An overview of heat stress in tomato (Solanum lycopersicum L.)

Heat stress has been defined as the rise of temperature for a period of time higher than a threshold level, thereby permanently affecting the plant growth and development. Day or night temperature is considered as the major limiting factor for plant growth. Earlier studies reported that night temperature is an important factor in the heat reaction of the plants. Tomato cultivars capable of setting viable fruits under night temperatures above 21 °C are considered as heat-tolerant cultivars. The development of breeding objectives is generally summarized in four points: (a) cultivars with higher yield, (b) disease resistant varieties in the 1970s, (c) long shelf-life in 1980s, and (d) nutritive and taste quality during 1990s. Some unique varieties like the dwarf “Micro-Tom”, and the first transgenic tomato (FlavrSavr) were developed through breeding; they were distributed late in the 1980s.High temperature significantly affects seed, pollen viability and root expansion. Researchers have employed different parameters to evaluate the tolerance to heat stress, including membrane thermo stability, floral characteristics (Stigma exertion and antheridia cone splitting), flower number, and fruit yield per plant. Reports on pollen viability and fruit set/plant under heat stress by comparing the pollen growth and tube development in heat-treated and non-heat-stressed conditions are available in literature. The electrical conductivity (EC) have been used to evaluate the tolerance of some tomato cultivars in vitro under heat stress conditions as an indication of cell damage due to electrolyte leakage; they classified the cultivars into three groups: (a) heat tolerant, (b) moderately heat tolerant, and (c) heat sensitive.It is important to determine the range in genetic diversity for heat tolerance in tomatoes. Heat stress experiments under field conditions offer breeders information to identify the potentially heat tolerant germplasm.     

Assessment of genetic variation in tomato （Solanum lycopersicum L.） inbred lines using SSR molecular markers

A study was conducted to determine the genetic diversity of 39 determinate and indeterminate tomato inbred lines collected from China, Japan, S. Korea, and USA. Using 35 SSR polymorphic markers, a total of 150 alleles were found with moderate levels of diversity, and a high number of unique alleles existing in these tomato lines. The mean number of alleles per locus was 4.3 and the average polymorphism information content (PIC) was 0.31. Unweighted Pair Group Method with Arithmetic Mean (UPGMA) clustering at genetic similarity value of 0.85 grouped the inbred lines into four groups, where one USA cultivar formed a separate and more distant cluster. The most similar inbred lines are from USA, both with determinate type, whereas the most different lines are from USA (Us-16) and Japan (Ja-2) with determinate and indeterminate growth habit, respectively. Clustering was consistent with the known information regarding geographical location and growth habit. The genetic distance information reported in this study might be used by breeders when planning future crosses among these inbred lines.     

Population structure and genetic differentiation associated with breeding history and selection in tomato (Solanum lycopersicum L.)

Tomato (Solanum lycopersicum L.) has undergone intensive selection during and following domestication. We investigated population structure and genetic differentiation within a collection of 70 tomato lines representing contemporary (processing and fresh-market) varieties, vintage varieties and landraces. The model-based Bayesian clustering software, STRUCTURE, was used to detect subpopulations. Six independent analyses were conducted using all marker data (173 markers) and five subsets of markers based on marker type (single-nucleotide polymorphisms, simple sequence repeats and insertion/deletions) and location (exon and intron sequences) within genes. All of these analyses consistently separated four groups predefined by market niche and age into distinct subpopulations. Furthermore, we detected at least two subpopulations within the processing varieties. These subpopulations correspond to historical patterns of breeding conducted for specific production environments. We found no subpopulation within fresh-market varieties, vintage varieties and landraces when using all marker data. High levels of admixture were shown in several varieties representing a transition in the demarcation between processing and fresh-market breeding. The genetic clustering detected by using the STRUCTURE software was confirmed by two statistics, pairwise F(st) (θ) and Nei's standard genetic distance. We also identified a total of 19 loci under positive selection between processing, fresh-market and vintage germplasm by using an F(st)-outlier method based on the deviation from the expected distribution of F(st) and heterozygosity. The markers and genome locations we identified are consistent with known patterns of selection and linkage to traits that differentiate the market classes. These results demonstrate how human selection through breeding has shaped genetic variation within cultivated tomato.     

