Flowering time QTL in natural populations of Arabidopsis thaliana and implications for their adaptive value

The genetic basis of phenotypic traits is of great interest to evolutionary biologists, but their contribution to adaptation in nature is often unknown. To determine the genetic architecture of flowering time in ecologically relevant conditions, we used a recombinant inbred line population created from two locally adapted populations of Arabidopsis thaliana from Sweden and Italy. Using these RILs, we identified flowering time QTL in growth chambers that mimicked the natural temperature and photoperiod variation across the growing season in each native environment. We also compared the genomic locations of flowering time QTL to those of fitness (total fruit number) QTL from a previous three‐year field study. Ten total flowering time QTL were found, and in all cases, the Italy genotype caused early flowering regardless of the conditions. Two QTL were consistent across chamber environments, and these had the largest effects on flowering time. Five of the fitness QTL colocalized with flowering time QTL found in the Italy conditions, and in each case, the local genotype was favoured. In contrast, just two flowering time QTL found in the Sweden conditions colocalized with fitness QTL and in only one case was the local genotype favoured. This implies that flowering time may be more important for adaptation in Italy than Sweden. Two candidate genes (FLC and VIN3) underlying the major flowering time QTL found in the current study are implicated in local adaptation.     

Genomic regions controlling vernalization and photoperiod responses in oat

Oat genotypes vary for photoperiod and vernalization responses. Vernalization often promotes earlier flowering in fall-sown but not spring-sown cultivars. Longer photoperiods also promote earlier flowering, and the response to longer photoperiods tends to be greater in cultivars from higher latitudes. To investigate the genetic basis of photoperiod and vernalization responses in oat, we mapped QTLs for flowering time under four combinations of photoperiod and vernalization treatments in the Ogle 2 TAM O-301 mapping population in growth chambers. We also mapped QTLs for flowering time in early spring and late-spring field plantings to determine the genetic basis of response to early spring planting in oat. Three major flowering-time QTLs (on linkage groups OT8, OT31 and OT32) were detected in most conditions. QTLs with smaller effects on flowering were less-consistently observed among treatments. Both vernalization-sensitive and insensitive QTLs were discovered. Longer photoperiod or vernalization alone tended to decrease the effects of flowering-time QTLs. Applied together, longer photoperiod and vernalization interacted synergistically, often on the same genomic regions. Earlier spring planting conferred an attenuated vernalization treatment on seeds. The major flowering-time QTLs mapped in this study matched those mapped previously in the Kanota 2 Ogle oat mapping population. Between these two studies, we found a concordance of flowering-time QTLs, segregation distortion, and complex genetic linkages. These effects may all be related to chromosomal rearrangements in hexaploid oat. Comparative mapping between oat and other grasses will facilitate molecular analysis of vernalization response in oat.     

Seasonal and plant-density dependency for quantitative trait loci affecting flowering time in multiple populations of Arabidopsis thaliana

Multiple environmental cues regulate the transition to flowering. In natural environments, plants perceive seasonal progression by changes in day length and growth temperature, and plant density is monitored by changes in the light quality reflected from neighbouring vegetation. To understand the seasonal and plant-density dependence associated with natural allelic variation in flowering time, we conducted a quantitative trait loci (QTL) mapping study in Ler x Cvi, Bay x Sha and Ler x No-0 recombinant inbred line (RIL) populations of Arabidopsis thaliana. Days and total leaf number to bolting were examined under low and high plant density (200 or 1600 plants m(-2)) in autumn-winter and spring seasons. We found between 4 and 10 QTLs associated with seasonal and density variations in each RIL population. For Ler x Cvi and Bay x Sha RIL populations, a major proportion of QTLs showed seasonal and density interaction (up to 63%) and four QTLs were common to all environments (21%). Only three QTLs showed seasonal or density dependency. By aligning the linkage maps onto a common physical map, we detected at least one QTL at chromosome 2 and two QTLs at chromosome 5 that overlap between the three RIL populations, suggesting that these QTLs play a crucial role in the adaptive control of flowering time.     

