stability across environments\u003csup\u003e\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e\u003c/sup\u003e\u003c/p\u003e \u003cp\u003eThe current study revealed that the mean square due to GCA for lines and SCA for crosses showed a significant difference for all traits measured, indicating that both additive and non-additive gene actions control the inheritance of most traits studied\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e,\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e\u003c/sup\u003e, suggesting that improvement in these traits could be achieved through the selection of parents and their crosses for hybrid maize improvement. However, the ratio of SCA variance was higher than the GCA variance components for all traits studied, indicating that the predominance of non-additive gene action was more important in controlling the inheritance of traits, which differed from results of other studies\u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e,\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e, but it was consistent with\u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e,\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u003c/sup\u003e who reported the non-additive gene effect for governing grain yield and other traits. Such contradictory result reported on the mode of inheritance of traits highlighted that might be due to the type of germplasm used and the environmental condition under which the study carried out\u003csup\u003e\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e\u003c/sup\u003e. Thus, further testing is required on Pro-vitamin A enriched with germplasm having diverse genetic background across multiple environments.\u003c/p\u003e \u003cp\u003eThe GCA effect of lines is important for the selection of parent and hybrid development in the direction of yield improvements and other traits. In the current study, L2, L6, L8, L10, and L12 showed positive contributions to grain yield and other traits, suggesting the presence of additive and additive x additive gene action in the inheritance of studied traits\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e. Also, these inbred lines that are within heterotic groups can be used for further inbred line development and hybridization\u003csup\u003e\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAmong the lines, L8 and L12 showed the highest GCA effect for grain yield and seemed to have the best average performance for yield when crossed with other lines; had positive contribution to height and negative to GLS and TLB, suggesting the development of hybrid resistance to disease. Moreover, crosses of L8 and L12 exhibited one of the top average performances for hybrid yield and other traits. The choice of tester in maize hybrid breeding programs is determined by the capacity of the tester to discriminate among maize inbred lines, GCA effects for traits and average testcross performance for yield\u003csup\u003e\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e; \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u003c/sup\u003e. This result was consistent with the findings of other studies\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e; \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u003c/sup\u003e. Thus, L8 (Tester A) and L12 (Tester B) can be used as alternative opposite testers to evaluate new inbred lines and/or progenies across optimum conditions.\u003c/p\u003e \u003cp\u003eTwenty crosses showed yield performance ranging from 99.30\u0026ndash;685.59% compared to mid-parent heterosis. This result is in line with that of\u003csup\u003e\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e\u003c/sup\u003e who reported that the top crosses for grain yield ranged from 488 to 539% of mid-parent heterosis. Similarly, \u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e\u003c/sup\u003ewho reported cross hybrids that expressed significantly higher than mid-parent heterosis for grain yield and other traits. In this study, the top three crosses (L10 / L12 (685.59%), L9/LL10 (509.55%), and L9/LL11 (505.05%) supplementary Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e) showed the highest mid-parent heterosis for grain yield. Moreover, their single-cross combination was also found to be the top hybrid performance for yield which might be due to difference in the origins of parents, genetic divers and dominant gene action.\u003c/p\u003e \u003cp\u003eThe aim of maize breeders is to develop and select stable superior maize hybrids that are better than checks in terms of grain yield and other desirable traits. Our result showed that only one cross (L9 / L12) had significant standard heterosis over BH 549, a normal maize hybrid check, whereas there was a non-significant positive heterosis with BHQPY 545, a yellow quality protein maize hybrid check for grain yield. On the other hand, cross L7 and L12 had zero heterosis with BHQPY 545 for grain yield, indicating that this cross had the same yield as BHQPY 545. Thus, further PVA breeding is required with comprehensive allelic variants and genetic divergence of parents for crosses combination in heterosis manifestation for yield and other agronomic traits.\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eThe current study showed that F1 hybrids (L9 / L12), (L10/LL12), (L9/LL10), (L9/LL11), L9/L11, L5/L11, L5/L9, and L8/L12 exhibited better yield performance over checks and it need further testing across locations. Inbred lines L2, L6, L8, L10, and L12 had the highest GCA effects for grain yield, suggesting the transfer of favorable alleles to their progenies; thus, these inbred lines within the heterotic group could be used as new sources of PVA inbred line development. Out of these, L8 and L12 showed the highest GCA effect for grain yield, better average performance for yield when crossed with other lines, and negative contributions for GLS and TLB diseases. Thus, L8 (Tester A) and L12 (Tester B) can be used as alternative opposite testers to evaluate new PVA inbred lines and/or progenies across optimum conditions.