Figures (5)  Tables (2)
    • Figure 1. 

      The pathway of carotenoid biosynthesis and its transcriptional regulation. The pathway in the rectangle above represents the production of substrates for carotenoid synthesis through the MEP pathway. The pathway in the lower ellipsis represents the synthesis and decomposition of carotenoids in plastids. Enzymes are shown in blue. Blue arrows denote positive regulation, whereas red arrows denote negative regulation. DXS, 1-deoxy-D-xylulose-5-phosphate synthase; GGPP, geranylgeranyl pyrophosphate; DXR, 1-deoxy-d-xylulose 5-phosphate reductoisomerase; MCT, 2-C-methyl-d-erythritol 4-phosphate cytidylyltransferase; CMK, 4-(cytidine 5′-diphospho)-2-C-methyl-d-erythritol kinase; MDS, 2-C-methyl-d-erythritol 2,4-cyclodiphosphate synthase; HDS, (E)-4-hydroxy-3-methylbut-2-enyl diphosphate synthase; HDR, (E)-4-hydroxy-3-methylbut-2-enyl diphosphate reductase; IDI, isopentenyl diphosphate isomerase; GPPS, geranyl diphosphate synthase; PSY, phytoene synthase; PDS, phytoene desaturase; ZDS, ζ-carotene desaturase gene; ZISO, ζ-carotene isomerase; CrtISO, carotene isomerase; LCY, lycopene beta-cyclase; HYDB, β-carotene hydroxylase; CYP97C, cytochrome P450-type monooxygenase 97C; VDE, violaxanthin de-epoxidase; ZEP, zeaxanthin epoxidase; NXS, neoxanthin synthase.

    • Figure 2. 

      Overview of the synthesis and metabolism framework of polyphenols in tomato. Enzymes are shown in red. Different colored rectangles represent the synthesis of different polyphenols. PAL, phenylalanine ammonia-lyase; C4H, cinnamate-4-hydroxylase; C3H, coumarate 3-hydroxylase; 4CL, 4-coumarate CoA ligase; COMT, catechol-Omethyl transferase; CHS, chalcone synthase; CHI, chalcone isomerase; F3H, flavanone 3-hydroxylase; DFR, dihydroflavonol reductase; ANS, anthocyanidin synthase; FOMT, flavonoid O-methyltransferase; F3'5'H, flavonoid-3,5-hydroxylase; A3GlcT, anthocyanin-3–O-glucosyltransferase; A3G6''RhaT, anthocyanin-3–O-glucoside-6′′–O-rhamnosyltransferase; ACT, Acylase; HCT, hydroxycinnamoyl-CoA shikimate/quinate hydroxycinnamoyltransferase; HCGQT, hydroxycinnamoyl glucose quinate hydroxycinnamoyl transferase; SGT, Solanidine: UDP-glucosyltransferase; HQT, hydroxycinnamoyl-Co A quinate hydroxycinamoyl transferase; F3'H, flavonoid 3-Hydroxylase; FLS, flavonol synthase.

    • Figure 3. 

      Biosynthetic pathways of L-ascorbic acid in plants. PMI, mannose-6-phosphate isomerase; PMM, phosphomannomutase; GMP, GDP-mannose pyrophosphorylase (mannose-1-phosphate guanylyltransferase); GME, GDP-mannose-3′,5′-epimerase; GGP, GDP-L-galactose phosphorylase; L-GalDH, L-galactose dehydrogenase; L-GalLDH, L-galactono-1,4-lactone to produce ascorbate; AO, ascorbate oxidase; APX, ascorbate peroxidase; DHAR, dehydroascorbate reductase; MDHAR, monodehydroascorbate reductase; GPP, L-galactose-1-phosphate phosphatase.

    • Figure 4. 

      Important intermediates in plant sugar metabolism. RuBP, ribulose-1,5-bisphosphate; 1,3-BPG, 1,3-bisphosphoglycerate; GAP, glyceraldehyde3-phosphate; Ru5P, Ribulose 5-phosphate; PRK, phosphoribulokinase; Glu, glucose; Fru, fructose; SAI, soluble acid invertase; NI, neutral invertase; CWI, cellwall invertase; G6P, glucose-6-phosphate; SUS, sucrose synthase; G1P, glucose-1-phosphate; F6P, fructose-6-phosphate; F1,6BP, fructose-1,6-bisphosphate; 6PG, 6-phosphogluconate; G6PH, glucose 6-phosphatedehydrogenase; 6PGH, 6-phosphogluconate dehydrogenase; PGM, Phosphoglucomutase; ADP-Glu, ADP-glucose; UDP-Glu, UDP-glucose; SS, Starch synthase; SPS, sucrose phosphate synthase; SPP, sucrose phosphate phosphatase;

    • Figure 5. 

      Citric acid and malic acid metabolic pathways in fruit. TCA, tricarboxylic acid cycle; ACO, aconitase; ATP-CL, ATP-citrate lyase; CS, citrate synthase; ICL, isocitrate lyase; MS, malate synthase; NAD-MDH, NAD-malate; dehydrogenase; NAD-ME, NAD-malic enzyme; NAD-IDH, NAD-isocitrate dehydrogenase; NADP-ME, NADP-malic enzyme; OAA, oxaloacetate; NADP-IDH, NADP-isocitrate dehydrogenase; PDH, pyruvate dehydrogenase; PEPC, phosphoenolpyruvate carboxylase; PEPCK, phosphoenolpyruvate carboxykinase; PPDK, pyruvate orthophosphate dikinase.

