
NSF Org: |
IOS Division Of Integrative Organismal Systems |
Recipient: |
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Initial Amendment Date: | December 15, 2015 |
Latest Amendment Date: | September 4, 2018 |
Award Number: | 1564366 |
Award Instrument: | Continuing Grant |
Program Manager: |
Diane Jofuku Okamuro
dokamuro@nsf.gov (703)292-4508 IOS Division Of Integrative Organismal Systems BIO Directorate for Biological Sciences |
Start Date: | September 16, 2015 |
End Date: | April 30, 2022 (Estimated) |
Total Intended Award Amount: | $4,993,470.00 |
Total Awarded Amount to Date: | $5,018,470.00 |
Funds Obligated to Date: |
FY 2016 = $1,137,201.00 FY 2017 = $2,378,545.00 FY 2018 = $947,908.00 |
History of Investigator: |
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Recipient Sponsored Research Office: |
310 E CAMPUS RD RM 409 ATHENS GA US 30602-1589 (706)542-5939 |
Sponsor Congressional District: |
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Primary Place of Performance: |
500 D. W. Brooks Dr. Athens GA US 30602-0001 |
Primary Place of
Performance Congressional District: |
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Unique Entity Identifier (UEI): |
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Parent UEI: |
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NSF Program(s): | Plant Genome Research Project |
Primary Program Source: |
01001617DB NSF RESEARCH & RELATED ACTIVIT 01001718DB NSF RESEARCH & RELATED ACTIVIT 01001819DB NSF RESEARCH & RELATED ACTIVIT |
Program Reference Code(s): |
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Program Element Code(s): |
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Award Agency Code: | 4900 |
Fund Agency Code: | 4900 |
Assistance Listing Number(s): | 47.074 |
ABSTRACT
PI: Esther van der Knaap (University of Georgia)
CoPIs: Ana Caicedo (University of Massachusetts); Denise Tieman (University of Florida); Lukas Mueller (Boyce Thompson Institute)
Collaborators: Joaquin Cañizares, Maria Jose Diez, Jose Blanca (Universitat Politecnica De Valencia); Sofia Visa, Dean Fraga, Simon Gray (College of Wooster)
A high quality tomato is expected to have good flavor, size, color and firmness. Years of selective breeding have produced the modern tomato, and yet the quality and production efficiency of the crop still needs improvement. Despite this need, desirable traits that will improve fruit quality are often hard to find, partly because the underlying factors that specify fruit traits are not fully known. Another impediment is that some beneficial traits were eliminated or hidden during breeding, thus reducing the available genetic diversity in modern tomato genomes. Project scientists have discovered an untapped resource to find new traits from wild tomato relatives and from ancestral populations growing in diverse geographic locations. These plants may appear undesirable by having small, seedy, soft or unpalatable fruits. However, these relatives are actually genetic gold mines: they hold within their DNA useful and untapped traits that were lost during tomato domestication. New technologies and genetic methods are now available to mine these wild tomato genomes for new suites of breeding traits. This project uses the selected wild populations to explore and capture quality traits associated with fruit weight, firmness, flavor and color. The project will identify the molecular basis of these fruit traits and will deliver new traits to improve the breeding potential of the modern tomato. The project provides interdisciplinary training in genomics, computational data analysis and breeding to all involved, including post-doctoral researchers and graduate students. Through summer workshops, high school and college students will be trained in tomato genomics and breeding, will interact directly with scientists, and will gain hands-on skills in research. These trained students will be critical additions to a workforce that advances agriculture through scientific discovery.
