Genome-wide association mapping and genomic prediction of stalk rot in two mid-altitude tropical maize populations

Maize stalk rot reduces grain yield and quality.Information about the genetics of resistance to maize stalk rot could help breeders design effective breeding strategies for the trait.Genomic prediction may be a more effective breeding strategy for stalk-rot resistance than marker-assisted selection.We performed a genome-wide association study(GWAS)and genomic prediction of resistance in testcross hybrids of 677 inbred lines from the Tuxpe?o and non-Tuxpe?o heterotic pools grown in three environments and genotyped with 200,681 single-nucleotide polymorphisms(SNPs).Eighteen SNPs associated with stalk rot shared genomic regions with gene families previously associated with plant biotic and abiotic responses.More favorable SNP haplotypes traced to tropical than to temperate progenitors of the inbred lines.Incorporating genotype-by-environment(G×E)interaction increased genomic prediction accuracy.

Comparison of sequencing-based and array-based genotyping platforms for genomic prediction of maize hybrid performance

Genomic selection(GS)is a powerful tool for improving genetic gain in maize breeding.However,its routine application in large-scale breeding pipelines is limited by the high cost of genotyping platforms.Although sequencing-based and array-based genotyping platforms have been used for GS,few studies have compared prediction performance among platforms.In this study,we evaluated the predictabilities of four agronomic traits in 305 maize hybrids derived from 149 parental lines subjected to genotyping by sequencing(GBS),a 40K SNP array,and target sequence capture(TSC)using eight GS models.The GBS marker dataset yielded the highest predictabilities for all traits,followed by TSC and SNP array datasets.We investigated the effect of marker density and statistical models on predictability among genotyping platforms and found that 1K SNPs were sufficient to achieve comparable predictabilities to 10K and all SNPs,and BayesB,GBLUP,and RKHS performed well,while XGBoost performed poorly in most cases.We also selected significant SNP subsets using genome-wide association study(GWAS)analyses in three panels to predict hybrid performance.GWAS facilitated selecting effective SNP subsets for GS and thus reduced genotyping cost,but depended heavily on the GWAS panel.We conclude that there is still room for optimization of the existing SNP array,and using genotyping by target sequencing(GBTS)techniques to integrate a few functional markers identified by GWAS into the 1K SNP array holds great promise of being an effective strategy for developing desirable GS breeding arrays.

Heat-tolerant maize for rainfed hot,dry environments in the lowland tropics:From breeding to improved seed delivery

Climate change-induced heat stress combines two challenges:high day-and nighttime temperatures,and physiological water deficit due to demand-side drought caused by increase in vapor-pressure deficit.It is one of the major factors in low productivity of maize in rainfed stress-prone environments in South Asia,affecting a large population of smallholder farmers who depend on maize for their sustenance and livelihoods.The International Maize and Wheat Improvement Center(CIMMYT)maize program in Asia,in partnership with public-sector maize research institutes and private-sector seed companies in South Asian countries,is implementing an intensive initiative for developing and deploying heat-tolerant maize that combines high yield potential with resilience to heat and drought stresses.With the integration of novel breeding tools and methods,including genomics-assisted breeding,doubled haploidy,fieldbased precision phenotyping,and trait-based selection,new maize germplasm with increased tolerance to heat stress is being developed for the South Asian tropics.Over a decade of concerted effort has resulted in the successful development and release of 20 high-yielding heat-tolerant maize hybrids in CIMMYT genetic backgrounds.Via public–private partnerships,eight hybrids are presently being deployed on over 50,000 ha in South Asian countries,including Bangladesh,Bhutan,India,Nepal,and Pakistan.

Identification and validation of stable quantitative trait loci for yield component traits in wheat

