Quantitative trait loci increasing acylsugars in tomato breeding lines and their impacts on silverleaf whiteflies

Solanum pennellii LA716, a wild relative of tomato, produces acylsugars, an insect resistance compound with activity against many tomato insect pests. Breeding of cultivated tomato using S. pennellii LA716 as a donor parent has led to the development of the elite acylsugar-producing tomato breeding line CU071026. CU071026 contains five introgressed S. pennellii genomic regions, and produces acylsugars at moderate levels that are effective against insect pests. A BC1F1 population was created by crossing the F1 CU071026?×?S. pennellii LA716 with CU071026 as the recurrent parent; this BC1F1 population was used to identify additional regions of the S. pennellii genome important for further improvement of acylsugar production. This population was genotyped with 94 markers in the segregating regions and phenotyped for level of acylsugar production. Using QTLNetwork 2.1 for the detection of quantitative trait loci (QTL) and epistatic interactions, this study identified five QTL for total acylsugar level. Additionally, two epistatic interactions between QTL were found to control significant levels of total acylsugar production. Two of the QTL identified were further evaluated in silverleaf whitefly (Bemisia tabaci) field cage trials using acylsugar breeding lines that differ for the presence/absence of these QTL. While high levels of silverleaf whitefly resistance were observed in all acylsugar breeding lines, lines containing the additional QTL on either chromosomes 6 or 10 had increased levels of total acylsugar production and reduced incidence of whitefly. Acylsugar lines containing the chromosome 6 QTL also had increased density of the type IV glandular trichomes which produce and exude acylsugars.     

Inheritance of acylsugar contents in tomatoes derived from an interspecific cross with the wild tomato Lycopersicon pennellii and their effect on spider mite repellence

Acylsugars present in Lycopersicon pennellii are responsible for the high levels of pest resistance often found in this wild tomato taxon. We investigated the inheritance of acylsugar contents in segregating populations of the interspecific tomato cross L. esculentum x L. pennellii and estimated correlations between leaflet acylsugar contents and the levels of mite repellence. Acylsugar contents were quantified with the Sommogy-Nelson colorimetric method in the acessions L. esculentum 'TOM-584' (P(1), low acylsugars), L. pennellii 'LA-716' (P(2), high acylsugars), in the interspecific F(1) (P(1) x P(2)) and in the F(2 )(P(1) x P(2)) generations. Mite resistance was assessed by a repellence test. Broad-sense heritability of acylsugar contents was moderately high (h(2)(b) = 0.476). Frequency distributions in the P(1), P(2), F(1) and F(2) can be explained by the action of a single major locus, with near-complete dominance of the L. esculentum allele for low-acylsugar content over the L. pennellii allele for high content. Indirect selection for high levels of acylsugars in leaflets led to correlated increases in the levels of mite repellency, indicating that acylsugars may be the main factor involved in mite resistance.     

Identification of quantitative trait loci associated with acylsugar accumulation using intraspecific populations of the wild tomato, Lycopersicon pennellii

 Lycopersicon pennellii LA716, a wild relative of tomato, is resistant to a number of insect pests due to the accumulation of acylsugars exuded from type IV trichomes. These acylsugars are a class of compounds including both acylglucoses and acylsucroses. Intraspecific populations between L. pennellii LA716 and L. pennellii LA1912, the latter an accession that assorts for low-level acylsugar accumulation, were created to study the inheritance of type IV trichome density, acylsugar accumulation levels, percentage of acylsugars that are acylglucoses, and leaf area. The F₂ population was subsequently used to determine genomic regions associated with these traits. The relative proportion of acylglucoses and acylsucroses was found to be largely controlled by a single locus near TG549 on chromosome 3. One locus on chromosome 10 showed significant associations with acylsugar levels. In addition, 1 locus on chromosome 4 showed significant associations with leaf area. Ten additional loci showed modest associations with one or more of the traits examined, 5 of which have been previously reported. Received: 13 March 1997 / Accepted: 19 September 1997     

Quantitative trait loci regulating the fatty acid profile of acylsugars in tomato