The extent and genetic basis of phenotypic divergence in life history traits in Mimulus guttatus

Differential natural selection acting on populations in contrasting environments often results in adaptive divergence in multivariate phenotypes. Multivariate trait divergence across populations could be caused by selection on pleiotropic alleles or through many independent loci with trait‐specific effects. Here, we assess patterns of association between a suite of traits contributing to life history divergence in the common monkey flower, Mimulus guttatus, and examine the genetic architecture underlying these correlations. A common garden survey of 74 populations representing annual and perennial strategies from across the native range revealed strong correlations between vegetative and reproductive traits. To determine whether these multitrait patterns arise from pleiotropic or independent loci, we mapped QTLs using an approach combining high‐throughput sequencing with bulk segregant analysis on a cross between populations with divergent life histories. We find extensive pleiotropy for QTLs related to flowering time and stolon production, a key feature of the perennial strategy. Candidate genes related to axillary meristem development colocalize with the QTLs in a manner consistent with either pleiotropic or independent QTL effects. Further, these results are analogous to previous work showing pleiotropy‐mediated genetic correlations within a single population of M. guttatus experiencing heterogeneous selection. Our findings of strong multivariate trait associations and pleiotropic QTLs suggest that patterns of genetic variation may determine the trajectory of adaptive divergence.     

Prediction of flowering time in Brassica oleracea using a quantitative trait loci-based phenology model

Uniformly developing plants with a predictable time to harvest or flowering under unfavourable climate conditions are a major breeding goal in crop species. The main flowering regulators and their response to environmental signals have been identified in Arabidopsis thaliana and homologues of flowering genes have been mapped in many crop species. However, it remains unclear which genes determine within and across genotype flowering time variability in Brassica oleracea and how genetic flowering time regulation is influenced by environmental factors. The goal of this study is model-based prediction of flowering time in a B. oleracea DH-line population using genotype-specific and quantitative trait loci (QTL) model input parameters. A QTL-based phenology model accounting for genotypic differences in temperature responses during vernalisation and non-temperature-sensitive durations from floral transition to flowering was evaluated in two field trials. The model was parameterised using original genotype-specific model input parameters and QTL effects. The genotype-specific model parameterisation showed accurate predictability of flowering time if floral induction was promoted by low temperature (R(2) = 0.81); unfavourably high temperatures reduced predictability (R(2) = 0.65). Replacing original model input parameters by QTL effects reduced the capability of the model to describe across-genotype variability (R(2) = 0.59 and 0.50). Flowering time was highly correlated with a model parameter accounting for vernalisation effects. Within-genotype variability was significantly correlated with the same parameter if temperature during the inductive phase was high. We conclude that flowering time variability across genotypes was largely due to differences in vernalisation response, although it has been shown elsewhere that the candidate FLOWERING LOCUS C (FLC) did not co-segregate with flowering time in the same population. FLC independent vernalisation pathways have been described for several species, but not yet for B. oleracea.     

