Ghorbel et al. 2023 Role of jasmonic acid in plants_ the molecular point of view

Plant Cell Reports (2021) 40:1471–1494
https://doi.org/10.1007/s00299-021-02687-4
REVIEW
Role of jasmonic acid in plants: the molecular point of view

Mouna Ghorbel1,2 · Faiçal Brini2 · Anket Sharma3 · Marco Landi4
Received: 15 January 2021 / Accepted: 23 March 2021 / Published online: 5 April 2021
© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2021
Abstract
Key message Recent updates in JA biosynthesis, signaling pathways and the crosstalk between JA and others phy-
tohormones in relation with plant responses to different stresses.
Abstract In plants, the roles of phytohormone jasmonic acid (JA), amino acid conjugate (e.g., JA-Ile) and their derivative
emerged in last decades as crucial signaling compounds implicated in stress defense and development in plants. JA has
raised a great interest, and the number of researches on JA has increased rapidly highlighting the importance of this phyto-
hormone in plant life. First, JA was considered as a stress hormone implicated in plant response to biotic stress (pathogens
and herbivores) which confers resistance to biotrophic and hemibiotrophic pathogens contrarily to salicylic acid (SA) which
is implicated in plant response to necrotrophic pathogens. JA is also implicated in plant responses to abiotic stress (such as
soil salinity, wounding and UV). Moreover, some researchers have recently revealed that JA controls several physiological
processes like root growth, growth of reproductive organs and, finally, plant senescence. JA is also involved in the biosynthesis
of various metabolites (e.g., phytoalexins and terpenoids). In plants, JA signaling pathways are well studied in few plants
essentially Arabidopsis thaliana, Nicotiana benthamiana, and Oryza sativa L. confirming the crucial role of this hormone
in plants. In this review, we highlight the last foundlings about JA biosynthesis, JA signaling pathways and its implication
in plant maturation and response to environmental constraints.
Keywords Environmental stress · Jasmonates · MYC2 · Pathogen stress · Plant defense
Introduction messengers such as C a2+, reactive oxygen species (ROS),

hydrogen sulfide ( H2S), inositol trisphosphate (IP3), nitro-
Plants are threatened by several biotic and abiotic stresses gen oxide (NO) which interacts with several proteins to
that endanger their life. As sessile beings, evolution has enhance plant response to a given stress (Agurla et al. 2018).
helped plants to develop many mechanisms and behaviors ROS-Ca2+ hub have been extensively studied in plants in
to cope with predators and unfavorable life conditions. Per- response to many environmental constraints and controls lots
ception of threat by cellular receptors is transduced into of mechanisms such as osmotic adjustment, hormonal sig-
signals within seconds. Those signals are called second nal transduction (HST), osmoregulation, mineral nutrition,
cell elongation and development as well as programmed cell
death (Demidchik and Shabala 2017). It has been suggested
Communicated by Aryadeep Roychoudhury. that ROS-Ca2+ hub/phytohormone signaling simultaneously
activates the plant stress signaling cascades (Sewelam et al.
* Marco Landi 2016). Moreover, ROS, as the main source of signaling in
marco.landi@unipi.it
plants under unfavorable conditions, interacts with other sec-
1
Biology Department, Faculty of Science, University of Ha’il, ond messengers and signaling components such as Mito-
P.O. box, Ha’il 2440, Saudi Arabia gen Activated protein Kinases (MAPKs), plant hormones,
2
Laboratory of Biotechnology and Plant Improvement, Center and transcription factors (Sewelam et al. 2016). In another
of Biotechnology of Sfax, B.P ‘1177’, 3018 Sfax, Tunisia hand, plant growth, development, and survival are strictly
3
State Key Laboratory of Subtropical Silviculture, Zhejiang interconnected in a complex biological network that is dis-
A&F University, Hangzhou 311300, China turbed by environmental conditions, such as soil condition
4
Department of Agriculture, Food and Environment and salinity, extreme temperatures, and pathogen attacks
- University of Pisa, 56124 Pisa, Italy
13
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1472 Plant Cell Reports (2021) 40:1471–1494
(Altaf-Ul-Amin et al. 2015). Hence, a relevant number of spp., tomato (Lycopersicum esculentum), and monocotyle-
intermediate compounds, such as ROS and their derivatives, dons (Delker et al. 2006; Kazan and Manners 2013). Many
might accumulate in the cytoplasm and disturb signaling groups described that wounding and systemin increases the
pathways especially when they are converted into toxic endogenous level of JAs including the metabolic precursor,
compounds and affect cell survival (Ayala et al. 2014). Phy- 12-oxo-phytodienoic acid (12-OPDA) and its derivatives JA
tohormones which are involved in ROS-Ca2+ mediated sig- and MeJA to activate the defense response machinery (Wang
nals include abscisic acid (ABA), cytokinins (CKs), auxins et al. 2016).
(IAA), gibberellins (GA), ethylene (ET), salicylic acid (SA),
brassinosteroids (BRs), jasmonic acid (JA) and strigolac- Reactions in the chloroplast
tones (Pottosin et al. 2014; Deng et al. 2018). All those hor-
mones have different effect and can act with an antagonist The JA biosynthetic pathway was studied in Arabidopsis
or synergistic action in response to a stressor. (Ishiguro et al. 2001; Schaller 2001; Feussner and Wast-
Among those phytohormones, JA and its derivatives, ernack 2002; Turner et al. 2002). JA is originally synthe-
such as jasmonyl isoleucine (JA-Ile), methyl jasmonate tized from a ω-3 fatty acid called linoleinc acid (ALA)
(MeJA), 12-hydroxyjasmonic acid sulfate (12-HSO4-JA), (C18:39,12,15). Briefly, the reaction starts when phospho-
cis-jasmone, JA-glucosyl ester, JA-Ile methyl ester, jas- lipases (PLDs) separates phospholipids such as linolenic
monoyl-amino acid, 12-carboxy-JA-IIe, 12-O-glucosyl-JA- (18:2) and α-linolenic (18:3) acid from chloroplast mem-
IIe, 12-O-glucosyl-JA, JA-Ile glucosyl ester, are known as branes (Bargmann et al. 2009). ALA constitutes the most
jasmonates (JAs). JAs are lipid-derived signaling molecules abundant leaf lipids and can be found as galactolipids form-
that control many developmental processes. They are impli- ing the main lipid components of chloroplasts: monogalac-
cated in plant response to biotic and abiotic stress and they tosyldiacylglycerol (MGDG) and digalactosyldiacylglycerol
are detected in almost all plants (Sun et al. 2011). JAs are (DGDG) of the chloroplast (Nilsson et al. 2016). Though the
fatty acids that belong to oxidized lipids family. Those mol- first substrate of the pathway is yet to be identified, it has
ecules are synthesized from cyclo-pentanones and known as been suggested that an esterification then oxygenation of
oxylipins (Wasternack and Feussner 2018). The oxylipins ALA into galactolipid occurs via 13-lipoxygenase (13-LOX;
signaling molecules are biologically active components Fig. 1; (Nilsson et al. 2016)). Moreover, other findings sug-
produced either through auto-oxidation of poly-unsaturated gest that the ALA could be hydrolyzed then converted into
fatty acids or by the activity of lipoxygenases enzymes or 13-HPL (Wang et al. 2019). Allene oxide synthase (AOS)
alpha-dioxygenases enzymes (Göbel and Feussner 2009). is the next acting enzyme in the octadecanoid pathway.
JAs regulates many crucial biological processes in planta, Allene oxide synthase (AOS) is a cytochrome P450 (CYP)
including stamen and trichome development, stomatal open- enzyme that belongs to the CYP74A family. AOS converts
ing, leaf expansion, hook formation, cell cycle regulation, galactolipids into 12,13-epoxylinolenic acid which is rapidly
and glucose transport (Wasternack and Hause 2007; Yoshida transformed to cis (+)-12-oxo-phytodienoic acid (OPDA)
et al. 2009). by allene oxide cyclase (AOC) (Wasternack and Feussner
In this review, we intend to describe the most recent 2018). In tomato, systemin activates a phospholipase A2
updates in JA biosynthesis and signaling pathways, and we (PLA2) that operates the release of LA from membrane
focus on the crosstalk between JA and others phytohormones lipids after leaves are wounded (Narvaez-Vasquez et al.
to understand the mechanism of plant responses different 1999). For example, systemin a peptide which is involved in
stresses. the expression of JA-, wound- and insect-inducible protein-
ase inhibitors (PIs) was found in tomato but not in tobacco
or Arabidopsis (Sun et al. 2011). Mechanisms of JA biosyn-
JA metabolism thesis in Arabidopsis are summarized in Fig. 1.
In monocotyledons, JA biosynthetic pathway is almost
JAs are ubiquitous molecules detected in higher plant spe- conserved with dicotyledons. In fact, some works confirmed
cies (Yang et al. 2019). The endogen level of JA in plants the involvement of many enzymes such as PLDs in JA bio-
is higher in reproductive organs as well as in flowers but synthetic (Nahar et al. 2011). Knockdown of PLD enzymes
it is very low in roots and mature leaves (Dar et al. 2015). involved in JA biosynthetic pathway such as OsPLDα4
In recent years, the role of JA as an important phytohor- (Os06g0604200) and OsPLDα5 (Os06g0 604300) con-
mone implicated in plant development under normal and fers herbivory susceptibility to rice (Qi et al. 2011). As
stress conditions has emerged (Jang et al. 2019, 2020). enzymes implicated in the reduction of ODPA are located
Different studies have been conducted on plants belonging in the peroxisomes, this cyclopentenone must be exported
to different families such as Brassicacea (the model plant from chloroplast and transported to peroxisomes. This is
Arabidopsis thaliana), Solanaceae (tobacco: Nicotiana ensured by an ABC transporter located in the peroxisomal
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Plant Cell Reports (2021) 40:1471–1494 1473
Fig. 1 A proposed model of JA biosynthesis in Arabidopsis. AOS and allene oxide cyclase (AOC) led to the production of 12-oxo-
enzyme is encoded by a single gene while four genes encode for the phytodienoic acid (12-OPDA). 12-OPDA is then translocated from
cyclase (AOC1–4). Growth and development processes as well as chloroplasts via JASSY proteins and penetrates into peroxysome via
extracellular stimuli activate the octadecanoid pathway for JA bio- CTS channels where it is reduced to 3-oxo-2(29[Z]-pentenyl)-cyclo-
synthesis with a so-far unknown mechanism. The first step in JA bio- pentane-1-octanoic acid (OPC:8). OPC:8 undergoes three rounds of
synthesis takes place in the choloroplast where we notice a sequential b-oxidation to yield JA. After biosynthesis, JA is conjugates with ILE
activity of several enzymes involved in the initial steps of JA biosyn- and transported to the cytoplasm where is could be transported all
thesis including lipoxygenase (LOX), allene oxide synthase (AOS), over the plant or could be inactivated via CYP94B3 proteins
membrane called COMATOSE (CTS) and a possible impli- 2019). In jassy mutant plants, ODPA synthesis is normal
cation of an ion trapping mechanism was proposed (Theo- but its export out of the chloroplast was defected. However,
doulou et al. 2005). Recently, a novel protein called JASSY evidence that ODPA pass through a JASSY channel with
was described and it presented an export activity toward OPDA-regulated voltage-dependent gating is still missing.
OPDA (Guan et al. 2019) and was found in the chloroplast Furthermore, little information is available about specific-
outer envelope (OE) membranes (Simm et al. 2013). JASSY ity for OPDA-related compounds (Wasternack and Hause
proteins, which are exclusively located in the chloroplastic 2019). Thus, a link between JASSY proteins and JA biosyn-
OE, could have a role in binding and/or transport of hydro- thesis genes was suggested.
phobic molecules as it contains a steroid acute regulatory
protein-related lipid transfer (START) domain. JASSY Reactions in peroxisomes
mutants represent an altered phenotype (growth inhibition
and decrease in cold stress tolerance) is correlated with a The OPDA cyclopentenone component (12-oxo-phytodien-
decreased level of transcription factors INDUCER OF CBF oic acid) is scaled down to cyclopentanone using the OPDA
EXPRESSION1 (ICE1) and C-REPEAT BINDING FAC- reductase (OPDR; 12-oxo-phytodienoic acid reductase). The
TOR 3 (CBF3) involved in the cold acclimation response). cyclopentenone ring is reduced by the AOC enzyme (Fig. 1).
Moreover, jassy mutant had an increased susceptibility to The (9S,13S)-OPDA [cis-(+)-OPDA] is the only natural
infection by the necrotrophic pathogen Botrytis cinerea and component synthetized in nature (Wasternack and Feussner
decreased expression of corresponding marker genes, such 2018). In another hand, OPDA could act as a signaling mol-
as PLANT DEFENSIN1.2 (PDF1.2). In conclusion, after ecule and thus activates gene transcription independently of
wounding, a lack of JA response (associated with reduced its conversion into JA (Taki et al. 2005). Another JA syn-
level of JA biosynthesis) and a reduced JA biosynthesis thetic pathway was also revealed using Arabidopsis OPDA
genes activation, as well as transcription factors involved in reductase 3 (OPR3) mutant which revealed another inde-
JA responses, were observed in the jassy mutant (Guan et al. pendent JA pathway using 4,5-didehydrojasmonate (Fig. 1;
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Chini et al. 2018). OPR could also regulate seed germina- of the ω-end of the pentenyl sidechain catalyzed by specific
tion in Arabidopsis (Dave et al. 2011). In maize, it has been enzyme, CYP94 (Kitaoka et al. 2014). In the first step, JA-Ile
shown that OPR proteins are crucial for plant development is hydroxylated to 12-hydroxy JA-Ile. This can then either
and immunity (to pathogens and insects; Dave et al. 2011). undergo further oxidation to produce the 12-carboxy-JA-Ile
Once synthetized, JA (or its active forms, e.g. MeJA) are (Koo et al. 2014), or else can be glycosylated to generate
transported all over the plant and in the phloem and xylem 12-O-glycosyl-JA-Ile (Fig. 2; Haroth et al. 2019).
of vascular bundles. This transport is ensured via a protein
transporter called GTR1 (Ishimaru et al. 2017; Matsu et al. JA conversion into signaling molecules
2017).
Me‑JA
Reactions in cytoplasm
Me-JA is synthesized via the acceptation of methyl group
In Arabidopsis, works conducted on jar1-1, a JA-insensitive from S-adenosyl-l-methionine (SAM) to JA. This reaction
mutant revealed that the active form of JA was an amino acid is catalyzed by the SAM: JA carboxyl methyl-transferase
conjugate. The isoleucine is the principal amino acid that enzyme. This esteric compound formed is volatile in con-
conjugates with JA to form JA (JA-Ile) synthetized in the trast to JA (Sanchez-Aguayo et al. 2004). Me-JA has been
cytoplasm (Fig. 1). Other amino acid could rarely conjugate detected in plant leaves after an herbivore attack. Interest-
with JA such as valine, leucine, and phenylalanine (Staswick ingly, this molecule is detected only in leaves that have not
and Tiryaki 2004). been predated. Thus, this molecule can serve as an airborne
warning molecule signal (Wasternack and Feussner 2018).
JA catabolism In plants like potato and tomato, the production of PIN II
protease inhibitor is one of the common markers for JA
JA-Ile metabolism is ensured by two major routes: hydro- pathway activation (Singh et al. 2020). Those inhibitors act
lyzation and oxidation (Fig. 2; Bruckhoff et al. 2016). JA-Ile as an anti-feedant compound as it interferes with digestive
hydrolyzation allows the liberation JA and Ile from JA-Ile. proteins in the insect gut leading to lower food availability
This is controlled by an aminohydrolase (AMH) enzyme. for the insect and hence the insect take more time to grow
After that, JA is subjected to a hydroxylation, controlled by and complete its life cycle (Singh et al. 2020).
a P450 enzyme to generate 12-hydroxy-JA-Ile that could The consequence of insect growth delay is that weather
be oxidized later to generate carboxy-derivative. In another changes may occur and hence reduce the survival of the
hand, JA-Ile could also be subjected to different oxidations insect (Bayram and Tonga 2018). In addition, Me-JA is one
Fig. 2 Catabolism of jasmonic