Grain weight and grain number are important yield component traits in wheat and identification of underlying genetic loci is helpful for improving yield.Here,we identified eight stable quantitative trait loci(QTL)for yield component traits,including five loci for thousand grain weight(TGW)and three for grain number per spike(GNS)in a recombinant inbred line population derived from cross Yangxiaomai/Zhongyou 9507 across four environments.Since grain size is a major determinant of grain weight,we also mapped QTL for grain length(GL)and grain width(GW).QTGW.caas-2D,QTGW.caas-3B,QTGW.caas-5A and QTGW.caas-7A.2 for TGW co-located with those for grain size.QTGW.caas-2D also had a consistent genetic position with QGNS.caas-2D,suggesting that the pleiotropic locus is a modulator of trade-off effect between TGW and GNS.Sequencing and linkage mapping showed that TaGL3-5A and WAPO-A1 were candidate genes of QTGW.caas-5A and QTGW.caas-7A.2,respectively.We developed Kompetitive allele specific PCR(KASP)markers linked with the stable QTL for yield component traits and validated their genetic effects in a diverse panel of wheat cultivars from the Huang-Huai River Valley region.KASP-based genotyping analysis further revealed that the superior alleles of all stable QTL for TGW but not GNS were subject to positive selection,indicating that yield improvement in the region largely depends on increased TGW.Comparative analyses with previous studies showed that most of the QTL could be detected in different genetic backgrounds,and QTGW.caas-7A.1 is likely a new QTL.These findings provide not only valuable genetic information for yield improvement but also useful tools for marker-assisted selection.

QTL mapping for pre-harvest sprouting in a recombinant inbred line population of elite wheat varieties Zhongmai 578 and Jimai 22

Pre-harvest sprouting(PHS)is one of the serious global issues in wheat production.Identification of quantitative trait loci(QTL)and closely-linked markers is greatly helpful for wheat improvement.In the present study,a recombinant inbred line(RIL)population derived from the cross of Zhongmai 578(ZM578)/Jimai 22(JM22)and parents were phenotyped in five environments and genotyped by the wheat 50 K single-nucleotide polymorphism(SNP)array.Two QTL of germination index(GI),QGI.caas-3A and QGI.caas-5A,were detected,explaining 4.33%–5.58%and 4.43%–8.02%of the phenotypic variances,respectively.The resistant effect of QGI.caas-3A was contributed by JM22,whereas that of QGI.caas.5A was from ZM578.The two QTL did not correspond to any previously identified genes or genetic loci for PHSrelated traits according to their locations in the Chinese Spring reference genome,indicating that they are likely to be new loci for PHS resistance.Four kompetitive allele-specific PCR(KASP)markers K_AX-109605367and K_AX-179559687 flanking QGI.caas-3A,and K_AX-111258240 and K_AX-109402944flanking QGI.caas-5A,were developed and validated in a natural population of 100 wheat cultivars.The distribution frequency of resistance alleles at Qphs.caas-3A and Qphs.caas-5A loci were 82.7%and57.1%,respectively,in the natural population.These findings provide new QTL and tightly linked KASP markers for improvement of PHS resistance in wheat.

The wheat sucrose synthase gene TaSus1 is a determinant of grain number per spike

Some haplotypes of the sucrose synthase gene TaSus1 are associated with thousand-grain weight(TGW)in wheat(Triticum aestivum L.).However,no mutations have been identified within the gene to test this association.The effects of TaSus1 on grain number per spike(GNS)also are largely unknown.Our previous genome-wide association study identified TaSus-A1 as a candidate gene controlling fertile spikelet number per spike(FSN).In the present study,we generated two independent mutants for the three TaSus1 homoeologs by CRISPR/Cas9-mediated genome editing.The triple mutants displayed lower FSN,GNS,grain number per spikelet(GNST),and TGW than wild-type plants.In 306 hexaploid wheat accessions,two single-nucleotide polymorphisms in TaSus-A1 contributed differently to GNS.Introgression of the two alleles into a wheat genetic background confirmed their effects.The alleles differed in geographical distribution among the accessions.

Remote sensing of quality traits in cereal and arable production systems:A review

Cereal is an essential source of calories and protein for the global population.Accurately predicting cereal quality before harvest is highly desirable in order to optimise management for farmers,grading harvest and categorised storage for enterprises,future trading prices,and policy planning.The use of remote sensing data with extensive spatial coverage demonstrates some potential in predicting crop quality traits.Many studies have also proposed models and methods for predicting such traits based on multiplatform remote sensing data.In this paper,the key quality traits that are of interest to producers and consumers are introduced.The literature related to grain quality prediction was analyzed in detail,and a review was conducted on remote sensing platforms,commonly used methods,potential gaps,and future trends in crop quality prediction.This review recommends new research directions that go beyond the traditional methods and discusses grain quality retrieval and the associated challenges from the perspective of remote sensing data.

