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식물에서 shikimate 및 방향족 아미노산 생합성 경로와 이의 대사공학적 응용

Biosynthetic pathway of shikimate and aromatic amino acid and its metabolic engineering in plants

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초록

The aromatic amino acids, which are composed of ʟ-phenylalanine, ʟ-tyrosine and ʟ-tryptophan, are general components of protein synthesis as well as precursors for a wide range of secondary metabolites. These aromatic amino acids-derived compounds play important roles as ingredients of diverse phenolics including pigments and cell walls, and hormones like auxin and salicylic acid in plants. Moreover, they also serve as the natural products of alkaloids and glucosinolates, which have a high potential to promote human health and nutrition. The biosynthetic pathways of aromatic amino acids share a chorismate, the common intermediate, which is originated from shikimate pathway. Then, tryptophan is synthesized via anthranilate and the other phenylalanine and tyrosine are synthesized via prephenate, as intermediates. This review reports recent studies about all the enzymatic steps involved in aromatic amino acid biosynthetic pathways and their gene regulation on transcriptional/posttranscriptional levels. Furthermore, results of metabolic engineering are introduced as efforts to improve the production of the aromatic amino acids-derived secondary metabolites in plants.

Abstract

The aromatic amino acids, which are composed of ʟ-phenylalanine, ʟ-tyrosine and ʟ-tryptophan, are general components of protein synthesis as well as precursors for a wide range of secondary metabolites. These aromatic amino acids-derived compounds play important roles as ingredients of diverse phenolics including pigments and cell walls, and hormones like auxin and salicylic acid in plants. Moreover, they also serve as the natural products of alkaloids and glucosinolates, which have a high potential to promote human health and nutrition. The biosynthetic pathways of aromatic amino acids share a chorismate, the common intermediate, which is originated from shikimate pathway. Then, tryptophan is synthesized via anthranilate and the other phenylalanine and tyrosine are synthesized via prephenate, as intermediates. This review reports recent studies about all the enzymatic steps involved in aromatic amino acid biosynthetic pathways and their gene regulation on transcriptional/posttranscriptional levels. Furthermore, results of metabolic engineering are introduced as efforts to improve the production of the aromatic amino acids-derived secondary metabolites in plants.

참고문헌 (141)

  1. Anton IA, "Sequencing and overexpression of the Escherichia coli aroE gene encoding shikimate dehydrogenase", Biochem J, 249, pp.319~326 (1988)
  2. Arabidopsis Genome Initiat, "Analysis of the genome sequence of the flowering plant Arabidopsis thaliana", Nature, 408, pp.796~815 (2000)
  3. Barends TRM, "Tryptophan synthase, an allosteric molecular factory", Curr Opin Chem Biol, 12, pp.593~600 (2008)
  4. Bartee L, "Two Arabidopsis methylation-deficiency mutations confer only partial effects on a methylated endogenous gene family", Nucleic Acids Res, 29, pp.2127~2134 (2001)
  5. Basset G, "Folate synthesis in plants: the p-aminobenzoate branch is initiated by a bifunctional PabA-PabB protein that is targeted to plastids", Proc Natl Acad Sci USA, 101, pp.1496~1501 (2004)
  6. Benesova M, "Chorismate mutase isoforms from seeds and seedlings of Papaver somniferum", Phytochemistry, 31, pp.2983~2987 (1992)
  7. Benfey PN, "Regulated genes in transgenic plants", Science, 244, pp.174~181 (1989)
  8. Bentley R, "The shikimate pathway a metabolic tree with many branches", Crit Rev Biochem Mol Biol, 25, pp.307~384 (1990)
