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PubMed2004

A mammalian steroid action inhibitor spironolactone retards plant growth by inhibition of brassinosteroid action and induces light-induced gene expression in the dark.

We screened steroid derivatives and found that spironolactone, an inhibitor of both 17beta-hydroxysteroid dehydrogenase (17beta-HSD) and aldosterone receptor, is an inhibitor of phytohormone brassinosteroid (BR) action in plants. Under both dark and light growing conditions, spironolactone induced morphological changes in Arabidopsis, characteristic of brassinosteroid-deficient mutants. Spironolactone-treated plants were also nearly restored to the wild-type phenotype by treatment with additional BRs. In the spironolactone-treated Arabidopsis, the CPD gene in the BR biosynthesis pathway was up-regulated, probably due to feedback regulation caused by BR-deficiency. Spironolactone-treated tobacco plants grown in the dark showed expression of light-regulated genes as was observed in the deficient mutant. These data suggest that spironolactone inhibits brassinosteroid action probably due to the blockage of biosynthesis and exerts its activity against plants. Thus, spironolactone, in conjunction with brassinosteroid-deficient mutants, can be used to clarify the function of BRs in plants and characterize mutants. The spironolactone action site was also investigated by feeding BR biosynthesis intermediates to Arabidopsis grown in the dark, and the results are discussed.

Animals↗
PubMed2001

Accumulation of 6-deoxocathasterone and 6-deoxocastasterone in Arabidopsis, pea and tomato is suggestive of common rate-limiting steps in brassinosteroid biosynthesis.

To gain a better understanding of brassinosteroid biosynthesis, the levels of brassinosteroids and sterols related to brassinolide biosynthesis in Arabidopsis, pea, and tomato plants were quantified by gas chromatography-selected ion monitoring. In these plants, the late C-6 oxidation pathway was found to be the predominant pathway in the synthesis of castasterone. Furthermore, all these plant species had similar BR profiles, suggesting the presence of common biosynthetic control mechanisms. The especially high levels of 6-deoxocathasterone and 6-deoxocastasterone may indicate that their respective conversions to 6-deoxoteasterone and castasterone are regulated in planta and hence are important rate-limiting steps in brassinosteroid biosynthesis. Other possible rate-limiting reactions, including the conversion of campestanol to 6-deoxocathasteonre. are also discussed. Tomato differs from Arabidopsis and pea in that tomato contains 28-norcastasterone as a biologically active brassinosteroid, and that its putative precursors, cholesterol and its relatives are the major sterols.

Arabidopsis↗
PubMed2003

The chemical characteristic and distribution of brassinosteroids in plants.

Brassinosteroids represent a class of plant hormones with high-growth promoting activity. They are found at low levels in pollen, anthers, seeds, leaves, stems, roots, flowers, grain, and young vegetative tissues throughout the plant kingdom. Brassinosteroids are a family of about 60 phytosteroids. The article gives a comprehensive survey on the hitherto known brassinosteroids isolated from plants. The chemical characteristic of brassinosteroids is also presented.

Phytosterols↗
PubMed1998

Brassinosteroids in Arabidopsis thaliana.

From the seeds and siliques of Arabidopsis thaliana, six brassinosteroids, brassinolide, castasterone, typhasterol, 6-deoxocastasterone, 6-deoxotyphasterol and 6-deoxoteasterone, were identified by GC-mass spectrometry or GC-selected ion monitoring. As the occurrence of castasterone, typhasterol, 6-deoxocastasterone and 6-deoxotyphasterol in the shoots of A. thaliana has already been reported, this study provides evidence for the occurrence of the above four brassinosteroids in different organs, seeds and siliques, and the first evidence for the occurrence of brassinolide and 6-deoxoteasterone in A. thaliana. All brassinosteroids identified in this study belong to important components of both the early and late C-6 oxidation pathways, which were established in the cultured cells of Catharanthus roseus. This suggests that both pathways are operating in A. thaliana to produce the most biologically active brassinosteroid, brassinolide, which is responsible for growth and development of the plant.

Arabidopsis↗
PubMed2005

Activation of the cytochrome P450 gene, CYP72C1, reduces the levels of active brassinosteroids in vivo.

