Feeding behavior of potato aphid affected by glandular trichomes of wild tomato

Mortality of the potato aphid, Macrosiphum euphorbiae (Thomas), on Lycopersicon pennellii (Corr.) D'Arcy and its F₁ hybrid with Lycopersicon esculentum Mill. was significantly greater than that on L. esculentum. Physical entrapment was not the sole mechanism of resistance in L. pennellii since few late instar aphids were found trapped in the sticky glandular exudate of the type IV trichomes; entrapment could, however, affect survival of early instars. Aphid settling on L. pennellii was dramatically less than that on L. esculentum, suggesting that starvation may have contributed to high mortality. Compared to L. esculentum, aphid feeding behavior on L. pennellii and the F₁ was characterized by a delay in the time to first probe, a reduction in the number of probes, and a decrease in the total proportion of time spent feeding. Removal of the glandular exudate of the type IV trichomes from L. pennellii resulted in a decrease in preprobe time and an increase in both the number of probes and the percent of time spent probing. Transfer of glandular trichome exudate of L. pennellii to leaflets of L. esculentum resulted in an increase in resistance as measured by these three parameters.

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Zusammenfassung Die Absterberate der Kartoffellaus, Macrosiphum euphorbiae Thomas, auf Lycopersicon pennellii (Corr.) D'Arcy, sowie auf der Kreuzung L. esculentum Mill. und L. pennellii, war deutlich grösser als auf L. esculentum. Das mechanische Verfangen der Läuse war nicht der Hauptgrund der Resistenz von L. pennellii. Wenige tote Läuse wurden in dem klebrigen Sekret der Typus IV Trichome gefunden. Auf L. pennellii siedelten sich die Läuse in viel geringerer Zahl an als auf L. esculentum. Dies führte zum Schluss, dass Verhungern eine der Ursachen der hohen Mortalität der Läuse war. Im Vergleich zum Saugverhalten auf L. esculentum war das Saugverhalten auf L. pennellii, wie auch auf F₁, durch Folgendes gekennzeichnet 1) Verspätung des ersten Stichversuchs, 2) Verminderung der Stichversuche pro Zeiteinheit und 3) Verminderung des Zeitanteils, der zum Saugen verwendet wurde. Die Entfernung des Sekrets der Typus IV Trichomen auf L. pennellii verursachte 1) eine kürzere Zeitspanne vor dem ersten Stichversuch, 2) eine Vergrösserung der Anzahl Stichversuche pro Zeiteinheit, 3) eine Verlängerung der Saugzeit. Die Uebertragung des Sekretes von L. pennellii auf Blätter von L. esculentum verbesserte deren Resistenz gegen Blattläuse gemessen mit den genannten drei Kriterien.

     

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     

fw 2.2:a major QTL controlling fruit weight is common to both red- and green-fruited tomato species

We have shown that a major QTL for fruit weight (fw2.2) maps to the same position on chromosome 2 in the green-fruited wild tomato species, Lycopersicon pennellii and in the red-fruited wild tomato species, L. pimpinellifolium. An introgression line F₂ derived from L. esculentum (tomato) x L. pennellii and a backcross 1 (BC₁) population derived from L. esculentum x L. pimpinellifolium both place fw2.2 near TG91 and TG167 on chromosome 2 of the tomato highdensity linkage map. fw2.2 accounts for 30% and 47% of the total phenotypic variance in the L. pimpinellifolium and L. pennellii populations, respectively, indicating that this is a major QTL controlling fruit weight in both species. Partial dominance (d/a of 0.44) was observed for the L. pennellii allele of fw 2.2 as compared with the L. esculentum allele. A QTL with very similar phenotypic affects and gene action has also been identified and mapped to the same chromosomal region in other wild tomato accessions: L. cheesmanii and L. pimpinellifolium. Together, these data suggest that fw2.2 represents an orthologous QTL (i.e., derived by speciation as opposed to duplication) common to most, if not all, wild tomato species. High-resolution mapping may ultimately lead to the cloning of this key locus controlling fruit development in tomato.     