应用中国和欧洲油菜建立的双二倍体群体对3个重要农艺性状的QTL定位以及与环境的互作分析

以中国的高油分自交系“高油”和欧洲高含油量品种“Sollux”的F1产生的282个株系组成的双二倍体(DH)群体为材料,在125个SSR标记座位构建的连锁图谱基础上,根据在中国和欧洲四个不同环境下的表型鉴定结果,采用混合线性模型基础上的QTL分析软件,对油菜3个重要农艺性状:株高,开花期和成熟期进行了数量性状基因座位(QTL)的联合定位分析,估测了这些QTL的加性、上位性以及与环境的互作效应。结果表明各性状均受多个加性、加加上位以及与环境互作的QTL控制。株高受多个QTL影响(12个位点具有加性或兼有环境互作效应,5个位点具有互作效应),以加性效应为主,加性效应总和可解释定位群体表型变异的75%左右,并多兼有上位性效应。12个主效QTL中,9个是“高油”等位基因相对“Sollux”有降低株高的作用,大多数加性×环境互作QTL的有效等位基因具有环境选择特异性。7个ae基因座位中,5个“高油”等位基因在杭州种植环境下,除一例外所有在德国环境下的互作基因座中,“Sollux”等位基因起着增加株高的作用,加加上位性主效总和为加性主效总和的三分之一。7个控制花期和8个控制成熟期的主效QTL中,分别有6个和5个是来自“高油”的等位基因相对“Sollux”具有提前开花和成熟的效应,这些QTL的效应总和占到性状表型变异的60%左右。5个位于第2和第12连锁群中的2个大效应QTL可能和已多次报导的VFN1和VFN3基因相近或相同。开花期和成熟期两性状均检测到显著的ae互作效应,双亲等位基因的效应在各环境下呈离散分布。位于14和19连锁群上的两个主效株高QTL同时也是控制开花期和油分含量的基因位点,因而利用这两个位点进行标记辅助筛选时要考虑到对油分含量的影响。控制成熟期的8个主效QTL中有3个同时也是控制开花期的基因座位,证实了开花期和成熟期高度正相关的遗传基础,两个生育性状均表现有较弱的QTL间加加上位互作,但以主效QTL的作用为主。     

Genetic variation in flowering phenology and avoidance of seed predation in native populations of Ulex europaeus

The genetic variation in flowering phenology may be an important component of a species’ capacity to colonize new environments. In native populations of the invasive species Ulex europaeus, flowering phenology has been shown to be bimodal and related to seed predation. The aim of the present study was to determine if this bimodality has a genetic basis, and to investigate whether the polymorphism in flowering phenology is genetically linked to seed predation, pod production and growth patterns. We set up an experiment raising maternal families in a common garden. Based on mixed analyses of variance and correlations among maternal family means, we found genetic differences between the two main flowering types and confirmed that they reduced seed predation in two different ways: escape in time or predator satiation. We suggest that this polymorphism in strategy may facilitate maintain high genetic diversity for flowering phenology and related life‐history traits in native populations of this species, hence providing high evolutionary potential for these traits in invaded areas.     

Habitat-specific natural selection at a flowering-time QTL is a main driver of local adaptation in two wild barley populations

Understanding the genetic basis of local adaptation requires insight in the fitness effects of individual loci under natural field conditions. While rapid progress is made in the search for genes that control differences between plant populations, it is typically unknown whether the genes under study are in fact key targets of habitat-specific natural selection. Using a quantitative trait loci (QTL) approach, we show that a QTL associated with flowering-time variation between two locally adapted wild barley populations is an important determinant of fitness in one, but not in the other population's native habitat. The QTL mapped to the same position as a habitat-specific QTL for field fitness that affected plant reproductive output in only one of the parental habitats, indicating that the genomic region is under differential selection between the native habitats. Consistent with the QTL results, phenotypic selection of flowering time differed between the two environments, whereas other traits (growth rate and seed weight) were under selection but experienced no habitat-specific differential selection. This implies the flowering-time QTL as a driver of adaptive population divergence. Our results from phenotypic selection and QTL analysis are consistent with local adaptation without genetic trade-offs in performance across environments, i.e. without alleles or traits having opposing fitness effects in contrasting environments.     

Genetic Variation for Life History Sensitivity to Seasonal Warming in Arabidopsis thaliana

Climate change has altered life history events in many plant species; however, little is known about genetic variation underlying seasonal thermal response. In this study, we simulated current and three future warming climates and measured flowering time across a globally diverse set of Arabidopsis thaliana accessions. We found that increased diurnal and seasonal temperature (1°–3°) decreased flowering time in two fall cohorts. The early fall cohort was unique in that both rapid cycling and overwintering life history strategies were revealed; the proportion of rapid cycling plants increased by 3–7% for each 1° temperature increase. We performed genome-wide association studies (GWAS) to identify the underlying genetic basis of thermal sensitivity. GWAS identified five main-effect quantitative trait loci (QTL) controlling flowering time and another five QTL with thermal sensitivity. Candidate genes include known flowering loci; a cochaperone that interacts with heat-shock protein 90; and a flowering hormone, gibberellic acid, a biosynthetic enzyme. The identified genetic architecture allowed accurate prediction of flowering phenotypes (R² > 0.95) that has application for genomic selection of adaptive genotypes for future environments. This work may serve as a reference for breeding and conservation genetic studies under changing environments.     

