Endogenous opiates: 1981

This paper is the fourth of an annual series reviewing the research concerning the endogenous opiate peptides. This installment covers only work published during 1981 and attempts to provide a comprehensive, but not exhaustive, survey of the area. Previous papers in the series have dealt with research done before 1981. Topics concerning endogenous opiates reviewed here include a delineation of their receptors, their distribution, their precursors and degradation, behavioral effects resulting from their administration, their possible involvement in physiological responses, and their interactions with other peptides and hormones. Due to the burgeoning literature in this field, the comprehensive nature of this review in the future will be limited to considerations of behavioral phenomena related to the endogenous opiates.     

Brassinosteroid-Mediated Stress Responses

Brassinosteroids (BRs) are a group of naturally occurring plant steroidal compounds with wide-ranging biological activity that offer the unique possibility of increasing crop yields through both changing plant metabolism and protecting plants from environmental stresses. In recent years, genetic and biochemical studies have established an essential role for BRs in plant development, and on this basis BRs have been given the stature of a phytohormone. A remarkable feature of BRs is their potential to increase resistance in plants to a wide spectrum of stresses, such as low and high temperatures, drought, high salt, and pathogen attack. Despite this, only a few studies aimed at understanding the mechanism by which BRs promote stress resistance have been undertaken. Studies of the BR signaling pathway and BR gene-regulating properties indicate that there is cross-talk between BRs and other hormones, including those with established roles in plant defense responses such as abscisic acid, jasmonic acid, and ethylene. Recent studies aimed at understanding how BRs modulate stress responses suggest that complex molecular changes underlie BR-induced stress tolerance in plants. Analyses of these changes should generate exciting results in the future and clarify whether the ability of BRs to increase plant resistance to a range of stresses lies in the complex interactions of BRs with other hormones. Future studies should also elucidate if BRI1, an essential component of the BR receptor, directly participates in stress response signaling through interactions with ligands and proteins involved in plant defense responses.     

Brassinosteroid-mediated stress tolerance in Arabidopsis shows interactions with abscisic acid,ethylene and salicylic acid pathways

Background  

Brassinosteroids (BRs) play crucial roles in plant development and also promote tolerance to a range of abiotic stresses. Although much has been learned about their roles in plant development, the mechanisms by which BRs control plant stress responses and regulate stress-responsive gene expression are not fully known. Since BR interacts with other plant hormones, it is likely that the stress tolerance conferring ability of BR lies in part in its interactions with other stress hormones.     

生长素与油菜素甾醇相互作用机制的研究进展

植物生长发育是一个复杂、精细的调控过程,涉及细胞、组织和器官间多层次的信息交流,激素在其间发挥了重要调控作用．生长素和油菜素甾醇（BR）都能促进植物伸长,随着对其作用机制研究的深入,人们发现它们协同调控很多生理过程,对二者作用机制和信号转导的相互作用研究也成为激素研究领域的热点之一．对生长素和BR转导途径的揭示及直接下游基因的大规模发掘为二者通过相互作用调控不同生理过程的机制研究提供了重要线索．生长素和BR的相互作用涉及到下游基因转录的调控、信号组分互作以及合成代谢和极性运输等多层次的调控．     

Evaluation of biological activity of new synthetic brassinolide analogs

The responses of plants to exogenous treatment with new synthetic brassinosteroids (BRs) were assessed and compared with the activity of natural 24-epibrassinolide (24-EPI). Morphological experiments on plants of pea and flax showed that the boundary between stimulatory and inhibitory concentrations of individual BRs and 24-EPI used is very narrow and differs also with the plant species. Moreover brassinosteroids can exhibit effects similar to various other plant hormones. This was proven also in our experiments, where auxin, anti-auxin and cytokinin like effects were achieved by BRs application. One of the explanations of the different morphological effects could be the influence of brassinosteroid application on the level of endogenous hormones. There are changes in the levels of indole-3-acetic acid, 6-benzylaminopurine, trans-zeatin and dihydrozeatin in rape and wheat plants caused by BR 4 and 24-EPI application, but there is no general trend explaining unequivocally their influence. The fact that all tested BRs significantly increased the dry weight accumulation in comparison with non-treated reference rape plants can be accounted for the known BRs characteristics to avoid biotic stresses.     