Genome-wide association mapping of vitamins B1 and B2 in common wheat

Vitamin B is essential for maintaining normal life activities in humans and animals who have to intake the microelement from the outside, especially from cereal products. In the present study 166 Chinese and foreign wheat cultivars planted in two environments were characterized for variation in vitamin B1 and B2 contents. A genome-wide association study(GWAS) using the wheat 90 K SNP assay identified 17 loci for vitamin B1 and 7 for vitamin B2 contents. Linear regression analysis showed a significantly positive correlation of the number of favorable alleles with vitamin B1 and B2 contents. Marker-trait associations(MTAs) at IWB43809(6AS, 0cM) and IWB69903(6AS, 13cM) were new and stable, and significantly associated with vitamin B1 content across two environments. The loci identified in this study and associated SNP markers could be used for improvement of vitamin B1 and B2 contents to obtain superior quality along with grain yield in wheat.

Three co-located resistance genes confer resistance to leaf rust and stripe rust in wheat variety Borlaug 100

Leaf rust(LR) and stripe rust(YR) are important diseases in wheat producing areas worldwide and cause severe yield losses under favorable environmental conditions when susceptible varieties are grown. We determined the genetic basis of resistance to LR and YR in variety Borlaug 100 by developing and phenotyping a population of 198 F6 recombinant inbred lines derived from a cross with the susceptible parent Apav#1. LR and YR phenotyping were conducted for 4 and 3 seasons, respectively, at CIMMYT research stations in Mexico under artificial epidemics. Mendelian segregation analyses indicated that 3–5 LR and 2 YR genes conferred resistance in Borlaug 100. Lr46/Yr29(1 BL), Yr17(2 AS) and Yr30(3 BS) were present in the resistant parent and segregated in the RIL population based on characterization by molecular markers linked to these genes. When present alone, Lr46/Yr29 caused average 13% and 16% reductions in LR and YR severities, respectively, in RILs. Similarly, Yr17 and Yr30 reduced YR severities by 57% and 11%, respectively. The Yr30 and the Yr17 translocation were also associated with 27% and 14% reductions, respectively, in LR severity, indicating that the 3 BS and 2 AS chromosomal regions likely carry new slow rusting LR resistance genes, temporarily designated as Lr B1 and Lr B2, respectively. Additive effects of Yr30*Yr17, Yr29*Yr17 and Yr29*Yr30 on YR and LR were significant and reduced YR severities by 56%,55%, and 45%, respectively, and LR severities by 34%, 40%, and 45%, respectively. Furthermore, interaction between the three genes was also significant, with mean reductions of 70% for YR and 54% for LR severities. Borlaug 100, or any one of the 21 lines with variable agronomic traits but carrying all three colocated resistance loci, can be used as resistance sources in wheat breeding programs.

Genome-wide association study and genomic prediction of Fusarium ear rot resistance in tropical maize germplasm

Fusarium ear rot(FER)is a destructive maize fungal disease worldwide.In this study,three tropical maize populations consisting of 874 inbred lines were used to perform genomewide association study(GWAS)and genomic prediction(GP)analyses of FER resistance.Broad phenotypic variation and high heritability for FER were observed,although it was highly influenced by large genotype-by-environment interactions.In the 874 inbred lines,GWAS with general linear model(GLM)identified 3034 single-nucleotide polymorphisms(SNPs)significantly associated with FER resistance at the P-value threshold of 1×10^(-5),the average phenotypic variation explained(PVE)by these associations was 3%with a range from 2.33%to 6.92%,and 49 of these associations had PVE values greater than 5%.The GWAS analysis with mixed linear model(MLM)identified 19 significantly associated SNPs at the P-value threshold of 1×10^(-4),the average PVE of these associations was 1.60%with a range from 1.39%to 2.04%.Within each of the three populations,the number of significantly associated SNPs identified by GLM and MLM ranged from 25 to 41,and from 5 to 22,respectively.Overlapping SNP associations across populations were rare.A few stable genomic regions conferring FER resistance were identified,which located in bins 3.04/05,7.02/04,9.00/01,9.04,9.06/07,and 10.03/04.The genomic regions in bins 9.00/01 and 9.04 are new.GP produced moderate accuracies with genome-wide markers,and relatively high accuracies with SNP associations detected from GWAS.Moderate prediction accuracies were observed when the training and validation sets were closely related.These results implied that FER resistance in maize is controlled by minor QTL with small effects,and highly influenced by the genetic background of the populations studied.Genomic selection(GS)by incorporating SNP associations detected from GWAS is a promising tool for improving FER resistance in maize.

