Genetics of resistance to early blight (Alternaria solani Sorauer) in tomato (Lycopersicon esculentum L.)

The genetic nature of early blight resistance in tomato was studied in three crosses at seedling and adult plant stages. A six generation mean analysis of the cross Arka Saurabh (susceptible) × IHR1939 (resistance) and its reciprocal cross revealed that the resistance to early blight was conferred by recessive polygenes at both seedling and adult plant stages. This polygenic early blight resistance revealed the importance of additive and additive × additive gene effects at seedling stage and higher magnitude of dominance and dominance× dominance gene effects at adult plant stage. Evaluation of parents, F₁, F₂ and backcross generations of IHR1816 (resistance) × IHR1939 (resistance) revealed that the early blight resistance genes in IHR1816 (Lycopersicon esculentum NCEBR-1) and IHR1939 (Lycopersicon pimpinellifolium L4394) are independent. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance in two Lycopersicon species to an Arkansas isolate of tomato spotted wilt virus

Summary Tomato spotted wilt virus (TSWV) was obtained from infected tomatoes in commercial fields in Arkansas in 1985. A greenhouse screening procedure for identifying tomatoes resistant to TSWV was established using an enzyme-linked immunosorbent assay (ELISA) to detect infected plants. Symptom expression was variable and symptom expression was not reliable for identifying infected plants. Germplasm evaluated for resistance to one typical Arkansas isolate (85–9) of TSWV included: twenty cultivars and breeding lines of Lycopersicon esculentum Mill, 52 accessions of L. pimpinellifolium (Jusl.) Mill and 8 accessions of L. peruvianum (L.) Mill. All cultivated accessions and breeding lines evaluated were susceptible. Some individual plants in several accessions of L. pimpinellifolium were resistant and nearly all plants of the L. peruvianum accessions that were evaluated were resistant to isolate 85–9.Dept. of Plant Pathology     

Zingiberene-mediated resistance to the South American tomato pinworm derived from Lycopersicon hirsutum var. hirsutum

The Lycopersicon hirsutum var. hirsutum accession PI 127826 is recognized as a good source of resistance to arthropod pests due to the action of the allelochemical zimgiberene, a sesquiterpene present in its glandular trichomes. Five genotypes were selected from the F₂ generation of the interspecific cross Lycopersicon esculentum ‘TOM-556’ × Lycopersicon hirsutum var. hirsutum ‘PI 127826’, based on their low levels (BPX-368-clone#56) or high levels(BPX-368-clone#92, BPX-368-clone#105,BPX-368-clone#179, BPX-368-clone#250) of zingiberene. The five F₂ genotypes were tested for resistance to the South American tomato pinworm Tuta absolutaalong with accession L. esculentum ‘TOM-556’ (pinworm susceptible), and the accessions L. hirsutum var. hirsutum ‘PI 127826’ and L. pennellii ‘LA716’ (resistant). The F₂ clones selected for high foliar zingiberene levels showed lower scores for leaflet lesion type(LLT), percent leaflets attacked (PLA) and overall plant damage (OPD) than the low zingiberene genotypes. The results indicated that zingiberene mediates resistance to the South American pinworm, based on feeding and on ovipositing deterrence, in populations derived from the interspecific cross between Lycopersicon. esculentum and Lycopersicon hirsutum var. hirsutum. Indirect selection for high foliar zingiberene content is suggested as an efficient technique for breeding tomatoes for resistance to the South American tomato pinworm. This revised version was published online in July 2006 with corrections to the Cover Date.     

