Silvana Grandillo

Fleshy Fruit Expansion and Ripening Are Regulated by the Tomato SHATTERPROOF Gene TAGL1

The maturation and ripening of fleshy fruits is a developmental program that synchronizes seed maturation with metabolism, rendering fruit tissues desirable to seed dispersing organisms. Through RNA interference repression, we show that Tomato AGAMOUS-LIKE1 (TAGL1), the tomato (Solanum lycopersicum) ortholog of the duplicated SHATTERPROOF (SHP) MADS box genes of Arabidopsis thaliana, is necessary for fruit ripening. Tomato plants with reduced TAGL1 mRNA produced yellow-orange fruit with reduced carotenoids and thin pericarps. These fruit are also decreased in ethylene, indicating a comprehensive inhibition of maturation mediated through reduced ACC Synthase 2 expression. Furthermore, ectopic expression of TAGL1 in tomato resulted in expansion of sepals and accumulation of lycopene, supporting the role of TAGL1 in ripening. In Arabidopsis, the duplicate SHP1 and SHP2 MADS box genes regulate the development of separation layers essential for pod shatter. Expression of TAGL1 in Arabidopsis failed to completely rescue the shp1 shp2 mutant phenotypes, indicating that TAGL1 has evolved distinct molecular functions compared with its Arabidopsis counterparts. These analyses demonstrate that TAGL1 plays an important role in regulating both fleshy fruit expansion and the ripening process that together are necessary to promote seed dispersal of fleshy fruit. From this broad perspective, SHP1/2 and TAGL1, while distinct in molecular function, regulate similar activities via their necessity for seed dispersal in Arabidopsis and tomato, respectively.

Biochemical and Molecular Analysis of Pink Tomatoes: Deregulated Expression of the Gene Encoding Transcription Factor SlMYB12 Leads to Pink Tomato Fruit Color

The color of tomato fruit is mainly determined by carotenoids and flavonoids. Phenotypic analysis of an introgression line (IL) population derived from a cross between Solanum lycopersicum ‘Moneyberg’ and the wild species Solanum chmielewskii revealed three ILs with a pink fruit color. These lines had a homozygous S. chmielewskii introgression on the short arm of chromosome 1, consistent with the position of the y (yellow) mutation known to result in colorless epidermis, and hence pink-colored fruit, when combined with a red flesh. Metabolic analysis showed that pink fruit lack the ripening-dependent accumulation of the yellow-colored flavonoid naringenin chalcone in the fruit peel, while carotenoid levels are not affected. The expression of all genes encoding biosynthetic enzymes involved in the production of the flavonol rutin from naringenin chalcone was down-regulated in pink fruit, suggesting that the candidate gene underlying the pink phenotype encodes a regulatory protein such as a transcription factor rather than a biosynthetic enzyme. Of 26 MYB and basic helix-loop-helix transcription factors putatively involved in regulating transcription of genes in the phenylpropanoid and/or flavonoid pathway, only the expression level of the MYB12 gene correlated well with the decrease in the expression of structural flavonoid genes in peel samples of pink- and red-fruited genotypes during ripening. Genetic mapping and segregation analysis showed that MYB12 is located on chromosome 1 and segregates perfectly with the characteristic pink fruit color. Virus-induced gene silencing of SlMYB12 resulted in a decrease in the accumulation of naringenin chalcone, a phenotype consistent with the pink-colored tomato fruit of IL1b. In conclusion, biochemical and molecular data, gene mapping, segregation analysis, and virus-induced gene silencing experiments demonstrate that the MYB12 transcription factor plays an important role in regulating the flavonoid pathway in tomato fruit and suggest strongly that SlMYB12 is a likely candidate for the y mutation.