Acylsugars are secondary metabolites with proven insect resistance properties that are produced by many Solanaceous species including Solanum pennellii, a wild relative of tomato. The acylsugar chemotypes of S. pennellii varies greatly within its natural range and might be the product of plant/insect coevolution. The S. pennellii accession LA716 was used to transfer increased levels of acylsugar production into the cultivated tomato, resulting in the acylsugar tomato breeding line CU071026. S. pennellii accession LA716 produces high levels of acylsugars with chemotypes that differ greatly from those produced by CU071026 or the trace acylsugars of cultivated tomato. Understanding the genetic regulation of acylsugar chemistry will aid efforts to breed acylsugar production into cultivated tomato, allowing for alteration of both acylsugar base moieties and fatty acid profiles. This study uses a BC1F1 population produced from the cross of S. pennellii LA716 and CU071026 with CU071026 as the recurrent parent to identify QTL that change the fatty acid profile of acylsugars. Multiple QTL and epistatic interactions between QTL were detected including three QTL on chromosomes 2, 5, and 7, which significantly alter the percentage of extended iso-odd branched fatty acids and straight chain fatty acids on the acylsugars. We also report the introgression of one of these QTL, FA 2, into CU071026, resulting in a new tomato line with significantly increased i11:0 as a percentage of the fatty acids in its acylsugars. Candidate genes for these QTL and the impacts of altering acylsugar fatty acid are discussed.     

QTL analysis of pest resistance in the wild tomato Lycopersicon pennellii: QTLs controlling acylsugar level and composition

Some accessions of Lycopersicon pennellii, a wild relative of the tomato Lycopersicon esculentum, are resistant to a number of important pests of cultivated tomato due to the accumulation of acylsugars, which constitute 90% of the exudate of type-IV trichomes in L. pennellii LA716. An interspecific F₂ population, created by the cross L. esculentum x L. pennellii LA 716, was surveyed for acylsugar accumulation and subjected to RFLP/QTL analysis to determine the genomic regions associated with the accumulation of acylglucoses, acylsucroses, and total acylsugars, as well as with acylglucoses as a percentage of total acylsugars (mole percent acylglucoses). Data were analyzed using MAPMAKER/QTL with and without a log₁₀ transformation. A threshold value of 2.4 (default value for MAPMAKER/QTL) was used, as well as 95% empirically derived threshold values. Five genomic regions, two on chromosome 2 and one each on chromosomes 3, 4 and 11, were detected as being associated with one or more aspects of acylsugar production. The L. esculentum allele is partially dominant to the L. pennellii allele in the regions on chromosomes 2 and 11, but the L. pennellii allele is dominant in the region on chromosome 3. Throughout this study, we report the comparative effects of analytical methodology on the identification of acylsugar QTLs. Similarities between our results and published results for the genus Solanum are also discussed.R. W. Doerge · S.-C. Liu · J. P. Kuai contributed equally to the paper, and we ordered randomly     

Quantitative trait loci regulating sugar moiety of acylsugars in tomato

Acylsugars are broad-spectrum insect resistance sugar esters produced at very high levels by some accessions of the wild tomato, Solanum pennellii. Transferring acylsugar production from S. pennellii LA716 to cultivated tomato through traditional breeding developed the benchmark acylsugar breeding line CU071026. The base moiety of acylsugars (sucrose vs. glucose) can vary among S. pennellii accessions. Additionally the accession S. pennellii LA716 produces almost exclusively acylglucoses, but the breeding line CU071026 derived from S. pennellii LA716 produces exclusively acylsucroses. This study uses a BC1F1 and a BC1F2 population derived from the cross CU071026 × (CU071026 × S. pennellii LA716) to identify and confirm the action of three quantitative trait loci (QTL) on chromosomes 3, 4, and 11. The QTL on chromosomes 3 and 11 are both required for acylglucose production, while addition of the chromosome 4 QTL affects the level of acylglucose produced in the presence of the QTL on chromosomes 3 and 11. A three-way interaction between these acylglucose QTL was confirmed with a post hoc ANOVA. Identification of these three QTL provides a blueprint for breeding to shift acylsucrose production to acylglucose production in tomato breeding lines. The implications of these QTL and two additional QTL affecting total acylsugar level in the BC1F2 are discussed.     