acid (JA). JA could be inacti-
vated via two major pathways:
hydrolysation (I) and oxydation
(II). Hydrolyzes is unsured
by amidohydrolase (AMH) to
release Ile and JA. Then JA
could be either hydroxylated (to
12-hydroxy-JA), glycosylated,
methylated to form methyl JA
(Me-JA) or decarboxylated to
synthesize cis-jasmone (CJ).
JA-Ile could also undergo
sequential oxidation reaction to
generate 12-carboxy-JA-Ile. JA
could also be glycosylated to
generate 12-O-glycosyl-JA-Ile.
Enzymes that catalyses the reac-
tions are shown in red (colour
figure online)
13
of complex volatiles compounds released during plant infec- tobacco, tomato, and soybean (Scheer and Ryan 2002; Li
tion that could play the role of chemical attractor for insect et al. 2004).
predators such as wasps as demonstrated for Triticum aes-
tivum (Bayram and Tonga 2018). Moreover, treatment with The AtMYC2: master regulator of JA signaling
Me-JA stimulates phytoalexins compounds such as alkaloids
terpenoids and glycosteroids which have an antimicrobial MYC2 structure
activity (Shi et al. 2015), enhances carotenoid biosynthe-
sis as demonstrated for maize (Luo et al. 2019, 2020) or Several works have classified the basic helix-loop-helix
stimulates sucrose degradation. Moreover, MeJA changes (bHLH) transcription factor (TF) as the greater JA signaling
enzymatic activity of many enzymes of the phenylpropanoid pathway in Arabidopsis (Browse 2009; Chini et al. 2009a;
pathway (Sadeghnezhad et al. 2020). Me-JA also decreased Howe 2010; Pauwels and Goossens 2011; Wager and Browse
the amino acid biosynthesis rate thus it controls the cell 2012). The best characterized TF in this group and the most
growth rate (Sadeghnezhad et al. 2020). multifunctional in the JA signaling pathway in Arabidopsis
is MYC2 (Kazan and Manners 2013). This nucleolar protein
cis‑Jasmone (CJ) (Chini et al. 2009b) presents the general characteristics of
bHLH TF protein family (Toledo-Ortiz et al. 2003; Pires
CJ is used in perfume industry as an important ingredient. and Dolan 2010) such as the conserved bHLH domain at its
This natural compound is liberated from flowers of differ- carboxyl domain. Those domains are formed by two amphi-
ent species such as jasmine, neroli (Citrus bigaradia; Matsu pathic alpha-helices consisting of hydrophobic amino acids
et al. 2017), jonquil (Narcissus jonquilla L.; Dabrowska and united by a loop region important to form homo/heterodi-
Boland 2007), bergamot (Citrus bergamia), and members of mers with other TFs like the most closed TFs MYC3 and
the Pittosporum family (Griffiths 2020). After plant infec- MYC4 (Fernandez-Calvo et al. 2011). It has been suggested
tion, CJ is released, and it could act like JA. It can act as an that the C-terminal leucine zipper domain could function
insect repellent or it could participate to attract insect preda- as a supplementary dimerization domain to increase the
tors (Matthes et al. 2010). In fact, exogenous application of specificity of cooperation with other TFs (Amoutzias et al.
CJ to wheat was effective against cereal aphids and some 2008). Moreover, the 15–20 mostly basic amino acids are
phytophagous thrips species as well as wheat stem sawflies involved in linking to the G-box (5′-CACGTG-3′) found in
Bayram and Tonga 2018). Moreover, foliar application of CJ MYC2 target promoters (Toledo-Ortiz et al. 2003; Pires and
increased volatile organic compounds (VOCs) production as Dolan 2010; Fernandez-Calvo et al. 2011; Amoutzias et al.
well as the upregulation of genes encoding them. Besides, 2008; Carretero-Paulet et al. 2010). In the N-terminal part of
CJ negatively affect tomato pest Spodoptera exigua repro- the MYC2 TF, a putative transcriptional activation domain
duction (Disi et al. 2017). (TAD) is found, which is important for transcription initia-
tion (Fernandez-Calvo et al. 2011; Amoutzias et al. 2008;
Carretero-Paulet et al. 2010). Moreover, MYC2 and their
The signaling pathway of JA closely related proteins, MYC3, and MYC4, interplay with
the C-terminal part of JAZ proteins (JAS domain) through
The SCFCOI1 protein and JA signaling JID domain (JAZ Interaction Domain) (Fernandez-Calvo
et al. 2011; Amoutzias et al. 2008; Carretero-Paulet et al.
Experiments conducted with the Arabidopsis mutant coi1 2010; Chini et al. 2007). Some works proposed that protein
(coronatine insensitive 1) add knowledge on the comprehen- phosphorylation is involved in the modulation of JA signal-
sion of the JA response pathway (Feys et al. 1994). COI1 ing (Hiruma et al. 2011). Though MYC2 was shown to be
was identified as an F-box protein; it is closely related to the phosphorylated, the identity of kinase protein implicated in
auxin receptor/F-box protein TIR1 (Xie et al. 1998). Thus, this phosphorylation is still unknown (Zhai et al. 2013).
the authors suggested that it could act as part of an SCF
(Skp/Cullin/F-box) E3 ubiquitin ligase to mediate JA signal- MYC2 and JA response
ing (Xie et al. 1998). In fact, COI1 was demonstrated to be
associated with components of the SCF complex, including MYC2 orchestrates the JA pathway by controlling several
ASK1, ASK2, CUL1, and RBX1 (Xu et al. 2002). Moreover, mechanisms such as plant response to pathogen infection,
in Arabidopsis, other works suggest that SCFCOI1 protein abiotic stress applications and developmental processes
targets the repressor proteins of JA signaling for degrada- (Kazan and Manners 2013). It tightly regulates the expres-
tion and, therefore, activates JA-modulated changes at tran- sion of downstream transcriptional activators as well as JAZ
scriptional level. The same founding was also reported in repressors that, in turn, act upstream from MYC2. In fact,
myc2 mutant Arabidopsis seedlings had reduced expression
13
of JAZ genes comparing with wild-type proteins in response at least 100 proteins as revealed by The Arabidopsis thali-
to JA treatment (Chini et al. 2007; Grunewald et al. 2009). ana Promoter Analysis Net, network (AtPAN) (http://atpan.
Later, it has been demonstrated that MYC2 directly binds to itps.ncku.edu.tw/), conclude more than 100 different MYC2
the promoter of JAZ3 (Chini et al. 2007) and JAZ2 (Figueroa cooperator (Chen et al. 2012). The most important studied
and Browse 2012) confirming that the MYC2 TF is a regu- interactome are resumed in Table 1.
lator of JAZ gene expression. MYC2-mediated transcrip-
tional activation and JAZ repressors synthesis coordinate Jasmonate ZIM‑domain proteins: inhibitor of JA
to degrade the JAZ proteins (Chini et al. 2007; Demianski response
et al. 2012). MYC2 TF controls the expression of a high
number of JA-responsive genes including TFs (Guo et al. Eight different JAZ proteins (jasmonate ZIM-domain pro-
2012; Dombrecht et al. 2007). In fact, several researchers teins) were identified in opr3 mutants during JA response
have identified that MYC2 presented a dual effect on the JA (Thines et al. 2007). Those proteins harbor two conserved
signaling pathways as it acts as a transcriptional activator as interaction domains, called ZIM and Jas (Thireault et al.
well as a repressor in the regulation of JA signaling network 2015). The Jas domain controls JAZ interaction with COI1
(Fig. 3). In fact, it has been demonstrated that a comparative and with other TFs as well (Gimenez-Ibanez et al. 2014).
microarray analysis using JA-treated and untreated myc2 and Moreover, the ZIM (TIFY) domain regulates the JAZ
wild-type plants. The analysis identified many JA-respon- dimerization and interaction with other interactives of JAZ
sive genes that are directly or indirectly managed by MYC2 (NINJA) proteins which requires TOPLESS (TPL) proteins,
all along JA signaling (Dombrecht et al. 2007). Moreover, transcriptional co-suppressor, via the interaction with the
bioinformatic analyzes showed that MYC2 interacts with conserved EAR domain (Zhang et al. 2015). Importantly,
Fig. 3 A simplified schematic presentation of jasmonate (JA) signal- ily proteins interaction. This leads to JAZ ubiquitination and degrada-
ing. After its biosynthesis in plastids, JA is conjugated with aa, espe- tion with the 26S proteasome. MYC2 TFs interacts with the MED25
cially Ile by JAR under the control of JAR enzymes in the cytoplasm. subunit of the moderator complex then interacts with the G-box motif
In some conditions, JAZ interacts with NINJA, TOPLESS (TPL), and of the target promoters, which allows the activation of JA-responsive
histone deacetylase (HAD) proteins to repress JA gene expression via genes. In another hand, IIId bHLH factors antagonize MYC2 action
inhibiting MYC2 TF. It also inhibits many negative regulators of via competitive binding to the G-box motif and inhibiting JA-respon-
JA signaling such as IIId bHLH factors. In presence of extracellular sive genes. Thus, succeeding transcription factors are stimulated,
stimuli or in growth and development processes, JA-Ile enhances the allowing them to enhance JA-responsive early genes activities and JA
CORONATINE INSENSITIVE1 (COI1)/JA ZIM-domain (JAZ) fam- responses
13
Table 1 The most identified MYC2 interacting proteins in Arabidopsis thaliana

