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.     

Mutations at the Asc locus of tomato confer resistance to the fungal pathogen Alternaria alternata f. sp. lycopersici

The fungal pathogen Alternaria alternata f. sp. lycopersici produces host-selective AAL-toxins that cause Alternaria stem canker in tomato. Susceptibility to the disease is based on the relative sensitivity of the host to the AAL-toxins and is controlled by the Asc locus on chromosome 3L. Chemical mutagenesis was employed to study the genetic basis of sensitivity to AAL-toxins and susceptibility to fungal infection. Following the treatment of seeds of a susceptible line with ethyl methanesulphonate (EMS), resistant M₂ mutants were obtained. Most plants with induced resistances showed toxin-sensitivity responses that were comparable to those of resistant control lines carrying the Asc locus. In addition, genetic analysis of the mutagenised plants indicated that the mutations occurred at the Asc locus. Furthermore, novel mutants were identified that were insensitive to the AAL-toxins at the seedling stage but toxin-sensitive and susceptible to fungal infection at mature stages. No AAL-toxin-insensitive insertion mutants were identified following a transposon mutagenesis procedure. Molecular mechanisms involved in host defence against A a. lycopersici are discussed.     

Tomato resistance to Alternaria stem canker: localization in host genotypes and functional expression compared to non-host resistance

Summary The Alternaria stem canker resistance locus (Asc-locus), involved in resistance to the fungal pathogen Alternaria alternata f. sp. lycopersici and in insensitivity to host-specific toxins (AAL-toxins) produced by the pathogen, was genetically mapped on the tomato genome. Susceptibility and resistance were assayed by testing a segregating F₂ population for sensitivity to AAL-toxins in leaf bioassays. Linkage was observed to phenotypic markers solanifolium and sunny, both on chromosome 3. For the Asc-locus, a distance of 18 centiMorgan to solanifolium was calculated, corresponding to position 93 on chromosome 3. This map position of the resistance locus turned out to be the same in three different resistant tomato accessions, one Dutch and two American, that are at least 40 years apart. AAL-toxin sensitivity in susceptible and resistant tomato genotypes was compared with AAL-toxin sensitivity in a non-host Nicotiana tabacum during different levels of plant cell development. In susceptible and resistant tomato genotypes, inhibitory effects were demonstrated at all levels, except for leaves of resistant genotypes. However, during pollen and root development, inhibitory effects on susceptible genotypes were larger than on resistant genotypes. In the non-host Nicotiana tabacum, hardly any effects of AAL-toxins were demonstrated. Apparently, a cellular target site is present in tomato, but not in Nicotiana tabacum. It was concluded that three levels of AAL-toxin sensitivity exist: (1) a susceptible host sensitivity, (2) a resistant host sensitivity, (3) a non-host sensitivity, and that the resistance mechanism operating in tomato is different from that operating in Nicotiana tabacum.     

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.     

Genetic and physiological response to fumonisin and AAL-toxin by intact tissue of a higher plant

The differential phytotoxicity of purified AAL-toxin to lines of tomato isogenic for the Asc gene parallels resistance to Alternaria alternata f.sp. lycopersici. This relationship, as reported earlier, is consistent with the role of AAL-toxin as a host-specific toxin with the role of a primary chemical determinant of Alternaria stem canker. Current results indicate the pathogen and the AAL-toxin also can be recovered from ripe fruit with symptoms of the disease known as black mold. Fumonisins are structurally similar to the AAL-toxins but are secreted by Fusarium moniliforme which is taxonomically distinct from A. alternata. F. moniliforme, is not pathogenic on living tomato tissues but was recovered from ripe tomato fruit with symptoms of black mold. The penetration of ripe fruit and subsequent colonization by both fungi appears to be saprophytic. Fumonisins and AAL-toxins express equivalent genotype-specific activity against the isogenic Asc lines of tomato and produce equivalent necrotic symptoms in tomato leaflet bioassays. Evidence was obtained that the biosynthetic pathway for production of these toxins is present in several species of both Alternaria and Fusarium. Toxin biosynthesis was sensitive to nutritional regulation in both genera. However, pathogenicity on tomato was not altered by the medium used for inoculum production in either genera and remained restricted to A. alternata f.sp. lycopersici in the studies reported here. Differences in the amount of toxin produced were found among isolates of both genera while the magnitude of the differences was defined by the substrate on which the fungi were grown.     

Effects of Alternaria alternata f.sp. lycopersici toxins on pollen

Summary Effects of the phytotoxic compounds (AAL-toxins) isolated from cell-free culture filtrates of Alternaria alternata f.sp. lycopersici on in vitro pollen development were studied. AAL-toxins inhibited both germination and tube growth of pollen from several Lycopersicon genotypes. Pollen from susceptible genotypes, however, was more sensitive for AAL-toxins than pollen from resistant plants, while pollen of species not belonging to the host range of the fungus was not significantly affected by the tested toxin concentrations. AAL-toxins elicit symptoms in detached leaf bioassays indistinguishable from those observed on leaves of fungal infected tomato plants, and toxins play a major role in the pathogenesis. Apparently, pathogenesis-related processes and mechanisms involved in disease resistance are expressed in both vegetative and generative tissues. This overlap in gene expression between the sporophytic and gametophytic level of a plant may be advantageously utilized in plant breeding programmes. Pollen may be used to distinguish susceptible and resistant plants and to select for resistances and tolerances against phytotoxins and other selective agents.     

