Monogenic resistance in tomato to Fusarium oxysporum f. sp. lycopersici race 3

Summary Resistance to fusarium wilt, incited by Fusarium oxysporum (Schlecht.) f. sp. lycopersici (Sacc.) Snyder & Hansen race 3 in tomato (Lycopersicon esculentum Mill.) was discovered in LA 716, a L. pennellii accession. A resistant BC₁F₃ breeding line, E427, was developed from LA 716. E427 was crossed with the susceptible cv. Suncoast and F₁, BCP₁, BCP₂ (to Fla 7155, a susceptible parent) F₂, F₃, and BCP₂S₁ seeds were obtained. Segregation for resistance following root dip inoculation over three experiments indicated a single dominant gene controlled resistance. Five of the 12 BCP₁S₁'s segregated more susceptible plants, whereas one of the 12 segregated more resistant plants than expected (P<0.05). Three of 23 F₃ lines segregated more susceptible plants than expected while 1 of the 23 had more resistant plants than expected (P<0.05). Segregation in all other lines fit expected ratios. Five of the 23 F₃'s were homozygous resistant which was an acceptable fit to expectations (P=0.1–0.5). The gene symbol I ₃ is proposed for resistance to race 3 of the wilt pathogen. Deviations from expected ratios in data reported here and for other breeding lines indicate an effect of modifier genes and/or incomplete penetrance. Plant age at inoculation and seed dormancy did not affect results.Florida Agricultural Experiment Station Journal Series No. 8101.     

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.     

Inheritance of black leaf mold resistance in tomato

Summary Inheritance of black leaf mold (BLM) (caused by Pseudocercospora fuligena) resistance was studied in four crosses involving two resistant Lycopersicon accessions (PI134417, L. hirsutum and PI254655, L. esculentum) and four susceptible Asian Vegetable Research and Development Center tomato lines (CLN657BC₁F₂-267-0-3-12-7, CL143-0-10-3-0-1-10, CLN698BC₁F₂-358-4-13 and CL5915-93D₄-1-0-3). For each cross, six generations, i.e. P₁, P₂, F₁, F₂, BC₁F₁ and BC₁F₂ were evaluated following inoculations with isolate Pf-2 of P. fuligena. Chi-square analyses of the data based on the ratio of resistant to susceptible plants in the F₂ in three of four crosses gave a good fit to a segregation ratio of 1 R : 15 S, and BC₁F₂ data in three of four crosses gave an acceptable fit to the segregation ratio of 1 R : 63 S. The results indicate that resistance to BLM may be conditioned by two recessive genes acting epistatically in both PI134417 and PI254655.     

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.     

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.     

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.     

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).     

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.     

Inheritance of resistance to fruit rot (Phytophthora parasiticaDast.) in tomato (Lycopersicon esculentum Mill.)

Summary Fruit rot disease caused by Phytophthora parasiticaDast. is a limiting factor in tomato production in Himachal Pradesh. 30 to 60 per cent fruits are damaged by this disease. Crosses were made between EC 54725 (Lycopersicon pimpinellifolium), a small tyuited type, resistant to fruit rot and four highly susceptible tomato commercial cultivars (Gola, Sioux, S12, and Lalmani). Studies of F₁'s, F₂'s and back crosses indicated that EC 54725 carries a dominant gene imparting resistance to fruit rot.     

Introgression of resistance to Columbia and Northern root-knot nematodes fromSolanum bulbocastanum into cultivated potato

Summary Resistance toMeliodogyne chitwoodi races 1 (MC1) and 2 (MC2) andM. hapla (MH) derived fromSolanum bulbocastanum was introduced into the cultivated potato gene pool through somatic fusion. The initial F₁ hybrids showed resistance to the three nematodes. Resistance to reproduction on roots by MC1 was accompanied by resistance to tuber damage in F₁ clones. Tuber damage sometimes occurred, however, in hybrids of BC₁ progeny resistant to reproduction on roots when MC2 and MH were the challenging nematodes. Resistance to reproduction was transferred into BC₁ individuals, but a greater proportion of BC₁ progeny was resistant to MC1 than to MC2 or MH. Resistance to MC1 appears to be dominant and discretely inherited. F₁ and BC₁ progeny were pollen sterile, but seed were produced from crosses using cultivated tetraploid pollen sources. Approximately 11 and 33 per cent of pollinations produced berries on F₁ and BC₁ pistillate parents, respectively. Seed yield increased fourfold overall in crosses with F₁ compared to BC₁ individuals.Abbreviations MC1 Meloidogyne chitwoodi race 1 - MC2 Meloidogyne chitwoodi race 2 - MH Meloidogyne hapla - Rf Reproductive factor     

