Resistance of wild Lycopersicon species to the potato moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae)

Abstract  Trichome-based host plant resistance of Lycopersicon species to potato moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae), was examined in a laboratory bioassay. Neonate P. operculella were placed on the foliage of accessions of L. hirsutum , L. hirsutum f. glabratum , L. pennellii , L. cheesmanii f. minor and L. esculentum for 48 h . At the end of this period, larval mortality, the numbers of larvae emigrating from, and mining into, the leaf surface were recorded as were leaf area and the densities of trichomes. Analysis by categorical logistic regression found accessions differed in levels of resistance to P. operculella ( P  < 0.001). Further analysis using generalised linear models showed mortality was associated with high densities of type VI and low densities of type V trichomes; emigration from the leaf was associated with high densities of type I and type VII trichomes; and the numbers of mines associated with low densities of type I and type VI trichomes and increased leaf area. Although results indicate that certain accessions of L. hirsutum and L. hirsutum f. glabratum may be most appropriate for inclusion in a breeding program aimed at introducing trichome-based host plant resistance of wild Lycopersicon species into the tomato, further research is necessary to fully understand the mechanisms.     

Galactolipase activity and free fatty acid levels in chloroplasts of domestic and wild tomatoes with different chilling tolerance

In order to establish differences in the chilling sensitivity of domestic and wild Lycopersicon species, galactolipase (EC 3.1.1.26) activity, free fatty acid (FFA) level and Hill reaction activity were measured in chloroplasts isolated from control and cold treated leaves of L. esculentum Mill., cv. Norton, L. hirsutum Humb. and Bonpl., L. peruvianum var. glandulosum Mill. Galactolipase activity was higher in chloroplasts from Lycopersicon species with high chilling sensitivity than in chloroplasts of more chilling-resistant ones. A similar relationship was observed for FFA level in chloroplasts from both cold-stored and control leaves. Decrease in Hill reaction activity due to cold stress was greater in chloroplasts of more chilling-sensitive species. The changes are accompanied by a decline of photochemical activity. Considering the changes in the three parameters noted above, an increasing order of chilling tolerance was established: L. esculentum < L. hirsutum (700 m) < L. hirsutum (3100 m) < L. peruvianum (3400 m). It is suggested that measurements of galactolipase activity and FFA may be useful in an evaluation of differences in resistance to chilling injury of closely related species.     

Variation in Response of Wild Lycopersicon and Solanum spp. against Tomato Powdery Mildew (Oidium lycopersici)

A set of 154 accessions of nine wild Lycopersicon spp. and five accessions of three closely related Solanum spp. were tested for resistance to tomato powdery mildew ( Oidium lycopersici ). Screening revealed valuable sources of resistance, mainly among L. hirsutum, L. pennellii, L. cheesmanii, L. chilense, L. peruvianum and L. parviflorum. L. esculentum (all ssp.) and L. pimpinellifolium expressed high susceptibility to O. lycopersici inoculation. Results of variance and cluster analysis of responses to O. lycopersici coincide with recent taxonomic classification and genetic relationships within genus Lycopersicon .     

Relationships, origin, and diversity of Galapagos tomatoes: implications for the conservation of natural populations