Protein Role References
All known JAZ repressors (ex: JAZ3 and JAZ2) Repression activity of MYC2 Chini et al. (2007), Figuerosa
and Browse (2012)
MED25 subunit of the plant Mediator complex Both transcriptional activation and Chen et al. (2012)
repression activity of MYC2
bHLH TFs (MYC3 and MYC4) ? Fernandez-Calvo et al. (2011)
uncharacterized (e.g. At5g57150, At1g68920, At3g61950, At1g01260, ? Chen et al. (2012)
and At3g57800) TF
PIF4 (PHYTOCHROME INTERACTING FACTOR4) ? Chen et al. (2012)
PAP3 (PHYTOCHROME-ASSOCIATED PROTEIN3) ? Chen et al. (2012)
CIB5 (CRYPTOCHROME-INTERACTING BASICHELIX-LOOP- A regulator of cryptochrome signaling Chen et al. (2012)
HELIX5)
ICE1 (INDUCER OF CBF EXPRESSION1) A regulator of abiotic stress tolerance Chen et al. (2012)
BIM1 A regulator of brassinosteroid signaling Chen et al. (2012)
ALC (ALCATRAZ) A regulator of plant development Chen et al. (2012)
in vitro and in vivo assays demonstrated that JA-Ile, not of JA signaling. Those proteins play a crucial role in plant
other derivatives, mediates direct JAZ1 and COI1 interac- extension and maturing as well as its reply to environmental
tion (Thines et al. 2007). In response to JA, many JAZ pro- stresses (Gimenez-Ibanez et al. 2014; Dubois et al. 2018).
teins, such as JAZ3, are fixed with SCFCOI1 to mediate their JAZ interact with many TFs and specifically regulate many
proteasome degradation (Chini et al. 2007). Moreover, in responses (Chini et al. 2016). In Arabidopsis thaliana, when
absence of JA, JAZ directly interacts with AtMYC2 which endogenous jasmonate concentration increase, JAZ proteins
leads to transcriptional repression of JA-responsive genes interact with SCFCOI1 ubiquitin ligase which leads to JAZ
(Chini et al. 2007). degradation by proteasome 26S (Chini et al. 2007). Thus,
degradation of JAZ protein liberates COI1 and AtMYC2 to
Identification of NINJA proteins enhance jasmonate-dependent gene expression (Chini et al.
2007; Thines et al. 2007). Defense compounds increases
As discussed above, physical interplay between JAZ repres- in presence of JAZ-MYC module. Thus, this could stimu-
sor proteins and AtMYC2 regulates JA-responsive gene late defense response against pathogen infection or inhibit
expression. This requires the interaction between NINJA plant growth (Havko et al. 2016). Moreover, COI1/JAZ2/
proteins and Groucho/Tup1-type co-repressor TOPLESS MYC2, 3,4/ANAC19,55,72 and other modules have been
(TPL) and TPL-related proteins (TPRs) with the presence identified to enhance plant response to endogenous or exog-
of unknown adaptor protein. TPL and NINJA proteins act as enous stimuli (Gimenez-Ibanez et al. 2014; Mao et al. 2017).
negative regulators in the JA signaling pathway. NINJA acts MYCs–JAZs interactions could trigger the hormonal cross
as a transcriptional repressor whose activity is mediated by talk pathways like ET-mediated cell division throughout ET
a functional TPL-binding EAR repression motif (Pauwels RESPONSE Transcription factor (ERF) (Gimenez-Ibanez
et al. 2010). et al. 2014; Zhang et al. 2015; Dubois et al. 2018).
SCFCOI1/JAZ/ AtMYC2 complex: the JA Core