  9. Bischoff M, "Cloning of a cDNA encoding a 3-dehydroquinate synthase from a higher plant, and analysis of the organ-specificand elicitor-induced expression of the corresponding gene", Plant Mol Biol, 31, pp.69~76 (1996)
  10. Bischoff M, "Molecular characterization of tomato 3-dehydroquinate dehydratase-shikimate: NADP oxidoreductase", Plant Physiol, 125, pp.1891~1900 (2001)
  11. Bohlmann J, "Purification and cDNA cloning of anthranilate synthase from Ruta graveolens: modes of expression and properties of native and recombinant enzymes", Plant J, 7, pp.491~501 (1995)
  12. Bonner CA, "Novel features of prephenate aminotransferase from cell cultures of Nicotiana silvestris", Arch Biochem Biophys, 238, pp.237~246 (1985)
  13. Chibani K, "Proteomic analysis of seed dormancy in Arabidopsis", Plant Physiol, 142, pp.1493~1510 (2006)
  14. Cho HJ, "Increasing tryptophan synthesis in a forage legume Astragalus sinicus by expressing the tobacco feedback-insensitive anthranilate synthase (ASA2) gene", Plant Physiol, 123, pp.1069~1076 (2000)
  15. Cho M, "Phenylalanine biosynthesis in Arabidopsis thaliana identification and characterization of Arogenate dehydratases", J Biol Chem, 282, pp.30827~30835 (2007)
  16. Chook YM, "Crystal structures of the monofunctional chorismate mutase from Bacillus subtilis and its complex with a transition state analog", Proc Natl Acad Sci USA, 90, pp.8600~8603 (1993)
  17. Colquhoun TA, "A petunia chorismate mutase specialized for the production of floral volatiles", Plant J, 61, pp.145~155 (2010)
  18. Colquhoun TA, "Petunia floral volatile benzenoid", phenylpropanoid genes are regulated in a similar manner, 2010, pp.71, 158~167
  19. Connelly JA, "Tyrosine biosynthesis in Sorghum bicolor: isolation and regulatory properties of arogenate dehydrogenase", Z Naturforsch C, 41, pp.69~78 (1986)
  20. d’Amato T, "Subcellular localization of chorismate-mutase isoenzymes in protoplasts from mesophyll and suspension-cultured cells of Nicotiana silvestris", Planta, 162, pp.104~108 (1984)
  21. Devoto A, "Expression profiling reveals COI1 to be a key regulator of genes involved in wound- and methyl jasmonateinduced secondary metabolism, defence, and hormone interactions", Plant Mol Biol, 58, pp.497~513 (2005)
  22. Ding L, "Functional analysis of the essential bifunctional tobacco enzyme 3-dehydroquinate dehydratase", shikimate dehydrogenase in transgenic tobacco plants, 2007, pp.58, 2053~2067
  23. Djamei A, "Metabolic priming by a secreted fungal effector", Nature, 478, pp.395~398 (2011)
  24. Dombrecht B, "MYC2 differentially modulates diverse jasmonate-dependent functions in Arabidopsis", Plant Cell, 19, pp.2225~2245 (2007)
  25. Doong RL, "Plastid-localized 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase (DS-Mn):the early-pathway target of sequential feedback inhibition in higher plants", Plant Cell Environ, 16, pp.393~402 (1993)
  26. Dornfeld C, "Phylobiochemical characterization of class-Ib aspartate", prephenate aminotransferases reveals evolution of the plant arogenate phenylalanine pathway, 2014, pp.26, 3101~3114
  27. Duncan K, "The pentafunctional arom enzyme of Saccharomyces cerevisiae is a mosaic of monofunctional domains", Biochem J, 246, pp.375~386 (1987)
  28. Dyer WE, "A cDNA encoding 3-deoxy-d-arabino-heptulosonate 7-phosphate synthase from Solanum tuberosum L", J Biol Chem, 265, pp.1608~1614 (1990)
  29. Eberhard J, "Cytosolic and plastidic chorismate mutase isozymes from Arabidopsis thaliana: molecular characterization and enzymatic properties", Plant J, 10, pp.815~821 (1996)
  30. Ehlting J, "Global transcript profiling of primary stems from Arabidopsis thaliana identifies candidate genes for missing links in lignin biosynthesis and transcriptional regulators of fiber differentiation", Plant J, 42, pp.618~640 (2005)