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003ePlants materials involved in this research are used for scientific research. This article did not contain any studies with human participants or animals and did not involve any protected species.\u003c/p\u003e\n\u003ch2\u003e\u003cstrong\u003eCompeting interest\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eThe authors declare that no conflict of interest\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eBG: Writing \u0026ndash; original draft, Investigation, Data curation, Formal analysis, Software. GA: Writing \u0026ndash; editing draft, Methodology, Visualization and Project administration; KB: Supervision, Writing \u0026ndash; review \u0026amp; editing, AM: Writing \u0026ndash; review and editing, Project administration, Funding acquisition.\u003c/p\u003e\n\u003ch2\u003eAcknowledgment\u003c/h2\u003e\n\u003cp\u003eThe authors highly acknowledge the Harvest Plus Challenge Program for funding this work, particularly for carotenoid analyses. The author would like to thank EIAR/BNMRC for funding this activity, particularly for testing trials across locations. The authors also express appreciation to all BNMRC and EIAR collaboration staff who participated in planting and managing trials. The authors also gratefully acknowledge the help of anonymous editorials and reviewers for improving this article\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eall relevant collected data are included in this article. Additional information is presented in the Supplementary Tables S1 and S2. Furthermore, direct request to corresponding author on data available\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eRanum, P., Pe\u0026ntilde;a-Rosas, J. P. \u0026amp; Garcia-Casal, M. N. Global maize production, utilization, and consumption. \u003cem\u003eAnn. N. Y. Acad. Sci.\u003c/em\u003e \u003cb\u003e1312\u003c/b\u003e (1), 105\u0026ndash;112 (2014).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEkpa, O., Palacios-Rojas, N., Kruseman, G., Fogliano, V. \u0026amp; Linnemann, A. R. Sub-Saharan African maize-based foods: technological perspectives to increase the food and nutrition security impacts of maize breeding programme. \u003cem\u003eGlobal food Secur.\u003c/em\u003e \u003cb\u003e17\u003c/b\u003e, 48\u0026ndash;56 (2018).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGupta, H. S., Hossain, F. \u0026amp; Muthusam, V. 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Performance of testers with different genetic structure for evaluation of maize inbred lines. \u003cem\u003eCi\u0026ecirc;ncia Rural\u003c/em\u003e. \u003cb\u003e42\u003c/b\u003e, 770\u0026ndash;7769 (2012).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Combing ability, yield, heterosis, tester, inbred lines, maize","lastPublishedDoi":"10.21203/rs.3.rs-4839825/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4839825/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eCombining ability of enriched Pro-vitamin A yellow maize inbred lines can provide information for breeders to develop biofortified maize hybrids with high yield and desirable traits. Sixty-eight F1 hybrids along with two checks were arranged in alpha lattice design, replicated twice and evaluated across three locations at mid-altitude agro-ecology of Ethiopia. The combined analysis of variance revealed a significant difference (P\u0026thinsp;\u0026le;\u0026thinsp;0.01) in mean square due to PVA hybrids, environment and hybrid interaction with environments for grain yield and most of the agronomic traits and it indicates that genotypes performed differently across environments. Seven PVA hybrids were selected based on average yield performance ranging from 7.11 to 7.95 t/ha and 18.69\u0026ndash;29.04% yield advantage over the best check and these hybrids should be further evaluated across environments. The mean squares analysis of variance for GCA and SCA showed a significant difference (p\u0026thinsp;\u0026le;\u0026thinsp;0.001) for all traits studied, indicating that both additive and non-additive gene effects were important for controlling these traits. Significant positive SCA effect on grain yield was recorded for crosses of L7/L12 (3.16 t/ha), L7/L9 (2.43 t/ha) and L10 x L12 (2.05 t/ha). Only one cross (L9 x L12) displayed significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) positive standard heterosis (34.52%) over BH 549, whereas non-significant positive heterosis was detected over the BHQPY 545 check. Inbred lines L12, L8, L6, L10, and L2 had the highest positive and significant GCA effects on grain yield and other traits. Out of these lines, L8 and L12 were good combiners for most traits and will be used as alternative testers as they had the highest GCA effect for yield as well as other traits, and also their cross combination were among the top performing hybrids for yield. Our study highlighted that those inbred lines that had a high GCA effect within the heterotic group, could be used as a new source of PVA inbred line development. Furthermore, L8 and L12 can be used as alternative testers for future PVA maize breeding programs.\u003c/p\u003e","manuscriptTitle":"Combining Ability and Heterosis for Grain Yield and Other Agronomic Traits in Provitamin A Enriched Maize Inbred Lines","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-12-04 05:31:22","doi":"10.21203/rs.3.rs-4839825/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"a7a4cce7-74aa-493d-8898-fd4ce56e55b5","owner":[],"postedDate":"December 4th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":41046139,"name":"Biological sciences/Genetics"},{"id":41046140,"name":"Biological sciences/Plant sciences"}],"tags":[],"updatedAt":"2025-07-21T04:23:18+00:00","versionOfRecord":[],"versionCreatedAt":"2024-12-04 05:31:22","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4839825","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4839825","identity":"rs-4839825","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}