Genetic basis and mapping of the resistance to rice yellow mottle virus. II. Evidence of a complementary epistasis between two QTLs

 The genetic basis of resistance to rice yellow mottle virus (RYMV) was studied in a doubled-haploid (DH) population derived from a cross between the very susceptible indica variety ‘IR64’ and the resistant upland japonica variety Azucena. As a quantitative trait locus (QTL) involved in virus content estimated with an ELISA test has been previously identified on chromosome 12, we performed a wide search for interactions between this QTL and the rest of the genome, and between this QTL and morphological traits segregating in the population. Multiple regression with all identified genetic factors was used to validate the interactions. Significant epistasis accounting for a major part of the total genetic variation was observed. A complementary epistasis between the QTL located on chromosome 12 and a QTL located on chromosome 7 could be the major genetic factor controlling the virus content. Resistance was also affected by a morphology-dependent mechanism since tillering was interfering with the resistance mechanism conditioned by the epistasis between the two QTLs. Marker-assisted backcross breeding was developed to introgress the QTLs of chromosome 7 and chromosome 12 in the susceptible ‘IR64’ genetic background. First results confirmed that if both QTLs do not segregate in a backcross-derived F₂ population, then the QTL of chromosome 12 cannot explain differences in virus content. A near-isogenic line (NIL) approach is currently being developed to confirm the proposed genetic model of resistance to RYMV. Received: 20 April 1990 / Accepted: 30 April 1998     

Genetic basis and mapping of the resistance to rice yellow mottle virus. I. QTLs identification and relationship between resistance and plant morphology

 Rice yellow mottle virus (RYMV) resistance QTLs were mapped in a doubled-haploid population of rice, ‘IR64/Azucena’. Disease impact on plant morphology and development, expression of symptoms and virus content were evaluated in field conditions. Virus content was also assessed in a growth chamber. RYMV resistance was found to be under a polygenic determinism, and 15 QTLs were detected on seven chromosomal fragments. For all of the resistance QTLs detected, the favourable allele was provided by the resistant parent ‘Azucena’. Three regions were determined using different resistance parameters and in two environments. On chromosome 12, a QTL of resistance that had already been detected in this population and another indica/japonica population was confirmed both in the field and under controlled conditions. Significant correlations were observed between resistance and tillering ability, as measured on control non-inoculated plants. In addition, the three genomic fragments involved in resistance were also involved in plant architecture and development. In particular, the semi-dwarfing gene sd-1, on chromosome 1, provided by the susceptible parent, ‘IR64’, mapped in a region where resistance QTLs were detected with most of the resistance parameters. In contrast, the QTL of resistance mapped on chromosome 12 was found to be independent of plant morphology. Received: 20 April 1998 / Accepted: 30 April 1998     

Three QTLs from Lycopersicon peruvianum confer a high level of resistance to Clavibactermichiganensis ssp. michiganensis

Lycopersicon peruvianum LA2157 originates from 1650 m above sea level and harbours several beneficial traits for cultivated tomatoes such as cold tolerance, nematode resistance and resistance to bacterial canker (Clavibacter michiganensis ssp. michiganensis). In order to identify quantitative trait loci (QTLs) for bacterial canker resistance, a QTL mapping approach was carried out in an F₂ population derived from the interspecific F₁ between Lycopersicon esculentum cv Solentos and L. peruvianum LA2157. Three QTLs for resistance mapped to chromosomes 5, 7 and 9 respectively. The resistance loci were additive and co-dominant with the QTL on chromosome 7 explaining the largest part of the variation for resistance in the F₂ population. The combination of this QTL with either of the other two QTLs conferred a resistance similar to the level in the resistant parent L. peruvianum. Some RFLP markers flanking this QTL on chromosome 7 were converted into SCAR markers allowing efficient marker-assisted selection of plants with high resistance to bacterial canker. Received: 26 February 1999 / Accepted: 12 March 1999     

Identification of QTL regions and SSR markers associated with resistance to reniform nematode in <Emphasis Type="Italic">Gossypium barbadense</Emphasis> L. accession GB713

