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European traditional tomato varieties have been selected by farmers given their consistent performance and adaptation to local growing conditions. Here we developed a multipurpose core collection, comprising 226 accessions representative of the genotypic, phenotypic, and geographical diversity present in European traditional tomatoes, to investigate the basis of their phenotypic variation, gene×environment interactions, and stability for 33 agro-morphological traits. Comparison of the traditional varieties with a modern reference panel revealed that some traditional varieties displayed excellent agronomic performance and high trait stability, as good as or better than that of their modern counterparts. We conducted genome-wide association and genome-wide environment interaction studies and detected 141 quantitative trait loci (QTLs). Out of those, 47 QTLs were associated with the phenotype mean (meanQTLs), 41 with stability (stbQTLs), and 53 QTL-by-environment interactions (QTIs). Most QTLs displayed additive gene actions, with the exception of stbQTLs, which were mostly recessive and overdominant QTLs. Both common and specific loci controlled the phenotype mean and stability variation in traditional tomato; however, a larger proportion of specific QTLs was observed, indicating that the stability gene regulatory model is the predominant one. Developmental genes tended to map close to meanQTLs, while genes involved in stress response, hormone metabolism, and signalling were found within regions affecting stability. A total of 137 marker-trait associations for phenotypic means and stability were novel, and therefore our study enhances the understanding of the genetic basis of valuable agronomic traits and opens up a new avenue for an exploitation of the allelic diversity available within European traditional tomato germplasm.
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Solanum lycopersicum , Mapeo Cromosómico , Solanum lycopersicum/genética , Estudio de Asociación del Genoma Completo , Sitios de Carácter Cuantitativo , FenotipoRESUMEN
A comprehensive collection of 1254 tomato accessions, corresponding to European traditional and modern varieties, early domesticated varieties, and wild relatives, was analyzed by genotyping by sequencing. A continuous genetic gradient between the traditional and modern varieties was observed. European traditional tomatoes displayed very low genetic diversity, with only 298 polymorphic loci (95% threshold) out of 64 943 total variants. European traditional tomatoes could be classified into several genetic groups. Two main clusters consisting of Spanish and Italian accessions showed higher genetic diversity than the remaining varieties, suggesting that these regions might be independent secondary centers of diversity with a different history. Other varieties seem to be the result of a more recent complex pattern of migrations and hybridizations among the European regions. Several polymorphic loci were associated in a genome-wide association study with fruit morphological traits in the European traditional collection. The corresponding alleles were found to contribute to the distinctive phenotypic characteristic of the genetic varietal groups. The few highly polymorphic loci associated with morphological traits in an otherwise a low-diversity population suggests a history of balancing selection, in which tomato farmers likely maintained the morphological variation by inadvertently applying a high selective pressure within different varietal types.
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Solanum lycopersicum , Alelos , Agricultores , Variación Genética , Estudio de Asociación del Genoma Completo , Humanos , Solanum lycopersicum/genética , Fenotipo , Polimorfismo de Nucleótido SimpleRESUMEN
MAIN CONCLUSION: The introgression of the A ft allele into domesticated tomato induced a shift from flavonol to anthocyanin production in response to UV-B radiation, while the hp - 1 allele negatively influenced the response of flavonoid biosynthesis to UV-B. Introgression of the dominant allele Anthocyanin fruit (Aft) from Solanum chilense induces anthocyanin accumulation in the peel of tomato (Solanum lycopersicum L.) fruit. UV-B radiation can influence plant secondary metabolism regulating the expression of several genes, among which those involved in flavonoid biosynthesis. Here, we investigated whether post-harvest UV-B treatment could up-regulate flavonoid production in tomato fruits and whether the Aft allele could affect flavonoid biosynthesis under UV-B radiation. Mature green fruits of an anthocyanin-rich tomato mutant line (SA206) and of its wild-type reference, cv. Roma, were daily subjected to post-harvest UV-B treatment until full ripening. Up-regulation of CHS and CHI transcription by UV-B treatment induced flavonoid accumulation in the peel of cv. Roma. Conversely, UV-B decreased the total flavonoid content and CHS transcript levels in the SA206 peel. SA206 being a double mutant containing also hp-1 allele, we investigated also the behavior of hp-1 fruit. The decreased peel flavonoid accumulation and gene transcription in response to UV-B suggest that hp-1 allele is involved in the marked down-regulation of the flavonoid biosynthesis observed in SA206 fruit. Interestingly, in SA206, UV-B radiation promoted the synthesis of delphinidin, petunidin, and malvidin by increasing F3'5'H and DFR transcription, but it decreased rutin production, suggesting a switch from flavonols to anthocyanins. Finally, although UV-B radiation does not reach the inner fruit tissues, it down-regulated flavonoid biosynthesis in the flesh of both genotypes. This study provides, for the first time, evidence that the presence of the functional Aft allele, under UV-B radiation, redirects flavonoid synthesis towards anthocyanin production and suggests that the hp-1 allele negatively influences the response of flavonoid biosynthesis to UV-B.
