Rice Science ›› 2023, Vol. 30 ›› Issue (1): 15-35.DOI: 10.1016/j.rsci.2022.08.002
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M. Iqbal R. Khan1(
), Sarika Kumari1, Faroza Nazir1, Risheek Rahul Khanna1, Ravi Gupta2, Himanshu Chhillar1
Received:2022-05-04
Accepted:2022-08-04
Online:2023-01-28
Published:2022-11-11
Contact:
M. Iqbal R. Khan
About author:First author contact:This is an open access article under the CC BY-NC-ND license (
Peer review under responsibility of China National Rice Research Institute
M. Iqbal R. Khan, Sarika Kumari, Faroza Nazir, Risheek Rahul Khanna, Ravi Gupta, Himanshu Chhillar. Defensive Role of Plant Hormones in Advancing Abiotic Stress-Resistant Rice Plants[J]. Rice Science, 2023, 30(1): 15-35.
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| Plant hormone | Abiotic stress and its condition | Morphological, physiological, and molecular effects | Reference |
| Auxin | Drought stress SMC between 7.3% ± 0.5% and 6.7% ± 0.2% Heat stress (40.3 ºC ± 0.36 ºC and 37.7 ºC± 0.35 ºC) | Improve pollen viability, spikelet fertility, grain number per panicle, grain weight per panicle, 1000-grain weight, and grain yield per panicle; enhance chlorophyll content, delay senescence, and maintain greenness; lower lipid peroxidation and ROS accumulation while improving membrane stability; improve the expression levels of auxin signalling genes (TIR1, AFB2, SPL, TFL2 and HAF) | Sharma et al, |
| As stress (NaAsO2, 150 µmol/L) Se stress (Na2SeO4, 20 µmol/L) | I. Auxin (As and Se stress): Enhance root and shoot growths II. Auxin + Se (As stress): Improve chlorophyll content, proline and cysteine; lower protein content inhibition and DNA damage; reduce lipid peroxidation | Pandey and Gupta, | |
| Gibberellic acid (GA) | Drought stress (PEG6000, 15%) | Improve germination vigor, shoot length, root length, α-amylase activity, and soluble sugar content; induce the expression levels of α-amylase and expansion-related genes | Li J Z et al, 2019 |
| Salt stress (NaCl: 0, 50, 100, 150 and 200 mmol/L) | Enhance contents of proline, soluble sugar and protein as well as inorganic nutrients (K, Mg and Ca); increase panicle number, panicle length, grain yield per plant, 1000-grain weight, filled grain number per panicle, and harvest index | Misratia et al, | |
| Cold stress (10 ºC to 20 ºC) | Improve cold resistance and increase yield traits; promote seedling emergence and vigor | Chen et al, | |
| Abscisic acid (ABA) | Drought stress (PEG6000, 15%) | Enhance net photosynthesis, stomatal conductance, and transpiration rate; improve expression levels of chloroplast genes (OsPsbD1 and OsPsbD2); up-regulate expression levels of ABA biosynthesis genes (OsNCED2, OsNCED3, OsNCED4 and OsNCED5) | Teng et al, |
| Cold stress at 5 ºC | Improve the survival ratio | Lee et al, | |
| Salicylic acid (SA) | Salt stress (NaCl: 0, 100, 200, 300 and 400 mmol/L) | Enhance germination rate, root length, shoot length and plant height; improve the number of grains per panicle, number of filled grains per panicle, 1000-grain weight, the number of panicles per plant, and yield; increase contents of carbohydrate and protein along with minerals (Ca, P and K) while decreasing Na and Cl contents as well as Na+ and Cl- accumulation; enhance endogenous SA content | Jini and Joseph, |
| Cd stress (CdCl2, 50 µmol/L) | Improve root growth; reduce ROS level, and membrane damage; enhance SOD, POD and CAT activities as well as GSH, and AsA contents; improve non-protein thiols’ concentration | Guo et al, | |
| Pb stress (0.05, 0.15 and 0.25 mmol/L) | Increase seedling shoot, and root length; improve chlorophyll content; reduce peroxide levels; alter SOD and APX activities | Chen et al, | |
| Heat stress (40 ºC for 10 d) | Improve pollen viability, and seed setting; lower ROS generation, and MDA content in anthers; differently alter SOD, POD, CAT and APX activities in anthers | Zhang et al, | |
