Inheritance of growth characters of tomato (Lycopersicon esculentum Mill.) under low energy conditions

Summary To study the mode of inheritance of W, RGR, NAR, LAR, SLA and LWR, the growth of 15 tomato genotypes and 104 of their F1's was analyzed. The plants were grown at a 19°C day temperature and a 10°C night temperature under a light intensity of 24 Wm^-2 visible radiation and a daylength of 8 hours. Combining ability analyses of variance revealed highly significant differences in GCA values between parents for all characters studied and significant SCA values for some of the characters. GCA values for NAR and LAR, and for NAR and SLA were strongly negatively correlated. A strong positive correlation existed between GCA values for LAR and SLA. The possibilities for improving growth under low energy conditions by using growth characters in breeding are discussed.Abbreviations W dry weight of the plant - RGR relative growth rate - NAR net assimilation rate - LAR leaf area ratio - SLA specific leaf area - LWR leaf weight ratio - MANOVA multivariate analysis of variance - GCA general combining ability - SCA specific combining ability     

Inheritance of characters determining growth and development of tomato (Lycopersicon esculentum Mill.) under low energy conditions

Summary Diallel crosses were made between 15 tomato genotypes with varying performance under a low energy regime. It appeared that differences between genotypes for 11 vegetative and generative plant characters are determined mainly by additive genetic variation. For breeding cultivars adapted to low energy conditions, crossing genotypes with a high GCA for weight of trusses and fruits (WTF) and genotypes with a high GCA for weight of vegetative plant parts (WVP) seems most promising. Complications may arise from the negative correlation between GCA values for WTF and WVP, and the strong positive correlation between GCA values for fruit number and WTF, causing small fruits.     

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.     

Inheritance of net photosynthesis,dark respiration,stomatal resistance and related characters in tomato (Lycopersicon esculentum Mill.) under low energy conditions

Summary Inheritance of net photosynthesis, dark respiration, specific leaf fresh weight, and stomatal resistance was studied in a diallel cross without reciprocals with 15 tomato genotypes. The plants were grown under low light intensity (24 Wm^-2), short days (8 h) and low temperature (19 °C/10 °C day/night temperature). Significant f-values for GCA were observed for all characters. Smaller, but for most characters also significant f-values for SCA were found. This indicated that the characters were mainly controlled by additive gene action. The relationships between the physiological characters and breeding aspects are discussed.     

Differences between tomato genotypes in net photosynthesis and dark respiration under low light intensity and low night temperatures

Summary Net photosynthesis and dark respiration of sixteen selected tomato genotypes grown under low light intensity and low night temperatures were measured. Differences between genotypes in net photosynthesis and dark respiration were present. In all genotypes an increase of net photosynthesis with time was observed. Significant genotypic differences in this respect were established. The rate of dark respiration remained constant throughout the experimental period. The photosynthesis on a leaf area basis and dark respiration were not affected by the night temperature. Net photosynthesis on a fresh weight basis was lower under the lower night temperatures.     

Growth analyses of tomato genotypes grown under low night temperatures and low light intensity

Summary In three experiments the growth of 16 genotypes of tomato was examined. The plants were raised in growth rooms at a day temperature of 19°C and night temperatures of 14, 10 or 6°C respectively under a light intensity of 24 Wm^-2 visible radiation and a daylength of 8 hours. The results were analysed by a multivariate analysis of variance.Relative growth rate (RGR), leaf area ratio (LAR), specific leaf area (SLA) and leaf weight ratio (LWR) decreased with time. The change in net assimilation rate (NAR) was small. The decrease in RGR was mainly due to the decrease in LAR.RGR, LAR and SLA were lower and LWR slightly higher at lower night temperatures. NAR was hardly affected. The lower RGR at lower night temperatures was mainly due to a lower SLA.Significant differences between genotypes were found in total dry weight, NAR, LAR, SLA and LWR, but hardly in RGR. Significant genotype × temperature interactions occurred.LAR and SLA were positively, NAR and LAR, and NAR and SLA negatively correlated.     

Morphological and physiological characters affecting flower drop and fruit set of tomatoes at high temperatures

Summary Seven tomato cultivars and lines were studied under high temperature conditions. Fruit set varied between 77.3% in the heat tolerant cv. Hotset, 62% in cv. Gamad and 16.3% in the most sensitive cv. Hosen-Eilon. The characters contributing to low fruit set were bud drop, splitting of the antheridial cone, style exsertion and reduction of the quantity and/or functionality of the gametes. Employing the above characters as criteria for selection, fruit set of an F₄ line, phenotypically similar to the sensitive parent, was improved to 63.1%. Improved fruit set, 87.6%, was also obtained in an F₁ hybrid between Hotset and Gamad. The importance is discussed of various easily recognizable flower components contributing to satisfactory fruit set under high temperatures and their possible use in breeding is elaborated.     