Brassinosteroids Regulate Growth in Plants Under Stressful Environments and Crosstalk with Other Potential Phytohormones

Brassinosteroids (BRs) are an important group of plant steroidal hormones that are actively involved in a myriad of key growth and developmental processes from germination to senescence. Moreover, BRs are known for their effective role in alleviation of stress-induced changes in normal metabolism via the activation of different tolerance mechanisms. Efforts to improve plant growth through exogenous application of BRs (through different modes such as foliar spray, presowing seed treatment, or through root growing medium) have gained considerable ground world over. It has been widely demonstrated that the exogenous application of BRs to stressed plants imparts the stress tolerance mechanisms. In BR-induced regulation of physio-biochemical processes in plants, interaction (crosstalk) of BRs with other phytohormones has been reported. This crosstalk may fine-tune the effective roles of other hormones in regulating stress tolerance. The multifaceted role of BRs consolidated so far has reflected their immense potential to help plants in counteracting the stress-induced changes. The effects of introgression and up- and down-regulation of BR-related genes reported so far to improve crop productivity have been presented here. Strong evidence exists that BRs are involved in signal transduction particularly in the regulation of the mitogen-activated protein kinase (MAPK) cascade, which in turn is involved in controlled development, cell death, and the perception of pathogen-associated molecular pattern (PAMP) signaling. How far BRs are involved in signal transduction pathways operative under stressful environments has also been comprehensively discussed in this review.     

Physiological and molecular mechanisms of brassinosteroid-induced tolerance to high and low temperature in plants

Brassinosteroids (BRs) are plant hormones that were isolated for the first time in the 1970s. This group currently includes more than 70 compounds that differ in their structure and physiological activity. BRs are present in plants in a free form or in the form of conjugates. BRs are known as plant growth regulators, but they also play a role in the plant response to environmental stresses. In the case of plants that are exposed to low/high temperature, exogenous BRs can counteract growth inhibition and reduce biomass losses as well as increase plant survival. BRs show a multidirectional activity in regulating the metabolism of plants exposed to extreme temperatures. The following BRs actions can be distinguished: changes in membrane physicochemical properties, regulation of the expression of selected genes (including stress-responsive genes), as well as indirect effects on metabolism through other hormones or signalling molecules (such as hydrogen peroxide). This review summarizes the current knowledge about the effects of BRs on the physiological and biochemical processes that occur in plants during exposure to low or high temperatures.     

Brassinosteroids function in spikelet differentiation and degeneration in rice

Brassinosteroids(BRs) play crucial roles in many aspects of plant development. However, their function in spikelet differentiation and degeneration in rice(Oryza sativa L.) remains unclear. Here, we investigated the roles of these phytohormones in spikelet development in fieldgrown rice subjected to five different nitrogen(N)fertilization treatments during panicle differentiation. BR levels and expression of genes involved in BR biosynthesis and signal transduction were measured in spikelets. Pollen fertility and the number of differentiated spikelets were closely associated with 24-epicastasterone(24-epiCS) and28-homobrassinolide(28-homoBL) levels in spikelets.Enhanced BR biosynthesis and signal transduction, in response to N treatment, enhanced spikelet differentiation, reduced spikelet degeneration, and increased grain yield. Increases in proton-pumping ATPase activity, ATPconcentration, energy charge, and antioxidant system(AOS) levels were consistent with 24-epiCS and28-homoBL concentrations. Exogenous application of24-epiCS or 28-homoBL on young panicles induced a marked increase in endogenous 24-epiCS or 28-homoBL levels, energy charge, AOS levels, spikelet differentiation, and panicle weight. The opposite effects were observed following treatment with a BR biosynthesis inhibitor. Our findings indicate that, in rice, BRs mediate the effects of N fertilization on spikelet development and play a role in promoting spikelet development through increasing AOS levels and energy charge during panicle development.     

Human cytochromes P450

The cytochrome P450 proteins (CYPs) are a family of haem proteins resulting from expression of a gene super-family that currently contains around 1000 members in species ranging from bacteria through to plants and animals. In humans, about 40 different CYPs are present and these play critical roles by catalyzing reactions in: (a) the metabolism of drugs, environmental pollutants and other xenobiotics; (b) the biosynthesis of steroid hormones; (c) the oxidation of unsaturated fatty acids to intracellular messengers; and (d) the stereo- and regio-specific metabolism of fat-soluble vitamins. This review deals with aspects of cytochrome P450s of relevance to human physiology, biochemistry, pharmacology and medicine. Topics reviewed include: pharmacogenetics of CYPs, induction and inhibition of these haem proteins, their role in metabolism of endogenous compounds such as steroids and eicosanoids, the effect of disease on CYP function, CYPs and cancer, and CYPs as targets of antibodies in immune-mediated diseases.     