| Brassinosteroid (BR) | Salt stress (NaCl: 0, 75, 100 and 125 mmol/L) | Enhance shoot length, root length, root number, fresh weight and dry weight; improve total chlorophyll, protein, and proline contents; lower lipid peroxidation while enhance activities of enzymatic antioxidants (SOD, APX, CAT, GR, GPX, DHAR and MDHAR); improve expression levels of antioxidant enzyme genes | Sharma I et al, |
| Auxin precursor | Cd stress (30 mg/kg) | Enhance plant height and yield; increase the numbers of tillers, panicles and grain weight; decrease Cd content in grains | Farooq et al, |
| Methyl jasmonate | Cd stress [Cd(NO3)2, 50 µmol/L] | Improve growth of root and shoot, and chlorophyll content; lower electrolyte leakage, lipid peroxidation, and ROS generation; decrease Cd uptake and accumulation; enhance GSH content, and activities of CAT, GR, POD and SOD | Singh and Shah, |
| As stress (NaAsO2, 25 µmol/L) | Improve root growth, and seedling biomass; decrease As accumulation; enhance photosynthetic rates, and chlorophyll content; lower membrane damage, electrolyte leakage, and lipid peroxidation; alter gene expression levels of JA signalling genes (OsCOI1 and OsJAZ3), JA responsive transport factor (OsMYC2), as well as As uptake, translocation and detoxification genes (OsLsi1, OsLsi2, OsNIP1;1, OsNIP3;1, OsLsi6, OsINT5, OsNRAMP1, OsPCS2 and OsABCC2) | Verma et al, | |
| BR and BR mimic | Heat stress (47 ºC for 2 h) | Improve shoot and root biomass; enhance photosynthetic pigment content, RWC and sugar content; improve stomatal conductivity and CO2 assimilation; reduce ROS generation and membrane damage | Thussagunpanit et al, |
| Brassinolide | Cold stress for 15 ºC | Promote cell elongation | Fujii and Saka, |
| As, Arsenic; AsA, Ascorbate; APX, Ascorbate peroxidase; Ca, Calcium; CAT, Catalase; Cd, Cadmium; DHAR, Dehydroascorbate reductase; GR, Glutathione reductase; GPX, Glutathione peroxidase; GSH, Glutathione; IAA, Indole-3-acetic acid; MDHAR, Monodehydroascorbate reductase; MDA, Malondialdehyde; Mg, Magnesium; P, Phosphorous; Pb, Plumbum; PEG, Polyethylene glycol; POX, Peroxidase; ROS; Reactive oxygen species; Se, Selenium; SMC, Soil moisture content; SOD, Superoxide dismutase; POD, Peroxidase; RWC, Relative water content. | |||
Table 1. Exogenous application of various plant hormones and their impacts on abiotic stress amelioration in rice.
| Plant hormone | Abiotic stress and its condition | Morphological, physiological, and molecular effects | Reference |
| Auxin | Drought stress SMC between 7.3% ± 0.5% and 6.7% ± 0.2% Heat stress (40.3 ºC ± 0.36 ºC and 37.7 ºC± 0.35 ºC) | Improve pollen viability, spikelet fertility, grain number per panicle, grain weight per panicle, 1000-grain weight, and grain yield per panicle; enhance chlorophyll content, delay senescence, and maintain greenness; lower lipid peroxidation and ROS accumulation while improving membrane stability; improve the expression levels of auxin signalling genes (TIR1, AFB2, SPL, TFL2 and HAF) | Sharma et al, |
| As stress (NaAsO2, 150 µmol/L) Se stress (Na2SeO4, 20 µmol/L) | I. Auxin (As and Se stress): Enhance root and shoot growths II. Auxin + Se (As stress): Improve chlorophyll content, proline and cysteine; lower protein content inhibition and DNA damage; reduce lipid peroxidation | Pandey and Gupta, | |
| Gibberellic acid (GA) | Drought stress (PEG6000, 15%) | Improve germination vigor, shoot length, root length, α-amylase activity, and soluble sugar content; induce the expression levels of α-amylase and expansion-related genes | Li J Z et al, 2019 |
| Salt stress (NaCl: 0, 50, 100, 150 and 200 mmol/L) | Enhance contents of proline, soluble sugar and protein as well as inorganic nutrients (K, Mg and Ca); increase panicle number, panicle length, grain yield per plant, 1000-grain weight, filled grain number per panicle, and harvest index | Misratia et al, | |
| Cold stress (10 ºC to 20 ºC) | Improve cold resistance and increase yield traits; promote seedling emergence and vigor | Chen et al, | |
| Abscisic acid (ABA) | Drought stress (PEG6000, 15%) | Enhance net photosynthesis, stomatal conductance, and transpiration rate; improve expression levels of chloroplast genes (OsPsbD1 and OsPsbD2); up-regulate expression levels of ABA biosynthesis genes (OsNCED2, OsNCED3, OsNCED4 and OsNCED5) | Teng et al, |
