Polyamines (PAs) are small, positively charged, organic molecules that are ubiquitous in all living organisms. These are considered as one of the oldest group of substances known in biochemistry. There are three common types of polyamines, putrescine, spermidine, hermospermine according to structure, universal distribution in all cellular compartments, and presumed involvement in physiological activities. Polyamine is found in all cellular compartments and physiological activities due to their simple structures. The function of polyamine is very diverse in that it performs a key macromolecule to cellular membrane. Because of their essential roles in plant, mutation of polyamines can cause critical damage on plants. [1] Furthermore, some polyamines like putrescine inhibit biosynthetic activities in plants. The activity of polyamines can be categorized to some parts due to its signalling and growing activity.
Polyamines, which are the key compounds in plant physiology, play essential role in cell proliferation, growth and differentiation. They also carry out significant effects on embryo/fetus proliferation, implantation, embryonic diapause, placentation, angiogenesis and fetal development.
Even though polyamines carry out important roles like soothing and ameliorating in rice physiology, [2] its catabolic products can cause stress damage. Putrescine, a kind of polyamine, decreaseS phytochelatin synthesis at enzymatic and gene expression levels by increasing cadmium toxicity in rice. Polyamines provide protection against heavy metals such as inhibiting Cadmium uptake or its entry into the cells because exogenous polyamines are mainly allocated to the apoplast. Spermidine and spermine, kinds of polyamine, are found in reducing cadmium toxicity in rice. Nevertheless, putrescine leads to increasing hydrogen peroxide accumulation and inhibits root elongation in rice. In this case, huge amounts of polyamine-cadmium complexes form, so higher density of cadmium and putrescine decreases phytochelatin synthesis as well as its gene expression. Therefore, plants with decreasing phytochelatin activities can struggle with detoxification of metal ions like Zn, Cu, and Ni.
Polyamines are compounds that are involved in a complex signaling system and have a key role in the regulation of stress tolerance. [3] In the past, however, the main role of polyamines was originally assumed to be as direct protective compounds important under stress conditions because of their cationic nature (-) at physiological pH. This feature of polyamine makes it possible to interact with negatively charged macromolecules, stabilizing and protecting compounds under stress condition. These polyamines can bind to diverse types of proteins non-specifically, and this make it possible to the conjugation of polyamines to photosynthetic complexes and proteins is catalyzed by transglutaminase and leads to enhanced photosynthetic activity under stress conditions. [4]
In the case of Arabidopsis , [5] since polyamines perform essential roles for growth in all organisms, and most polyamine biosynthetic genes are present in at least two copies in plants, break-out of single gene mutants can cause critical stress in plant. For example, Arabidopsis showed much lower activity in seed development when a specific gene of was impaired. With both genes of pair impaired (acl5/spms), arabidopsis cannot perform enzyme activities anymore because it compromises thermospermine-spermine biosynthesis and causes hypersensitivity to NaCl and KCl. The mutant level of thermospermine-spermine can also disturb homeostasis of calcium ions leading to a differential response to salts.
With ubiquitous presence, absolute necessity for cell survival, and their presence in relatively large quantities, [6] it is difficult to explain the accurate roles of polyamines in plant stress. Especially, the mechanism of polyamines in plant stress is not fully identified. problems relating to the role of PAs in abiotic stress is the lack of our understanding of the mechanisms underlying their function. Furthermore, it is hard to generalize the polyamines are directly associated to stress condition because [7] there is often an immediate increase in polyamine levels in response to stress, but after a while polyamine levels decrease and resemble those of non-stressed plants, even if the stress conditions persist.
Putrescine is an organic compound with the formula (CH2)4(NH2)2. It is a colorless solid that melts near room temperature. It is classified as a diamine. Together with cadaverine, it is largely responsible for the foul odor of putrefying flesh, but also contributes to other unpleasant odors.
Glutathione is an organic compound with the chemical formula HOCOCH(NH2)CH2CH2CONHCH(CH2SH)CONHCH2COOH. It is an antioxidant in plants, animals, fungi, and some bacteria and archaea. Glutathione is capable of preventing damage to important cellular components caused by sources such as reactive oxygen species, free radicals, peroxides, lipid peroxides, and heavy metals. It is a tripeptide with a gamma peptide linkage between the carboxyl group of the glutamate side chain and cysteine. The carboxyl group of the cysteine residue is attached by normal peptide linkage to glycine.
The enzyme ornithine decarboxylase catalyzes the decarboxylation of ornithine to form putrescine. This reaction is the committed step in polyamine synthesis. In humans, this protein has 461 amino acids and forms a homodimer.
Plant hormones 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 size, pathogen defense, stress tolerance and reproductive development. Unlike in animals each plant cell is capable of producing hormones. Went and Thimann coined the term "phytohormone" and used it in the title of their 1937 book.
Brassinosteroids 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 cancer cells and inhibiting cancerous growth. These brassinosteroids were first explored during the 1970s when Mitchell et al. reported promotion in stem elongation and cell division by the treatment of organic extracts of rapeseed pollen. Brassinolide was the first brassinosteroid to be isolated in 1979, when pollen from Brassica napus 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.