Combination of QTL affecting acylsugar chemistry reveals additive and epistatic genetic interactions to increase acylsugar profile diversity

The tomato breeding line, CU071026, was bred to accumulate high levels of the insect control compounds called acylsugars, which are exuded from glandular trichomes. The acylsugars of CU071026 exhibit a characteristic profile of acylsugar composition and constituent fatty acid acyl groups that is distinct from that of the progenitor wild tomato, Solanum pennellii LA716. A prior study reported the transfer of three QTL (FA2, FA7, and FA8), from S. pennellii LA716, that are associated with changes in acylsugar chemistry into CU071026 and demonstrated that the resulting lines, each of which possesses one of these QTL, displayed a unique acylsugar and fatty acid profile distinct from that characteristic of the acylsugars of CU071026 and each other. The current study utilized marker-assisted backcrossing to combine pairs of two of these QTL or all three of these QTL. This created a new set of lines, which allowed evaluation of the combinatory effects of FA2QTL, FA7QTL, and FA8QTL, on acylsugar level and acylsugar and fatty acid profile and diversity. Analysis of the new tomato lines revealed that these QTL interacted additively and epistatically to alter acylsugar level and chemistry, increasing the diversity of fatty acid constituents and/or acylsugar chemotypes present in the exudates of some of the lines. Extensive characterization of the lines for acylsugar level, through a spectrophotometric invertase assay, and acylsugar chemistry, through gas and liquid chromatography-mass spectrometry, allowed association of the QTL interactions with aspects of acylsugar chemotype. The evaluated fatty acids and acylsugars accumulated by the set of lines generally displayed high heritability and minimal environmental effect, which is discussed. The QTL interactions that govern a more diverse acylsugar and fatty acid profile provide valuable information for the generation of tomato lines with improved acylsugar efficacy against pests of tomato.     

Introgression of acylsugar chemistry QTL modifies the composition and structure of acylsugars produced by high-accumulating tomato lines

Acylsugars are important insect defense compounds produced at high levels by glandular trichomes of the wild tomato, Solanum pennellii. The ability to produce acylsugars at elevated levels was bred into the tomato line CU071026. This study utilized a marker-assisted backcross approach to individually introgress into CU071026 and to fine map the three quantitative trait loci (QTL) fatty acid 5 (FA5QTL), fatty acid 7 (FA7QTL), and fatty acid 8 (FA8QTL), which were previously associated with changes in acylsugar chemistry. Additional breeding with and fine mapping the previously introgressed QTL, fatty acid 2 (FA2QTL), was also conducted. The effect of these four QTL on acylsugar quality and quantity in the presence of the five introgressions of CU071026 was evaluated. Incorporation of the QTL altered acylsugar chemotype by modulating the length, orientation, and/or relative proportion of fatty acid acyl groups. The resulting quantities of acylsugar produced in most of the new lines were similar to that of CU071026; however, introgression of FA5QTL reduced acylsugar levels. The acylsugar lines containing each QTL were characterized for acylsugar level, trichome abundance, and acylsugar chemistry through gas chromatography/mass spectrometry and liquid chromatography/mass spectrometry. The novel acylsugar chemotype lines created can contribute to elucidation of the mechanism of insect resistance mediated by acylsugars and help with identification of yet-unknown genes contributing to acylsugar synthesis and diversity.     

Combination of Acylglucose QTL reveals additive and epistatic genetic interactions and impacts insect oviposition and virus infection