A novel source of resistance to the two-spotted spider mite in Lycopersicon pimpinellifolium (Jusl.) Mill.: its genetics as affected by interplot interference

We have previously found an accession of Lycopersicon pimpinellifolium (Jusl.) Mill. (`TO-937') that appeared to resist attack by the two-spotted spider mite (Tetranychus urticae Koch). L. pimpinellifolium is a very close relative of the cultivated tomato (Lycopersicon esculentum Mill.) and thereby a potential source of desirable traits that could be introgressed to the crop species. The objective of this study was to investigate the genetics of the resistance present in `TO-937'. Resistance to infestation by the spider mite was quantified in 24-plant plots of L. pimpinellifolium accessions `TO-937' and `PE-10', L. pennellii accession `PE-45', L. esculentum cultivars `Moneymaker', `Roma' and `Kalohi' (reported to be partially resistant: Stoner & Stringfellow, 1967), and the interspecific F₁ cross, L. esculentum `Moneymaker' × L. pimpinellifolium `TO-937'. Only `TO-937', the F<sub>1</sub>, and`PE-45' were found to be resistant. Resistance of `TO-937' was complete when evaluated in two small greenhouses completely planted with `TO-937' so as to simulate the genotypic homogeneity usual in commercial crops. Generations (P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂) of a P₁ (susceptible) × P₂ (resistant) cross (`Moneymaker' × `TO-937') were studied for resistance in a single-plant per plot design. Resistance of `TO-937' was inherited with complete dominance and appeared to be controlled by either two or four genes according to whether segregation in the F₂ or the BC₁P₁, respectively, were considered. However, calculation of the number of genes involved in the resistance was complicated by negative interplot interference due to the high frequency of resistant genotypes within most of the generations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Trichomes on leaves of Lycopersicon hirsutum,L. esculentum and their hybrids

Summary Density of certain types of trichomes were characterized on leaflets of Lycopersicon hirsutum, L. esculentum and their hybrids. Trichome density covaried with leaflet development, and depending on the species and type of trichome, density also covaried with leaflet surface. Type IV trichomes were not observed on leaflets of L. esculentum and Type V trichomes were not observed on leaflets of L. hirsutum. Type IV and V trichomes were present on F₁ hybrids and densities of these two types of trichomes segregated in the F₂ population. When corrected for differences of leaflet size between L. hirsutum and L. esculentum, densities of Type VI trichomes were more similar between species than uncorrected densities. There was little difference in Type VI density among F₂ individuals. The appearance, lipid and phenol staining properties, and native fluorescence of the multicellular tip of the Type VI trichome differed between L. hirsutum and L. esculentum. F₁ hybrids were intermediate for these characters and these characters segregated in the F₂ generation.The investigation reported in the paper (84-10-94) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director. This material is based upon work supported, in part, by the U.S. Department of Agriculture, Agreement No. 59-2213-1-1-717-0 (Competitive Research Grants Program).     

Introgression of in vitro regeneration capability of Lycopersicon pimpinellifolium Mill. into recalcitrant tomato cultivars

The goal of this research was to study the introgression of the high regeneration capacity of Lycopersicon pimpinellifolium Mill line WV-700, in recalcitrant tomato cultivars (L. esculentum Mill cvs. Petomech, Santa Rita and VFN-8) using backcrossing. Hypocotyl explants of in vitro germinated seeds were cultivated in half strength MS medium supplemented with 5mg/l 6-BA to assess their shoot regeneration capacity. The apical shoot of the in vitro germinated seeds were cultured on a separate medium. Apical shoots from genotypes showing high regeneration rates were acclimated in a glasshouse and used as pollen donors for the next backcrossing. After four backcrossings, the material showed a similar mean fruit weight for the cultivated tomato and a high regeneration capacity similar to the wild species. It is shown that L. pimpinellifolium can be used with success as donor parent to introgress in vitro regeneration capacity to recalcitrant tomato cultivars. This revised version was published online in August 2006 with corrections to the Cover Date.     

Breeding tomato for pollen tolerance to low temperatures by gametophytic selection

Haploid selection for traits related to pollen cold tolerance in tomato was performed in segregating populations derived from a Lycopersicon esculentum × L. pennellii hybrid. BC₁ populations were obtained by combining normal and low temperature treatments on two stages of pollen development: pollen formation, and germination and pollen tube growth. F₁ hybrids were cultivated under low and normal temperatures and their pollen was used to pollinate L. esculentum plants at low and normal temperatures. The four BC₁ populations obtained were tested for the quality and quantity of pollen produced at low temperatures. The population obtained by cold treatment at both stages had a significantly improved pollen germination ability at low temperatures. The two other coldselected BC₁ populations showed no differences compared with the unselected BC population. A second cycle of pollen selection, corresponding to BC₂, was applied in order to test its persistence in the subsequent generations and the possibility to further improve the character. This second cycle showed no improvement although some plants retained the high pollen germination ability at low temperatures that was observed in the first cycle. Hence, gametophytic selection of some characters related with tomato pollen performance may be feasible, at least for the first cycle of selection.     

