If an enzyme name is shown in bold, there is experimental evidence for this enzymatic activity.
Some taxa known to possess this pathway include : Arabidopsis thaliana col
Expected Taxonomic Range: Viridiplantae
Brassinosteroids (BRs) are steroidal hormones that regulate the growth and development of plants. The most abundant and widely occurring BRs are C28 steroids, and among them, brassinolide (BL) is the most biologically active one. The biosynthesis pathway of BL was initially elucidated by extensive metabolic studies which suggested parallel-branched pathways, namely the early and late C-6 oxidation pathways ( brassinosteroid biosynthesis I). In all plant species examined to date, the 6-deoxo intermediates are predominant indicating that the late C-6 oxidation branch is the main route. Following the identification and biochemical characterization of CYP P450s involved in BR biosynthesis, an early C-22 oxidation branch ( brassinosteroid biosynthesis II) was proposed which was indicated to be the dominant route leading to BL biosynthesis. More recently, a new shortcut via C-23 hydroxylation following C-22 oxidation was demonstrated ( brassinosteroid biosynthesis II). Additional modifications (for example, lactonization) of the intermediates have also been reported [Katsumata08]. The BL biosynthesis pathway may be best represented as a grid where substantial interconnections between the different proposed routes exist [Ohnishi06, Shimada03].
Katsumata08: Katsumata T, Hasegawa A, Fujiwara T, Komatsu T, Notomi M, Abe H, Natsume M, Kawaide H (2008). "Arabidopsis CYP85A2 catalyzes lactonization reactions in the biosynthesis of 2-deoxy-7-oxalactone brassinosteroids." Biosci Biotechnol Biochem 72(8);2110-7. PMID: 18685225
Ohnishi06: Ohnishi T, Szatmari AM, Watanabe B, Fujita S, Bancos S, Koncz C, Lafos M, Shibata K, Yokota T, Sakata K, Szekeres M, Mizutani M (2006). "C-23 hydroxylation by Arabidopsis CYP90C1 and CYP90D1 reveals a novel shortcut in brassinosteroid biosynthesis." Plant Cell 18(11);3275-88. PMID: 17138693
Shimada03: Shimada Y, Goda H, Nakamura A, Takatsuto S, Fujioka S, Yoshida S (2003). "Organ-specific expression of brassinosteroid-biosynthetic genes and distribution of endogenous brassinosteroids in Arabidopsis." Plant Physiol 131(1);287-97. PMID: 12529536
Asami01: Asami T, Mizutani M, Fujioka S, Goda H, Min YK, Shimada Y, Nakano T, Takatsuto S, Matsuyama T, Nagata N, Sakata K, Yoshida S (2001). "Selective interaction of triazole derivatives with DWF4, a cytochrome P450 monooxygenase of the brassinosteroid biosynthetic pathway, correlates with brassinosteroid deficiency in planta." J Biol Chem 276(28);25687-91. PMID: 11319239
Asami03: Asami T, Mizutani M, Shimada Y, Goda H, Kitahata N, Sekimata K, Han SY, Fujioka S, Takatsuto S, Sakata K, Yoshida S (2003). "Triadimefon, a fungicidal triazole-type P450 inhibitor, induces brassinosteroid deficiency-like phenotypes in plants and binds to DWF4 protein in the brassinosteroid biosynthesis pathway." Biochem J 369(Pt 1);71-6. PMID: 12350224
Bancos02: Bancos S, Nomura T, Sato T, Molnar G, Bishop GJ, Koncz C, Yokota T, Nagy F, Szekeres M (2002). "Regulation of transcript levels of the Arabidopsis cytochrome p450 genes involved in brassinosteroid biosynthesis." Plant Physiol 130(1);504-13. PMID: 12226529