Endemic Galápagos tomatoes (Lycopersicon cheesmanii) are of great value for cultivated tomato (L. esculentum) breeding, and therefore their conservation is of significance. Although within L. cheesmanii there is heterogeneity for many traits and formal infraspecific classification is not justified, here we distinguish three forms, without taxonomic significance, of L. cheesmanii that are of interest to breeders because of their distinctive morphology and habitat preferences: L. cheesmanii 'short' (one- to two-pinnate leaves, short internodes, and coastal habitats), L. cheesmanii 'long' (one- to two-pinnate leaves, long internodes, and inland habitats), and L. cheesmanii forma minor (three- to four-pinnate leaves, short internodes, and coastal habitats). In a recent survey of tomato populations in the Galápagos Islands, we found that several populations of L. cheesmanii reported 30-50 years earlier had disappeared, mostly as a consequence of human activity. In addition, a previously unreported invasive wild red-fruited form, which we named L. esculentum 'Gal cer,' was found on the island of Santa Cruz. The total diversity (estimated with amplified fragment length polymorphisms [AFLPs]) within L. cheesmanii (H(T) = 0.051) is almost as high as that for the mainland wild species L. pimpinellifolium (H(T) = 0.072). Lycopersicon esculentum 'Gal cer,' on the other hand, has a much lower diversity (H(T) = 0.014). Comparison of AFLP fragments shared by L. esculentum 'Gal cer' with other species showed that it is closely related to weedy tomato L. esculentum var. cerasiforme and, therefore, likely of recent origin. Genetic differentiation among the three native L. cheesmanii forms is low (G(ST) = 0.235), indicating that they share a common genetic background. Nonetheless, L. cheesmanii 'short' is about twice as diverse as L. cheesmanii 'long' or L. cheesmanii f. minor. UPGMA cluster and principal components analysis distinguish four groups within Eulycopersicon: L. pimpinellifolium, cultivated L. esculentum, L. esculentum var. cerasiforme including L. esculentum 'Gal cer,' and L. cheesmanii. The geographic distance and genetic distance in the wild forms of Galápagos tomatoes were not correlated. Apart from the pressure of humans, some native L. cheesmanii populations, especially L. cheesmanii 'long,' might be displaced by invasive L. esculentum 'Gal cer' because they share a similar habitat. We did not find evidence of intercrossing of L. cheesmanii with introduced L. esculentum, but occasional hybridization that contributes to loss of genetic integrity of L. cheesmanii cannot be ruled out. Establishment of reserves of L. cheesmanii to protect this species from introduced herbivorous animals and from hybridization with L. esculentum 'Gal cer' would help to conserve L. cheesmanii. Furthermore, accessions collected by C. M. Rick and others in the 1950s-1970s and now stored in germplasm banks could be used to reinstate some extinct populations.     

A set of simple PCR markers converted from sequence specific RFLP markers on tomato chromosomes 9 to 12

A set of 24 simple PCR markers was generated for tomato chromosomes 9, 10, 11 and 12. Polymorphism was sought for between Lycopersicon esculentum and one of six other Lycopersicon species (L. parviflorum, L. cheesmanii, L. hirsutum, L. pennellii, L. peruvianum, and L. chilense). PCR primers, which were designed from mapped RFLP sequences, were used to amplify genomic DNA of the different species and the PCR amplification products were screened for polymorphism by testing restriction enzymes. With this approach, 24 (71%) of the 34 selected RFLPs were converted into simple PCR markers. By using a reference population, the map positions of these markers relative to the original RFLP markers were verified. These markers are locus specific and can be efficiently used for alignment of linkage maps, mapping target genes and marker assisted selection.     

Effect of fertilization levels, age and canopy height of Lycopersicon hirsutum on the resistance to Myzus persicae

The effects of different levels of fertilization with nitrogen and potassium (NK), height within the canopy and plant age of Lycopersicon hirsutum f. glabratum (PI 134417) C. H. Mull and L. esculentum on the density of foliar cristalliferous idioblasts and trichomes, and on the levels of 2- tridecanone (2-TD) and 2-undecanone (2-UD) and their influence on the resistance to Myzus persicae (Sulzer) were investigated. NK levels were inversely related to densities of trichomes and cristalliferous idioblasts in L. esculentum, but only to densities of cristalliferous idioblasts in L. hirsutum. Trichome density increased with increasing height within the canopy and with plant age in L. esculentum and L. hirsutum. Density of cristalliferous idioblasts in L. hirsutum increased with canopy height reaching a maximum at mid-height within the canopy in opposition to L. esculentum in which a small decrease in density with height within the canopy was found. There was also a small decrease of idioblast density with increasing plant age for L. esculentum, while a greater and opposite variation took place in L. hirsutum. L. esculentum has mainly non- glandular leaf trichomes (90%), while L. hirsutum has mainly glandular trichomes (97%). The main glandular trichomes of L. hirsutum are producers of 2-TD and 2-UD, whose leaf content increases with increase of trichome density. A negative effect of 2-TD concentration and cristalliferous idioblast density on the survival and longevity of M. persicae was observed. Since L. hirsutum presented much higher levels of 2-TD and cristalliferous idioblast density than L. esculentum, L. hirsutum seems a promising source of resistance to M. persicae for tomato breeding programs.     