  31. Emanuelsson O, "ChloroP, a neural network-based method for predicting chloroplast transit peptides and their cleavage sites", Protein Sci, 8, pp.978~984 (1999)
  32. Entus R, "Redox regulation of Arabidopsis 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase", Plant Physiol, 129, pp.1866~1871 (2002)
  33. Falk J, "Constitutive overexpression of barley 4-hydroxyphenylpyruvate dioxygenase in tobacco results in elevation of the vitamin E content in seeds but not in leaves", FEBS Lett, 540, pp.35~40 (2003)
  34. Fiedler E, "Localization, purification, and characterization of shikimate oxidoreductase dehydroquinate hydrolyase from stroma of spinach chloroplasts", Plant Physiol, 79, pp.212~218 (1985)
  35. Forlani G, "5-Enol-pyruvyl-shikimate-3-phosphate synthase from Zea mays cultured cells (purification and properties)", Plant Physiol, 105, pp.1107~1114 (1994)
  36. Fucile G, "Structural and biochemical investigation of two Arabidopsis shikimate kinases: The heat-inducible isoform is thermostable", Protein Sci, 20, pp.1125~1136 (2011)
  37. Funke T, "Molecular basis for the herbicide resistance of Roundup Ready crops", Proc Natl Acad Sci USA, 103, pp.13010~13015 (2006)
  38. Galili G, "Metabolic engineering of amino acids and storage proteins in plants", Metab Eng, 4, pp.3~11 (2002)
  39. Garg B, "In-silico analysis and expression profiling implicate diverse role of EPSPS family genes in regulating developmental and metabolic processes", BMC Res Notes, 7, pp.58~ (2014)
  40. Gasser CS, "Structure, expression, and evolution of the 5-enolpyruvylshikimate-3-phosphate synthase genes of petunia and tomato", J Biol Chem, 263, pp.4280~4287 (1988)
  41. Gilchrist DG, "Regulation of aromatic amino acid biosynthesis in higher plants: I. Evidence for a regulatory form of chorismate mutase in etiolated mung bean seedlings", Plant Physiol, 49, pp.52~57 (1972)
  42. Gorlach J, "Temporally distinct accumulation of transcripts encoding enzymes of the prechorismate pathway in elicitor-treated, cultured tomato cells", Proc Natl Acad Sci USA, 92, pp.3166~3170 (1995)
  43. Gorlach J, "Differential expression of tomato (Lycopersicon esculentum L.) genes encoding shikimate pathway isoenzymes. II. Chorismate synthase", Plant Mol Biol, 23, pp.707~716 (1993)
  44. Gross J, "A plant locus essential for phylloquinone (vitamin K1) biosynthesis originated from a fusion of four eubacterial genes", J Biol Chem, 281, pp.17189~17196 (2006)
  45. Hamberger B, "Comparative genomics of the shikimate pathway in Arabidopsis, Populus trichocarpa and Oryza sativa: shikimate pathway gene family structure and identification of candidates for missing links in phenylalanine biosynthesis", Recent Adv Phytochem, 40, pp.85~113 (2006)
  46. He Y, "Differential expression of triplicate phosphoribosylanthranilate isomerase isogenes in the tryptophan biosynthetic pathway of Arabidopsis thaliana (L.) Heynh", Planta, 212, pp.641~647 (2001)
  47. Herrmann KM, "The shikimate pathway", Annu Rev Plant Physiol Plant Mol Biol, 50, pp.473~503 (1999)
  48. Holding DR, "Identification and characterization of the maize arogenate dehydrogenase gene family", J Exp Bot, 61, pp.3663~3673 (2010)
  49. Huang L, "Purification and characterization of shikimate kinase enzyme activity in Bacillus subtilis", J Biol Chem, 250, pp.7675~7681 (1975)
  50. Hughes EH, "Metabolic engineering of the indole pathway in Catharanthus roseus hairy roots and increased accumulation of tryptamine and serpentine", Metab Eng, 6, pp.268~276 (2004)
  51. Huisman OC, "Regulation of aromatic amino acid biosynthesis in higher plants. II. 3-deoxy-arabino-heptulosonic acid 7-phosphate synthetase from cauliflower", J Biol Chem, 249, pp.6842~6848 (1974)
