Divergent recurrent selection for cold tolerance in two improved maize populations

Maize (Zea mays L.) production has significantly expanded into very short-season environments where germination and growth in cooler environments is essentially a pre-requisite. Therefore, an important goal for maize breeders is to improve local germplasm sources of inbred lines that are able to grow under these challenging conditions. The objective of this research was to evaluate direct and correlated responses in two improved early maturing maize populations [NDSCD(M)C10 and BS22(R)C7] after two cycles of S₁ and full-sib intra-population recurrent selection for cold tolerance. The S₁ and full-sib progenies were obtained by self-pollinating 100 random plants and by intercrossing 200 random plants, respectively. Ten percent of the families were selected, based on an index that included emergence percentage, seedling vigor, and root lodging percentage, and recombined at the same time in a summer nursery based upon data across northern North Dakota locations. The essential benefit of this breeding methodology was to achieve one year per cycle of selection based upon progenies. However, direct response to selection was not significant while some correlated responses were significant. We decided to report these results in order to encourage other scientists the evaluation of additional sources of germplasm, the screening at various dates, and the selection of target environments with more intensive cold stress before initiating long-term selection programs for cold tolerance. In addition, further research on the current and alternative long-term selection methods for cold tolerance is recommended for continuous genetic improvement of advanced cycles in the northern U.S. Corn Belt. Part of the thesis submitted by B. Sezegen in partial fulfillment of the requirements for a MS degree at North Dakota State University.     

Evaluation of three cycles of full-sib reciprocal recurrent selection in two maize populations from the Northeast of Spain

In Europe a heterotic pattern commonly used in maize breeding is “American Dent × European Flint”. Maize breeding programs generally use only a small portion of the useful genetic variability present in the local open-pollinated varieties which, otherwise, have a poor performance that hampers their use. Two maize composites, EZS33 formed by open-pollinated flint varieties from dry or Mediterranean Spain, and EZS34 from USA dent populations, were developed in Zaragoza (Spain). Both were subjected to three cycles of full-sib reciprocal recurrent selection (RRS) for yield. The objective of our study was to evaluate after three cycles of selection the effect of RRS. The selection significantly increased yield in the population crosses (3.0 % per cycle) and in the crosses of the populations with testers of different origin. The correlated responses for the population crosses in other agronomic traits like flowering and early vigor were in the desired direction, but plant height was reduced. We propose that stover yield or related traits could be included in selection programs as selection criteria to prevent their decline. The adapted population increased the frequency of favorable alleles for yield, but the inbreeding depression counteracted their effect on the mean. RRS had a positive effect on other traits. According to our data, RRS can be useful to develop improved populations from which it would be possible to develop lines with improved both specific and general combining ability with different heterotic groups. We conclude that the heterotic pattern “Mediterranean Spain × US Dent” is potentially very useful for maize breeding for adaptation to Mediterranean conditions and an interesting source of cultivars for low-input agriculture.     

Genetic-diversity assessed by microsatellites in tropical maize populations submitted to a high-intensity reciprocal recurrent selection

BR-105 and BR-106 are important tropical maize populations, which were submitted to a high-intensity reciprocal recurrent selection, generating the IG-3 and IG-4synthetics. Using 30-microsatellite loci,we measured and compared the genetic diversity of these populations and their synthetics. The populations did not differ significantly regarding the amount of genetic diversity. As a consequence of selection, genetic variability losses, in terms of mean number of alleles per locus,proportion of polymorphic loci, and gene diversity did occur and were greater in the synthetic IG-3 than IG-4. In the synthetics, the number of loci in adherence to Hardy-Weinberg proportions was superior to that observed in the populations. The Wright's mean fixation index was higher than the mean value expected for outcrossing species (5%) indicating as light excess of homozygotic individuals in both populations. The genetic distances confirmed the favourable effects of one cycle of recurrent selection, as the synthetics became more isolated in comparison to the original populations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Mass selection for improved cold and density tolerance of two maize populations

