Summary: Cholestanes substituted in any position with one or more hydroxy groups. They are found in feces and bile. In contrast to bile acids and salts, they are not reabsorbed.

Top Publications

  1. Albrecht C, Russinova E, Kemmerling B, Kwaaitaal M, de Vries S. Arabidopsis SOMATIC EMBRYOGENESIS RECEPTOR KINASE proteins serve brassinosteroid-dependent and -independent signaling pathways. Plant Physiol. 2008;148:611-9 pubmed publisher
    ..We propose that, in plants, closely related receptor kinases have a minimal homo- or heterodimeric configuration to achieve specificity. ..
  2. Bao F, Shen J, Brady S, Muday G, Asami T, Yang Z. Brassinosteroids interact with auxin to promote lateral root development in Arabidopsis. Plant Physiol. 2004;134:1624-31 pubmed
  3. Wang X, Kota U, He K, Blackburn K, Li J, Goshe M, et al. Sequential transphosphorylation of the BRI1/BAK1 receptor kinase complex impacts early events in brassinosteroid signaling. Dev Cell. 2008;15:220-35 pubmed publisher
    ..This model suggests both conservation and distinct differences between the molecular mechanisms regulating phosphorylation-dependent kinase activation in plant and animal receptor kinases. ..
  4. Hong Z, Ueguchi Tanaka M, Shimizu Sato S, Inukai Y, Fujioka S, Shimada Y, et al. 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. 2002;32:495-508 pubmed
    ..On the basis of these findings, we discuss the biological function of BRs in rice plants. ..
  5. Bancos S, Nomura T, Sato T, Molnar G, Bishop G, Koncz C, et al. Regulation of transcript levels of the Arabidopsis cytochrome p450 genes involved in brassinosteroid biosynthesis. Plant Physiol. 2002;130:504-13 pubmed
    ..Inverse correlation between CYP90A1/CPD transcript levels and the amounts of the CYP90A1 substrate 6-deoxocathasterone in shoots and roots suggests that transcriptional regulation plays an important role in controlling BR biosynthesis. ..
  6. Li J, Wen J, Lease K, Doke J, Tax F, Walker J. BAK1, an Arabidopsis LRR receptor-like protein kinase, interacts with BRI1 and modulates brassinosteroid signaling. Cell. 2002;110:213-22 pubmed
    ..Expression of a dominant-negative mutant allele of BAK1 causes a severe dwarf phenotype, resembling the phenotype of null bri1 alleles. These results indicate BAK1 is a component of BR signaling. ..
  7. Tang W, Kim T, Oses Prieto J, Sun Y, Deng Z, Zhu S, et al. BSKs mediate signal transduction from the receptor kinase BRI1 in Arabidopsis. Science. 2008;321:557-60 pubmed publisher
    ..These results demonstrate that BSKs are the substrates of BRI1 kinase that activate downstream BR signal transduction. ..
  8. Jin H, Hong Z, Su W, Li J. A plant-specific calreticulin is a key retention factor for a defective brassinosteroid receptor in the endoplasmic reticulum. Proc Natl Acad Sci U S A. 2009;106:13612-7 pubmed publisher
    ..Our study revealed not only a functional role for a plant-specific CRT, but also functional diversity among the 3 Arabidopsis CRT paralogues. ..
  9. Yu J, Huang L, Hu W, Zhou Y, Mao W, Ye S, et al. A role for brassinosteroids in the regulation of photosynthesis in Cucumis sativus. J Exp Bot. 2004;55:1135-43 pubmed
    ..It was concluded that EBR increases the capacity of CO(2) assimilation in the Calvin cycle, which was mainly attributed to an increase in the initial activity of Rubisco. ..

More Information


  1. Kinoshita T, Caño Delgado A, Seto H, Hiranuma S, Fujioka S, Yoshida S, et al. Binding of brassinosteroids to the extracellular domain of plant receptor kinase BRI1. Nature. 2005;433:167-71 pubmed
    ..Our results demonstrate that brassinosteroids bind directly to the 94 amino acids comprising ID-LRR22 in the extracellular domain of BRI1, and define a new binding domain for steroid hormones. ..
  2. Wang Z, Wang Q, Chong K, Wang F, Wang L, Bai M, et al. The brassinosteroid signal transduction pathway. Cell Res. 2006;16:427-34 pubmed
    ..The BR signaling pathway has become a paradigm for both receptor kinase signaling in plants and steroid signaling by cell surface receptors in general. ..