Effects of Penicillium extracts on germination vigour in subsequent seedling growth of tomato (Solanum lycopersicum L.)

Penicillium spp. are well known to produce a variety of beneficial metabolites for plant growth and survival, as well as they defend their hosts from attack of certain pathogens. In this study, effects of culture filtrate of different Penicillium spp. were tested on tomato seeds. On the whole, presoaking of seeds in the filtrates of the nine Penicillium isolates tested significantly increased seed germination when compared with the control seeds. Cultural extracts of P. expansum and P. bilaii were highly effective in growth promotion up to 90%. It was also observed that P. implicatum and P. oxalicum significantly enhanced the root growth in tomato seedling as compared to other species. In case of shoot length, P. verrucosum (3.38 cm), P. granulatum (2.81 cm) and P. implicatum (2.62 cm) were effective. P. implicatum was quite promising in increasing shoot and root length in tomato seedlings, while P. simplicissimum and P. citrinum were less effective on seedling growth. The plant growth-promoting ability of Penicillium strains may help in growth promotion in other plants and crops. Penicillium spp. are already known for producing various mycotoxins and enzymes. Plant growth-promoting ability of Penicillium spp. will open new aspects of research and investigations. The role of Penicillium spp. in tomato plant growth requires further exploration.     

Piriformospora indica inoculation alleviates the adverse effect of NaCl stress on growth,gas exchange and chlorophyll fluorescence in tomato (Solanum lycopersicum L.)

• Salinity is now an increasingly serious environmental issue that affects the growth and yield of many plants.
• In the present work, the influence of inoculation with the symbiotic fungus, Piriformospora indica, on gas exchange, water potential, osmolyte content, Na/K ratio and chlorophyll fluorescence of tomato plants under three salinity levels (0, 50, 100 and 150 mm NaCl) and three time periods (5, 10 and 15 days after exposure to salt) was investigated.
• Results indicate that P. indica inoculation improved growth parameters of tomato under salinity stress. This symbiotic fungus significantly increased photosynthetic pigment content under salinity, and more proline and glycine betaine accumulated in inoculated roots than in non‐inoculated roots. P. indica further significantly improved K⁺ content and reduced Na⁺ level under salinity treatment. After inoculation with the endophytic fungus, leaf physiological parameters, such as water potential, net photosynthesis, stomatal conductance and transpiration, were all higher under the salt concentrations and durations compared with controls without P. indica. With increasing salt level and salt treatment duration, values of F₀ and qP increased but F_m, F_v/F_m, F′_v/F′_m and NPQ declined in the controls, while inoculation with P. indica improved these values.
• The results indicate that the negative effects of NaCl on tomato plants were alleviated after P. indica inoculation, probably by improving physiological parameters such as water status and photosynthesis.

     

Overexpression of osmotin gene confers tolerance to salt and drought stresses in transgenic tomato (Solanum lycopersicum L.)

Abiotic stresses, especially salinity and drought, are major limiting factors for plant growth and crop productivity. In an attempt to develop salt and drought tolerant tomato, a DNA cassette containing tobacco osmotin gene driven by a cauliflower mosaic virus 35S promoter was transferred to tomato (Solanum lycopersicum) via Agrobacterium-mediated transformation. Putative T0 transgenic plants were screened by PCR analysis. The selected transformants were evaluated for salt and drought stress tolerance by physiological analysis at T1 and T2 generations. Integration of the osmotin gene in transgenic T1 plants was verified by Southern blot hybridization. Transgenic expression of the osmotin gene was verified by RT-PCR and northern blotting in T1 plants. T1 progenies from both transformed and untransformed plants were tested for salt and drought tolerance by subjecting them to different levels of NaCl stress and by withholding water supply, respectively. Results from different physiological tests demonstrated enhanced tolerance to salt and drought stresses in transgenic plants harboring the osmotin gene as compared to the wild-type plants. The transgenic lines showed significantly higher relative water content, chlorophyll content, proline content, and leaf expansion than the wild-type plants under stress conditions. The present investigation clearly shows that overexpression of osmotin gene enhances salt and drought stress tolerance in transgenic tomato plants.     

