Brassinolide
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Brassinolide is a
plant hormone Plant hormones (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of Organ (anat ...
. The first isolated
brassinosteroid Brassinosteroids (BRs or less commonly BS) are a class of polyhydroxysteroids that have been recognized as a sixth class of plant hormones and may have utility as anticancer drugs for treating endocrine-responsive cancers by inducing apoptosis of ...
, it was discovered when it was shown that pollen from
rapeseed Rapeseed (''Brassica napus'' subsp. ''napus''), also known as rape and oilseed rape and canola, is a bright-yellow flowering member of the family Brassicaceae (mustard or cabbage family), cultivated mainly for its oil-rich seed, which naturall ...
(''Brassica napus'') could promote stem elongation and cell division. The biologically active component was isolated and named brassinolide.


Biosynthesis

The production of brassinolide begins with a closely related sterol called campesterol, which is found in the cell membrane. Initially, it is reduced by an enzyme called DET2. This is followed by a series of oxidation reactions, facilitated by cytochrome P-450 enzymes, which add hydroxyl groups to the molecule. The most biologically significant of these reactions is the C6 oxidation, where a ketone is formed at the C6 carbon position. This single reaction increases the biological activity of the molecule by a factor of 200. Depending on when this C6 oxidation occurs, it is referred to as either the early or late C6 oxidation pathway. Both of these synthetic pathways have been observed in
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organ ...
seedlings. It appears that the late C6 oxidation pathway predominates when the seedlings are exposed to light, while the early pathway is active in the absence of light. If the plant cannot perform C6 oxidation, it results in the "Dwarf phenotype," characterized by severe growth deficits. Finally, in Arabidopsis, the Baeyer-Villiger lactonization process occurs through the action of the two homologous enzymes CYP85A1 and CYP85A2, leading to the formation of brassinolide. Alternatively, there is a suggested synthetic pathway that starts from cholesterol, giving rise to C27 brassinosteroids.


Mechanism of Action


Signal Transduction

Brassinosteroids, particularly the potent brassinolide, play a crucial role in controlling various plant processes such as germination, aging, and the ability to withstand environmental and biological stresses. Because of this, researchers from around the world have extensively studied model organisms like
Catharanthus roseus ''Catharanthus roseus'', commonly known as bright eyes, Cape periwinkle, graveyard plant, Madagascar periwinkle, old maid, pink periwinkle, rose periwinkle, is a perennial species of flowering plant in the family Apocynaceae. It is native and en ...
and
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organ ...
since they were first isolated in 1979. These organisms have been thoroughly examined, from how they receive brassinolide signals to how these signals affect gene expression. In Arabidopsis, the process begins with the BRI1 receptor (Brassinosteroid Insensitive 1 receptor). This receptor is a type of protein called a leucine-rich receptor
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 ...
and allows brassinolide to attach to it from outside the cell. This binding causes a change in the receptor's shape, and it then interacts with another protein called BRI1 associated receptor kinase 1 (BAK1). This interaction results in both proteins being chemically modified by the addition of phosphate groups in a process called phosphorylation. This, in turn, sets off a chain reaction within the cell, activating some proteins and inhibiting others, including various kinases,
phosphatase In biochemistry, a phosphatase is an enzyme that uses water to cleave a phosphoric acid Ester, monoester into a phosphate ion and an Alcohol (chemistry), alcohol. Because a phosphatase enzyme catalysis, catalyzes the hydrolysis of its Substrate ...
s, and
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription (genetics), transcription of genetics, genetic information from DNA to messenger RNA, by binding t ...
s. Among the activated proteins are the BR signaling kinases (BSK1, BSK2, and BSK3). Their activation, in turn, activates the phosphatase BRI1 suppressor1 (BSU1), which removes a phosphate group from another protein called brassinosteroid insensitive 2 (BIN2). Removing this phosphate group inactivates BIN2, an important enzyme. As a result, protein phosphotase 2A (PP2A) can remove phosphate groups from two transcription factors, brassinazole-resistant-1 (BZR1) and BRI1-EMS-suppressor-1 (BES1), allowing them to accumulate within the cell's
nucleus Nucleus (: nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucleu ...
. There, they control the expression of specific target genes, which are involved in various cellular processes. However, when there's no brassinolide around, a regulator called BRI1 kinase inhibitor (BKI1) prevents the BRI1 receptor from interacting with the BAK1 co-receptor. This prevents the activation of BIN2, causing BZR1 and BES1 to be chemically modified by adding phosphate groups. These modified transcription factors then interact with a protein called 14-3-3 and accumulate in the cell's cytoplasm. Eventually, they are broken down and degraded by a 26S proteasome. In this way, BIN2 kinase serves as an essential negative regulator, dampening the activity of the central transcription factors BES1 and BZR1.