  3. Mori M, Nomura T, Ooka H, Ishizaka M, Yokota T, Sugimoto K, et al. Isolation and characterization of a rice dwarf mutant with a defect in brassinosteroid biosynthesis. Plant Physiol. 2002;130:1152-61 pubmed
  4. Chono M, Honda I, Zeniya H, Yoneyama K, Saisho D, Takeda K, et al. A semidwarf phenotype of barley uzu results from a nucleotide substitution in the gene encoding a putative brassinosteroid receptor. Plant Physiol. 2003;133:1209-19 pubmed
    ..The uzu gene is being introduced into all hull-less barley cultivars in Japan as an effective dwarf gene for practical use, and this is the first report about an agronomically important mutation related to BRs. ..
  5. Nomura T, Kushiro T, Yokota T, Kamiya Y, Bishop G, Yamaguchi S. The last reaction producing brassinolide is catalyzed by cytochrome P-450s, CYP85A3 in tomato and CYP85A2 in Arabidopsis. J Biol Chem. 2005;280:17873-9 pubmed publisher
    ..We postulate that castasterone is the major active brassinosteroid during vegetative growth in tomato, whereas brassinolide may play an organ-specific role in fruit development in this species...
  6. Wang Z, He J. Brassinosteroid signal transduction--choices of signals and receptors. Trends Plant Sci. 2004;9:91-6 pubmed
    ..Recent studies demonstrated that tomato BRI1 (tBRI1) perceives both BR and the peptide hormone systemin, raising new questions about the molecular mechanism and evolution of receptor-ligand specificity. ..
  7. Pullman G, Zhang Y, Phan B. Brassinolide improves embryogenic tissue initiation in conifers and rice. Plant Cell Rep. 2003;22:96-104 pubmed
    ..61 mg/l kinetin, 3.4 mg/l silver nitrate, 10 micro M cGMP, 0.1 micro M brassinolide, and 2 g/l Gelrite. Across 12 open-pollinated families of loblolly pine, initiation percentages ranged from 2.5% to 50.7%, averaging 22.5%...
  8. He J, Fujioka S, Li T, Kang S, Seto H, Takatsuto S, et al. Sterols regulate development and gene expression in Arabidopsis. Plant Physiol. 2003;131:1258-69 pubmed
  9. Choe S, Schmitz R, Fujioka S, Takatsuto S, Lee M, Yoshida S, et al. Arabidopsis brassinosteroid-insensitive dwarf12 mutants are semidominant and defective in a glycogen synthase kinase 3beta-like kinase. Plant Physiol. 2002;130:1506-15 pubmed
  10. Zhao J, Peng P, Schmitz R, Decker A, Tax F, Li J. Two putative BIN2 substrates are nuclear components of brassinosteroid signaling. Plant Physiol. 2002;130:1221-9 pubmed
    ..We propose that BES1/BZR1 are two nuclear components of BR signaling that are negatively regulated by BIN2 through a phosphorylation-initiated process. ..
  11. Kagale S, Divi U, Krochko J, Keller W, Krishna P. Brassinosteroid confers tolerance in Arabidopsis thaliana and Brassica napus to a range of abiotic stresses. Planta. 2007;225:353-64 pubmed
    ..Both det2-1 and dwf4 mutants still expressed heat shock proteins (hsps) to high levels during HS, indicating that although BR augments thermotolerance in plants, it is not necessary for hsp expression during HS. ..
  12. Yan Z, Zhao J, Peng P, Chihara R, Li J. BIN2 functions redundantly with other Arabidopsis GSK3-like kinases to regulate brassinosteroid signaling. Plant Physiol. 2009;150:710-21 pubmed publisher
  13. Sekimata K, Kimura T, Kaneko I, Nakano T, Yoneyama K, Takeuchi Y, et al. A specific brassinosteroid biosynthesis inhibitor, Brz2001: evaluation of its effects on Arabidopsis, cress, tobacco, and rice. Planta. 2001;213:716-21 pubmed
    ..Brz2001 can be used to clarify the function of BRs in dicots as a complement to BR-deficient mutants, and to elucidate the different roles of BRs in monocots and dicots. ..
  14. Zurek D, Clouse S. Molecular cloning and characterization of a brassinosteroid-regulated gene from elongating soybean (Glycine max L.) epicotyls. Plant Physiol. 1994;104:161-70 pubmed
    ..The elevated levels of BRU1 transcripts in elongating tissue and the homology with a xyloglucan endotransglycosylase suggest a possible role for the BRU1 protein in brassinosteroid-stimulated elongation. ..