    • Precursor Volatile compound Genes
      (abbreviation)
      Full nameReference
      Fatty acidHexanalTomloxC[72], [85]
      Trans-2-Heptenal
      1-Penten-3-ol
      Hexanol
      Nonylaldehyde
      Heptaldehyde
      cis-3-Hexenol
      l-Penten-3-one
      Lsobutyl cyanideSlBCAT1Tomato branched-chain amino acid aminotransferase[86]
      Trans-2-PentenalSlscADH1tomato short chain dehydrogenase-reductase[87]
      3-Methylbutanol
      cis-3-hexenylacetate
      Hexylacetate
      Pentanol
      Isoleucine/Leucine2/3-MethylbutanalSlscADH1Tomato short chain dehydrogenase-reductase[87]
      Trans-2-HexenalLeHPLHydroperoxidelyase[88]
      PhenylalanineMethyl salicylateSlSAMTtomato salicylic acid methyl transferase[75]
      PhenylacetaldehydeLeAADCAromatic amino acid decarboxylase[54]
      2-Phenylethanol
      l-nitro-2-Phenylethane
      Benzylcyanide
      Benzaldehyde
      MethionineEugenol
      Leucine2-Isobutylthiazole
      lycopeneβ-CyclocitralLeCCD4Carotenoid cleavage dioxygenases[31]
      β-caroteneβ-ionone
      CarotenoidPseudoiononeLeCCD1BCarotenoid cleavage dioxygenases[31]
      ζ-carotenoGeranylacetone
      Carotenoidβ-Damasccnonc
      Lycopene6-Methyl-5-hepten-2-one
      CarotenoidEpoxy-B-ionone
      GeranialSlADH2Tomato alcohol dehydrogenases[89]
      Neral
      Salicylaldehyde
      Trans-3-hexen-1-olMiADH1Mango alcohol dehydrogenase[90]
      MonoterpenoidsLinaloolSlGPPS, LeMTS1Geranyl diphosphate synthases, linalool synthase[91]

      Table 1. 

      List of selected volatile compounds related to the flavor of tomatoes and the genes that regulate their accumulation.

    • CategoriesGenes
      (abbreviation)
      Full nameFunctionReference
      Soluble sugarSlCIF1/2Tomato cell wall invertase inhibitor geneIt can inhibit the activity of CWIN at the post-translational level[60]
      INH1Invertase inhibitor geneIt can inhibit the activity of CWIN at the post-translational level
      SlHSP17.7Tomato heat shock proteins encoding geneIt can increase the accumulation of sugar[48]
      SlVPE1/2/3/4/5Tomato vacuolar processing enzyme encoding geneIt can promote acid invertases synthesis and improve hexose level[76]
      SlPEPCKTomato phosphoenolpyruvate carboxykinase encoding geneIt can regulate gluconeogenesis and enchance high sugar/acid ratio[77]
      LeLINFruit apoplastic invertase encoding geneIncrease the activity of invertases[78]
      SlARF4Abaxial encoding genesInhibite the activity of AGPase and decreasing starch levels[79]
      TFT1, TFT1014-3-3 protein encoding geneDown-regulated sucrose phosphate synthase activity[80]
      SlHXK1Hexokinase encoding geneIt can inhibit the accumulate of starch in tomato leaves[81]
      LeHT1/2/3Tomato hexose transporter encoding geneEncode the hexose transporters and concentrate hexoses in storage parenchyma cells[82]
      SlSUT1Tomato sucrose transporter encoding genePhloem loader of sucrose in Solanaceae/
      RINRipening inhibitorRegulate sugar metabolism[61]
      SlMBP22MADS-box subfamily, Bsister (Bs) genesIt positively modulates the levels of starch and soluble sugar in tomato leaves[81]
      SlVIFVacuolar invertase inhibitor geneIt can inhibit the activity of VIN at the post-translational level[61]
      Organic acidsSlPEPC2Allosteric enzyme phosphoenolpyruvate carboxylase encoding genePromote the decomposition of malic acid[83]
      TRXL1Thioredoxin-like 1 encoding geneActivate NADP-MDH and increases malate[8]
      IDH3Isocitrate dehydrogenases encoding genePromote citric acid degradation[66]
      AST2Aspartate aminotransferase encoding gene
      LeACS2Ethylene biosynthesis geneEnhance the metabolization of malic and citric acid
      HMGCR13-hydroxy-3-methylglutaryl-CoA reductase encoding geneInhibit the accumulation of critic acid
      MDHMalate dehydrogenase encoding geneIncrease the content of malic acid
      PDC3phosducin-like 3 encoding geneDecrease the content of malic acid
      SlAREB1ABA-response element binding factorMediate ABA signal and increase citric acid and malic acid level[84]
      AP2/ERFAPETALA2/Ethylene Response FactorRegulate the organic acid levels[66]
      SlWRKYsEthylene-responsive genes
      SNACStress-responsive NAC transcription factors

      Table 2. 

      Overview of the genes that affect the accumulation of soluble sugar and organic acids in tomatoes.