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Antocianinas/metabolismo , Flavonoles/metabolismo , Regulación de la Expresión Génica de las Plantas/efectos de la radiación , Proteínas de Plantas/metabolismo , Solanum lycopersicum/genética , Solanum/genética , Alelos , Regulación hacia Abajo , Flavonoides/metabolismo , Genes Dominantes , Genotipo , Solanum lycopersicum/metabolismo , Solanum lycopersicum/efectos de la radiación , Proteínas de Plantas/genética , Metabolismo Secundario , Regulación hacia ArribaRESUMEN
KEY MESSAGE: We characterized the photoperiod-sensitive 7B - 1 male-sterile mutant in tomato, showing its allelism with stamenless - 2 . Mapping experiments indicated SlGLO2 , a B-class MADS-box family member, as a strong candidate to underlie the 7B - 1 mutation. The interest in male sterility (MS) dates back to a long time due to its perspective use in hybrid seed production. Here, we characterize 7B-1, a photoperiod-sensitive male-sterile (ms) mutant in tomato (Solanum lycopersicum L.), in which stamens are restored to fertility by permissive growth conditions in short days (SD). This system represents a useful strategy to facilitate the maintenance of the ms line. Examination of 7B-1 and other structural mutants, vms, sl, sl-2 and tap3, showed carpellization of stamens in the third floral whorl. 7B-1 exhibits strong expressivity in long days (LD), producing 100% aberrant anthers and virtually no seed production under open pollination, whereas it recovered fertility in SD. By genetic analysis, we demonstrate that 7B-1 is not allelic to sl nor to vms; instead it shows allelism to sl-2. Because the homeotic phenotype of the mutation resembles lesions to members of the B-class MADS-box transcription factor family, that specify petal and stamen identity, we pursued a candidate gene approach towards these targets. Using an interspecific backcross mapping population and markers linked to B-class MADS-box genes, significant linkage was found between 7B-1 and the SlGLO2 gene on Chr6. This result was supported by the 7B-1 phenotype that is similar to that of SlGLO2 knock outs and by the strong downregulation of the gene in the mutant. Although the lesion underlying the mutant phenotype is still elusive, our results pave the way for the final demonstration that SlGLO2 underlies 7B-1 and further the use of 7B-1 mutant in tomato hybrid seed production schemes.
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Infertilidad Vegetal/genética , Solanum lycopersicum/genética , Alelos , Mapeo Cromosómico , Cruzamientos Genéticos , Flores/genética , Ligamiento Genético , Marcadores Genéticos , Solanum lycopersicum/fisiología , Mutación , Fenotipo , Polen/genéticaRESUMEN
The tomato parthenocarpic fruit (pat) mutation associates a strong competence for parthenocarpy with homeotic transformation of anthers and aberrancy of ovules. To dissect this complex floral phenotype, genes involved in the pollination-independent fruit set of the pat mutant were investigated by microarray analysis using wild-type and mutant ovaries. Normalized expression data were subjected to one-way ANOVA and 2499 differentially expressed genes (DEGs) displaying a >1.5 log-fold change in at least one of the pairwise comparisons analyzed were detected. DEGs were categorized into 20 clusters and clusters classified into five groups representing transcripts with similar expression dynamics. The "regulatory function" group (685 DEGs) contained putative negative or positive fruit set regulators, "pollination-dependent" (411 DEGs) included genes activated by pollination, "fruit growth-related" (815 DEGs) genes activated at early fruit growth. The last groups listed genes with different or similar expression pattern at all stages in the two genotypes. qRT-PCR validation of 20 DEGs plus other four selected genes assessed the high reliability of microarray expression data; the average correlation coefficient for the 20 DEGs was 0.90. In all the groups were evidenced relevant transcription factors encoding proteins regulating meristem differentiation and floral organ development, genes involved in metabolism, transport and response of hormones, genes involved in cell division and in primary and secondary metabolism. Among pathways related to secondary metabolites emerged genes related to the synthesis of flavonoids, supporting the recent evidence that these compounds are important at the fruit set phase. Selected genes showing a de-regulated expression pattern in pat were studied in other four parthenocarpic genotypes either genetically anonymous or carrying lesions in known gene sequences. This comparative approach offered novel insights for improving the present molecular understanding of fruit set and parthenocarpy in tomato.