The identification of molecular markers that are closely linked to gene(s) in Gossypium barbadense L. accession GB713 that confer a high level of resistance to reniform nematode (RN), Rotylenchulus reniformis Linford & Oliveira, would be very useful in cotton breeding programs. Our objectives were to determine the inheritance of RN resistance in the accession GB713, to identify SSR markers linked with RN resistance QTLs, and to map these linked markers to specific chromosomes. We grew and scored plants for RN reproduction in the P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂ generations from the cross of GB713 × Acala Nem-X. The generation means analysis using the six generations indicated that one or more genes were involved in the RN resistance of GB713. The interspecific F₂ population of 300 plants was genotyped with SSR molecular markers that covered most of the chromosomes of Upland cotton (G. hirsutum L.). Results showed two QTLs on chromosome 21 and one QTL on chromosome 18. One QTL on chromosome 21 was at map position 168.6 (LOD 28.0) flanked by SSR markers, BNL 1551_162 and GH 132_199 at positions 154.2 and 177.3, respectively. A second QTL on chromosome 21 was at map position 182.7 (LOD 24.6) flanked by SSR markers BNL 4011_155 and BNL 3279_106 at positions 180.6 and 184.5, respectively. Our chromosome 21 map had 61 SSR markers covering 219 cM. One QTL with smaller genetic effects was localized to chromosome 18 at map position 39.6 (LOD 4.0) and flanked by SSR markers BNL 1721_178 and BNL 569_131 at positions 27.6 and 42.9, respectively. The two QTLs on chromosome 21 had significant additive and dominance effects, which were about equal for each QTL. The QTL on chromosome 18 showed larger additive than dominance effects. Following the precedent set by the naming of the G. longicalyx Hutchinson & Lee and G. aridum [(Rose & Standley) Skovsted] sources of resistance, we suggest the usage of Ren ^barb1 and Ren ^barb2 to designate these QTLs on chromosome 21 and Ren ^barb3 on chromosome 18.     

QTL analysis and mapping of pi21, a recessive gene for field resistance to rice blast in Japanese upland rice

Field resistance is defined as the resistance that allows effective control of a parasite under natural field conditions and is durable when exposed to new races of that parasite. To identify the genes for field resistance to rice blast, quantitative trait loci (QTLs) conferring field resistance to rice blast in Japanese upland rice were detected and mapped using RFLP and SSR markers. QTL analysis was carried out in F₄ progeny lines from the cross between Nipponbare (moderately susceptible, lowland) and Owarihatamochi (resistant, upland). Two QTLs were detected on chromosome 4 and one QTL was detected on each of chromosomes 9 and 12. The phenotypic variation explained by each QTL ranged from 7.9 to 45.7% and the four QTLs explained 66.3% of the total phenotypic variation. Backcrossed progeny lines were developed to transfer the QTL with largest effect using the susceptible cultivar Aichiasahi as a recurrent parent. Among 82 F₃ lines derived from the backcross, resistance segregated in the expected ratio of resistant 1 : heterozygous 2 : susceptible 1. The average score for blast resistance measured in the field was 4.2 ± 0.67, 7.5 ± 0.51and 8.2 ± 0.66, for resistant, heterozygous and susceptible groups, respectively. The resistance gene, designated pi21, was mapped on chromosome 4 as a single recessive gene between RFLP marker loci G271 and G317 at a distance of 5.0 cM and 8.5 cM, respectively. The relationship to previously reported major genes and QTLs conferring resistance to blasts, and the significance of marker-assisted selection to improve field resistance, are discussed. Received: 8 June 2000 / Accepted: 24 November 2000     

Mapping quantitative trait loci (QTLs) for resistance to Cercospora leaf spot disease (Cercospora beticola Sacc.) in sugar beet (Beta vulgaris L.)

The breeding of sugar beet varieties that combine resistance to Cercospora and high yield under non-diseased conditions is a major challenge to the breeder. The understanding of the quantitative trait loci (QTLs) contributing to Cercospora resistance offers one route to solving this problem. A QTL analysis of Cercospora resistance in sugar beet was carried out using a linkage map based on AFLP and RFLP markers. Two different screening methods for Cercospora resistance (a field test at Copparo, Italy, under natural infection, and a newly-developed leaf disc test) were used to estimate the level of Cercospora resistance; the correlation between scores from the field (at 162 days after sowing) and the leaf disc test was significant. QTL analysis was based on F₂ and F₃ (half-sib family) generations derived from crosses between diploid single plants of 93164P (resistant to Cercospora leaf spot disease) and 95098P (susceptible). Four QTLs associated with Cercospora resistance (based on Lsmean data of the leaf disc test) on chromosomes III, IV, VII and IX were revealed using Composite interval mapping. To produce populations segregating for leaf spot resistance as a single Mendelian factor, we selected for plants heterozygous for only one of the QTLs (on chromosome IV or IX) but homozygous for the others. Received: 1 September 1999 / Accepted 7 October 1999     