| Cold stress at 5 ºC | Improve the survival ratio | Lee et al, | |
| Salicylic acid (SA) | Salt stress (NaCl: 0, 100, 200, 300 and 400 mmol/L) | Enhance germination rate, root length, shoot length and plant height; improve the number of grains per panicle, number of filled grains per panicle, 1000-grain weight, the number of panicles per plant, and yield; increase contents of carbohydrate and protein along with minerals (Ca, P and K) while decreasing Na and Cl contents as well as Na+ and Cl- accumulation; enhance endogenous SA content | Jini and Joseph, |
| Cd stress (CdCl2, 50 µmol/L) | Improve root growth; reduce ROS level, and membrane damage; enhance SOD, POD and CAT activities as well as GSH, and AsA contents; improve non-protein thiols’ concentration | Guo et al, | |
| Pb stress (0.05, 0.15 and 0.25 mmol/L) | Increase seedling shoot, and root length; improve chlorophyll content; reduce peroxide levels; alter SOD and APX activities | Chen et al, | |
| Heat stress (40 ºC for 10 d) | Improve pollen viability, and seed setting; lower ROS generation, and MDA content in anthers; differently alter SOD, POD, CAT and APX activities in anthers | Zhang et al, | |
| Brassinosteroid (BR) | Salt stress (NaCl: 0, 75, 100 and 125 mmol/L) | Enhance shoot length, root length, root number, fresh weight and dry weight; improve total chlorophyll, protein, and proline contents; lower lipid peroxidation while enhance activities of enzymatic antioxidants (SOD, APX, CAT, GR, GPX, DHAR and MDHAR); improve expression levels of antioxidant enzyme genes | Sharma I et al, |
| Auxin precursor | Cd stress (30 mg/kg) | Enhance plant height and yield; increase the numbers of tillers, panicles and grain weight; decrease Cd content in grains | Farooq et al, |
| Methyl jasmonate | Cd stress [Cd(NO3)2, 50 µmol/L] | Improve growth of root and shoot, and chlorophyll content; lower electrolyte leakage, lipid peroxidation, and ROS generation; decrease Cd uptake and accumulation; enhance GSH content, and activities of CAT, GR, POD and SOD | Singh and Shah, |
| As stress (NaAsO2, 25 µmol/L) | Improve root growth, and seedling biomass; decrease As accumulation; enhance photosynthetic rates, and chlorophyll content; lower membrane damage, electrolyte leakage, and lipid peroxidation; alter gene expression levels of JA signalling genes (OsCOI1 and OsJAZ3), JA responsive transport factor (OsMYC2), as well as As uptake, translocation and detoxification genes (OsLsi1, OsLsi2, OsNIP1;1, OsNIP3;1, OsLsi6, OsINT5, OsNRAMP1, OsPCS2 and OsABCC2) | Verma et al, | |
| BR and BR mimic | Heat stress (47 ºC for 2 h) | Improve shoot and root biomass; enhance photosynthetic pigment content, RWC and sugar content; improve stomatal conductivity and CO2 assimilation; reduce ROS generation and membrane damage | Thussagunpanit et al, |
| Brassinolide | Cold stress for 15 ºC | Promote cell elongation | Fujii and Saka, |
| As, Arsenic; AsA, Ascorbate; APX, Ascorbate peroxidase; Ca, Calcium; CAT, Catalase; Cd, Cadmium; DHAR, Dehydroascorbate reductase; GR, Glutathione reductase; GPX, Glutathione peroxidase; GSH, Glutathione; IAA, Indole-3-acetic acid; MDHAR, Monodehydroascorbate reductase; MDA, Malondialdehyde; Mg, Magnesium; P, Phosphorous; Pb, Plumbum; PEG, Polyethylene glycol; POX, Peroxidase; ROS; Reactive oxygen species; Se, Selenium; SMC, Soil moisture content; SOD, Superoxide dismutase; POD, Peroxidase; RWC, Relative water content. | |||
