Transcription factors in abiotic stress response
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1 Transcription factors in abiotic stress response Nelson Saibo ITQB Outline 1 - Plant responses to adverse environmental conditions 2 - Transcription factors and transcriptional regulation 3 - Transcription factors involved in abiotic stress responses 4 -TFs and phtosynthetic responses to abiotic stress 5 - Identification and characterization of novel TFs 1
2 Plants and the environmental conditions Environment altered beyond its normal range of variation to adversely affect the individual physiology of the organism in a significant way Plant stress Plant response to abiotic stress Cold Drought Salinity Chemical pollution Heat Cell damage Secondary stress: Osmotic stress Oxidative stress Signal sensing, perception and transduction ABIOTIC STRESS 5% crop loss world wide Osmosensor, second mensagers, MAPKs, Ca 2+ sensors, CDPKs Transcription control TFs: CBF/DREB, bzib, MYC/MYB Gene activation Osmoprotection, water and ion movement, detoxification, and chaperone functions Recovery of cellular homeostasis, functional and structural protection of proteins and membranes Stress tolerance 2
3 Abiotic stress tolerance is a multigenic trait Environmental stimuli or stress TF Stress responsive genes STRESS TOLERANCE Transcription Factors Transcription factors (TFs) - proteins that show sequence-specific DNA-binding and that are capable acivating or repressing gene transcription. +1 Basic Transcription machinery GGCATGGC TATA gene mrna Promoter Transcription coregulators (coactivators/corepressors), chromatin remodelers, histone acetylases, kinases, and methylases play crucial roles in gene regulation, but lack DNA biding domains and therefore are not classified as TFs. 3
4 Transcription in Eukaryotes a d b e c f Transcription in Eukaryotes Basic transcriptional machinery 4
5 Schematic diagram of a prototypical transcription factor TFs contain DNA-binding domain (DBD), signal sensing domain (SSD), and a transactivation domain (TAD) The order of placement and the number of domains may differ in various types of TFs The transactivation and signal sensing functions are frequently contained within the same domain Transcription Factor Families DNA binding domain FAMILY DNA binding domain(s) + (protein-protein interaction domains) Subfamily 5
6 Arabidopsis Transcription Factors Relationships and domain shuffling among the different Arabidopsis transcription factor families Riechmann (2) Science 29, 215 Eukaryotic Transcriptional Regulators 6
7 Plant TF families and their function AP2/ERF (144) Development (flower/seed/root); metabolic pathways; stress response; hormone response (ABA/C 2 H 4 ) bhlh (139) Development (trichome/root/carpel) abiotic stress; secondary metabolism; light responses; MYB (19) Development; secondary metabolism; defence response; abiotic stress; hormone response (ABA/GA 3 ); cell cycle; light C2H2(Zn) (112) Flower/seed development; abiotic stress; light NAC (19) Development (meristem); auxin-response; virus resistance; * HB (9) Development (several); sucrose signalling; cell death; * MADS (82) Reproductive organs development; flowering time/abscission; * bzip (77) Flower/leaf/photomorphogenic development; seed-storage; WRKY (72) defence response; hormone response/biosynthesis; * Defence response; * C2C2(Zn) (14) Seed development/metabolism; flowering time; circadian rhythm; * * Abiotic stress Riechmann 22 Transcriptional regulation: an overview. The Arabidopsis Book Transcriptional regulatory network Environmental stimuli or stress Developmental signals Induction P Modification U S A TF1 Activ. B TF2 P C TF3 U S promoter TFIID TFIIA TF TF TF II IIF II H P E TFIIB RNA pol II Basic machinery Transcription TATA +1 ATGNNNNN gene mrna 7