  15. Duan K, Li L, Hu P, Xu S, Xu Z, Xue H. A brassinolide-suppressed rice MADS-box transcription factor, OsMDP1, has a negative regulatory role in BR signaling. Plant J. 2006;47:519-31 pubmed
  16. Silvente Poirot S, Poirot M. Cholesterol epoxide hydrolase and cancer. Curr Opin Pharmacol. 2012;12:696-703 pubmed publisher
  17. Hu Y, Poh H, Chua N. The Arabidopsis ARGOS-LIKE gene regulates cell expansion during organ growth. Plant J. 2006;47:1-9 pubmed
    ..Ectopic expression of ARL in bri1-119 partially restores cell growth in cotyledons and leaves. Our results suggest that ARL acts downstream of BRI1 and partially mediates BR-related cell expansion signals during organ growth. ..
  18. Kim B, Fujioka S, Takatsuto S, Tsujimoto M, Choe S. Castasterone is a likely end product of brassinosteroid biosynthetic pathway in rice. Biochem Biophys Res Commun. 2008;374:614-9 pubmed publisher
    ..However, only a single copy of this gene is found in the currently available genome sequences of graminaceous plants; this is a likely explanation for the absence of an endogenous pool of brassinolide in rice plants. ..
  19. Brunel J, Salmi C, Loncle C, Vidal N, Letourneux Y. Squalamine: a polyvalent drug of the future?. Curr Cancer Drug Targets. 2005;5:267-72 pubmed
    ..All these results led to a question: could we consider squalamine as a polyvalent drug of the future? ..
  20. Tanaka A, Nakagawa H, Tomita C, Shimatani Z, Ohtake M, Nomura T, et al. BRASSINOSTEROID UPREGULATED1, encoding a helix-loop-helix protein, is a novel gene involved in brassinosteroid signaling and controls bending of the lamina joint in rice. Plant Physiol. 2009;151:669-80 pubmed publisher
    ..These results indicate that BU1 may participate in some other unknown processes modulated by BR in rice. ..
  21. de Medina P, Paillasse M, Payre B, Silvente Poirot S, Poirot M. Synthesis of new alkylaminooxysterols with potent cell differentiating activities: identification of leads for the treatment of cancer and neurodegenerative diseases. J Med Chem. 2009;52:7765-77 pubmed publisher
  22. Rozhon W, Mayerhofer J, Petutschnig E, Fujioka S, Jonak C. ASKtheta, a group-III Arabidopsis GSK3, functions in the brassinosteroid signalling pathway. Plant J. 2010;62:215-23 pubmed publisher
    ..ASKtheta phosphorylated BEH2 both in vitro and in vivo. Overall, these data provide strong evidence that ASKtheta is a novel component of the BR signalling cascade, targeting the transcription factors BES1, BZR1 and BEH2. ..
  23. Li J, Nagpal P, Vitart V, McMorris T, Chory J. A role for brassinosteroids in light-dependent development of Arabidopsis. Science. 1996;272:398-401 pubmed
    ..Thus, DET2 may encode a reductase in the brassinolide biosynthetic pathway, and brassinosteroids may constitute a distinct class of phytohormones with an important role in light-regulated development of higher plants. ..
  24. Bouquin T, Meier C, Foster R, Nielsen M, Mundy J. Control of specific gene expression by gibberellin and brassinosteroid. Plant Physiol. 2001;127:450-8 pubmed
    ..Reporter transgene analyses and RNA-blot analysis showed that BR and GA modulate GA5 expression, at least in part, at the transcriptional level, and that the signals are independent and subtractive. ..
  25. Bishop G, Koncz C. Brassinosteroids and plant steroid hormone signaling. Plant Cell. 2002;14 Suppl:S97-110 pubmed
  26. Jonak C, Hirt H. Glycogen synthase kinase 3/SHAGGY-like kinases in plants: an emerging family with novel functions. Trends Plant Sci. 2002;7:457-61 pubmed
    ..Analysis of the Arabidopsis genome revealed the existence of ten GSK genes that fall into four distinct subfamilies. We discuss the functions and mechanisms of GSK action in plants and other organisms. ..
  27. Goda H, Sawa S, Asami T, Fujioka S, Shimada Y, Yoshida S. Comprehensive comparison of auxin-regulated and brassinosteroid-regulated genes in Arabidopsis. Plant Physiol. 2004;134:1555-73 pubmed
  28. Jiang X, Sidhu R, Porter F, Yanjanin N, Speak A, Te Vruchte D, et al. A sensitive and specific LC-MS/MS method for rapid diagnosis of Niemann-Pick C1 disease from human plasma. J Lipid Res. 2011;52:1435-45 pubmed publisher
    ..The LC-MS/MS assay was able to discriminate with high sensitivity and specificity between control and NPC1 subjects, and offers for the first time a noninvasive, rapid, and highly sensitive method for diagnosis of NPC1 disease. ..