Genomic prediction of the performance of hybrids and the combining abilities for line by tester trials in maize

The two most important activities in maize breeding are the development of inbred lines with high values of general combining ability(GCA)and specific combining ability(SCA),and the identification of hybrids with high yield potentials.Genomic selection(GS)is a promising genomic tool to perform selection on the untested breeding material based on the genomic estimated breeding values estimated from the genomic prediction(GP).In this study,GP analyses were carried out to estimate the performance of hybrids,GCA,and SCA for grain yield(GY)in three maize line-by-tester trials,where all the material was phenotyped in 10 to 11 multiple-location trials and genotyped with a mid-density molecular marker platform.Results showed that the prediction abilities for the performance of hybrids ranged from 0.59 to0.81 across all trials in the model including the additive effect of lines and testers.In the model including both additive and non-additive effects,the prediction abilities for the performance of hybrids were improved and ranged from 0.64 to 0.86 across all trials.The prediction abilities of the GCA for GY were low,ranging between-0.14 and 0.13 across all trials in the model including only inbred lines;the prediction abilities of the GCA for GY were improved and ranged from 0.49 to 0.55 across all trials in the model including both inbred lines and testers,while the prediction abilities of the SCA for GY were negative across all trials.The prediction abilities for GY between testers varied from-0.66 to 0.82;the performance of hybrids between testers is difficult to predict.GS offers the opportunity to predict the performance of new hybrids and the GCA of new inbred lines based on the molecular marker information,the total breeding cost could be reduced dramatically by phenotyping fewer multiple-location trials.

Identification of QTL for adult-plant resistance to powdery mildew in Chinese wheat landrace Pingyuan 50

Powdery mildew caused by Blumeria graminis f. sp. tritici is one of the major wheat diseases worldwide. The Chinese wheat landrace Pingyuan 50 has shown adult-plant resistance(APR)to powdery mildew in the field for over 60 years. To dissect the genetic basis of APR to powdery mildew in this cultivar, a mapping population of 137 double haploid(DH) lines derived from Pingyuan 50/Mingxian 169 was evaluated in replicated field trials for two years in Beijing(2009–2010 and 2010–2011) and one year in Anyang(2009–2010). A total of 540 polymorphic SSR markers were genotyped on the entire population for construction of a linkage map and QTL analysis. Three QTL were mapped on chromosomes 2BS(QPm.caas-2BS.2), 3BS(QPm.caas-3BS),and 5AL(QPm.caas-5AL) with the resistance alleles contributed by Pingyuan 50 explaining 5.3%,10.2%, and 9.1% of the phenotypic variances, respectively, and one QTL on chromosome 3BL(QPm.caas-3BL) derived from Mingxian 169 accounting for 18.1% of the phenotypic variance.QPm.caas-3BS, QPm.caas-3BL, and QPm.caas-5AL appear to be new powdery mildew APR loci.QPm.caas-2BS.2 and QPm.caas-5AL are possibly pleiotropic or closely linked resistance loci to stripe rust resistance QTL. Pingyuan 50 could be a potential genetic resource to facilitate breeding for improved APR to both powdery mildew and stripe rust.

Characterization of A-and B-type starch granules in Chinese wheat cultivars

Starch is the major component of wheat flour and serves as a multifunctional ingredient in food industry. The objective of the present study was to investigate starch granule size distribution of Chinese wheat cultivars, and to compare structure and functionality of starches in four leading cultivars Zhongmai 175, CA12092, Lunxuan 987, and Zhongyou 206. A wide variation in volume percentages of A- and B-type starch granules among genotypes was observed. Volume percentages of A- and B-type granules had ranges of 68.4–88.9% and 9.7–27.9% in the first cropping seasons, 74.1–90.1% and 7.2–25.3% in the second. Wheat cultivars with higher volume percentages of A- and B-type granules could serve as parents in breeding program for selecting high and low amylose wheat cultivars, respectively. In comparison with the B-type starch granules, the A-type granules starch showed difference in three aspects：（1） higher amount of ordered short-range structure and a lower relative crystallinity,（2） higher gelatinization onset（To） temperatures and enthalpies（ΔH）, and lower gelatinization conclusion temperatures（Tc）,（3） greater peak, though, and final viscosity, and lower breakdown viscosity and pasting temperature. It provides important information for breeders to develop potentially useful cultivars with particular functional properties of their starches suited to specific applications.