8 Abiotic stress transcriptional network Biotic stress and wounding Drought, High salinity Signal perception Cold Jasmonic acid ABA ABA-independent ICE1 HOS1 SIZ1 MYB15 ICE1 MYB MYC AREB/ABF ZF-HD NAC DREB2? CBF4/DREB1D CBF3/DREB1A CBF1/DREB1B? DREB2 ZAT12 AREB/ABF CBF2/DREB1C MYCR RD22 MYBR ABRE rps-1like NACR RD29B ERD1 CAB? STZ/ZAT1 DRE/CRT RD29A?? Annals Botany 29 13, 69 Manipulation of TFs to improve abiotic stress tolerance Nature Biotechnology , 287 8
9 DREB1A driven by the 35 S CaMV vs stress inducible rd29a promoter 35S:TF growth retardation Nature Biotechnology , 287 Both 35S:DREB1A and rd29a:dreb1a show enhanced stress tolerance Nature Biotechnology , 287 9
10 DREB1A target genes are strongly expressed under control conditions OX mimics i acclimation Higher stress tolerance Nature Biotechnology , 287 Overexpression of HvCBF4 in rice enhances drought tolerance NT Ubq:: ::HvCBF4 12 days drought 1 week recovering Survival rate: Plants overexpressing HvCBF4-9% Non transformed plants 19% 1
11 Overexpression of HvCBF4 in rice enhances drought tolerance Ubi::HvCBF4 AtRD29A::HvCBF4 NT Days after withhold water Days after withhold water AtDREB1A overexpressed in chrysanthemum enhances tolerance to heat stress Survival rate: 7,8% 16,3% 36h at 45ºC 3 weeks 22ºC Plant Mol Biol DOI 1.17/s z 11
12 Higher photosynthetic capacity and elevated activity of Rubisco Plant Mol Biol DOI 1.17/s z TF overexpression can improve photosynthetic performance under abiotic stress 12
13 Photosynthetic responses to abiotic stress ABIOITIC STRESS Stomatal control of CO 2 diffusion Photosystem II repair Electron transport Rubisco activity Scavenging of ROS Photorespiration Photosynthetic efficiency is greatly decreased Are the photosynthetic responses to abiotic stress modulated at transcriptional level? Annals Botany 29 13, 69 13
14 Gene expression in rice plants under drought stress Drought stress causes down-regulation of rice genes coding for proteins involved in the photosynthetic light reactions Plant Mol Biol 29 69, 133 TFs involved in the photosynthetic response to abiotic stress Fast responses Long term responses MYB6 Protodermal Cell Meristemoid Mother Cell Guard Mother Cell Mature Guard Cells ABI3 MMC GMC Stomatal MYB61 ABA SPCH ICE1+SCRM2 ABA MAP KINASE PATHWAY Abiotic stress MUTE ICE1+SCRM2 FLP MYB88 FAMA ICE1+SCRM2 Annals Botany 29 13, 69 14
15 TFs involved in the photosynthetic response to abiotic stress Abiotic stress LIGHT Non-stomatal Annals Botany 29 13, 69 MYB PSII PSI Chloroplast genes STZ AZF2 Glycine betaine Chloroplast bzip Rubisco Calvin cycle MYB? σ factor Mesophyll cell PpcI and GapI Nuclear genes Cytosol DOF PEPCase Bundle sheath NOVEL TRANSCRIPTION FACTORS REGULATING ABIOTIC STRESS TOLERANCE IN RICE ( (ORYZA SATIVA L.) 15
16 Rice in Portugal Consumption: 17 kg/capita/year Mondego Production: 6% Tejo Sorraia Sado PRESENT PROBLEMS: Pests and diseases Low production Infestants Cold Salinity What makes rice an ideal model organism? Important crop main source of energy for 2/3 of the world population Rice genome fully sequenced ~ 4 Mb, (maize ~ 25 Mb, barley ~ 5 Mb, wheat ~ 16 Mb) High-efficiency genetic transformation Genetic and physical maps of high density High degree of synteny among genes in cereal genomes Insertion knockout mutants available Microarrays for the whole genome ~5. transcripts RICE - AN IDEAL MODEL ORGANISM FOR MONOCOTS AND CEREAL CROPS 16
17 Low temperature signalling pathway COLD ICE1 [Ca 2+ ] cyt U HOS1 P ICE1 S Kinases SIZ1 ICE1-like U ICE1 MYB15 Proteolysis DREB1A/ CBF3 DREB1C/ CBF2 DREB1B/ CBF1 CRT/DRE COR genes ACCLIMATION Yeast one-hybrid screening to isolate TFs or other DNA-binding proteins 2μ ori pact II ' PADH1 GAL4-AD cdna library Transformation Yeast reporter strain (leu2, his3) LEU2 pint1 vector library protein GAL4 AD Transcription if hybrid protein interacts with bait sequence promoter X TATA HIS3 selection gene Selection for growth on histidine-lacking medium Ouwerkerk PBF, Meijer AH (21) Cur. Prot. Mol. Bio.,