Summary Data of planting and plant density are two cultural practices influencing grain yield of maize (Zea mays L.). Our study was designed to evaluate the usefulness of a mass selection scheme to improve cold and density tolerance of the BS2 and BS3 maize populations. Populations were planted at an early planting date and a high plant density, and three cycles of mass selection for well-filled ears on erect plants were conducted at earch of three Corn Belt locations (i.e., Waseca, MN; Ames, IA; Portageville, MO).Results showed that selection improved cold tolerance traits of BS3, but not of BS2. Mass selection did not increase density tolerance of either population at any location. Selection did not improve response to planting dates, although the early planting date did improve agronomic performance and grain yield of all entries. We concluded that mass selection at high plant densities and early planting dates at diverse geographical locations did not produce cycles adapted to specific environmental conditions.     

Responses to selection and changes in combining ability after three cycles of a modified reciprocal recurrent selection in maize

This research reports responses to selection and changes in general (GCA) and specific (SCA) combining abilities after three cycles of a modified reciprocal recurrent selection procedure (MRRS) in EPB-4 and EPB-5 maize populations. In the MRRS procedure a cycle can be completed in 1 or 2 years depending on the availability of winter breeding nurseries. The original and the three selection cycles of the two populations per se (eight entries) and the partial diallel developed from the crosses between them (16 entries) were evaluated in six environments. Realized response to selection on the population cross was 7.25% cycle⁻¹ for grain yield, −13.63% cycle⁻¹ for plant lodging, and 11.93% cyle⁻¹ for prolificacy, whereas plant and ear heights remained unchanged. GCA estimates increased with selection cycles for both populations for grain yield and prolificacy, and decreased for plant lodging, indicating that the frequency of favorable alleles with additive effects for these traits increased with the MRRS cycles in both populations. SCA estimates increased for grain yield and prolificacy indicating that the frequency of favorable complementary alleles at loci with non-additive effects in the reciprocal populations increased with the MRRS cycles. For grain yield, SCA effects increased more than GCA effects with selection cycles, indicating that MRRS exploited more the non-additive effects than the additive effects for the improvement of this trait. The overall results showed that the MRRS procedure was highly effective in improving the population cross, exploiting both GCA and SCA effects.     

Six cycles of selection for adaptation in two exotic populations of maize

Summary The maturity rating of maize (Zea mays L.) grown in Galicia (Northwestern Spain) varies from FAO 200 to 600. Later germplasm has usually too much moisture content in the kernel at normal harvesting time so serious problems with storing could appear. Besides, farmers have tended to use earlier varieties in the last few years. All this imposes limitations on the amount of germplasm available to develop varieties adapted to this area.To study the possibility of adapting late, exotic material to the environment of the Atlantic coast of Galicia we carried out six cycles of individual selection on two non-adapted populations of maize (Purdue A and Purdue B). The criterion of selection was early silking and the criterion of response was moisture content of kernel at harvesting.The original populations and the populations obtained after each cycle of selection were crossed to the hybrids CM105 × CM109 (tester Reid) and H99 × H95 (tester Lancaster) and were evaluated in four environments to study the efficiency of the selection scheme. There were significant reductions in days to silking (6.1 and 6.7 days from cycle 0 to cycle 6 for Purdue A and Purdue B, respectively) and grain moisture at harvesting (3.0 and 3.9% from cycle 0 to cycle 6 for Purdue A and Purdue B, respectively). There were also reductions in plant height and yield in both populations.In general, the crosses population × Lancaster were higher for yield than the crosses population × Reid. Yield of the population crosses by both testers decreased after the six cycles of selection probably because of the earlier maturity of the selected populations. Some inbreeding depression may also have occurred.     

Effects of recurrent selection for grain yield on plant and ear traits of five maize populations

Summary Maize (Zea mays L.) breeders are interested in the effects of recurrent selection for grain yield on other traits. Changes in plant traits could alter agronomic acceptability of the populations under selection, and observed improvements in grain yield could be explained by changes in ear traits. We evaluated changes in combining ability for plant and ear traits of BS10(FR), BS11(FR), BSSS(R), BSCB1(R), and Lancaster Surecrop associated with recurrent selection for grain yield.Recurrent selection procedures generally did not change plant and ear heights or date of silking of testcrosses of populations or of the population crosses, BS10(FR)×BSS11(FR) and BSSS(R)×BSCB1(R). Grain yield improvements, however, were associated with increases in ear-sink size.Journal Paper No. J 9517 of the Iowa Agriculture and Home Economics Exp. Stn., Ames, Iowa. Project 2152.     