Signaling Module
Involvement of JA in plant growth
Generally, JA-Ile endogenous level is low in normal condi- and development
tions, but they can rapidly increase in presence of a given
stimuli. This increase in sensed by a JA receptor called COI1 Phytohormones control internal developmental interactions
which constitute a fundamental component of SCFCOI1. with entourage stimuli to regulate plant maturation and sur-
COI1 lie to JAZ proteins and enhances JAZ ubiquitination vival. In the last decade, a huge body of studies focused on
and degradation through the 26S proteasome pathway. Thus, understanding the role of JA in plant maturation and devel-
downstream TFs (e.g.: MYCs) can be released to activate opment. An important number of works have shown that
JA responses (Gimenez-Ibanez et al. 2014; Dubois et al. jasmonates are involved in a huge number of plants expand-
2018). As a result, many works identified MYC, JAZ, and ing events, such as primary root growth, leaf senescence,
COI1 protein as the core signal transduction mechanism and reproductive development (Wasternack and Hause 2007;
13
Kim et al. 2015). Moreover, JA is implicated into the regula- (MYC2 and its homologs MYC3/4/5), interact with JAZ pro-
tion of various metabolites production, such as phytoalexins teins (Qi et al. 2015). MYC2/3/4 are in the primary root
and terpenoids. apex; make place to the primary root growth inhibition
by JAs (Gasperini et al. 2015). Moreover, MYC2 inhibits
Effects on seed germination PLETHORA genes (PLT1 and PLT2) expression to minimize
the activity of root meristematic cells and suppress primary
It has been described that some phytohormones like ABA, root growth (Chen et al. 2011). Besides, MYC2 interacts
IAA, and JA play important roles during seed germination with MED25 subunit to unsure this inhibitory effect by
(Xiao et al. 2018). Both ABA and JA inhibit seed germi- inhibiting RNA polymerase II (Chen et al. 2011). It has been
nation but their interactions during this process are still demonstrated that JAZ9 handicaps MYC3–MED25 interac-
elusive (Tang et al. 2020). In Arabidopsis, JA inhibition tions (Gimenez-Ibanez et al. 2014). MYC2 ubiquitination
occurs in a COI1-independent manner (Dave et al. 2011). via PLANT U-BOX PROTEIN10 and its phosphorylation of
In Triticum aestivum plants, cold-stimulated germination MYC2 in presence of mitogen-activated protein kinase 3/6
of seeds resulted in an increase in endogenous level of JA suppresses JA-mediated root maturation inhibition (Chico
after upregulation of JA biosynthesis-related gene and JA et al. 2014; Sethi et al. 2014).
promotes cold-stimulated germination (Xu et al. 2016; Avra- In Arabidopsis, ETHYLENE INSENSITIVE (EIN) 3 and
mova 2017). Recently, a novel SAPK10-bZIP72-AOC’ path- EIN3-LIKE1 (EIL1) are primordial TFs acting in ethylene
way was identified in rice. In this pathway, ABA stimulates signaling. Those TFs interact with JAZ proteins and posi-
JA biosynthesis to synergically inhibit the germination of tively mediate the JA-dependent primary root growth inhi-
rice seeds (Wang et al. 2020). Some authors demonstrated bition as well as JA-induced root hair formation (Zhu et al.
that SAPK10 was activated by auto-phosphorylation, then 2011). In Arabidopsis, JA stimulates also the lateral root
this molecule stabilizes by phosphorylation bZIP72 TF and formation as it upregulates ERF109 expression which stimu-
promotes it’s binding to the G-box cis-element of AOC pro- lates ANTHRANILATE SYNTHASE Α1 (ASA1) and YUCCA2
moter to enhance AOC transcription in presence of elevated genes implicated in auxin biosynthesizes (Cai et al. 2014).
concentrations of JA. Interestingly, alleviation of ABA sensi- In contrast, JA negatively regulates adventitious root forma-
tivity was recorded after blocking of JA biosynthesis (Wang tion via the COI1-MYC2/3/4 cascade (Gutierrez et al. 2012).
et al. 2020).
Inhibition of root growth Regulation of stomatal closure
Mutations in the co-receptor for JA-Ile COI1 make plants Stomata control gas exchange, water loss in plants. It also
insensitive to the prevention of primary root maturation regulates plant immunity against phytopathogens. Stomatal
by JAs (Yan et al. 2009). Moreover, inositol pentakispho- closure is also mediated by JA. In fact, Methyl-JA/ COI1
sphate (InsP5) enhances COI1–JAZ9 interactions and thus activate a plasma membrane H+-ATPase. Membrane depo-
the blocking effect of JAs on root expansion and maturation larization generates an influx of Ca2+ and an efflux of H+
(Mosblech et al. 2011). In another hand, coronatine-O-meth- (Yin et al. 2016). Methyl-JA/ COI1 induces also the produc-
yloxime, a competitive JA antagonist obstruct COI1–JAZs tion of ROS, the stimulation of Cl– channels (which engen-
interactions and thus suppress the inhibitory outcome of der an efflux of C l–) and the stimulation of K
+ efflux (via
+
coronatine on primitive root growth (Monte et al. 2014). In K channels), causing turgor guard cell loss and stomatal
Arabidopsis, most JAZ proteins (13 members) lack the ERF- closure (Yan et al. 2015) Drought and salt stress conditions
associated amphiphilic repression (EAR) domain. Thus, they induce stomatal closure in plants. It has been proven that JA
should interact with TPL and TPL-related proteins (TPRs) and ABA pathways control stomatal closure in Arabidop-
via the EAR of NINJA proteins to crush JA responses. Inter- sis through the stimulation of OPEN STOMATA1 protein
estingly, the non-canonical JAZ proteins, JAZ8 and JAZ13 kinase (Yin et al. 2016). Moreover, in drought conditions,
do not require NINJA protein to interact with TPLs /TPRs plants synthesizes OPDA rather than jasmonate. This mol-
but use directly their one EAR domain (Thireault et al. 2015; ecule is more efficient than JA in such conditions. OPDA
Chini et al. 2016). Interestingly, overexpressed NINJA pro- interplay with ABA to enhance stomatal closure in Arabi-
teins or modified JAZ proteins (carrying a deletion, muta- dopsis (Savchenko et al. 2014).
tion) in Jas domain suppressed the inhibitory effect of JA on Plant infection with Pseudomonas syringae induces also
primary root growth. But this inhibitory effect was stimu- stomatal closure to limit bacterial invasion (Melotto et al.
lated by the NINJA/TPL or combined mutations in JAZ7, 2006). To protect themselves, many P. syringae strains syn-
JAZ8, JAZ10, and JAZ13 (Thireault et al. 2015; Thatcher thesize coronatine which interferes with COI1-JAZ-MYC
et al. 2016). In Arabidopsis, bHLH type transcription factors signaling and prohibits SA intensification via NAC019/
13
NAC055/NAC072 and JA2L, enhancing stomatal reopen- but not in JA signaling mutants (Jewell and Browse 2016).
ing to help bacterial infection (Zhu et al. 2015). Moreover, Moreover, in coi1 background, re-imposed COI1 expression
HopX1 and HopZ1a, effectors of P. syringae, deplete JAZ in many tissues such as filaments epidermis or anthers can
proteins in a COI1-independent manner, and maintain sto- retrieve anther dehiscence, filament elongation, and pollen
matal apertures to promote infection (Gimenez-Ibanez et al. maturation (Jewell et al. 2016). R2R3-MYB TFs such as
2014). MYB21, MYB24, and MYB57 interact directly with JAZs
(Song et al. 2011). The myb21 myb24 double mutant rep-
Delay of flowering resents a delayed anther dehiscence, a non-viable pollen
grains and short filaments. Coi1-1 plants overexpressing
In Arabidopsis, vegetative-reproductive maturation transi- MYB21 proteins represent a restored stamen development
tion is inhibited by JA. COI1–JAZ interaction inhibits flow- (Song et al. 2011). MYB21 and MYB24 connect physically
ering. In fact, coi1 mutant as well as JAZ1Δ3A transgenic with MYC2, MYC3, MYC4, and MYC5 to control stamen
plants presented an early flowering. Moreover, TARGET OF development (Qi et al. 2015). Interestingly, overexpression
EAT TFs (TOE1 and TOE2), APETALA2/ERF domain TFs of MYC5 and MYC3 in coi1-1 plants can restore stamen
type, control plant flowering. In fact, their interaction with maturation and productivity (Qi et al. 2015).
JAZ proteins inactivates the transcription of FLOWERING
LOCUS T and thus inhibits flowering. In another hand, over- Inhibition of petal expansion
expression of TOE1 and/or TOE2 inhibits coi1 early flower-
ing phenotype (Zhai et al. 2015). In Arabidopsis, JA inhibits petal expansion via COI1 path-
way. In fact, coi1 mutants represent larger petals compar-
Inhibition of hypocotyl growth ing with wild type at plant anthesis. The same observation
was registered for Arabidopsis JA-deficient mutants’ aos
In Arabidopsis, JA inhibits hypocotyl elongation in many and opr3 (Reeves et al. 2012; Brioudes et al. 2009). JA
light stress conditions such as far-red and blue wavelengths. represses MYB21 and MYB24 proteins during sexual organs
This is ensured through COI1 (Chen et al. 2013). Studies formation which lead to the restriction of petal growth con-
showed that JA-deficient mutant jar1 plants grown in far-red firming that MYB21 and MYB24 are important for petal
light conditions represented elongated hypocotyls (Robson expansion (Reeves et al. 2012). Interestingly, aos and coi1
et al. 2010). Moreover, coi1 mutants have longer hypocotyls plants showed an increased MYB21 expression level in petals
compared to WT plants when they were grown in dark or leading to continuous petal extension and thus grand petals
under fared lights conditions (Chen et al. 2013) or under (Reeves et al. 2012). Moreover, opr3 plants represent an
red light or low R/FR light ratio (Chen et al. 2013; Robson increased bHLH TF BIGPETALp expression level. This TF
et al. 2010). The same observation was registered for the controls post-mitotic cell expansion and unsure larger petals
myc2/jin1 mutant grown under far-red light and a low R/ and larger cells in this mutant (Brioudes et al. 2009).
FR light ratio (Robson et al. 2010), but those plants show
a shorter hypocotyl under blue light (Yadav et al. 2005). Promotion of trichome formation
Authors suggest that bHLH type TF MYC2 plays a role in
plant response to red or far-red light and positively regu- Trichomes are barriers or sensors that protect plants from
lates the repression of hypocotyls expansion, but it nega- herbivore attack. They can also release volatile compounds.
tively controls the inhibition of hypocotyls elongation by In Arabidopsis, it has been demonstrated that defect in JA
blue light. In rice, JA represses coleoptiles maturation and biosynthesis or perception can block trichome formation
plant height. In Zea mays, JA inhibits ear shoot maturation after wounding (Yoshida et al. 2009). At low endogen JA
(Yan et al. 2012; Yang et al. 2012). level in cells, JAZ proteins physically connect with WD-
repeat/bHLH/MYB complexes to inhibit their transcriptional
Stamen development in Arabidopsis activity (Qi et al. 2011). After wounding or pathogen attack,
endogenous JA concentration increase leading to JAZ pro-
In Arabidopsis, many male sterile plants were described. teins turn over. In such situations, WD-repeat/bHLH/MYB
Among those mutants, JA-deficient mutants were described. complexes enhance trichome formation. Subgroup IIIdb-
In fact, coi1, lox3 lox4, aos, opr3; fad3 fad7 fad8, defec- HLH TFs inhibits WD-repeat/bHLH/MYB complexes action
tive in anther dehiscence1 (dad1), JAZ1Δ3A and JAZ10.4 by binding to the promoters of their putative target genes
mutants as well as CYP94B3 overexpression lines repre- competitively leading to the inhibition of trichome formation
sented incomplete stamen development (Song et al. 2013). (Nakata et al. 2013).
Interestingly, exogenous application of JA restores stamen
development only in plants deficient in JA biosynthesis
13
Inhibition of apical hook formation control plants (Awan et al. 2020). Exogenous application of
ABA and JA weaken drought stress effects on pearl millet
COI1-JAZs-MYC2/3/4 cascade inhibits the formation of plantlet by increasing the fresh and dry weight of seedlings,
apical hook of dark grown plants (Song et al. 2014). In root length and shoot length with a pronounced effect of JA
dark stress conditions, MYC2, MYC3, and MYC4 TFs are comparing to ABA (Awan et al. 2020) as also shown for
