Allozyme frequencies, heterozygosity and genetic distances following S1 recurrent selection in two synthetic maize populations

 Three cycles of S₁ recurrent selection for yield were carried out in two synthetic maize populations, EPS6 from humid Spain and EPS7 from arid Spain. One-hundred S₁ lines were evaluated from each cycle of selection and the ten highest-yielding S₁ lines were recombined to produce the next cycle. Changes in variability and genetic distances in two synthetic maize populations, following three cycles of recurrent selection, recombining ten S₁ lines in each cycle, were determined. Isozyme analysis was performed on 125 seedlings per cycle of selection (four cycles in each of two populations). Regressions of each allozyme frequency on cycles of selection were performed, genetic distances between populations were determined, and simple correlations between genetic distances and heterosis were calculated. The average heterozygosity per locus was also calculated for each population. Regression analysis did not reveal any common trend between EPS6 and EPS7 for changes in allele frequencies presumably due to selection. The number of polymorphic loci, the mean alleles per locus, and the mean heterozygosity did not show any reduction in variability. Finally, selection did not affect genetic distances among cycles of selection. The agronomic evaluation of the selection program, after three cycles of selection, revealed that the genetic variance was not significantly reduced for most traits, and that the heterosis among cycles of selection of both populations had not changed significantly. The conclusions based on isozyme data supported the deductions made from agronomic data. Three cycles of selection neither caused relevant changes on variability nor on genetic distance among cycles of selection of both maize synthetic populations. These data did not indicate any basis for increasing the number of S₁ lines recombined for recurrent selection. Received: 28 April 1997 / Accepted: 23 June 1997     

Use of SSRs for establishing heterotic groups in subtropical maize

Heterotic groups and patterns are of fundamental importance in hybrid breeding. The objectives of our research were to: (1) investigate the relationship of simple sequence repeats (SSR) based genetic distances between populations and panmictic midparent heterosis (PMPH) in a broad range of CIMMYT maize germplasm, (2) evaluate the usefulness of SSR markers for defining heterotic groups and patterns in subtropical germplasm, and (3) examine applications of SSR markers for broadening heterotic groups by systematic introgression of other germplasm. Published data of two diallels and one factorial evaluated for grain yield were re-analyzed to calculate the PMPH in population hybrids. Additionally, 20 pools and populations widely used in CIMMYT's breeding program were assayed with 83 SSR markers covering the entire maize genome. Correlations of squared modified Roger's distance (MRD²) and PMPH were mostly positive and significant, but adaption problems caused deviations in some cases. For intermediate- and early-maturity subtropical germplasm, two heterotic groups could be suggested consisting of a flint and dent composite. We concluded that the relationships between the populations obtained by SSR analyses are in excellent agreement with pedigree information. SSR markers are a valuable complementation to field trials for identifying heterotic groups and can be used to introgress exotic germplasm systematically.Communicated by F. Salamini     

An experimental comparison of selection alternatives to plateaued response

Three alternative selection methods for extending selection limits or breaking response plateaus were compared over ten generations in a replicated model experiment using two unrelated populations of Drosophila melanogaster that no longer responded to purebred selection for high egg number, a heterotic polygenic trait. The three methods were: (1) reciprocal recurrent selection (RRS) with selection within each of the plateaued populations based solely on crossbred performance, (2) a modification of reciprocal recurrent selection (MRRS) with selection within each population based on both purebred and crossbred performance, and (3) purebred selection within a new synthetic population formed by crossing the two plateaued populations.--Conflicting estimates were obtained for heritability of purebred egg number in each of the plateaued populations. The realized heritability values and estimates from diallel analyses indicated an absence of additive genetic variation for both populations; however, estimates from conventional intraclass correlation methods were positive. The diallel analyses revealed significant amounts of nonadditive gene effects for purebred egg number in each population, while the significant gene effects for crossbred egg numbers were additive. Estimates of the genetic correlation between purebred and crossbred egg number were negative (-0.85 +/- 0.68 and -0.32 +/- 0.25) for the two base populations.--All three alternatives to continued purebred selection gave significant responses, with the average gain per generation from MRRS being significantly superior to the other two methods. Observed purebred and crossbred responses under RRS were in agreement with quantitative genetic theory. Such was not the case for MRRS, which suggested the possibility of major gene segregation.--Evidence supporting a negative genetic correlation between purebred and crossbred performance and the possibility of overdominance is presented and discussed.     