A pool of Arabidopsis lines transformed with the activation vector was screened for short hypocotyl mutants under dim far-red light, and three mutant lines designated chibi1-3 (chi) were isolated. Among the chi mutants, chi2 was dominant. The chi2 seedlings were short, regardless of the light conditions. The chi2 mature plants exhibited phenotypic features such as dwarfism, reduced male fertility and dark green, rounded epinastic leaves, which are characteristics of brassinosteroid-deficient mutants such as det2, cpd, and dwf4. Furthermore, the hypocotyl phenotype was restored by the addition of brassinolide to the culture medium, suggesting that brassinosteroids had been affected in this mutant. The molecular analysis of the chi2 mutant revealed that the CYP72C1 gene was overexpressed by the enhancing activity of the inserted DNA. Wild-type plants that were transformed with a vector containing a chimeric gene between the 35S promoter and the CYP72C1 genomic DNA exhibited a similar phenotype. Consistent with the morphological and physiological phenotype, the levels of active brassinosteroids were reduced in the chi2 mutant. Hence, CYP72C1, together with BAS1/CYP72B1, is speculated to regulate active brassinosteroid levels in plants. Expression analysis suggested that wild-type CYP72C1 transcript levels increased after exposure to white light, although the physiological significance of such a response remains obscure.

Arabidopsis↗
PubMed1994

Investigation of Gene Expression, Growth Kinetics, and Wall Extensibility during Brassinosteroid-Regulated Stem Elongation.

Brassinosteroids promote stem elongation in a variety of plants but little is known about the mechanism of action of these plant growth regulators. We investigated a number of physiological and molecular parameters associated with brassinosteroid-enhanced elongation. Continuous growth recordings of soybean (Glycine max L. cv Williams 82) epicotyls showed that there was a 45-min lag before 0.1 [mu]M brassinolide (BR) exerted a detectable effect on elongation. BR caused a marked increase in Instron-measured plastic extensibility, suggesting that BR may promote elongation in part by altering mechanical properties of the cell wall (wall loosening). Structure-function studies suggested that the dimensions of the brassinosteroid side chain were critical for promotion of elongation and expression of BRU1, a gene regulated specifically by active brassinosteroids. Auxin-BR interactions were examined by using small auxin up RNA (SAUR) gene probes and the auxin-insensitive diageotropica (dgt) mutant of tomato (Lycopersicon esculentum Mill.). We have shown that in wild-type tomato, which elongates in response to exogenous auxin, a transcript of identical size to the soybean SAUR 15A is strongly induced within 1 h by 50 [mu]M 2,4-dichlorophenoxyacetic acid or indoleacetic acid, whereas in the dgt mutant, which does not elongate in response to auxin, no transcript is expressed. Furthermore, BR promotes equal elongation of hypocotyls in both wild-type and dgt tomatoes but does not rapidly induce the SAUR 15A homolog in either genotype. BR does not cause rapid induction of SAUR 6B in elongating soybean epicotyls but does lead to increased expression after 18 h. This late BR activation of SAUR 6B is controlled, at least in part, at the transcriptional level and is not accompanied by an increase of free indoleacetic acid in the tissue. We conclude that although both BR and auxin affect wall relaxation processes, BR-promoted elongation in soybean and tomato stems acts via a mechanism that most likely does not proceed through the auxin signal transduction pathway.

Journal Article↗
PubMed2006

The regulation of DWARF4 expression is likely a critical mechanism in maintaining the homeostasis of bioactive brassinosteroids in Arabidopsis.

Mutants that are defective in brassinosteroid (BR) biosynthesis or signaling display severely retarded growth patterns due to absence of growth-promoting effects by BRs. Arabidopsis (Arabidopsis thaliana) DWARF4 (DWF4) catalyzes a flux-determining step in the BR biosynthetic pathways. Thus, it is hypothesized that the tissues of DWF4 expression may represent the sites of BR biosynthesis in Arabidopsis. Here we show that DWF4 transcripts accumulate in the actively growing tissues, such as root, shoot apices with floral clusters, joint tissues of root and shoot, and dark-grown seedlings. Conforming to the RNA gel-blot analysis, DWF4:beta-glucuronidase (GUS) histochemical analyses more precisely define the tissues that express the DWF4 gene. Examination of the endogenous levels of BRs in six and seven different tissues of wild type and brassinosteroid insensitive1-5 mutant, respectively, revealed that BRs are significantly enriched in roots, shoot tips, and joint tissues of roots and shoots. In addition, DWF4:GUS expression was negatively regulated by BRs. DWF4:GUS activity was increased by treatment with brassinazole, a BR biosynthetic inhibitor, and decreased by exogenous application of bioactive BRs. When DWF4:GUS was expressed in a different genetic background, its level was down-regulated in brassinazole resistant1-D, confirming that BRASSINAZOLE RESISTANT1 acts as a negative regulator of DWF4. Interestingly, in the brassinosteroid insensitive2/dwf12-1D background, DWF4:GUS expression was intensified and delocalized to elongating zones of root, suggesting that BRASSINOSTEROID INSENSITIVE2 is an important factor that limits DWF4 expression. Thus, it is likely that the DWF4 promoter serves as a focal point in maintaining homeostasis of endogenous bioactive BR pools in specific tissues of Arabidopsis.