A Dispersed Family of Repetitive DNA Sequences Exhibits Characteristics of a Transposable Element in the Genus Lycopersicon

A segment of DNA 5' to the transcribed region of an auxin-regulated gene, ARPI, from Lycopersicon esculentum Mill. cv. VFN8 contains a sequence with the structural characteristics of a transposable element. The putative element (Lyt1) is 1340 bp long, has terminal inverted repeats of approximately 235 bp and is flanked by 9-bp direct repeats. Lyt1 has a structure similar to the Robertson's Mutator (Mu) family from maize. The terminal inverted repeats are 80% AT-rich, are 96.6% identical, and define a larger family of repetitive elements. Southern analysis and genomic dot-blot reconstructions detected at least 41 copies of Lyt1-hybridizing sequences in red-fruited Lycopersicon spp. (L. esculentum, L. pimpinellifolium and L. cheesmanii), and 2-8 copies in the green-fruited species (L. hirsutum, L. pennellii, L. peruvianum, L. chilense and L. chmielewskii). There were two to four copies in the Solanum spp. closely allied with the genus Lycopersicon (S. lycopersicoides, S. ochranthum and S. juglandifolium), while the more distantly related Solanum spp. showed little (one to two copies in S. tuberosum) to no (S. quitoense) detectable hybridization under stringent conditions. Linkage analysis in the F(2) progeny of a cross between L. esculentum and L. cheesmanii indicated that at least six loci that hybridize to the Lyt1 sequence are dispersed in the genome. Polymerase chain reaction and Southern analyses revealed that some red-fruited accessions and L. chmielewskii lacked Lyt1 5' to the transcribed region of ARPI. Subsequent sequence analysis indicated that only one copy of the 9-bp direct repeat (target site) was present, suggesting that transposition of the element into the ARPI gene occurred after the divergence of the red-fruited and green-fruited Lycopersicon species.     

Chemical and principal-component analyses of the essential oils of Apioideae taxa (Apiaceae) from central Balkan

The volatile constituents of the essential oils of 23 taxa belonging to the Apioideae subfamily were studied in detail. The investigated taxa were Pimpinella serbica (Vis.) Bentham & Hooker, Libanotis montana Cr., Cnidium silaifolium (Jacq.) Simk. ssp. orientale (Boiss.) Tutin, Bupleurum praealtum L., B. sibthorpianum S. S. var. diversifolium (Roch.) Hay, Aegopodium podagraria L., Torilis anthriscus (L.) Gmel., Orlaya grandiflora (L.) Hoffm., Laserpitium siler L., Laser trilobum (L.) Brokh., Chaerophyllum aureum L., C. hirsutum L., C. temulum L., Pastinaca sativa L., P. hirsuta Pancic., Tordylium maximum L., Physospermum cornubiense (L.) DC., Peucedanum alsaticum L., P. oreoselinum (L.) Moench, P. cervaria (L.) Cuss., P. austriacum (Jacq.) Koch, P. longifolium W. et K., and P. officinale L. All of these species grow wild in the central part of the Balkan Peninsula. The essential oils were found to be complex mixtures of various compounds, more than 100 constituents being in each taxon, with contributions of main products never exceeding 25% of the total content. Sesquiterpene hydrocarbons were found to be the main group of constituents of all taxa, except for Peucedanum species, where monoterpene hydrocarbons were identified as the main components. The chemotaxonomic value of the essential-oil composition is discussed according to results of principal-component analysis (PCA). The essential-oil composition mainly reflects current taxonomic relationships between the investigated taxa.     

Toxicity of methyl ketones from tomato trichomes to Tetranychus urticae Koch

The two methyl ketones, 2-tridecanone and 2-undecanone, are constituents of type VI glandular trichomes of the wild tomato, Lycopersicon hirsutum f. glabratum (PI 134417). They are known to cause mortality in several herbivorous insect species. In this study we investigated the effects of these chemicals on two strains of the two-spotted spider mite, Tetranychus urticae Koch, collected from tomato and cucumber crops in Dutch greenhouses. The two ketones were tested separately, in combination in the ratio found in L. hirsutum f. glabratum and in several other ratios to detect any synergistic interaction between them. We measured both the direct contact and residual toxicity, as well as the viability of the eggs produced by ketone-treated females. The methyl ketones can be considered as quite toxic to spider mites because their LC50 values were comparable to that found for the formulated acaricide amitraz. 2-Tridecanone was slightly more toxic than 2-undecanone but only in the tomato strain. Synergistic effects were not detected. In the bio assays for the residual effects of the two methyl ketones no significant mortality occurred but the mites avoided feeding on the treated surface and the eggs were laid almost exclusively on the untreated area. Furthermore, there was no significant egg viability for most of the treatments and when some egg viability occurred, it was not correlated to the dose of the chemical. Based on published data on the levels of methyl ketones in the trichome tips of wild and cultivated tomato (L. hirsutum f. glabratum and Lycopersicon esculentum Mill, respectively) it was calculated how many contacts are required to reach the LD50 (ng a.i. per mite). It appeared that on wild tomato just a few contacts suffice, but many are needed on cultivated tomato. Nevertheless, it seems justified to conclude that even in cultivated tomato, 2-tridecanone represents a naturally occurring acaricide. The cucumber and tomato strains of the two-spotted spider mite did not differ in their response to the methyl ketones. Because this is surprising, given the selection pressures on their respective host plants, we formulated two hypotheses, one explaining the absence of a difference due to cross-resistance and another one proposing that bioaccumulation obscures the underlying differences between the strains. This revised version was published online in November 2006 with corrections to the Cover Date.     