Brassinosteroids interact with auxin to promote lateral root development in Arabidopsis

Plant hormone brassinosteroids (BRs) and auxin exert some similar physiological effects likely through their functional interaction, but the mechanism for this interaction is unknown. In this study, we show that BRs are required for lateral root development in Arabidopsis and that BRs act synergistically with auxin to promte lateral root formation. BR perception is required for the transgenic expression of the beta-glucuronidase gene fused to a synthetic auxin-inducible promoter (DR5::GUS) in root tips, while exogenous BR promotes DR5::GUS expression in the root tips and the stele region proximal to the root tip. BR induction of both lateral root formation and DR5::GUS expression is suppressed by the auxin transport inhibitor N-(1-naphthyl) phthalamic acid. Importantly, BRs promote acropetal auxin transport (from the base to the tip) in the root. Our observations indicate that BRs regulate auxin transport, providing a novel mechanism for hormonal interactions in plants and supporting the hypothesis that BRs promote lateral root development by increasing acropetal auxin transport.     

Physiological Roles of Brassinosteroids in Early Growth of <Emphasis Type="Italic">Arabidopsis:</Emphasis> Brassinosteroids Have a Synergistic Relationship with Gibberellin as well as Auxin in Light-Grown Hypocotyl Elongation

We examined the physiological effects of brassinosteroids (BRs) on early growth of Arabidopsis. Brassinazole (Brz), a BR biosynthesis inhibitor, was used to elucidate the significance of endogenous BRs. It inhibited growth of roots, hypocotyls, and cotyledonous leaf blades dose-dependently and independent of light conditions. This fact suggests that endogenous BRs are necessary for normal growth of individual organs of Arabidopsis in both photomorphogenetic and skotomorphogenetic programs. Exogenous brassinolide (BL) promoted hypocotyl elongation remarkably in light-grown seedlings. Cytological observation disclosed that BL-induced hypocotyl elongation was achieved through cell enlargement rather than cell division. Furthermore, a serial experiment with hormone inhibitors showed that BL induced hypocotyl elongation not through gibberellin and auxin actions. However, a synergistic relationship of BL with gibberellin A₃ (GA₃) and indole-3-acetic acid (IAA) was observed on elongation growth in light-grown hypocotyls, even though gibberellins have been reported to be additive to BR action in other plants. Taken together, our results show that BRs play an important role in the juvenile growth of Arabidopsis; moreover, BRs act on light-grown hypocotyl elongation independent of, but cooperatively with, gibberellins and auxin.     

YCZ-18 Is a New Brassinosteroid Biosynthesis Inhibitor

Plant hormone brassinosteroids (BRs) are a group of polyhydroxylated steroids that play critical roles in regulating broad aspects of plant growth and development. The structural diversity of BRs is generated by the action of several groups of P450s. Brassinazole is a specific inhibitor of C-22 hydroxylase (CYP90B1) in BR biosynthesis, and the application use of brassinazole has emerged as an effective way of complementing BR-deficient mutants to elucidate the functions of BRs. In this article, we report a new triazole-type BR biosynthesis inhibitor, YCZ-18. Quantitative analysis the endogenous levels of BRs in Arabidopsis indicated that YCZ-18 significantly decreased the BR contents in plant tissues. Assessment of the binding affinity of YCZ-18to purified recombinant CYP90D1 indicated that YCZ-18 induced a typical type II binding spectrum with a K_d value of approximately 0.79 μM. Analysis of the mechanisms underlying the dwarf phenotype associated with YCZ-18 treatment of Arabidopsis indicated that the chemically induced dwarf phenotype was caused by a failure of cell elongation. Moreover, dissecting the effect of YCZ-18 on the induction or down regulation of genes responsive to BRs indicated that YCZ-18 regulated the expression of genes responsible for BRs deficiency in Arabidopsis. These findings indicate that YCZ-18 is a potent BR biosynthesis inhibitor and has a new target site, C23-hydroxylation in BR biosynthesis. Application of YCZ-18 will be a good starting point for further elucidation of the detailed mechanism of BR biosynthesis and its regulation.     

Arabidopsis brassinosteroid biosynthetic mutant <Emphasis Type="Italic">dwarf7-1</Emphasis>exhibits slower rates of cell division and shoot induction

Background  

Plant growth depends on both cell division and cell expansion. Plant hormones, including brassinosteroids (BRs), are central to the control of these two cellular processes. Despite clear evidence that BRs regulate cell elongation, their roles in cell division have remained elusive.     

