Late blight (LB), caused by Phytophthora infestans, is a destructive disease of the cultivated tomato, Solanum lycopersicum. Environmental concerns and pathogen resistance have propelled research towards developing host resistance. The current LB-resistant cultivars of tomato exhibit susceptibility under severe disease pressure, necessitating the identification, characterization, and incorporation of additional resistance genes into new tomato cultivars. Recently, we identified Solanum pimpinellifolium accession PI 270443 with strong resistance to LB and developed a RIL population from its cross with an LB-susceptible tomato breeding line. In the present study, we constructed a high-density genetic map of the RIL population, using 8,470 SNP markers set into 1,195 genomic bins, with a total genetic distance of 1232 cM and an average bin size of 1 cM. We identified 2 major adjoining LB-resistance QTLs on chromosome 10 and a few minor QTLs on chromosomes 1 and 12 of PI 270443. While one of the QTLs on chromosome 10 colocalized with the known LB-resistance gene Ph- 2 and a LB-resistance QTL previously identified in an F2 population of the same cross, the present study allowed marker saturation of the region, fine mapping of the QTL, and identification of candidate resistance genes in the region. One of the 2 major QTLs on chromosome 10 and the 3 QTLs on chromosomes 1 and 12 were not previously reported in S. pimpinellifolium for LB resistance. These results will expedite transferring of LB resistance from PI 270443 into the tomato cultigen via MAS and discovering the underpinning LB-resistance genes in PI 270443.
Late blight (LB), caused by the oomycete Phytophthora infestans, is one of the most devastating diseases of the cultivated tomato (Solanum lycopersicum) worldwide. Attempts to control the disease through fungicide applications are becoming less effective, as new and aggressive genotypes of the pathogen emerge. Further, some new P. infestans genotypes overcome the currently available resistance in tomato, necessitating the identification, characterization, and utilization of new sources of host resistance. In the present study, to detect QTLs underlying LB resistance in a recently-identified LB-resistant S. pimpinellifolium accession (PI 270442), an F2 population (n = 1,175) of a cross between PI 270442 and LB-susceptible tomato breeding line Fla. 8059 was screened for LB resistance and subjected to selective genotyping. A total of 19,839 single-nucleotide polymorphisms (SNPs) were identified from reduced representation libraries (RRLs) constructed from the parents, of which 212 were used to build a genetic linkage map and locate QTLs. Ten LB-resistance QTLs were identified in PI 270442 on chromosomes 1, 2, 5, 6, 10, 11 and 12, of which those on chromosomes 6, 10 and 11 were the strongest and co-localized with previously-reported LB-resistance QTLs. Genomic locations of the QTLs were compared with the tomato physical map, which resulted in the identification of several candidate genes that might be underpinning the LB resistance in PI 270442. The identified QTL-linked markers can be used in breeding programs to transfer resistance from PI 270442 into the cultivated tomato via marker-assisted breeding and to develop near-isogenic lines for fine mapping of the QTLs.
Late blight (LB), caused by oomycete Phytophthora infestans, is one of the most destructive diseases of the cultivated tomato, Solanum lycopersicum. Since new and aggressive clonal lineages of P. infestans, many of which overcoming formerly effective fungicides or host resistance genes, have continued to emerge, it is crucial to identify, characterize, and utilize new sources of host resistance in tomato breeding. A recent screening of tomato germplasm identified Solanum pimpinellifolium accession PI 224710 with very strong resistance to several current P. infestans clonal lineages. The present study aimed to identify and characterize QTLs associated with LB resistance in PI 224710. Disease screening of a large F2 population (n = 1721), derived from a cross between PI 224710 and LB-susceptible tomato breeding line Fla. 8059, followed by F3 progeny testing, resulted in the identification of 43 highly-resistant and 27 highly-susceptible F2 individuals. A selective genotyping approach, using 469 non-identical SNP markers, resulted in the construction of a genetic linkage map and identification of three LB-resistance QTLs on chromosomes 6, 9 and 10 of PI 224710. A comparison of the QTLs genomic locations with the tomato physical map resulted in the identification of several candidate genes, which might be underpinning the LB-resistance QTLs in PI 224710. The identified markers associated with the LB-resistance QTLs can be utilized in breeding programs to transfer resistance from PI 224710 into tomato breeding lines and hybrid cultivars via marker-assisted breeding; they also can be used to develop near-isogenic lines for fine mapping of the QTLs.