The Asc locus for resistance to Alternaria stem canker in tomato does not encode the enzyme aspartate carbamoyltransferase

The fungal disease resistance locus Alternaria stem canker (Asc) in tomato has been suggested to encode the enzyme aspartate carbamoyltransferase (AC Tase). To test this hypothesis a segment of the tomato ACTase gene was amplified by the polymerase chain reaction (PCR) using degenerate primers. The PCR product obtained was subsequently used to isolate an ACTase cDNA clone. Restriction fragment length polymorphism (RFLP) linkage analysis showed that the ACTase gene and the Asc locus do not cosegregate. RFLP mapping positioned the ACTase gene on chromosome 11, while the Asc locus is located on chromosome 3. These results exclude the possibility that the ACTase protein is encoded by the Asc locus.     

Effects of Alteria alternata f.sp. lycopersici toxins at different levels of tomato plant cell development

Since the host-specific toxins of Alternaria alternata f. sp. lycopersici play an important role in pathogenesis, they potentially could be applied as selective agents in in vitro selection at the cellular level for disease resistance. Prerequisite for this is that sensitivity to the Alternaria alternata f.sp. lycopersici pathotoxins is manifest at the cellular level. To gain insight into cellular effects of AAL-toxins and into the mechanisms of plant insensitivity to AAL-toxins, effects of AAL-toxins on leaves, leaf discs, roots, calli, suspension cells, minicalli and protoplasts of susceptible and resistant tomato genotypes were studied. In leaves of susceptible genotypes, toxins cause severe necrosis, while in leaves of resistant genotypes necrosis was never observed. Inhibition effects of toxins were observed at all other levels in susceptible and resistant genotypes: toxins inhibited shoot induction on leaf discs, root growth and growth of calli, suspension cells and protoplasts. This indicates a cellular site for AAL-toxins. Differences in sensitivity to AAL-toxins between susceptible and resistant genotypes were observed in leaves and roots, but were not observed during shoot induction on leaf discs, in calli, suspension cells and protoplasts. However, differences in sensitivity to AAL-toxins in roots were at least 20 times less than in leaves. Therefore insensitivity seems related to a higher level of tomato plant differentiation and is most pronounced in leaves.     

Genetic and physical analysis of a YAC contig spanning the fungal disease resistance locus Asc of tomato (Lycopersicon esculentum)

The Alternaria stem canker disease of tomato is caused by the necrotrophic fungal pathogen Alternaria alternata f. sp. lycopersici (AAL). The fungus produces AAL toxins that kill the plant tissue. Resistance to the fungus segregates as a single locus, called Asc, and has been genetically mapped on chromosome 3 of tomato. We describe here the establishment of a 1383-kb YAC contig covering the Asc locus and a series of plants selected for recombination events around the Asc locus. It was shown that the YAC contig corresponds to a genetic distance of at least 11.2 cM. Thus, the recombination rate in the Asc region is six times higher (123 kb/cM) than the average for the tomato genome. Furthermore, the Asc locus could be localised to a 91-kb fragment, thus paving the way for the cloning and identification of the Asc gene(s) by complementation. Received: 31 July 1998 / Accepted: 6 October 1998     

Trichome-borne and artificially applied acylsugars of wild tomato deter feeding and oviposition of the leafminer Liriomyza trifolii

Oviposition and adult feeding of the leafminer Liriomyza trifollii (Burgess) (Diptera, Agromyzidae) on Lycopersicon pennellii (Corr.) D'Arcy and its F₁ hybrid with Lycopersicon esculentum (Mill.) was significantly less than that on the cultivated tomato, L. esculentum. The resistance of L. pennellii and the F₁ was reduced following rinsing of foliage with ethanol. Resistant attributes of L. pennellii were transferred to L. esculentum through appression of L. pennellii foliage to L. esculentum leaflets. Application of purified 2,3,4-tri-O-acylglucoses (the principal component of type IV glandular trichome exudate of L. pennellii) to L. esculentum significantly decreased feeding and oviposition on L. esculentum leaflets by 61–99%. Therefore the principal mechanism of resistance to this leafminer by L. pennellii is the secretion of these acylglucoses. Dose response analysis of acylglucoses applied to L. esculentum shows that dosages as low as 10% those found on L. pennellii provide large reductions (91%) in leaf punctures and mines.     