C-23 hydroxylation by Arabidopsis CYP90C1 and CYP90D1 reveals a novel shortcut in brassinosteroid biosynthesis

Brassinosteroids (BRs) are biosynthesized from campesterol via several cytochrome P450 (P450)-catalyzed oxidative reactions. We report the functional characterization of two BR-biosynthetic P450s from Arabidopsis thaliana: CYP90C1/ROTUNDIFOLIA3 and CYP90D1. The cyp90c1 cyp90d1 double mutant exhibits the characteristic BR-deficient dwarf phenotype, although the individual mutants do not display this phenotype. These data suggest redundant roles for these P450s. In vitro biochemical assays using insect cell-expressed proteins revealed that both CYP90C1 and CYP90D1 catalyze C-23 hydroxylation of various 22-hydroxylated BRs with markedly different catalytic efficiencies. Both enzymes preferentially convert 3-epi-6-deoxocathasterone, (22S,24R)-22-hydroxy-5alpha-ergostan-3-one, and (22S,24R)-22-hydroxyergost-4-en-3-one to 23-hydroxylated products, whereas they are less active on 6-deoxocathasterone. Likewise, cyp90c1 cyp90d1 plants were deficient in 23-hydroxylated BRs, and in feeding experiments using exogenously supplied intermediates, only 23-hydroxylated BRs rescued the growth deficiency of the cyp90c1 cyp90d1 mutant. Thus, CYP90C1 and CYP90D1 are redundant BR C-23 hydroxylases. Moreover, their preferential substrates are present in the endogenous Arabidopsis BR pool. Based on these results, we propose C-23 hydroxylation shortcuts that bypass campestanol, 6-deoxocathasterone, and 6-deoxoteasterone and lead directly from (22S,24R)-22-hydroxy-5alpha-ergostan-3-one and 3-epi-6-deoxocathasterone to 3-dehydro-6-deoxoteasterone and 6-deoxotyphasterol.     

Brassinosteroid-regulated GSK3/Shaggy-like Kinases Phosphorylate Mitogen-activated Protein (MAP) Kinase Kinases,Which Control Stomata Development in Arabidopsis thaliana

Brassinosteroids (BRs) are steroid hormones that coordinate fundamental developmental programs in plants. In this study we show that in addition to the well established roles of BRs in regulating cell elongation and cell division events, BRs also govern cell fate decisions during stomata development in Arabidopsis thaliana. In wild-type A. thaliana, stomatal distribution follows the one-cell spacing rule; that is, adjacent stomata are spaced by at least one intervening pavement cell. This rule is interrupted in BR-deficient and BR signaling-deficient A. thaliana mutants, resulting in clustered stomata. We demonstrate that BIN2 and its homologues, GSK3/Shaggy-like kinases involved in BR signaling, can phosphorylate the MAPK kinases MKK4 and MKK5, which are members of the MAPK module YODA-MKK4/5-MPK3/6 that controls stomata development and patterning. BIN2 phosphorylates a GSK3/Shaggy-like kinase recognition motif in MKK4, which reduces MKK4 activity against its substrate MPK6 in vitro. In vivo we show that MKK4 and MKK5 act downstream of BR signaling because their overexpression rescued stomata patterning defects in BR-deficient plants. A model is proposed in which GSK3-mediated phosphorylation of MKK4 and MKK5 enables for a dynamic integration of endogenous or environmental cues signaled by BRs into cell fate decisions governed by the YODA-MKK4/5-MPK3/6 module.     

Brassinosteroid levels increase drastically prior to morphogenesis of tracheary elements

As the first step toward understanding the involvement of endogenous brassinosteroids (BRs) in cytodifferentiation, we analyzed biosynthetic activities of BRs in zinnia (Zinnia elegans L. cv Canary Bird) cells differentiating into tracheary elements. The results of feeding experiments suggested that both the early and late C6-oxidation pathways occur during tracheary element differentiation. Gas chromatography-mass spectrometry analysis revealed that five BRs, castasterone, typhasterol, 6-deoxocastasterone, 6-deoxotyphasterol, and 6-deoxoteasterone, actually existed in cultured zinnia cells and culture medium. Quantification of endogenous BRs in each stage of tracheary element differentiation by gas chromatography-mass spectrometry exhibited that they increased dramatically prior to the morphogenesis, which was consistent with the idea that BRs are necessary for the initiation of the final stage of tracheary element differentiation. Moreover, the proportion of each BR in culture medium was quite different from that in cells, suggesting that specific BRs are selectively secreted into medium and may function outside the cells.