Acylsugars are important insect defense compounds produced at high levels by glandular trichomes of the wild tomato, Solanum pennellii. Marker-assisted selection was used to select for plants containing the three AGQTL named AG3QTL, AG4QTL, and AG11QTL from self-pollinated populations derived from an interspecific backcross population of CU071026 x (CU071026 x S. pennellii LA716). High acylglucose-accumulating lines were selected from these populations that possess these three AGQTL and the fewest number of extraneous S. pennellii LA716 introgressions. Incorporation of these three acylglucose QTL in the presence of the five standard S. pennellii introgressions of CU071026 altered acylsugar level and sugar moiety, demonstrating epistatic interactions between the acylglucose QTL on both of these traits. Comparison of the lines generated from the two breeding techniques indicated the three acylglucose QTL are essential but not necessarily sufficient for the production of elevated levels of acylglucose acylsugars. Fine-mapping of AG3QTL, AG4QTL, and AG11QTL resulted in less than 1 Mbp intervals for the locations of AG4QTL and AG11QTL; proposals of the causal genes underlying these acylglucose QTL are discussed. Characterization of the fatty acid profile of lines selected out of the interspecific backcross populations revealed an increase in the proportion of acylsugar n-C10 fatty acid acyl chains, possibly governed by one or more of the three acylglucose QTL. Characterization of the acylsugar profile of acylglucose lines selected from the interspecific backcross populations also demonstrated interactions among the acylglucose QTL to further modulate the diversity of acylsugars accumulated. Evaluation of an acylglucose line and controls against the tomato insect pest Frankliniella occidentalis demonstrated that levels of resistance differed among these lines and that the acylsugars accumulated by the acylglucose line were effective at reducing both F. occidentalis oviposition and incidence of Tomato spotted wilt orthotospovirus. However, of some of the acylglucose lines and hybrids tested against Spodoptera exigua did not indicate differences for larval weight gain and survival.     

Use of introgression lines and zonal mapping to identify RAPD markers linked to QTL

RAPD primers were identified as giving parent-specific bands when screened with a set of introgression lines containing introgressed regions of Lycopersicon pennellii that encompass 5 quantitative trait loci affiliated with the production and composition of acylsugars, compounds associated with insect resistance. Primers giving L. pennellii introgression specific bands were zonally mapped to identify bands affiliated with the quantitative trait target and flanking regions using subsets of 7 to 16 F2 individuals which contained small overlapping segments (zones) of the L. pennellii genome spanning those regions. Seventeen RAPD primers, agt-related primers, and an agt clone were then used in mapping the complete F2 population of 144 individuals. This work resulted in the identification of RAPD markers for three of the 5 quantitative trait loci and the construction of an integrated RAPD/RFLP genomic map for tomato (Lycopersicon esculentum x L. pennellii LA716) of 111 RAPD and 8 acylglucose transferase related markers added to a framework map of 150 RFLP markers.     

Constitutive activation of the jasmonate signaling pathway enhances the production of secondary metabolites in tomato

The phytohormone jasmonic acid (JA) regulates the synthesis of secondary metabolites in a wide range of plant species. Here, we show that exogenous methyl-JA (MeJA) elicits massive accumulation of caffeoylputrescine (CP) in tomato leaves. A mutant (jai1) that is defective in jasmonate perception failed to accumulate CP in flowers and MeJA-treated leaves. Conversely, a transgenic tomato line (called 35S::PS) that exhibits constitutive JA signaling accumulated high levels of leaf CP in the absence of jasmonate treatment. RNA blot analysis showed that genes encoding enzymes in the phenylpropanoid and polyamine pathways for CP biosynthesis are upregulated in MeJA-treated wild-type plants and in untreated 35S::PS plants. These results indicate that CP accumulation in tomato is tightly controlled by the jasmonate signaling pathway, and provide proof-of-concept that the production of some plant secondary metabolites can be enhanced by transgenic manipulation of endogenous JA levels.     

Recessive resistance genes against potyviruses are localized in colinear genomic regions of the tomato ( Lycopersicon spp.) and pepper ( Capsicum spp.) genomes

Resistance against both Potato virus Y (PVY) and Tobacco etch virus (TEV) was identified in the wild tomato relative Lycopersicon hirsutum PI247087. Analysis of the segregation ratio in F(2)/F(3) and BC(1) interspecific progenies indicated that a single recessive gene, or two very tightly linked recessive loci, are involved in resistance to both potyviruses. This locus was named pot-1. Using amplified fragment length polymorphism markers and a set of L. hirsutum introgression lines, pot-1 was mapped to the short arm of tomato chromosome 3, in the vicinity of the recessive py-1 locus for resistance to corky root rot. Because of the occurrence of phenotypically similar genes in pepper ( Capsicum spp.), the comparative genetics of resistance to potyviruses between tomato and pepper was investigated. Unlike most of the comparative genetic studies on resistance genes, pot-1 was tightly flanked by the same restriction fragment length polymorphism (RFLP) markers than the pvr2/pvr5 locus for resistance to PVY and TEV from pepper. These results may indicate that recessive resistance genes against potyviruses evolve less rapidly than the majority of the dominant genes cloned so far, and consequently may belong to a different family of resistance genes.     