Mapping the Sw-5 locus for tomato spotted wilt virus resistance in tomatoes using RAPD and RFLP analyses

The Sw-5 locus confers dominant resistance to tomato spotted wilt virus (TSWV). To map the location and facilitate the identification of markers linked to Sw-5 we developed a pair of near-isogenic lines (NILs) and an F₂ Lycopersicon esculentum x L. pennellii population segregating for resistance to TSWV. DNA from the NILs was analyzed using 748 random 10-mer oligonucleotides to discern linked molecular markers using a random amplified polymorphic DNA (RAPD) approach. One random primer (GAGCACGGGA) was found to produce a RAPD band of about 2200 bp that demonstrates linkage to Sw-5. Data from co-segregation of resistance and restriction fragment length polymorphisms (RFLPs) in a F₂ interspecific population position Sw-5 between the markers CT71 and CT220 near the telomere of the long arm of chromosome 9.     

Inhibition of Telenomus sphingis an egg parasitoid of Manduca spp. by trichome/2-tridecanone-based host plant resistance in tomato

The resistance of accession PI 134417 of the wild tomato Lycopersicon hirsutum f. glabratum C. H. Mull to Manduca sexta (L.) (Lepidoptera: Spingidae) and Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) is conditioned by the high densities of 2-tridecanone-containing, glandular trichomes associated with the foliage. In laboratory experiments, rates of parasitism of M. sexta eggs by Telenomus sphingis (Ashmead) (Hymenoptera: Scelionidae) were lower among eggs on PI 134417 foliage than among eggs on foliage of the cultivated tomato L. esculentum Mill. (cv. Better Boy). The latter is characterized by a significantly lower density of type VI glandular trichomes than PI 134417 and an absence of 2-tridecanone. Parasitism by T. sphingis was also reduced among eggs on foliage of the F₁ hybrid between PI 134417 and L. esculentum. The hybrid foliage lacks 2-tridecanone but has a density of type VI glandular trichomes that is intermediate between those of PI 134417 and L. esculentum, indicating that elevated densities of type VI glandular trichomes adversely affect T. sphingis. This conclusion was further substantiated by the finding that there were no differences among plant lines in the levels of parasitism of M. sexta eggs when the eggs were on foliage that had been divested of glandular trichomes.In bioassays in which T. sphingis adults or immatures in host eggs were exposed to filter paper treated with 2-tridecanone at rates comparable to those associated with PI 134417 foliage, 2-tridecanone was acutely toxic and caused high levels of mortality. In addition, at high concentrations, 2-tridecanone vapors were repellent to T. sphingis adults. However, when exposed to PI 134417 foliage, few T. sphingis adults were killed.Parasitism of M. sexta eggs was unaffected when the eggs were deposited by moths reared as fifth instar larvae on diet containing 2-tridecanone and/or 2-undecanone at levels comparable to those associated with PI 134417 foliage.     

Resistance to powdery mildew (Oidium lycopersicum) in Lycopersicon hirsutum is controlled by an incompletely-dominant gene Ol-1 on chromosome 6

The inheritance of resistance to powdery mildew (Oidium lycopersicum) in Lycopersicon hirsutum was investigated by disease tests in segregating populations obtained by hybridising tomato (L. esculentum) cv Moneymaker with the wild relative L. hirsutum G1.1560. One incompletely dominant gene Ol-1 was found to largely control resistance to the disease. To map Ol-1, DNA pools from seven resistant and ten susceptible F₂ plants were analyzed for random amplified polymorphic DNA (RAPD). With 32 primers tested, one RAPD, primed with the sequence 5-GACGTGGTGA-3, was observed between the susceptible and the resistant bulks, which cosegregated with resistance in the F₂ population of L. esculentum × L. hirsutum G1.1560. This RAPD was mapped on chromosome 6 by using an F₂ (L. esculentum × L. pennellii) already mapped for 49 RFLPs. RFLP analysis of the F₂ from L. esculentum cv Moneymaker × L. hirsutum G1.1560 demonstrated that Ol-1 maps near the Aps-1 region on chromosome 6, in the vicinity of the resistance genes to Meloidogyne spp. (Mi) and to Cladosporium fulvum (Cf-2/Cf-5).     