Tomato Spotted Wilt Virus (TSWV) resistance in tomato derived from Lycopersicon chilense Dun. LA 1938

Tomato spotted wilt virus (TSWV) resistance wasidentified in Y118 (Fla 925-2), an F₁BC₁S₆ tomato line(Lycopersicon esculentum Mill.), derived from a crosswith L. chilense Dun. (LA 1938). This line waspreviously selected for tomato mottle virus (ToMoV)resistance in Florida. Progeny from crosses betweenFla 925-2 and three different TSWV susceptible L.esculentum parents were used in TSWV resistancestudies. A total of 75 F₁ and 596 F₂ plants from allthree crosses were screened for TSWV resistance. ForF₂ plants free of TSWV symptoms, evaluations were madeusing enzyme-linked immunosorbent assay (ELISA). TenF₃ populations used for further greenhouse and fieldscreenings were selected from F₂ plants found to befree of the virus using visual and ELISA criteria ateach evaluation. One F₁ and four F₃ lines werestudied under field conditions (Stellenbosch, SouthAfrica) in which 100% of the `Flora-Dade' susceptiblecontrols were severely infected with TSWV. Theresults of the field study clearly establish that TSWVfield resistance is present in the Fla 925-2 (Y118)derived lines. Studies conducted on these linesrevealed that this resistance has the distinctcharacteristic of often `recovering' from initiallyhigh levels of virus titer in the tissue to levelsbelow detection with ELISA.     

Interactions of BA, GA3, NAA, and surfactant on interspecific hybridization of Lycopersicon esculentum × L. chilense

Time consuming and expensive tissue culture type techniques are currently used for achieving Lycopersicon esculentum × L. chilense interspecific hybrids. The objective of this study was to determine if the number of viable hybrid seed produced directly from this wide cross could be improved by optimizing female/male parent selection when treated with various hormones in the presence or absence of a surfactant (Triton X-100). Individual or combination treatments of BA(6-benzylaminopurine), GA₃ (gibberellic acid), and/or NAA(α-naphthaleneacetic acid) were applied (both with and without surfactant)to the ovaries of two L. esculentum female parents (Fla 7613 and89S) one day following pollination by nineL. chilense accessions. Hormone applications over 1-, 3-, and 5-dayperiods were made and all treatments were compared to a distilled water control. Across all experiments 21 viable hybrids from 1920 fruits (1.1%) were obtained from2128 pollinations. There were five consistent trends observed across three experiments. First, treatments utilizing NAA produced more hybrids than any other treatment hormone. Second, eight of the total 21 hybrids resulted from five consecutive days of hormone treatments (P-value ≤0.01), and six of those eight were realized when NAA was involved. Third, the use of Triton X-100 in combination with the hormone treatments significantly reduced the number of fruits/pollination (P-value ≤ 0.01), but did not significantly reduce the number of overall viable hybrids (P-value > 0.4). Fourth, the L. chilense male parent LA2759 significantly produced the most interspecific hybrids (P-value ≤ 0.04),seven hybrids from 295 fruits, compared to the male parent LA130 which did not produce any hybrids (from 293 fruits) -other accessions were intermediate. Fifth, nearly ten times more hybrids were produced from crosses with Fla 7613 than with 89S. These experiments provide a foundation for future studies to discern specific concentrations, timing and frequency of application of NAA and possibly, GA₃, to efficiently produce interspecific hybrids of L. esculentum × L. chilense. This revised version was published online in August 2006 with corrections to the Cover Date.     

