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Genetic engineering for increasing tolerance to abiotic stresses has come in due to the progress made in exploring and understanding plant abiotic stress responses at whole-plant( physiological, biochemical, cellular and molecular levels). Plant molecular biology is a fast-expanding research frontier of our times. This important branch of science has given several clues in understanding how plants respond under stressful regimes. A great deal of success has been achieved in unveiling gene/protein alterations associated with preparation of plants against the abiotic stresses. In parallel, major progress has been made in the characterization of stress-related promoters and transcription factors as well as stress signalling components.
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2.local response and gene inactivation:
Alterations in secondary metabolites/pathways Cessation of cell cycle Synthesis of pathogen related proteins Accumulation of benzoic and salicylic acid Production of ethylene and jasmonic acid Fortification of cell-walls( lignin,PGIPs,HRGPs)
121,000 20,100
42,600 2,830
61,300 16,200
50.7 80.6
Adaptations/Aclimation evolutionary improvements/adjustment of individual organisms by changing its homeostasis (e.g. Spinach (Osmotic adjustment) ,Black spruce (freezing tolerance) Desert Ephemerals escape drought by germinating and completing their life-cycle while enough water is available (short lived-4-8 weeks)
Saguaro
Pictorial-adaptaions/aclimation
Spinach
Black spruce
The word ephemeral means transitory or quickly fading (e.g.-Trillium grandifolium &Thalictrum thalicroides)
Plants
Physiological & developmental events
Cont
Transcriptional activation of gene expression suggests that accumulation of gene products is also influenced by post transcriptional regulatory mechanisms that increase the amount of specific m RNAs,enhance translation,stabilizing proteins,alter protein activity or some combination of these Using molecular genetic techniques,scientists have started to dissect plant responses associated with exposure to specific stresses.
Pressure Potential
Pressure potential is based on mechanical pressure, and is an important component of the total water potential within plant cells. Pressure potential is increased as water enters a cell. As water passes through the cell wall and cell membrane, it increases the total amount of water present inside the cell, which exerts an outward pressure that is retained by the structural rigidity of the cell wall. By creating this pressure, the plant can maintain turgor, which allows the plant to keep its rigidity. Without turgor, plants lose structure and wilt. The pressure potential in a living plant cell is usually positive. In plasmolysed cells, pressure potential is almost zero. Negative pressure potentials occur when water is pulled through an open system such as a plant xylem vessel. Withstanding negative pressure potentials (frequently called tension) is an important adaptation of xylem vessels. Defined in units of pressure(megapascals) rather than energy
Solute potential
Pure water is usually defined as having a solute potential () of zero, and in this case, solute potential can never be positive. The relationship of solute concentration (in molarity) to solute potential is given by the van 't Hoff equation: = MiRT where M is the concentration in molarity of the solute, i is the van 't Hoff factor, the ratio of amount of particles in solution to amount of formula units dissolved, R is the ideal gas constant, and T is the absolute temperature. Cont..
when a solute is dissolved in water, water molecules are less likely to diffuse away via osmosis than when there is no solute. A solution will have a lower and hence more negative water potential than that of pure water. Furthermore, the more solute molecules present, the more negative the solute potential is. Solute potential has important implication for many living organisms. If a living cell with a smaller solute concentration is surrounded by a more concentrated solution, the cell will tend to lose water to the more negative water potential (w) of the surrounding environment. This is often the case for marine organisms living in sea water and halophytic plants growing in saline environments. In the case of a plant cell, the flow of water out of the cell may eventually cause the plasma membrane to pull away from the cell wall, leading to plasmolysis. It can be measured in plant cells using the Pressure bomb. Most plants, however, have the ability to increase solute inside the cell to drive the flow of water into the cell and maintain turgor. This effect can be used to power an osmotic power plant
Cont..
When water is in contact with solid particles (e.g., clay or sand particles within soil), adhesive intermolecular forces between the water and the solid can be large and important. The forces between the water molecules and the solid particles in combination with attraction among water molecules promote surface tension and the formation of menisci within the solid matrix. Force is then required to break these menisci. The magnitude of matrix potential depends on the distances between solid particlesthe width of the menisci and the chemical composition of the solid matrix. In many cases, matrix potential can be quite large and comparable to the other components of water potential . It is worth noting that matrix potentials are very important for plant water relations. Strong (very negative) matrix potentials bind water to soil particles within very dry soils. Plants then create even more negative matrix potentials within tiny pores in the cell walls of their leaves to extract water from the soil and allow physiological activity to continue through dry periods. Germinating seeds have a very negative matrix potential. This causes water uptake in even somewhat dry soils and hydrates the dry seed.