Fig. 1. Plant hormone involvement in amelioration of abiotic stress in rice. Green lines with inhibitory heads signify repression mediated by hormonal action. Red lines with inhibitory heads indicate inhibition/repression. Black lines with arrowheads indicate activation/course of signalling. Blue cross indicates failure of inhibition. Aux/IAA, Auxin/indole-3-acetic acid; ARF, Auxin response factor; ABP1, Auxin-binding protein 1; AFB, Auxin signalling F-box protein; Aux-RE, Auxin response DNA element; ACS, 1-aminocyclopropane-1-carboxylic acid (AC) synthase; ACO, AC oxidase; AOS, Allene oxide synthase; AOC, Allene oxide cyclase; ADP, Adenosine diphosphate; AMP, Adenosine monophosphate; ATP, Adenosine triphosphate; AREB, bZIP transcription factors; As, Arsenic; ABRE, Abscisic acid (ABA) response element; ABI, ABA insensitive; BZR1, Brassinazole resistant 1; BRI1, Brassinosteroid insensitive 1; BAK1, BRI1-associated receptor kinase 1; BES1, BRI1-EMS-suppressor 1; CTR1, Constitutive triple response 1; CYP, Cytochrome P450 monooxygenase; CKX, Cytokinin oxidase/dehydrogenase; Cr, Chromium; CRE1, Cyclic AMP response element 1; Cd, Cadmium; DMAPP, Dimethylallyl pyrophosphate; ER, Endoplasmic reticulum; ETR, Ethylene receptor; EIN, Ethylene-insensitive; ERF, Ethylene response factor; HMBPP, Hydroxymethylbutenyl pyrophosphate; IPT, Isopentenyl transferase; JAZ, Jasmonate-ZIM domain proteins; JAV1, Jasmonate-associated VQ domain protein 1; NCED, 9-cis-epoxycarotenoid dioxygenase; OPDA, 12-oxophytodienoic acid; OPR, OPDA reductase; Pb, Lead; PM, Plasma membrane; PYR/PYL/RCAR, ABA receptors; PP2C, 2C-type protein phosphatase; SnRK2, SNF1-related kinase 2; SCF-COI1, Ubiquitin-ligase complex; TIR1, Transport inhibitor response 1; YUCCA, A flavin monooxygenase gene; ZEP, Zeaxanthin epoxidase.
| Plant hormone | Transgenic line/ gene-targeted | Abiotic stress | Response of plant | Reference |
| Auxin | OsIAA6 (Aux/IAA gene) | Drought | Upon over-expression, transgenic lines show reduced leaf rolling, improved recovery upon rehydration, better chlorophyll fluorescence, and lower transcript levels of DIP1 (molecular dehydration marker). Knock-down mutants exhibit irregular tiller outgrowth | Jung et al, |
| OsPIN3t (auxin efflux carrier gene) | Drought | Over-expression seedlings show improved shoot and root growth, better tillering, greater seed setting and adventitious root formation; improve the expression levels of drought-responsive genes (OsDREB2A and OsAP37); result in improved recovery upon rehydration. RNAi knock-down seedlings exhibit stunted growth and lower seed setting, tillering and yield per plant | Zhang et al, | |
| OsMYB-R1 (TF) | Cr6+ and drought | Over-expression lines show greater auxin accumulation and enhanced lateral root formation, leading to enhanced enzymatic antioxidant defence (CAT, APX, SOD and GPX); show improved revival efficiency from drought stress conditions. Loss-of-function mutants exhibit greater stress sensitivity showing severe wilting in drought and ineffective recovery patterns. Knock-down lines experience greater membrane damage and electrolyte leakage | Tiwari et al, | |
| Auxin and ABA | OsGH3-2 | Cold | Over-expression lines show delayed wilting, improve recovery rates after removal of stress conditions, lower membrane damage, and better oxidative stress tolerance | Du et al, |
| OsMADS25 | Salt | Over-expression lines show greater primary root length, lateral root density, and root cell elongation; show improved chlorophyll content as well as lower lipid peroxidation, membrane damage, and ROS accumulation in roots, shoots, leaves and bracts; exhibit significantly reduced water loss upon ABA application; result in better enzymatic antioxidant (CAT, APX, GPX and GR) activities in roots exposed to H2O2 as well as improved expression of ROS scavenger genes (OsCATB and OsGST4); show enhanced transcript levels of ABA-dependent stress-responsive genes (OsZIP23, OsZIP46, OsTRAB1, OsLEA3, OsABI5, OsP5CS1 and OsP5CR); result in enhanced transcript levels of auxin biosynthesis and signalling genes (OsYUC4, OsARF1 and OsARF16). Knock-down lines result in lowering of primary root elongation and root cell elongation; show enhanced membrane damage, chlorosis and leaf wilting as well as hampered ROS homeostasis in roots, shoots, leaves and bracts; hamper the root enzymatic antioxidant activity after H2O2 exposure and expression of ROS scavenger genes | Xu N et al, | |