Genetic and physiological analysis of tomato fruit weight and composition: influence of carbon availability on QTL detection

Throughout tomato domestication, a large increase in fruit size was associated with a loss of dry matter and sugar contents. This study aims to dissect the contributions of genetic variation and the physiological processes underlying the relationships between fruit growth and the accumulation of dry matter and sugars. Fruit quality traits and physiological parameters were measured on 20 introgression lines derived from the introgression of Solanum chmielewskii into S. lycopersicum, under high (HL, unpruned trusses) and low (LL, trusses pruned to one fruit) fruit load conditions. Inter- and intra-genotypic correlations among traits were estimated and quantitative trait loci (QTL) for size, composition, and physiological traits were mapped. LL increased almost all traits, but the response of sugar content was genotype-dependent, involving either dilution effects or differences in carbon allocation to sugars. Genotype×fruit load interactions were significant for most traits and only 30% of the QTL were stable under both fruit loads. Many QTL for fresh weight and cell or seed numbers co-localized. Eleven clusters of QTL for fresh weight and dry matter or sugar content were detected, eight with opposite allele effects and three with negative effects. Two genotypic antagonistic relationships, between fresh weight and dry matter content and between cell number and cell size, were significant only under HL; the second could be interpreted as a competition for carbohydrates among cells. The role of cuticular conductance, fruit transpiration or cracking in the relationship between fruit fresh weight and composition was also emphasized at the genetic and physiological levels.

Structural and Functional Genomics of Tomato

Tomato (Solanum lycopersicum L.) is the most intensively investigated Solanaceous species both in genetic and genomics studies. It is a diploid species with a haploid set of 12 chromosomes and a small genome (950 Mb). Based on the detailed knowledge on tomato structural genomics, the sequencing of the euchromatic regions started in the year 2005 as a common effort of different countries. The manuscript focuses on markers used for tomato, on mapping efforts mainly based on exploitation of natural biodiversity, and it gives an updated report on the international sequencing activities. The principal tools developed to explore the function of tomato genes are also summarized, including mutagenesis, genetic transformation, and transcriptome analysis. The current progress in bioinformatic strategies available to manage the overwhelming amount of data generated from different tomato “omics” approaches is reported, and emphasis is given to the effort of producing a computational workbench for the analysis of the organization, as well as the functionality and evolution of the Solanaceae family.

Resolution by recombination: breaking up Solanum pennellii introgressions

Quantitative trait locus (QTL) genetics retains an important role in the study of biological and agronomic processes; however, its genetic resolution is often comparatively low. Community-based strategies are thus required to address this issue. Here we detail such a strategy wherein the widely used Solanum pennellii introgression lines (ILs) in the genetic background of the cultivated tomato (Solanum lycopersicum) are broken up into molecular marker-defined sublines as a community resource for map-based cloning.

Solanum sect. Lycopersicon

In this review, we examine the plant group Solanum sect. Lycopersicon – a clade of 13 species, including the domesticated tomato (Solanum lycopersicum L.) and its wild relatives – along with four allied species in the immediate outgroups Solanum sects. Juglandifolia and Lycopersicoides. We summarize the geographic distribution and morphological characters of these plant groups, describing their evolutionary relationships in the context of a new taxonomic revision at the species level of all these groups. We provide an overview of the role that wild tomato species have played in the development of cytogenetic stocks, in classical and molecular genetic studies as well as in crop improvement through traditional and advanced tools. We discuss how the very narrow genetic basis of cultivated tomato germplasm has forced tomato geneticists and breeders to rely on the wealth of genetic variation present in the wild relatives to address the many breeding challenges. The numerous molecular mapping studies conducted using interspecific crosses have clearly demonstrated that the breeding value of exotic (wild) tomato germplasm goes far beyond its phenotype. These studies also show that we are still far from being able to fully exploit the breeding potential of the thousands of accessions stored in seed banks around the world, in addition to those that may still be found in natural habitats. Over the past decades, tomato breeders have been at the forefront of establishing new principles for crop breeding based on the use of wild species to improve modern cultivars. In this respect, among all model systems, the wild and domesticated species of the tomato clade have pioneered development of novel populations such as “exotic libraries.” These genetic resources, combined with the increasing knowledge deriving from the many “omics” tools, including the tomato genome sequence, are expected to further improve the efficiency with which wild tomato relatives will contribute to the improvement of this important crop.