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Frutas/crecimiento & desarrollo , Regulación de la Expresión Génica de las Plantas/fisiología , Solanum lycopersicum/genética , Transcriptoma/fisiología , Análisis por Conglomerados , Flores/genética , Flores/metabolismo , Solanum lycopersicum/metabolismo , Proteínas de Plantas/genética , Proteínas de Plantas/metabolismoRESUMEN
Pomaces obtained from three San Marzano tomato genotypes including the wild type (WT), Sun Black (SB), and colorless fruit epidermis (CL) were dried at 50 °C and analyzed for nutritional composition, total polyphenol (TPC), flavonoid (TFC) content, polyphenol qualitative profile, total antioxidant capacity (TAC), and antimicrobial activity. Commercial dried tomato powder (CTRP) was included as a control. No differences were detected nutritionally, in TPC and antimicrobial activity, but significant changes were observed for TFC and TAC, underlying variation in the phenolic profile. SB pomace (SBP) had the highest TFC and TAC. LC-HRMS analysis showed a flavonoid-enriched profile in SBP besides the exclusive presence of anthocyanins, with petanin and negretein as the most abundant. Among flavonoids, quercetin-hexose-deoxyhexose-pentose, naringenin, and rutin were the major. Overall, we showed the potential of dried tomato pomace, especially SBP, as an extremely valuable waste product to be transformed into a functional ingredient, reducing the food industry waste.
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Antioxidantes , Flavonoides , Frutas , Solanum lycopersicum , Residuos , Solanum lycopersicum/química , Antioxidantes/química , Residuos/análisis , Frutas/química , Flavonoides/química , Flavonoides/análisis , Polifenoles/química , Polifenoles/farmacología , Polifenoles/análisis , Extractos Vegetales/química , Extractos Vegetales/farmacología , Alimento Perdido y DesperdiciadoRESUMEN
Parthenocarpy allows fruit set independently of fertilization. In parthenocarpic-prone tomato genotypes, fruit set can be achieved under pollen-limiting environmental conditions and in sterile mutants. Parthenocarpy is also regarded as a quality-related trait, when seedlessness is associated with positive fruit quality aspects. Among the different sources of genetic parthenocarpy described in tomato, the parthenocarpic fruit (pat) mutation is of particular interest because of its strong expressivity, high fruit set, and enhanced fruit quality. The complexity of the pat "syndrome" associates a strong competence for parthenocarpy with a complex floral phenotype involving stamen and ovule developmental aberrations. To understand the genetic basis of the phenotype, we mapped the pat locus within a 0.19-cM window of Chr3, comprising nine coding loci. A non-tolerated missense mutation found in the 14th exon of Solyc03g120910, the tomato ortholog of the Arabidopsis HD-Zip III transcription factor HB15 (SlHB15), cosegregated with the pat phenotype. The role of SlHB15 in tomato reproductive development was supported by its expression in developing ovules. The link between pat and SlHB15 was validated by complementation and knock out experiments by co-suppression and CRISPR/Cas9 approaches. Comparing the phenotypes of pat and those of Arabidopsis HB15 mutants, we argued that the gene plays similar functions in species with fleshy and dry fruits, supporting a conserved mechanism of fruit set regulation in plants.
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Studies on the reproductive dynamics under heat stress are crucial to breed more tolerant cultivars. In tomato, cultivars, breeding lines, and wild species have been evaluated for their response to heat stress. Here, we addressed the study to a panel of selected landraces representing traditional genotypes that usually show high adaptation to local environments. In two experiments, spaced by 12 years, we set-up an identical experimental design with plants transplanted at two different dates to expose the second field to thermic stress with natural fluctuations. Such a strategy resulted in both a mild and severe stress in the two years. The landraces showed wide variation for both vegetative and reproductive traits; all traits were affected by heat, mostly with a significant Genotype*Environment interaction. A high broad-sense heritability was estimated for plant height, stigma position, pollen viability, and fruit weight. Low heritability estimates were found for the number of flowers, fruit set, and yield. Despite the interaction, traits recorded under control and heat conditions were positively correlated. Multivariate analysis located the genotypes in a topography that was stable under all conditions, except under the harshest temperatures. The study revealed that landraces present a wide variability for the response of reproductive traits to thermic challenges and that such a variation could be useful to dissect the traits with higher heritability and identify quantitative trait loci for breeding more resilient varieties.
