Association of barley photoperiod and vernalization genes with QTLs for flowering time and agronomic traits in a BC2DH population and a set of wild barley introgression lines

The control of flowering time has important impacts on crop yield. The variation in response to day length (photoperiod) and low temperature (vernalization) has been selected in barley to provide adaptation to different environments and farming practices. As a further step towards unraveling the genetic mechanisms underlying flowering time control in barley, we investigated the allelic variation of ten known or putative photoperiod and vernalization pathway genes between two genotypes, the spring barley elite cultivar ‘Scarlett’ (Hordeum vulgare ssp. vulgare) and the wild barley accession ‘ISR42-8’ (Hordeum vulgare ssp. spontaneum). The genes studied are Ppd-H1, VRN-H1, VRN-H2, VRN-H3, HvCO1, HvCO2, HvGI, HvFT2, HvFT3 and HvFT4. ‘Scarlett’ and ‘ISR42-8’ are the parents of the BC₂DH advanced backcross population S42 and a set of wild barley introgression lines (S42ILs). The latter are derived from S42 after backcrossing and marker-assisted selection. The genotypes and phenotypes in S42 and S42ILs were utilized to determine the genetic map location of the candidate genes and to test if these genes may exert quantitative trait locus (QTL) effects on flowering time, yield and yield-related traits in the two populations studied. By sequencing the characteristic regions of the genes and genotyping with diagnostic markers, the contrasting allelic constitutions of four known flowering regulation genes were identified as ppd-H1, Vrn-H1, vrn-H2 and vrn-H3 in ‘Scarlett’ and as Ppd-H1, vrn-H1, Vrn-H2 and a novel allele of VRN-H3 in ‘ISR42-8’. All candidate genes could be placed on a barley simple sequence repeat (SSR) map. Seven candidate genes (Ppd-H1, VRN-H2, VRN-H3, HvGI, HvFT2, HvFT3 and HvFT4) were associated with flowering time QTLs in population S42. Four exotic alleles (Ppd-H1, Vrn-H2, vrn-H3 and HvCO1) possibly exhibited significant effects on flowering time in S42ILs. In both populations, the QTL showing the strongest effect corresponded to Ppd-H1. Here, the exotic allele was associated with a reduction of number of days until flowering by 8.0 and 12.7%, respectively. Our data suggest that Ppd-H1, Vrn-H2 and Vrn-H3 may also exert pleiotropic effects on yield and yield-related traits.     

Earliness per se QTLs and their interaction with the photoperiod insensitive allele Ppd-D1a in the Cutler × AC Barrie spring wheat population

Earliness per se regulates flowering time independent of environmental signals and helps to fine tune the time of flowering and maturity. In this study, we aimed to map earliness per se quantitative trait loci (QTLs) affecting days to flowering and maturity in a population developed by crossing two spring wheat cultivars, Cutler and AC Barrie. The population of 177 recombinant inbred lines (RILs) was genotyped for a total of 488 SSR and DArT polymorphic markers on all 21 chromosomes. Three QTLs of earliness per se affecting days to flowering and maturity were mapped on chromosomes 1B (QEps.dms-1B1 and QEps.dms-1B2) and 5B (QEps.dms-5B1), in individual environments and when all the environments were combined. A QTL affecting flowering time (QFlt.dms-4A1) was identified on chromosome 4A. Two grain yield QTLs were mapped on chromosome 5B, while one QTL was mapped on chromosome 1D. The population segregated for the photoperiod insensitive gene, Ppd-D1a, and it induced earlier flowering by 0.69 days and maturity by 1.28 days. The photoperiod insensitive allele Ppd-D1a interacted in an additive fashion with QTLs for flowering and maturity times. The earliness per se QTL QFlt.dms-5B.1 inducing earlier flowering could help to elongate grain filling duration for higher grain yield. Hence, chromosome 5B possesses promising genomic regions that may be introgressed for higher grain yield with earlier maturity through marker-assisted selection in bread wheat.     

Quantitative trait loci for growing degree days to flowering and photoperiod response in Sunflower (Helianthus annuus L.)