activated by JA and physically interact with and repress the soybean and wheat under drought stress (Ruan et al. 2019).
transcriptional activity of EIN3/EIL1. This leads to down- Authors showed that JA have a more pronounced effect of JA
regulation of HOOKLESS1 gene that controls apical hook comparing with ABA in plant stress alleviation (Awan et al.,
formation and to the inhibition of apical hook curvature 2020). This indicates the signaling interaction between ABA
(Song et al. 2014; Zhang et al. 2014). Besides, MYC2 acti- and JA in response to environmental stresses.
vates EIN3 BINDING F-BOX PROTEIN1 expression which Moreover, JA intensifies the antioxidant enzyme (SOD,
leads to EIN3 degradation (Zhang et al. 2014). CAT, and APX) activities with a pronounced effect compar-
ing with ABA hormone treatment in pearl millet (Awan et al.
Induction of leaf senescence and chlorophyll 2020). In contrast, in the absence of allene oxide cyclase
degradation (AOC) gene, which encodes the enzyme involved in JA bio-
synthesis, rice showed enhanced tolerance to water deficit
JA stimulates leaf senescence in Arabidopsis. This effect stress in term of stomatal conductance, water content, and
occurs in COI1-dependent manner (Qi et al. 2015). Moreo- shoot ABA content compared to the wild type (Dhakarey
ver, JAZ7 represses leaf senescence in dark grown plants (Yu et al. 2017). Besides, both hormones, ABA and JA, reveal
et al. 2015). Many NAC TFs members (such as NAC019, drought stress effects by minimizing oxidative damage (seri-
NAC055, and NAC072) act downstream of MYC2/3/4 to ous oxidative impairment of cell membrane), in the leaves.
enhance chlorophyll degradation (Melotto et al. 2006). In fact, exogenously applied ABA and JA improved drought
In another hand, Subgroup IIIdbHLH TFs antagonize stress in Arabidopsis (de Ollas et al. 2015), wheat (Siddiqi
MYC2/3/4 actions as they bind competitively to their tar- and Husen 2019), and pearl millet (Awan et al. 2020) which
get promoters and inhibit leaf senescence (Qi et al. 2015). positively raise MDA and H2O2 contents in those plants.
WRKY57 interacts physically with JAZ4/8 and act as a neg-
ative regulator of leaf senescence in presence of JA (Jiang JA alleviates negative impacts of heavy metals
et al. 2014). YABBY1 and YABBY3 proteins interact with stress
JAZs leading to chlorophyll degradation promotion (Boter
et al. 2015). Heavy metals [manganese (Mn), nickel (Ni), cadmium (Cd),
and lead (Pb)] damage the environment and reduce plant
growth and maturation (Zhang et al. 2018). Many found-
Involvement of JA in plant abiotic and biotic ings revealed that JA controls plant responses to heavy
stress responses metal applications by modulating their antioxidant systems
(Noriega et al. 2012; Carvalho et al. 2013; Ali et al. 2018).
Role of JA in plant drought and oxidative stress In tomato, it has been demonstrated that spr2, a JA-deficient
response mutant accumulated more Cd in roots and leaves compar-
ing with wild-type plants (Zhao et al. 2016). Furthermore,
JAs and MeJAs are known to take part in various physiologi- in Vicia faba, external JA supplementation enhanced plant
cal processes. Exogenous application of JAs so far tested on growth and biomass yielding, restores total chlorophyll con-
different plants under abiotic stresses particularly salinity, tent and carotenoid concentrations, enhances glutathione
drought, and temperature (low/high) conditions have proved reductase activity and proline accumulation (Ahmad et al.
effective in improving plant stress tolerance (Ahmad et al. 2017), reduces Cd accumulation in roots, shoots, and leaves
2016). Several studies conducted on Arabidopsis, barley, by stimulating antioxidant activity and inhibiting MDA and
wheat, maize, and pearl millet showed that both ABA and H2O2 accumulation (Carvalho et al. 2013; Ahmad et al.
JA positively control plant tolerance to drought stress (Awan 2017). Moreover, JA activated ascorbate or glutathione
et al. 2020; Wang et al. 2021). In rice and Arabidopsis, both antioxidant machinery in soybean to inhibit lipid peroxidase
ABA and JA-connected genes were controlled in the first activity (Noriega et al. 2012). In Brassica napus, JA mini-
drought constraint subjection, but showed different figure mized the Cd uptake in leaves, which ensured a reduction in
under the next drought stress cycles, underlying a potential membrane damage and MDA content. This facilitated also
role of those hormones in stress memory (Li et al. 2019). In essential nutrient uptake by roots and increases gas exchange
pearl millet, water deficit conditions diminish root and shoot and photosynthetic pigment contents by protecting chloro-
length as well as their fresh- and dry weight comparing with plasts from damaging effects of Cd (Ali et al. 2018).
13
Exogenous JA application on Glycine max seeding before lights in several plants (Mutar and Fattah 2013; Yang et al.
NiCl2 stress could enhance tolerance to N i2+ stress. This 2015). Recent founding showed that the incidence of Root
positive effect of JA is ensured by protecting total protein Knot nematodes in watermelon was inhibited by diurnal
contents and controlling the antioxidant machinery (Sirhindi red lights comparing to white lights. This plant defense was
et al. 2015). In Zea mays, external application of JA protects closely related to a remarkable increase of many genes (AOS,
plant from Ni toxicity by stimulating antioxidant enzyme PRI, ICS, WRKY70, and LOX) in association with a pro-
activity (Azeem 2018). nounced increase in SA, JA, and H2O2 contents in various
Jasmonic acid alleviates also the nefast effect of lead tissues of watermelon plants (Yang et al. 2018). JA and SA
(Pb) stress. Pb minimizes macro and micronutrients uptake participate to plant defense against nematodes such as M.
by plants roots (Sharma and Dubey 2005). It prevents the incognita (Kammerhofer et al. 2015). This pathogen forces
penetration of many essential cations (such as K +, Zn2+, plants to activate SA pathways and to inhibit JA pathway in
Mn , Mg , Cu , C a , Fe ), and anions (NO3−) in
2+ 2+ 2+ 2+ 3+
leaves to allow a successful pathogen invasion (Martìnez-
the roots (Bali et al. 2019a). Pb causes also a decline in Medina et al. 2017). Interestingly, JA has a higher accumu-
roots and shoots lengths in many plants such as tobacco lation in roots after root knot nematodes infection which is
((Alkhatib et al. 2012) and tomato (Bali et al. 2019a). Bali transported later to the leaves which helps plants to trigger
et al. (2018) showed that JA alleviates Pb effects by enhanc- their defense against nematodes (Zhang and Baldwin 1997).
ing tomato growth. External JA application minimizes Furthermore, plant treatment by red lights enhances JA accu-
Pb uptake by tomato roots. Moreover, this hormone helps mulation in roots and leaves (Yang et al. 2018) confirming
plants to improve their photosynthetic efficiency and gase- the importance of JA signaling in rhizo-bacteria-induced
ous exchange. Furthermore, JA-treated plants exhibits an systemic resistance (Fujimoto et al. 2011). Besides, studies
increased content of osmolytes (glycine betaine, proline, pol- conducted on tomato and watermelon confirms that exog-
yamine, and total carbohydrates). Besides, JA-treated tomato enous MeJA application reduced root knot nematodes infec-
cultivated under Pb stress presented an enhanced accumula- tions (Fujimoto et al. 2011; Yang et al. 2018). Moreover,
tion of metal chelating compounds (total thiols, non-protein due to the antagonistic roles between SA and JA, red lights
thiols, and protein thiols) (Bali et al. 2018, 2019b). inhibit SA accumulation in leaves (Yang et al. 2018) but
stimulated its accumulation in roots after nematode infec-
Role of JA in plant resistance against nematodes tion underlying the positive effect of red lights on SA accu-
mulation on roots (Branch et al. 2004; Yang et al. 2018).
Recently, the infection of watermelon plants (Citrul- Meanwhile, red lights stimulate JA accumulation in roots
lus lanatus L.) by a soil-dwelling, microscopic nematode demonstrating a positive regulation of JA biosynthesis genes
called Meloidogyne incognita was investigated. Plants by red lights; such results could indicate a synergic interac-
infection leads to apparition of galls on roots which can tion (van Wees et al. 2000; Yang et al. 2018). This is shown
enhance plant sensitivity to infection by other pathogens at the molecular level by transcript accumulation of ICS
(Kyndt et al. 2013). After nematodes invasion, plants acti- gene (for SA biosynthesis) and LOX and AOS genes (for JA
vate different defense strategies such as ROS, hormones, biosynthesis). In parallel, red lights induces accumulation
and activation of appropriate defense-related genes (Melillo of WRKY70 transcript and H2O2 particles after root knot
et al. 2006; Martìnez-Medina et al. 2017; Song et al. 2017). nematode infections in both roots and shoots of watermelon
In Arabidopsis thaliana, ROS production is mediated by (Yang et al. 2018) suggesting a pivotal role of WRKY70
RbohD and RbohF genes after nematodes infections lead- TFs and ROS signaling in red lights-induced plant defense
ing to galls formation and cell death (Siddique et al. 2014). against Root Knot nematodes in watermelon plants. Accu-
Moreover, SA, JA, ET, BRs, and ABA are involved in plant mulation of WRKY70 in roots only could suggest that this
response to nematodes (Nahar et al. 2011; Kammerhofer TF acts as a point of convergence between JA-mediated and
et al. 2015; Kyndt et al. 2017; Song et al. 2017). Notably, JA SA-mediated signals in plant defense (Li et al. 2004; Yang
is implicated in systemic defense induction of rice (Nahar et al. 2018).
et al. 2011), watermelon (Yang et al. 2018) and tomato (Sun
et al. 2010) against nematodes. Recently, it has been demon-
strated that implication of JA in plant defense against Root Interaction between JA and other plant
Knot nematode is related to the alterations of antioxidative growth regulators
protection and photosynthetic processes (Bali et al. 2017).
Moreover, application of exogenous SA alleviates plant Plant genomes encodes for plant growth regulators which
infections by nematodes (Molinari et al. 2014; Martìnez- are implicated in plant growth and development and are also
Medina et al. 2017). In another hand, biotrophic and necro- implicated in plant response to biotic and abiotic stresses.
trophic pathogens infections are positively regulated by red Thousands of works have described biosynthesis of those
13
endogenous organic compounds (Liu et al. 2017; Chan- orchestrate to upregulate almost 859 genes in Arabidopsis
dler and Werr 2015; Kazan 2015; Li et al. 2019). They also as revealed by transcriptomic studies (Ding et al. 2013). In
underline the way to modulate plant response at molecular this section, we describe the crosstalk between JA and other
and physiological levels. Interactions between phytohor- phytohomones GA, CKs, SA, ABA, BR, ET, and auxin to
mones are crucial for prompt and efficient plant responses modulate plant growth and development.
to biotic and abiotic stress factors. The endogenous concen-
trations of those natural compounds are low, and it changes The JA–GA interaction
depending of the applied stress. Those observations suggest
that plant hormones interact with each other to modulate GAs are crucial hormones controlling many aspects of plant
plant development as revealed for IAA/CKs interplay (Liu growth and development, such as leaf expansion, stem elon-
et al. 2017; Chandler and Werr 2015). Some works demon- gation, root development, seed germination, and stamen and
strated that JA interacts with salicylic acid (Liu et al. 2016a), flower development (Um et al. 2018). In another hand, sev-
ethylene (Kazan 2015; Li et al. 2019), GA (Wasternack eral studies suggest that GA is crucial for growth inhibi-
and Hause 2007), IAA (Chen et al. 2011), BR (Choudhary tion in stress conditions (Liu and Hou 2018). In unfavorable
et al. 2012) and ABA (Aleman et al. 2016) to unsure plant stress conditions, plants inhibit their growth by activating
response to biotic and abiotic stress. Moreover, it has been their defense systems and inhibiting developmental pro-
