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Jasmonates and Biotic Stress

               By

          Dipesh Dhakal




  DEPARTMENT OF BIOTECHNOLOGY
   GURU NANAK DEV UNIVERSITY
           AMRITSAR
Auxins


        Salicylic acid                      Gibberellins




Abscisic acid            Plant hormones              Cytokinins




      Jasmonates                             Ethylene


                         Brassinosteroids
JASMONIC ACID
•    Jasmonates are cyclopentanone
     compound or plant hormones
     derived from α-linolenic acid.
•    It includes group of oxygenated
     fatty acids collectively called
                                                                                             A
     oxylipins and Jasmonic Acid is
     main precursor        to different
     compounds to this group.
•    Methyl jasmonate was first
     isolated from the essential oil of
     Jasminum grandiflorum
     Demole et. al.,(1962).Helv. Chim.Acta.45:675-695

•   They are ubiquitous in plant
    kingdom and are also produced by
    certain fungi.                                                                           B
•   First isolated in culture filtrate of
                                                            A. Jasmonic acid
    fungi Lasiodiplodia theobromae.                         B. Jasminum grandiflorum plant
     Aldridge et. al.,(1971).J.Chem.Soc.Chem. Comm. 1623-
    1627
BIOSYNTHESIS OF JASMONATES
                                                      Jasmonic acid synthesized from fatty acid(α-
                                                       linolenic acid)

                                                      Lipoxygenase (LOX), AOS , AOC are key
                                                       enzymes of JA biosynthesis in Chloroplast,
                                                       and they form OPDA.

                                                      OPDA is transported to peroxisome through
                                                       ABC transporter COMATOSE(CTS)

                                                      Reduction of cyclopentanone ring of OPDA is
                              CTS
                                                       catalyzed by peroxisomal OPR enzyme

                                                      Three cycles of β-oxidation occurs to give
                                                       finally Jasmonic Acid. Enzymes involved are:

                                                      •         ACX1 (Acyl-CoA oxidase in tomato)
                                                      •         MFP (Multifunctional Protein)
                                                      •         KAT (L-3-ketoacyl CoA thiolase)

                                                       The JA    and   its metabolic derivatives are

  Fig : Biosynthetic Pathway of Jasmonic Acid             collectively called JASMONATES

Source: Wasternack et .al.,(2002).PNARMB.72:165-221
Derivatives of Jasmonic acid – THE JASMONATES

                                                               Carboxylic acid side chain
                                                                conjugated to ACC(1-amino
                                                                cyclopropane-1-carboxylic acid)

                                                               Methylated    form    by    JA
                                                                methyltransferase

                                                               Decarboxylated to cis-jasmone

                                                               Conjugated to AA such as Ile by
                                                                JA amino acid synthase

                                                               Reduction of keto group of
                                                                pentanone ring to cucurbic acid

                                                               Pentenyl      side      chain
                                                                hydroxylated in position C-11
                                                                and C-12

    Fig : Different Metabolites Produced from Jasmonic Acid

Source: Wasternack, C. ,(2007).Annals of Botany.100:681-697
DIFFERENT PHYSIOLOGICAL ROLES OF JA
           S. No       Process                          Putative Signals   Alteration/Species

           1           Root Growth                      JA,JA-Ile          Inhibition

           2           Seed Germination                 JA                 Inhibition

           3           Tuber Formation                  JA-OH,JA           Induction/Potato

           4           Tendril Coiling                  OPDA               Stimulation/Bryonia

           5           Nyctinasty                       JA                 Stimulation/Albizzia

           6           Trichome formation               JA                 Induction/Tomato

           7           Senescence                       JA                 Stimulation

           8           Flower Development               JA

                       Anther Development and                              Induction/Arabidopsis
                       dehiscence
                                                                           Induction/Tomato
                       Female Organ
                       Development                                         Induction/Arabidopsis

                       Filament elongation
           12          Biotic Stress                    JA, JME,OPDA       Resistance/Ubiqutitous

Source: Wasternack, C. (2007).Annals of Botany.100:681-697
STRESS AND ITS TYPES

   Stress is any change in the environmental
   condition that may adversely affect the
   plant’s    growth,   development     and
   adaptability



                                 STRESS



 BIOTIC                                                          ABIOTIC

            Bacteria

              Virus        Drought        Salinity    Temperature

              Fungi       Heavy metal     Pesticide      Light
            Herbivores

Nematodes
Fungal pathogen      Wounding by herbivores
 Bacteria                                  Insect attack



Virus              Biotic stress                Nematodes
                    perception




                       Stress
                      signaling

                                 Jasmonates ?