重金属镉在番茄及其害虫西花蓟马中的积累与传递

重金属镉(Cd)是农田土壤中的重要污染源,可在植物和植食性昆虫中积累与传递。本文采用水培法,研究了不同浓度的Cd在番茄Solanum lycopersicum不同组织和在其重要害虫西花蓟马Frankliniella occidentalis体内的积累量。结果表明,随着水培营养液中Cd浓度的增加,番茄植株的根、茎和叶中Cd含量呈增长趋势。根中Cd的积累量远高于茎和叶,当水培溶液中Cd含量为20 mg/L时,根、茎和叶中的积累量分别达19 333.67±233.38、122.67±6.84和147.33±2.96 mg/Kg(干重)。随着Cd浓度的增加,番茄根、茎和叶的鲜重和干重均显著下降。西花蓟马取食Cd处理的番茄叶片后,体内Cd显著积累,最高达1.95±0.36 mg/Kg。同时,Cd积累量的提高进一步影响了以番茄叶片为食的西花蓟马的适合度,降低了其存活率。除对照外,番茄茎-叶的转移系数和叶片对Cd的富集系数均大于1,叶片表现出较强的富集能力。而在所有的试验浓度处理中,西花蓟马对Cd的富集系数和转移系数均小于1,表明Cd未在其体内产生生物放大作用。研究结果明确了Cd在番茄各组织及其害虫中的积累和传递水平,为揭示重金属在农业生态系统食物链中的富集效应提供了基础数据。     

A genetic map of tomato based on BC(1) Lycopersicon esculentum x Solanum lycopersicoides reveals overall synteny but suppressed recombination between these homeologous genomes

F(1) hybrids between the cultivated tomato (Lycopersicon esculentum) and the wild nightshade Solanum lycopersicoides are male sterile and unilaterally incompatible, breeding barriers that impede further crosses to tomato. Meiosis is disrupted in 2x hybrids, with reduced chiasma formation and frequent univalents, but is normal in allotetraploid hybrids, indicating the genomes are homeologous. In this study, a partially male-fertile F(1) was backcrossed to tomato, producing the first BC(1) population suitable for genetic mapping from this cross. BC(1) plants were genotyped at marker loci to study the transmission of wild alleles and to measure rates of homeologous recombination. The pattern of segregation distortion, in favor of homozygotes on chromosomes 2 and 5 and heterozygotes on chromosomes 6 and 9, suggested linkage to a small number of loci under selection on each chromosome. Genome ratios nonetheless fit Mendelian expectations. Resulting genetic maps were essentially colinear with existing tomato maps but showed an overall reduction in recombination of approximately 27%. Recombination suppression was observed for all chromosomes except 9 and 12, affected both proximal and distal regions, and was most severe on chromosome 10 (70% reduction). Recombination between markers on the long arm of this chromosome was completely eliminated, suggesting a lack of colinearity between S. lycopersicoides and L. esculentum homeologues in this region. Results are discussed with respect to phylogenetic relationships between the species and their potential use for studies of homeologous pairing and recombination in a diploid plant genome.     