Influence of genetic background and heterozygosity on meiotic recombination in Arabidopsis thaliana.

Plant breeding relies on genetic variability generated by meiotic recombination. Control of recombination frequencies is not yet possible, but would significantly extend the options for plant-breeding strategies. A prerequisite would be variability of recombination frequencies. In this study, 15 transgenic kanamycin (KR) and hygromycin (HR) resistance gene insertions mapping to the five Arabidopsis thaliana chromosomes were used as genetic markers. Recombination frequencies were determined from the frequencies of resistance phenotypes within populations segregating for linked KR and HR markers. Recombination frequencies of marker pairs were compared among these four ecotypes, among F1s in both reciprocal forms derived from these ecotypes, and between F1s and their parent lines. On average, the recombination frequencies in F1 crosses were substantially higher (up to 2-fold) than in the homozygous parental ecotypes. A strong negative correlation between genetic similarities of ecotypes and recombination frequencies was detected for two adjacent marker pairs located on the long arm of chromosome 3, but not for marker pairs in other genomic regions. Our results suggest that heterozygosity influences recombination in plant breeding, and cannot be ignored in genetic mapping of genomes.     

The Genomic Impacts of Drift and Selection for Hybrid Performance in Maize

Although maize is naturally an outcrossing organism, modern breeding utilizes highly inbred lines in controlled crosses to produce hybrids. The U.S. Department of Agriculture’s reciprocal recurrent selection experiment between the Iowa Stiff Stalk Synthetic (BSSS) and the Iowa Corn Borer Synthetic No. 1 (BSCB1) populations represents one of the longest running experiments to understand the response to selection for hybrid performance. To investigate the genomic impact of this selection program, we genotyped the progenitor lines and >600 individuals across multiple cycles of selection using a genome-wide panel of ∼40,000 SNPs. We confirmed previous results showing a steady temporal decrease in genetic diversity within populations and a corresponding increase in differentiation between populations. Thanks to detailed historical information on experimental design, we were able to perform extensive simulations using founder haplotypes to replicate the experiment in the absence of selection. These simulations demonstrate that while most of the observed reduction in genetic diversity can be attributed to genetic drift, heterozygosity in each population has fallen more than expected. We then took advantage of our high-density genotype data to identify extensive regions of haplotype fixation and trace haplotype ancestry to single founder inbred lines. The vast majority of regions showing such evidence of selection differ between the two populations, providing evidence for the dominance model of heterosis. We discuss how this pattern is likely to occur during selection for hybrid performance and how it poses challenges for dissecting the impacts of modern breeding and selection on the maize genome.     

Linkage disequilibrium in two European F<Subscript>2</Subscript> flint maize populations under modified recurrent full-sib selection

According to quantitative genetic theory, linkage disequilibrium (LD) can hamper the short- and long-term selection response in recurrent selection (RS) programs. We analyzed LD in two European flint maize populations, KW1265 x D146 (A x B) and D145 x KW1292 (C x D), under modified recurrent full-sib selection. Our objectives were to investigate (1) the decay of initial parental LD present in F(2) populations by three generations of intermating, (2) the generation of new LD in four (A x B) and seven (C x D) selection cycles, and (3) the relationship between LD changes and estimates of the additive genetic variance. We analyzed the F(2) and the intermated populations as well as all selection cycles with 104 (A x B) and 101 (C x D) simple sequence repeat (SSR) markers with a uniform coverage of the entire maize genome. The LD coefficient D and the composite LD measure Delta were estimated and significance tests for LD were performed. LD was reduced by intermating as expected from theory. A directional generation of negative LD between favorable alleles could not be observed during the selection cycles. However, considerable undirectional changes in D were observed, which we attributed to genetic sampling due to the finite population size used for recombination. Consequently, a long-term reduction of the additive genetic variance due to negative LD was not observed. Our experimental results support the hypothesis that in practical RS programs with maize, LD generated by selection is not a limiting factor for obtaining a high selection response.     