The number of days from seedling emergence to flowering (DTF) is a major consideration in sunflower breeding programs. This is a complex trait determined by the genotype, environmental conditions and interactions. Photoperiod and temperature have major effects on DTF and could be important sources of genotype× environment interaction. The objectives of this study were to locate quantitative trait loci (QTLs) associated with growing degree days (GDD) to flowering and photoperiod (PP) response in an elite sunflower population. Two hundred and thirty five F₂-generation plants and their F_2:3 and F_2:4 progenies of a single-cross population of two divergent inbred lines were evaluated in six environments (locations, years and sowing dates) with photoperiods known to elicit a PP response between the inbred lines. Detection of QTLs was facilitated with a genetic linkage map of 205 RFLP loci and composite interval mapping. The 205 restriction fragment length polymorphism (RFLP) loci covered 1380 cM and were arranged in 17 linkage groups, which is the haploid number of chromosomes in this species. The average interval size was 5.9 cM. Six QTLs in linkage groups A, B, F, I, J and L were associated with GDD to flowering and accounted for 76% of the genotypic variation in the mean environment. QTLs in linkage groups A and B accounted for 72% of the genetic variation. QTL×environment (QTL×E) interactions were highly significant for linkage groups A, B, F and J (P<0.01). QTLs in linkage groups A and B were highly dependent on PP. Also, QTL mapping of the ratio of the GDD required by a progeny to flower at a PP of 12.1 and 15.0 h, defined as the photoperiod response (PPR), suggested that alleles at QTLs in linkage groups A and B were responsive to PP. QTLs in linkage groups F and J showed QTL×E interaction but the LOD values were not associated with PP. QTL×E interactions for additive effects were highly significant (P<0.01) for linkage groups A, B and F. QTL×E interactions for QTLs with dominant effects were significant (P<0.01) for linkage groups A, B and J. The dominant effect of QTLs in linkage group B increased in environments with a longer PP. The knowledge of how these QTLs influence the GDD for flowering and how they interact with the environment will facilitate marker- assisted selection and backcross conversion of photoperiod-sensitive germplasm. Received: 7 February 2000 / Accepted: 13 June 2000     

Mapping of two genes for resistance to clubroot (Plasmodiophora brassicae) in a population of doubled haploid lines of Brassica oleracea by means of RFLP and AFLP markers

A genetic map covering 615 cM in 12 linkage groups was assembled based on 92 RFLP and AFLP markers segregating in a population of 107 doubled haploid lines (DH lines) of Brassica oleracea. The DH-line population was obtained through microspore culture from the of two homozygous parents: DH-line Bi derived from the cabbage landrace Bindsachsener, and DH-line Gr from broccoli cv ‘Greenia’. Sixty-five percent of the loci, and in some cases complete linkage groups, displayed distorted segregation ratios, a frequency much higher than that observed in populations of the same species. DH-line Bi was resistant to clubroot, which is caused by a Dutch field isolate of Plasmodiophora brassicae. Resistance in the DH-line population was determined in two ways: by assigning symptom grades to each plant, and by measuring the fresh weights of the healthy and affected parts of the root system of each plant. Using a multiple QTL mapping approach to analyze the fresh weight data, we found two loci for clubroot resistance; these were designated pb-3 and pb-4. The additive effects of these loci were responsible for 68% of the difference between the parents and for 60% of the genetic variance among DH-line means. Also, indications for the presence of two additional, minor QTLs were found. Analysis of symptom grades revealed the two QTLs pb-3 and pb-4, as well as one of the two minor QTLs indicated by analysis of the fresh weight data. Received: 29 April 1996 / Accepted: 10 May 1996     

Genotype-by-environment interaction in genetic mapping of multiple quantitative trait loci

The interval mapping method is widely used for the genetic mapping of quantitative trait loci (QTLs), though true resolution of quantitative variation into QTLs is hampered with this method. Separation of QTLs is troublesome, because single-QTL is models are fitted. Further, genotype-by-environment interaction, which is of great importance in many quantitative traits, can only be approached by separately analyzing the data collected in multiple environments. Here, we demonstrate for the first time a novel analytic approach (MQM mapping) that accommodates both the mapping of multiple QTLs and genotype-by-environment interaction. MQM mapping is compared to interval mapping in the mapping of QTLs for flowering time in Arabidopsis thaliana under various photoperiod and vernalization conditions.     