Effect of cropping season and possible compounds involved in the resistance of Lycopersicon hirsutum f. typicum to Tuta absoluta (Meyrick) (Lep., Gelechiidae)

The effect of cropping season on the resistance of the accession LA 1777 of Lycopersicon hirsutum f. typicum to the tomato leafminer Tuta absoluta (Meyrick) and the preliminary identification of the possible compounds involved in this trait were the objectives of this study. Two experiments were carried out with the accession LA1777 and the commercial cultivar Santa Clara of Lycopersicon esculentum . The first experiment was carried out during the summer season (January to April 1997) and the second was carried out during the autumn–winter season (May to September 1997). The accession LA 1777 was significantly more resistant to T. absoluta than the commercial cultivar, especially during the autumn–winter cropping season. Either antibiosis or antixenosis was probably responsible for the higher larvae mortality (54.2 and 91.2% for the summer and autumn–winter seasons, respectively) and longer larval development of T. absoluta (14.2 and 25.9 days for summer and autumn–winter seasons, respectively) on L. hirsutum f. typicum as compared with L. esculentum (11.0 and 19.6 days for summer and autumn–winter seasons, respectively). Insect fecundity parameters such as oviposition period, number of eggs/female, and rate of larvae eclosion were also negatively affected by L. hirsutum f. typicum . Fourteen main peaks were observed in total ion chromatograms of both tomato species. Eleven of these were present only in chromatograms of the accession, one was only present in chromatograms of the commercial cultivar, and two peaks were present in chromatograms from both when hexane extracts of leaves were subjected to a gas chromatographic–mass spectrometric analysis (GC–MS). However, the matches obtained between the peak spectra and the MS database were too low to justify even preliminary identification of the compounds.     

The inheritance of chilling tolerance in tomato (Lycopersicon spp.)

During the past 25 years, chilling tolerance of the cultivated (chilling-sensitive) tomato Lycopersicon esculentum and its wild, chilling-tolerant relatives L. peruvianum and L. hirsutum (and, less intensively studied, L. chilense) has been the object of several investigations. The final aim of these studies can be seen in the increase in chilling tolerance of the cultivated genotypes. In this review, we will focus on low-temperature effects on photosynthesis and the inheritance of these traits to the offspring of various breeding attempts. While crossing L. peruvianum (male symbol) to L. esculentum (female symbol) so far has brought the most detailed insight with respect to physiological questions, for practical purposes, e.g., the readily cross ability, crossing programmes with L. hirsutum as pollen donor at present seem to be a promising way to achieve higher chilling-tolerant genotypes of the cultivated tomato. This perspective is due to the progress that has been made with respect to the genetic basis of chilling tolerance of Lycopersicon spp. over the past five years.     

Phylogeny and systematics of the genus Lophozia s. str. (Dumort.) Dumort. (Hepaticae) and related taxa from nuclear ITS1-2 and chloroplast trnL-F sequences

Nuclear ITS1-2 and chloroplast trnL-F were sequenced for 21 taxa of Lophozia s. str., two species of Protolophozia, five species of Schistochilopsis, three species of Barbilophozia and Obtusifolium obtusum. The topologies of phylogenetic trees for 49 taxa constructed from combined sequences of these regions by maximum parsimony, maximum likelihood and Bayesian methods are similar. The species of Lophozia s. str., excluding Lophozia sudetica, combine into two main clades and these contradict subdivisions of Lophozia s. str. based on morphology. The species status of Lophozia lantratoviae is confirmed, whereas Lophozia austro-sibirica is almost identical to Lophozia ventricosa var. guttulata. The genus Schistochilopsis is paraphyletic and occupies basal position to Lophozia s. str., while O. obtusum is clearly separated from Schistochilopsis. A low level of divergence was found between L. sudetica and Protolophozia debiliformes, which are closer to Barbilophozia than to Lophozia s. str. Molecular divergence between geographically remote populations of L. sudetica, Lophozia silvicoloides and Protolophozia debiliformis are low as opposed to those of Lophozia polaris, Lophozia pellucida or Lophozia excisa. Consideration of the trnL intron P8 region indels alone can adequately assign some clades revealed by tree building. A consensus secondary structure of the trnL intron P8 region could not be inferred for taxa studied mainly due to high sequence length diversity originated from deletions.     