demonstrated that stress imposition involves usually the acti- grams. Many works suggest an antagonistic role between
vation/inhibition of a high number of genes as a response GA and JA to coordinate plant development under normal
to the biosynthesis of phytohormones. For example, in and stress conditions (Jang et al. 2017). This is ensured
response to drought stress ABA, IAA, JA, GA, SA, and ET by the direct interaction between DELLA (GA signaling
Fig. 4 Schematic representation of antagonistic action between PIF TFs to interact with responsive gene inducing GA responses. b
JA and GA. a In Free JA conditions, the GA concentrations in cell In GA-free situation, JA concentration increases. Ile-JA/JAZ1/COI1
increase to unsure growth programs. The direct interaction between complex is released by the ubiquitation and degradation of JAZs by
JAZs and DELLA proteins is abolished by degradation of DELLAs, proteosome 26. Thus, the MYC2 TF is activated and enhanced tran-
and JAZ proteins inhibit the activity of MYC2 TF. This activates scription of JA-induced genes
13
repressor) and JAZ (JA signaling repressor) proteins (Fig. 4; The JA–auxin interaction
Yang et al. 2012; Hou et al. 2010). It has recently shown
that rice OsJAZ9 acts as the key JAZ protein in mediating Auxin is a crucial phytohormone implicated in plant growth
crosstalk between JA and GA (Um et al. 2018). It interacts and development (Figueiredo and Köhler 2018). In plants,
with a DELLA protein called SLENDER RICE 1 (SLR1). the predominant form is called IAA (Indole-3-acetic-acid).
Overexpression of OsJAZ9 enhanced rice GA response. IAA biosynthesis could take place through two different
Thus, we can speculate that JA acts as an essential hormone pathways: tryptophan-dependent and -independent path-
that modulates plant growth under stress conditions due to ways (Wang et al. 2019). The JA–IAA interaction controls
its antagonistic interaction with GA. many aspects of plant physiology and development such as
production of secondary metabolites, tendril coiling and
The JA–CKs interaction cell elongation (Saniewski et al. 2002). Unfortunately, those
interactions have never been demonstrated at the molecular
Cytokinin (CKs) control a plethora of aspects related to plant level. The most studied aspect of JA–IAA interaction is the
growth and development by maintaining stem cell integ- regulation of plant roots development. In fact, the apical
rity as well as cell proliferation. In higher plants, zeatin is roots growth was inhibited by JA. This was demonstrated by
the most abundant form of CKs and occurs as two isomers, wild-type plants treated by JA. They form much shorter roots
trans-zeatin (tZ), the active form and cis-zeatin (cZ); less comparing with non-treated plants. In contrast, JA defec-
active form. Trans-zeatin is synthetized under the control of tive mutant plants presents roots with similar length com-
several hormones such as Isopentenyl transferases (IPTs), paring with wild type mainly in presence of JA treatment
and cytochrome P450 CYP735A1 and CYP735A2 (Kieber (Jang et al. 2017). Meanwhile, auxin is crucial hormone
and Schaller 2018). JA and JA-dependent stress response implicated in the control of root growth. Auxin deficient
genes affect the expression of CKs gene response (Argueso mutants or signaling mutants develop shorter roots in com-
et al. 2009). CKs deficient plant mutants are tolerant to stress parison with wild-type plants as demonstrated for (trp2-12)
and respond to stress application in a manner like trans- and auxin resistant 3 (arx3-1) (Zhang et al. 2019). Those
genic plants having high JA concentrations (Qi et al. 2014). findings suggest that JA–IAA interaction could mediate
In higher plants, CKs receptors, called histidine phospho- root growth inhibition as revealed by (Chen et al. 2011).
transferase-proteins transcription factors, regulates CKs The group showed that JA inhibits root growth by reducing
responses in plants. Many works suggest that JA and CKs root meristem activity as exogenous JA application inhibits
act antagonistically controlling various plant development PLETHORAs (PLTs) expression, auxin-responsive tran-
aspects. In fact, in soybean, the CKs induced callus Growth scription factors controlling cell proliferation (Mahonem
is inhibited by JA (Ueda and Kato 1982). Furthermore, JA et al. 2014). Interestingly, PLTs expression level was not
has an antagonistic effect on CKs as JA inhibits the expres- modified in JA signaling mutants, such as coi1-1 and myc2.
sion of genes implicated in chlorophyll development (Liu Authors suggest then that COI1 could be implicated in con-
et al. 2016b). In another hand, it has been demonstrated that trolling root phenotype, whereas MYC2 directly binds to the
JA positively controls xylem differentiation in Arabidopsis PTLS promoters and inhibits their expression. This suggests
roots and eliminates the specific procambium response to that auxin and JA antagonistically interact in the regulation
CKs application (Jang et al. 2017). In contrast, CKs acts as of apical root growth (Fig. 5).
a negative regulator of xylem differentiation in Arabidopsis The development of floral organs is another physiologi-
roots. Taken together, those results approve the idea that cal process controlled by JA–IAA interaction (Reeves et al.
CKs interacts antagonistically with JA in xylem develop- 2012). An interaction between IAA and JA modulates petals
ment in Arabidopsis roots. Interestingly, myc2 mutant did and stamens development under the control of R2R3 MYB
not show extra xylem formation in response to exogenous transcription factors especially MYB21 and MYB24 TFs
JA. Moreover, the expression of CKs signaling inhibitor, which are considered as key regulators of petal and stamen
AHP6, was inhibited in this mutant. Those results suggest growth (Reeves et al. 2012). Moreover, ARF6 and ARF8 TFs
that MYC2 transcription factor is involved in this process control the regulation of expression of MYB21 and MYB24
by promoting AHP6 expression. Nevertheless, it has been JA-responsive TFs (Fig. 5). Nevertheless, the regulation of
demonstrated that JA and CKs antagonistically interact in leaves senescence is also controlled by IAA–JA interaction.
the plant response to circadian stress (Nitschke et al. 2016). In fact, JA acts as a positive regulator of leaf senescence as
In fact, plants with reduced or defective CKs levels present a JAZ7 proteins suppress dark-induced leaf senescence and
JA-dependent cell death phenotype in response to circadian MYCs TFs, such as MYC2, promote senescence by activat-
modification, in contrast to wild types (Nitschke et al. 2016). ing the senescence-associated genes expression as well as
chlorophyll degradating-related genes, demonstrating that
JA stimulates leaf senescence pathway in a COI1-dependent
13
Fig. 5 Role of JA/Auxin crosstalk in plant growth and development. JA and IAA signaling pathways positively regulate flowering via MYB21/
MYB24 Tfs but negatively regulate root growth through JAZs-MYC2
manner (Yu et al. 2015). In this pathway, JAZ4, JAZ8 and DWARF4 restored JA sensitivity in the coi1 mutant back-
WRKY57 act as negative regulators contrarily to JAZ7 and ground and showed JA hypersensitivity in the wild-type
the auxin signaling repressor IAA29 functions as a positive background. Furthermore, expression of DWARF4 was
regulator. Moreover, WRKY57 negatively regulates senes- down regulated by JA in a COI1-dependent manner, and
cence-associated gene expression by interacting with JAZ4/8 exogenous BR treatment attenuated the effects of JA on root
and IAA29. Authors speculate that competition between the growth inhibition (Ren et al. 2009). These results indicate
WRKY57–JAZ4/8 WRKY57-IAA29 implies the adequate that a BR–JA interaction is involved in the modulation of
response to the physiological stimulus. Thus, an antagonistic JA signaling. A low BR concentration activates BR signal-
JA–IAA interaction is involved in leaf senescence (Fig. 5; ing pathways such as BR receptor BRI1, BR-related kinase
Jiang et al. 2014). BAK1, and BR-related TFs. This induces the activation of
several downstream genes such as BES1 and BZR1, which
JA–Brassinosteroids crosstalk activate plant response to many abiotic stresses (Yang et al.
2019).
Brassinosteroids (BR) are implicated in different aspects of
plant growth and development. They are also involved in The JA–SA crosstalk
the modulation of JA signaling and JA-dependent growth
inhibition. After stress perception, JA rapidly regulates plant gene
In contrast to JA, BR promotes above-ground plant defense (Genva et al. 2019). The A. thaliana coronatine
growth. Thus, JA-BR crosstalk is insured by the bal- insensitive 1 (coi1) mutants presented a blocked JA response
ance between plant growth and defense resistance (Yang (Feys et al. 1994). COI1 is a JA receptor that belongs to
et al. 2019). For example, low BR concentrations induce the F-box protein family. It acts as an E3-ubiquitin ligase-
OsDI1 and OsDWARF expression at early and late stages of mediated proteolysis of target proteins (Hiruma et al. 2011)
BR biosynthesis, which activate the defense pathway (Yang such as JASMONATE ZIM-DOMAIN (JAZ) proteins.
et al. 2019). Moreover, DWARF4 encodes a key enzyme JA is essentially involved in plant response against necro-
responsible for BR biosynthesis and a leaky mutation of trophic pathogens. SA mediates plant resistance against
13
biotrophic and hemibiotrophic pathogens (Fu et al. 2012). Zander et al. 2010). A schematic presentation of JA–SA
JA signaling modulates many NAC Tfs activities (such as crosstalk component is presented in Fig. 6.
ANAC019/055/072) to block SA accumulation through In addition to its crucial role in ensuring plant defense
modulation of multiple NAC TFs. Briefly, physical interac- against pathogens, other important roles of SA in regulat-
tion between MYC2 and several NACs promoter activates ing many physiological processes and responding to adverse
their transcription. Expressed NAC TFs inhibits the expres- environmental situations have emerged. SA is implicated in
sion of SA biosynthesis gene called ISOCHORISMATE plant response to salinity (Khan et al. 2012), light (Tuteja
SYNTHASE 1 (ICS1) but stimulate the expression of a and Gill 2013) and cold (Gornik et al. 2014). In fact, plant
SA methylation gene, BENZOIC ACID/SA CARBOXYL treatment with JA or SA improves plant resistance to chilling
METHYLTRANSFERASE 1 (BSMT1) (Zheng et al. 2012). (Gornik et al. 2014). Moreover, MeJA and MeSA present
In addition, many other components are involved in SA-JA a protective effect against chilling injury in pomegranates
crosstalk such as MYC2, TGAs, and PDF 1.2 (Gatz 2013); fruits (Sayyari et al. 2010). SA and JA are also implicated
mitogen-activated protein kinase (MAPK) (Suarez-Rodri- in plant tolerance to drought (Ilyas et al. 2017). In another
guez et al. 2010), redox regulators glutathione (GRX) and hand, several studies concentrated on the role of SA and
thioredoxin (TRX) (Sopel and Gary 2011) and WRKY70 JA in plant tolerance to salinity. In fact, some researches
(Shim et al. 2013). GRXs genes can block TGA-mediated demonstrated that JA and SA protect plants from negative
JA response gene expression, such as ORA59 confirming effects of salinity by the induction of gene encoding effective
SA–JA antagonism (Zander et al. 2010). MPK4 is a posi- proteins expression (Khan et al. 2012). Moreover, it has been
tive regulator of GRX480 (SA signaling pathway) and acts found that exogenous SA and JA application decreased Na+
as a negative regulator of MYC2 (JA signaling pathway) concentrations in soybean cells under different salt stress
(Wasternack and Hause 2007). Meanwhile, TRX enzyme levels and that JA has a greater effect on Na+ reduction than
transforms NPR1 polymers to monomers in presence of SA. SA (Farhangi-Abriz and Ghassemi-Golezani 2018). There-
Those monomers such as GRX480 are transported to the fore, a regulatory protein called glutaredoxin GRX480 is
nucleus and specifically bind to TGAs, which also directly a key protein in JA and SA signaling. This protein medi-
regulate the expression of PR1 (Fu et al. 2012; Gatz 2013; ates redox regulation of proteins by catalyzing disulfide
Fig. 6 JA-SA crosstalk in plants