QTL analysis of malting quality in barley based on the doubled-haploid progeny of two elite North American varieties representing different germplasm groups

Characterization of the determinants of economically important phenotypes showing complex inheritance should lead to the more effective use of genetic resources. This study was conducted to determine the number, genome location and effects of QTLs determining malting quality in the two North American barley quality standards. Using a doubled-haploid population of 140 lines from the cross of Harrington×Morex, malting quality phenotype data sets from eight environments, and a 107-marker linkage map, QTL analyses were performed using simple interval mapping and simplified composite interval mapping procedures. Seventeen QTLs were associated with seven grain and malting quality traits (percentage of plump kernels, test weight, grain protein percentage, soluble/total protein ratio, α-amylase activity, diastatic power and malt-extract percentage). QTLs for multiple traits were coincident. The loci controlling inflorescence type [vrs1 on chromosome 2(2H) and int-c on chromosome 4(4H)] were coincident with QTLs affecting all traits except malt-extract percentage. The largest effect QTLs, for the percentage of plump kernels, test weight protein percentage, S/T ratio and diastatic power, were coincident with the vrs1 locus. QTL analyses were conducted separately for each sub-population (six-rowed and two-rowed). Eleven new QTLs were detected in the subpopulations. There were significant interactions between the vrs1 and int-c loci for grain-protein percentage and S/T protein ratio. Results suggest that this mating of two different germplasm groups caused a disruption of the balance of traits. Information on the number, position and effects of QTLs determining components of malting quality may be useful for maintaining specific allele configurations that determine target quality profiles. Received: 28 May 1999 / Accepted: 9 November 1999     

Looking into flowering time in almond (Prunus dulcis (Mill) D. A. Webb): the candidate gene approach

Blooming time is one of the most important agronomic traits in almond. Biochemical and molecular events underlying flowering regulation must be understood before methods to stimulate late flowering can be developed. Attempts to elucidate the genetic control of this process have led to the identification of a major gene (Lb) and quantitative trait loci (QTLs) linked to observed phenotypic differences, but although this gene and these QTLs have been placed on the Prunus reference genetic map, their sequences and specific functions remain unknown. The aim of our investigation was to associate these loci with known genes using a candidate gene approach. Two almond cDNAs and eight Prunus expressed sequence tags were selected as candidate genes (CGs) since their sequences were highly identical to those of flowering regulatory genes characterized in other species. The CGs were amplified from both parental lines of the mapping population using specific primers. Sequence comparison revealed DNA polymorphisms between the parental lines, mainly of the single nucleotide type. Polymorphisms were used to develop co-dominant cleaved amplified polymorphic sequence markers or length polymorphisms based on insertion/deletion events for mapping the candidate genes on the Prunus reference map. Ten candidate genes were assigned to six linkage groups in the Prunus genome. The positions of two of these were compatible with the regions where two QTLs for blooming time were detected. One additional candidate was localized close to the position of the Evergrowing gene, which determines a non-deciduous behaviour in peach.     

Location of genes involved in ear compactness in wheat (Triticum aestivum) by means of molecular markers

Quantitative trait loci (QTLs) for three traits related to ear morphology (spike length, number of spikelets, and compactness as the ratio between number of spikelets and spike length) in wheat (Triticum aestivum L.) were mapped in a doubled-haploid (DH) population derived from the cross between the cultivars Courtot and Chinese Spring. A molecular marker linkage map of this cross that had previously been constructed based on 187 DH lines and 380 markers was used for QTL mapping. The genome was well covered (85%) except chromosomes 1D and 4D and a set of anchor loci regularly spaced (one marker each 15.5 cM) were chosen for marker regression analysis. The presence of a QTL was declared at a significance threshold = 0.001. The population was grown in one location under field conditions during three years (1994, 1995 and 1998). For each trait, 4 to 6 QTLs were identified with individual effects ranging between 6.9% and 21.8% of total phenotypic variation. Several QTLs were detected that affected more than one trait. Of the QTLs 50% were detected in more than one year and two of them (number of spikelets on chromosome 2B, and compactness on chromosome 2D) emerged from the data from the three years. Only one QTL co-segregated with the gene Q known to be involved in ear morphology, namely the speltoid phenotype. However, this chromosome region explained only a minor part of the variation (7.5–11%). Other regions had a stronger effect, especially two previously unidentified regions located on chromosomes 1A and 2B. The region on the long arm of chromosome 1A was close to the locus XksuG34-1A and explained 12% of variation in spike length and 10% for compactness. On chromosome 2B, the QTL was detected for the three traits near the locus Xfbb121-2B. This QTL explained 9% to 22% of variation for the traits and was located in the same region as the gene involved in photoperiod response (Ppd2). Other regions were located at homoeologous positions on chromosomes 2A and 2D.     