Wild relatives and semi-domesticated germplasm of cultivated plants provide a significant reservoir of genetic and epigenetic diversity for key regulators of agronomic traits. Future crop improvement relies on harnessing this diversity. However, mining semi-domesticated and wild germplasm for beneficial alleles of agriculturally important traits is not straightforward because fruit quality is quantitatively inherited. Consequently, visual inspection of unselected germplasm does not readily lead to the identification of accessions that have desirable characteristics to improve modern germplasm. The association of traits with genes controlling fruit quality and the identification of beneficial alleles that may have been lost during domestication should provide a model for studying how to efficiently mine germplasm of the closest wild relatives for quantitative trait loci leading to tangible crop improvement. To identify genes and pathways that control complex tomato fruit quality traits, this project will (1) assemble and phenotype a tomato population (Solanum spp.) constituting the continuum of wild, semi-domesticated and ancestral landraces; (2) identify loci underlying fruit quality traits through genome-wide association studies (GWAS) and differentially expressed small RNAs; (3) confirm genetically the traits associated with candidate regions to genes, and (4) analyze the developmental and biochemical pathways that control fruit quality. The project will result in genome sequence data for 150 tomato accessions, including 20 from the closest but fully wild relative of cultivated tomato, 110 from wild and semi-domesticated direct ancestors of domesticated tomatoes and 20 from the earliest landraces of cultivated tomatoes. In addition, the project will generate small RNA sequence data from different stages of tomato fruit development from a subset of this population, and will provide detailed fruit quality information about flavor, firmness, weight and palatability for each of the 150 accessions. The information will be available through a public resource, the Sol Genomic Network (SGN, http://www.sgn.cornell.edu/), and seeds of the accessions will be available from germplasm repository sites (TGRC, http://tgrc.ucdavis.edu/; and COMAV, http://www.comav.upv.es/).
PUBLICATIONS PRODUCED AS A RESULT OF THIS RESEARCH
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PROJECT OUTCOMES REPORT
Disclaimer
This Project Outcomes Report for the General Public is displayed verbatim as submitted by the Principal Investigator (PI) for this award. Any opinions, findings, and conclusions or recommendations expressed in this Report are those of the PI and do not necessarily reflect the views of the National Science Foundation; NSF has not approved or endorsed its content.
This project focused on the consequences of tomato domestication upon genetic diversity and fruit quality. The goals were to uncover tomato’s recent evolutionary history and to link them to characteristics that contributed to the extensive size and flavor variation among the accessions. This knowledge will provide insights into the regulation of the size and flavor in produce, while also offering tools to breeding programs. One of the most important contributions is that this population is ready to be explored for variation in many other traits such as nutrient efficiency, drought and heat tolerance, and others. Pertinent information is publicly available on the Solanaceae Genomics Network Varitome page (https://solgenomics.net/projects/varitome/), in addition to publications in scientific journals; the seeds are available through the TGRC (https://tgrc.ucdavis.edu/) and COMAV (http://www.comav.upv.es).
Discerning favorable traits in wild and semi-domesticated plants is difficult because their effects are often masked by many characteristics that are considered unfavorable in modern agriculture. However, a combination of approaches and integration of the findings have led to novel discoveries as summarized below. Contrary to expectations, our findings demonstrated that a species considered to be semi-domesticated (Solanum lycopersicum var. cerasiforme) actually diverged from a wild relative over 70,000 years ago, prior to human intervention. The location of where the direct ancestor of tomato emerged has led us to propose two possible scenarios for the migration and origin of modern tomato in Mesoamerica. Another unique finding was the epigenetic impact resulting from domestication. Research on small RNAs (sRNA) showed differential selection pressures over distinct classes, and large-scale genomic deletions and insertions as a major driving force for the dynamic expression of these sRNAs as tomato evolved. We also determined that though cultivated tomato lost genetic diversity during the domestication process, a greater proportion of mutations in cultivated than wild genomes are potentially deleterious; this mutational load can constrain adaptability of cultivated tomatoes.
Our studies also demonstrated that semi-domesticated tomatoes from Ecuador and Peru provided the largest source of variation for fruit flavor and size. With respect to fruit size, we found at least eleven new regions in the tomato genome that contribute to weight, representing novel genes. For three of them, the analyses suggested that the traits were left behind as tomato evolved into modern types, contrary to expectations. For one, the most likely gene controlling weight is of unknown function, single copy and highly conserved among other plant species. Therefore, it is plausible that this gene could affect the weight of produce in other crops. With respect to flavor, we focused on multiple pathways and regions of the genome. Many candidate genes have been identified and some have been validated. Genome analyses showed that gene duplications, large deletions or insertions are often found around the genes of interest, further solidifying that this variant type is common in crop plants. The most advanced investigations are for a methyl esterase that controls methyl salicylate levels in ripe fruits. One to four copies of methyl esterase gene(s) underlie the levels of this volatile in tomato. Gene knockouts showed that inactivation of one of them leads to higher levels of the volatile and enzyme assays supports its predicted function. The volatile is not appreciated in human consumption but required for signaling and defense responses. Therefore, a careful balance of its levels will create a tasty tomato with the desired agronomic traits.
Last Modified: 08/30/2022
Modified by: Esther Van Der Knaap
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