Overcoming unilateral breeding barriers between Lycopersicon peruvianum and cultivated tomato,Lycopersicon esculentum

Summary Treatment of Lycopersicon peruvianum stigmas with an artificial medium analogous to stigmatic exudate allowed pollen germination and growth on immature pistils. Growth of Lycopersicon esculentum pollen tubes to L. peruvianum ovules, an otherwise incompatible cross, was achieved following such treated bud-pollinations. No plantlets were recovered, although a few embryos from this cross at the globularity heart stage of development were excised at 22 days after pollination, indicating the presence of crossing failures as severe as in the reciprocal cross. Hybrid plants were obtained from the reciprocal cross, using as pollen parent an L. peruvianum line selected for congruity with L. esculentum. Bud pollinations to L. peruvianum, using these interspecific F₁ hybrids as the pollen parent, allowed viable embryo development and plantlet recovery. Resulting backcross plants, which possess approximately 1/4 L. esculentum genome in a L. peruvianum cytoplasm, may facilitate further introgression of the L. esculentum nuclear genes into in foreign cytoplasm.     

Effects of root systems of tomato genotypes on growth and earliness,studied in grafting experiments at low temperature

Summary The contribution of the roots to growth and earliness of 29 genotypes of tomato cultivated under low energy conditions, was analysed in two grafting experiments.There were significant effects of the rootstocks on the vegetative growth, earliness and fruit production of the different scions, but not on plant length or number of leaves or trusses. There was hardly any interaction between rootstock and scion.The commercial rootstock KNVF and a commercial breeding line especially promoted vegetative growth of all scions, but the former delayed flowering of the scion by 6–7 days. The breeding line also stimulated early flowering. Most of the IVT breeding lines selected for growth and early production under low energy conditions scored very poorly as rootstock.Implications for breeding tomatoes with efficient root systems are discussed.     

Resistance at the TM-2 locus in the tomato to tomato mosaic virus

Summary There are three known tomato mosaic virus (TMV) resistance factors, Tm-1, Tm-2 and Tm-2 ², in the tomato. Tm-2 ² is currently the most widely utilised in glasshouse cultivars. Both Tm-2 and Tm-2² can induce systemic necroses in response to virus infection. These are considered to be hypersensitive resistance reactions in view of the low virus concentrations in affected plants and because sub-inoculation usually fails to infect all plants possessing the same resistance gene. The literature relating to TMV resistance at the Tm-2 locus in the tomato is reviewed.Virulent strains may readily establish when Tm-1 or Tm-2 are used, but Tm-2 ² confers more effective resistance. The possible development of aggressive isolates capable of affecting Tm-2 ²/Tm-2² plants is discussed. The establishment of virus types which cause systemic necrosis at normal growing temperatures is considered more likely than widespread infection from fully virulent strain 2² mutants. However, the growing of crops isolated from the TMV reservoirs in the soil considerably reduces the likelihood of even this occurring.     

The effect of the lateral suppressor gene on seed germination in the tomato

Summary A reduction in germination of tomato seed due to the lateral suppressor mutant (ls) is demonstrated. This is shown to be an effect of the maternal genotype rather than of the embryo. Poor germination is not determined until comparatively late in seed development and the effect can be avoided by embryo culture. Cuticular necrosis of the mutant seed appears to be associated with poor germination but the former symptom may be suppressed independently of the latter.     

Genetics of parthenocarpy in tomato under a low temperature regime: I. Line RP 75/59

Summary Genetics of parthenocarpy in line RP 75/59 was tested under natural low temperature conditions, under which only seedless fruits were produced. Results were consistent with the hypothesis that three recessive genes with additive effects are responsible for parthenocarpy. Linkage studies, using 40 morphological marker genes located among all tomato chromosomes except chromosome seven, revealed linkage of one gene to diageotropica (dgt) located on chromosome 1 L site 152, and a second gene to yellow verescent (yv) located on chromosome 6 L site 34. Location of the third gene involved in parthenocarpy could not be determined. To calculate the power of the linkage tests, a simulation was carried out for three genetic models; the results are presented graphically in two figures.     