Genetic divergence in two tropical maize composites after four cycles of reciprocal recurrent selection

Two tropical maize composites were subjected to four cycles of reciprocal recurrent selection to develop divergent inbred lines with good combining ability. This study was conducted to examine the extent of genetic diversity, changes in allele composition and genetic structure, of 100 randomly selected S₁ lines each from the original (C₀) and advanced (C₄) selection cycles of TZL COMP3 and TZL COMP4, genotyped using single nucleotide polymorphism (SNP) markers. Results revealed that the proportion of alleles at both low and high frequencies decreased from C₀ to C₄, whereas those at intermediate frequencies increased at C₄ in the two composites. More unique and other alleles were lost at C₄ in TZL COMP3 relative to those in TZL COMP4. The changes in different measures of genetic diversity were either small or negligible with selection in the two composites. The proportion of markers departing from Hardy–Weinberg equilibrium (HWE) decreased with selection, whereas the total number of pairs of markers in linkage disequilibrium increased with selection in the two composites. Examination of changes in population structures using a model‐based approach as well as cluster and multivariate analyses found a high degree of concordance in stratifying the 400 S₁ lines into four non‐overlapping groups corresponding to the two selection cycles each within the reciprocal composites. The observed molecular‐based divergence between cycles within the same composite and the clear differentiation between the complementary composites highlight the importance of reciprocal recurrent selection for preserving genetic diversity for long‐term selection. This increases the potential of the advanced selection cycles to sustain genetic gain in productivity of hybrids adapted to the savannas in West and Central Africa.     

Originality of M3S maize population and changes in allele frequencies revealed by SSR markers after two cycles of selfed progeny recurrent selection

Maksimir 3 Synthetic (M3S) maize population was developed at the Faculty of Agriculture University of Zagreb by intercrossing inbred lines, whose origins trace back to several open-pollinated varieties and local populations from different regions of the former Yugoslavia. The population was subjected to two cycles of selfed progeny recurrent selection for grain yield. The objectives of this study were: (i) to determine genetic distances among the parental inbred lines of the M3S population (M3S progenitors), the M3S population before and after two cycles of recurrent selection, and elite inbred lines representing the BSSS and Lancaster heterotic group; and (ii) to examine the effect of two cycles of recurrent selection on allele frequency changes in the population. Nine M3S progenitors, three BSSS lines, and three Lancaster lines were genotyped at 24 SSR loci, out of which nine randomly chosen loci were used for genotyping 96 individuals from both C0 (the M3S population before selection) and from C2 (M3S population after two cycles of selection). A total of 101 alleles were detected across 24 loci in the 15 lines, whereas 83 alleles were found in the nine M3S progenitors. Among the latter 83 alleles 31 were unique, i.e. found only in one of the progenitors. Mean genetic distance among nine M3S progenitors was 0.61 indicating a broad genetic base of the M3S population. High mean genetic distance was found between M3S progenitors and BSSS lines (0.69) and M3S progenitors and Lancaster lines (0.71). This indicates that the M3S population represents a germplasm source unrelated to both the BSSS and Lancaster germplasm. Mean genetic distance between the M3S population and BSSS as well as Lancaster lines decreased slightly after two cycles of recurrent selection suggesting the need to introduce testers from both groups in future selection in the M3S population in order to maintain heterotic complementarity of the M3S population to these groups. A test of selective neutrality identified several non-neutral loci in the population whose allele frequency changes from the C0 to the C2 cannot be explained by genetic drift. The majority of non-neutral alleles, whose frequency increased after two cycles of selection, were present in at least one line from the BSSS or Lancaster heterotic group.     

