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Brassinosteroids (BRs or less commonly BS) are a class of polyhydroxysteroids that have been recognized as a sixth class of
plant hormones Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pat ...
and may have utility as an anticancer drug for endocrine-responsive cancers to induce apoptosis and inhibit growth. These brassinosteroids were first explored during the 70s, when Mitchell et al. reported promotion in stem elongation and cell division by the treatment of organic extracts of rapeseed (''
Brassica napus Rapeseed (''Brassica napus ''subsp.'' napus''), also known as rape, or oilseed rape, is a bright-yellow flowering member of the family Brassicaceae (mustard or cabbage family), cultivated mainly for its oil-rich seed, which naturally contains ...
'') pollen. Brassinolide was the first isolated brassinosteroid in 1979, when pollen from ''
Brassica napus Rapeseed (''Brassica napus ''subsp.'' napus''), also known as rape, or oilseed rape, is a bright-yellow flowering member of the family Brassicaceae (mustard or cabbage family), cultivated mainly for its oil-rich seed, which naturally contains ...
'' was shown to promote stem elongation and cell divisions, and the biologically active molecule was isolated. The yield of brassinosteroids from 230 kg of ''Brassica napus'' pollen was only 10 mg. Since their discovery, over 70 BR compounds have been isolated from plants.


Biosynthesis

The BR is biosynthesised from
campesterol Campesterol is a phytosterol whose chemical structure is similar to that of cholesterol, and is one of the ingredients for E number E499. Natural occurrences Many vegetables, fruits, nuts, and seeds contain campesterol, but in low concentrat ...
. The biosynthetic pathway was elucidated by Japanese researchers and later shown to be correct through the analysis of BR biosynthesis mutants in ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter ...
'', tomatoes, and peas. The sites for BR synthesis in plants have not been experimentally demonstrated. One well-supported hypothesis is that all tissues produce BRs, since BR biosynthetic and signal transduction genes are expressed in a wide range of plant organs, and short distance activity of the hormones also supports this. Experiments have shown that long distance transport is possible and that the flow is from the base to the tips (acropetal), but it is not known if this movement is biologically relevant.


Hormonal activity

BRs have been shown to be involved in numerous plant processes: *Promotion of cell expansion and cell elongation; works with
auxin Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essenti ...
to do so. *It has an unclear role in cell division and cell wall regeneration. *Promotion of
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
differentiation; BR
signal transduction Signal transduction is the process by which a chemical or physical signal is transmitted through a cell as a series of molecular events, most commonly protein phosphorylation catalyzed by protein kinases, which ultimately results in a cellula ...
has been studied during vascular differentiation. *Is necessary for pollen elongation for
pollen tube A pollen tube is a tubular structure produced by the male gametophyte of seed plants when it germinates. Pollen tube elongation is an integral stage in the plant life cycle. The pollen tube acts as a conduit to transport the male gamete cells fr ...
formation. *Acceleration of
senescence Senescence () or biological aging is the gradual deterioration of functional characteristics in living organisms. The word ''senescence'' can refer to either cellular senescence or to senescence of the whole organism. Organismal senescence invol ...
in dying tissue cultured cells; delayed senescence in BR mutants supports that this action may be biologically relevant. *Can provide some protection to plants during chilling and drought stress. Extract from the plant ''Lychnis viscaria'' contains a relatively high amount of Brassinosteroids. ''
Lychnis viscaria ''Viscaria vulgaris'', the sticky catchfly or clammy campion, is a flowering plant in the family Caryophyllaceae. It is an upright perennial growing to in height. The leaves are lanceolate. The flowers, which are 20 mm across and bright rosy ...
'' increases the disease resistance of surrounding plants. 24-Epibrassinolide (EBL), a brassinosteroid isolated from ''
Aegle marmelos ''Aegle marmelos'', commonly known as bael (or ''bili'' or ''bhel''), also Bengal quince, golden apple, Japanese bitter orange, stone apple or wood apple, is a rare species of tree native to the Indian subcontinent and Southeast Asia. It is pre ...
'' Correa (Rutaceae), was further evaluated for the antigenotoxicity against
maleic hydrazide Maleic hydrazide, often known by the brand name Fazor is a plant growth regulator that reduces growth through preventing cell division but not cell enlargement. It is applied to the foliage of potato, onion, garlic and carrot crops to prevent sprou ...
(MH)-induced
genotoxicity Genotoxicity is the property of chemical agents that damage the genetic information within a cell causing mutations, which may lead to cancer. While genotoxicity is often confused with mutagenicity, all mutagens are genotoxic, but some genotoxic ...
in '' Allium cepa'' chromosomal aberration assay. It was shown that the percentage of chromosomal aberrations induced by maleic hydrazide (0.01%) declined significantly with 24-epibrassinolide treatment. BRs have been reported to counteract both abiotic and biotic stress in plants. Application of brassinosteroids to cucumbers was demonstrated to increase the
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run ...
and removal of pesticides, which could be beneficial for reducing the human ingestion of residual pesticides from non-organically grown vegetables. BRs have also been reported to have a variety of effects when applied to rice seeds (Oryza sativa L.). Seeds treated with BRs were shown to reduce the growth inhibitory effect of salt stress. When the developed plants fresh weight was analyzed the treated seeds outperformed plants grown on saline and non-saline medium however when the dry weight was analyzed BR treated seeds only outperformed untreated plants that were grown on saline medium. When dealing with tomatoes (''Lycopersicon esculentum'') under salt stress the concentration of chlorophyll a and chlorophyll b were decreased and thus pigmentation was decreased as well. BR treated rice seeds considerably restored the pigment level in plants that were grown on saline medium when compared to non-treated plants under the same conditions.