Exploitation of Natural Biodiversity Through Genomics

The genetic improvement of crop plants is the most viable approach to meeting the increasing demand for agricultural output. This goal may be achieved by using the wealth of genetic variation provided by nature. Until now, scientists have been unable to exploit the genetic potential warehoused in plant germplasm repositories for quantitative traits associated with agricultural yield. Here we review the development and application of the advanced-backcross and introgression-line breeding populations for the identification of wild species derived chromosome segments that improve agricultural performance of elite germplasm. The results of studies in a wide range of crops indicate that, unlike their domestic relatives, which are often depleted in genetic variation, wild populations of plants carry a tremendous wealth of potentially valuable alleles, many of which would not have been predicted from the phenotype of the wild plants. The results from these studies may help open up new sources of genetic variation for plant breeding and biotechnology and shed light on the nature of quantitative trait variation.

Identification, introgression, and validation of fruit volatile QTLs from a red-fruited wild tomato species.

Highlight Over 100 fruit volatile QTLs were identified in a RIL population derived from the red-fruited wild species Solanum pimpinellifolium and a fresh market tomato variety ‘Moneymaker’ and subsequently confirmed in introgression lines.

Identification of Loci Affecting Accumulation of Secondary Metabolites in Tomato Fruit of a Solanum lycopersicum × Solanum chmielewskii Introgression Line Population

Semi-polar metabolites such as flavonoids, phenolic acids, and alkaloids are very important health-related compounds in tomato. As a first step to identify genes responsible for the synthesis of semi-polar metabolites, quantitative trait loci (QTLs) that influence the semi-polar metabolite content in red-ripe tomato fruit were identified, by characterizing fruits of a population of introgression lines (ILs) derived from a cross between the cultivated tomato Solanum lycopersicum and the wild species Solanum chmielewskii. By analyzing fruits of plants grown at two different locations, we were able to identify robust metabolite QTLs for changes in phenylpropanoid glycoconjugation on chromosome 9, for accumulation of flavonol glycosides on chromosome 5, and for alkaloids on chromosome 7. To further characterize the QTLs we used a combination of genome sequencing, transcriptomics and targeted metabolomics to identify candidate key genes underlying the observed metabolic variation.

Whole-genome re-sequencing of two Italian tomato landraces reveals sequence variations in genes associated with stress tolerance, fruit quality and long shelf-life traits

Abstract Tomato is a high value crop and the primary model for fleshy fruit development and ripening. Breeding priorities include increased fruit quality, shelf life and tolerance to stresses. To contribute towards this goal, we re-sequenced the genomes of Corbarino (COR) and Lucariello (LUC) landraces, which both possess the traits of plant adaptation to water deficit, prolonged fruit shelf-life and good fruit quality. Through the newly developed pipeline Reconstructor, we generated the genome sequences of COR and LUC using datasets of 65.8 M and 56.4 M of 30–150 bp paired-end reads, respectively. New contigs including reads that could not be mapped to the tomato reference genome were assembled, and a total of 43, 054 and 44, 579 gene loci were annotated in COR and LUC. Both genomes showed novel regions with similarity to Solanum pimpinellifolium and Solanum pennellii. In addition to small deletions and insertions, 2, 000 and 1, 700 single nucleotide polymorphisms (SNPs) could exert potentially disruptive effects on 1, 371 and 1, 201 genes in COR and LUC, respectively. A detailed survey of the SNPs occurring in fruit quality, shelf life and stress tolerance related-genes identified several candidates of potential relevance. Variations in ethylene response components may concur in determining peculiar phenotypes of COR and LUC.