Segregation Distortion Of Markers Research Articles

Perennial Rye: Genetics of Perenniality and Limited Fertility.

Perenniality, the ability of plants to regrow after seed set, could be introgressed into cultivated rye by crossing with the wild relative and perennial Secale strictum. However, studies in the past showed that Secale cereale × Secale strictum-derived cultivars were also characterized by reduced fertility what was related to so called chromosomal multivalents, bulks of chromosomes that paired together in metaphase I of pollen mother cells instead of only two chromosomes (bivalents). Those multivalents could be caused by ancient translocations that occurred between both species. Genetic studies on perennial rye are quite old and especially the advent of molecular markers and genome sequencing paved the way for new insights and more comprehensive studies. After a brief review of the past research, we used a basic QTL mapping approach to analyze the genetic status of perennial rye. We could show that for the trait perennation 0.74 of the genetic variance in our population was explained by additively inherited QTLs on chromosome 2R, 3R, 4R, 5R and 7R. Fertility on the other hand was with 0.64 of explained genetic variance mainly attributed to a locus on chromosome 5R, what was most probably the self-incompatibility locus S5. Additionally, we could trace the Z locus on chromosome 2R by high segregation distortion of markers. Indications for chromosomal co-segregation, like multivalents, could not be found. This study opens new possibilities to use perennial rye as genetic resource and for alternative breeding methods, as well as a valuable resource for comparative studies of perennation across different species.

Development of sub-tropically adapted diverse provitamin-A rich maize inbreds through marker-assisted pedigree selection, their characterization and utilization in hybrid breeding.

Malnutrition has emerged as one of the major health problems worldwide. Traditional yellow maize has low provitamin-A (proA) content and its genetic base in proA biofortification breeding program of subtropics is extremely narrow. To diversify the proA rich germplasm, 10 elite low proA inbreds were crossed with a proA rich donor (HP702-22) having mutant crtRB1 gene. The F2 populations derived from these crosses were genotyped using InDel marker specific to crtRB1. Severe marker segregation distortion was observed. Seventeen crtRB1 inbreds developed through marker-assisted pedigree breeding and seven inbreds generated using marker-assisted backcross breeding were characterized using 77 SSRs. Wide variation in gene diversity (0.08 to 0.79) and dissimilarity coefficient (0.28 to 0.84) was observed. The inbreds were grouped into three major clusters depicting the existing genetic diversity. The crtRB1-based inbreds possessed high β-carotene (BC: 8.72μg/g), β-cryptoxanthin (BCX: 4.58μg/g) and proA (11.01μg/g), while it was 2.35μg/g, 1.24μg/g and 2.97μg/g in checks, respectively. Based on their genetic relationships, 15 newly developed crtRB1-based inbreds were crossed with five testers (having crtRB1 gene) using line × tester mating design. 75 experimental hybrids with crtRB1 gene were evaluated over three locations. These experimental hybrids possessed higher BC (8.02μg/g), BCX (4.69μg/g), proA (10.37μg/g) compared to traditional hybrids used as check (BC: 2.36 μg/g, BCX: 1.53μg/g, proA: 3.13μg/g). Environment and genotypes × environment interaction had minor effects on proA content. Both additive and dominance gene action were significant for proA. The mean proportion of proA to total carotenoids (TC) was 44% among crtRB1-based hybrids, while 11% in traditional hybrids. BC was found to be positively correlated with BCX (r = 0.68) and proA (r = 0.98). However, no correlation was observed between proA and grain yield. Several hybrids with >10.0 t/ha grain yield with proA content >10.0 μg/g were identified. This is the first comprehensive study on development of diverse proA rich maize hybrids through marker-assisted pedigree breeding approach. The findings provides sustainable and cost-effective solution to alleviate vitamin-A deficiency.

A new genetic locus for self-compatibility in the outcrossing grass species perennial ryegrass (Lolium perenne).