The cultivated tomato, Solanum lycopersicum, is highly sensitive to cold stress (CS), resulting in significant losses during cultivation and postharvest fruit storage. Previously, we demonstrated the presence of substantial genetic variation in fruit chilling tolerance in a tomato recombinant inbred line (RIL) population derived from a cross between a chilling-sensitive tomato line and a chilling-tolerant accession of the wild species S. pimpinellifolium. Here, we investigated molecular and biochemical components associated with chilling tolerance in fruit and leaves, using contrasting groups of "chilling tolerant" and "chilling sensitive" RI lines. Transcriptomic analyses were conducted on fruit exposed to CS, and gene expressions and biochemical components were measured in fruit and leaves. The analyses revealed core responding genes specific to either the cold-tolerant or cold-sensitive RI lines, which were differentially regulated in similar fashion in both leaves and fruit within each group. These genes may be used as markers to determine tomato germplasm cold tolerance or sensitivity. This study demonstrated that tomato response to CS in different developmental stages, including seedling and postharvest fruit, might be mediated by common biological/genetic factors. Therefore, genetic selection for cold tolerance during early stages of plant development may lead to lines with greater postharvest fruit chilling tolerance.
Whole-genome QTL mining and meta-analysis in tomato for resistance to bacterial and fungal diseases identified 73 meta-QTL regions with significantly refined/reduced confidence intervals. Tomato production is affected by a range of biotic stressors, causing yield losses and quality reductions. While sources of genetic resistance to many tomato diseases have been identified and characterized, stability of the resistance genes or quantitative trait loci (QTLs) across the resources has not been determined. Here, we examined 491 QTLs previously reported for resistance to tomato diseases in 40 independent studies and 54 unique mapping populations. We identified 29 meta-QTLs (MQTLs) for resistance to bacterial pathogens and 44 MQTLs for resistance to fungal pathogens, and were able to reduce the average confidence interval (CI) of the QTLs by 4.1-fold and 6.7-fold, respectively, compared to the average CI of the original QTLs. The corresponding physical length of the CIs of MQTLs ranged from 56 kb to 6.37 Mb, with a median of 921 kb, of which 27
Late blight (LB), caused by the oomycete Phytophthora infestans, is one of the most destructive diseases of the cultivated tomato (Solanum lycopersicum L.) and potato (Solanum tuberosum L.) worldwide. Genetic changes in the pathogen have resulted in the emergence of new genotypes, overcoming formerly effective fungicides or host resistance genes. We previously reported the identification of a LB-resistant accession (PI 270441) of the wild tomato species S. pimpinellifolium L. and the high heritability of its resistance. In the present study, an F-2 population (n = 1,209), derived from a cross between PI 270441 and a LB-susceptible tomato breeding line (Fla. 8059), was screened for response to LB infection. Extreme resistant (n = 44) and susceptible (n = 39) F-2 individuals were selected and used in a trait-based marker analysis (TBA; a.k.a selective genotyping) to identify and map quantitative trait loci (QTLs) conferring LB resistance. Reduced representation libraries (RRLs) of Fla. 8059 and PI 270441 were constructed, sequenced, and mapped to the tomato genome. A total of 13,054 single-nucleotide polymorphisms (SNPs) were identified, of which, 200 were used to construct a genetic linkage map and locate QTLs. Four LB resistance QTLs were identified on chromosomes 1, 10, and 11 of PI 270441. The markers associated with these QTLs can be used to transfer LB resistance from PI 270441 into new tomato cultivars and to develop near-isogenic lines for fine mapping of the QTL.
Storage at low temperatures is a common practice to prolong postharvest life of fruit and vegetables with a minimal negative impact on human/environmental health. Storage at low temperatures, however, can be restricted due to produce susceptibility to non-freezing chilling temperatures, when injuries such as physiological disorders and decays may result in unmarketable produce. We have investigated tomato fruit response to postharvest chilling stress in a recombinant inbred line (RIL) population developed from a cross between a chilling-sensitive cultivated tomato (Solanum lycopersicum L.) breeding line and a chilling-tolerant inbred accession of the tomato wild species S. pimpinellifolium L. Screening of the fruit of 148 RILs under cold storage (1.5°C) indicated presence of significant variations in chilling tolerance, manifested by varying degrees of fruit injury. Two extremely contrasting groups of RILs were identified, chilling-tolerant and chilling-sensitive RILs. The RILs in the two groups were further investigated under chilling stress conditions, and several physiological parameters, including weight loss, chlorophyll fluorescence parameters Fv/Fm, and Performance Index (PI), were determined to be efficient markers for identifying response to chilling stress in postharvest fruit. The Fv/Fm values reflected the physiological damages endured by the fruit after cold storage, and PI was a sensitive marker for early changes in photosystem II function. These two parameters were early indicators of chilling response before occurrence of visible chilling injuries. Antioxidant activities and ascorbic acid content were significantly higher in the chilling-tolerant than the chilling-sensitive lines. Further, the expression of C-repeat/DREB binding factors (CBFs) genes swiftly changed within 1-hr of fruit exposure to the chilling temperature, and the SlCBF1 transcript level was generally higher in the chilling-tolerant than chilling-sensitive lines after 2-hr exposure to the low temperature. This research demonstrates the presence of potential genetic variation in fruit chilling tolerance in the tomato RIL population. Further investigation of the RIL population is underway to better understand the genetic, physiological, and biochemical mechanisms involved in postharvest fruit chilling tolerance in tomato.