Fine mapping of quantitative trait loci Hd-1, Hd-2 and Hd-3, controlling heading date of rice, as single Mendelian factors

 Fine mapping was carried out on three putative QTLs (tentatively designated as Hd-1 to Hd-3) of five such QTLs controlling heading date in rice that had been earlier identified using an F₂ population derived from a cross between a japonica variety, ‘Nipponbare’, and an indica variety, ‘Kasalath’, using progeny backcrossed with ‘Nipponbare’ as the recurrent parent. One BC₃F₂ and two BC₃F₁ plants, in which the target QTL regions were heterozygous and most other chromosomal regions were homozygous for the ‘Nipponbare’ allele, were selected as the experimental material. Self-pollinated progeny (BC₃F₂ and BC₃F₃) of the BC₃F₁ or BC₃F₂ showed continuous variation in days to heading. By means of progeny testing based on BC₃F₃ or BC₃F₄ lines, we determined the genotypes of each BC₃F₂ or BC₃F₃ individual at target QTLs. Their segregation patterns fitted Mendelian inheritance ratios. When the results obtained by RFLP analysis and progeny tests were combined, Hd-1, Hd-2 and Hd-3 were mapped precisely on chromosomes 6, 7 and 6, respectively, of a rice RFLP linkage map. The results demonstrated that QTLs can be treated as Mendelian factors. Moreover, these precise locations were in good agreement with the regions estimated by QTL analysis of the initial F₂ population, demonstrating the high reliability of QTL mapping using a high-density linkage map. Received: 5 November 1997 / Accepted: 10 February 1998     

Identification of a new major QTL associated with resistance to soybean cyst nematode (Heterodera glycines)

Resistance of soybean [Glycine max (L.) Merr.] to cyst nematode (SCN) (Heterodera glycines Ichinohe), one of the most destructive pathogens affecting soybean, involves a complex genetic system. The identification of QTLs associated with SCN resistance may contribute to the understanding of such system. The objective of this work was to identify and map QTLs for resistance to SCN Race 14 with the aid of molecular markers. BC₃F_2:3 and F_2:3 populations, both derived from an original cross between resistant cv. Hartwig and the susceptible line BR-92–31983 were screened for resistance to SCN Race 14. Four microsatellite (Satt082, Sat_001, Satt574 and Satt301) and four RAPD markers (OPAA-11₇₉₅, OPAE-08₈₃₇, OPR-07₅₄₈ and OPY-07₂₀₃₀) were identified in the BC₃F_2:3 population using the bulked segregant analysis (BSA) technique. These markers were amplified in 183 F_2:3 families and mapped to a locus that accounts for more than 40% of the resistance to SCN Race 14. Selection efficiency based on these markers was similar to that obtained with the conventional method. In the case of the microsalellite markers, which identify homozygous resistant genotypes, the efficiency was even higher. This new QTL has been mapped to the soybean linkage group D2 and, in conjunction with other QTLs already identified for SCN resistance, will certainly contribute to our understanding of the genetic basis of resistance of this important disease in soybean. Received: 12 October 1999 / Accepted: 14 April 2000     

SSR mapping and confirmation of the QTL from PI96354 conditioning soybean resistance to southern root-knot nematode