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Tomatoes are among the most consumed vegetables worldwide and represent a source of health-beneficial substances. Our study represents the first investigating the peel-associated epiphytic bacteria of red and purple (anthocyanin-rich) tomatoes subjected to organic and conventional farming systems. Proteobacteria was the dominant phylum (relative abundances 79-91%) in all experimental conditions. Enterobacteriaceae represented a large fraction (39.3-47.5%) of the communities, with Buttiauxella and Atlantibacter as the most represented genera. The core microbiota was composed of 59 operational taxonomic units (OTUs), including the majority of the most abundant ones. The occurrence of the most abundant OTUs differed among the experimental conditions. OTU 1 (Buttiauxella), OTU 2 (Enterobacteriales), and OTU 6 (Bacillales) were higher in red and purple tomatoes grown under organic farming. OTU 5 (Acinetobacter) had the highest abundance in red tomatoes subjected to organic farming. OTU 3 (Atlantibacter) was among the major OTUs in red tomatoes under both farming conditions. OTU 7 (Clavibacter) and OTU 8 (Enterobacteriaceae) had abundances ≥1% only in red tomatoes grown under conventional farming. PCA and clustering analysis highlighted a high similarity between the bacterial communities of red and purple tomatoes grown under organic farming. Furthermore, the bacterial communities of purple tomatoes grown under organic farming showed the lowest diversity and evenness. This work paves the way to understand the role of nutritional superior tomato genotypes, combined with organic farming, to modulate the presence of beneficial/harmful bacteria and supply healthier foods within a sustainable agriculture.
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Italy is a recognized secondary center of diversification for cultivated tomato (Solanum lycopersicum L.). The study of phenotypic and genetic diversity in landrace collections is important for germplasm conservation and valorization. Here, we set up to study the tomato germplasm collected in the region of Lazio in Central Italy, with a focus on the distinctiveness among landraces and the attribution of membership to unnamed accessions. Our regional collection included 32 accessions belonging to eight different locally recognized landraces and 19 unnamed accessions. All accessions were gathered from local farmers and are preserved in the collection held at the Regional Agency for the Development and the Innovation of Lazio Agriculture (ARSIAL) and at the University of Tuscia. We included 13 control genotypes comprising nine landraces from neighbor regions and four reference cultivars. The collection showed wide phenotypic variability for several qualitative and quantitative traits, such as leaf border and shape, inflorescence type, fruit shape, green shoulder, fruit weight (range 14-277 g), locule number (2-12), shape index (0.54-2.65), yield (0.24-3.08 kg/plant), and soluble solids (3.4-7.5°B). A few landraces showed uncommon phenotypes, such as potato leaf, colorless fruit epidermis, or delayed ripening. Multivariate analysis of 25 cardinal phenotypic variables separated the accessions into two distinct groups; accessions showing a flattened-ribbed fruit were distinguished from those with round to elongate fruits with smooth structure. Genotyping analysis of 7,720 SNPs was performed using the tomato array platform SolCAP, to point out the genetic relationship among the studied accessions. A neighbor-joining tree analysis allowed to confirm or deny phenotypic data and to assign some of the unnamed accessions to recognized groups. Allelic status at marker loci linked to resistance genes commonly used in breeding identified accessions putatively derived from modern material or commercial hybrids, thus not classifiable as landraces. Overall, this study provided the information useful to preserve, valorize, and juridically protect tomato local landraces from the Lazio region and will in addition be helpful to their improvement by breeding.