Abstract Vegetables are an integral part of the human diet worldwide. Traditional breeding approaches have been used extensively to develop new cultivars of vegetables with desirable characteristics, including resistance/tolerance to biotic and abiotic stresses, high yield, and an elevated content of compounds beneficial to human health. The technological progress since the early 1980s has revolutionized our ability to study and manipulate genetic variation in crop plants. The development of high-throughput sequencing platforms and accompanying analytical methods have led to sequencing and assembly of a large number of plant genomes, construction of dense and ultra-dense molecular linkage maps, identification of structural variants, and application of molecular markers in breeding programs. Linkage mapping and genome-wide association mapping studies have been used to identify chromosomal locations of genes and QTLs associated with plant phenotypic variations important for crop improvement. This review provides up-to-date information on the status of genomics and marker-assisted improvement of vegetable crops with the focus on tomato, pepper, eggplant, lettuce, spinach, cucumber, and chicory. For each vegetable crop, we present the most recent information on genetic resources, mapping populations, genetic maps, genome sequences, mapped genes and QTLs, the status of marker-assisted selection and genomic selection, and discuss future research prospects and application of novel techniques and approaches.
Late blight (LB), caused by Phytophthora infestans, is one of the most devastating diseases of tomato (Solanum lycopersicum) worldwide. Aggressive pathogen isolates resistant to fungicides have driven research in favour of finding new sources of host resistance for tomato breeding. Recently, we reported S. pimpinellifolium accession PI 270443 exhibiting LB resistance stronger than all commercial LB-resistant tomato cultivars. The purpose of this study was to examine the inheritance of LB resistance conferred by this accession. An interspecific cross was made between PI 270443 and a LB-susceptible tomato breeding line and advanced to F-10 generation. A total of 166 F-9 and corresponding F-10 recombinant inbred lines (RILs) were evaluated for response to LB in four replicated greenhouse experiments. Estimates of heritability (h(2)) of LB resistance, determined by parent-offspring (F-9:F-10) correlation analysis, ranged from 0.66 to 0.81, with an average of 0.76. The moderately high h(2) of LB resistance in PI 270443 suggests the utility of this accession for tomato breeding. Molecular mapping and RNA-sequencing efforts are underway to identify genes underlying LB resistance in PI 270443.
Genotyping-by-sequencing (GBS) was employed to construct a highly saturated genetic linkage map of a tomato ( L.) recombinant inbred line (RIL) population, derived from a cross between cultivar NC EBR-1 and the wild tomato L. accession LA2093. A pipeline was developed to convert single nucleotide polymorphism (SNP) data into genomic bins, which could be used for fine mapping of quantitative trait loci (QTL) and identification of candidate genes. The pipeline, implemented in a python script named SNPbinner, adopts a hidden Markov model approach for calculation of recombination breakpoints followed by genomic bins construction. The total length of the newly developed high-resolution genetic map was 1.2-fold larger than previously estimated based on restriction fragment length polymorphism (RFLP) and polymerase chain reaction (PCR)-based markers. The map was used to verify and refine QTL previously identified for two fruit quality traits in the RIL population, fruit weight (FW) and fruit lycopene content (LYC). Two well-described FW QTL ( and ) were localized precisely at their known underlying causative genes, and the QTL intervals were decreased by two- to tenfold. A major QTL for LYC content () was verified at high resolution and its underlying causative gene was determined to be ζ (). The RIL population, the high resolution genetic map, and the easy-to-use genotyping pipeline, SNPbinner, are made publicly available.