| ABA | JERF1 (an ERF) | Drought | Over-expression lines lower wilting and yellowing while improved leaf production, fresh weight, tillering and recovery upon rehydration as well as greater root length; result in improved proline content and leaf RWC while lower water loss; exhibit greater expression of stress-responsive genes (OsP5CS, OsSPDS2, OsCDPK13 and OsLTP1); enhance expression of ABA synthesis genes (OsABA2 and Os03g0810800), and up-regulate ABA synthesis causing improved ABA content | Zhang et al, |
| SAPK6 and OsbZIP46 (Co-over- expressed) | Heat, cold and drought | Co-over-expression lines result in improved biomass, grain number, spikelet number, average panicle number, and yield per plant under drought conditions; show lesser water loss rate, drought symptoms such as leaf rolling, improved survival under heat and cold stress conditions | Chang et al, | |
| CK | OsCKX2 | Salt | Knock-down lines result in enhanced endogenous CK content; show improved growth, photosynthetic pigment content, net photosynthetic rate, stomatal conductance, intercellular CO2 concentration, RWC and PSII efficiency while reduce electrolyte leakage; show greater spikelet number per panicle, panicle branching, grain number per panicle, grain weight and overall panicle number | Joshi et al, |
| ET | OsARD1 | Submergence, drought and salt | Over-expression lines result in greater endogenous ethylene content, coleoptile elongation upon germination under submergence accompanying anoxia; exhibit longer shoot under submergence and greater induction of Ahd1 and Sub1C genes; show delayed symptoms of drought and improved green color retention lines; expose to salt stress show longer shoot length, improved maintenance of leaf structure, green color retention, and delayed withering; show greater trichome, and stomatal density as well as improved water holding capacity; exhibit enhanced expression of ethylene biosynthesis genes (OsACS2 and OsACS4), ERFs (AP37 and AP59), TFs (OsbZIP23) and stress-responsive genes (OsNCED4, OsNCED5, OsPP108 and OsSalT) | Liang et al, |
| ABA, Abcisic acid; APX, Ascorbate peroxidase; Aux/IAA, Auxin/indole-3-acetic acid; CAT, Catalase; Cr, Chromium; CK, Cytokinin; ET, Ethylene; ERF, Ethylene response factor; GR, Glutathione reductase; GPX, Glutathione peroxidase; GSH, Glutathione; K, Potassium; POX, Peroxidase; ROS, Reactive oxygen species; RNAi, RNA interference; RWC, Relative water content; SOD, Superoxide dismutase; TF, Transcription factor. | ||||
Table 2. Representative studies of rice genes targeted at plant hormone for abiotic stress mitigation.
| Plant hormone | Transgenic line/ gene-targeted | Abiotic stress | Response of plant | Reference |
| Auxin | OsIAA6 (Aux/IAA gene) | Drought | Upon over-expression, transgenic lines show reduced leaf rolling, improved recovery upon rehydration, better chlorophyll fluorescence, and lower transcript levels of DIP1 (molecular dehydration marker). Knock-down mutants exhibit irregular tiller outgrowth | Jung et al, |
| OsPIN3t (auxin efflux carrier gene) | Drought | Over-expression seedlings show improved shoot and root growth, better tillering, greater seed setting and adventitious root formation; improve the expression levels of drought-responsive genes (OsDREB2A and OsAP37); result in improved recovery upon rehydration. RNAi knock-down seedlings exhibit stunted growth and lower seed setting, tillering and yield per plant | Zhang et al, | |
| OsMYB-R1 (TF) | Cr6+ and drought | Over-expression lines show greater auxin accumulation and enhanced lateral root formation, leading to enhanced enzymatic antioxidant defence (CAT, APX, SOD and GPX); show improved revival efficiency from drought stress conditions. Loss-of-function mutants exhibit greater stress sensitivity showing severe wilting in drought and ineffective recovery patterns. Knock-down lines experience greater membrane damage and electrolyte leakage | Tiwari et al, | |
| Auxin and ABA | OsGH3-2 | Cold | Over-expression lines show delayed wilting, improve recovery rates after removal of stress conditions, lower membrane damage, and better oxidative stress tolerance | Du et al, |