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     

Molecular analysis of the extent of asymmetry in two asymmetric somatic hybrids of tomato

Summary Two somatic hybrid plants generated from a single fusion event between Lycopersicon esculentum and irradiated L. pennellii protoplasts have been analyzed at the molecular level. Over 30 loci have been analyzed using isozymes and RFLPs. All loci tested on chromosomes 2–10 were heterozygous, while those loci on chromosome 12 were homozygous L. pennellii in both somatic hybrids. In one of the somatic hybrids, 2850, loci on chromosome 1 were also homozygous L. pennellii. The other somatic hybrid, 28F5, was heterozygous at all chromosome 1 loci tested, but exhibited altered stoichiometry of parental bands as compared to the sexual hybrid. Loci on chromosome 2 from both somatic hybrids have altered stoichiometry, with L. pennellii alleles being four times more abundant than expected. Both somatic hybrids contain the L. esculentum chloroplast genome, while only L. pennellii polymorphisms have been detected in the mitochondrial genome.     

Heat Stress and Spermidine: Effect on Chlorophyll Fluorescence in Tomato Plants

Two tomato (Lycopersicon esculentum L.) cultivars: Robin (tolerant) and Roma (sensitive to heat stress) were studied. Chlorophyll fluorescence induction parameters (F_v/F_p, A_max, and Rfd) at 25 °C showed that the PS2 activity was similar for both cultivars. The parameters, measured at 38 °C, decreased in both cultivars, but more in cv. Roma. Exogenous application of 4 mM spermidine improved the plant heat-resistance in both cultivars, and especially in cv. Roma. Analysis of chlorophyll fluorescence changes during linear increase in temperature showed that cv. Robin plants have higher ability to hardening and higher resistance to thermal damage of the pigment-protein complexes structure and the activity of PS2 than cv. Roma.     

RFLP mapping of I1, a new locus in tomato conferring resistance against Fusarium oxysporum f. sp. lycopersici race 1

Summary The inheritance and linkage relationships of a gene for resistance to Fusarium oxysporum f. sp. lycopersici race 1 were analyzed. An interspecific hybrid between a resistant Lycopersicon pennellii and a susceptible L. esculentum was backcrossed to L. esculentum. The genotype of each backcross-1 (BC₁) plant with respect to its Fusarium response was determined by means of backcross-2 progeny tests. Resistance was controlled by a single dominant gene, I1, which was not allelic to I, the traditional gene for resistance against the same fungal pathogen that was derived from L. pimpinellifolium. Linkage analysis of 154 molecular markers that segregated in the BC₁ population placed I1 between the RFLP markers TG20 and TG128 on chromosome 7. The flanking markers were used to verify the assignment of the I1 genotype in the segregating population. The results are discussed with reference to the possibility of cloning Fusarium resistance genes in tomato.     

Expression of unilateral incompatibility in pollen of Lycopersicon pennellii is determined by major loci on chromosomes 1, 6 and 10