Identification and validation of QTLs conferring resistance to sorghum downy mildew (Peronosclerospora sorghi) and Rajasthan downy mildew (P. heteropogoni) in maize

We have mapped the quantitative trait loci (QTLs) conferring resistance to sorghum downy mildew (Peronosclerospora sorghi; SDM) and Rajasthan downy mildew (P. heteropogoni; RDM), two species of DM prevalent throughout India. QTL mapping was carried out on a backcross population of 151 individuals derived from a cross between CM139 (susceptible parent) and NAI116 (highly resistant to both SDM and RDM). Heritability estimates were 0.74 for SDM and 0.67 for RDM. Composite interval mapping combined with a linkage map constructed with 80 simple sequence repeat (SSR) markers resulted in the identification of three QTLs (one each on chromosomes 2, 3 and 6) for SDM resistance and two QTLs (one each on chromosomes 3 and 6) for RDM resistance, all of which were contributed by NAI116. The significance of the major QTL on chromosome 6 (bin 6.05) that confers resistance to diverse DMs in tropical Asia, including SDM and RDM in India, was also verified. The results confirmed that some common QTLs contribute to both SDM and RDM resistance, while additional loci might specifically govern resistance to SDM. The QTL information generated in this study provide information that will aid in undertaking an integrated breeding strategy for the transfer of resistance to SDM and RDM in maize lines using marker-assisted selection.     

Identification of quantitative trait loci controlling sucrose content in soybean (Glycine max)

Sucrose is a primary constituent of soybean (Glycine max) seed; however, little information concerning the inheritance of seed sucrose in soybean is available. The objective of this research was to use molecular markers to identify genomic regions significantly associated with quantitative trait loci (QTL) controlling sucrose content in a segregating F₂ population. DNA samples from 149 F₂ individuals were analyzed with 178 polymorphic genetic markers, including RFLPs, SSRs, and RAPDs. Sucrose content was measured on seed harvested from each of 149 F_2:3 lines from replicated field experiments in 1993 and 1995. Seventeen marker loci, mapping to seven different genomic regions, were significantly associated with sucrose variation at P<0.01. Individually, these markers explained from 6.1% to 12.4% of the total phenotypic variation for sucrose content in this population. In a combined analysis these genomic regions; explained 53% of total variation for sucrose content. No significant evidence of epistasis among QTLs was observed. Comparison of our QTL mapping results for sucrose content and those previously reported for protein and oil content (the other major seed constituents in soybean), suggests that seed quality traits are inherited as clusters of linked loci or that `major' QTLs with pleiotropic effects may control all three traits. Of the seven genomic regions having significant effects on sucrose content, three were associated with significant variation for protein content and three were significantly associated with oil content.     

Intraspecific evolution ofMicroseris pygmaea (Asteraceae,Lactuceae) analyzed by cosegregation of phenotypic characters (QTLs) and molecular markers (RAPDs)

The Chilean annual,Microseris pygmaea, has differentiated in distinct coastal and inland series of populations after long-distance dispersal from western North America. Two plants from the most diverse biotypes were crossed, a large F₂ was raised and analysed for segregation of 30 phenotypic characters. Segregation of molecular markers (47 RAPDs, 1 RFLP, 2 isozymes) was determined in a subpopulation of 45 plants which include all extremes for the phenotypic characters. 32 marker/character cosegregations were significant at the 1% level in t-tests between dominant and homozygous recessive marker genotypes. Considering linkage among markers and pleiotropy of certain marker loci, the number of independent quantitative trait loci (QTLs) is reduced to about 18. Interactions among 2 or 3 QTLs affecting one character have been characterized. The phenotypic differentiation ofM. pygmaea during its evolution from a single founder individual begins to be understood at the level of single-gene mutants.     