Modern tomatoes have narrow genetic diversity limiting their improvement potential. We present a tomato pan-genome constructed using genome sequences of 725 phylogenetically and geographically representative accessions, revealing 4,873 genes absent from the reference genome. Presence/absence variation analyses reveal substantial gene loss and intense negative selection of genes and promoters during tomato domestication and improvement. Lost or negatively selected genes are enriched for important traits, especially disease resistance. We identify a rare allele in the TomLoxC promoter selected against during domestication. Quantitative trait locus mapping and analysis of transgenic plants reveal a role for TomLoxC in apocarotenoid production, which contributes to desirable tomato flavor. In orange-stage fruit, accessions harboring both the rare and common TomLoxC alleles (heterozygotes) have higher TomLoxC expression than those homozygous for either and are resurgent in modern tomatoes. The tomato pan-genome adds depth and completeness to the reference genome, and is useful for future biological discovery and breeding.
Crop domestication and selective breeding have altered plant defense mechanisms, influencing insect-plant interactions. A reduction in plant resistance/tolerance against herbivory is generally expected in domesticated species, however, limited efforts have been made to compare inducibility of plant defenses between wild and domesticated genotypes. In the present study, the inducibility of several plant defense mechanisms (e.g. defensive chemicals, trichomes, plant volatiles) were investigated, and the performance and preference of the herbivore Helicoverpa zea were measured in three different tomato genotypes; a) wild tomato, Solanum pimpinellifolium L. (accession LA 2093), b) cherry tomato, S. lycopersicum L. var. cerasiforme (accession Matts Wild Cherry), and c) cultivated tomato, S. lycopersicum L. var. Better Boy). Enhanced inducibility of defensive chemicals, trichomes, and plant volatiles in the cultivated tomato, and a higher level of constitutive plant resistance against herbivory in the wild genotype was observed. When comparing the responses of damaged vs. undamaged leaves, the percent reduction in larval growth was higher on damaged leaves from cultivated tomato, suggesting a higher induced resistance compared to other two genotypes. While all tomato genotypes exhibited increased volatile organic compound (VOCs) emissions in response to herbivory, the cultivated variety responded with generally higher levels of VOCs. Differences in VOC patterns may have influenced the ovipositional preferences, as H. zea female moths significantly preferred laying eggs on the cultivated versus the wild tomato genotypes. Selection of traits during domestication and selective breeding could alter allocation of resources, where plants selected for higher yield performance would allocate resources to defense only when attacked.
Late blight (LB), caused by Phytophthora infestans, is a destructive disease of tomato (Solanum lycopersicum) worldwide. Currently, there are few commercial cultivars of tomato with resistance to LB, and the disease is mainly controlled by heavy use of fungicides. Due to the emergence of fungicide-resistant pathogen isolates, there is a concerted effort to identify new genetic sources of resistance and breed new resistant cultivars. A recent screening identified several new tomato accessions with strong resistance to LB. Here, we report on the genetic basis of LB resistance in S.pimpinellifolium accession PI 270441, as determined by generation means analysis and analysis of response to selection, using populations derived from crosses with LB-susceptible breeding line Fla. 8059. Heritability of LB resistance ranged from 0.76 to 0.78, and the minimum number of genes was estimated 1few. These results suggest that transfer of LB resistance from PI 270441 to the cultivated tomato should be feasible via a traditional backcross breeding approach. Genetic mapping studies are underway to identify molecular markers associated with resistance in this accession.
Headspace solid-phase microextraction (HS-SPME) coupled to gas chromatography–mass spectrometry (GC-MS) is widely employed for volatile analyses of plants, including mapping populations used in plant breeding research. Studies often employ a single internal surrogate standard, even when multiple analytes are measured, with the assumption that any relative changes in matrix effects among individuals would be similar for all compounds, i.e., matrix effects do not show Compound × Individual interactions. We tested this assumption using individuals from two plant populations: an interspecific grape (Vitis spp.) mapping population (n = 140) and a tomato (Solanum spp.) recombinant inbred line (RIL) population (n = 148). Individual plants from the two populations were spiked with a cocktail of internal standards (n = 6, 9, respectively) prior to HS-SPME-GC-MS. Variation in the relative responses of internal standards indicated that Compound × Individual interactions exist but were different between the two populations. For the grape population, relative responses among pairs of internal standards varied considerably among individuals, with a maximum of 249% relative standard deviation (RSD) for the pair of [U13C]hexanal and [U13C]hexanol. However, in the tomato population, relative responses of internal standard pairs varied much less, with pairwise RSDs ranging from 8% to 56%. The approach described in this paper could be used to evaluate the suitability of using surrogate standards for HS-SPME-GC-MS studies in other plant populations.