Summary We have previously described gene introgression from the wild nightshade Solanum lycopersicoides into tomato (Lycopersicon esculentum) through the use of either diploid or sesquidiploid hybrids (the latter consisting of two genomes of L. esculentum and one genome of S. lycopersicoides). Both types of intergeneric hybrids display pollen sterility, but workable ovule fertility. Unilateral incompatibility prevents their direct hybridization with staminate L. esculentum. Pollen of a self-compattible form of the related wild species L. pennellii is compatible with pistils of L. esculentum x S. lycopersicoides hybrids. This trait was backcrossed from L. pennellii to L. esculentum in order to develop bridging lines that could be used to obtain progeny from the intergeneric hybrids and to study the inheritance of bridging ability. In progeny of L. esculentum x S. lycopersicoides hybrids pollinated with L. pennellii-derived bridging lines, preferential transmission of L. pennellii alleles was observed for certain isozyme and RFLP markers on chromosomes 1, 6 and 10. The skewed segregations suggest linkage to three major pollen-expressed compatibility loci. This was confirmed by observations of pollen tube growth, which indicated that compatibility with pistils of the diploid intergeneric hybrid occurred only in bridging lines at least heterozygous for the L. pennellii markers on chromosomes 1, 6 and 10. Compatibility with the sesquidiploid hybrid required only the chromosome 1 and 6 loci, indicating an apparent effect of gene dosage on expression of incompatibility in the pistil. In an F₂ L. esculentum x L. pennellii population, preferential transmission of L. pennellii alleles was observed for the same markers on chromosomes 1 and 10, as well as other markers on chromosomes 3, 11, and 12, but not 6. The chromosome 1 pollen compatibility locus maps to or near the S-locus, which determines S-allele specificity. The results are discussed in relation to existing genetic models for unilateral incompatibility, including the possible involvement of the S-locus.     

Inheritance of resistance to the two-spotted spider mite from L. pimpinellifolium (Jusl.) Mill. accession ‘TO-937’

The two-spotted spider mite (Tetranychus urticae Koch) is an important pest of tomato (Lycopersicon esculentum Mill.) crops in temperate regions as this spider mite has a very large capacity for population increase and causes severe tomato yield losses. There is no described tomato cultivar fully resistant to this pest, although resistant accessions have been reported within the green-fruited tomato wild species L. pennellii (Corr.) D’Arcy and L. hirsutum Humb. & Bonpl. We observed a L. pimpinellifolium (Jusl.) Mill. accession, ‘TO-937’, which seemed to be completely resistant to mite attacks and we crossed it with the susceptible L. esculentum cultivar. ‘Moneymaker’ to obtain a family of generations consisting of the two parents, the F₁, the F₂, the BC₁ to L. esculentum, and the BC₁ to L. pimpinellifolium. This family was evaluated for mite resistance in a polyethylene greenhouse using an experimental design in 60 small complete blocks distributed along 12 double rows. Each block consisted of five F₂ plants in one row and one plant of each of the two parents, the F₁, the BC₁ to L. esculentum, and the BC₁ to L. pimpinellifolium in the adjacent row. Plants at the 10–15 leaf stage were artificially infested by putting on them two pieces of French bean leaf heavily infested with T. urticae. After two months, evaluations of infestation were made by visual observation of mite nets and leaf damage. Plants that were free of signs of mite reproduction on the top half were considered as resistant, plants with silky nets only on their basal leaves, intermediate, and plants with mite reproduction on both basal and top canopies were scored as susceptible. Dominance for resistance appeared because all the ‘To-937’, BC₁ to L. pimpinellifolium, and F₁ plants were resistant. Not all ‘Moneymaker’ plants behaved as susceptible because 35% of plants were intermediate. In the BC₁ to L. pimpinellifolium and the F₂, most plants were scored as resistant, only 7 % BC₁ and 3 % F₂ plants were intermediate, and a single F₂ plant (0.3 %) was susceptible. With these figures, resistance seemed to be controlled by either four or two genes according to whether segregation in the BC₁ or in the F₂, respectively, were considered. These results could in part be explained because of appearance of negative interplot interference due to the high frequency of resistant genotypes within most of the generations. Therefore, the family was evaluated again but using a different experimental design. In the new experiment, 16 ‘TO-937’, 17 ‘Moneymaker’, 17 F₁, 37 BC₁ to L. pimpinellifolium, 38 BC₁ to L. esculentum, and 125 F₂ plants were included. Each of these test plants was grown besides a susceptible ‘Moneymaker’ auxilliary plant that served to keep mite population high and homogeneous in the greenhouse. Negative interplot interference was avoided with this design and all the ‘TO-937’, F₁, and BC₁ to L. pimpinellifolium plants were resistant, all ‘Moneymaker’ test plants were susceptible, and 52 % BC₁ to L. esculentum and 25 % F₂ plants were susceptible, which fitted very well with the expected for resistance governed by a single dominant gene. The simple inheritance mode found will favour sucessful introgression of mite resistance into commercial tomatoes from the very close relative L. pimpinellifolium.     