The effect of semi-dwarf genes on root system size in field-grown barley

Root system size (RSS) was measured in 12 diverse barley genotypes and 157 double-haploid lines (DHs), using electric capacitance. The parents of the DHs, Derkado and B83-12/21/5, carry different semi-dwarfing genes, sdw1 and ari-e.GP, respectively. Estimates of RSS were taken in the field thrice during plant development: stem elongation (RSS1), heading (RSS2) and grain filling (RSS3). The 12 barley genotypes were assessed over 3 years and at two or three locations each year; the DH mapping population was assessed at two locations in 2002. Among the 12 barley genotypes, those with the semi-dwarf genes had greater RSS values in all 3 years (28.9, 24.6 and 15.0% in years 1, 2 and 3, respectively) compared to non-semi-dwarf controls. The DH population showed transgressive segregation on both sides of the parent means, indicating polygenic control of RSS. Quantitative trait loci (QTLs) for RSS were found on five of the seven chromosomes: 1H, 3H, 4H, 5H and 7H and these were compared with previously mapped agronomic traits. The TotalRSS QTL on 3H was associated with sdw1 and QTLs for height, plant yield and plant weight. The RSS3 QTL on 5H was associated with ari-e.GP and QTLs for height, plant yield, plant weight, harvest index and tiller number. The RSS3 QTL on 7H was also associated with a TotalRSS QTL and QTLs for plant weight and harvest index. Other RSS QTLs were not associated with any other trait studied. RSS is considered to be a polygenic trait linked to important traits, in particular to yield. The study highlights the effects of semi-dwarfing genes and discusses the potential for breeding for root traits.     

Detection of quantitative trait loci on chromosome 5R of rye (Secale cereale L.)

 Progenies of an F₂ mapping population were analyzed for quantitative traits to detect QTLs by using marker information from F₂ plants for chromosome 5R. The mapping population was segregating for the major dwarfing gene Ddw1 and the gene Hp1 for hairy peduncle. The only QTL determining plant height was located between HP1 and Ddw1 on the distal part of chromosome 5RL. At the same position a QTL for peduncle length was found, and this trait was closely related to plant height (r=0.895). Since Hp1 and Ddw1 are dominant marker loci, no dominance effect could be estimated. The QTLs for spike length and the number of florets were located near the centromere on 5RL. These two traits were correlated with r=0.824 and showed partial dominance, but these traits were not correlated to plant height and peduncle length. Homoeologous relationships between the QTLs mapped for the first time in rye and those mapped in other Triticeae members are discussed. Received: 8 June 1998 / Accepted: 8 October 1998     

QTLs influencing panicle size detected in two reciprocal introgressive line (IL) populations in rice (Oryza sativa L.)

Two sets of reciprocal introgression line (IL) populations, i.e., ILs with Lemont as recurrent parent (IL_LT) and ILs with Teqing as recurrent parent (IL_TQ), were developed and evaluated for traits representing panicle size, including primary branch number (PBN), secondary branch number (SBN), and spikelet number per panicle (SNP). Together with the regression to recurrent parent by advanced backcross, transgressive segregations were observed for all traits. Correlation and regression analysis showed that SBN had much higher contribution to SNP than PBN. It was confirmed by the QTL analysis that many common loci were detected between SBN and SNP, in comparison with single common locus between PBN and SNP. One and three main effect QTLs (M-QTLs) were detected for PBN in IL_LT and IL_TQ, respectively. Six M-QTLs per trait per populations were associated with SBN and SNP. Less number and lower contribution of epistasis were detected in IL populations in comparison with mapping result from F ₂ or RI population. There were only four QTLs in fourteen loci (near 30%) commonly detected in both reciprocal IL populations implying the large impact of genetic background on QTLs expression. H. W. Mei and J. L. Xu contributed equally to this work.     

RFLP mapping of five major genes and eight quantitative trait loci controlling flowering time in a winter x spring barley (Hordeum vulgare L.) cross.

A genetic map of 92 RFLP loci and two storage protein loci was made using 94 doubled-haploid lines from a cross between the winter barley variety Igri and the spring variety Triumph. The markers were combined with data from two field experiments (one spring sown and one autumn (fall) sown) and a glasshouse experiment to locate a total of 13 genes (five major genes and eight quantitative trait loci (QTL)) controlling flowering time. Two photoperiod response genes were found; Ppd-H1 on chromosome 2(2H)S regulated flowering time under long days, while Ppd-H2 on chromosome 5(1H)L was detected only under short days. In the field experiments Ppd-H1 strongly affected flowering time from spring and autumn sowings, while Ppd-H2 was detected only in the autumn sowing. The glasshouse experiment also located two vernalization response genes, probably Sh and Sh2, on chromosomes 4(4H)L and 7(5H)L, respectively. The vernalization response genes had little effect on flowering time in the field. Variation in flowering time was also affected by nine additional genes, whose effects were not specifically dependent on photoperiod or vernalization. One was the denso dwarfing gene on chromosome 3(3H)L. The remaining eight were QTLs of smaller effect. One was located on chromosome 2(2H), one on 3(3H), one on 4(4H), one on 7(5H), two on 6(6H), and two on 1(7H). Model fitting showed that the 13 putative genes, and their interactions, could account for all the observed genetical variation from both spring and autumn sowings, giving a complete model for the control of flowering time in this cross.     