in response to pathogen infec-
tion. Plant infection by a patho-
gen leads to the increase in the
endogenous SA concentration
and the activation of NONEX-
PRESSOR OF PR GENES1
(NPR1), which induces
WRKY70 transcriptional activ-
ity. In turn, WRKY70 promotes
PR1 gene expression and thus
enhancing pathogen response
13
transitions (Meldau et al. 2012). In Nicotiana attenuata PDF1.2 expression by activating ERF1 and ORA59 expres-
plants, researches demonstrated that mitogen-activated pro- sion. Thus, JA and ET control plant response to nectrophic
tein kinase 4 (MPK4) have a dual role in plant signaling. In pathogens through JAZs-MYC2 and EIN3/EIL1 (Pieterse
fact, in response to light stress, this kinase acts as a negative 2012). Moreover, JA enhances the degradation of JAZ, so
regulator during the SA signaling pathway, but it plays a MYC2 is activated to regulate ORA59/ERF1 expression and
positive role during the JA signaling pathway (Tuteja and VSP2, a wound responsive gene to respond to herbivorous
Gill 2013). To coordinate regulation between SA and JA, insect attacks (Yang et al. 2019). Besides, JAZ inhibits EIL2/
cells involve several factors such as GRX480 or MAPK4. EIN3 transcriptional activity downstream ORA59/ERF1 that
Moreover, exogenous application of SA and/or JA can targets the promoter of PLANT DEFENSIN 1.2 (PDF1.2)
enhance abiotic stress responses (Tuteja and Gill 2013). and induces its expression thereby resisting the infection of
necrotrophic pathogens and hemibiotrophic pathogens (Piet-
JA–ABA crosstalk erse 2012). In Arabidopsis, JA inhibits apical hook forma-
tion via activation of MYC2:3:4 TFs. ET stabilizes EIN3/
Upon external stimuli application on plants, an immense EIL1 complex leading to the inhibition of the transcriptional
signaling network of events occurs in cells. C a2+ sensors function of MYCs, and it attenuates MYC-mediated plant
act in the first line of defense to decode and then transducer defenses against insect attack (Song et al. 2014).
signals through the suitable stress hormone pathways to acti- In the lights of those findings, we can conclude that hor-
vate the appropriate signaling cascade. ABA is a well-stud- monal interactions play a critical role for insuring plant
ied phytohormone implicated in plant response to abiotic physiology and growth (Yang et al. 2019). The cross talk
stress. Many studies described that ABA and JA presents an of JA with other hormones (IAA, GA, CKs, BR, JA ABA
antagonistic role in plant growth and development (Chen and SA) ensures plant growth and defense. Direct JA–GA,
et al. 2011). In another hand, those hormones act synergisti- JA–IAA, JA–BR, and JA–CKs interactions play important
cally in response to environmental stress and MYC2 unsure role in modulating physiological role such as root and floral
that crosstalk (Chen et al. 2011). In A. thaliana as well as organs development and can help plant to optimize their
tobacco plants, PYRABACTIN RESISTANCE1-Like pro- growth and development in stressful conditions. However,
teins (PYLs) known as ABA receptor proteins control meta- this interaction is almost antagonistic but in some cases it is
bolic reprogramming via JA signaling pathway. Thus, JA/ synergic. In fact, despite JA–GA interaction antagonistically
ABA crosstalk can control plant metabolism and growth (Per control stem elongation, it plays a positive role to control
et al. 2018). When endogenous ABA concentration increases stamen development (Qi et al. 2014). In the same way of
in response to external (salt stress, drought…) or internal idea, JA–IAA interaction negatively regulates apical roots
(growth and development) conditions, ABA interacts with growth but positively regulate lateral root growth (Cai et al.
many members of the PYL/RCAR ABA receptor family to 2014).
inhibit 2C protein phosphatases (PP2C proteins) thus to
initiate signal transduction. Recently, a direct interaction
between the ABA receptor PYL6 RCAR9 and MYC2 TF Conclusion and prospects
has been demonstrated (Aleman et al. 2016). This interaction
is enhanced in presence of ABA. Moreover, ABA receptor Jasmonic acid and its derivatives have pivotal roles in the
PYL6 forms a complex with JAZ6 and JAZ8 to stimulate the establishing plant defense and resistance in response to
transcriptional activity of MYC2. In fact, MYC2 stimulate environmental stimuli. The mechanisms of action of JA are
VSP2 expression, inhibits PTL1 and PTL2 expression and specific when plants are subjected to different environmental
root development. constraints due to the diversification of plant hormone inter-
play. Several genes (e.g. JAZ, AOS1, AOC, LOX2, and COI1)
JA–ET crosstalk in plants and TFs (MYC2 and bHLH) are involved in the core JA
signaling pathway as activators or repressors, thereby modu-
Many researchers found that JA and ET could antagonize lating the responses to stimuli received by the plants from
or coordinate to regulate plant stress response (Zhu and the surrounding environment. The JAZ-MYC module has a
Lee 2015). In fact, MYC2 (JA signaling) and (EIN3-like key role in the JA signaling pathway through the assimilation
1 (EIL1) and ET INSENSITIVE3 (EIN3) involved in ET of regulatory TFs and associated genes. Plant hormone sign-
signaling pathway are implicated in JA/ET crosstalk (Zhang aling systems are complicated and unstable, especially under
et al. 2014). Recent findings showed that when plants are complex environmental conditions. Over the years, JA cross-
exposed to a necrotrophic or hemibiotrophic pathogen talk with other phytohormones has been elucidated. How-
attacks, JA signaling pathway act in synergy with the ET ever, information in plants is still limited. Crosstalk nodes
signaling pathway to activate defense proteins such as between those hormones have to be investigated. In presence
13
of abiotic constraints, JA orchestrate plant stress responses of the PYL6 ABA receptor with MYC2 transcription factor: a
by stimulating the plant’s defense apparatus, which mainly putative link of ABA and JA signaling. Sci Rep 6:28941. https://
doi.org/10.1038/srep28941
involves phytohormones and other defensive compounds Ali E, Hussain N, Shamsi IH, Jabeen Z, Siddiqui MH, Lx J (2018) Role
such as antioxidative enzymes. JA functions synergistically of jasmonic acid in improving tolerance of rapeseed (Brassica
and antagonistically with ABA, ET, SA, and other plant napus L.), to Cd toxicity. J Zhejiang Univ Sci B 19:130–146.
hormones to accomodate environmental constraints. The https://doi.org/10.1631/jzus.B1700191
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to background and characteristics of major molecular biological
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the molecular control of JA interaction with its transporter Choose your partners, dimerization in eukaryotic transcription
to control it’s moving through plants. Furthermore, cross- factors. Trends Biochem Sci 33:220–229. https://doi.org/10.
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talk between JA and other phytohormones is not linear and Argueso CT, Ferreira FJ, Kieber JJ (2009) Environmental percep-
researches should focus on the spatio-temporal relationships tion avenues: The interaction of cytokinin and environmental
between hormones using new chemicals that act as targets response pathways. Plant Cell Environ 32:1147–1160. https://
of JA in vivo or using high-throughput nucleotide sequenc- doi.org/10.1111/j.1365-3040.2009.01940.x
Avramova Z (2017) The jasmonic acid signaling and abscissic acid
ing techniques as well CRISPR-CAS technology to generate signaling pathways during one, but not repeated dehydratation
specific mutant that allows underlying mechanisms of plant stress: a nonspecific ‘paniky’ or a meaningful response? Plant
tolerance to unfavorable environments. Thus, understanding Cell Env 40:1704–1710. https://doi.org/10.1111/pce.12967
the signal transduction mechanism of JA under environmen- Awan SA, Khan I, Rizwan M, Zhang X, Brestic M, Khan A, El-Sheikh
MA, Alyemeni MN, Ali S, Huang L, Huang L (2020) Exogenous
tal stress is essential to promote plant resistance, with poten- abscisic acid and jasmonic acid restrain polyethylene glycol-
tial improvements in plant survival and crop yield. induced drought by improving the growth and antioxidative
enzyme activities in pearl millet. Physiol Plant. https://doi.org/
10.1111/ppl.13247
Author contributions MG and FB wrote the manuscript. AS and ML Ayala A, Muñoz MF, Argüelles S (2014) Lipid peroxidation: produc-
read and revised the manuscript. All the authors approved the final tion, metabolism, and signaling mechanisms of malondialdehyde
version of the manuscript. and 4-hydroxy-2-nonenal. Oxid Med Cell Longev. https://doi.
org/10.1155/2014/360438
Azeem U (2018) Ameliorating Nickel stress by Jasmonic acid treat-
Funding This review received no external funding. ment in Zea mays L. Russ Agric Sci 44:209–215
Bali S, Kaur P, Sharma A, Ohri P, Bhardwaj R, Alyemeni MN et al
Declarations (2017) Jasmonic acid-induced tolerance to root-knot nematodes
in tomato plants through altered photosynthetic and antioxidative
Conflict of interest The authors declare no conflicts of interest. defense mechanisms. Protoplasma 255:471–484. https://doi.org/
10.1007/s00709-017-1160-6
Ethical approval This review does not contain any studies with human Bali S, Kaur P, Kohli SK, Ohri P, Thukral AK, Bhardwaj R, Wijaya L,
participants or animals performed by any of the authors. Alyemeni MN, Ahmad P (2018) Jasmonic acid induced changes
in physio-biochemical attributes and ascorbate-glutathione path-
way in Lycopersicon esculentum under lead stress at different
growth stages. Sci Total Environ 645:1344–1360. https://d oi.o rg/
10.1016/j.scitotenv.2018.07.164
Bali S, Jamwal VL, Kaur P, Kohli SK, Ohri P, Gandhi SG, Bardwaj R,
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Ghorbel et al. 2023 Role of jasmonic acid in plants_ the molecular point of view