  29. Fujioka S, Noguchi T, Sekimoto M, Takatsuto S, Yoshida S. 28-Norcastasterone is biosynthesized from castasterone. Phytochemistry. 2000;55:97-101 pubmed
    ..Moreover, the natural occurrence of 28-norcastasterone and 28-nortyphasterol in seedlings of A. thaliana has been demonstrated. This is the first report of the natural occurrence of 28-nortyphasterol in plants. ..
  30. Sharma A, Matsuoka M, Tanaka H, Komatsu S. Antisense inhibition of a BRI1 receptor reveals additional protein kinase signaling components downstream to the perception of brassinosteroids in rice. FEBS Lett. 2001;507:346-50 pubmed
    ..The data obtained suggest that MAPK and Ca(2+)-dependent protein kinase in rice are discrete but parallel signaling cascades and might involve receptors other than BRI1 in response to BR stimulus. ..
  31. Belkhadir Y, Chory J. Brassinosteroid signaling: a paradigm for steroid hormone signaling from the cell surface. Science. 2006;314:1410-1 pubmed
    ..BRs bind a small family of leucine-rich repeat receptor kinases at the cell surface, thereby initiating an intracellular signal transduction cascade that results in the altered expression of hundreds of genes. ..
  32. Wang X, Chory J. Brassinosteroids regulate dissociation of BKI1, a negative regulator of BRI1 signaling, from the plasma membrane. Science. 2006;313:1118-22 pubmed
    ..BKI1 is a substrate of BRI1 kinase and limits the interaction of BRI1 with its proposed coreceptor, BAK1, suggesting that BKI1 prevents the activation of BRI1. ..
  33. Kim T, Chang S, Lee J, Takatsuto S, Yokota T, Kim S. Novel biosynthetic pathway of castasterone from cholesterol in tomato. Plant Physiol. 2004;135:1231-42 pubmed
    ..In conclusion, the biosynthesis of 28-norBRs appears to play a physiologically important role in maintaining homeostatic levels of CS in tomato seedlings. ..
  34. Vidya Vardhini B, Rao S. Acceleration of ripening of tomato pericarp discs by brassinosteroids. Phytochemistry. 2002;61:843-7 pubmed
    ..Fruit ripening as induced by brassinosteroids was associated with increase in ethylene production. The study revealed the ability of brassinosteroids in accelerating fruit-senescence. ..
  35. Müssig C. Brassinosteroid-promoted growth. Plant Biol (Stuttg). 2005;7:110-7 pubmed
    ..This review focuses on physiology and molecular mechanisms underlying BR-promoted growth in the different plant organs. Interactions with other phytohormones are discussed. ..
  36. Castle J, Szekeres M, Jenkins G, Bishop G. Unique and overlapping expression patterns of Arabidopsis CYP85 genes involved in brassinosteroid C-6 oxidation. Plant Mol Biol. 2005;57:129-40 pubmed
  37. Turk E, Fujioka S, Seto H, Shimada Y, Takatsuto S, Yoshida S, et al. BAS1 and SOB7 act redundantly to modulate Arabidopsis photomorphogenesis via unique brassinosteroid inactivation mechanisms. Plant J. 2005;42:23-34 pubmed
  38. Ahima R, Patel H, Takahashi N, Qi Y, Hileman S, Zasloff M. Appetite suppression and weight reduction by a centrally active aminosterol. Diabetes. 2002;51:2099-104 pubmed
    ..These findings indicate that MSI-1436 acts in the brain to regulate food intake and energy expenditure, likely through suppression of orexigenic hypothalamic pathways. ..
  39. Poppenberger B, Fujioka S, Soeno K, George G, Vaistij F, Hiranuma S, et al. The UGT73C5 of Arabidopsis thaliana glucosylates brassinosteroids. Proc Natl Acad Sci U S A. 2005;102:15253-8 pubmed
    ..These data support the hypothesis that 23-O-glucosylation of BL is a function of UGT73C5 in planta, and that glucosylation regulates BR activity. ..
  40. Hong Z, Ueguchi Tanaka M, Umemura K, Uozu S, Fujioka S, Takatsuto S, et al. A rice brassinosteroid-deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell. 2003;15:2900-10 pubmed
    ..Based on these results, we conclude that D2/CYP90D2 catalyzes the steps from 6-deoxoteasterone to 3-dehydro-6-deoxoteasterone and from teasterone to 3-dehydroteasterone in the late BR biosynthesis pathway. ..