                Downstream activation
                        of stress
               related/resistance genes
CERTAIN FACTS TO PONDER OVER



 In India, fungal diseases are rated either the most
  important or second most important factor contributing to
  yield loss
  Grover A. et al., 2003. Current science 84:330-340
 Global loss because of pathogens is estimated to be
  12% of potential crop production
  Shah D.M. et al., 1997. Current Opinion in Biotechnology B:208-214

 The highest losses, estimated at more than $42 billion per
  year, occur in vegetables, fruits and rice
   Shah D.M. et al., 1997. Current Opinion in Biotechnology B:208-214
FUNGAL DISEASES AND ECONOMIC LOSSES
                                                                          Total yield
   Crop                    Pathogen                        Disease         loss(%)

Rice           Paricularia oryzae                           Blast             21

Wheat          Puccinia recondita                       Brown leaf rust       30

Maize          Helminthosporium maydis                    Leaf blight         30

Sorghum        Sphacelotheca reiliaria                   Grain mould          18

Pigeonpea      Fusarium udum                                 Wilt             24

Chickpea       Fusarium oxysorium                            Wilt             23

Brassica       Alternaria brassiceae                        Blight            30

Soyabeen       Phakospora packyrhizi                         Rust             23

Potato         Phytopthora infestans                      Late blight         31
 Source: Current science, vol.84, No. 3, 10 Feb. 2003
ROLE OF JA IN RESISTANCE
       Increased endogenous level of JA on treatment of Arabidopsis with necrotrophic fungus
        A. brassicola    Penninckx et al.(1996).Plant Cell.8:2309-2323

       Biosynthetic and Signaling Arabidopsis mutants were more sensitive to attack by
        necrotrophic fungal pathogen Phytium (jar1,coi1 and fad3-2fad7-1fad8 mutants) and
        necrotrophic bacteria Erwinia carotovora (coi1 mutants)
         Staswick et al.,(1998).Plant Journal 15:747-754
         Vijayan et al.,(1998).PNAS USA 95:7209-7214

       Transgenic plants overexpressing JMT and thus higher levels of
        methyl jasmonate more resistant to necrotrophic pathogen
        Botrytis cinera
         Walling (2000 ). Journal of Plant Growth Regulators 19: 195-216
       The fad3-2fad7-1fad8 mutant more sensitive to attack
        by fungal gnat Bradysia and coi1 is more sensitive to
        Diamond –back moth
         McConn et al.,(1997). PNAS USA.94:5473-5477;
         Xie et al (1998). Science 280: 1091-1094
                                                                                                Source: Berger,(2002).Planta 214:497-504
Mutant       Biochemical Significance          Role of Mutant    Expression of JA responsive   Effects
                                                                 genes

coi          Coronatine insensitive            JA Signaling      Reduced                       Sensitivity to insect and necrotrophic pathogens increased

fad          Defective Fatty Acid Desaturase   JA Biosynthetic   Reduced                       Sensitivity to insect and necrotrophic pathogens increased

cev          Constitutive Expression of VSP    JA Signaling      Increased                     Increased sensitivity to biographic pathogens

jar          JA Resistant                      JA Signaling      Reduced                       Increased sensitivity to necrotrophic pathogens

opr          Defective OPDA Reductase          JA Biosynthetic   Reduced                       Increased sensitivity to insects

jin          JA Insensitive                    JA Signaling      Reduced                       Increased sensitivity to pathogens
Mechanism of action of JAs in biotic stress




                                                        Source:Wasternack (2007).Annals of Botany.100:681-697
Source:Farmer and Ryan(1992).The Plant Cell.4:129-134
OVERALL MECHANISM OF SYSTEMIC RESPONSE
     Wounded leaves produce an 18-amino acid peptide called systemin from
      carboxyl terminal of prosystemin (200 AA precursor) in PP and it elicits
      production of JA in companion cell-sieve element complex.
     JA moves throughout the plant in the phloem. Covalently modified JA (JA-x)
      play important role in systemic signaling.
     signal is recognized at the target cell e.g. mesophyll (leaf)
     Jasmonic acid turns on defense related genes (genes for proteinase inhibitor
      etc.) in target cells.