Root-knot nematodes (Meloidogyne spp.) can cause severe yield loss of soybean [Glycine max (L.) Merr.] in the southern production region of the USA. Planting root-knot nematode-resistant cultivars is the most effective method of preventing yield loss. DNA marker-assisted breeding may accelerate the development of root-knot nematode-resistant cultivars. RFLP markers have previously been used to identify quantitative trait loci (QTLs) conferring resistance to southern root-knot nematode [Meloidogyne incognita (Kofoid and White) Chitwood] (Mi) in a F_2:3 soybean population created by crossing the resistant PI96354 and the susceptible ’Bossier.’ A major QTL on linkage group (LG) O conditioning 31% of the variation in Mi gall number and a minor QTL on LG-G conditioning 14% of the gall variation were reported. With the development of SSR markers for soybean improvement, a higher level of mapping resolution and semi-automated detection has become possible. The objectives of this research were: (1) to increase the marker density in the genomic regions of the QTLs for Mi resistance on LG-O and LG-G with SSR markers; and (2) to confirm the effect of the QTLs in a second population and a different genetic background. With SSR markers, the QTL on LG-O was flanked by Satt492 and Satt358, and on LG-G by Satt012 and Satt505. Utilizing SSR markers flanking the two QTLs, marker-assisted selection was performed in a second F_2:3 population of PI96354× Bossier. Results confirmed the effectiveness of marker-assisted selection to predict the Mi phenotypes. By screening the BC₂F₂ population of Prichard (3)×G93–9009 we confirmed that selection for the minor QTL on LG-G with flanking SSR markers would enhance the resistance of lines containing the major QTL (which is most-likely Rmi1). Received: 29 September 2000 / Accepted: 17 April 2001     

Marker-assisted introgression of blackmold resistance QTL alleles from wild Lycopersicon cheesmanii to cultivated tomato (L. esculentum) and evaluation of QTL phenotypic effects

Blackmold, caused by the fungus Alternaria alternata, is a major ripe fruit disease of processing tomatoes. Previously, we found blackmold resistance in a wild tomato (Lycopersicon cheesmanii) and quantitative trait loci (QTL) for resistance were mapped in an interspecific population. Five QTLs were selected for introgression from L. cheesmanii into cultivated tomato using marker-assisted selection (MAS). Restriction fragment length polymorphism and PCR-based markers flanking, and within, the chromosomal regions containing QTLs were used for MAS during backcross and selfing generations. BC₁ plants heterozygous at the QTLs, and subsequent BC₁S₁ and BC₁S₂ lines possessing different homozygous combinations of alleles at the target QTLs, were identified using DNA markers. Field experiments were conducted in 1998 (with 80 marker-selected BC₁S₂ lines) and 1999 (with 151 marker-selected BC₁S₂ and BC₁S₃ lines) at three California locations. Blackmold resistance was assessed during both years, and horticultural traits were evaluated in 1999. The BC₁S₂ and BC₁S₃ lines containing L. cheesmanii alleles at the QTLs were associated with a large genetic variance for resistance to blackmold and moderate heritability, suggesting that significant genetic gain may be achieved by selection in this genetic material. L. cheesmanii alleles at three of the five introgressed QTLs showed a significant, positive effect on blackmold resistance. A QTL on chromosome 2 had the largest positive effect on blackmold resistance, alone and in combination with other QTLs, and was also associated with earliness, a positive horticultural trait. The other four QTLs were associated primarily with negative horticultural traits. Fine mapping QTLs using near isogenic lines could help determine if such trait associations are due to linkage drag or pleiotropy.     

Mapping quantitative trait loci controlling seed dormancy and heading date in rice, Oryza sativa L., using backcross inbred lines

 To detect quantitative trait loci (QTLs) controlling seed dormancy, 98 BC₁F₅ lines (backcross inbred lines) derived from a backcross of Nipponbare (japonica)/Kasalath (indica)//Nipponbare were analyzed genetically. We used 245 RFLP markers to construct a framework linkage map. Five putative QTLs affecting seed dormancy were detected on chromosomes 3, 5, 7 (two regions) and 8, respectively. Phenotypic variations explained by each QTL ranged from 6.7% to 22.5% and the five putative QTLs explained about 48% of the total phenotypic variation in the BC₁F₅ lines. Except for those of the QTLs on chromosome 8, the Nipponbare alleles increased the germination rate. Five putative QTLs controlling heading date were detected on chromosomes 2, 3, 4, 6 and 7, respectively. The phenotypic variation explained by each QTL for heading date ranged from 5.7% to 23.4% and the five putative QTLs explained about 52% of the total phenotypic variation. The Nipponbare alleles increased the number of days to heading, except for those of two QTLs on chromosomes 2 and 3. The map location of a putative QTL for heading date coincided with that of a major QTL for seed dormancy on chromosome 3, although two major heading-date QTLs did not coincide with any seed dormancy QTLs detected in this study. Received: 10 October 1997 / Accepted: 12 January 1998     