Late blight (LB), caused by the oomycete (Mont.) de Bary, is one of the most devastating diseases of tomato ( L.) and potato ( tuberosum L. worldwide. The importance of LB on tomato has increased due to the occurrence of aggressive and fungicide-resistant clonal lineages of . Consequently, identification and characterization of new sources of genetic resistance to LB has become a priority in tomato breeding. Previously, we reported accession PI 163245 as a promising source of highly heritable LB resistance for tomato breeding. The purpose of this study was to identify and map quantitative trait loci (QTLs) associated with LB resistance in this accession using a trait-based marker analysis (a.k.a. selective genotyping). An F mapping population ( = 560) derived from a cross between a LB-susceptible tomato breeding line (Fla. 8059) and PI 163245 was screened for LB resistance, and the most resistant ( = 39) and susceptible ( = 35) individuals were selected for genotyping. Sequencing and comparison of the reduced representation libraries (RRLs) derived from genomic DNA of the two parents resulted in the identification of 33,541 putative single nucleotide polymorphism (SNP) markers, of which, 233 genome-wide markers were used to genotype the 74 selected F individuals. The marker analysis resulted in the identification of four LB resistance QTLs conferred by PI 163245, located on chromosomes 2, 3, 10, and 11. Research is underway to develop near-isogenic lines (NILs) for fine mapping the QTLs and develop tomato breeding lines with LB resistance introduced from PI 163245.
Late blight (LB), caused by Phytophthora infestans, is one of the most devastating diseases of tomato (Solanum lycopersicum) worldwide. Due to the emergence of new and aggressive P.infestans isolates, identifying new genetic resistance to LB is a priority in tomato breeding. Recently, we reported the identification of several Solanum pimpinellifolium accessions with strong LB resistance. In this study, we investigated the utility of resistant accession PI 163245 for tomato breeding by examining heritability (h(2)) of resistance and the response to selection for resistance. Estimates of h(2) based on F-2:F-3 and F-3:F-4 parent:offspring correlation analyses averaged 0.79 and 0.94, respectively, suggesting the heritable nature of LB resistance in PI 163245. Analysis of response to selection for resistance from F-2 to F-4 generations indicated a realized h(2) of 0.63, confirming the utility of this resistance in tomato breeding. Two methods of estimating the minimum number of loci involved indicated the presence of one major resistance locus. Currently, genetic mapping and breeding efforts are underway to further confirm the viability of this accession for improving tomato LB resistance.
Most commercial cultivars of tomato (Solanum lycopersicum) are susceptible to early blight (EB), a devastating fungal (Alternaria solani and A. tomatophila) disease of tomato in the US and elsewhere. Common measures of disease control currently include sanitation, crop rotation and heavy use of fungicides. Use of resistant cultivars, however, is the most economically acceptable and environmentally sound approach to controlling EB disease in tomato. Sources of genetic resistance to EB have been identified within the related wild species of tomato, including S. habrochaites, S. peruvianum and S. pimpinellifolium. Characterization of genetic resistance to EB would facilitate transfer of resistance to elite breeding lines and hybrid cultivars of tomato. We used a recombinant inbred line (RIL) population of tomato, previously developed from a cross between S. pimpinellifolium accession LA 2093 and S. lycopersicum tomato breeding line NCEBR-1, to determine the inheritance of EB resistance as well as
A recombinant inbred line (RIL) population of tomato was developed from a cross between a tomato breeding line (NC 84173) and an accession (LA 0722) of the tomato wild species S. pimpinellifolium. NC 84173 is a horticulturally-superior, multiple-disease resistant inbred line that has been used as a parent in production of several commercial tomato hybrid cultivars. LA 0722 is a self-compatible, inbred accession, which was previously identified as a genetic source for fruit quality, disease resistance, and abiotic stress tolerance. The RIL population is in the F9 generation and consists of 145 lines. A genetic linkage map of the population was developed with 191 molecular markers, including 129 RFLPs and 62 RGAs. The genetic map spans 1505 cM of the 12 tomato chromosomes with an average inter-marker distance of 7.9 cM. The RFLP markers were chosen from the high-density map of tomato, previously developed based on a S. lycopersicum S. pennellii F2 population. The RGA markers were derived using degenerate oligonucleotide primers designed based on conserved leucine-rich repeat (LRR),