18 TFs controlling the expression of OsHOS1 ERF1-15pb -1pb -5pb ERF2 ATG OsHOS h 2h 3h 5h ERF1 Actin ERF1 is down-regulated at low temperature (5ºC) ERF2 is not regulated at transcriptional level Low temperature signalling pathway COLD ERF1 U HOS1 P ICE1 ICE1 S [Ca 2+ ] cyt Kinases SIZ1 ICE1-like U ICE1 MYB15 Proteolysis DREB1A/ CBF3 DREB1C/ CBF2 DREB1B/ CBF1 CRT/DRE COR genes ACCLIMATION 18
19 OsDREB1B gene is induced by cold and drought Mock Control 5ºC Control 1ºC Control Drought Control 1 2 Shoot 4 1h 2h 5h 1h 1 2 Root 4 1h 2h 5h 1h TFs controlling OsDREB1B expression: GFP::TF ZF-HD ZF-HD ZF-HD ZF-HD C2H2 bhlh C2H2 C2H2 ATG OsDREB1B -15pb -1pb -5pb OsSTZ (C2H2) binds to OsDREB1B promoter OsSTZ is induced by cold, salt, and drought Mock 5ºC 1ºC ABA NaCl Drought 1 2 Shoot 4 1h 2h 5h 1h 1 2 Root 4 1h 2h 5h 1h Mock 5ºC 1ºC ABA NaCl Drought 1 Internal control 2 4 1h 2h 5h 1h 1 Internal control 2 4 1h 2h 5h 1h 19
20 OsPIF (bhlh) binds to OsDREB1B promoter The control of OsPIF expression by cold involves alternative splicing Mock 1 2 Shoot 4 1h 2h 5h 1h 1 2 Root 4 1h 2h 5h 1h 5ºC 1ºC ABA NaCl Drought Mock 5ºC 1ºC ABA NaCl Drought 1 Internal control 2 4 1h 2h 5h 1h 1 Internal control 2 4 1h 2h 5h 1h A C2H2 that binds to OsDREB1B promoter C2H2 transcription is induced at a higher level in the roots Mock 5ºC 1ºC ABA NaCl Drought 1 2 Shoot 4 1h 2h 5h 1h 1 2 Root 4 1h 2h 5h 1h Mock 5ºC 1ºC ABA NaCl Drought 1 Internal control 2 4 1h 2h 5h 1h 1 Internal control 2 4 1h 2h 5h 1h 2
21 Low temperature signalling pathway COLD SALINITY DROUGHT ERF1 U HOS1 U ICE1 ICE1 P ICE1 S MYB15 [Ca 2+ ] cyt Kinases SIZ1 ICE1-like C2H2 bhlh STZ Proteolysis DREB1A/ CBF3 DREB1C/ CBF2 DREB1B/ CBF1 CRT/DRE COR genes ACCLIMATION OsDREB1A gene is induced by cold 28ºC 5ºC h Shoot 2h 3h 5h 7h 12h OsDREB1A Control Control OsActin1 Control of OsDREB1A expression: bzip ATG -15pb -1pb -5pb OsDREB1A bzip Known to be involved in biotic stress responses 21
22 Low temperature signalling pathway COLD SALINITY DROUGHT ERF1 U HOS1 U ICE1 ICE1 P ICE1 S MYB15 [Ca 2+ ] cyt Kinases SIZ1 ICE1-like C2H2 bhlh STZ Proteolysis bzip DREB1A/ CBF3 DREB1C/ CBF2 DREB1B/ CBF1 CRT/DRE COR genes BIOTIC ACCLIMATION Plant cell responses to salinity SALINITY High Na + RMC Root Meander Curling - perception / response biotic stress RMC Na + H + Ca + SOS3 SOS2 P? ABI2 H + SOS2 CBL1 Ca + Vacuole CAX1 Ca + Ca + CBL1 SOS2 H + NHX1 Na + H + P? Li + SOS3 SOS2 NHX1 -Na Na + /H + antiport tonoplast - response to salt stress 22
23 The expression of OsRMC and OsNHX1 is induced by NaCl 1mM NaCl 2mM NaCl OsRMC h 2h 5h 12h 2h 5h 12h OsNHX1 OsActin1 Transcriptional regulation of OsRMC -15pb ERF3-1pb ERF4-5pb ATG OsRMC ERF3 Homologue to the Arabidopsis ABR1, a negative regulator of ABA responses ERF4 Phosphorylated in vitro by MPK12/BWMK1 23
24 Salt signalling pathway SALINITY BIOTIC ERF3 P ERF4 RMC Receptor? TF? TF? Responsive Elements Stress-responsiveresponsive genes Preliminary conclusions The genes analysed are controlled by several TFs, differentially regulated by different stresses Regulation Transcriptional Post-transcriptional transcriptional (and post-translational translational?) Transcriptional regulation of OsDREB1A and OsRMC cold and salinity cross talk with biotic stress signalling, light sensing 24
25 ACKNOWLEDGEMENTS Margarida Oliveira EGP Group Pieter Ouwerkerk Leiden University Duarte Figueiredo (DREB1B) Tânia Serra (RMC) Tiago Lourenço (HOS1) Subhash Chander (DREB1A) Pedro Barros (cdna library) Roberto van Maanen (NHX1) FCT for the PhD and fellowships, and the project POCTI/BIA-BCM/5663/24 ADONIS Marie Curie Action (training and mobility) 25
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