Identification of QTLs involved in the resistance to South American leaf blight (Microcyclus ulei) in the rubber tree

South American leaf blight (SALB) is a disease of the rubber tree caused by the fungus Microcyclus ulei. Quantitative trait loci (QTLs) for resistance were mapped using 195 F₁ progeny individuals derived from the cross between a susceptible cultivated clone, PB260, and a resistant clone, RO38, derived from interspecific hybridization. The resistance level of the progeny individuals was evaluated in controlled conditions. The reaction type (RT) and the lesion diameter (LD) were measured on immature leaves after artificial inoculation of the fungus. Five different strains of the fungus were used, all highly sporulating on PB260. Among those, four did not sporulate and one sporulated partially on RO38. Both pseudo-testcross parental genetic maps and the consensus map were constructed. The search for QTLs was performed using the Kruskal-Wallis marker-by-marker test and the Interval-Mapping method for the three maps. Eight QTLs for resistance were identified on the RO38 map. Only one QTL was detected on the PB260 map. The analysis of the F₁ consensus map confirmed results obtained with the parental maps. A common QTL was detected for resistance to the five strains for both RT and LD. Two QTLs were common for complete resistance to four strains, for RT and LD respectively. Resistance determinism for complete and partial resistance, and perspectives for breeding for durable resistance to SALB are discussed. Received: 1 August 1999 / Accepted: 27 August 1999     

QTL analysis of an advanced backcross of Lycopersicon peruvianum to the cultivated tomato and comparisons with QTLs found in other wild species

 A BC₃ population previously developed from a backcross of Lycopersicon peruvianum, a wild relative of tomato, into the cultivated variety L. esculentum was analyzed for QTLs. Approximately 200 BC₄ families were scored for 35 traits in four locations worldwide. One hundred and sixty-six QTLs were detected for 29 of those traits. For more than half of those 29 traits at least 1 QTL was detected for which the presence of the wild allele was associated with an agronomically beneficial effect despite the inferior phenotype of the wild parent. Eight QTLs for fruit weight could be followed through the BC₂, BC₃, and BC₄, generations, supporting the authenticity of these QTLs. Comparisons were made between the QTLs found in this study and those found in studies involving two other wild species; the results showed that while some of these QTLs can be presumed to be allelic, most of the QTLs detected in this study are ones not previously discovered. Received: 9 April 1997 / Accepted: 20 May 1997     

Nature and type of gene action governing resistance to Helminthosporium turcicum leaf blight in maize (Zea mays L.)

Six generations, consisting of three resistant parents, three susceptible parents, their 15 possible F₁ crosses, 15 F₂'s, 15 BC₁'s (F₁ x resistant female parent) and 15 BC₂'s (F₁ x susceptible male parent) were analysed following Hayman (Heredity 12: 371–390, 1958) to evaluate the nature and type of gene action governing resistance to H. turcicum. The results showed that all types of gene effects, viz., additive, dominance and epistasis (i.e., additive x additive, additive x dominance and dominance x dominance) were operating in one cross or the other in controlling resistance. However, it was additive gene action and dominance x dominance type of epistasis with duplicate nature that were important in controlling resistance in most crosses. Depending upon the final objectives, one of the breeding methods, viz., recurrent selection, heterosis breeding, back-cross method or full-sib selection (bi-parental mating) may be followed.     

A major QTL for resistance to Gibberella stalk rot in maize

Fusarium graminearum Schwabe, the conidial form of Gibberella zeae, is the causal fungal pathogen responsible for Gibberella stalk rot of maize. Using a BC₁F₁ backcross mapping population derived from a cross between ‘1145’ (donor parent, completely resistant) and ‘Y331’ (recurrent parent, highly susceptible), two quantitative trait loci (QTLs), qRfg1 and qRfg2, conferring resistance to Gibberella stalk rot have been detected. The major QTL qRfg1 was further confirmed in the double haploid, F₂, BC₂F₁, and BC₃F₁ populations. Within a qRfg1 confidence interval, single/low-copy bacterial artificial chromosome sequences, anchored expressed sequence tags, and insertion/deletion polymorphisms, were exploited to develop 59 markers to saturate the qRfg1 region. A step by step narrowing-down strategy was adopted to pursue fine mapping of the qRfg1 locus. Recombinants within the qRfg1 region, screened from each backcross generation, were backcrossed to ‘Y331’ to produce the next backcross progenies. These progenies were individually genotyped and evaluated for resistance to Gibberella stalk rot. Significant (or no significant) difference in resistance reactions between homozygous and heterozygous genotypes in backcross progeny suggested presence (or absence) of qRfg1 in ‘1145’ donor fragments. The phenotypes were compared to sizes of donor fragments among recombinants to delimit the qRfg1 region. Sequential fine mapping of BC₄F₁ to BC₆F₁ generations enabled us to progressively refine the qRfg1 locus to a ~500-kb interval flanked by the markers SSR334 and SSR58. Meanwhile, resistance of qRfg1 to Gibberella stalk rot was also investigated in BC₃F₁ to BC₆F₁ generations. Once introgressed into the ‘Y331’ genome, the qRfg1 locus could steadily enhance the frequency of resistant plants by 32–43%. Hence, the qRfg1 locus was capable of improving maize resistance to Gibberella stalk rot.     