Source: Schilmiller and Howe (2005). Current Opinion in Plant Biology . 8:369-377
Jasmonates and Biotic Stress
AMPLIFICATION OF SYSTEMIN SIGNAL
 Amplification of wound signaling is a major
  event in systemic defense
                                                                   Wounding
 Systemin activates AOC which in turn
  activates Prosystemin through OPDA/JA.                         Prosystemin
  OPDA/JA again activates back the AOC.

 In systemic response once JA is                                  Systemin
  synthesized, its cyclic production of JA can
  occur by positive feedback mechanism
                                                                       AOC
 JA is a systemic signal that leads to
  systemic expression of genes encoding                           OPDA/JA
  Proteinase inhibitors(PINs)

 The plant may develop some resistance                          Prosystemin
  against subsequent herbivore attack.
                                                                  Systemin
 Thus JA signaling is necessary in systemic
  response in leaf.
                                                 Fig: Amplification of wound signaling of tomato


                                                        Source:Wasternack et al(2006).163: 297-306
SYSTEMIC RESISTANCE AND ROLE OF JAs
                                Systemic Acquired Resistance (SAR)
   Systemic resistance
        in plants                Induced Systemic Resistance (ISR)
SAR:
 Initial infection result in formation of necrotic lesions
 SA concentrations increase and methyl-SA is released in distal plant tissue
 PR proteins in the non-invaded parts of the plants are synthesised resulting in
  reduction in disease symptoms after subsequent infection of many pathogenic
  species.

ISR:
 Non pathogenic root colonizing
  rhizobacteria e.g. P. fluorescens or
  wounding initiates ISR
 ISR does not depend on SA and PR
  protein
 ISR requires both JA and ethylene
  signalling
 SAR regulatory protein NPRI is
  required
 Systemin is involved in ISR which is
  absent in SAR.

                                     Source:Harman et. al.(2004) .Nature Reviews Microbiology. 2:43-56
Overall Mechanism of Defence mediated by different signaling compounds
    Source: Piertse et.al.(2009).Nature Chemical Biology.5:308-316
TRANSCRIPTIONAL REGULATION OF JA
                                                                   Four major of interacting players of:
                                                                   1. a JA signal
                                                                   2. the SCF-type E3 ubiquitin ligase
                                                                      SCFCOI1
                                                                   3. Jasmonate ZIM-domain (JAZ)
                                                                      repressor proteins that are targeted by
                                                                      SCFCOI1 for degradation by the
                                                                      ubiquitin/26S proteasome pathway ,and
                                                                   4. transcription factors (TFs)
                                                                   Howe et. al.(2008)Current Opinion in Plant Biology 11:428-235


                                                                    Transcriptional regulation mechanism:
                                                                    1. Expression of regulatory TFs for
                                                                       JAs
                                                                    2. Cross talk with the TFs responsible
                                                                       for expression of other hormones.




Fig: Transcription factors involved in signalling pathways of JA

                                                                         Source: Wasternack(2007). Annals of Botany100:681-697
Cross Talk with Ethylene and JA
                                                            •The antagonistic action of MYC2
                                                            and ERF1 may cause independence
                                                            between wound signaling and
                                                            pathogen defence signaling
                                                            Lorenzo et al (2004)Plant Cell 16:1938-1950)
                                                            •JA alone induce the expression
                                                            of AtMYC2 responsible for the
                                                            activation of wound response genes
                                                            and for the repression of pathogen
                                                            response genes.
                                                            •The cooperation of ET and JA
                                                            signals through the induction of
                                                            ERF1 leads activation of PR genes
                                                            and to the repression of WR genes
                                                            •Therefore, the interplay between
                                                            ERF1 and AtMYC2 allows the plant
                                                            selection of the correct set of genes
                                                            in response to these two stresses
Schematic Representation TF in response to Pathogens and Wounding