GENETIC CONSTRAINTS ON LIFE-HISTORY EVOLUTION: QUANTITATIVE-TRAIT LOCI INFLUENCING GROWTH AND FLOWERING IN ARABIDOPSIS THALIANA

We have mapped genes causing life-history trade-offs, and they behave as predicted by ecological theory. Energetic and quantitative-genetic models suggest a trade-off between age and size at first reproduction. Natural selection favored plants that flower early and attain large size at first reproduction. Response to selection was opposed by a genetic trade-off between these two components of fitness. Two quantitative-trait loci (QTLs) influencing flowering time were mapped in a recombinant inbred population of Arabidopsis. These QTLs also influenced size at first reproduction, but did not affect growth rate (resource acquisition). Substitutions of small chromosomal segments, which may represent allelic differences at flowering time loci, caused genetic trade-offs between life-history components. One QTL explained 22% of the genetic variation in flowering time. It is within a few centiMorgans (cM) of the gigantea (GI) locus, and may be allelic with GI. Sixteen percent of the genetic variation was explained by another QTL, FDR1, near 18 cM on chromosome II, which does not correspond to any previously identified flowering-time locus. These life-history genes regulate patterns of resource allocation and life-history trade-offs in this population.     

QTL studies reveal little relevance of chilling-related seedling traits for yield in maize

Prolonged low temperature phases and short-term cold spells often occur in spring during the crucial stages of early maize (Zea mays L.) development. The effect of low temperature-induced growth retardation at the seedling stage on final yield is poorly studied. Therefore, the aim was to identify genomic regions associated with morpho-physiological traits at flowering and harvest stage and their relationship to previously identified quantitative trait loci (QTLs) for photosynthesis and morpho-physiological traits from the same plants at seedling stage. Flowering time, plant height and shoot biomass components at harvest were measured in a dent mapping population for cold tolerance studies, which was sown in the Swiss Midlands in early and late spring in two consecutive years. Early-sown plants exhibited chilling stress during seedling stage, whereas late-sown plants grew under favorable conditions. Significant QTLs, which were stable across environments, were found for plant height and for the time of flowering. The QTLs for flowering were frequently co-localized with QTLs for plant height or ear dry weight. The comparison with QTLs detected at seedling stage revealed only few common QTLs. A pleiotropic effect was found on chromosome 3 which revealed that a good photosynthetic performance of the seedling under warm conditions had a beneficial effect on plant height and partially on biomass at harvest. However, a high chilling tolerance of the seedling seemingly had an insignificant or small negative effect on the yield.     

QTL mapping with near-isogenic lines in maize

A set of 89 near-isogenic lines (NILs) of maize was created using marker-assisted selection. Nineteen genomic regions, identified by restriction fragment length polymorphism loci and chosen to represent portions of all ten maize chromosomes, were introgressed by backcrossing three generations from donor line Tx303 into the B73 genetic background. NILs were genotyped at an additional 128 simple sequence repeat loci to estimate the size of introgressions and the amount of background introgression. Tx303 introgressions ranged in size from 10 to 150 cM, with an average of 60 cM. Across all NILs, 89% of the Tx303 genome is represented in targeted and background introgressions. The average proportion of background introgression was 2.5% (range 0–15%), significantly lower than the expected value of 9.4% for third backcross generation lines developed without marker-assisted selection. The NILs were grown in replicated field evaluations in two years to map QTLs for flowering time traits. A parallel experiment of testcrosses of each NIL to the unrelated inbred, Mo17, was conducted in the same environments to map QTLs in NIL testcross hybrids. QTLs affecting days to anthesis, days to silking, and anthesis-silk interval were detected in both inbreds and hybrids in both environments. The testing environments differed dramatically for drought stress, and different sets of QTLs were detected across environments. Furthermore, QTLs detected in inbreds were typically different from QTLs detected in hybrids, demonstrating the genetic complexity of flowering time. NILs can serve as a valuable genetic mapping resource for maize breeders and geneticists. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The genetic basis of adaptive population differentiation: a quantitative trait locus analysis of fitness traits in two wild barley populations from contrasting habitats

