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Genomics and metabolomics converge for tomato flavor improvement Presented by: J. Erron Haggard To: HRT 221 April 28, 2008 Flavor in tomato results from the combination of sugars (glucose and fructose) and acids (citric and malic) perceived by receptors in the tongue and ~400 volatile compounds perceived by olfactory receptors ~30 of these volatiles contribute significantly , some positively, some negatively Many of these compounds are derived from amino acids, lipids, and carotenoids Many of the synthesis pathways are known, but some are mysteries, with no genes identified for synthesis or accumulation. In this study, the authors attempt to identify loci affecting the composition of chemicals related to flavor in a genetically diverse, but well-defined introgression line population: 75 lines, each with a single introgression from Solanum pennellii in an otherwise fixed S. esculentum background. Obviously, variation does exist. Nine lines exhibited significant season x line interactions (i.e. were higher or lower in spring than in autumn) Overall, volatiles were generally lower in autumn than in spring, with a few exceptions 107 bins Additional points: • In most cases, except for some apocarotenoids, the introgressions produced more of the metabolite than M82 • Correlations between loci controlling certain compounds are indicative of a putative pathway • While citrate-associated loci were identified, no loci were consistently significantly associated with malate concentration • The precursor-product relationship in carotenoidapocarotenoid biosynthesis was confirmed by these data GC/MS used to quantify 74 metabolites Identified 889 singlemetabolite QTL, but cannot establish causality Figure 1: Heat map of metabolite profiles by introgression line Red = increase Blue = decrease Purple = increase in one harvest and decrease in another (9.9% of results) Whole-plant phenotypic traits and phosphorylated intermediates Figure 2: Cartographic representation of the combined metabolic and morphological network of the tomato Amino Acids Figure 2: Cartographic representation of the combined metabolic and morphological network of the tomato Sugars and organic acids Figure 2: Cartographic representation of the combined metabolic and morphological network of the tomato Additional points: • Harvest index had the greatest number of significant associations (n=20), and a high degree of connectivity to the other modules (n=7) • Only Brix showed a similar pattern of relation outside its module, all others maintained their connections within their module • Most traits defined as “whole plant phenotype associated” belong to central metabolic pathways and seem to be more stable • It seems that the traditional breeding goal, Brix, is negatively correlated with the assimilation of photosynthate toward nutritional compounds, such as vitamins Figure 3: Morphologically associated and independent metabolites Red = correlated significantly to at least one true morphological trait (p < 0.005) Orange = correlated significantly to at least one true morphological trait (p < 0.05) Gray = no significant correlation Pale = not tested Figure 4: Genomic regions containing QTL a – malate is morphologically independent (fig. 3) – recall the previous authors’ failure to locate malate QTL b – IL6-3 -> 50% reduction in harvest index Concluding Thoughts The utility of the IL approach lies in negating all epistatic effects to elucidate only additive gene action. Will this approach necessarily bring about the “best” tomato? Is there some benefit to epistasis and heterosis with regard to breeding for tomato flavor? This approach also removes the aspect of human preference. It provides mechanisms, but doesn’t address desires. How much malate/α-tocopherol/2-methylbutanal do you like in your tomatoes? Concluding Thoughts What is the real benefit of these QTL? There exist a plethora of QTL mapping papers. Wouldn’t it be more interesting to see these results applied in a MAS program to produce an actual crop with some improvement? Until they are applied to some advantage, QTL remain a statistical fiction. References: Giovannoni, JJ (2006) Breeding new life into plant metabolism. Nature Biotech 24:418-9 Schauer et al. (2006) Comprehensive metabolic profiling and phenotyping of interspecific introgression lines for tomato improvement Nature Biotech 24:447-54 Tieman et al. (2006) Identification of loci affecting flavour volatiles emissions in tomato fruits. J. Exp Bot 57:887896