  52. Huynb QK, "5-enolpyruvyl shikimate-3-phosphate synthase from Escherichia coli: identification of Lys-22 as a potential active site residue", J Biol Chem, 263, pp.735~739 (1998)
  53. Ishihara A, "Metabolic changes in Arabidopsis thaliana expressing the feedback-resistant anthranilate synthase alpha subunit gene OASA1D", Phytochemistry, 67, pp.2349~2362 (2006)
  54. Job C, "Patterns of protein oxidation in Arabidopsis seeds and during germination", Plant Physiol, 138, pp.790~802 (2005)
  55. Jones PR, "A new member of plant CS-lyases: a cystine lyase from Arabidopsis thaliana", J Biol Chem, 278, pp.10291~10296 (2003)
  56. Jung E, "Chloroplasts of higher plants synthesize L-phenylalanine via Larogenate", Proc Natl Acad Sci USA, 83, pp.7231~7235 (1986)
  57. Kaminaga Y, "Plant phenylacetaldehyde synthase is a bifunctional homotetrameric enzyme that catalyzes phenylalanine decarboxylation and oxidation", J Biol Chem, 281, pp.23357~23366 (2006)
  58. Kasai K, "Identification of three shikimate kinase genes in rice: characterization of their differential expression during panicle development and of the enzymatic activities of the encoded proteins", Planta, 222, pp.438~447 (2005)
  59. Kaur H, "R2R3-NaMYB8 regulates the accumulation of phenylpropanoidpolyamine conjugates, which are essential for local and systemic defense against insect herbivores in Nicotiana attenuata", Plant Physiol, 152, pp.1731~1747 (2010)
  60. Keith B, "Differential induction of 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase genes in Arabidopsis thaliana by wounding and pathogenic attack", Proc Natl Acad Sci USA, 88, pp.8821~8825 (1991)
  61. Kim H, "The AAE14gene encodes the Arabidopsis o-succinylbenzoyl-CoA ligase that is essential for phylloquinone synthesis and photosystem-I function", Plant J, 54, pp.272~283 (2008)
  62. Klee HJ, "Cloning of an Arabidopsis thaliana gene encoding 5-enolpyruvylshikimate-3-phosphate synthase: sequence analysis and manipulation to obtain glyphosate tolerant plants", Mol Gen Genet, 210, pp.437~442 (1987)
  63. Kleeb AC, "A monofunctional and thermostable prephenate dehydratase from the archaeon Methanocaldococcus jannaschii", Biochemistry, 45, pp.14101~14110 (2006)
  64. Knaggs AR, "The biosynthesis of shikimate metabolites", Nat Prod Rep, 18, pp.334~355 (2001)
  65. Kriechbaumer V, "Characterisation of the tryptophan synthase alpha subunit in maize", BMC Plant Biol, 8, pp.44~ (2008)
  66. Kuroki GW, "Differential activities of chorismate mutase isozymes in tubers and leaves of Solanum tuberosum L", Plant Physiol, 89, pp.472~476 (1989)
  67. Lee AY, "Atomic structure of the buried catalytic pocket of Escherichia coli chorismate mutase", J Am Chem Soc, 117, pp.3627~3628 (1995)
  68. Legrand P, "Biochemical characterization and crystal structure of Synechocystis arogenate dehydrogenase provide insights into catalytic reaction", Structure, 14, pp.767~776 (2006)
  69. Leonard E, "Opportunities in metabolic engineering to facilitate scalable alkaloid production", Nat Chem Biol, 5, pp.292~300 (2009)
  70. Lepiniec L, "Genetics and biochemistry of seed flavonoids", Annu Rev Plant Biol, 57, pp.405~430 (2006)
  71. Li J, "The Arabidopsis thaliana trp5 mutant has a feedback-resistant anthranilate synthase and elevated soluble tryptophan", Plant Physiol, 110, pp.51~59 (1996)
  72. Li J, "Arabidopsis phosphoribosylanthranilate isomerase: molecular genetic analysis of triplicate tryptophan pathway genes", Plant Cell, 7, pp.447~461 (1995)
  73. Li J, "Isolation of cDNAs encoding the tryptophan pathway enzyme indole-3-glycerol phosphate synthase from Arabidopsis thaliana", Plant Physiol, 1088(1), pp.877~878 (1995)
  74. Lopukhina A, "Cloning and characterization of a coronatine-regulated tyrosine aminotransferase from Arabidopsis", Plant Physiol, 126, pp.1678~1687 (2001)