Arabidopsis↗
PubMed2005

Grapes on steroids. Brassinosteroids are involved in grape berry ripening.

Fruit ripening is a unique plant developmental process with direct implications for our food supply, nutrition, and health. In contrast to climacteric fruit, where ethylene is pivotal, the hormonal control of ripening in nonclimacteric fruit, such as grape (Vitis vinifera), is poorly understood. Brassinosteroids (BRs) are steroidal hormones, essential for normal plant growth and development but not previously implicated in the ripening of nonclimacteric fruit. Here we show that increases in endogenous BR levels, but not indole-3-acetic acid (IAA) or GA levels, are associated with ripening in grapes. Putative grape homologs of genes encoding BR biosynthesis enzymes (BRASSINOSTEROID-6-OXIDASE and DWARF1) and the BR receptor (BRASSINOSTEROID INSENSITIVE 1) were isolated, and the function of the grape BRASSINOSTEROID-6-OXIDASE gene was confirmed by transgenic complementation of the tomato (Lycopersicon esculentum) extreme dwarf (dx/dx) mutant. Expression analysis of these genes during berry development revealed transcript accumulation patterns that were consistent with a dramatic increase in endogenous BR levels observed at the onset of fruit ripening. Furthermore, we show that application of BRs to grape berries significantly promoted ripening, while brassinazole, an inhibitor of BR biosynthesis, significantly delayed fruit ripening. These results provide evidence that changes in endogenous BR levels influence this key developmental process. This may provide a significant insight into the mechanism controlling ripening in grapes, which has direct implications for the logistics of grape production and down-stream processing.

Cloning, Molecular↗
PubMed2025

Rice ETHYLENE RESPONSE FACTOR 101 increases leaf angle by upregulating BRASSINOSTEROID UPREGULATED 1.

The leaf angle (LA) is a critical component of plant architecture that directly influences photosynthetic efficiency and grain yield. In the present study, we found that ETHYLENE RESPONSE FACTOR 101 (OsERF101), an APETALA2/ethylene response factor, plays a role in LA formation. A null mutation in OsERF101 resulted in reduced LA, whereas transgenic plants overexpressing OsERF101 (OsERF101-OEs) exhibited increased LA. OsERF101 increased the development of the adaxial lamina joint (LJ). Transactivation assays and reverse transcription-quantitative polymerase chain reaction (RT-qPCR) analysis indicated that OsERF101 activated BRASSINOSTEROID UPREGULATED 1 (OsBU1) transcription by directly binding to its promoter. However, OsERF101 expression was suppressed by exogenous brassinosteroid (BR) treatment and elevated endogenous brassinolide (BL) levels during LJ development. Additionally, OsERF101 downregulated the expression of BR biosynthesis genes, including Brassinosteroid-deficient dwarf2 (OsBRD2) and CYP90B2/OsDWARF4, leading to reduced levels of endogenous BL, the most active BR, in OsERF101-OEs. These findings suggested that OsERF101 mediates a negative feedback loop that balances endogenous BR levels and signaling. Collectively, rice plants have evolved diverse regulatory mechanisms involving OsERF101 to tune LA formation and optimize plant architecture finely.

Oryza↗
PubMed1997

Synthesis and molecular modeling: related approaches to progress in brassinosteroid research.