Bishop99: Bishop GJ, Nomura T, Yokota T, Harrison K, Noguchi T, Fujioka S, Takatsuto S, Jones JD, Kamiya Y (1999). "The tomato DWARF enzyme catalyses C-6 oxidation in brassinosteroid biosynthesis." Proc Natl Acad Sci U S A 96(4);1761-6. PMID: 9990098
Choe01: Choe S, Fujioka S, Noguchi T, Takatsuto S, Yoshida S, Feldmann KA (2001). "Overexpression of DWARF4 in the brassinosteroid biosynthetic pathway results in increased vegetative growth and seed yield in Arabidopsis." Plant J 26(6);573-82. PMID: 11489171
Choe98: Choe S, Dilkes BP, Fujioka S, Takatsuto S, Sakurai A, Feldmann KA (1998). "The DWF4 gene of Arabidopsis encodes a cytochrome P450 that mediates multiple 22alpha-hydroxylation steps in brassinosteroid biosynthesis." Plant Cell 1998;10(2);231-43. PMID: 9490746
Choe99: Choe S, Noguchi T, Fujioka S, Takatsuto S, Tissier CP, Gregory BD, Ross AS, Tanaka A, Yoshida S, Tax FE, Feldmann KA (1999). "The Arabidopsis dwf7/ste1 mutant is defective in the delta7 sterol C-5 desaturation step leading to brassinosteroid biosynthesis." Plant Cell 1999;11(2);207-21. PMID: 9927639
Ephritikhine99: Ephritikhine G, Pagant S, Fujioka S, Takatsuto S, Lapous D, Caboche M, Kendrick RE, Barbier-Brygoo H (1999). "The sax1 mutation defines a new locus involved in the brassinosteroid biosynthesis pathway in Arabidopsis thaliana." Plant J 18(3);315-20. PMID: 10377996
Finsterbusch99: Finsterbusch A, Lindemann P, Grimm R, Eckerskorn C, Luckner M (1999). "Delta(5)-3beta-hydroxysteroid dehydrogenase from Digitalis lanata Ehrh. - a multifunctional enzyme in steroid metabolism?." Planta 209(4);478-86. PMID: 10550629
Fujioka97: Fujioka S, Li J, Choi YH, Seto H, Takatsuto S, Noguchi T, Watanabe T, Kuriyama H, Yokota T, Chory J, Sakurai A (1997). "The Arabidopsis deetiolated2 mutant is blocked early in brassinosteroid biosynthesis." Plant Cell 9(11);1951-62. PMID: 9401120
Fujita06: Fujita S, Ohnishi T, Watanabe B, Yokota T, Takatsuto S, Fujioka S, Yoshida S, Sakata K, Mizutani M (2006). "Arabidopsis CYP90B1 catalyses the early C-22 hydroxylation of C27, C28 and C29 sterols." Plant J 45(5);765-74. PMID: 16460510
Hong02: Hong Z, Ueguchi-Tanaka M, Shimizu-Sato S, Inukai Y, Fujioka S, Shimada Y, Takatsuto S, Agetsuma M, Yoshida S, Watanabe Y, Uozu S, Kitano H, Ashikari M, Matsuoka M (2002). "Loss-of-function of a rice brassinosteroid biosynthetic enzyme, C-6 oxidase, prevents the organized arrangement and polar elongation of cells in the leaves and stem." Plant J 32(4);495-508. PMID: 12445121
Kim05: Kim GT, Fujioka S, Kozuka T, Tax FE, Takatsuto S, Yoshida S, Tsukaya H (2005). "CYP90C1 and CYP90D1 are involved in different steps in the brassinosteroid biosynthesis pathway in Arabidopsis thaliana." Plant J 41(5);710-21. PMID: 15703058
Kim05a: Kim TW, Hwang JY, Kim YS, Joo SH, Chang SC, Lee JS, Takatsuto S, Kim SK (2005). "Arabidopsis CYP85A2, a cytochrome P450, mediates the Baeyer-Villiger oxidation of castasterone to brassinolide in brassinosteroid biosynthesis." Plant Cell 17(8);2397-412. PMID: 16024588
Koka00: Koka CV, Cerny RE, Gardner RG, Noguchi T, Fujioka S, Takatsuto S, Yoshida S, Clouse SD (2000). "A putative role for the tomato genes DUMPY and CURL-3 in brassinosteroid biosynthesis and response." Plant Physiol 122(1);85-98. PMID: 10631252
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