Inheritance of tomato leaf curl virus resistance in Lycopersicon hirsutum f. glabratum

Summary Inheritance of resistance to tomato leaf curl virus (TLCV) was studied in the progenies derived from interspecific crosses between TLCV resistant Lycopersicon hirsutum f. glabratum line B 6013 and five susceptible cultivars (HS 101, HS 102, HS 110, Pusa Ruby and Punjab Chhuhara) of L. esculentum. P₁, P₂, F₁, F₂, B₁ and B₂ progenies of the five crosses were artificially inoculated with local strains of TLCV by means of the vector whitefly, Bemisia tabaci (Genn.). and the disease reaction was studied in all the crosses. Reaction of parents, F₁, F₂ and backcrosses suggests that resistance derived from L. hirsutum f. glabratum B 6013 is based on two epistatic genes, one from the wild parent and one from the cultivated one, resulting in a 13:3 segragation in the F₂.     

Variation for aphid resistance and insecticidal acyl sugar expression among and within Lycopersicon pennellii-derived inbred backcross lines of tomato and their F1 progeny

Five inbred backcross lines (IBL) were selected for higher relative expression of insecticidal acyl sugars (rank average) from an inbred backcross population derived from the cross Lycopersicon esculentum cultivar ‘Peto 84’×Lycopersicon pennellii accession LA716. These five BC₂S₅ IBLs were crossed in a partial diallel design (Method II), and their self and F₁ progeny and three control cultivars were tested at two California field locations in 1996. Counts of potato aphids, Macrosiphum euphorbiae Thomas, on leaves, as a measure of plant infestation, revealed significant general combining ability (GCA) for lower aphid numbers with IBL44 and IBL59; the F₁ hybrid IBL44 × IBL59 had significantly fewer aphids per leaflet than the susceptible cultivar ‘Alta’. GCA for acyl sugars was associated with IBL59 only. Of all the IBL and IBL × IBL F₁ hybrids, only IBL59 produced significant levels of acyl sugars. Significant within IBL59 variation for acyl sugars was observed, but not for aphid resistance. Our results suggest that factors other than acyl sugars contributed to L. pennellii-derived aphid resistance in IBL × IBL F₁ hybrids and IBL that do not produce significant amounts of acyl sugars. IBL59 and IBL44 may be useful for breeding for aphid resistance in cultivated tomato.     

Frequency distributions and linkage relationships of 2-tridecadone in interspecific segregating generations of tomato

Summary The levels of the naturally occurring insecticide 2-tridecadone (2TD) were measured in leaves of Lycopersicon hirsutum f. glabratum, L. esculentum, the interspecific F₁ hybrid, the F₂ and backeross generations. The wild species contains 50 times more 2TD than the cultivated tomato and the frequency distribution of the substance indicates that there is dominance for low levels of the compound. The genotype of F₂ plants was determined with respect to 6 codominant isozyme markers and 4 dominant/recessive markers. Significant associations were detected between 5 of the marker genes and the level of 2TD. We interpret these results as implying linkage between marker genes and genes that control the level of 2TD. The behavior of the gene for the determinant growth habit of the plants suggests that it has a pleiotropic effect on the level of 2TD.     

Generation means analysis of right-censored response-time traits: Low temperature seed germination in tomato

Summary In studies to determine the inheritance of response-time traits, such as time to seed germination, some viable individuals may fail to respond during an experiment. If these right-censored observations are ignored, sample means and variances will be underestimated. This is illustrated using data from time to seed germination at 9°C for Lycopersicon esculentum (Mill.) fast germinating PI 120256, slow-germinating T3 and their reciprocal F₁, F₂ and backcross progeny. This paper presents methods to detect and to accommodate right-censored data in generation means analysis. Genetic interpretations derived from corrected and uncorrected estimates of generation means and variances are compared. Correction for right-censoring increased estimates of environmental and phenotypic variances, and decreased heritability estimates.     

A random amplified polymorphic DNA (RAPD) molecular marker for the Tm-2 a gene in tomato

Summary A RAPD marker, linked to the Tm-2 ^agene engendering TMV resistance in tomatoes, was identified. The validity of the RAPD marker was corroborated by screening several tomato varieties, and correctly identifying those which carried Tm-2 ^a, as well as by F₂ segregation analysis. All tested resistant varieties descending from a common Lycopersicon peruvianum/esculentum ancestor, LA1791, exhibited this marker.     