                                                               Source: Wasternack(2007). Annals of Botany100:681-697
Relevance of JAs studies
     Plant lack an immune system like in animals but posses mechanism
that recognizes potential pathogens and initiate defense responses.
During their biochemical evolution, the plants are devised with certain
magic molecules of defense (secondary metabolites) like JAs.
     Recent insights into the JAs mediated plant defense cascade and
knowledge of key regulators of this will help us to design future crops with
increased biotic stress resistance and better adaptability.
     Higher crop yields might be achieved by increasing the
pathogen/insect resistance which can be achieved by manipulating the
expression of the key genes involved in JAs biosynthesis and signaling
cascades.
Feeding the ever increasing human population is biggest
           social problem/challenge after all.
Overpopulation
‘The Problem’
Hunger
‘The Consequence’
Science
‘The answer’
THANK YOU

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Jasmonates and Biotic Stress

  • 1. Jasmonates and Biotic Stress By Dipesh Dhakal DEPARTMENT OF BIOTECHNOLOGY GURU NANAK DEV UNIVERSITY AMRITSAR
  • 2. Auxins Salicylic acid Gibberellins Abscisic acid Plant hormones Cytokinins Jasmonates Ethylene Brassinosteroids
  • 3. JASMONIC ACID • Jasmonates are cyclopentanone compound or plant hormones derived from α-linolenic acid. • It includes group of oxygenated fatty acids collectively called A oxylipins and Jasmonic Acid is main precursor to different compounds to this group. • Methyl jasmonate was first isolated from the essential oil of Jasminum grandiflorum Demole et. al.,(1962).Helv. Chim.Acta.45:675-695 • They are ubiquitous in plant kingdom and are also produced by certain fungi. B • First isolated in culture filtrate of A. Jasmonic acid fungi Lasiodiplodia theobromae. B. Jasminum grandiflorum plant Aldridge et. al.,(1971).J.Chem.Soc.Chem. Comm. 1623- 1627
  • 4. BIOSYNTHESIS OF JASMONATES Jasmonic acid synthesized from fatty acid(α- linolenic acid) Lipoxygenase (LOX), AOS , AOC are key enzymes of JA biosynthesis in Chloroplast, and they form OPDA. OPDA is transported to peroxisome through ABC transporter COMATOSE(CTS) Reduction of cyclopentanone ring of OPDA is CTS catalyzed by peroxisomal OPR enzyme Three cycles of β-oxidation occurs to give finally Jasmonic Acid. Enzymes involved are: • ACX1 (Acyl-CoA oxidase in tomato) • MFP (Multifunctional Protein) • KAT (L-3-ketoacyl CoA thiolase)  The JA and its metabolic derivatives are Fig : Biosynthetic Pathway of Jasmonic Acid collectively called JASMONATES Source: Wasternack et .al.,(2002).PNARMB.72:165-221
  • 5. Derivatives of Jasmonic acid – THE JASMONATES  Carboxylic acid side chain conjugated to ACC(1-amino cyclopropane-1-carboxylic acid)  Methylated form by JA methyltransferase  Decarboxylated to cis-jasmone  Conjugated to AA such as Ile by JA amino acid synthase  Reduction of keto group of pentanone ring to cucurbic acid  Pentenyl side chain hydroxylated in position C-11 and C-12 Fig : Different Metabolites Produced from Jasmonic Acid Source: Wasternack, C. ,(2007).Annals of Botany.100:681-697
  • 6. DIFFERENT PHYSIOLOGICAL ROLES OF JA S. No Process Putative Signals Alteration/Species 1 Root Growth JA,JA-Ile Inhibition 2 Seed Germination JA Inhibition 3 Tuber Formation JA-OH,JA Induction/Potato 4 Tendril Coiling OPDA Stimulation/Bryonia 5 Nyctinasty JA Stimulation/Albizzia 6 Trichome formation JA Induction/Tomato 7 Senescence JA Stimulation 8 Flower Development JA Anther Development and Induction/Arabidopsis dehiscence Induction/Tomato Female Organ Development Induction/Arabidopsis Filament elongation 12 Biotic Stress JA, JME,OPDA Resistance/Ubiqutitous Source: Wasternack, C. (2007).Annals of Botany.100:681-697