利用SSR标记分析玉米轮回选择群体的遗传多样性

利用SSR标记技术分析了玉米基础群体DC0及其选择两轮后的群体HSC2和MSC2以及基础群体XFC0和其选择一轮后的群体XFC1的遗传多样性。结果表明：49对引物在5个玉米群体中共扩增出185个等位位点,每个SSR座位的等位基因数目为1～7个,平均为3．8个;基础群体多态性位点总数和多态性位点比例比其改良群体的略高;DC0、HSC2和MSC2 3个群体的基因平均杂合度相似,XFC0与XFC1的基因平均杂合度相似;基础群体与改良群体的平均遗传距离也相似;累加各引物扩增的基因型种类,改良群体的基因型种类偏少,但这些差异均不显著。以上结果说明轮回选择的基础群体与其改良后群体的遗传变异相似,轮回选择可以保持群体的遗传变异范围,改变群体的遗传组成,增加群体内个体间的异质性。     

Association mapping in an elite maize breeding population

Association mapping (AM) is a powerful approach to dissect the genetic architecture of quantitative traits. The main goal of our study was to empirically compare several statistical methods of AM using data of an elite maize breeding program with respect to QTL detection power and possibility to correct for population stratification. These models were based on the inclusion of cofactors (Model A), cofactors and population effect (Model B), and SNP effects nested within populations (Model C). A total of 930 testcross progenies of an elite maize breeding population were field-evaluated for grain yield and grain moisture in multi-location trials and fingerprinted with 425 SNP markers. For grain yield, population stratification was effectively controlled by Model A. For grain moisture with a high ratio of variance among versus within populations, Model B should be applied in order to avoid potential false positives. Model C revealed large differences among allele substitution effects for trait-associated SNPs across multiple plant breeding populations. This heterogeneous SNP allele substitution effects have a severe impact for genomic selection studies, where SNP effects are often assumed to be independent of the genetic background.     

Evaluation of marker-assisted selection through computer simulation

Computer simulation was used to evaluate responses to marker-assisted selection (MAS) and to compare MAS responses with those typical of phenotypic recurrent selection (PRS) in an allogamous annual crop species such as maize (Zea mays L.). Relative to PRS, MAS produced rapid responses early in the selection process; however, the rate of these responses diminished greatly within three to five cycles. The gains from MAS ranged from 44.7 to 99.5% of the maximum potential, depending on the genetic model considered. Linkage distance between markers and quantitative trait loci (QTLs) was the factor which most limited the responses from MAS. When averaged across all models considered, flanking QTLs within two marker loci produced 38% more gain than did selection based on single markers if markers were loosely-linked to a QTL (20% recombination). Flanking markers were much less advantageous when markers were closely-linked to a QTL (5% recombination), producing an advantage over single markers of only 11%. Markers were most effective in fully exploiting the genetic potential when fewer QTLs controlled the trait. Large QTL numbers exacerbated the problem of marker-QTL recombination by requiring more generations for fixation. In annual crop species, MAS may offer a primary advantage of enabling two selection cycles per year versus the 2 years per cycle required by most PRS schemes for the evaluation of testcross progeny. MAS thus appears to allow very rapid gains for the first 2–3 years of recurrent selection, after which time conventional methods might replace MAS to achieve further responses.Publication number 19, 330 of the Minnesota Agricultural Experiment Station     

Predicted and realized grain yield responses to full-sib family selection in CIMMYT maize (Zea mays L.) populations

Summary The maize (Zea mays L.) improvement program of the International Maize and Wheat Improvement Center (CIMMYT) develops broad-based maize populations and, until recently, improved all of them through full-sib family selection with international testing. The purpose of this study was to estimate the genetic and genetic × environment variance components for ten of those populations and to measure expected yield improvement from full-sib selection. Mean yield ranged from 3.35–6.81 t ha^–1. For five populations the average yield in the last cycle was higher than in the initial cycles. Several populations showed no improvement or yielded less in the final cycle of selection, either because selection intensity was low or because strong selection pressure was applied simultaneously for several traits. Variation resulting from differences among family means within cycles and from interaction between families and locations within cycles were significant in all populations and cycles. Results indicate that variability among full-sib families was maintained throughout the cycles for all populations. The large _ge ² / _g ² ratio shown by most populations suggests that yield response per cycle could be maximized if the environments in which progenies are tested were subdivided and classified into similar subsets. The proportion of the predicted response realized in improved yield varied for each population.Journal Paper No. 8640, Nebraska Agric. Res. Div. Project No. 12-159. Research was supported in part by USAID/USDA/ CSRS Research Grant No. 86-CRSR-2-2789     