Identification of two closely linked quantitative trait loci for cold tolerance on chromosome 4 of rice and their association with anther length

Norin-PL8 is a cold-tolerant variety of rice (Oryza sativa L.) that was developed by introgressing chromosomal segments from a cold-tolerant javanica variety, Silewah. We previously detected quantitative trait loci (QTLs) for cold tolerance of Norin-PL8 in the introgressions on chromosomes 3 and 4. We provide fine mapping of the QTLs on chromosome 4 and the association between the QTLs and anther length, which has been reported to be a major component of cold tolerance. Interval mapping using a segregating population derived from an advanced backcross progeny indicated that a QTL for cold tolerance is probably located from the center to the proximal end of the introgression. For fine mapping, we developed a set of near-isogenic lines (NILs) from recombinants in the segregating population. Comparison of cold tolerance between the NILs indicated that either the proximal end or the center of the introgression is necessary for cold tolerance. From these results, we concluded that there are at least two QTLs for cold tolerance, tentatively designated as Ctb-1 and Ctb-2, in the introgression on chromosome 4. The map distance between Ctb-1 and Ctb-2 is estimated to be 4.7–17.2 cM. In order to investigate the mechanism underlying cold tolerance by the QTLs, we compared anther lengths of the NILs. The results indicate that both Ctb-1 and Ctb-2 are associated with anther length. Received: 17 July 2000 / Accepted: 1 February 2001     

Genetic architecture of adiposity in the cross of LG/J and SM/J inbred mice

The genetic basis of variation in obesity in human populations is thought to be owing to many genes of relatively small effect and their interactions. The LG/J by SM/J intercross of mouse inbred strains provides an excellent model system in which to investigate multigenic obesity. We previously mapped a large number of quantitative trait loci (QTLs) affecting adult body weight in this cross. We map body composition traits, adiposity, and skeletal size, in a replicate F₂ intercross of the same two strains containing 510 individuals. Using interval-mapping methods, we located eight QTLs affecting adiposity (Adip1–8). Two of these adiposity loci also affected tail length (Adip4 and Adip6) along with seven additional tail length QTLs (Skl1–7). A further four QTLs (Wt1–4) affect adult weight but not body composition. These QTLs have relatively small effects, typically about 0.2–0.4 standard deviation units, and account for between 3% and 10% of the variance in individual characters. All QTLs participated in epistatic interactions with other QTLs. Most of these interactions were due to additive-by-additive epistasis, which can nullify the apparent effects of single loci in our population. Adip8 interacts with all the other adiposity QTLs and seems to play a central role in the genetic system affecting obesity in this cross. Only two adiposity QTLs, Adip4 and Adip6, also affect tail length, indicating largely separate genetic control of variation in adiposity and skeletal size. Body size and obesity QTLs in the same locations as those discovered here are commonly found in mapping experiments with other mouse strains. Received: 11 January 2000 / Accepted: 17 August 2000     

A SEARCH FOR QUANTITATIVE TRAIT LOCI AFFECTING ASYMMETRY OF MANDIBULAR CHARACTERS IN MICE

An interval mapping procedure was used to search for and describe the effects of any quantitative trait loci (QTLs) for directional asymmetry (DA) and fluctuating asymmetry (FA) of 10 bilateral mandible characters in house mice. It was hypothesized that more QTLs would be found for DA than for FA, but that any discovered for FA should tend to exhibit dominance. All mandible characters were triply measured and 76 microsatellite markers were scored in an average of 471 mice from the F₂ intercross of the Large (LG/J) and Small (SM/J) inbred strains. A total of 16 QTLs significantly affected DA in nine of the 10 mandible characters, and this was more than the 9.5 expected by chance alone. These QTLs were found on seven of the 19 chromosomes, often at or near locations of QTLs affecting the mean of the two sides for various dimensions on the mandible. It was concluded that there is genetical variability for DA in these characters, although its level was low (4.4% of the total variation in this particular F₂ population). Eleven QTLs were detected for FA, suggesting that there is very little genetic variability for FA, at least as seen in the mandible characters in this particular F₂ population. As hypothesized, however, these QTLs did tend to exhibit dominance.     