| OsMADS25 | Salt | Over-expression lines show greater primary root length, lateral root density, and root cell elongation; show improved chlorophyll content as well as lower lipid peroxidation, membrane damage, and ROS accumulation in roots, shoots, leaves and bracts; exhibit significantly reduced water loss upon ABA application; result in better enzymatic antioxidant (CAT, APX, GPX and GR) activities in roots exposed to H2O2 as well as improved expression of ROS scavenger genes (OsCATB and OsGST4); show enhanced transcript levels of ABA-dependent stress-responsive genes (OsZIP23, OsZIP46, OsTRAB1, OsLEA3, OsABI5, OsP5CS1 and OsP5CR); result in enhanced transcript levels of auxin biosynthesis and signalling genes (OsYUC4, OsARF1 and OsARF16). Knock-down lines result in lowering of primary root elongation and root cell elongation; show enhanced membrane damage, chlorosis and leaf wilting as well as hampered ROS homeostasis in roots, shoots, leaves and bracts; hamper the root enzymatic antioxidant activity after H2O2 exposure and expression of ROS scavenger genes | Xu N et al, | |
| ABA | JERF1 (an ERF) | Drought | Over-expression lines lower wilting and yellowing while improved leaf production, fresh weight, tillering and recovery upon rehydration as well as greater root length; result in improved proline content and leaf RWC while lower water loss; exhibit greater expression of stress-responsive genes (OsP5CS, OsSPDS2, OsCDPK13 and OsLTP1); enhance expression of ABA synthesis genes (OsABA2 and Os03g0810800), and up-regulate ABA synthesis causing improved ABA content | Zhang et al, |
| SAPK6 and OsbZIP46 (Co-over- expressed) | Heat, cold and drought | Co-over-expression lines result in improved biomass, grain number, spikelet number, average panicle number, and yield per plant under drought conditions; show lesser water loss rate, drought symptoms such as leaf rolling, improved survival under heat and cold stress conditions | Chang et al, | |
| CK | OsCKX2 | Salt | Knock-down lines result in enhanced endogenous CK content; show improved growth, photosynthetic pigment content, net photosynthetic rate, stomatal conductance, intercellular CO2 concentration, RWC and PSII efficiency while reduce electrolyte leakage; show greater spikelet number per panicle, panicle branching, grain number per panicle, grain weight and overall panicle number | Joshi et al, |
| ET | OsARD1 | Submergence, drought and salt | Over-expression lines result in greater endogenous ethylene content, coleoptile elongation upon germination under submergence accompanying anoxia; exhibit longer shoot under submergence and greater induction of Ahd1 and Sub1C genes; show delayed symptoms of drought and improved green color retention lines; expose to salt stress show longer shoot length, improved maintenance of leaf structure, green color retention, and delayed withering; show greater trichome, and stomatal density as well as improved water holding capacity; exhibit enhanced expression of ethylene biosynthesis genes (OsACS2 and OsACS4), ERFs (AP37 and AP59), TFs (OsbZIP23) and stress-responsive genes (OsNCED4, OsNCED5, OsPP108 and OsSalT) | Liang et al, |
| ABA, Abcisic acid; APX, Ascorbate peroxidase; Aux/IAA, Auxin/indole-3-acetic acid; CAT, Catalase; Cr, Chromium; CK, Cytokinin; ET, Ethylene; ERF, Ethylene response factor; GR, Glutathione reductase; GPX, Glutathione peroxidase; GSH, Glutathione; K, Potassium; POX, Peroxidase; ROS, Reactive oxygen species; RNAi, RNA interference; RWC, Relative water content; SOD, Superoxide dismutase; TF, Transcription factor. | ||||
Fig. 2. Plant hormones modulate signaling mechanism in plant system. Red lines with inhibitory heads indicate inhibition/repression; Black lines with arrowheads indicate activation/course of signalling. ABA, Abscisic acid; ET, Ethylene; ACC, Aminocyclopropane-1-carboxylic acid; JA, Jasmonic acid; GA, Gibberellic; SA, Salicylic acid.
Fig. 3. Approaches for improving abiotic stress tolerance in rice by targeted regulation of plant hormone genes. gRNA, guide RNA; RISC, RNA-induced silencing complex; RNAi, RNA interference; siRNA, Small interfering RNA.
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