  75. Macheroux P, "A unique reaction in a common pathway: mechanism and function of chorismate synthase in the shikimate pathway", Planta, 207, pp.325~334 (1999)
  76. Maclean J, "The structure of chorismate synthase reveals a novel flavin binding site fundamental to a unique chemical reaction", Structure, 11, pp.1499~1511 (2003)
  77. Maeda H, "The shikimate pathway and aromatic amino acid biosynthesis in plants", Annu Rev Plant Biol, 63, pp.73~105 (2012)
  78. Maeda H, "RNAi suppression of Arogenate Dehydratase1 reveals that phenylalanine is synthesized predominantly via the arogenate pathway in petunia petals", Plant Cell, 22, pp.832~849 (2010)
  79. Maeda H, "Prephenate aminotransferase directs plant phenylalanine biosynthesis via arogenate", Nat Chem Biol, 7, pp.19~21 (2011)
  80. Malitsky S, "The transcript and metabolite networks affected by the two clades of Arabidopsis glucosinolate biosynthesis regulators", Plant Physiol, 148, pp.2021~2049 (2008)
  81. Matsuda F, "Characterization of tryptophan overproducing potato transgenic for a mutant rice anthranilate synthase alpha-subunit gene (OASA1D)", Planta, 222, pp.535~545 (2005)
  82. Melquist S, "Transcription from an upstream promoter controls methylation signaling from an inverted repeat of endogenous genes in Arabidopsis", Genes Dev, 17, pp.2036~2047 (2003)
  83. Mitsuda N, "NAC transcription factors, NST1 and NST3, are key regulators of the formation of secondary walls in woody tissues of Arabidopsis", Plant Cell, 19, pp.270~280 (2007)
  84. Mobley E, "Identification, characterization and comparative analysis of a novel chorismate mutase gene in Arabidopsis thaliana", Gene, 240, pp.115~123 (1999)
  85. Morino K, "Metabolic profiling of tryptophanoverproducing rice calli that express a feedback-insensitive alpha subunit of anthranilate synthase", Plant Cell Physiol, 46, pp.514~521 (2005)
  86. Morollo AA, "Structure of the cooperative allosteric anthranilate synthase from Salmonella typhimurium", Nat Struct Biol, 8, pp.243~247 (2001)
  87. Mousdale DM, "Detection and subcellular localization of a higher plant chorismate synthase", FEBS Lett, 205, pp.328~332 (1986)
  88. Niyogi KK, "Two anthranilate synthase genes in Arabidopsis: defense-related regulation of the tryptophan pathway", Plant Cell, 4, pp.721~733 (1992)
  89. Nobe Y, "The novel substrate recognition mechanism utilized by aspartate aminotransferase of the extreme thermophile Thermus thermophilus HB8", J Biol Chem, 273, pp.29554~29564 (1998)
  90. Oliva M, "Enhanced formation of aromatic amino acids increases fragrance without affecting flower longevity or pigmentation in Petunia× hybrida", Plant J, 13, pp.125~136 (2015)
  91. Ouyang J, "Indole-3-glycerol phosphate, a branch point of indole-3-acetic acid biosynthesis from the tryptophan biosynthetic pathway in Arabidopsis thaliana", Plant J, 24, pp.327~333 (2000)
  92. Pagnussat GC, "Genetic and molecular identification of genes required for female gametophyte development and function in Arabidopsis", Development, 132, pp.603~614 (2005)
  93. Pingsha H, "Functional contribution of chorismate synthase, anthranilate synthase, and chorismate mutase to penetration resistance in barley–powdery mildew interactions", MPMI, 22, pp.311~320 (2009)
  94. Pinto JE, "3-Deoxy-D-arabinoheptulosonate 7-phosphate synthase from potato tuber (Solanum tuberosum L.)", Plant Physiol, 82, pp.1040~1044 (1986)
  95. Prabhu PR, "Identification and partial charac terization of an L-tyrosine aminotransferase (TAT) from Arabidopsis thaliana", Biochem Res Int, 2010, pp.549~572 (2010)
  96. Quevillon-Cheruel S, "Crystal structure of the bifunctional chorismate synthase from Saccharomyces cerevisiae", J Biol Chem, 279, pp.619~625 (2004)
  97. Radwanski ER, "Characterization of tryptophan synthase alpha subunit mutants of Arabidopsis thaliana", Mol Gen Genet, 253, pp.353~361 (1996)