Efficient hybridization between Lycopersicon esculentum and L. peruvianum via 'embryo rescue' and in vitro propagation

For the transfer of valuable traits from wild species into the cultivated tomato, excised globular-stage embryos 13 and 15 days after pollination (DAP) were cultured in vitro. Plants were regenerated from interspecific crosses of Lycopersicon esculentum cv. ‘Early pink’×L. peruvianum PI270435, and backcrosses of L. esculentum‘Giban (JF) No. 1’× (‘Early pink’× PI270435). Somatic embryos and single cotyledons emerged on hypocotyl sections of the embryos. Five to nine plantlets per embryo were obtained by clonal propagation. The hybrid nature of the plants is confirmed by comparing hybrids and parents in their ability to regenerate shoots from leaf segments in vitro, by comparing plant morphology and characteristics and by chromosome number. This study describes an efficient ‘embryo rescue’ method, as well as somatic embryogenesis by clonal propagation. A novel and simple method for the characterization of the interspecific hybrids is also reported.     

Low temperature seed germination of Lycopersicon species evaluated by survival analysis

Summary Low temperature germination responses were evaluated for 18 high altitude accessions representing five wild Lycopersicon species and 19 accessions of L. esculentum which have reputed ability to germinate in the cold. Survival analysis indicated that one accession of L. chilense germinates better at 10°C than PI 120256, the fastest-germinating L. esculentum genotype, and that PI 120256 germinates as well as PI 126435 (L. peruvianum). Additional wild ecotypes exhibiting rapid germination at 10°C were identified from L. peruvianum and L. hirsutum. These ecotypes may possess genetic potential for introgressing cold germination ability into L. esculentum cultivars.     

Low-temperature-related growth and photosynthetic performance of alloplasmic tomato (Lycopersicon esculentum Mill.) with chloroplasts from L. hirsutum Humb. & Bonpl.

Growth and photosynthetic performance were analyzed in alloplasmic tomato at a high- (25/17 ^°C; HTR) and low-temperature regime (12/6 ^°C; LTR) in order to establish the role of cytoplasmic variation on low-temperature tolerance of tomato (Lycopersicon esculentum Mill.). Four alloplasmic tomato lines, containing the nuclear genome of tomato and the plastome of L. hirsutum LA 1777 Humb. & Bonpl., an accession collected at high-altitude in Peru, were reciprocally crossed with 11 tomato entries with a high inbreeding level and a wide genetic variation, resulting in a set of 44 reciprocal crosses. Irrespective of growth temperature, alloplasmic families with alien chloroplasts of L. hirsutum (h) were on average characterized by a high shoot biomass, a large leaf area, and a low specific leaf area in comparison with their euplasmic counterparts. These results do not directly point to an advantageous effect of h-chloroplasts on biomass accumulation at low temperature but rather towards a small general beneficial effect on growth and/or distribution of assimilates. Significant chloroplast-related differences in photosynthetic performance, however, were not detected at both temperature regimes, indicating that h-chloroplasts can properly function in a variable nuclear background of L. esculentum. It is concluded that chloroplast substitution is not an effective method for breeding tomato plants with improved low-temperature tolerance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of resistance against Phytophthora infestans in Lycopersicon pimpenellifolium L3707

Summary Lycopersicon pimpenellifolium L3707, resistant to the late blight oomycete Phytophthora infestans was crossed with the susceptible Lycopersicon pimpenellifolium 14377 or the susceptible Lycopersicon esculentum ZH. Progeny F1 and F2 generations were scored at the 5-leaf stage for resistance against 175 field and recombinant isolates of the pathogen. F1 plants exhibited various levels of moderate resistance and F2 plants segregated 3:6:7 resistant/moderately resistant/susceptible. The data support the hypothesis that race-non-specific resistance in L3707 is controlled by two independent genes: a partially-dominant gene and a dominant epistatic gene.     

High salt tolerance potential in Lycopersicon species during germination

Summary The potential to improve seed germination responses to salinity was evaluated for 13 accessions representing six wild Lycopersicon species and 20 accessions of L. esculentum. Germination response times increased in all accessions at 100 mM NaCl. Analysis indicated that one accession of L. peruvianum (PI126435) germinated faster under high salinity than all other accessions and was closely followed by L. pennellii (LA716). The fastest germinating L. esculentum accession, PI174263, ranked third. Additional wild ecotypes exhibiting rapid germination at 100 mM NaCl were identified among L. pimpinellifolium and L. peruvianum.     