Genetic diversity, structure and marker-trait associations in a collection of Italian tomato (Solanum lycopersicum L.) landraces

The study of phenotypic and genetic diversity in landrace collections is important for germplasm conservation. In addition, the characterisation of very diversified materials with molecular markers offers a unique opportunity to define significant marker-trait associations of biological and agronomic interest. Here, 50 tomato landraces (mainly collected in central Italy), nine vintage and modern cultivars, and two wild outgroups were grown at two locations in central Italy and characterised for 15 morpho-physiological traits and 29 simple sequence repeat (SSR) loci. The markers were selected to include a group of loci in regions harbouring reported quantitative trait loci (QTLs) that affect fruit size and/or shape (Q-SSRs) and a group of markers that have not been mapped or shown to have a priori known linkage (NQ-SSRs). As revealed by univariate and multivariate analyses of morphological data, the landraces grouped according to vegetative and reproductive traits, with emphasis on fruit size, shape and final destination of the product. Compared to the low molecular polymorphism reported in tomato modern cultivars, our data reveal a high level of molecular diversity in landraces. Such diversity has allowed the inference of the existence of a genetic structure that was factored into the association analysis. As the proportion of significant associations is higher between the Q-SSR subset of markers and the subset of traits related to fruit size and shape than for all of the other combinations, we conclude that this approach is valid for establishing true-positive marker-trait relationships in tomato. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Improved growth,productivity and quality of tomato (Solanum lycopersicum L.) plants through application of shikimic acid

A field experiment was conducted to investigate the effect of seed presoaking of shikimic acid (30, 60 and 120 ppm) on growth parameters, fruit productivity and quality, transpiration rate, photosynthetic pigments and some mineral nutrition contents of tomato plants. Shikimic acid at all concentrations significantly increased fresh and dry weights, fruit number, average fresh and dry fruit yield, vitamin C, lycopene, carotenoid contents, total acidity and fruit total soluble sugars of tomato plants when compared to control plants. Seed pretreatment with shikimic acid at various doses induces a significant increase in total leaf conductivity, transpiration rate and photosynthetic pigments (Chl. a, chl. b and carotenoids) of tomato plants. Furthermore, shikimic acid at various doses applied significantly increased the concentration of nitrogen, phosphorus and potassium in tomato leaves as compared to control non-treated tomato plants. Among all doses of shikimic acid treatment, it was found that 60 ppm treatment caused a marked increase in growth, fruit productivity and quality and most studied parameters of tomato plants when compared to other treatments. On the other hand, no significant differences were observed in total photosynthetic pigments, concentrations of nitrogen and potassium in leaves of tomato plants treated with 30 ppm of shikimic acid and control plants. According to these results, it could be suggested that shikimic acid used for seed soaking could be used for increasing growth, fruit productivity and quality of tomato plants growing under field conditions.     

Genome-wide and molecular evolution analyses of the KT/HAK/KUP family in tomato (Solanum lycopersicum L.)

Potassium transporters belonging to the KT/HAK/KUP family play an important role in plant growth, development, mineral nutrition, and stress adaptation. In this study, we identified 19 KT/HAK/KUP family genes in tomato, distributed on 10 chromosomes, by using bioinformatics methods. A complete overview of the KT/HAK/KUP (SlHAK) genes in tomato is presented, including chromosome location, phylogeny, gene structure, and evolution pattern. Phylogenetic analysis of 19 SlHAK proteins suggested that group IV of the KT/HAK/KUP family is absent in the tomato genome. In addition, five pairs of segmental duplicated paralogs and two pairs of tandem duplicated paralogs were identified in the tomato KT/HAK/KUP family. This suggests that segmental duplication is predominant for the expansion of the SlHAK genes. Calculation of the approximate dates of duplication events using the synonymous substitution rate indicated that the segmental duplication of the KT/HAK/KUP genes in tomato originated 35.89–62.77 million years ago. Adaptive evolution analysis showed that purifying selection contributed to the evolution of segmental duplicated pairs. Furthermore, Tajima’s relative rate test indicated that all segmental duplicated pairs evolved at similar rates. As a first step toward a genome-wide analysis of the KT/HAK/KUP gene family in tomato, our results provide valuable information for understanding the function and evolution of the KT/HAK/KUP gene family in tomato and other species.     