QTLs and epistasis for aluminum tolerance in rice (Oryza sativa L.) at different seedling stages

To investigate the genetic background for aluminum (Al) tolerance in rice, a recombinant inbred (RI) population, derived from a cross between an Al-sensitive lowland indica rice variety IR1552 and an Al-tolerant upland japonica rice variety Azucena, was used in culture solution. A molecular linkage map, together with 104 amplified fragment length polymorphism (AFLP) markers and 103 restriction fragment length polymorphism (RFLP) markers, was constructed to map quantitative trait loci (QTLs) and epistatic loci for Al tolerance based on the segregation for relative root length (RRL) in the population. RRL was measured after stress for 2 and 4 weeks at a concentration of 1mM of Al³⁺ and a control with a pH 4.0, respectively. Two QTLs were detected at both the 2nd and the 4th weeks on chromosomes 1 and 12 from unconditional mapping, while the QTL on chromosome 1 was only detected at the 2nd stress week from conditional mapping. The effect of the QTL on chromosome 12 was increased with an increase of the stress period from 2 to 4 weeks. The QTL on chromosome 1 was expressed only at the earlier stress, but its contribution to tolerance was prolonged during growth. At least one different QTL was detected at the different stress periods. Mean comparisons between marker genotypic classes indicated that the positive alleles at the QTLs were from the Al-tolerant upland rice Azucena. An important heterozygous non-allelic interaction on Al tolerance was found. The results indicated that tolerance in the younger seedlings was predominantly controlled by an additive effect, while an epistatic effect was more important to the tolerance in older seedlings; additionally the detected QTLs may be multiple allelic loci for Al tolerance and phosphorus-uptake efficiency, or for Al and Fe²⁺ tolerance. Received: 29 July 1999 / Accepted: 13 October 1999     

Construction of introgression lines carrying wild rice (Oryza rufipogon Griff.) segments in cultivated rice (Oryza sativa L.) background and characterization of introgressed segments associated with yield-related traits

Introgression lines (ILs) are useful tools for precise mapping of quantitative trait loci (QTLs) and the evaluation of gene action or interaction in theoretical studies. A set of 159 ILs carrying variant introgressed segments from Chinese common wild rice (Oryza rufipogon Griff.), collected from Dongxiang county, Jiangxi Province, in the background of Indica cultivar (Oryza sativa L.), Guichao 2, was developed using 126 polymorphic simple sequence repeats (SSR) loci. The 159 ILs represented 67.5% of the genome of O. rufipogon. All the ILs have the proportions of the recurrent parent ranging from 92.4 to 99.9%, with an average of 97.4%. The average proportion of the donor genome for the BC₄F₄ population was about 2.2%. The mean numbers of homozygous and heterozygous donor segments were 2 (ranging 0–8) and 1 (ranging 0–7), respectively, and the majority of these segments had sizes less than 10 cM. QTL analysis was conducted based on evaluation of yield-related traits of the 159 ILs at two sites, in Beijing and Hainan. For 6 out of 17 QTLs identified at two sites corresponding to three traits (panicles per plant, grains per panicle and filled grains per plant, respectively), the QTLs derived from O. rufipogon were usually associated with an improvement of the target trait, although the overall phenotypic characters of O. rufipogon were inferior to that of the recurrent parent. Of the 17 QTLs, 5 specific QTLs strongly associated with more than one trait were observed. Further analysis of the high-yielding and low-yielding ILs revealed that the high-yielding ILs contained relatively less introgressed segments than the low-yielding ILs, and that the yield increase or decrease was mainly due to the number of grain. On the other hand, low-yielding ILs contained more negative QTLs or disharmonious interactions between QTLs which masked trait-enchancing QTLs. These ILs will be useful in identifying the traits of yield, tolerance to low temperature and drought stress, and detecting favorable genes of common wild rice.