High genetic variability of herbivore-induced volatile emission within a broad range of maize inbred lines

Maize plants (Zea mays) attacked by caterpillars release a mixture of odorous compounds that attract parasitic wasps, natural enemies of the herbivores. We assessed the genetic variability of these induced volatile emissions among 31 maize inbred lines representing a broad range of genetic diversity used by breeders in Europe and North America. Odors were collected from young plants that had been induced by injecting them with caterpillar regurgitant. Significant variation among lines was found for all 23 volatile compounds included in the analysis: the lines differed enormously in the total amount of volatiles emitted and showed highly variable odor profiles distinctive of each genotype. Principal component analysis performed on the relative quantities of particular compounds within the blend revealed clusters of highly correlated volatiles, which may share common metabolic pathways. European and American lines belonging to established heterotic groups were loosely separated from each other, with the most clear-cut difference in the typical release of (E)-beta-caryophyllene by European lines. There was no correlation between the distances among the lines based on their odor profiles and their respective genetic distances previously assessed by neutral RFLP markers. This most comprehensive study to date on intraspecific variation in induced odor emission by maize plants provides a further example of the remarkably high genetic diversity conserved within this important crop plant. A better understanding of the genetic control of induced odor emissions may help in the development of maize varieties particularly attractive to parasitoids and other biological control agents and perhaps more repellent for herbivores.     

Linkage disequilibrium in two European F<Subscript>2</Subscript> flint maize populations under modified recurrent full-sib selection

According to quantitative genetic theory, linkage disequilibrium (LD) can hamper the short- and long-term selection response in recurrent selection (RS) programs. We analyzed LD in two European flint maize populations, KW1265 × D146 (A × B) and D145 × KW1292 (C × D), under modified recurrent full-sib selection. Our objectives were to investigate (1) the decay of initial parental LD present in F₂ populations by three generations of intermating, (2) the generation of new LD in four (A × B) and seven (C × D) selection cycles, and (3) the relationship between LD changes and estimates of the additive genetic variance. We analyzed the F₂ and the intermated populations as well as all selection cycles with 104 (A × B) and 101 (C × D) simple sequence repeat (SSR) markers with a uniform coverage of the entire maize genome. The LD coefficient D and the composite LD measure Δ were estimated and significance tests for LD were performed. LD was reduced by intermating as expected from theory. A directional generation of negative LD between favorable alleles could not be observed during the selection cycles. However, considerable undirectional changes in D were observed, which we attributed to genetic sampling due to the finite population size used for recombination. Consequently, a long-term reduction of the additive genetic variance due to negative LD was not observed. Our experimental results support the hypothesis that in practical RS programs with maize, LD generated by selection is not a limiting factor for obtaining a high selection response.     

Relationships among maize inbred lines and populations from European and North-American origins as estimated using RFLP markers

RFLP markers have proven to be a reliable and highly informative tool for characterizing genetic diversity in maize. Joint analysis of inbred lines and populations should provide valuable information with respect to (1) a better understanding of the genetic basis of present elite germplasm and (2) the identification of populations that may prove to be useful sources of genetic diversity for breeding programs. Sixty-two inbred lines of known heterotic groups and ten maize populations, some of them significant contributors to the genetic basis of the heterotic groups, were assayed at 28 RFLP loci. Joint data analyses first underlined that the populations displayed a large number of alleles that were absent in the set of inbred lines. Associations among inbreds and populations further proved consistent with pedigree data of the inbreds and provided new information on the genetical basis of heterotic groups. In particular, European flint inbreds were revealed to be as close to the Northeastern U.S. flint population studied as to the typical European populations. These results advocate the analysis of larger sets of populations by means of molecular markers in order to (1) gain insight into the history of maize germplasm and (2) set up appropriate strategies for the use of genetic resources in breeding programs. Received: 23 February 1998 / Accepted: 5 February 1999     