Genetic relationships among cold, salt and drought tolerance during seed germination in an interspecific cross of tomato

Seed of BC₁ progeny of an interspecific cross between a slow germinating Lycopersicon esculentum breeding line(NC84173; maternal and recurrent parent) and a fast germinating L.pimpinellifolium accession (LA722) were evaluated for germination under cold stress, salt stress and drought stress, and in each treatment the most rapidly germinating seeds (first 2%) were selected. Selected individuals were grown to maturity and self-pollinated to produce BC₁S₁ progeny families. The selected BC₁S₁ progeny from each experiment were evaluated for germination rate in each of a non stress (control),cold-, salt- and drought-stress treatment, and their performances were compared with those of a non selected BC₁S₁population in the same treatments. Results indicated that selection for rapid seed germination in each of the three stress treatments was effective and significantly improved progeny germination rate under all three stress conditions. The results support the suggestion that same genes might control the rate of seed germination under cold, salt and drought stress. Furthermore, selection in each of the three stress treatments resulted in improved progeny seed germination rate under nonstress conditions, suggesting that genetic mechanisms that facilitate rapid seed germination under stress conditions might also contribute to rapid germination under nonstress conditions. In practice, therefore, selection for rapid seed germination under a single stress environment may result in progeny with improved seed germination under a wide range of environmental conditions. Furthermore, to improve germination rate under nonstress conditions, it might be more efficient to make selections under stress conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Resistance to potato leafroll virus derived from Solanum chacoense: characterization and inheritance

Summary Resistance to potato leafroll virus (PLRV) was detected in an accession of Solanum chacoense. Inoculations with viruliferous aphids and subsequent graft challenges using Datura tatula and potato as PLRV sources determined that resistance appears to be of an extreme type. Virus was not detectable using enzyme-linked immunosorbent assay (ELISA) in S. chacoense, and in resistant F₁ and BC₁ progenies after attempts to transmit the virus through grafting. The segregation ratios of BC₁ progenies for positive and negative ELISA tests are consistent with simple dominant inheritance.     

Pollen selection in breeding tomato (lycopersicon esculentum Mill.) for adaptation to low temperature

Summary F₂ and BC₁ progenies from crosses between tomato (Lycopersicon esculentum) varieties differing for growth capacity at low temperature were produced under controlled conditions by hand pollination under two temperature regimes (22°C D/15°C N, normal temperature (N), and 15°C D/8°C N, low temperature (L)) with pollen formed under both regimes, resulting in four pollination treatments: NN, NL, LN and LL. Vegetative growth of the offspring was compared under a rather low temperature regime (19°C D/10°C N). Populations from different treatments within the progeny of a particular F₁ often differed significantly for average dry weight of 7 weeks old plants, the average of the NN population always being higher. Variances for dry weight were sometimes larger for LN populations, but this never resulted from a larger number of vigorous plants. Differences between populations within each progeny seemed to result in part from differences in the conditions for embryo development. Pollen selection at low temperature did not appear to be efficient for sporophyte breeding in this experiment.     

Genetics of the viability of pollen grain produced at low temperatures inLycopersicon Mill.

Summary The different viabilities of pollen produced at low temperatures in intra- and inter-specific crosses of tomato were studied. Cultivars Red Top, Moneymaker, and Marroqui were crossed with cultivar E-15 and these four cultivars were hybridized with lines of the wild speciesL. pimpinellifolium PE-13,L. parviflorum PE-52,L. pennellii PE-47, andL. hirsutum PE-37 and PE-41. A six-generation family of the Moneymaker x PE-47 cross was obtained to carry out a more detailed genetical study of pollen grain viability at low temperatures. Pollen grain viability was evaluated during the winter via acetocarmine staining. When the parents were compared with their F₁, the intra-specific tomato crosses showed dominance to better-quality pollen, theL. esculentum x L. pimpinellifolium inter-specific crosses showed positive heterosis, while the crosses ofL. esculentum with the tolerant speciesL. pennellii andL. hirsutum showed intermediate inheritance. However, in theL. esculentum x L. pennellii family, the dominance and the non-allelic interactions (homozygosis x homozygosis) were the most important factors, so that dominance to better viability at low temperatures appeared to be the general mode of inheritance. Genetical control of pollen grain viability at low temperatures seemed to be polygenetic.