Immunolocalization of plasma-membrane H+-ATPase and tonoplast-type pyrophosphatase in the plasma membrane of the sieve element-companion cell complex in the stem of Ricinus communis L

Plasma-membrane-located primary pumps were investigated in the sieve element (SE)-companion cell complex in the transport phloem of 2-week-old stems of Ricinus communis L. and, for comparison, in stems of Cucurbita pepo L. and in the secondary phloem of Agrobacterium tumefaciens-induced crown galls as a typical sink tissue. The plasma-membrane (PM) H⁺-ATPase and the tonoplast-type pyrophosphatase (PPase) were immunolocalized by epifluorescence and confocal laser scanning microscopy (CLSM) upon single or double labeling with specific monoclonal and polyclonal antibodies. Quantitative fluorescence evaluation by CLSM revealed both pumps in one membrane, the sieve-element PM. Different PM H⁺-ATPase antibody clones, raised against the PM H⁺-ATPase of Zea mays coleoptiles, induced in mouse and produced in mouse hybridoma cells, discriminated between different phloem cell types. Clones 30D5C4 and 44B8A1 labeled sieve elements and clone 46E5B11D5 labeled companion cells, indicating the existence of different phloem PM H⁺-ATPase isoforms. The results are discussed in terms of energization of SE transporters for retrieval of leaking sucrose, K⁺ and amino acids, as one of the unknown roles of ATP found in SEs. The function of the PPase could be related to phloem sucrose metabolism in support of ATP-requiring processes. Received: 3 July 2000 / Accepted: 12 October 2000     

Alleviation of cadmium phytotoxicity through melatonin modulated physiological functions,antioxidants, and metabolites in tomato (Solanum lycopersicum L.)

BioMetals - The rising concentration of cadmium (Cd) builds a harmful effect on human and plant health associated with food chain contagion. Melatonin (MT) is an indole compound. Hence, the...     

Exploring the interacting effect of soil texture and bulk density on root system development in tomato (Solanum lycopersicum L.)

Knowledge of the responses of root systems in horizoned heterogeneous soil is vital to optimise uptake of water and nutrients to maximise crop productivity. We explored the interacting effects of soil bulk density and texture on the development of root systems in tomato.Two main techniques were employed, X-ray micro-Computed Tomography (μCT), to provide non-destructive, three-dimensional (3D) images of root systems in situ and destructive root washing followed by WinRHIZO^® scanning. Solanum lycopersicum L. cv. Ailsa Craig plants were grown in soil columns for 10 days to measure the effect of soil compaction on selected root traits. Treatments included bulk density (1.2–1.6 Mg m⁻³), soil texture (loamy sand and clay loam) and the effects of layering.The effect of bulk density on root growth was greatest 3 days after transplanting (DAT) in both soil types. The effect of soil texture was not apparent at this stage, but was significant at 10 DAT for most root and shoot variables. The influence of bulk density differed between soil types as increasing compaction promoted plant growth in clay loam but retarded root growth in loamy sand.We observed that at 3 DAT root growth is primarily influenced by bulk density but by 10 DAT a switch in the processes regulating root growth occurs and the texture of the soil becomes very influential. Future investigations of root growth must consider soil physical properties individually and at specific time points, as their importance changes as the root system becomes established. Here we have demonstrated both positive and negative impacts across a wide range of bulk density treatments in different soil textures on root growth. This illustrates the importance of understanding the complex nature of root–soil interactions, especially for agricultural practices such as seedbed preparation.     