Inheritance and genetic mapping of resistance to Alternaria alternata f. sp. lycopersici in Lycopersicon pennellii

The fungal pathogen Alternaria alternata f. sp. lycopersici produces AAL-toxins that function as chemical determinants of the Alternaria stem canker disease in the tomato (Lycopersicon esculentum). In resistant cultivars, the disease is controlled by the Asc locus on chromosome 3. Our aim was to characterize novel sources of resistance to the fungus and of insensitivity to the host-selective AAL-toxins. To that end, the degree of sensitivity of wild tomato species to AAL-toxins was analyzed. Of all members of the genus Lycopersicon, only L. cheesmanii was revealed to be sensitive to AAL-toxins and susceptible to fungal infection. Besides moderately insensitive responses from some species, L. pennellii and L. peruvianum were shown to be highly insensitive to AAL-toxins as well as resistant to the pathogen. Genetic analyses showed that high insensitivity to AAL-toxins from L. pennellii is inherited in tomato as a single complete dominant locus. This is in contrast to the incomplete dominance of insensitivity to AAL-toxins of L. esculentum. Subsequent classical genetics, RFLP mapping and allelic testing indicated that high insensitivity to AAL-toxins from L. pennellii is conferred by a new allele of the Asc locus.     

Lycopersicon esculentum lines containing small overlapping introgressions from L. pennellii

Summary The objective of this project was to introgress small overlapping chromosome segments which cover the genome of L. pennellii into Lycopersicon esculentum lines. The interspecific hybrid was backcrossed to L. esculentum, and a map of 981 cM, based on 146 molecular markers covering the entire genome, was produced. A similar backcross 1 population was selfed for six generations, under strong selection for cultivated tomato phenotypes, to produce 120 introgression lines. The introgression lines were assayed for the above-mentioned molecular markers, and 21 lines covering 936 cM of L. pennellii, with an average introgression of 86 cM, were selected to provide a resource for the mapping of new DNA clones. The rest of the lines have shorter introgressions consisting of specific regions with an average size of 38 cM. The proportion of the L. pennellii genome in the introgression lines was lower than expected (252 cM) because of strong selection against the wild-parent phenotype. The mean introgression rate for ends of linkage groups in the 120 lines was 3 times higher than for other regions of the genome. The introgression lines can assist in RFLP-based gene cloning by allowing the rapid selection of DNA markers that map to specific chromosome segments. The introgression lines also provide a base population for the mapping and breeding for quantitative traits such as salt and drought tolerance that characterize the wild species L. pennellii.     

Inheritance of potato aphid resistance in hybrids between Lycopersicon esculentum and L. pennellii