Identification of QTLs for early blight ( Alternaria solani) resistance in tomato using backcross populations of a Lycopersicon esculentum x L. hirsutum cross

Most commercial cultivars of tomato, Lycopersicon esculentum Mill., are susceptible to early blight (EB), a devastating fungal ( Alternaria solani Sorauer) disease of tomato in the northern and eastern parts of the U.S. and elsewhere in the world. The disease causes plant defoliation, which reduces yield and fruit quality, and contributes to significant crop loss. Sources of resistance have been identified within related wild species of tomato. The purpose of this study was to identify and validate quantitative trait loci (QTLs) for EB resistance in backcross populations of a cross between a susceptible tomato breeding line (NC84173; maternal and recurrent parent) and a resistant Lycopersicon hirsutum Humb. and Bonpl. accession (PI126445). Sixteen hundred BC(1) plants were grown to maturity in a field in 1998. Plants that were self-incompatible, indeterminant in growth habit, and/or extremely late in maturity, were discarded in order to eliminate confounding effects of these factors on disease evaluation, QTL mapping, and future breeding research. The remaining 145 plants (referred to as the BC(1) population) were genotyped for 141 restriction fragment length polymorphism (RFLP) markers and 23 resistance gene analogs (RGAs), and a genetic linkage map was constructed. BC(1) plants were evaluated for disease symptoms throughout the season, and the area under the disease progress curve (AUDPC) and the final percent defoliation (disease severity) were determined for each plant. BC(1) plants were self-pollinated and produced BC(1)S(1) seed. The BC(1)S(1) population, consisting of 145 BC(1)S(1) families, was grown and evaluated for disease symptoms in replicated field trials in two subsequent years (1999 and 2000) and AUDPC and/or final percent defoliation were determined for each family in each year. Two QTL mapping approaches, simple interval mapping (SIM) and composite interval mapping (CIM), were used to identify QTLs for EB resistance in the BC(1) and BC(1)S(1) populations. QTL results were highly consistent across generations, years and mapping approaches. Approximately ten significant QTLs (LOD >/= 2.4, P 57% of the total phenotypic variation. All QTLs had the positive alleles from the disease-resistant parent. The good agreement between results of the BC(1) and 2 years of the BC(1)S(1) generations indicated the stability of the identified QTLs and their potential usefulness for improving tomato EB resistance using marker-assisted selection (MAS). Further inspections using SIM and CIM indicated that six of the ten QTLs had independent additive effects and together could account for up to 56.4% of the total phenotypic variation. These complementary QTLs, which were identified in two generations and 3 years, should be the most useful QTLs for MAS and improvement of tomato EB resistance using PI126445 as a gene resource. Furthermore, the chromosomal locations of 10 of the 23 RGAs coincided with the locations of three QTLs, suggesting possible involvement of these RGAs with EB resistance and a potential for identifying and cloning genes which confer EB resistance in tomato.     

Induction of defense-related proteins in tomato roots treated with Pseudomonas fluorescens Pf1 and Fusarium oxysporum f. sp. lycopersici

Pseudomonas fluorescens isolate Pf1 was found to protect tomato plants from wilt disease caused by Fusarium oxysporum f. sp. lycopersici. Induction of defense proteins and chemicals by P. fluorescens isolate Pf1 against challenge inoculation with F. oxysporum f. sp. lycopersici in tomato was studied. Phenolics were found to accumulate in bacterized tomato root tissues challenged with F. oxysporum f. sp. lycopersici at one day after pathogen challenge. The accumulation of phenolics reached maximum at the 5^th day after pathogen challenge. In pathogen-inoculated plants, the accumulation started at the 2^nd day and drastically decreased 4 days after the pathogen inoculation. Activities of phenylalanine ammonia-lyase (PAL), peroxidase (PO) and polyphenol oxidase (PPO) increased in bacterized tomato root tissues challenged with the pathogen at one day after pathogen challenge and activities of PAL and PO reached maximum at the 4^th day while activity of PPO reached maximum at the 5^th day after challenge inoculation. Isoform analysis revealed that a unique PPO1 isoform was induced and PO1 and PPO2 isoforms were expressed at higher levels in bacterized tomato root tissues challenge inoculated with the pathogen. Similarly, -1,3 glucanase, chitinase and thaumatin-like proteins (TLP) were induced to accumulate at higher levels at 3-5 days of challenge inoculation in bacterized plants. Western blot analysis showed that chitinase isoform Chi2 with a molecular weight of 46 kDa was newly induced due to P. fluorescens isolate Pf1 treatment challenged with the pathogen. TLP isoform with molecular weight of 33 kDa was induced not only in P. fluorescens isolate Pf1-treated root tissues challenged with the pathogen but also in roots treated with P. fluorescens isolate Pf1 alone and roots inoculated with the pathogen. These results suggest that induction of defense enzymes involved in phenylpropanoid pathway and accumulation of phenolics and PR-proteins might have contributed to restriction of invasion of F. oxysporum f. sp. lycopersici in tomato roots.     