  41. Wang Z, Seto H, Fujioka S, Yoshida S, Chory J. BRI1 is a critical component of a plasma-membrane receptor for plant steroids. Nature. 2001;410:380-3 pubmed
  42. Nakamura A, Higuchi K, Goda H, Fujiwara M, Sawa S, Koshiba T, et al. Brassinolide induces IAA5, IAA19, and DR5, a synthetic auxin response element in Arabidopsis, implying a cross talk point of brassinosteroid and auxin signaling. Plant Physiol. 2003;133:1843-53 pubmed
  43. Dhaubhadel S, Browning K, Gallie D, Krishna P. Brassinosteroid functions to protect the translational machinery and heat-shock protein synthesis following thermal stress. Plant J. 2002;29:681-91 pubmed
  44. Nakaya M, Tsukaya H, Murakami N, Kato M. Brassinosteroids control the proliferation of leaf cells of Arabidopsis thaliana. Plant Cell Physiol. 2002;43:239-44 pubmed
    ..Restoration of leaf size could not be explained solely on the basis of an increase in individual cell volume, thus suggesting that brassinosteroids play a dual role in regulating cell expansion and proliferation. ..
  45. De Rybel B, Audenaert D, Vert G, Rozhon W, Mayerhofer J, Peelman F, et al. Chemical inhibition of a subset of Arabidopsis thaliana GSK3-like kinases activates brassinosteroid signaling. Chem Biol. 2009;16:594-604 pubmed publisher
    ..The opportunity to generate multiple and conditional knockouts in key regulators in the BR signaling pathway by bikinin represents a useful tool to further unravel regulatory mechanisms. ..
  46. Yun H, Bae Y, Lee Y, Chang S, Kim S, Li J, et al. Analysis of phosphorylation of the BRI1/BAK1 complex in arabidopsis reveals amino acid residues critical for receptor formation and activation of BR signaling. Mol Cells. 2009;27:183-90 pubmed publisher
    ..These results suggest that BAK1 is a critical component regulating the duration of BR efficacy, even though it cannot directly bind BRs in plants. ..
  47. Alhanout K, Malesinki S, Vidal N, Peyrot V, Rolain J, Brunel J. New insights into the antibacterial mechanism of action of squalamine. J Antimicrob Chemother. 2010;65:1688-93 pubmed publisher
    ..The new insights into the mechanism of action of squalamine highlight the importance of aminosterols in the design of a new class of antibacterial compounds that could be used as disinfectants and detergents. ..
  48. Selinsky B, Zhou Z, Fojtik K, Jones S, Dollahon N, Shinnar A. The aminosterol antibiotic squalamine permeabilizes large unilamellar phospholipid vesicles. Biochim Biophys Acta. 1998;1370:218-34 pubmed
    ..At higher squalamine/phospholipid ratios, these structures release from the bilayers and aggregate to form either new vesicles or squalamine/phospholipid mixed micelles. ..
  49. Bajguz A. Brassinosteroid enhanced the level of abscisic acid in Chlorella vulgaris subjected to short-term heat stress. J Plant Physiol. 2009;166:882-6 pubmed publisher
    ..The present work also demonstrates that Chlorella vulgaris can synthesize ABA. This is the first evidence of ABA detection in Chlorella vulgaris cells...
  50. Bishop G. Refining the plant steroid hormone biosynthesis pathway. Trends Plant Sci. 2007;12:377-80 pubmed
    ..2006) indicates the role of CYP90C1 and CYP90D1 in the synthesis of the most bioactive plant steroid hormone, brassinolide. These results highlight the need for refining the brassinolide biosynthesis pathway. ..
  51. Bajguz A. Metabolism of brassinosteroids in plants. Plant Physiol Biochem. 2007;45:95-107 pubmed
    ..Metabolism of brassinosteroids can be divided into two categories: i) structural changes to the steroidal skeleton; and ii) structural changes to the side-chain. ..
  52. Fujioka S, Yokota T. Biosynthesis and metabolism of brassinosteroids. Annu Rev Plant Biol. 2003;54:137-64 pubmed
    ..Various metabolic reactions of BRs including epimerization, oxidation, and conjugation are also summarized. ..
  53. Kim T, Guan S, Sun Y, Deng Z, Tang W, Shang J, et al. Brassinosteroid signal transduction from cell-surface receptor kinases to nuclear transcription factors. Nat Cell Biol. 2009;11:1254-60 pubmed publisher
    ..This study establishes a fully connected BR signalling pathway and provides new insights into the mechanism of GSK3 regulation. ..