In the field of brassinosteroids, which are potent plant growth regulators, we have developed a quantitative structure-activity relationship study to develop knowledge from a structural point of view and to find out new requirement definitions. This will help identify other suitable active brassinosteroid derivatives with a good activity/synthetic cost ratio for further application in agriculture. The methodology used to achieve this goal represents a multidisciplinary study involving synthesis, molecular modeling calculations, and bioactivity evaluation. The influence of different molecular properties in the bioactivity of a set of synthetic compounds (i.e., molecular electrostatic potential and the ability to form H bonds) is discussed. The molecular electrostatic potential is expressed in terms of the electrostatic Carbó similarity index (CI) between brassinolide (1) and other brassinosteroids. We have found that the electrostatic charges of the functional groups play an important role in the description of the activity, as evidenced by its good correlation with the CI in most cases. Deviation from this rule could be explained by the H bonding abilities of some of these compounds, which we believe may play an essential role in binding to the natural receptors.

Brassinosteroids↗
PubMed1997

A putative leucine-rich repeat receptor kinase involved in brassinosteroid signal transduction.

Brassinosteroids are a class of growth-promoting regulators that play a key role throughout plant development. Despite their importance, nothing is known of the mechanism of action of these steroid hormones. We describe the identification of 18 Arabidopsis dwarf mutants that are unable to respond to exogenously added brassinosteroid, a phenotype that might be expected for brassinosteroid signaling mutants. All 18 mutations define alleles of a single previously described gene, BRI1. We cloned BRI1 and examined its expression pattern. It encodes a ubiquitously expressed putative receptor kinase. The extracellular domain contains 25 tandem leucine-rich repeats that resemble repeats found in animal hormone receptors, plant disease resistance genes, and genes involved in unknown signaling pathways controlling plant development.

Arabidopsis↗
PubMed2002

BAK1, an Arabidopsis LRR receptor-like protein kinase, interacts with BRI1 and modulates brassinosteroid signaling.

Brassinosteroids regulate plant growth and development through a protein complex that includes the leucine-rich repeat receptor-like protein kinase (LRR-RLK) brassinosteroid-insensitive 1 (BRI1). Activation tagging was used to identify a dominant genetic suppressor of bri1, bak1-1D (bri1-associated receptor kinase 1-1Dominant), which encodes an LRR-RLK, distinct from BRI1. Overexpression of BAK1 results in elongated organ phenotypes, while a null allele of BAK1 displays a semidwarfed phenotype and has reduced sensitivity to brassinosteroids (BRs). BAK1 is a serine/threonine protein kinase, and BRI1 and BAK1 interact in vitro and in vivo. 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.

Alleles↗
PubMed2002

Action of brassinosteroids in the cotton leafworm Spodoptera littoralis.

The two native plant hormones 24-epibrassinolide and 24-epicastasterone showed 50% competition for binding at IC(50) of 1-3.6 microM with [(3)H]ponasterone A using cultured imaginal wing discs from last-instar larvae of the cotton leafworm, Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae). However, culture of imaginal wing discs in different concentrations of brassinosteroids, even up to 100 microM, demonstrated no induction of evagination. In contrast, 20E and the non-steroidal agonist RH-5992 competed respectively about 23- and 42-fold more effectively with labeled ponasterone A, and their ability (EC(50)) to induce disc evagination in vitro was 158 and 87 nM, respectively. Injection of 10 microg of brassinosteroids in newly-moulted last-instar larvae did not cause mortality above controls; higher mortalities were scored when brassinosteroids were injected late in the last instar.

Animals↗
PubMed2003

Brassinosteroid and systemin: two hormones perceived by the same receptor.

Brassinosteroids, coordinating developmental events, and systemin, inducing systemic wound responses to attacks by insect pests, are newly recognized plant hormones that are perceived by plasma membrane-localized leucine-rich repeat receptor kinases. The recent characterization of the brassinosteroid receptor BRI1 from tomato revealed that this protein is identical to the previously isolated SR160 systemin receptor, strongly suggesting that both brassinosteroid and systemin signalling use the same surface receptor.

Animals↗
PubMed2000

Brassinosteroid signal transduction: still casting the actors.

Significant advances in the genetic dissection of brassinosteroid biosynthesis and signaling have been made during the past few years. Genetic and biochemical data have helped to elucidate the pathways of biosynthesis of brassinolide, the most active brassinosteroid. In addition, several models have been put forward for the perception of brassinolide by its putative receptor, BRI1, a ubiquitously expressed plasma membrane localized protein kinase. These studies provide the basic framework for future analysis of brassinosteroid signaling.

Brassinosteroids↗
PubMed1999

The sax1 dwarf mutant of Arabidopsis thaliana shows altered sensitivity of growth responses to abscisic acid, auxin, gibberellins and ethylene and is partially rescued by exogenous brassinosteroid.