Kinetics of leaf extension in maize: Parameterization for two tropically adapted cultivars planted on two dates at Gatton

The kinetics of extension of individual leaves and component laminae and sheaths in maize were studied in two field experiments at Gatton (Australia) using two cultivars, Pioneer 3527 and Pioneer C87, in one experiment and Pioneer C87 only in the second. Destructive sampling based on non-destructive observations to ensure accurate representation of the population was used to establish timing of leaf initiation, leaf tip appearance, ligule being able to be discerned and full extension of organs. The appearance and extension of leaves were related to thermal time, calculated from temperature measured in the region of cell division and extension. Extension of leaves and sheaths was analysed using the first three phases of a four phase framework consisting of two exponential phases, rapid (linear) extension and transition to final organ length. The last phase was not examined as there were insufficient data. Final organ length was related to leaf position, and linear extension rates of leaves and laminae were linearly related to final organ length, with differing equations for upper and lower leaves. The study has proposed extensions to an existing analytical approach, and argues that regular patterns of extension of leaves and sheaths can be explained in relation to the position of the ear using a physiological approach.     

Computer simulation of a selection strategy to accommodate genotype environment interactions in a wheat recurrent selection programme

Multi-environment trials (METs) are used in plant breeding programmes to evaluate genotypes (lines/families) as a basis for selection on expected performance (yield and/or quality) in a target population of environments (TPE). When a large component of the genotype environment (G × E) interactions results from crossover interactions, samples of environments in METs that deviate from the TPE provide a suboptimal basis for selection of genotypes on performance expected in the TPE. To adjust for the negative effects of these deviations, a selection strategy that weights the data from the MET according to their expected frequency of occurrence in the TPE (i.e. a weighted selection strategy) was investigated. Computer simulation methodology was used to obtain preliminary information on the weighted selection strategy and compare it to the traditional unweighted selection strategy for a range of MET scenarios and G × E interaction models. The evaluation of the weighted selection strategy was conducted in context with the germplasm enhancement programme (GEP) of the Northern Wheat Improvement Programme in Australia. The results indicated that when the environments sampled in the MET matched those expected in the TPE, the unweighted and weighted selection strategies achieved a similar response to selection in the TPE. However, when the environments sampled in the MET did not match the expectations in the TPE and a large component of the G × E interactions resulted from crossover interactions, the weighted selection strategy achieved a greater response to selection in the TPE. The advantage of the weighted strategy increased as the amount of crossover G × E interaction increased or fewer environments were sampled in the METs.     

Methods of increasing short term response to full-sib family recurrent selection in small populations

Accelerated recurrent selection (ARS), in which selection is carried out on the predicted value of the progeny rather than on the observed performance of the parents, has been proposed as a method of increasing response to selection and of reducing cycle time. ARS schemes based on test cross evaluation of full-sib families have been compared by stochastic computer simulation. The difference in genetic and economic time scales is emphasised, with the economic long term (21 years) being only 21 or fewer cycles of selection. ARS schemes are shown frequently to offer improvements over standard recurrent selection methods under these circumstances, since they allow more cycles of selection in a given time frame. Schemes with very low effective population sizes often give the greatest response to selection over the time scales considered here. It is suggested that evaluation of cumulative responses to selection over defined periods of time, either by Monte Carlo simulation or by stochastic theory, is the best method of ranking alternative selection schemes. Evaluating response to selection by deterministic methods, or by attempting to take drift into account by evaluating schemes with identical effective population sizes can be misleading. This revised version was published online in August 2006 with corrections to the Cover Date.     

Diallel analysis among 16 maize populations adapted to the northern U.S. Corn Belt for grain yield and grain quality traits