Potential of doubled-haploid lines and localization of quantitative trait loci (QTL) for partial resistance to bacterial leaf streak (Xanthomonas campestris pv. hordei) in barley

 Genetic variability for partial resistance to bacterial leaf streak in barley, caused by Xanthomonas campestris pv. hordei, was investigated in 119 doubled-haploid lines (DH) developed by the Hordeum bulbosum method from the F₁ progeny of the cross between two cultivars, ‘Morex’ (resistant) and ‘Steptoe’ (susceptible). Two experiments were undertaken in a randomized complete block design with three replicates, in a controlled growth chamber. Twenty seeds per replicate were planted in plastic containers (60×40×8 cm) containing moistened vermiculite. At the two-leaf stage seedlings were inoculated with an Iranian strain of the pathogen. Genetic variability was observed among the 119 DH lines for partial resistance to the disease. Some DH lines were significantly more resistant than ‘Morex’ (resistant parent) to bacterial leaf streak. Genetic gain in percentage of resistant parent for 5% of the selected DH lines was significant (47.70% and 33.72% in the first and the second experiment, respectively). A QTL analysis of bacterial leaf streak resistance showed that three QTLs were detected on chromosomes 3 and 7. Multilocus allelic effects of the three QTLs account for almost 54% of the mean difference between the parents and nearly 30% of the phenotypic variation of the trait in the mean experiment. The resistance locus on chromosome 3, near ABG377, apprears to be a major gene. Received: 15 July 1997 / Accepted: 4 August 1997     

Inheritance and genetic mapping of cucumber mosaic virus resistance introgressed from Lycopersicon chilense into tomato

Cucumber mosaic virus (CMV) infects a wide variety of crop plants and in tomato (Lycopersicon esculentum Mill.) causes significant economic losses in many growing regions, particularly the Mediterranean. The objective of the present study was to identify the number and map locations of genes controlling resistance to CMV in breeding lines (BC₁–inbreds) derived from the related wild species L. chilense. These lines also carried the gene Tm-2 ^a for resistance to ToMV, which facilitated the interpretation of disease symptoms. The segregation for CMV resistance in the BC₂F₁ and BC₂F₂ generations, following mechanical inoculation with subgroup-I isolates, was consistent with expectations for a single dominant gene, for which the symbol Cmr (cucumber mosaic resistance) was given. Resistant and susceptible BC₁-inbreds were analyzed with RFLP and isozyme markers to identify genomic regions introgressed from L. chilense. The only L. chilense-specific markers found were on chromosome 12; some resistant lines contained a single introgression comprising the entire short arm and part of the long arm of this chromosome, while others contained a recombinant derivative of this introgression. The chromosome 12 markers were significantly associated with CMV resistance in both qualitative and quantitative models of inheritance. The qualitative analysis, however, demonstrated that CMV resistance was not expressed as a reliable monogenic character, suggesting a lack of penetrance, significant environmental effects, or the existence of additional (undetected) resistance factors. In the quantitative analysis, the marker interval TG68 – CT79 showed the most significant association with CMV resistance. No association between CMV resistance and the Tm-2 ^a gene was observed. These breeding lines are potentially useful sources of CMV resistance for tomato improvement, in which context knowledge of the map location of Cmr should accelerate introgression by marker-assisted selection. Received: 9 August 1999 / Accepted: 22 December 1999     

Mapping of one major gene and of QTLs involved in resistance to clubroot in Brassica napus