Genetic structure of wild bean populations in their South-Andean centre of origin

 The genetic structure of wild common bean populations was studied in the South-Andean centre of origin of the species. Plants were collected from 21 populations in Argentina and genetic variability was assessed for molecular and resistance markers. Polymorphism was weak for phaseolin, the major seed-storage protein, and for RAPD markers, while a high level of polymorphism was observed for resistance to anthracnose, one of the most important diseases of common bean. For the three traits, within-population variability was important and represented between 43.6% and 67.5% of the total variation. Although among-population differentiation was significant for all the traits, no correlation was found between the population distances calculated from RAPDs and resistance. These results indicate that pathogen selection pressure may be an important factor influencing the distribution of variability within and among host plant populations. Received: 28 October 1997 / Accepted: 25 November 1997     

Genome-Wide High-Resolution Mapping by Recurrent Intermating Using Arabidopsis Thaliana as a Model

We demonstrate a method for developing populations suitable for genome-wide high-resolution genetic linkage mapping, by recurrent intermating among F(2) individuals derived from crosses between homozygous parents. Comparison of intermated progenies to F(2) and ``recombinant inbred' (RI) populations from the same pedigree corroborate theoretical expectations that progenies intermated for four generations harbor about threefold more information for estimating recombination fraction between closely linked markers than either RI-selfed or F(2) individuals (which are, in fact, equivalent in this regard). Although intermated populations are heterozygous, homozygous ``intermated recombinant inbred' (IRI) populations can readily be generated, combining additional information afforded by intermating with the permanence of RI populations. Intermated populations permit fine-mapping of genetic markers throughout a genome, helping to bridge the gap between genetic map resolution and the DNA-carrying capacity of modern cloning vectors, thus facilitating merger of genetic and physical maps. Intermating can also facilitate high-resolution mapping of genes and QTLs, accelerating map-based cloning. Finally, intermated populations will facilitate investigation of other fundamental genetic questions requiring a genome-wide high-resolution analysis, such as comparative mapping of distantly related species, and the genetic basis of heterosis.     

Potential causes of linkage disequilibrium in a European maize breeding program investigated with computer simulations

Knowledge about the forces generating and conserving linkage disequilibrium (LD) is important for drawing conclusions about the prospects and limitations of association mapping. The objectives of our research were to examine the importance of (1) selection, (2) mutation, and (3) genetic drift for generating LD in a typical maize breeding program. We conducted computer simulations based on genotypic data of Central European maize open-pollinated varieties which have played an important role as founders of the European flint heterotic group. The breeding scheme and the dimensioning underlying our simulations reflect essentially the maize breeding program of the University of Hohenheim. Results suggested that in a plant breeding program of the examined dimension and breeding scheme, genetic drift and selection are major forces generating LD. The currently used population-based association mapping tests do not explicitly correct for LD caused by these two forces. Therefore, increased type I error rates are expected if these tests are applied to plant breeding populations. As a consequence, we recommend to use family-based association tests for association mapping approaches in plant breeding populations.     

The relationship between parental genetic or phenotypic divergence and progeny variation in the maize nested association mapping population

Appropriate selection of parents for the development of mapping populations is pivotal to maximizing the power of quantitative trait loci detection. Trait genotypic variation within a family is indicative of the family's informativeness for genetic studies. Accurate prediction of the most useful parental combinations within a species would help guide quantitative genetics studies. We tested the reliability of genotypic and phenotypic distance estimators between pairs of maize inbred lines to predict genotypic variation for quantitative traits within families derived from biparental crosses. We developed 25 families composed of ~200 random recombinant inbred lines each from crosses between a common reference parent inbred, B73, and 25 diverse maize inbreds. Parents and families were evaluated for 19 quantitative traits across up to 11 environments. Genetic distances (GDs) among parents were estimated with 44 simple sequence repeat and 2303 single-nucleotide polymorphism markers. GDs among parents had no predictive value for progeny variation, which is most likely due to the choice of neutral markers. In contrast, we observed for about half of the traits measured a positive correlation between phenotypic parental distances and within-family genetic variance estimates. Consequently, the choice of promising segregating populations can be based on selecting phenotypically diverse parents. These results are congruent with models of genetic architecture that posit numerous genes affecting quantitative traits, each segregating for allelic series, with dispersal of allelic effects across diverse genetic material. This architecture, common to many quantitative traits in maize, limits the predictive value of parental genotypic or phenotypic values on progeny variance.     