Summary The potato aphid, Macrosiphum euphorbiae Thomas, is an important pest of tomato, Lycopersicon esculentum Mill., because it transmits tomato viruses and directly reduces crop yields by its feeding. This study was conducted to determine whether the wild tomato species, Lycopersicon pennellii (Corr.) D'Arcy, would be useful as a source of potato aphid resistance for tomato. Type IV trichome density and aphid resistance were assessed in six generations (P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂) from crosses between L. pennellii (LA 716) and two tomato cultivars, New Yorker and VF Vendor. Weighted leastsquares were used in joint scaling tests to estimate the relative importance of gene effects on type IV trichome density and potato aphid resistance of the hybrids. A simple additive-dominance model adequately explained the variation in type IV trichome density. Models which included digenic epistatic effects were required to explain the variation in aphid resistance. Standard unit heritability estimates of aphid resistance in the backcross to L. esculentum were obtained by regression of BC₁F₂ off-spring families on BC₁F₁ parents. Regression coefficients and heritability estimates varied between years with the level and uniformity of the aphid infestation. In the 1985–1986 growing seasons, when aphid infestations were uniform, aphid resistance exhibited a moderate level of heritability (29.8% ± 14.1% and 47.1% ± 11.5% in New Yorker and VF Vendor backcross populations, respectively). The non-uniform aphid infestation of 1984 resulted in lower heritability estimates in the 1984–1985 growing seasons (16.1% ± 15.7% and 21.9% ± 14.8% in the New Yorker and VF Vendor backcross populations, respectively). Selection for potato aphid resistance would probably be most efficient if it were delayed until gene combinations are fixed in later generations, because of the large epistatic effects and the low heritability of this trait in seasons with variable aphid infestations.     

Mapping of ripening-related or -specific cDNA clones of tomato (Lycopersicon esculentum)

Summary Nineteen ripening-related or -specific clones from Lycopersicon esculentum were mapped via RFLP analysis using an F₂ population from the cross L. esculentum x L. pennellii and cDNA or genomic clones of known map location. The map produced using cDNA and genomic clones of known map location corresponded well with previously published maps of tomato. The number of loci detected for each ripening-related or-specific clone varied from one to seven. These loci were located on all 12 chromosomes of the tomato genome. There was no significant clustering of ripening-related or-specific genes. Regions of very low recombination were observed. The clone for polygalacturonase (TOM6) mapped to a single region on chromosome 10, the same chromosome as the nor and alc ripening mutants. To fine map this chromosome, two backcross populations were produced from the cross of L. esculentum x L. pimpenillifolium, in which the esculentum parents used were homozygous for either the alc or the nor. The coding region for polygalacturonase is functionally unlinked to either of these two ripening mutants.     

Integration of the classical and RFLP linkage maps of the short arm of tomato chromosome 1

The classical map of the short arm of chromosome 1 of tomato (Lycopersicon esculentum) has been shown to contain inaccuracies while the RFLP map of this region is known to be generally accurate. Molecular analysis of populations derived from crosses between L. esculentum lines carrying chromosome 1 classical markers and L. pennellii has enabled us to produce an integrated classical and RFLP marker map of this region. New data concerning the linkage relationships between classical markers have also been combined with previous data to produce a new classical map of the short arm of chromosome 1. The orders of the classical markers on these two new maps are in almost complete agreement and are very different to that shown on the previous classical map.     

Osmotic adjustment in Lycopersicon esculentum and L. Pennellii under NaCl and polyethylene glycol 6000 iso–osmotic stresses

Changes in leaf solute contents in response to saline (NaCl) and osmotic (polyethylene glycol, PEG, 6000) stresses were measured in three different salt tolerant cultivars of Lycopersicon esculentum (L.) Mill. (Pera, P-73 and Volgogradskij), and its wild relative L. pennellii (Correll) D'Arcy accession PE-47. Iso-osmotic stresses (–0. 5 MPa) of NaCl (140 mM) and PEG 6000 (150 g l^-1) were applied to one-month old plants for 3 weeks. Decreasing leaf dry weight was similar in L. pennellii or L. esculentum cv. P-73 and Volgogradskij under both stresses, while leaf dry weight of L. esculentum cv. Pera decreased more under PEG stress than under NaCl stress. Water contents decreased in all the PEG treated populations, while their calculated solute potential (Ψ_s increased. Under osmotic stress, the total ion contents decreased in relation to control, whereas organic solutes (sugars, amino acids and organic acids) markedly increased in both tomato species, specially in the tomato cultivars, where these solutes represented 50% of the Ψ₅ calculated. Soluble sugar increase was three times higher in leaves of L. esculentum than in the leaves of L. pennellii. Free proline increased under both stresses and its content was highest in L. esculentum and in L. pennellii, respectively, under NaCl and PEG stresses. Nevertheless, the contribution of this metabolite to Ψ_s did not exceed 5%, irrespective of treatment and species. The greater organic solute accumulation in L. esculentum than in L. pennellii– which was not reflected in their Ψ₅ values – was not correlated with the tolerances of the two species to osmotic stress. Therefore, osmotic adjustment may not be the only process influencing salt and drought tolerances in tomato; the ability of plants to regulate their metabolic and physiological functions could also play an important role under these harmful conditions. The possible roles of inorganic solutes and metabolites in osmotic adjustment, energetic metabolism and redox regulation are discussed     