We used a quantitative trait locus (QTL) approach to study the genetic basis of population differentiation in wild barley, Hordeum spontaneum. Several ecotypes are recognized in this model species, and population genetic studies and reciprocal transplant experiments have indicated the role of local adaptation in shaping population differences. We derived a mapping population from a cross between a coastal Mediterranean population and a steppe inland population from Israel and assessed F3 progeny fitness in the natural growing environments of the two parental populations. Dilution of the local gene pool, estimated as the proportion of native alleles at 96 marker loci in the recombinant lines, negatively affected fitness traits at both sites. QTLs for fitness traits tended to differ in the magnitude but not in the direction of their effects across sites, with beneficial alleles generally conferring a greater fitness advantage at their native site. Several QTLs showed fitness effects at one site only, but no opposite selection on individual QTLs was observed across the sites. In a common-garden experiment, we explored the hypothesis that the two populations have adapted to divergent nutrient availabilities. In the different nutrient environments of this experiment, but not under field conditions, fitness of the F3 progeny lines increased with the number of heterozygous marker loci. Comparison of QTL-effects that underlie genotype x nutrient interaction in the common-garden experiment and genotype x site interaction in the field suggested that population differentiation at the field sites may have been driven by divergent nutrient availabilities to a limited extent. Also in this experiment no QTLs were observed with opposite fitness effects in contrasting environments. Our data are consistent with the view that adaptive differentiation can be based on selection on multiple traits changing gradually along ecological gradients. This can occur without QTLs showing opposite fitness effects in the different environments, that is, in the absence of genetic trade-offs in performance between environments.     

Assortative mating by flowering time and its effect on correlated traits in variable environments

Reproductive timing is a key life‐history trait that impacts the pool of available mates, the environment experienced during flowering, and the expression of other traits through genetic covariation. Selection on phenology, and its consequences on other life‐history traits, has considerable implications in the context of ongoing climate change and shifting growing seasons. To test this, we grew field‐collected seed from the wildflower Mimulus guttatus in a greenhouse to assess the standing genetic variation for flowering time and covariation with other traits. We then created full‐sib families through phenological assortative mating and grew offspring in three photoperiod treatments representing seasonal variation in daylength. We find substantial quantitative genetic variation for the onset of flowering time, which covaried with vegetative traits. The assortatively‐mated offspring varied in their critical photoperiod by over two hours, so that families differed in their probability of flowering across treatments Allocation to flowering and vegetative growth changed across the daylength treatments, with consistent direction and magnitude of covariation among flowering time and other traits. Our results suggest that future studies of flowering time evolution should consider the joint evolution of correlated traits and shifting seasonal selection to understand how environmental variation influences life histories.     

Selection on QTL and complex traits in complex environments

Understanding genetic variation for complex traits in heterogeneous environments is a fundamental problem in biology. In this issue of Molecular Ecology, Fournier‐Level et al. ( 2013 ) analyse quantitative trait loci (QTL) influencing ecologically important phenotypes in mapping populations of Arabidopsis thaliana grown in four habitats across its native European range. They used causal modelling to quantify the selective consequences of life history and morphological traits and QTL on components of fitness. They found phenology QTL colocalizing with known flowering time genes as well as novel loci. Most QTL influenced fitness via life history and size traits, rather than QTL having direct effects on fitness. Comparison of phenotypes among environments found no evidence for genetic trade‐offs for phenology or growth traits, but genetic trade‐offs for fitness resulted because flowering time had opposite fitness effects in different environments. These changes in QTL effects and selective consequences may maintain genetic variation among populations.     