Characterization of TaCOMT genes associated with stem lignin content in common wheat and development of a gene-specific marker

Stem lignin content(SLC) in common wheat(Triticum aestivum L.) contributes to lodging resistance. Caffeic acid 3-O-methyltransferase(COMT) is a key enzyme involved in lignin biosynthesis. Characterization of TaCOMT genes and development of gene-specific markers could enable marker-assisted selection in wheat breeding. In the present study, the full-length genomic DNA(gDNA) sequences of TaCOMT genes located on chromosomes 3 A, 3 B, and 3 D were cloned by homologous cloning. Two allelic variants, TaCOMT-3 Ba and TaCOMT-3 Bb, were identified and differed by a 222-bp insertion/deletion(InDel) in the 3′-untranslated region(3′-UTR). A co-dominant gene-specific marker based on this InDel was developed and designated as Ta COMT-3 BM. A total of 157 wheat cultivars and advanced lines grown in four environments were used to validate the associations between allelic patterns and SLC. The SLC of cultivars with TaCOMT-3 Ba was significantly(P<0.01) higher than that of those with TaCOMT-3 Bb, and the marker TaCOMT-3 BM could be effectively used in wheat breeding.

Performance and yield stability of maize hybrids in stress-prone environments in eastern Africa

Identification and deployment of high-yielding and stress-tolerant maize hybrids adapted to stress-prone agro-ecologies is important for improving the food security and livelihoods of smallholder farmers in eastern Africa.The objectives of this study were to(i)assess the performance of maize hybrids under well-watered and drought stress conditions;(ii)evaluate grain yield stability of 65 intermediate-maturing and 55 early-maturing hybrids in 24 well-watered locations and seven drought stress locations;and(iii)identify representative and/or discriminative testing locations for increasing genetic gains for the target traits.There were significant differences for grain yield among early-and intermediatematuring hybrids tested under well-watered and drought stress environments.Among the early-maturing hybrids,the top 10 hybrids produced 46.8%–73.9%and 31.2%–42.1%higher mean grain yields than the best commercial check under drought and well-watered conditions,respectively.Among the intermediate-maturing hybrids,the top 10 hybrids produced 25.2%–47.7%and 8.5%–13.5%higher grain yield than commercial checks under drought stress and well-watered conditions,respectively,suggesting improvement in the levels of drought tolerance in both early-and intermediate-maturing hybrids.GGE biplot analysis and a bi-segmented regression linear method identified specific early-maturing and intermediate-maturing hybrids that performed well under both well-watered and drought stress conditions.These hybrids could be recommended for commercial production in eastern Africa.Kakamega in Kenya was found to be the most representative and highly discriminating site among well-watered testing locations,while Kabuku in Tanzania was the least representative of test locations.For testing under drought stress conditions,Kiboko in Kenya was identified as the most representative location.This information could be useful for allocating resources and streamlining CIMMYT maize hybrid testing in eastern Africa.

Genome-wide association mapping for grain shape and color traits in Ethiopian durum wheat(Triticum turgidum ssp.durum)

Grain shape and color strongly influence yield and quality of durumwheat.Identifying QTL for these traits is essential for transferring favorable alleles based on selection strategies and breeding objectives.In the present study,192 Ethiopian durum wheat accessions comprising 167 landraces and 25 cultivars were genotyped with a high-density Illumina iSelect 90K singlenucleotide polymorphism(SNP)wheat array to conduct a genome-wide association analysis for grain width(GW),grain length(GL),CIE(Commission Internationale l'Eclairage)L*(brightness),CIE a*(redness),and CIE b*(yellowness)traits.The accessions were planted at Sinana Agricultural Research Center,Ethiopia in the 2015/2016 cropping season in a complete randomized block design with three replications.Twenty homogeneous and healthy seeds per replicate were used for trait measurement.Digital image analysis of seeds with GrainScan software package was used to generate the phenotypic data.Analysis of variance revealed highly significant differences between accessions for all traits.Atotal of 46 quantitative trait loci(QTL)were identified for all traits across all chromosomes.One novelmajor candidate QTL(−lg P≥4)with pleiotropic effects for grain CIE L*(brightness)and CIE a*(redness)was identified on the long armof chromosome 2A.Eighteen nominal QTL(−lg P≥3)and 26 suggestive QTL(−lg P≥2.5)were identified.Pleiotropic QTL influencing both grain shape and color were identified.