Chromosome association and pollen fertility of parental and interspecific hybrids of Lycopersicon esculentum X L. hirsutum and L. pennellii

Cytological studies were carried out on two wild species (L. hirsutum and L. pennellii) and the cultivated species (L. esculentum) of tomato and their F1 hybrids. Both parents and hybrids show a diploid chromosome number of 2n=24. The meiotic behaviour of the cultivated species showed a high degree of chromosome homology resulting in a high level of chiasmata frequency per bivalent. In contrast, the two wild species showed a slight increase in uniyalent frequency and a decrease in bivalent formation and chiasmata frequency. The meiotic behaviour of the hybrids showed a high level of univalents and low levels of bivalents as well as trivalents. Highly significant decreases in chiasmata frequency and increases in meiotic abnormalities, especially in the L. esculentum X L. pennellii hybrid, also were detected. The high meiotic irregularity and low chiasmata frequency recorded in the second hybrid indicated the disharmony and difference between its parental genomes and also served to predict its sterility. With regard to degree of pairing recorded in the hybrids, there is a possibility that sterility in such cases may refer to genetic factors in addition to the previously mentioned reasons. Pollen fertility showed no great difference between L. esculentum and L. hirsutum and their F1 hybrid, but a significant decrease was recorded in the L. esculentum X L. pennellii hybrid, which was clearly associated with high meiotic irregularity, low chiasmata frequency and chromosome association.     

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.     

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.     

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     

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.     

Responses to NaCl stress of cultivated and wild tomato species and their hybrids in callus cultures

Summary If in vitro culture is to be used for evaluating the salt tolerance of tomato hybrids and segregant populations in a breeding programme, it is previously necessary to get quick and reliable traits. In this work, growth and physiological responses to salinity of two interspecific hybrids between the cultivated tomato (Lycopersicon esculentum Mill) and its wild salt-tolerant species L pennellii are compared to those of their parents. The leaf callus of the first subculture was grown on media amended with 0, 35, 70, 105, 140, 175 and 210 mM NaCl for 40 days. Relative fresh weight growth of callus in response to increased salinity in the culture medium was much greater in L pennellii than in the tomato cultivars, and greater in the hybrids than in the wild species. Moreover, the different salt tolerance degree of hybrids was related to that of female parents. At high salt levels, only Cl^– accumulation was higher in L pennellii than in tomato cultivars, whereas in the hybrids both Cl^–, and Na⁺ accumulation were higher than in their parents. Proline increased with salinity in the callus of all genotypes; these increases were much higher in the tomato cultivars than in L pennellii, and the hybrids showed a similar response to that of the wild species. Salt-treated callus of the tomato cultivars showed significant increases in valine, isoleucine and leucine contents compared to control callus tissue. In contrast, these amino acids in callus tissues of the wild species and hybrids showed a tendency to decrease with increasing salinity.     

Genetic analysis of RFLPs, GATA microsatellites and RAPDs in a cross between L. esculentum and L. pimpinellifolium

A population of 257 BC₁ plants was developed from a cross between an elite processing line of tomato (Lycopersicon esculentum cvM82-1-7) and the closely related wild species L. pimpinellifolium (LA1589). The population was used to construct a genetic linkage map suitable for quantitative trait locus (QTL) analysis to be conducted in different backcross generations. The map comprises 115 RFLP, 3 RAPD and 2 morphological markers that span 1279 cM of the tomato genome with an average distance between markers of 10.7 cM. This map is comparable in length to that of the highdensity RFLP map derived from a L. esculentum x L. pennellii F₂ population. The order of the markers in the two maps is also in good agreement, however there are considerable differences in the distribution of recombination along the chromosomes. The segregation of six GATA-containing loci and 47 RAPD markers was also analyzed in subsets of the population. All of the microsatellite loci and 35 (75%) of the RAPDs mapped to clusters associated with centromeric regions.     

RAPD markers for constructing intraspecific tomato genetic maps

The existing molecular genetic maps of the tomato, Lycopersicon spp, are constructed based on isozyme and RFLP polymorphisms between tomato species. These maps are useful for certain applications but have few markers that exhibit sufficient polymorphisms for intraspecific analysis and manipulations within the cultivated tomato. The purpose of this study was to investigate the relative potential of RAPD technology, as compared to isozymes and RFLPs, to generate polymorphic DNA markers within cultivated tomatoes. Sixteen isozymes and 25 RFLP clones that were known to detect polymorphism between L. esculentum and L. pennellii, and 313 random oligonucleotide primers were examined. None of the isozymes and only four of the RFLP clones (i.e., 16%) revealed polymorphism between the cultivated varieties whereas up to 63% of the RAPD primers detected one or more polymorphic DNA fragments between these varieties. All RAPD primers detected polymorphism between L. esculentum and L. pennellii genotypes. These results clearly indicate that RAPD technology can generate sufficient genetic markers exploiting sequence differences within cultivated tomatoes to facilitate construction of intraspecific genetic maps.Abbreviations RFLP restriction fragments length polymorphism - RAPD random amplified polymorphic DNA - PCR polymerase chain reaction - QTLs quantitative trait loci     

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     

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