Molecular mechanisms for the photoperiodic regulation of flowering in soybean

Photoperiodic flowering is one of the most important factors affecting regional adaptation and yield in soybean (Glycine max). Plant adaptation to long-day conditions at higher latitudes requires early flowering and a reduction or loss of photoperiod sensitivity; adaptation to short-day conditions at lower latitudes involves delayed flowering, which prolongs vegetative growth for maximum yield potential. Due to the influence of numerous major loci and quantitative trait loci (QTLs), soybean has broad adaptability across latitudes. Forward genetic approaches have uncovered the molecular basis for several of these major maturity genes and QTLs. Moreover, the molecular characterization of orthologs of Arabidopsis thaliana flowering genes has enriched our understanding of the photoperiodic flowering pathway in soybean. Building on early insights into the importance of the photoreceptor phytochrome A, several circadian clock components have been integrated into the genetic network controlling flowering in soybean: E1, a repressor of FLOWERING LOCUS T orthologs, plays a central role in this network. Here, we provide an overview of recent progress in elucidating photoperiodic flowering in soybean, how it contributes to our fundamental understanding of flowering time control, and how this information could be used for molecular design and breeding of high-yielding soybean cultivars.     

Effects of genetic background and environment on QTLs and epistasis for rice (Oryza sativa L.) panicle number

A double-haploid (DH) population and a recombinant inbred (RI) line population, derived from a cross between a tropical japonica variety, Azucena, as male parent and two indica varieties, IR64 and IR1552, as female parents respectively, were used in both field and pot experiments for detecting QTLs and epistasis for rice panicle number in different genetic backgrounds and different environments. Panicle number (PN) was measured at maturity. A molecular map with 192 RFLP markers for the DH population and a molecular map with 104 AFLP markers and 103 RFLP markers for the RI population were constructed, in which 70 RFLP markers were the same. Six QTLs were identified in the DH population, including two detected from field experiments and four from pot experiments. The two QTLs, mapped on chromosomes 1 and 12, were identical in both field and pot experiments. In the RI population, nine QTLs were detected, five QTLs from field conditions and four from the pot experiments. Three of these QTLs were identical in both experimental conditions. Only one QTL, linked to CDO344 on chromosome 12, was detected across the populations and experiments. Different epistasitic interaction loci on PN were found under different populations and in different experimental conditions. One locus, flanked by RG323 and RZ801 on chromosome 1, had an additive effect in the DH population, but epistatic effects in the RI population. These results indicate that the effect of genetic background on QTLs is greater than that of environments, and epistasis is more sensitive to genetic background and environments than main-effect QTLs. QTL and epistatic loci could be interchangeable depending on the genetic backgrounds and probably on the environments where they are identified. Received: 26 May 2000 / Accepted: 19 October 2000     

Quantitative Trait Loci for Thermal Time to Flowering and Photoperiod Responsiveness Discovered in Summer Annual-Type Brassica napus L

Time of flowering is a key adaptive trait in plants and is conditioned by the interaction of genes and environmental cues including length of photoperiod, ambient temperature and vernalisation. Here we investigated the photoperiod responsiveness of summer annual-types of Brassica napus (rapeseed, canola). A population of 131 doubled haploid lines derived from a cross between European and Australian parents was evaluated for days to flowering, thermal time to flowering (measured in degree-days) and the number of leaf nodes at flowering in a compact and efficient glasshouse-based experiment with replicated short and long day treatments. All three traits were under strong genetic control with heritability estimates ranging from 0.85–0.93. There was a very strong photoperiod effect with flowering in the population accelerated by 765 degree-days in the long day versus short day treatments. However, there was a strong genetic correlation of line effects (0.91) between the long and short day treatments and relatively low genotype x treatment interaction indicating that photoperiod had a similar effect across the population. Bivariate analysis of thermal time to flowering in short and long days revealed three main effect quantitative trait loci (QTLs) that accounted for 57.7% of the variation in the population and no significant interaction QTLs. These results provided insight into the contrasting adaptations of Australian and European varieties. Both parents responded to photoperiod and their alleles shifted the population to earlier flowering under long days. In addition, segregation of QTLs in the population caused wide transgressive segregation in thermal time to flowering. Potential candidate flowering time homologues located near QTLs were identified with the aid of the Brassica rapa reference genome sequence. We discuss how these results will help to guide the breeding of summer annual types of B. napus adapted to new and changing environments.     