BackgroundSelf-incompatibility (SI) is a physiological mechanism that many flowering plants employ to prevent self-fertilization and maintain heterozygosity. In the grass family this is known to be controlled by a two locus (S-Z) system; however, the SI system is intrinsically leaky. Modifier genes of both the S and Z loci and a further locus, T, are known to override SI leading to self-fertilization and self-seed production. This has implications for the ecological and evolutionary success as well as the commercial breeding of grasses. Here we report a study where the genetic control of self-compatibility (SC) was determined from the results of self-pollinating an F2 population of perennial ryegrass from two independently derived inbred lines produced by single-seed descent.Methods In vitro self-pollinations of 73 fertile plants were analysed. A genetic association analysis was made with a panel of 1863 single-nucleotide polymorphism (SNP) markers, generated through genotype-by-sequencing methodology. Markers were placed on a recombination map of seven linkage groups (LGs) created using Joinmap v.5. The seed set on self- and open-pollinated inflorescences was determined on 143 plants, including the 73 plants analysed for self-pollination response.Key ResultsSelf-pollinations revealed a bimodal distribution of percentage SC with peaks at 50 and 100 %. A single quantitative trait locus (QTL) was identified with peak association for marker 6S14665z17875_11873 that mapped to LG 6. Peak position was associated with maximum marker segregation distortion. The self-compatible plants were equally fecund after self- and open pollination.ConclusionsThis is the first report in the Poaceae family of an SC locus located on LG 6. This new SC QTL discovery, as well as indicating the complex nature of the pollen–stigma recognition process and its evolutionary significance, provides an additional source of SC for breeding perennial ryegrass.

Effect of marker segregation distortion on high density linkage map construction and QTL mapping in Soybean (Glycine max L.)

Marker segregation distortion is a natural phenomenon. Severely distorted markers are usually excluded in the construction of linkage maps. We investigated the effect of marker segregation distortion on linkage map construction and quantitative trait locus (QTL) mapping. A total of 519 recombinant inbred lines of soybean from orthogonal and reciprocal crosses between LSZZH and NN493-1 were genotyped by specific length amplified fragment markers and seed linoleic acid content was measured in three environments. As a result, twenty linkage groups were constructed with 11,846 markers, including 1513 (12.77%) significantly distorted markers, on 20 chromosomes, and the map length was 2475.86 cM with an average marker-interval of 0.21 cM. The inclusion of distorted markers in the analysis was shown to not only improve the grouping of the markers from the same chromosomes, and the consistency of linkage maps with genome, but also increase genome coverage by markers. Combining genotypic data from both orthogonal and reciprocal crosses decreased the proportion of distorted markers and then improved the quality of linkage maps. Validation of the linkage maps was confirmed by the high collinearity between positions of markers in the soybean reference genome and in linkage maps and by the high consistency of 24 QTL regions in this study compared with the previously reported QTLs and lipid metabolism related genes. Additionally, linkage maps that include distorted markers could add more information to the outputs from QTL mapping. These results provide important information for linkage mapping, gene cloning and marker-assisted selection in soybean.

QTL Mapping of Important Agronomic Traits on Chromosomes Showing Marker Segregation Distortion in Interspecific Backcross Cotton Populations

QTL Mapping of Important Agronomic Traits on Chromosomes Showing Marker Segregation Distortion in Interspecific Backcross Cotton Populations

Marker assisted backcross breeding of Bg 358 (<em>Oryza sativa</em> L.) for the anaerobic germination tolerant QTL <em>AG1</em>

Sustainability of transplanting systems for rice crop establishment in Sri Lanka is threatened due to climatic changes and heavy rains. Tolerance to anaerobic germination (AG) stress is a key trait that will ensure good crop establishment in direct seeded rice (DSR) farming systems. In this study Marker Assisted backcross breeding was carried out to transfer the quantitative trait loci - QTL AG1 (qAG-9-2), in to the local variety Bg 358 for the development of AG stress tolerant elite lines by AG1 donors IR64-AG1 and Ciherang-Sub1AG1. Marker assisted selection (MAS) was carried out through foreground and recombinant selection. Molecular marker optimization for RM24161, RM8300 and RM553 to the QTL AG1 was carried out using BC1F1 populations. DNA was extracted and PCR was performed using simple sequence repeat (SSR) markers and visualized on 2.5% agarose gel. Based on polymorphism exhibited, RM24161 was chosen as the foreground selection marker, RM553 and RM8300 chosen as proximal and distal markers for recombinant selection. It was suggestive through this study to choose flanking markers RM105 & RM219, closer to the qAG-9-2 locus for recombinant selection as the selected recombinant makers were ineffective and exhibited marker segregation distortion. RM24161 was found to be a tightly linked marker to the AG tolerant QTL AG1 (qAG-9-2) and was successfully used for selection of lines by MAS in generation of anaerobic germination tolerant elite lines.