Non-destructive assessment of flesh firmness and dietary antioxidants of greenhouse-grown tomato (Solanum lycopersicum L.) at different fruit maturity stages

Non-destructive methods have been widely recognized for evaluating fruit quality traits of many horticultural crops and food processing industry. Destructive (analytical) test, and non-destructive evaluation of the quality traits were investigated and compared for ‘Red Rose’ tomato (Solanum lycopersicum L.) fruit grown under protected environment. Fresh tomato fruit at five distinctive maturity stages namely; breaker (BK), turning (TG), pink (PK), light-red (LR), and red (RD) were labeled and scanned using the handheld near infra-red (NIR) enhanced spectrometer at a wavelength range of 285–1200 nm. The labeled tomato samples were then measured analytically for flesh firmness, lycopene, β-carotene, total phenolic content (TPC) and total flavonoids content (TFC). The results revealed that quality traits could be estimated using NIR spectroscopy with a relatively high coefficient of determination (R²): 0.834 for total phenolic content, 0.864 for lycopene, 0.790 for total flavonoid content, 0.708 for β-carotene; and 0.679 for flesh firmness. The accumulation of Lyco and β-Car rapidly increased in tomatoes harvested between the TG and the LR maturity stages. Harvesting tomatoes at BK maturity stage resulted in significantly higher flesh firmness than harvesting at the later maturity stages. Tomato fruits had the lowest TPC and TFC contents at the earliest maturity stage (BK), while they had intermediate TPC and TFC levels at LR and RD maturity stages. NIR spectroscopic measurements of fruit firmness and lipophilic antioxidants in tomato fruit at various maturity stages were partially in accordance with those estimated by destructive (analytical) methods. Based on these findings, we recommend using non-destructive NIR spectroscopy as an effective tool for predicting tomato fruit quality during harvest stage and postharvest processing.     

Characterization of tomato (Solanum lycopersicum L.) mutants affected in their flowering time and in the morphogenesis of their reproductive structure

The impact of the season on flowering time and the organization and morphogenesis of the reproductive structures are described in three tomato mutants: compound inflorescence (s), single flower truss (sft), and jointless (j), respectively, compared with their wild-type cultivars Ailsa Craig (AC), Platense (Pl), and Heinz (Hz). In all environmental conditions, the sft mutant flowered significantly later than its corresponding Pl cultivar while flowering time in j was only marginally, but consistently, delayed compared with Hz. The SFT gene and, to a lesser extent, the J gene thus appear to be constitutive flowering promoters. Flowering in s was delayed in winter but not in summer compared with the AC cultivar, suggesting the existence of an environmentally regulated pathway for the control of floral transition. The reproductive structure of tomato is a raceme-like inflorescence and genes regulating its morphogenesis may thus be divided into inflorescence and floral meristem identity genes as in Arabidopsis. The s mutant developed highly branched inflorescences bearing up to 200 flowers due to the conversion of floral meristems into inflorescence meristems. The S gene appears to be a floral meristem identity gene. Both sft and j mutants formed reproductive structures containing flowers and leaves and reverting to a vegetative sympodial growth. The SFT gene appears to regulate the identity of the inflorescence meristem of tomato and is also involved, along with the J gene, in the maintenance of this identity, preventing reversion to a vegetative identity. These results are discussed in relation to knowledge accumulated in Arabidopsis and to domestication processes.     

Polysomes and intracisternal accumulations in enucleate sieve elements of rice (Oryza sativa L.)

Polysomes in sieve elements of rice (Oryza sativa L.) were studied with the electron microscope. The polysomes were found on the rough endoplasmic reticulum (ER) present in immature sieve elements and also on the cisternae of aggregated ER in the parietal layer of mature, enucleate sieve elements. In the immature sieve elements the ER cisternae existed as narrow profiles while in the mature sieve elements the ER cisternae were considerably dilated and contained a fibrillar material and, occasionally, electron-opaque inclusions. In addition to the aggregated ER, single profiles of ER were found applied to the lateral walls and also the sieve plates. These cisternae also bore ribosomes and were separated from the plasmalemma by a narrow, dense space. In the mature sieve elements much of the surface of the ER membranes was covered with polysomes. The dimensions of the polysomes are described and the possibility that they contribute to the formation of the fibrillar material in the intracisternal space is discussed.Abbreviations ER endoplasmic reticulum     

Effect of partial rootzone drying on the concentration of zeatin-type cytokinins in tomato (Solanum lycopersicum L.) xylem sap and leaves