  • 7. STRESS AND ITS TYPES Stress is any change in the environmental condition that may adversely affect the plant’s growth, development and adaptability STRESS BIOTIC ABIOTIC Bacteria Virus Drought Salinity Temperature Fungi Heavy metal Pesticide Light Herbivores Nematodes
  • 8. Fungal pathogen Wounding by herbivores Bacteria Insect attack Virus Biotic stress Nematodes perception Stress signaling Jasmonates ? Downstream activation of stress related/resistance genes
  • 9. CERTAIN FACTS TO PONDER OVER  In India, fungal diseases are rated either the most important or second most important factor contributing to yield loss Grover A. et al., 2003. Current science 84:330-340  Global loss because of pathogens is estimated to be 12% of potential crop production Shah D.M. et al., 1997. Current Opinion in Biotechnology B:208-214  The highest losses, estimated at more than $42 billion per year, occur in vegetables, fruits and rice Shah D.M. et al., 1997. Current Opinion in Biotechnology B:208-214
  • 10. FUNGAL DISEASES AND ECONOMIC LOSSES Total yield Crop Pathogen Disease loss(%) Rice Paricularia oryzae Blast 21 Wheat Puccinia recondita Brown leaf rust 30 Maize Helminthosporium maydis Leaf blight 30 Sorghum Sphacelotheca reiliaria Grain mould 18 Pigeonpea Fusarium udum Wilt 24 Chickpea Fusarium oxysorium Wilt 23 Brassica Alternaria brassiceae Blight 30 Soyabeen Phakospora packyrhizi Rust 23 Potato Phytopthora infestans Late blight 31 Source: Current science, vol.84, No. 3, 10 Feb. 2003
  • 11. ROLE OF JA IN RESISTANCE  Increased endogenous level of JA on treatment of Arabidopsis with necrotrophic fungus A. brassicola Penninckx et al.(1996).Plant Cell.8:2309-2323  Biosynthetic and Signaling Arabidopsis mutants were more sensitive to attack by necrotrophic fungal pathogen Phytium (jar1,coi1 and fad3-2fad7-1fad8 mutants) and necrotrophic bacteria Erwinia carotovora (coi1 mutants) Staswick et al.,(1998).Plant Journal 15:747-754 Vijayan et al.,(1998).PNAS USA 95:7209-7214  Transgenic plants overexpressing JMT and thus higher levels of methyl jasmonate more resistant to necrotrophic pathogen Botrytis cinera Walling (2000 ). Journal of Plant Growth Regulators 19: 195-216  The fad3-2fad7-1fad8 mutant more sensitive to attack by fungal gnat Bradysia and coi1 is more sensitive to Diamond –back moth McConn et al.,(1997). PNAS USA.94:5473-5477; Xie et al (1998). Science 280: 1091-1094 Source: Berger,(2002).Planta 214:497-504 Mutant Biochemical Significance Role of Mutant Expression of JA responsive Effects genes coi Coronatine insensitive JA Signaling Reduced Sensitivity to insect and necrotrophic pathogens increased fad Defective Fatty Acid Desaturase JA Biosynthetic Reduced Sensitivity to insect and necrotrophic pathogens increased cev Constitutive Expression of VSP JA Signaling Increased Increased sensitivity to biographic pathogens jar JA Resistant JA Signaling Reduced Increased sensitivity to necrotrophic pathogens opr Defective OPDA Reductase JA Biosynthetic Reduced Increased sensitivity to insects jin JA Insensitive JA Signaling Reduced Increased sensitivity to pathogens
  • 12. Mechanism of action of JAs in biotic stress Source:Wasternack (2007).Annals of Botany.100:681-697 Source:Farmer and Ryan(1992).The Plant Cell.4:129-134
  • 13. OVERALL MECHANISM OF SYSTEMIC RESPONSE  Wounded leaves produce an 18-amino acid peptide called systemin from carboxyl terminal of prosystemin (200 AA precursor) in PP and it elicits production of JA in companion cell-sieve element complex.  JA moves throughout the plant in the phloem. Covalently modified JA (JA-x) play important role in systemic signaling.  signal is recognized at the target cell e.g. mesophyll (leaf)  Jasmonic acid turns on defense related genes (genes for proteinase inhibitor etc.) in target cells. Source: Schilmiller and Howe (2005). Current Opinion in Plant Biology . 8:369-377