Hybrid seedling inviability locus (HIs1) mapped on linkage group 4 of the Japanese flowering cherry, Cerasus × yedoensis ‘Somei-yoshino’

Cerasus × yedoensis ‘Somei-yoshino’ (2n = 16), one of the most popular cultivars of Japanese flowering cherry, is considered a hybrid between Cerasus spachiana and Cerasus speciosa. In the present study, we observed segregation of the trait for seedling growth in F1 progenies of Somei-yoshino. In all three families crossed with ancestral wild species, C. spachiana, approximately half of the seedlings showed growth failure. To identify loci involved in seedling inviability, we constructed a genetic linkage map of Somei-yoshino. Both normal growth (N = 77) and growth failure seedlings (N = 101) of an F1 progenies crossed between Somei-yoshino (CY) and C. spachiana (E750) were used for map construction. In accordance to pseudo-testcross strategy, female (CY) and male (E750) linkage maps were constructed. The CY map consists of eight linkage groups with 59 simple sequence repeat (SSR) and 17 amplified fragment length polymorphism (AFLP) markers. The map length is 574.9 cM. In contrast, the E750 map reached 196.8 cM and included 17 SSRs. Subsequently, we explored positions of loci associated to seedling inviability along with linkage maps. Results of segregation distortion of markers, association analysis, and interval mapping consistently showed that an inviability gene was located between two SSR markers, EMPaS13 and BPPCT005 in the fourth linkage group (LG4) of the Somei-yoshino map with 99.6 % explanations for the inviability trait but not present in LG4 of the E750 map. From the genotypes and expressions of seedling inviability, we named the locus Hybrid Inviability of seedlings 1 (HIs1). Finally, HIs1 was mapped to LG4 as Mendelian trait loci and was completely linked with an SSR, EMPaS13. Although the mechanisms of seedling inviability cannot be explained by only one locus model of HIs1, one of the candidate genes involved in hybrid incompatibility between Somei-yoshino and C. spachiana is certainly located on LG4 of the CY map.

Mapping quantitative trait loci in selected breeding populations: A segregation distortion approach.

Quantitative trait locus (QTL) mapping is often conducted in line-crossing experiments where a sample of individuals is randomly selected from a pool of all potential progeny. QTLs detected from such an experiment are important for us to understand the genetic mechanisms governing a complex trait, but may not be directly relevant to plant breeding if they are not detected from the breeding population where selection is targeting for. QTLs segregating in one population may not necessarily segregate in another population. To facilitate marker-assisted selection, QTLs must be detected from the very population which the selection is targeting. However, selected breeding populations often have depleted genetic variation with small population sizes, resulting in low power in detecting useful QTLs. On the other hand, if selection is effective, loci controlling the selected trait will deviate from the expected Mendelian segregation ratio. In this study, we proposed to detect QTLs in selected breeding populations via the detection of marker segregation distortion in either a single population or multiple populations using the same selection scheme. Simulation studies showed that QTL can be detected in strong selected populations with selected population sizes as small as 25 plants. We applied the new method to detect QTLs in two breeding populations of rice selected for high grain yield. Seven QTLs were identified, four of which have been validated in advanced generations in a follow-up study. Cloned genes in the vicinity of the four QTLs were also reported in the literatures. This mapping-by-selection approach provides a new avenue for breeders to improve breeding progress. The new method can be applied to breeding programs not only in rice but also in other agricultural species including crops, trees and animals.

Genetic Map Construction and Detection of Genetic Loci Underlying Segregation Distortion in an Intraspecific Cross of Populus deltoides.

Based on a two-way pseudo-testcross strategy, high density and complete coverage linkage maps were constructed for the maternal and paternal parents of an intraspecific F2 pedigree of Populus deltoides. A total of 1,107 testcross markers were obtained, and the mapping population consisted of 376 progeny. Among these markers, 597 were from the mother, and were assigned into 19 linkage groups, spanning a total genetic distance of 1,940.3 cM. The remaining 519 markers were from the father, and were also were mapped into 19 linkage groups, covering 2,496.3 cM. The genome coverage of both maps was estimated as greater than 99.9% at 20 cM per marker, and the numbers of linkage groups of both maps were in accordance with the 19 haploid chromosomes in Populus. Marker segregation distortion was observed in large contiguous blocks on some of the linkage groups. Subsequently, we mapped the segregation distortion loci in this mapping pedigree. Altogether, eight segregation distortion loci with significant logarithm of odds supports were detected. Segregation distortion indicated the uneven transmission of the alternate alleles from the mapping parents. The corresponding genome regions might contain deleterious genes or be associated with hybridization incompatibility. In addition to the detection of segregation distortion loci, the established genetic maps will serve as a basic resource for mapping genetic loci controlling traits of interest in future studies.