Species and recombination effects on DNA variability in the tomato genus

Population genetics theory predicts that strong selection for rare, beneficial mutations or against frequent, deleterious mutations reduces polymorphism at linked neutral (or weakly selected) sites. The reduction of genetic variation is expected to be more severe when recombination rates are lower. In outbreeding species, low recombination rates are usually confined to certain chromosomal regions, such as centromeres and telomeres. In contrast, in predominantly selfing species, the rarity of double heterozygotes leads to a reduced effective recombination rate in the whole genome. We investigated the effects of restricted recombination on DNA polymorphism in these two cases, analyzing five Lycopersicon species with contrasting mating systems: L. chilense, L. hirsutum, L. peruvianum, L. chmielewskii, and L. pimpinellifolium, of which only the first three species have self-incompatibility alleles. In each species, we determined DNA sequence variation of five single-copy genes located in chromosomal regions with either high or low recombination rate. We found that the mating system has a highly significant effect on the level of polymorphism, whereas recombination has only a weak influence. The effect of recombination on levels of polymorphism in Lycopersicon is much weaker than in other well-studied species, including Drosophila. To explain these observations, we discuss a number of hypotheses, invoking selection, recombination, and demographic factors associated with the mating system. We also provide evidence that L. peruvianum, showing a level of polymorphism (almost 3%) that is comparable to the level of divergence in the whole genus, is the ancestral species from which the other species of the genus Lycopersicon have originated relatively recently.     

Susceptibility of Tuta absoluta (Meyrick) (Lep., Gelechiidae) reared on two species of Lycopersicon to Bacillus thuringiensis var. kurstaki

The mortality of Tuta absoluta (Meyrick) larvae that were fed on leaves of Lycopersicon hirsutum f. glabratum (PI 134417, insect resistant) and Lycopersicon esculentum (cultivar Santa Clara, susceptible) treated with Bacillus thuringiensis var. kurstaki (Btk) was evaluated. Feeding on untreated PI 134417 was detrimental to the survival of T. absoluta larvae. When Btk was applied to the two Lycopersicon plants, mortality occurred in all T. absoluta instars. Application of Btk on tomato leaves had synergistic or additive effects with the resistant genotype on larval survival. This effect was dependent on the instar at which the larvae were fed Btk-treated leaves. Delayed Btk application may cause higher insect mortality if the insects become more susceptible to the pathogen after a longer period of feeding on the resistant crop.     

Variation for resistance to aphids (Homoptera: Aphididae) among tomato inbred backcross lines derived from wild Lycopersicon species

Two tomato inbred backcross line (IBL) populations, derived from crosses between aphid-susceptible Lycopersicon esculentum Mill. 'Peto 95-43' X resistant wild L. pennellii Corr (D'arcy) accession LA716, and Peto 95-43 X resistant wild L. hirsutum f. glabratum Mull accession LA407, were evaluated in replicated field experiments for resistance to potato aphid, Macrosiphum euphorbiae (Thomas), and green peach aphid, Myzus persicae (Sulzer). Aphid infestation scores for each IBL and control (LA716, LA407, Peto 95-43, and susceptible 'Alta') plot were recorded weekly for 5 and 9 wk during the summers of 2000 and 2001, respectively. Aphid infestation scores from leaflets were used to calculate area under the infestation pressure curve (AUIPC), a measure of aphid infestation throughout the growing season, for each IBL and control. Score AUIPC was highly correlated with actual aphid count AUIPC, indicating that scores accurately reflected aphid infestation. Score AUIPC was also highly correlated across both years (2000 and 2001) and locations. Low score AUIPC was significantly correlated with larger plant size and sprawling, indeterminate plant growth habit. Seven IBLs, LA716, and LA407 were significantly more resistant to aphids (lower score AUIPC) than susceptible parent Peto 95-43 in both years. Two IBLs, 1034 and 1051, were not significantly different from resistant LA407 for score AUIPC in both years. The seven aphid-resistant IBLs identified here can be useful as donor parent material for resistance breeding efforts in cultivated tomato.