  • 15. AMPLIFICATION OF SYSTEMIN SIGNAL  Amplification of wound signaling is a major event in systemic defense Wounding  Systemin activates AOC which in turn activates Prosystemin through OPDA/JA. Prosystemin OPDA/JA again activates back the AOC.  In systemic response once JA is Systemin synthesized, its cyclic production of JA can occur by positive feedback mechanism AOC  JA is a systemic signal that leads to systemic expression of genes encoding OPDA/JA Proteinase inhibitors(PINs)  The plant may develop some resistance Prosystemin against subsequent herbivore attack. Systemin  Thus JA signaling is necessary in systemic response in leaf. Fig: Amplification of wound signaling of tomato Source:Wasternack et al(2006).163: 297-306
  • 16. SYSTEMIC RESISTANCE AND ROLE OF JAs Systemic Acquired Resistance (SAR) Systemic resistance in plants Induced Systemic Resistance (ISR) SAR:  Initial infection result in formation of necrotic lesions  SA concentrations increase and methyl-SA is released in distal plant tissue  PR proteins in the non-invaded parts of the plants are synthesised resulting in reduction in disease symptoms after subsequent infection of many pathogenic species. ISR:  Non pathogenic root colonizing rhizobacteria e.g. P. fluorescens or wounding initiates ISR  ISR does not depend on SA and PR protein  ISR requires both JA and ethylene signalling  SAR regulatory protein NPRI is required  Systemin is involved in ISR which is absent in SAR. Source:Harman et. al.(2004) .Nature Reviews Microbiology. 2:43-56
  • 17. Overall Mechanism of Defence mediated by different signaling compounds Source: Piertse et.al.(2009).Nature Chemical Biology.5:308-316
  • 18. TRANSCRIPTIONAL REGULATION OF JA Four major of interacting players of: 1. a JA signal 2. the SCF-type E3 ubiquitin ligase SCFCOI1 3. Jasmonate ZIM-domain (JAZ) repressor proteins that are targeted by SCFCOI1 for degradation by the ubiquitin/26S proteasome pathway ,and 4. transcription factors (TFs) Howe et. al.(2008)Current Opinion in Plant Biology 11:428-235 Transcriptional regulation mechanism: 1. Expression of regulatory TFs for JAs 2. Cross talk with the TFs responsible for expression of other hormones. Fig: Transcription factors involved in signalling pathways of JA Source: Wasternack(2007). Annals of Botany100:681-697
  • 19. Cross Talk with Ethylene and JA •The antagonistic action of MYC2 and ERF1 may cause independence between wound signaling and pathogen defence signaling Lorenzo et al (2004)Plant Cell 16:1938-1950) •JA alone induce the expression of AtMYC2 responsible for the activation of wound response genes and for the repression of pathogen response genes. •The cooperation of ET and JA signals through the induction of ERF1 leads activation of PR genes and to the repression of WR genes •Therefore, the interplay between ERF1 and AtMYC2 allows the plant selection of the correct set of genes in response to these two stresses Schematic Representation TF in response to Pathogens and Wounding Source: Wasternack(2007). Annals of Botany100:681-697
  • 20. Relevance of JAs studies Plant lack an immune system like in animals but posses mechanism that recognizes potential pathogens and initiate defense responses. During their biochemical evolution, the plants are devised with certain magic molecules of defense (secondary metabolites) like JAs. Recent insights into the JAs mediated plant defense cascade and knowledge of key regulators of this will help us to design future crops with increased biotic stress resistance and better adaptability. Higher crop yields might be achieved by increasing the pathogen/insect resistance which can be achieved by manipulating the expression of the key genes involved in JAs biosynthesis and signaling cascades. Feeding the ever increasing human population is biggest social problem/challenge after all.