Inheritance of sesquiterpene carboxylic acid synthesis in crosses of Lycopersicon hirsutum with insect-susceptible tomatoes

Sesquiterpene carboxylic acids (SCA) from Lycopersicon hirsutum f. typicum Humb, & Bonpl, accession LA 1777 confer host‐plant resistance to Helicoverpa zea (Boddie) and Spodoptera exigua (Hübner), two Lepidopteran (Noctuidae) insect pests of the cultivated tomato. L. esculcntum Mill. Hybrids were made between LA 1777 and two accessions of insect‐susceptible tomatoes that do not produce SCA, L. hirsutum LA 1033 and the L. L. esculentum cultivar Chico III, F₂ and backcross progeny of both crosses were evaluated for variation in amounts of total SCA in replicated field and greenhouse plantings. Analysis of variance of total SCA in a set of F₂ plants of the cross LA 1033 × LA 1777 demonstrated significant variation in SCA attributable to genotype, environment and their interaction. Segregation for high levels of SCA in the populations suggests polygenic inheritance. Broad sense heritability was 0.84, which indicates that phenotypic selection for plants with high SCA is feasible in a backcross‐breeding programme for the introgression of SCA‐mediated insect resistance into tomato cultivars.     

Evaluation of seven Lycopersicon species for resistance to tomato spotted wilt virus (TSWV)

Summary One hundred eighty-eight accessions of Lycopersicon cheesmanii, L. chilense, L. chmielewskii, L. hirsutum, L. parviflorum, L. pennellii, and L. peruvianum were screened for resistance to three isolates of tomato spotted wilt virus (TSWV). All plants in an accession were initially screened for resistance to TSWV using isolate 85–9 from Arkansas. Visual symptoms were used to cull obviously infected plants, followed by enzyme-linked immunosorbent assay (ELISA) to identify uninfected plants. Cuttings were taken from uninfected plants in the first screening and the resulting plants were inoculated with isolates Glox and T-2 from Texas and Hawaii, respectively. No resistance was identified in L. cheesmanii, L. chmielewskii, L. hirsutum, L. parviflorum, and L. pennellii. However, 33 of 63 L. chilense accession produced 91 of 1268 plants that were uninfected with isolate 85–9 and 20 accessions that produced 40 of 257 plants that were not infected with any of the isolates. After screening with isolate 85–9 9 of 12 L. peruvianum accessions tested had 38 plants uninfected and 8 accessions had 25 plants that were not infected with any of the isolates.     

Somatic hybridization as a tool for tomato breeding

Protoplast fusion can be used to produce somatic hybrids of species that cannot be obtained by sexual hybridization. The possibility to introgress genes from Solanum species into the cultivated tomato species Lycopersicon esculentum, and to obtain novel cytoplasm-nucleus combinations (cybrids) was considered as an important strategy to extend the genetic variation available for tomato breeding. Somatic hybrids between L. esculentum and other Lycopersicon species, as well as between L. esculentum and Solanum or Nicotiana species, have been produced. Specific mutants, genotypes with antibiotic resistances, and metabolic inhibition by iodoacetate or iodoacetamide and irradiation were used for the selection of hybrids. In addition, the improvement of protoplast culture techniques and the use of the favourable tissue culture traits derived from species such as L. peruvianum, which have been introduced into tomato by classical breeding, allowed the efficient recovery of somatic hybrids. However, the occurrence of somatic incongruity in fusion combinations of L. esculentum and Solanum and even more in L. esculentum and Nicotiana, did not allow the production of true cybrids and/or fertile hybrids, indicating the importance of both cytoplasm-nucleus and nucleus-nucleus interactions in somatic incongruity. Another problem with fusions between distantly related species is the strongly reduced fertility of the hybrids and the very limited homoeologous recombination between chromosomes of the parental species. Partial genome transfer from donor to recipient through microprotoplast (+) protoplast fusion, and the production of monosomic or disomic chromosome addition lines, light overcome some of these problems. In symmetric somatic hybrids between L. esculentum and S. tuberosum the occurrence of limited somatic and meiotic recombination was demonstrated. Fertile progeny plants could be obtained, though at a low frequency, when embryo rescue was performed on a large scale after backcrossing hexaploid somatic tomato (+) potato hybrids with a tetraploid potato genotype. The potential value of genomic in situ hybridization (GISH) and RFLPs for the analysis of the genome/chromosome composition of the hybrids has been demonstrated for intergeneric somatic hybrids between Lycopersicon and Solanum.Abbreviations cpDNA chloroplast DNA - mtDNA mitochondrial DNA     