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Gene editing has already proved itself as an invaluable tool for the generation of mutants for crop breeding, yet its ultimate impact on agriculture will depend on how crops generated by gene editing technologies are regulated, and on our ability to characterize the impact of mutations on plant phenotype. A starting operational strategy for evaluating gene editing-based approaches to plant breeding might consist of assessing the effect of the induced mutations in a crop- and locus-specific manner: this involves the analysis of editing efficiency in different cultivars of a crop, the assessment of potential off-target mutations, and a phenotypic evaluation of edited lines carrying different mutated alleles. Here, we targeted the GREENFLESH (GF) locus in two tomato cultivars ('MoneyMaker' and 'San Marzano') and evaluated the efficiency, specificity and mutation patterns associated with CRISPR/Cas9 activity for this gene. The GF locus encodes a Mg-dechelatase responsible for initiating chlorophyll degradation; in gf mutants, ripe fruits accumulate both carotenoids and chlorophylls. Phenotypic evaluations were conducted on two transgene-free T2 'MoneyMaker' gf lines with different mutant alleles (a small insertion of 1 nucleotide and a larger deletion of 123 bp). Both lines, in addition to reduced chlorophyll degradation, showed a notable increase in carotenoid and tocopherol levels during fruit ripening. Infection of gf leaves and fruits with Botrytis cinerea resulted in a significant reduction of infected area and pathogen proliferation compared to the wild type (WT). Our data indicates that the CRISPR/Cas9-mediated mutation of the GF locus in tomato is efficient, specific and reproducible and that the resulting phenotype is robust and consistent with previously characterized greenflesh mutants obtained with different breeding techniques, while also shedding light on novel traits such as vitamin E overaccumulation and pathogen resistance. This makes GF an appealing target for breeding tomato cultivars with improved features for cultivation, as well as consumer appreciation and health.
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The Mediterranean basin countries are considered secondary centres of tomato diversification. However, information on phenotypic and allelic variation of local tomato materials is still limited. Here we report on the evaluation of the largest traditional tomato collection, which includes 1499 accessions from Southern Europe. Analyses of 70 traits revealed a broad range of phenotypic variability with different distributions among countries, with the culinary end use within each country being the main driver of tomato diversification. Furthermore, eight main tomato types (phenoclusters) were defined by integrating phenotypic data, country of origin, and end use. Genome-wide association study (GWAS) meta-analyses identified associations in 211 loci, 159 of which were novel. The multidimensional integration of phenoclusters and the GWAS meta-analysis identified the molecular signatures for each traditional tomato type and indicated that signatures originated from differential combinations of loci, which in some cases converged in the same tomato phenotype. Our results provide a roadmap for studying and exploiting this untapped tomato diversity.
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Anthocyanins are naturally occurring pigments ubiquitously present in plants and, as such, part of the human diet. Owing to their biological activity, anthocyanins have beneficial health effects but, unfortunately, are not present in some important crop plants, such as tomatoes. Recently, a 'purple' tomato, highly enriched with anthocyanins, was produced by the ectopic expression of two selected transcription factors from the ornamental flower snapdragon. In addition to being enriched with anthocyanin, these fruits also prolonged the life of cancer-susceptible mice, suggesting that they have additional health-promoting effects.
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Antocianinas/metabolismo , Plantas Modificadas Genéticamente/metabolismo , Solanum lycopersicum/metabolismo , Animales , Antocianinas/genética , Solanum lycopersicum/genética , Ratones , Neoplasias/dietoterapia , Plantas Modificadas Genéticamente/genéticaRESUMEN
The health benefits of tomato, a vegetable consumed daily in human diets, have received great attention in the scientific community, and a great deal of experiments have tested their utility against several diseases. Herein, we present a scientometric analysis of recent works aimed to estimate the biological effects of tomato, focusing on bibliographic metadata, type of testers, target systems, and methods of analysis. A remarkably variable array of strategies was reported, including testers obtained by standard and special tomatoes, and the use of in vitro and in vivo targets, both healthy and diseased. In vitro, 21 normal and 36 cancer human cell lines derived from 13 different organs were used. The highest cytotoxic effects were reported on cancer blood cells. In vivo, more experiments were carried out with murine than with human systems, addressing healthy individuals, as well as stressed and diseased patients. Multivariate analysis showed that publications in journals indexed in the agriculture category were associated with the use of fresh tomatoes; conversely, medicine and pharmacology journals were associated with the use of purified and formulate testers. Studies conducted in the United States of America preferentially adopted in vivo systems and formulates, combined with blood and tissue analysis. Researchers in Italy, China, India, and Great Britain mostly carried out in vitro research using fresh tomatoes. Gene expression and proteomic analyses were associated with China and India. The emerging scenario evidences the somewhat dichotomic approaches of plant geneticists and agronomists and that of cell biologists and medicine researchers. A higher integration between these two scientific communities would be desirable to foster the assessment of the benefits of tomatoes to human health.