Decreased cytokinin (CK) export from roots in drying soil might provide a root-to-shoot signal impacting on shoot physiology. Although several studies show that soil drying decreases the CK concentration of xylem sap collected from the roots, it is not known whether this alters xylem CK concentration ([CK(xyl)]) in the leaves and bulk leaf CK concentration. Tomato (Solanum lycopersicum L.) plants were grown with roots split between two soil columns. During experiments, water was applied to both columns (well-watered; WW) or one (partial rootzone drying; PRD) column. Irrigation of WW plants aimed to replace transpirational losses every day, while PRD plants received half this amount. Xylem sap was collected by pressurizing detached leaves using a Scholander pressure chamber, and zeatin-type CKs were immunoassayed using specific antibodies raised against zeatin riboside after separating their different forms (free zeatin, its riboside, and nucleotide) by thin-layer chromatography. PRD decreased the whole plant transpiration rate by 22% and leaf water potential by 0.08 MPa, and increased xylem abscisic acid (ABA) concentration 2.5-fold. Although PRD caused no detectable change in [CK(xyl)], it decreased the CK concentration of fully expanded leaves by 46%. That [CK(xyl)] was maintained and not increased while transpiration decreased suggests that loading of CK into the xylem was also decreased as the soil dried. That leaf CK concentration did not decline proportionally with CK delivery suggests that other mechanisms such as CK metabolism influence leaf CK status of PRD plants. The causes and consequences of decreased shoot CK status are discussed.     

CO2 enrichment using CRAM fermentation improves growth,physiological traits and yield of cherry tomato (Solanum lycopersicum L.)

Carbon dioxide (CO₂) concentration in greenhouses is sub-optimal for vegetable production. Many techniques have been used to increase CO₂ concentration in greenhouses but most of them are expensive with certain limitations and drawbacks. We adopted a new strategy to elevate CO₂ concentration in the greenhouse throughout the day via crop residues and animal manure composting (CRAM). During the whole cultivation period, CRAM-treated greenhouse had doubled CO₂ concentration which significantly increased the yield of cherry tomatoes (Solanum lycopersicum L.), i.e., up to 38%. The influence of CRAM procedure on cherry tomato quality was also investigated and the concentrations of total soluble solids (TSS) and soluble sugar were found to be significantly higher in cherry tomatoes grown under composting greenhouse than that of non-composting greenhouse. Additionally, CRAM-CO₂ enrichment also resulted in increased concentrations of ascorbic acid (Vitamin C) and titrate acid as compared to control. In contrast, the concentration of nitrate was considerably decreased in cherry tomato grown under CO₂ enriched condition than that of control. The increase in active oxygen metabolisms such as POD, CAT and SOD while a decrease in MDA, as well as APX was observed for cherry tomatoes grown under CO₂ enriched condition. Hence, CO₂ fertilization by using CRAM in greenhouse significantly improved quality and increased the yield of cherry tomatoes.     

A genome‐scale metabolic network reconstruction of tomato (Solanum lycopersicum L.) and its application to photorespiratory metabolism

Tomato (Solanum lycopersicum L.) has been studied extensively due to its high economic value in the market, and high content in health‐promoting antioxidant compounds. Tomato is also considered as an excellent model organism for studying the development and metabolism of fleshy fruits. However, the growth, yield and fruit quality of tomatoes can be affected by drought stress, a common abiotic stress for tomato. To investigate the potential metabolic response of tomato plants to drought, we reconstructed iHY3410, a genome‐scale metabolic model of tomato leaf, and used this metabolic network to simulate tomato leaf metabolism. The resulting model includes 3410 genes and 2143 biochemical and transport reactions distributed across five intracellular organelles including cytosol, plastid, mitochondrion, peroxisome and vacuole. The model successfully described the known metabolic behaviour of tomato leaf under heterotrophic and phototrophic conditions. The in silico investigation of the metabolic characteristics for photorespiration and other relevant metabolic processes under drought stress suggested that: (i) the flux distributions through the mevalonate (MVA) pathway under drought were distinct from that under normal conditions; and (ii) the changes in fluxes through core metabolic pathways with varying flux ratio of RubisCO carboxylase to oxygenase may contribute to the adaptive stress response of plants. In addition, we improved on previous studies of reaction essentiality analysis for leaf metabolism by including potential alternative routes for compensating reaction knockouts. Altogether, the genome‐scale model provides a sound framework for investigating tomato metabolism and gives valuable insights into the functional consequences of abiotic stresses.