Spermine is a polyamine involved in cellular metabolism that is found in all eukaryotic cells. The precursor for synthesis of spermine is the amino acid ornithine. It is an essential growth factor in some bacteria as well. It is found as a polycation at physiological pH. Spermine is associated with nucleic acids and is thought to stabilize helical structure, particularly in viruses. It functions as an intracellular free radical scavenger to protect DNA from free radical attack. Spermine is the chemical primarily responsible for the characteristic odor of semen.
Spermidine is a polyamine compound found in ribosomes and living tissues and having various metabolic functions within organisms.
Spermidine synthase is an enzyme that catalyzes the transfer of the propylamine group from S-adenosylmethioninamine to putrescine in the biosynthesis of spermidine. The systematic name is S-adenosyl 3-(methylthio)propylamine:putrescine 3-aminopropyltransferase and it belongs to the group of aminopropyl transferases. It does not need any cofactors. Most spermidine synthases exist in solution as dimers.
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 in 1957 as the methyl ester of jasmonic acid by the Swiss chemist Édouard Demole and his colleagues.
Sulfur assimilation is the process by which living organisms incorporate sulfur into their biological molecules. In plants, sulfate is absorbed by the roots and then transported to the chloroplasts by the transipration stream where the sulfur are reduced to sulfide with the help of a series of enzymatic reactions. Furthermore, the reduced sulfur is incorporated into cysteine, an amino acid that is a precursor to many other sulfur-containing compounds. In animals, sulfur assimilation occurs primarily through the diet, as animals cannot produce sulfur-containing compounds directly. Sulfur is incorporated into amino acids such as cysteine and methionine, which are used to build proteins and other important molecules.
The enzyme adenosylmethionine decarboxylase catalyzes the conversion of S-adenosyl methionine to S-adenosylmethioninamine. Polyamines such as spermidine and spermine are essential for cellular growth under most conditions, being implicated in many cellular processes including DNA, RNA and protein synthesis. S-adenosylmethionine decarboxylase (AdoMetDC) plays an essential regulatory role in the polyamine biosynthetic pathway by generating the n-propylamine residue required for the synthesis of spermidine and spermine from putrescein. Unlike many amino acid decarboxylases AdoMetDC uses a covalently bound pyruvate residue as a cofactor rather than the more common pyridoxal 5'-phosphate. These proteins can be divided into two main groups which show little sequence similarity either to each other, or to other pyruvoyl-dependent amino acid decarboxylases: class I enzymes found in bacteria and archaea, and class II enzymes found in eukaryotes. In both groups the active enzyme is generated by the post-translational autocatalytic cleavage of a precursor protein. This cleavage generates the pyruvate precursor from an internal serine residue and results in the formation of two non-identical subunits termed alpha and beta which form the active enzyme.
Spermine synthase is an enzyme that converts spermidine into spermine. This enzyme catalyses the following chemical reaction
Diamine acetyltransferase 1 is an enzyme that in humans is encoded by the SAT1 gene found on the X chromosome.
S-Adenosylmethioninamine is a substrate that is required for the biosynthesis of polyamines including spermidine, spermine, and thermospermine. It is produced by decarboxylation of S-adenosyl methionine.
A polyamine is an organic compound having more than two amino groups. Alkyl polyamines occur naturally, but some are synthetic. Alkylpolyamines are colorless, hygroscopic, and water soluble. Near neutral pH, they exist as the ammonium derivatives. Most aromatic polyamines are crystalline solids at room temperature.
Spermine oxidase (EC 1.5.3.16, PAOh1/SMO, AtPAO1, AtPAO4, SMO) is an enzyme with systematic name spermidine:oxygen oxidoreductase (spermidine-forming). This enzyme catalyses the following chemical reaction
Non-specific polyamine oxidase (EC 1.5.3.17, polyamine oxidase, Fms1, AtPAO3) is an enzyme with systematic name polyamine:oxygen oxidoreductase (3-aminopropanal or 3-acetamidopropanal-forming). This enzyme catalyses the following chemical reaction
Thimet oligopeptidases, also known as TOPs, are a type of M3 metallopeptidases. These enzymes can be found in animals and plants, showing distinctive functions. In animals and humans, they are involved in the degradation of peptides, such as bradykinin, neurotensin, angiotensin I, and Aβ peptide, helping to regulate physiological processes. In plants, their role is related to the degradation of targeting peptides and the immune response to pathogens through Salicylic Acid (SA)-dependent stress signaling. In Arabidopsis thaliana—recognized as a model plant for scientific studies—two thimet oligopeptidases, known as TOP1 and TOP2, have been identified as targets for salicylic acid binding in the plant. These TOP enzymes are key components to understand the SA-mediated signaling where interactions exist with different components and most of the pathways are unknown.
2-Phosphoglycolate (chemical formula C2H2O6P3-; also known as phosphoglycolate, 2-PG, or PG) is a natural metabolic product of the oxygenase reaction mediated by the enzyme ribulose 1,5-bisphosphate carboxylase (RuBisCo).
Barley is known to be more environmentally-tolerant than other cereal crops, in terms of soil pH, mineral nutrient availability, and water availability. Because of this, much research is being done on barley plants in order to determine whether or not there is a genetic basis for this environmental hardiness.
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