Differences between tomato genotypes in stomatal resistance and specific leaf fresh weight in relation to differences in net photosynthesis under low light intensity and low night temperatures

Summary Differences between tomato genotypes (Lycopersicon esculentum Mill.) in stomatal resistance (r_s) and specific leaf fresh weight (SLFW) were observed under low light intensity and low night temperature. No relationship between r_s and previously published rates of net photosynthesis per dm² leaf area (P_A) was discovered. A negative relationship between SLFW and net photosynthesis per gram fresh weight (P_FW) was found but no clear relationship between SLFW and P_A. Genotypes with a low SLFW seem better adapted to a low light intensity.     

Evaluation of screening techniques and determination of criteria for assessing resistance to Corynebacterium michiganense in tomato

Summary When screening populations segregating for resistance to Corynebacterium michiganense a rapid and reliable inoculation technique is needed. From the several techniques tested, inoculation of the petiole was found to give better differentiation of resistant and susceptible plants than did inoculation of the root or leaf. When petiole-inoculation was compared with stem-inoculation the latter showed the lowest environmental variation and is suggested for use in genetic studies. Stem-inoculation involved clipping the stems of 4 week old seedlings 1 cm above the cotyledonary leaves with scissors which had been dipped in inoculum.Petiole inoculation and, to a lesser degree, stem inoculation failed to produce uniform infection, resulting in variation in the expression of the disease. The scoring was therefore based on the presence, and not the degree, of wilting and stunting, or the size of the cankers. Plants free of all these symptoms were rated resistant. The coefficient of variation for vascular discoloration on individual plants was calculated to be 50%, which in part may be explained by the histopathology of the host.The virulent isolates cm 21 caused death of seedlings resistant to isolate H. However, cultivars resistant to isolate H in the seedling stage did not decrease in fruit weight as compared to susceptible cultivars when inoculated with isolate cm 21 at anthesis.Michigan Agricultural Experiment Station Journal No. 7424.     

Perspectives of molecular marker assisted breeding for earliness in tomato

In a tomato (Lycopersicon esculentum) breeding programme aimed at earliness, a breeding line, designated IVT-KT₁, was developed by several rounds of crossing and selection. Among its ancestors were the two wild relatives L. pimpinellifolium and L. parviflorum. The breeding line flowered and set fruit one to four weeks earlier than old and modern cultivars. To identify QTLs for earliness, an F₂ population was obtained by crossing IVT-KT₁ with the late true breeding cultivar Premier. In winter and early spring, 690 plants of this F₂ population were evaluated for earliness of which 292 were selected for RFLP analysis. Only limited parts of the genomes of IVT-KT₁ and Premier were polymorphic and these polymorphisms were likely due to introgression from the ancestor wild species. By using the interval mapping method on the combined data of the greenhouse evaluations and RFLP analyses, three loci were identified that were associated with earliness. One was mainly associated with flowering time, another with fruitset time and a third one with ripening time. Two of these loci were also associated with fruit weight. F₃ progenies selected for earliness on the basis of the QTL-data did confirm the F₂ results. Two major earliness genes explained a difference in earliness of more than three weeks. The association with reduced fruit weight seemed to be due to pleiotropic effects rather than to linked genes. Consequently, the gain in earliness will cause a reduction in fruit weight that may not be acceptable for breeding applications.     

In vitro pollen germination and tube growth of tomato (Lycopersicon esculentum Mill.) and its relation with plant growth

Summary In vitro pollen germination at 10°C, 14°C and 22°C of four groups of two pure line tomato varieties was compared with their plant growth at 19°C D/10°C N under controlled environmental conditions. Generally, pollen germination was slow at 10°C but after 6 h the percentage of germination was similar to that at 22°C. Maximum germination was obtained at 14°C already after 4 h. The longest pollen tubes occurred at 22°C. The two varieties within each group differed significantly from each other for percentage of pollen germination. For one group, this difference was greater at 10°C than at 22°C. The varieties in two groups also differed significantly for pollen tube growth. These differences in pollen tube growth rate were greater at 22°C than at 10°C. There was no clear relationship between pollen germination, pollen tube growth and plant growth in any of the four pairs of varieties. The results are discussed with regard to the possibility of pollen selection at low temperature in order to improve the efficiency of breeding for growth at low temperature.     

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.     

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.     

The response to temperature of leaf area in tomato genotypes. I. cell size and number in relation to the area of a leaf

Summary An investigation was carried out into the genotypic variation in the response to temperature of leaf area in tomatoes. Differences between genotypes in the response of the area of a leaf to contrasting temperature regimes were found, and this was due to variation in the number of epidermal cells. The epidermal cell size showed little response to temperature. The results are discussed in relation to the potential for selection for improved leaf area and the development of a rapid screening test for cold tolerance.Guestworker Project Group Tomato Physiology.