Signalling mechanism

BRs are perceived at the cell membrane by a co-receptor complex, comprising brassinosteroid insensitive-1 (BRI1) and BRI1-associated receptor kinase 1 (BAK1). BRI1 acts as a
kinase In biochemistry, a kinase () is an enzyme that catalyzes the transfer of phosphate groups from high-energy, phosphate-donating molecules to specific substrates. This process is known as phosphorylation, where the high-energy ATP molecule don ...
, but in the absence of BR its action is inhibited by another protein, BRI1 kinase inhibitor 1 (BKI1). When BR binds to the BRI1:BAK1 complex, BKI1 is released, and a
phosphorylation cascade A phosphorylation cascade is a sequence of signaling pathway events where one enzyme phosphorylates another, causing a chain reaction leading to the phosphorylation of thousands of proteins. This can be seen in signal transduction of hormone messag ...
is triggered which results in the de-activation of another kinase, brassinosteroid insensitive 2 (BIN2). BIN2 and its close homologues inhibit several
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The f ...
s. The inhibition of BIN2 by BR releases these transcription factors to bind to DNA and to enact certain developmental pathways.


Agricultural uses

BR might reveal to have a prominent interest in the role of horticultural crops. Based on extensive research BR has the ability to improve the quantity and quality of horticultural crops and protect plants against many stresses that can be present in the local environment. With the many advances in technology dealing with the synthesis of more stable synthetic analogues and the genetic manipulation of cellular BR activity, using BR in the production of horticultural crops has become a more practical and hopeful strategy for improving crop yields and success. The application of BR successfully alleviate drought stress and improve wheat growth under deficit irrigation system. It had further positive impacts on increasing plant growth parameters via their integral role in decreasing oxidative stress indicators. BR application has demonstrated efficacy against ''
Phytophthora infestans ''Phytophthora infestans'' is an oomycete or water mold, a fungus-like microorganism that causes the serious potato and tomato disease known as late blight or potato blight. Early blight, caused by ''Alternaria solani'', is also often called "po ...
'',
mildew Mildew is a form of fungus. It is distinguished from its closely related counterpart, mould, largely by its colour: moulds appear in shades of black, blue, red, and green, whereas mildew is white. It appears as a thin, superficial growth consi ...
on
cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated creeping vine plant in the Cucurbitaceae family that bears usually cylindrical fruits, which are used as culinary vegetables.viral diseases Viral means "relating to viruses" (small infectious agents). Viral may also refer to: Viral behavior, or virality Memetic behavior likened that of a virus, for example: * Viral marketing, the use of existing social networks to spread a mar ...
, and various others. BR could also help bridge the gap of the consumers' health concerns and the producers need for growth. A major benefit of using BR is that it does not interfere with the environment because they act in a natural way. Since it is a “plant strengthening substance” and it is natural, BR application would be more favorable than pesticides and does not contribute to the co-evolution of pests. In Germany, extract from the plant is allowed for use as a "plant strengthening substance."gas chromatography mass spectrometry and
bioassay A bioassay is an analytical method to determine the concentration or potency of a substance by its effect on living animals or plants (''in vivo''), or on living cells or tissues(''in vitro''). A bioassay can be either quantal or quantitative, dir ...
s. There are some bioassays that can detect BRs in the plant such as the bean second internode elongation assay and the rice leaf lamina inclination test.


References


External links

{{Plant_hormones Steroids Plant hormones Lactones Secondary alcohols