Inheritance of foliar zingiberene contents and their relationship to trichome densities and whitefly resistance in tomatoes

The sesquiterpene zingiberene, present in leaf glandular trichomes, is reportedly responsible for the high level of arthropod resistance found in Lycopersicon hirsutum var. hirsutum. This paper reports on the inheritance of zingiberene contents and of the various types of glandular trichomes in the interspecific cross L. esculentum × Lycopersicon hirsutum var. hirsutum. Plants of L. esculentum ‘TOM-556’ (= P₁), L. hirsutum var. hirsutum ‘PI-127826’(= P₂), F₁ (P₁ × P₂) and F₂ (P₁ ×P₂) were evaluated for zingiberene contents and densitities of and glandular (types I, IV, VI and VII) trichomes. Broad sense heritabilities were high for all traits studied (0.678, 0.831, 0.996, 0.799 and 0.717 respectively for zingiberene and trichome types I, IV, VI, VII). There were significant positive genetic correlations between zingiberene contents and densities of trichomes types IV, VI and VII. Inheritance of zingiberenecontents can be explained mostly by the action of a single major locus, inwhich the allele from L. hirsutum that conditions high content is incompletely recessive over the allele from L. esculentum. Action of an incompleteley recessive allele in one major locus appears to be evident for densities of trichome types IV, VI and VII, but there is also evidence of the action of other epistatic loci for types IV and VI. F₂ genotypes selected for high zingiberene levels showed higher levels of resistance to the silverleaf whitefly Bemisia argentifolii than L. esculentum ‘TOM-556’, levels that were comparable those found in L. hirsutum var. hirsutum ‘PI-127826’ and other whitefly resistant accessions. This revised version was published online in August 2006 with corrections to the Cover Date.     

Resistance in Lycopersicon hirsutum f. glabratum to the greenhouse whitefly (Trialeurodes vaporariorum) increases with plant age

Summary In this study genetic variation for resistance to the greenhouse whitefly (Trialeurodes vaporariorum) of four genotypes of tomato (L. esculentum) and two subspecies of L. hirsutum was investigated. Resistance was quantified by the whitefly life history components adult survival, oviposition rate, pre-adult survival and developmental period, measured on plants inoculated with whiteflies in clip-on cages.The largest differences between species were found when life history components were measured on adult plants of about four months old. On L. hirsutum f. glabratum whiteflies had the lowest adult survival, oviposition rate and pre-adult survival. On L. hirsutum these components were intermediate whereas on all L. esculentum genotypes they were highest. The variation between plants was low compared to the variation within plants. These results indicate that single plant tests can be used to determine accurately genetic variation between individual plants in a segregating population.     

Resistance in Lycopersicon species to black leaf mold caused by Pseudocercospora Fuligena

Over 540 accessions of wild Lycopersicon species or their crosses with L. esculentum were screened for resistance in a series of trials. Forty-six accessions were selected for the final screening trial based on lower disease ratings in previous trials. Of these, L. hirsutum had the greatest number of resistant accessions, followed by L. esculentum and L. peruvianum. Twenty accessions were quantified for their levels of resistance based on leaf area infected, area under disease progress curve (AUDPC), and the degree of sporulation. There was a significant positive correlation between the AUDPC calculated from 20 accessions evaluated under growth room and field conditions. Five L. hirsutum accessions had no sporulation associated with leaf lesions, whereas L. esculentum accessions had an average of 1.6×10⁴ conidia/cm² of leaf tissue. There was significant positive correlation between the AUDPC values and the number of conidia per cm² of leaf tissue.