Multi-QTL mapping for quantitative traits using distorted markers

Marker segregation distortion is a common natural phenomenon. However, relatively little is known about utilizing distorted markers for detecting quantitative trait loci (QTL). Therefore, in this study we proposed a multi-QTL mapping approach that uses distorted markers. First, the information from all markers, including distorted markers, was used to detect segregation distortion loci (SDL). Second, the information from the detected SDL was used to correct the conditional probabilities of the QTL genotypes conditional on marker information, and these corrected probabilities were then incorporated into a multi-QTL mapping methodology. Finally, the proposed approach was validated by both Monte Carlo simulation studies and real data analysis. The results from the simulation studies show that as long as one or two SDL are placed around the simulated QTL, there are no differences between the new method and the ordinary interval mapping method in terms of the power of QTL detection or the estimates of the position and dominant effects of the QTL. However, the power of QTL detection is higher under the dominant genetic model of SDL than under the additive genetic model, and the estimate for the additive effect of QTL using the new method is significantly different from the estimate obtained using ordinary interval mapping. The above results were further confirmed by the detection of QTL for dried soymilk in 222 F2:4 families in soybean.

Comparative mapping in watermelon [Citrullus lanatus (Thunb.) Matsum. et Nakai

The first single-nucleotide polymorphism (SNP) maps for watermelon [Citrullus lanatus (Thunb.) Matsum. et Nakai] were constructed and compared. Three populations were developed from crosses between two elite cultivars, Klondike Black Seeded × New Hampshire Midget (KBS × NHM), an elite cultivar and wild egusi accession, Strain II × PI 560023 (SII × Egusi) and an elite cultivar and a wild citron accession, ZWRM50 × PI 244019 (ZWRM × Citroides). The SII × Egusi and ZWRM × Citroides F(2) populations consisted of 187 and 182 individuals respectively while the KBS × NHM recombinant inbred line (RIL) population consisted of 164 lines. The length of the genetic maps were 1,438, 1,514 and 1,144 cM with average marker distances of 3.8, 4.2, and 3.4 cM for the KBS × NHM, SII × Egusi and ZWRM × Citroides populations, respectively. Shared markers were used to align the three maps so that the linkage groups (LGs) represented the 11 chromosomes of the species. Marker segregation distortion were observed in all three populations, but was highest (12.7 %) in the ZWRM × Citroides population, where Citroides alleles were favored. The three maps were used to construct a consensus map containing 378 SNP markers with an average distance of 5.1 cM between markers. Phenotypic data was collected for fruit weight (FWT), fruit length (FL), fruit width (FWD), fruit shape index (FSI), rind thickness (RTH) and Brix (BRX) and analyzed for quantitative trait loci (QTL) associated with these traits. A total of 40 QTL were identified in the three populations, including major QTL for fruit size and shape that were stable across genetic backgrounds and environments. The present study reports the first SNP maps for Citrullus and the first map constructed using two elite parents. We also report the first stable QTL associated with fruit size and shape in Citrullus lanatus. These maps, QTL and SNPs should be useful for the watermelon community and represent a significant step towards the potential use of molecular tools in watermelon breeding.