Genetic approaches using Arabidopsis thaliana aimed at the identification of mutations affecting events involved in auxin signalling have usually led to the isolation of auxin-resistant mutants. From a selection screen specifically developed to isolate auxin-hypersensitive mutants, one mutant line was selected for its increased sensitivity to auxin (x 2 to 3) for the root elongation response. The genetic analysis of sax1 (hypersensitive to abscisic acid and auxin) indicated that the mutant phenotype segregates as a single recessive Mendelian locus, mapping to the lower arm of chromosome 1. Sax1 seedlings grown in vitro showed a short curled primary root and small, round, dark-green cotyledons. In the greenhouse, adult sax1 plants were characterized by a dwarf phenotype, delayed development and reduced fertility. Further physiological characterization of sax1 seedlings revealed that the most striking trait was a large increase (x 40) in ABA-sensitivity of root elongation and, to a lesser extent, of ABA-induced stomatal closure; in other respects, hypocotyl elongation was resistant to gibberellins and ethylene. These alterations in hormone sensitivity in sax1 plants co-segregated with the dwarf phenotype suggesting that processes involved in cell elongation are modified. Treatment of mutant seedlings with an exogenous brassinosteroid partially rescued a wild-type size, suggesting that brassinosteroid biosynthesis might be affected in sax1 plants. Wild-type sensitivities to ABA, auxin and gibberellins were also restored in sax1 plants by exogenous application of brassinosteroid, illustrating the pivotal importance of the BR-related SAX1 gene.

Abscisic Acid↗
PubMed2005

Physical restriction of pods causes seed size reduction of a brassinosteroid-deficient faba bean (Vicia faba).

BACKGROUND AND AIMS: A brassinosteroid-deficient mutant faba bean (Vicia faba 'Rinrei') shows dwarfism in many organs including pods and seeds. 'Rinrei' has normal-sized seeds together with dwarf seeds, suggesting that dwarfism in the seed may be indirectly caused by brassinosteroid deficiency. The mechanism of seed size reduction in this mutant was investigated. METHODS: The associations between seed orientation in the pod, seed numbers per pod and pod lengths with seed sizes were analysed in 'Rinrei' and the wild-type plant. KEY RESULTS: 'Rinrei' seeds are tightly arranged in pods containing two or three seeds. Seed size decreased as the number of seeds per pod increased or as the length of the pod decreased. Where no physical restriction occurred between seeds in a pod, the wild-type faba bean seeds had a nearly constant size regardless of seed number per pod or pod length. 'Rinrei' seeds in pods containing single seeds were the same size as wild-type seeds. Brassinolide treatment increased the seed size and the length of pods containing three seeds in 'Rinrei'. CONCLUSION: Seed size of 'Rinrei' is mainly regulated through a reduction of pod length due to brassinosteroid deficiency; physical restriction within pods causes a reduction in seed size. These results suggest a possible mechanism for increasing faba bean yields to optimal levels.

Brassinosteroids↗
PubMed2002

Promotion of transcript accumulation of novel Zinnia immature xylem-specific HD-Zip III homeobox genes by brassinosteroids.

We isolated three novel homeobox genes (ZeHB-10, -11 and -12) from Zinnia elegans to elucidate the molecular mechanism underlying vascular system formation. ZeHB-10, -11 and -12 encode for HD-Zip proteins of the class III to which Arabidopsis Athb-8, -9, -14, -15 and IFL1 belong. In situ hybridization analysis demonstrated that the ZeHB-10, -11 and -12 mRNAs accumulated preferentially in procambium and immature xylem cells in 14-day-old plants. Transcripts for the three genes also accumulated in cultured Zinnia cells in a xylogenesis-specific manner. The accumulation of transcripts for all of ZeHB-10, -11 and -12 in cultured Zinnia cells was suppressed strongly by uniconazole, an inhibitor of brassinosteroid synthesis, and such suppression was reversed by the addition of brassinolide, a biologically active brassinosteroid. Thus the expression of ZeHB-10, -11 and -12 may be regulated by endogenous levels of brassinosteroids. Taken together with the fact that ZeHB-10, -11 and -12 proteins can bind to each other in yeast, the roles of HD-Zip III genes in vascular development are discussed.

Arabidopsis Proteins↗