Osmotic stress
Under osmotic stress, accumulation of osmotically active compounds called osmolytes occur to lower the osmotic potential. These molecules are not highly charged, but are polar, highly soluble and have a larger hydration shell. Since s is a function of total no. of solutes in a given vol. of water, a remarkable array of organic compounds and inorganic ions contribute to the s In principle ,OA involve metabolic changes that alter rates of ion uptake, decrease assimilation of low mol.mass organic compounds, or enhance their synthesis
Glycine betain accumulation is regulated by the rates of its synthesis and transport
Ethanol amine ATPADP Ethanol amine kinase Phosphoethanolamine/phosphatidylcholine s-adenosylmethionine phospho base N-methyltransferase s-adenosylhomocysteine Phosphocholine -Pi Phosphocholine phosphatase Choline O2+2H+ +2Fdxox Choline monooxygenase Betain aldehyde NAD+---NADH Betain aldehyde dehydrogenase Glycine betain
Phosphomannose isomerase
Mannose-6-Po4 Mannose -6 Po4 reductase NADPH----NADP Mannitol-1 po4 Mannitol-1 Po4 phosphatse -Pi Mannitol
SOS signaling pathway for ion homeostasis under salt stress in Arabidopsis.
Salt stress elicited Ca2+ signals are perceived by SOS3, which activates the protein kinase SOS2. Activated SOS2 phosphorylates SOS1, a plasma membrane Na+ /H+ antiporter, which then transports Na + out of the cytosol. The transcript level of SOS1 is regulated by the SOS3-SOS2 kinase complex. SOS2 also activates the tonoplast Na + /H + antiporter that sequesters Na + into the vacuole. Na entry into the cytosol through the Na + transporter HKT1 may also be restricted by SOS2. ABI1 regulates the gene expression of NHX1, while ABI2 interacts with SOS2 and negatively regulates ion homeostasis either by inhibiting SOS2 kinase activity or the activities of SOS2 targets. Double arrow indicates SOS3-independent and SOS2-dependent pathway.
Generation and scavenging of superoxide radical and hydrogen peroxide, and hydroxyl radical-induced lipid peroxidation and glutathione peroxidase-mediated lipid (fatty acid) stabilization
Freezing stress
Some plants can acclimate to subfreezing temperatures- called cold acclimation A primary function of freeze- tolerance mechanism is membrane stabilization Roles of the osmolytes and antifreeze proteins that accumulate in promoting freezing tolerance remain poorly understood Freezing tolerance involves changes in gene expression
Wetland plants- sweet flag, rice grass, coral tree, rice, wild rice, golden dock, barnyard grass Flood-tolerant plants- Arabidopsis, barnyard grass,oat,potato,wheat,corn Flood- sensitive plants- soy bean,tomato,pea
Structural adaptations
Aerenchyma (maize)-continuous,columnar intracellular spaces formed in root cortical tissues Lenticels (marshell)-opening in the periderm that allow gaseous exchange Pneumatophores (mangrove)-shallow roots that grow with negative geotrophy out of aquatic environment
Oxidative stress
Tropospheric ozone is linked to oxidative stress in plants Ozone causes oxidative damage to biomolecules Chloroplasts are susceptible to ozone induced damage Increased synthesis of antioxidants and antioxidant enzymes can improve tolerance to oxidative stress Oxidative stress or ozone can interact with plant hormones such as SA and ethylene to produce plant responses
Heat Stress
Heat stress alters cellular functions Plants can acclimate to heat stress HSPs are conserved among different organisms Five classes of HSPs are defined according to size Expression of many HSPs is controlled by a transcription factor that recognizes a conserved promotor sequences
Classes of HSPs
Protein Class HSP100 HSP90 HSP70 Name Hsp104(yeast) ClpB &ClpA ( E.coli) Hsp90 GRP94 Dnak(E.coli) BiP & GRP SSA1-4(yeast) SSB1-2(yeast) SSC1(yeast) KAR2(yeast) Chaperonin 60 groEL(E.coli) GroES(E.coli) Size (kDa) 100-114 80-94 69-71 ER Cytoplasm Cytoplasm Mitochondria ER 60 57 10 15-30 Chloroplast & mitochondria Location cytoplasm Cytoplasm ER
HSP60
smHSP
Cytoplasm,chloroplast,ER, mitochondria
ROS-defense-mechanism
H2O2 is directly toxic to pathogen & in presence of iron give rise to highly reactive hydroxyl radicle or may contribute to structural reinforcement of plant cell walls either by cross- linking various hydroxy-proline and proline- rich glycoproteins to the polysaccharide matrix or by increasing the rate of lignin polymerization by peroxidase enzyme activity and both would make the plant cell wall more resistant to microbial penetration and enzymatic degradation Signaling role of ROS: H2O2 induces benzoic acid 2hydroxylase(BA 2-H) enzyme activity for SA biosynthesis H2O2 also induces genes for protein involved in certain cell protection mechanism(e.g.glutathione s- reductase. ROS production also alter the redox balance in responding cells (e.g.-specific plant transcription factors are redox regulated.