Genetic drift and selection effects of modified recurrent full-sib selection programs in two F2 populations of European flint maize

Selection response of a modified recurrent full-sib (FS) selection scheme conducted in two European flint F₂ maize (Zea mays L.) populations was re-evaluated. Our objectives were to (1) determine the selection response for per se and testcross performance in both populations and (2) separate genetic effects due to selection from those due to random genetic drift. Modified recurrent FS selection was conducted at three locations using an effective population size N _e = 32 and a selection rate of 25% for a selection index, based on grain yield and grain moisture. Recombination was performed according to a pseudo-factorial mating scheme. Selection response was assessed using a population diallel including the source population and advanced selection cycles, as well as testcrosses with unrelatesd inbred line testers and the parental F₁ generation. Selection response per cycle was significant for grain yield and grain moisture in both populations. Effects of random genetic drift caused only a small reduction in the selection response. No significant selection response was observed for testcrosses, suggesting that for heterotic traits, such as grain yield, a high frequency of favorable alleles in the elite tester masked the effects of genes segregating in the populations. We conclude that our modified recurrent FS selection is an alternative to other commonly applied intrapopulation recurrent selection schemes, and some of its features may also be useful for increasing the efficiency of interpopulation recurrent selection programs.     

Genomic location of the Fw gene for resistance to Fusarium wilt race 1 in peas

Resistance to fusarium wilt in peas (Pisumsativum L.) caused by Fusarium oxysporum Schlect. f. sp. pisi race 1 (van Hall) Snyd. & Hans. is conferred by a single dominant gene, Fw. The gene was located in the pea genome by analyzing progenies from crosses involving genetic markers across all pea linkage groups. Phenotyping of the progenies for reaction to race 1 of the fusarium wilt pathogen was determined by field screening in a "wilt-sick" plot in Pullman, Washington. Fw was shown to be located on linkage group III, about 13 map units from Lap-1 and b and 14 map units from Td. The relatively large distances between these markers and Fw precludes the use of the linked markers in marker-assisted selection for wilt resistance. Additional markers in this region of the pea genome will be required if marker-assisted selection for Fw is to be successful.     

Change in ribosomal DNA intergenic spacer-length composition in maize recurrent selection populations. 1. Analysis of BS13, BSSS,and BSCB1

Five different ribosomal DNA (rDNA) intergenic spacer-length variants (slvs) were detected among the maize inbreds which were the progenitors of Iowa Stiff Stalk Synthetic (BSSS). One rDNASstI restriction site polymorphism in the 3 region of the 26S gene was detected. Nine different rDNA intergenic spacer (IGS) hybridization fragment patterns (assigned letter designations A-I) were observed among the BSSS progenitors. Following 7 cycles of half-sib recurrent selection in BSSS using the Ia13 double cross as a tester, hybridization fragment pattern E became predominant in the population. In contrast, 11 cycles of reciprocal recurrent selection in BSSS with the Iowa Corn Borer Synthetic No. 1 (BSCB1) population resulted in hybridization pattern D becoming predominant. Hybridization pattern E is present in the elite inbreds B14, B37, B73, and B84, which were derived from different cycles of the BSSS half-sib recurrent selection program with Ia13. Hybridization pattern D is present in the elite inbreds B89 and B94, which were derived from different cycles of the BSSS reciprocal recurrent selection program with BSCB1. Therefore, two different forms of recurrent selection on BSSS resulted in different hybridization patterns becoming predominant in the selected populations and present in elite inbreds derived from the populations. These results also suggest that rDNA IGS hybridization fragment patterns D and E, which both have the longest slv detected, may have a selective or adaptive advantage in BSSS materials grown in the Corn Belt.