Localization of QTLs for in vitroplant regeneration in tomato

BackgroundLow regeneration ability limits biotechnological breeding approaches. The influence of genotype in the regeneration response is high in both tomato and other important crops. Despite the various studies that have been carried out on regeneration genetics, little is known about the key genes involved in this process. The aim of this study was to localize the genetic factors affecting regeneration in tomato.ResultsWe developed two mapping populations (F2 and BC1) derived from a previously selected tomato cultivar (cv. Anl27) with low regeneration ability and a high regeneration accession of the wild species Solanum pennellii (PE-47). The phenotypic assay indicated dominance for bud induction and additive effects for both the percentage of explants with shoots and the number of regenerated shoots per explant. Two linkage maps were developed and six QTLs were identified on five chromosomes (1, 3, 4, 7 and 8) in the BC1 population by means of the Interval Mapping and restricted Multiple QTL Mapping methods. These QTLs came from S. pennellii, with the exception of the minor QTL located on chromosome 8, which was provided by cv. Anl27. The main QTLs correspond to those detected on chromosomes 1 and 7. In the F2 population, a QTL on chromosome 7 was identified on a similar region as that detected in the BC1 population. Marker segregation distortion was observed in this population in those areas where the QTLs of BC1 were detected. Furthermore, we located two tomato candidate genes using a marker linked to the high regeneration gene: Rg-2 (a putative allele of Rg-1) and LESK1, which encodes a serine/threonine kinase and was proposed as a marker for regeneration competence. As a result, we located a putative allele of Rg-2 in the QTL detected on chromosome 3 that we named Rg-3. LESK1, which is also situated on chromosome 3, is outside Rg-3. In a preliminary exploration of the detected QTL peaks, we found several genes that may be related to regeneration.ConclusionsIn this study we have identified new QTLs related to the complex process of regeneration from tissue culture. We have also located two candidate genes, discovering a putative allele of the high regeneration gene Rg-1 in the QTL on chromosome 3. The identified QTLs could represent a significant step toward the understanding of this process and the identification of other related candidate genes. It will also most likely facilitate the development of molecular markers for use in gene isolation.

Quantitative Trait Locus Mapping Can Benefit From Segregation Distortion

Segregation distortion is a phenomenon that has been observed in many experimental systems. How segregation distortion among markers arises and its impact on mapping studies are the focus of this work. Segregation distortion of markers can be considered to arise from segregation distortion loci (SDL). I develop a theory of segregation distortion and show that the presence of only a few SDL can cause the entire chromosome to distort from Mendelian segregation. Segregation distortion is detrimental to the power of detecting quantitative trait loci (QTL) with dominance effects, but it is not always a detriment to QTL mapping for additive effects. When segregation distortion of a locus is a random event, the SDL is beneficial to QTL mapping approximately 44% of the time. If SDL are present and ignored, power loss can be substantial. A dense marker map can be used to ameliorate the situation, and if dense marker information is incorporated, power loss is minimal. However, other situations are less benign. A method that can simultaneously map QTL and SDL is discussed, maximizing both use of mapping resources and use by agricultural and evolutionary biologists.

Construction of a genetic map based on ILP markers in rice

Intron length polymorphism (ILP) is a new type of PCR-based molecular markers with many advantages. We had previously developed 172 ILP markers in rice using the published genome sequence data of indica cultivar 93-11 and ja-ponica cultivar Nipponbare. In order to examine the reliability and the applicability of these ILP markers to genetic map-ping, we constructed a rice genetic map consisting of 172 ILP and 13 SSR markers with a total length of 1 905.7 cM using a BC1F1 (Nipponbare/93-11//Nipponbare) population. Comparison showed that the order of the markers in this map was ex-actly the same as their physical order, verifying the feasibility and efficiency of using ILP markers for genetic mapping. We also investigated the phenomenon of marker segregation distortion and found a region of serious segregation distortion in the short arm of chromosome 6.

Refined mapping of the Pierce’s disease resistance locus, PdR1, and Sex on an extended genetic map of Vitis rupestris × V. arizonica

A framework genetic map based on genomic DNA-derived SSR, EST-derived SSR, EST-STS and EST-RFLP markers was developed using 181 genotypes generated from D8909-15 (female) x F8909-17 (male), the '9621' population. Both parents are half siblings with a common female parent, Vitis rupestris 'A. de Serres', and different male parents (forms of V. arizonica). A total of 542 markers were tested, and 237 of them were polymorphic for the female and male parents. The female map was developed with 159 mapped markers covering 865.0 cM with an average marker distance of 5.4 cM in 18 linkage groups. The male map was constructed with 158 mapped molecular markers covering 1055.0 cM with an average distance of 6.7 cM in 19 linkage groups. The consensus '9621' map covered 1154.0 cM with 210 mapped molecular markers in 19 linkage groups, with average distance of 5.5 cM. Ninety-four of the 210 markers on the consensus map were new. The 'Sex' expression locus segregated as single major gene was mapped to linkage group 2 on the consensus and the male map. PdR1, a major gene for resistance to Pierce's disease, caused by the bacterium Xylella fastidiosa, was mapped to the linkage group 14 between markers VMCNg3h8 and VVIN64, located 4.3 and 2.7 cM away from PdR1, respectively. Differences in segregation distortion of markers were also compared between parents, and three clusters of skewed markers were observed on linkage groups 6, 7 and 14.