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San Marzano (SM) is a traditional Italian landrace characterized by red elongated fruits, originating in the province of Naples (Italy) and cultivated worldwide. Three mutations, yellow flesh (r), green flesh (gf) and colorless fruit epidermis (y) were introduced into SM by backcross and the resulting introgression lines (ILs) produced the expected yellow, brown and pink fruit variants. In addition, ILs carrying double combinations of those mutations were obtained. The six ILs plus the SM reference were analyzed for volatile (VOC), non-polar (NP) and polar (P) metabolites. Sixty-eight VOCs were identified, and several differences evidenced in the ILs; overall gf showed epistasis over r and y and r over y. Analysis of the NP component identified 54 metabolites; variation in early carotenoids (up to lycopene) and chlorophylls characterized respectively the ILs containing r and gf. In addition, compounds belonging to the quinone and xanthophyll classes were present in genotypes carrying the r mutation at levels higher than SM. Finally, the analysis of 129 P metabolites evidenced different levels of vitamins, amino acids, lipids and phenylpropanoids in the ILs. A correlation network approach was used to investigate metabolite-metabolite relationships in the mutant lines. Altogether these differences potentially modified the hedonistic and nutritional value of the berry. In summary, single and combined mutations in gf, r and y generated interesting visual and compositional diversity in the SM landrace, while maintaining its original typology.
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Tomato (Solanum lycopersicum L.) is one of the most cultivated vegetable in the world and it represents a large source of bioactive compounds, including carotenoids and polyphenols (phenolic acids and flavonoids). However, the concentration of flavonoids in tomato is considered sub-optimal, particularly because anthocyanins are not generally present. Therefore, this crop has been the object of an intense metabolic engineering in order to obtain anthocyanin-enriched tomatoes by using either breeding or transgenic strategies. Some wild tomato species, such as S. chilense and S. cheesmaniae, biosynthesize anthocyanins in the fruit sub-epidermal tissue, and some alleles from those genotypes have been introgressed into a new developed purple tomato line, called "Sun Black" (SB). It is a tomato line with a purple skin color, both in green and in red fruit stages, due to the biosynthesis of anthocyanins in the peel, and a normal red color pulp, with a taste just like a traditional tomato. SB is the result of a breeding programme and it is not a genetically modified (GM) product. We report the chemical characterization and structure elucidation of the attractive anthocyanins found in the peel of SB tomato, as well as other bioactive compounds (carotenoids, polyphenols, vitamin C) of the whole fruit. Using one- and two-dimensional NMR experiments, the two main anthocyanins were identified to be petunidin 3-O-[6â³-O-(4â´-O-E-p-coumaroyl-α-rhamnopyranosyl) -ß-glucopyranoside]-5-O-ß-glucopyranoside (petanin) and malvidin 3-O-[6â³-O-(4â´-O-E-p-coumaroyl-α-rhamnopyranosyl)-ß-glucopyranoside]-5-O-ß-glucopyranoside (negretein). The total anthocyanins in the whole ripe fruit was 1.2 mg/g dry weight (DW); 7.1 mg/100 g fresh weight (FW). Chlorogenic acid (the most abundant phenolic acid) was 0.6 mg/g DW; 3.7 mg/100 g FW. The main flavonol, rutin was 0.8 mg/g DW; 5 mg/100 g FW. The total carotenoid content was 211.3 µg/g DW; 1,268 µg/100 g FW. The total phenolic content was 8.6 mg/g DW; 52.2 mg/100 g FW. The vitamin C content was 37.3 mg/100 g FW. The antioxidant activities as measured by the TEAC and ORAC assays were 31.6 and 140.3 µmol TE/g DW, respectively (193 and 855.8 µmol TE/100 g FW, respectively). The results show the unique features of this new tomato genotype with nutraceutical properties.