QTL mapping of adult plant resistance to stripe rust and leaf rust in a Fuyu 3/Zhengzhou 5389 wheat population

Stripe or yellow rust(YR)and leaf rust(LR)cause large losses in wheat production worldwide.Resistant cultivars curtail the levels of losses.The present study aimed to identify quantitative trait loci(QTL)for YR and LR resistance in 147 F2:6 recombinant inbred lines(RIL)derived from the cross Fuyu 3/Zhengzhou 5389.The RIL population and parents were genotyped with the Wheat55 K single nucleotide polymorphism(SNP)array and simple sequence repeat(SSR)markers.All materials were also phenotyped for YR severity at Mianyang in Sichuan province and Baoding in Hebei province in the 2015/2016,2016/2017,and 2017/2018 cropping seasons,and LR severity at Zhoukou in Henan province and at Baoding in 2017/2018.Eleven QTL for YR resistance and five for LR resistance were detected using inclusive composite interval mapping(Ici Mapping).Four of these QTL on chromosomes 1 BL,2 BS,3 AL,and 5 AL conferred resistance to both YR and LR.The QTL on 1 BL was Lr46/Yr29,and that on 7 BL might be Lr68.The QTL on chromosome 2 BS was detected at a similar position to previously detected loci.QYr.hebau-3 AL/QLr.hebau-3 AL,QYr.hebau-5 AL/QLr.hebau-5 AL,QYr.hebau-7 DL,QYr.hebau-4 BS,QYr.hebau-6 DL,and QYr.hebau-2 AS are likely to be new.An SSR marker for QYr.hebau-7 DL was developed and validated in a diverse wheat panel from China,suggesting effectiveness in different genetic backgrounds.These QTL with closely linked SNP and SSR markers could be useful for marker-assisted selection in wheat breeding programs targeting durable resistance to both diseases.

A wheat chromosome 5AL region confers seedling resistance to both tan spot and Septoria nodorum blotch in two mapping populations

Tan spot(TS) and Septoria nodorum blotch(SNB), caused by Pyrenophora tritici-repentis and Parastagonospora nodorum, respectively, are important fungal leaf-spotting diseases of wheat that cause significant losses in grain yield. In this study, two recombinant inbred line populations, ‘Bartai’ × ‘Ciano T79’(referred to as B × C) and ‘Cascabel’ × ‘Ciano T79’(C × C) were tested for TS and SNB response in order to determine the genetic basis of seedling resistance. Genotyping was performed with the DAr Tseq genotypingby-sequencing(GBS) platform. A chromosome region on 5 AL conferred resistance to TS and SNB in both populations, but the effects were larger in B × C(R^2= 11.2%–16.8%) than in C × C(R^2= 2.5%–9.7%). Additionally, the chromosome region on 5BL(presumably Tsn1)was significant for both TS and SNB in B × C but not in C × C. Quantitative trait loci(QTL)with minor effects were identified on chromosomes 1B, 2A, 2B, 3A, 3B, 4D, 5A, 5B, 5D, 6B,and 6D. The two CIMMYT breeding lines ‘Bartai’ and ‘Cascabel’ contributed resistance alleles at both 5AL and 5BL QTL mentioned above. The QTL on 5AL showed linkage with the Vrn-A1 locus, whereas the vrn-A1 allele conferring lateness was associated with resistance to TS and SNB.

Genetic gains in wheat in Turkey: Winter wheat for dryland conditions

Wheat breeders in Turkey have been developing new varieties since the 1920 s, but few studies have evaluated the rates of genetic improvement. This study determined wheat genetic gains by evaluating 22 winter/facultative varieties released for rainfed conditions between 1931 and2006. The study was conducted at three locations in Turkey during 2008-2012, with a total of 21 test sites. The experimental design was a randomized complete block with four replicates in2008 and 2009 and three replicates in 2010-2012. Regression analysis was conducted to determine genetic progress over time. Mean yield across all 21 locations was 3.34 t ha^(-1) but varied from 1.11 ha^(-1) to 6.02 t ha^(-1) and was highly affected by moisture stress. Annual genetic gain was 0.50% compared to Ak-702, or 0.30% compared to the first modern landmark varieties. The genetic gains in drought-affected sites were 0.75% compared to Ak-702 and0.66% compared to the landmark varieties. Modem varieties had both improved yield potential and tolerance to moisture stress. Rht genes and rye translocations were largely absent in the varieties studied. The number of spikes per unit area decreased by 10% over the study period,but grains spike^(-1) and 1000-kemel weight increased by 10%. There were no significant increases in harvest index, grain size, or spike fertility, and no significant decrease in quality over time. Future use of Rht genes and rye translocations in breeding programs may increase yield under rainfed conditions.