Improving QTL Mapping Resolution Based on Genotypic Sampling—a Case Using a RIL Population

The QTL mapping results were compared with the genotypically selected and random samples of the same size on the base of a RIL population. The results demonstrated that there were no obvious differences in the trait distribution and marker segregation distortion between the genotypically selected and random samples with the same population size. However, a significant increase in QTL detection power, sensitivity, specificity, and QTL resolution in the genotypically selected samples were observed. Moreover, the highly significant effect was detected in small size of genotypically selected samples. In QTL mapping, phenotyping is a more sensitive limiting factor than genotyping so that the selection of samples could be an attractive strategy for increasing genome-wide QTL mapping resolution. The efficient selection of samples should be more helpful for QTL maker assistant selection, fine mapping, and QTL cloning.

Recent approaches into the genetic basis of inbreeding depression in plants.

Predictions for the evolution of mating systems and genetic load vary, depending on the genetic basis of inbreeding depression (dominance versus overdominance, epistasis and the relative frequencies of genes of large and small effect). A distinction between the dominance and overdominance hypotheses is that deleterious recessive mutations should be purged in inbreeding populations. Comparative studies of populations differing in their level of inbreeding and experimental approaches that allow selection among inbred lines support this prediction. More direct biometric approaches provide strong support for the importance of partly recessive deleterious alleles. Investigators using molecular markers to study quantitative trait loci (QTL) often find support for overdominance, though pseudo-overdominance (deleterious alleles linked in repulsion) may bias this perception. QTL and biometric studies of inbred lines often find evidence for epistasis, which may also contribute to the perception of overdominance, though this may be because of the divergent lines initially crossed in QTL studies. Studies of marker segregation distortion commonly uncover genes of major effect on viability, but these have only minor contributions to inbreeding depression. Although considerable progress has been made in understanding the genetic basis of inbreeding depression, we feel that all three aspects merit more study in natural plant populations.

Construction of a RFLP genetic linkage map for oil palm (Elaeis guineensisJacq.)

We have developed a restriction fragment length polymorphism (RFLP) genetic map in oil palm (Elaeis guineensis Jacq.) for use in breeding programmes. A segregating population of 98 individuals was probed with 84 informative low copy clones (mainly PstI genomics). This yielded 103 scorable loci, of which 97 could be linked into 24 groups of two or more markers (n = 16 for oil palm), encompassing a total of 860 cM. The high level of linkage between markers (95%) suggests good genome coverage and very little segregation distortion of markers was observed. The mapping population, which was generated by the selfing of an important breeding palm (A 137/30), also segregated for the shell thickness character (Sh), enabling mapping of the gene conferring this major commercial trait. The nearest RFLP marker (pOPgSP1282) was located 9.8 cM from Sh in the mapping population and 6.6 cM away in a related (A137/30 x E80/29) smaller population (45 palms). Strategies for conversion of RFLP markers to a PCR format were explored and two important markers were shown to generate allelic variants following PCR amplification and restriction digestion of PCR products. The application of this map to oil palm breeding programmes is discussed.

Chromosomal regions associated with segregation distortion of molecular markers in F2, backcross, doubled haploid, and recombinant inbred populations in rice (Oryza sativa L.).

Chromosomal regions associated with marker segregation distortion in rice were compared based on six molecular linkage maps. Mapping populations were derived from one interspecific backcross and five inter-subspecific (indica/japonica) crosses, including two F2 populations, two doubled haploid (DH) populations, and one recombinant inbred (RI) population. Mapping data for each population consisted of 129-629 markers. Segregation distortion was determined based on chi-square analysis (P < 0.01) and was observed at 6.8-31.8%, of the mapped marker loci. Marker loci associated with skewed allele frequencies were distributed on all 12 chromosomes. Distortion in eight chromosomal regions bracketed previously identified gametophyte (ga) or sterility genes (S). Distortion in three other chromosomal regions was found only in DH populations, where japonica alleles were over-represented, suggesting that loci in these regions may be associated with preferential regeneration of japonica genotypes during anther culture. Three additional clusters of skewed markers were observed in more than one population in regions where no gametophytic or sterility loci have previously been reported. A total of 17 segregation distortion loci may be postulated based on this study and their locations in the rice genome were estimated.