Genetically diverse germplasm is needed to increase frequency of favorable alleles of economically important traits in maize improvement. The objectives of this study were to determine the genetic components involved in grain yield and grain quality traits, and provide preliminary assessment of useful heterotic groups and patterns from a large sample of maize populations adapted to the northern U.S. Corn Belt. Sixteen populations were used in the diallel mating design following Gardner–Eberhart Analysis II to estimate variety (v _i ) and heterosis (h _ij ) genetic effects for grain yield and grain quality traits. Specific heterosis (s _ij ) and predicted means of population crosses for grain yield were used to evaluate the heterotic relationships among the populations. Data for grain yield and grain quality traits were generated in partially balanced single lattice experiments across North Dakota (ND) locations in 2010, 2011, and 2012. Analyses of variance showed significant differences among genotypes. Heterosis effects explained most of the differences among diallel entries for grain yield, while v _i effects had greater influence on grain quality traits. NDL, EARLYGEM 21c, NDSCD(FS-CS)C2, NDSS, and NDSM(M-FS)C9 were identified as elite populations for grain quality improvement. NDSS × NDBS22(R-T1)C9 and NDBS1011 × EARLYGEM 21c showed high s _ij effects for grain yield with good grain quality. NDSS and EARLYGEM 21c represent stiff stalk synthetic (SSS) group, and NDBS1011 fall under non-SSS group. Further studies need to validate the heterotic group of NDBS22(R-T1)C9. Recurrent and pedigree selection programs will be established for selected populations and population crosses to integrate pre-breeding with cultivar development.     

Comparison of predicted and observed response to selection in two breeding populations of perennial ryegrass

To predict any genetic response to selection, two breeding populations of perennial ryegrass, consisting of half-sib families, were evaluated as spaced plants from 1987 to 1989 for several agronomical traits. In each population, the best plants selected through a multitrait Smith and Hazel's index were planted in a polycross to produce the second generation half-sib families. Both generations were evaluated together from 1991 to 1993 as spaced plants. The difference between generations was the observed genetic response, which was compared to that predicted. Rust tolerance, aftermath heading and persistence were significantly improved in both populations. In trial plots used to assess the yield, no effect of mass selection was noticed. Thus, it is concluded that mass selection failed to increase productivity but produced some improvements in disease tolerance and aftermath heading.     

Reciprocal recurrent selection applied to Coffea canephora Pierre. I. Characterization and evaluation of breeding populations and value of intergroup hybrids

Summary Two breeding populations have been identified with Coffea canephora, based on geographical and genetic differences: the Guinean group from West Africa and the Congolese group from central Africa. A reciprocal recurrent selection programme based on these groups was initiated in Côte d'Ivoire in 1984. Genotypes of both groups available in field collections are either of cultivated or wild origin.Genotypic and phenotypic variability was evaluated within and between the groups for the following characteristics: isozyme patterns, architecture, drought tolerance and vigour, technological and organoleptic traits, pest and disease resistance. Variation between and within groups was found to be large. Based on isozyme analysis and phenotypic observations, two subgroups were identified within the Congolese group. Phenotypic values of parents and testers used in the first cycle of reciprocal recurrent selection are presented.Results of two intergroup hybrid trials indicate high yield and vigour for these hybrids compared to intragroup hybrids, justifying the reciprocal selection approach. Furthermore, the variation observed indicates that recurrent selection would allow progress for selection traits for both populations and intergroup hybrids.     

Marker-assisted selection efficiency in multiple connected populations: a simulation study based on the results of a QTL detection experiment in maize

Many studies already investigated marker-assisted selection (MAS) efficiency but mainly in biparental populations. Connected multiparental populations address a broader diversity and confer a gain of power for QTL detection which must be beneficial for MAS. Our objective was to compare multiparental connected designs to biparental populations taken separately for MAS and phenotypic selection. We first detected QTL for flowering time and grain yield in an experimental maize design involving four parental inbred lines crossed to produce six different biparental populations and confirmed the advantage of multiparental connected designs over biparental populations for QTL detection. Based on these results we performed stochastic simulations to evaluate the expected efficiency of four generations of MAS and phenotypic selection. Different parameters were considered: trait heritability, genetic architecture and whether QTL were assumed to be known or have to be detected. Genetic gains were higher in the multiparental design than on average over the biparental populations considered separately, especially when favourable alleles were equally distributed among parental lines. When QTL detection was included in the simulation process, we found that type I error risk considered for declaring QTL as significant should be adapted to the design. Type I error risks leading to the best response were higher for the biparental populations than for the multiparental design. Besides addressing a broader diversity, multiparental designs increase the power of QTL detection, which reinforces their superiority over biparental designs for MAS. Application of MAS to multiparental designs therefore appears promising to accelerate genetic gain in plant breeding programs.