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The development of the ovary into a fruit depends on pollination and fertilization. It has been proposed that the restriction of ovary growth before pollination is because of the stamens acting as negative regulators. Accordingly, the silencing of genes responsible for stamen identity has been correlated with parthenocarpy in different species. The tomato (Solanum lycopersicum L.) parthenocarpic fruit (pat) mutation associates autonomous ovary development with homeotic transformation of the anthers and aberrancy of ovules in the ovary. In this study, we tested the hypothesis that stamen aberrations and parthenocarpy in pat are driven by cues coming from the altered expression of class B MADS box genes. The data showed that the Pat locus is not allelic to either of the two tomato mutations putatively involved in the B function, stamenless (sl)-2 and pistillate (pi) or to genes encoding class B transcription factors. Whereas pat pi double mutants were not recovered because of tight linkage, pat sl-2 double mutants showed mainly epistatic effects. The developmental regulation of the Sl DEFICIENS (DEF) gene in the wild-type (WT) at anthesis as well as its differential transcription in the pat ovary suggest that it plays a role in the control of ovary growth. Accordingly, when compared with the WT, the gene was also differentially expressed in the parthenocarpic fruit-2 (pat-2) mutant, that is not allelic to pat and has normal ovule development. Altogether the results indicate that in tomato SlDEF plays a role in the control of ovary growth and that the pat mutation is located upstream of this regulatory cascade.
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Proteína DEFICIENS/genética , Genes de Plantas , Mutación , Solanum lycopersicum/genética , Solanum lycopersicum/crecimiento & desarrolloRESUMEN
Parthenocarpy in a broad sense includes those processes that allow the production of seedless fruits. Such fruits are favorable to growers, because they are set independently of successful pollination, and to processors and consumers, because they are easier to deal with and to eat. Seedless fruits however represent a biological paradox because they do not contribute to offspring production. In this work, the occurrence of parthenocarpy in Angiosperms was investigated by conducting a bibliographic survey. We distinguished monospermic (single seeded) from plurispermic (multiseeded) species and wild from cultivated taxa. Out of 96 seedless taxa, 66% belonged to plurispermic species. Of these, cultivated species were represented six times higher than wild species, suggesting a selective pressure for parthenocarpy during domestication and breeding. In monospermic taxa, wild and cultivated species were similarly represented. The occurrence of parthenocarpy in wild species suggests that seedlessness may have an adaptive role. In monospermic species, seedless fruits are proposed to reduce seed predation through deceptive mechanisms. In plurispermic fruit species, parthenocarpy may exert an adaptive advantage under suboptimal pollination regimes, when too few embryos are formed to support fruit growth. In this situation, parthenocarpy offers the opportunity to accomplish the production and dispersal of few seeds, thus representing a selective advantage. Approximately 20 sources of seedlessness have been described in tomato. Excluding the EMS induced mutation parthenocarpic fruit (pat), the parthenocarpic phenotype always emerged in biparental populations derived from wide crosses between cultivated tomato and wild relatives. Following a theory postulated for apomictic species, we argument that wide hybridization could also be the force driving parthenocarpy, following the disruption of synchrony in time and space of reproductive developmental events, from sporogenesis to fruit development. The high occurrence of polyploidy among parthenocarpic species supported this suggestion. Other commonalities between apomixis and parthenocarpy emerged from genetic and molecular studies of the two phenomena. Such insights may improve the understanding of the mechanisms underlying these two reproductive variants of great importance to modern breeding.
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The 7B-1 tomato (Solanum lycopersicum L. cv Rutgers) is a male-sterile mutant with enhanced tolerance to abiotic stress, which makes it a potential candidate for hybrid seed breeding and stress engineering. To underline the molecular mechanism regulating the male-sterility in 7B-1, transcriptomic profiles of the 7B-1 male-sterile and wild type (WT) anthers were studied using mRNA sequencing (RNA-Seq). In total, 768 differentially expressed genes (DEGs) were identified, including 132 up-regulated and 636 down-regulated transcripts. Gene ontology (GO) enrichment analysis of DEGs suggested a general impact of the 7B-1 mutation on metabolic processes, such as proteolysis and carbohydrate catabolic process. Sixteen candidates with key roles in regulation of anther development were subjected to further analysis using qRT-PCR and in situ hybridization. Cytological studies showed several defects associated with anther development in the 7B-1 mutant, including unsynchronized anther maturation, dysfunctional meiosis, arrested microspores, defect in callose degradation and abnormal tapetum development. TUNEL assay showed a defect in programmed cell death (PCD) of tapetal cells in 7B-1 anthers. The present study provides insights into the transcriptome of the 7B-1 mutant. We identified several genes with altered expression level in 7B-1 (including beta-1,3 glucanase, GA2oxs, cystatin, cysteine protease, pectinesterase, TA29, and actin) that could potentially regulate anther developmental processes, such as meiosis, tapetum development, and cell-wall formation/degradation.