  98. Radwanski ER, "Tryptophan biosynthesis and metabolism: biochemical and molecular genetics", Plant Cell, 7, pp.921~934 (1995)
  99. Radwanski ER, "Arabidopsis thaliana tryptophan synthase alpha: gene cloning, expression, and subunit interaction", Mol Gen Genet, 248, pp.657~667 (1995)
  100. Rajjou L, "Proteomic investigation of the effect of salicylic acid on Arabidopsis seed germination and establishment of early defense mechanisms", Plant Physiol, 141, pp.910~923 (2006)
  101. Ramsay NA, "MYB-bHLH-WD40 protein complex and the evolution of cellular diversity", Trends Plant Sci, 10, pp.63~70 (2005)
  102. Reinink M, "3-Deoxy-D-arabino-heptulosonate 7-phosphate synthase from pea leaves: inhibition by L-tyrosine", Plant Sci Lett, 26, pp.167~171 (1982)
  103. Rippert P, "Purification and kinetic analysis of the two recombinant arogenate dehydrogenase isoforms of Arabidopsis thaliana", Eur J Biochem, 269, pp.4753~4761 (2002)
  104. Rippert P, "Engineering plant shikimate pathway for production of tocotrienol and improving herbicide resistance", Plant Physiol, 134, pp.92~100 (2004)
  105. Rippert P, "Tyrosine and phenylalanine are synthesized within the plastids in Arabidopsis", Plant Physiol, 149, pp.1251~1260 (2009)
  106. Romero RM, "Anthranilate synthase in microorganisms and plants", Phytochemistry, 39, pp.263~276 (1995)
  107. Romero RM, "Chorismate mutase in microorganisms and plants", Phytochemistry, 40, pp.1015~1025 (1995)
  108. Rose A, "A phosphoribosylanthranilate transferase gene is defective in blue fluorescent Arabidopsis thaliana tryptophan mutants", Plant Physiol, 100, pp.582~592 (1992)
  109. Rose AB, "Introns act post-transcriptionally to increase expression of the Arabidopsis thaliana tryptophan pathway gene PAT1", Plant J, 11, pp.455~464 (1997)
  110. Rose AB, "An allelic series of blue fluorescent trp1 mutants of Arabidopsis thaliana", Genetics, 145, pp.197~205 (1997)
  111. Wilczek), "Plant Physiol", 1979, 727~734
  112. Wilczek, "Plant Physiol", 1985, 711~718
  113. Sasaki-Sekimoto Y, "Coordinated activation of metabolic pathways for antioxidants and defense compounds by jasmonates and their roles in stress tolerance in Arabidopsis", Plant J, 44, pp.653~668 (2005)
  114. Schaller A, "Molecular cloning and analysis of a cDNA coding for chorismate synthase from the higher plant Corydalis sempervirens Pers", J Biol Chem, 266, pp.21434~21438 (1991)
  115. Schaller A, "Purification and characterization of chorismate synthase from Euglena gracilis: comparison with chorismate synthases of plant and microbial origin", Plant Physiol, 97, pp.1271~1279 (1991)
  116. Schmid J, "The in-vitro synthesized tomato shikimate kinase precursor is enzymatically active and is imported and processed to the mature enzyme by chloroplasts", Plant J, 2, pp.375~383 (1992)
  117. Siehl DL, "Kinetic and regulatory properties of arogenate dehydratase in seedlings of Sorghum bicolor (L.)Moench", Arch Biochem Biophys, 260, pp.822~829 (1988)
  118. Singh SA, "Structure of Arabidopsis dehydroquinate dehydratase-shikimate dehydrogenase and implications for metabolic channeling in the shikimate pathway", Biochemistry, 45, pp.7787~7796 (2006)
  119. Spitzer-Rimon B, "EOBII, a gene encoding a flower-specific regulator of phenylpropanoid volatiles’ biosynthesis in petunia", Plant Cell, 22, pp.1961~1976 (2010)
  120. Stenmark SL, "Blue-green bacteria synthesise L-tyrosine by the pre-tyrosine pathway", Nature, 247, pp.290~292 (1974)
  121. Stracke R, "Differential regulation of closely related R2R3-MYB transcription factors controls flavonol accumulation in different parts of the Arabidopsis thaliana seedling", Plant J, 50, pp.660~677 (2007)