Homeostasis

The two transcription factors BES1 and BZR1 regulate a large number of genes involved in the synthesis of hormones, growth processes and stress response. In addition, there is inhibition of BR biosynthesis, which is evident as early as 15 minutes after brassinolide treatment. Thus, the expression of several biosynthetic genes such as the CPD, DWF4 and CYP85A2 gene is inhibited. These encode for brassinolide biosynthetic enzymes, thus the CPD gene encodes for the cytochrome P450 monooxygenase, which in the late C6 oxidation pathway oxidizes 6-deoxo-cathasterone to 6-deoxo teasterone and in the early C6 oxidation step catalyzes the reaction of cathasterone to teasterone. The enzyme CYP85A2 catalyzes the final biosynthesis step namely the Baeyer-Villiger lactonization of castasterone to brassinolide. This negative feedback loop ensures
homeostasis In biology, homeostasis (British English, British also homoeostasis; ) is the state of steady internal physics, physical and chemistry, chemical conditions maintained by organism, living systems. This is the condition of optimal functioning fo ...
of the hormone brassinolide.


Cell growth

Brassinolide induces genes that promote growth such as TCH4 and SAUR-Ac. The gene SAUR-Ac stands for ''small auxin upregulated RNAs'', these belong to the auxin hormone induced genes and provide cell expansion. Thus, SAURs inhibit the phosphatase PP2C-D so that the plasma membrane H+-ATPase is not dephosphorylated. The active phosphorylated proton pump can thus establish an electrochemical gradient in the cell wall. The acidity increases from 6 to 4.5-6 and according to the acid growth hypothesis, it ensures the activation of expansins that cleave the bond of cellulose and
hemicellulose A hemicellulose (also known as polyose) is one of a number of heteropolymers (matrix polysaccharides), such as arabinoxylans, present along with cellulose in almost all embryophyte, terrestrial plant cell walls. Cellulose is crystalline, strong, an ...
. TCH4 was identified as xyloglucan endotransglycosylase (XETs) by sequence analysis and enzyme activity. Its main function is the modification of cell walls. Thus, hemicellulose is composed of xyloglucans, which is built from 1,4-β-linked glucose polymers with lateral 1,6-β-linked xylene residues. The xylogucans can form
hydrogen bond In chemistry, a hydrogen bond (H-bond) is a specific type of molecular interaction that exhibits partial covalent character and cannot be described as a purely electrostatic force. It occurs when a hydrogen (H) atom, Covalent bond, covalently b ...
s with the cellulose microfibrils and thus structurally stabilize the cell wall. This means XET can modify the cell wall structure. It cleaves xyloglucan molecules, stores some of the energy, and then consumes it again after expansion for linking. Thus, during cell migration expansion, XET can further loosen the cell wall, which provides for the absorption of water. The resulting internal pressure (
turgor Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall. It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilib ...
) is compensated for by the cell wall expansion, so that after re-linking the result is an expanded cell.


Cross talk with other phytohormones


Gibberellin

Brassinolide and
gibberellin Gibberellins (GAs) are plant hormones that regulate various Biological process, developmental processes, including Plant stem, stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. They are one of th ...
are both interdependent. BZR1 in Arabidsopsis inhibits DELLAs a negative regulator of gibberellin transduction. Thus, the activity at the promoter for binding is reduced, so gibberellin promotes cell elongation. And DELLAs interaction provides a decrease in BZR1 binding affinity to DNA.


Jasmonic acid

Brassinolide inhibits
jasmonic acid Jasmonic acid (JA) is an organic compound found in several plants including jasmine. The molecule is a member of the jasmonate class of plant hormones. It is biosynthesized from linolenic acid by the octadecanoid pathway. It was first isolated ...
(JA) induced pathogen defensive for an ideal trade-off between growth and defense. Here, the transcription factor BES1 inhibits gene expression of PDF1.2a and PDF1.2b to reduce that of antimicrobial protein defensins. Furthermore, BES1 interacts with the transcription factors MYBs and reduces their activity to reduce glucosinolate (GS) biosynthesis, which is an important precursor for defense substances against predators.


References

{{reflist Plant hormones Steroids Epsilon-lactones Plant growth regulators