Genetic loci controlling lethal cell death in tomato caused by viral satellite RNA infection

Cucumber mosaic virus (CMV) associated with D satellite RNA (satRNA) causes lethal systemic necrosis (LSN) in tomato (Solanum lycopersicum), which involves programmed cell death. No resistance to this disease has been found in tomato. We obtained a line of wild tomato, S. habrochaitis, with a homogeneous non-lethal response (NLR) to the infection. This line of S. habrochaitis was crossed with tomato to generate F1 plants that survived the infection with NLR, indicating that NLR is a dominant trait. The NLR trait was successfully passed on to the next generation. The phenotype and genotype segregation was analyzed in the first backcross population. The analyses indicate that the NLR trait is determined by quantitative trait loci (QTL). Major QTL associated with the NLR trait were mapped to chromosomes 5 and 12. Results from Northern blot and in situ hybridization analyses revealed that the F1 and S. habrochaitis plants accumulated minus-strand satRNA more slowly than tomato, and fewer vascular cells were infected. In addition, D satRNA-induced LSN in tomato is correlated with higher accumulation of the minus-strand satRNA compared with the accumulation of the minus strand of a non-necrogenic mutant D satRNA.     

Inheritance and genetic mapping of fruit sucrose accumulation in Lycopersicon chmielewskii

Fruit of the domestic tomato (Lycopersicon esculentum Mill.) accumulate soluble sugars primarily in the form of the hexoses, glucose and fructose. In contrast, the predominant sugar in fruit of the wild tomato relative, L. chmielewskii, is sucrose. In the present study, the inheritance and linkage relations of sucrose accumulation were examined in interspecific L. esculentum x L. chmielewskii populations. In backcrosses to either the wild or domestic tomato, segregation for sucrose accumulation permitted qualitative analysis of the trait and indicated monogenic recessive control, although deviations from Mendelian inheritance were observed in some populations. This major gene, designated sucr, was mapped in F₂, F₃, and BC₁F₂ populations using a set of 95 informative RFLP and isozyme markers covering the tomato genome. A map location near the centromere of chromosome 3 was established, with tight linkage to the genomic clone TG102. Association of sucrose accumulation with yellow fruit, encoded by an allele of the r gene, permitted alignment with the classical map, thereby confirming the map location of sucr. A linkage map of the region surrounding sucr was obtained by monitoring recombination between flanking markers in the back-crosses to tomato. A cDNA clone of tomato fruit acid invertase, TIV1, was mapped to TG102 and sucr, with no recombination between the two RFLP markers observed in over 1700 meiotic products. Despite the tight linkage, TG102 and TIV1 hybridize to distinct restriction fragments, hence do not represent the same gene. The genetic data strongly suggest that sucr is an allele of the invertase gene and thus support previous biochemical studies that demonstrated low invertase activity in sucrose-accumulating fruit. L. hisutum, another low-invertase, sucrose-accumulating species, was hybridized with L. chmielewskii and the resulting F₁ plants accumulated sucrose, indicating that genetic control of soluble sugar composition is conserved in these two species.     

Management of root-knot nematode in tomato Lycopersicon esculentum,Mill, with biogas slurry

The effect of biogas slurry application on the severity of root-knot nematode, Meloidogyne incognita, attack on tomato cv. Co-1, was tested in the green house with two levels of biogas slurry: 5% and 10% (w/w) added to soil. Both the number (3 fruits/plant) and fruit yield (35.2 g/plant) of tomato increased significantly with 10% (w/w) biogas slurry. The plants amended with biogas slurry put up more vegetative growth and tended to flower and fruit much earlier than did those of the control. The nematode population in the soil decreased thus decreasing the severity of nematode attack.