Identification of quantitative trait loci for yield and yield components in an advanced backcross population derived from the<Emphasis Type="Italic"> Oryza sativa</Emphasis> variety IR64 and the wild relative<Emphasis Type="Italic"> O. rufipogon</Emphasis>

A BC₂F₂ population developed from an interspecific cross between Oryza sativa (cv IR64) and O. rufipogon (IRGC 105491) was used in an advanced backcross QTL analysis to identify and introduce agronomically useful genes from this wild relative into the cultivated gene pool. The objectives of this study were: (1) to identify putative yield and yield component QTLs that can be useful to improve the elite cultivar IR64; (2) to compare the QTLs within this study with previously reported QTLs in rice as the basis for identifying QTLs that are stable across different environments and genetic backgrounds; and (3) to compare the identified QTLs with previously reported QTLs from maize to examine the degree of QTL conservation across the grass family. Two hundred eighty-five families were evaluated in two field environments in Indonesia, with two replications each, for 12 agronomic traits. A total of 165 markers consisting of 131 SSRs and 34 RFLPs were used to construct the genetic linkage map. By employing interval mapping and composite interval mapping, 42 QTLs were identified. Despite its inferior performance, 33% of the QTL alleles originating from O. rufipogon had a beneficial effect for yield and yield components in the IR64 background. Twenty-two QTLs (53.4%) were located in similar regions as previously reported rice QTLs, suggesting the existence of stable QTLs across genetic backgrounds and environments. Twenty QTLs (47.6%) were exclusively detected in this study, uncovering potentially novel alleles from the wild, some of which might improve the performance of the tropical indica variety IR64. Additionally, several QTLs for plant height, grain weight, and flowering time detected in this study corresponded to homeologous regions in maize containing previously detected maize QTLs for these traits.     

Drought stress and tropical maize: QTL-by-environment interactions and stability of QTLs across environments for yield components and secondary traits

A recombinant inbred line (RIL) population was evaluated in seven field experiments representing four environments: water stress at flowering (WS) and well-watered (WW) conditions in Mexico and Zimbabwe. The QTLs were identified for each trait in each individual experiment (single-experiment analysis) as well as per environment, per water regime across locations and across all experiments (joint analyses). For the six target traits (male flowering, anthesis-to-silking interval, grain yield, kernel number, 100-kernel fresh weight and plant height) 81, 57, 51 and 34 QTLs were identified in the four step-wise analyses, respectively. Despite high values of heritability, the phenotypic variance explained by QTLs was reduced, indicating epistatic interactions. About 80, 60 and 6% of the QTLs did not present significant QTL-by-environment interactions (QTL × E) in the joint analyses per environment, per water regime and across all experiments. The expression of QTLs was quite stable across years at a given location and across locations under the same water regime. However, the stability of QTLs decreased drastically when data were combined across water regimes, reflecting a different genetic basis of the target traits in the drought and well-watered trials. Several clusters of QTLs for different traits were identified by the joint analyses of the WW (chromosomes 1 and 8) and WS (chromosomes 1, 3 and 5) treatments and across water regimes (chromosome 1). Those regions are clear targets for future marker-assisted breeding, and our results confirm that the best approach to breeding for drought tolerance includes selection under water stress.