  122. Suzich JA, "3-Deoxy-D-arabinoheptulosonate 7-phosphate synthase from carrot root (Daucus carota) is a hysteretic enzyme", Plant Physiol, 79, pp.765~770 (1985)
  123. Tohge T, "Shikimate and phenylalanine biosynthesis in the green lineage", Front Plant Sci, 4, pp.62~ (2013)
  124. Tozawa Y, "Characterization of rice anthranilate synthase alpha-subunit genes OASA1 and OASA2: tryptophan accumulation in transgenic rice expressing a feedback-insensitive mutant of OASA1", Plant Physiol, 126, pp.1493~1506 (2001)
  125. Tsegaye Y, "Overexpression of the enzyme p-hydroxyphenolpyruvate dioxygenase in Arabidopsis and its relation to tocopherol biosynthesis", Plant Physiol Biochem, 40, pp.913~920 (2002)
  126. Tzin V, "New insights into the shikimate and aromatic amino acids biosynthesis pathways in plants", Mol Plant, 3, pp.956~972 (2010)
  127. Tzin V, "Expression of a bacterial bi-functional chorismate mutase", prephenate dehydratase modulates primary and secondary metabolism associated with aromatic amino acids in Arabidopsis, 2009, pp.60, 156~167
  128. Tzin V, "Expression of a bacterial feedback-insensitive 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase of the shikimate pathway in Arabidopsis elucidates potential metabolic bottlenecks between primary and secondary metabolism", New Phytologist, 194, pp.430~439 (2012)
  129. Tzin V, "Tomato fruits expressing a bacterial feedback-insensitive 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase of the shikimate pathway possess enhanced levels of multiple specialized metabolites and upgraded aroma", J Exp Bot, 64, pp.4441~4452 (2013)
  130. Waller JC, "Developmental and feed forward control of the expression of folate biosynthesis genes in tomato fruit", Mol Plant, 3, pp.66~77 (2010)
  131. Wang W, "A novel 5-enolpyruvoylshikimate-3-phosphate (EPSP) synthase transgene for glyphosate resistance stimulates growth and fecundity in weedy rice (Oryza sativa) without herbicide", New Phytologist, 202, pp.679~688 (2014)
  132. Warpeha KM, "G-protein-coupled receptor 1, G-protein Gα-subunit 1, and prephenate dehydratase 1 are required for blue light-induced production of phenylalanine in etiolated Arabidopsis", Plant Physiol, 140, pp.844~855 (2006)
  133. L-phenylalanine into 2-phenylethanol and its β-D-glucopyranoside during the flower opening of Rosa ‘Hoh-Jun’ and Rosa damascena Mill, "Biosci Biotechno Biochem", 2002, 943~947
  134. Wildermuth M, "Isochorismate synthase is required to synthesize salicylic acid for plant defense", Nature, 414, pp.562~565 (2001)
  135. Wu J, "New insights into the evolutionary links relating to the 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase subfamilies", J Biol Chem, 281, pp.4042~4048 (2006)
  136. Yamada T, "Mutation of a rice gene encoding a phenylalanine biosynthetic enzyme results in accumulation of phenylalanine and tryptophan", Plant Cell, 20, pp.1316~1329 (2008)
  137. Yan Y, "A downstream mediator in the growth repression limb of the jasmonate pathway", Plant Cell, 19, pp.2470~2483 (2007)
  138. Yoo H, "An alternative pathway contributes to phenylalanine biosynthesis in plants via a cytosolic tyrosine: phenylpyruvate aminotransferase", Nat Commun, 4, pp.28~33 (2013)
  139. Zamir LO, "Enzymatic and nonenzymatic dehydration reactions of L-arogenate", Biochemistry, 24, pp.1607~1612 (1985)
  140. Zhang XH, "Targeting a nuclear anthranilate synthase α-subunit gene to the tobacco plastid genome results in enhanced tryptophan biosynthesis: return of a gene to its pre-endosymbiotic origin", Plant Physiol, 127, pp.131~141 (2001)
  141. Zhao J, "Immunological characterization and chloroplast localization of the tryptophan biosynthetic enzymes of the flowering plant Arabidopsis thaliana", J Biol Cheam, 270, pp.6081~6087 (1995)

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