High-resolution linkage analysis and physical characterization of the EIX-responding locus in tomato

An ethylene-inducing xylanase (EIX) from Tricohoderma viride is a potent elicitor of ethylene biosynthesis, localized cell death and other defense responses in specific cultivars of tobacco (Nicotiana tabacum) and tomato (Lycopersicon esculentum). Wild species of tomato, such as Lycopersicon cheesmanii and Lycopersicon pennellii, do not respond to EIX treatment. The F₁ progeny of a L. esculentum×L. cheesmanii and a L. esculentum×L. pennellii cross responded to EIX treatment with an increase in ethylene biosynthesis and the induction of localized cell death. The F₂ progeny of the above mentioned crosses segregated 3:1 (responding:non-responding). We mapped the EIX-responding locus (Eix) to the short arm of chromosome 7 using a population of introgression lines (ILs), containing small RFLP-defined chromosome segments of L. pennellii introgressed into L. esculentum. RFLP analysis of 990 F₂ plants that segregated for the introgressed segment mapped the Eix locus 0.1 cM and 0.9 cM from the flanking markers TG61 and TG131, respectively. Using the marker TG61 we isolated a yeast artificial chromosome (YAC) clone that carries 300-kb DNA segments derived from the Eix region. By mapping the ends of this YAC clone we show that it spans the Eix locus. Thus, positional cloning of the Eix locus appears feasible. Received: 20 March 1999 / Accepted: 30 April 1999     

Alterations of the manifestations of hybrid breakdown in Lycopersicon esculentum L. pennellii F2 populations containing L. esculentum versus L. pennellii cytoplasm

Reproductive abnormalities reduced the percent stainable pollen, and fruit and seed set in interspecific F₂ populations derived from crosses of Lycopersicon esculentum and L. pennellii but were not observed in parental lines and interspecific F₁ populations. The degree to which these reproductive abnormalities were expressed in the interspecific F₂ populations was affected by cytoplasm. Reproduction was impeded in interspecific F₂ populations containing L. esculentum cytoplasm (F ₂ ^Le ) by reduction in pollen production, the lack of fruit set and a high proportion of parthenocarpic fruit among plants capable of fruit set. The F₂ populations containing L. pennellii cytoplasm (F ₂ ^Lp4 ) showed a reduced frequency of reproductive abnormalities at all stages of reproductive development, resulting in higher values for percent stainable pollen, fruit and seed set and higher proportions of the F ₂ ^Lp4 populations being capable of setting fruit or seed than F ₂ ^Le populations. The major barrier remaining in F ₂ ^Lp4 populations was reduced fruit set compared to parental lines. The barrier to fruit and seed set observed in the F ₂ ^Le populations, and to a lesser extent in the F ₂ ^Lp4 populations, occurs around the time of fertilization or early embryonic development. The effect of L. pennellii cytoplasm on barriers in the F ₂ ^Lp4 populations is proposed to be due to an interaction between cytoplasmic and nuclear genes during fertilization of the F₁ plants to produce F₂ populations and may also affect subsequent generations.