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Plant Cell Reports (2021) 40:1471–1494

Role of jasmonic acid in plants: the molecular point of view

Keywords Environmental stress · Jasmonates · MYC2 · Pathogen stress · Plant defense

Introduction messengers such as C a2+, reactive oxygen species (ROS),

Fig. 2 Catabolism of jasmonic

Table 1 The most identified MYC2 interacting proteins in Arabidopsis thaliana

SCFCOI1/JAZ/ AtMYC2 complex: the JA Core

Inhibition of root growth Regulation of stomatal closure

Fig. 6 JA-SA crosstalk in plants

Cell Environ 32:837–850. https://doi.org/10.1111/j.1365-3040. in jasmonate signaling. Nature 448:666–671. https://doi.org/10.

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Ghorbel et al. 2023 Role of jasmonic acid in plants_ the molecular point of view

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Ghorbel et al. 2023 Role of jasmonic acid in plants_ the molecular point of view

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Ghorbel et al. 2023 Role of jasmonic acid in plants_ the molecular point of view

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Plant Cell Reports (2021) 40:1471–1494

Role of jasmonic acid in plants: the molecular point of view

Keywords Environmental stress · Jasmonates · MYC2 · Pathogen stress · Plant defense

Introduction messengers such as C ­ a2+, reactive oxygen species (ROS),

Fig. 2 Catabolism of jasmonic

Table 1 The most identified MYC2 interacting proteins in Arabidopsis thaliana

SCFCOI1/JAZ/ AtMYC2 complex: the JA Core

Inhibition of root growth Regulation of stomatal closure

Fig. 6 JA-SA crosstalk in plants

Cell Environ 32:837–850. https://​doi.​org/​10.​1111/j.​1365-​3040.​ in jasmonate signaling. Nature 448:666–671. https://​doi.​org/​10.​

You might also like

Introduction messengers such as C a2+, reactive oxygen species (ROS),

Cell Environ 32:837–850. https://doi.org/10.1111/j.1365-3040. in jasmonate signaling. Nature 448:666–671. https://doi.org/10.