Clubroot, caused by Plasmodiophora brassicae, is a damaging disease of Brassica napus. Genetic control and mapping of loci involved in high and partial quantitative resistance expressed against two single spore isolates (Pb137–522 and K92–16) were studied in the F₁ and DH progenies of the cross Darmor-bzh (resistant) x Yudal (susceptible). The high level of resistance expressed by Darmor-bzh to isolate Pb137–522 was found to be mainly due to a major gene, which we have named Pb-Bn1, located on linkage group (LG) DY4. Partial quantitative resistance showed by Darmor-bzh to the K92–16 isolate arose from the association of at least two additive QTLs detected on LGs DY4 and DY15; the QTL on DY4, explaining 19% of the variance, was mapped at the same position as the major gene Pb-Bn1. Epistatic interactions between nine regions with or without additive effects were detected. The total phenotypic variation accounted for by additive and epistatic QTLs ranged from 62% to 81% depending on the isolate. For one isolate, the relative effect due to additivity was similar to that due to epistasis. Received: 10 November 1999 / Accepted:18 February 2000     

QTLs for a component of partial resistance to cucumber mosaic virus in pepper: restriction of virus installation in host-cells

 Ninety four doubled-haploid (DH) lines obtained from the F₁ between Perennial, a cucumber mosaic virus (CMV)-partially resistant Capsicum annuum line, and Yolo Wonder, a CMV-susceptible C. annuum line, were analysed with 138 markers including mostly RFLPs and RAPDs. Clustering of RAPD markers was observed on five linkage groups of the intraspecific linkage map. These clusters could correspond to the centromeric regions of pepper chromosomes. The same progenies were evaluated for restriction of CMV installation in pepper cells in order to map quantitative trait loci (QTLs) controlling CMV resistance. This component of partial resistance to CMV was quantitatively assessed using a CMV strain that induced necrotic local lesions on the inoculated leaves. The number of local lesions gave an estimation of the density of the virus-infection sites. Genotypic variance among the DH lines was highly significant for the number of local lesions, and heritability was estimated to be 0.94. Using both analysis of variance and non-parametric tests, three genomic regions significantly affecting CMV resistance were detected on chromosomes Noir, Pourpre and linkage group 3, together explaining 57% of the phenotypic variation. A digenic epistasis between one locus that controlled significant trait variation and a second locus that by itself had no demonstrable effect on the trait was found to have an effect on CMV resistance. For each QTL, the allele from Perennial was associated with an increased resistance. Implications of QTL mapping in marker-based breeding for CMV resistance are discussed. Received: 16 September 1996     

Identification and Validation of Quantitative Trait Loci for Agronomic Traits in Advanced Backcross Breeding Lines Derived from Oryza rufipogon × Oryza sativa Cultivar MR219

A backcross breeding strategy was used to identify quantitative trait loci (QTLs) associated with 14 traits in a BC₂F₂ population derived from a cross between MR219, an indica rice cultivar and an accession of Oryza rufipogon (IRGC 105491). A total of 261 lines were genotyped with 96 microsatellite markers and evaluated for plant morphology, yield components and growth period. The genetic linkage map generated for this population with an average interval size of 16.2?cM, spanning 1,553.4?cM (Kosambi) of the rice genome. Thirty-eight QTLs were identified with composite interval mapping (CIM), whereas simple interval mapping (SIM) resulted in 47 QTLs (LOD >3.0). The O. rufipogon allele was favourable for 59% of QTLs detected through CIM. Of 261 BC₂F₂ families, 26 advanced backcross breeding lines (BC₂F₅) were used for QTL validation. These lines were selected on the basis of the yield traits potentiality in BC₂F₃ and BC₂F₄ generations. The field trial was conducted at three different locations in Malaysia using randomized complete block design with three replications. Trait based marker analysis was done for QTL determination. Twenty-five QTLs were detected in BC₂F₅ generation whereas 29 QTLs were detected in BC₂F₂ generation of the same population. Two QTLs (qPL-1 and qSPL-7) were not considered for validation due to their low R ² values and two QTLs (qPSS-3-2 and qGW-3-2) were not detected in the BC₂F₅ population. Fifteen QTLs showed the beneficial effect to enhance the trait value of the breeding lines. QTL validation aided to select the promising lines for further utilization.