Regulation of Stomatal Immunity by Interdependent …...2017/03/02 · 1 1 RESEARCH ARTICLE 2...

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RESEARCH ARTICLE 1

Regulation of Stomatal Immunity by Interdependent Functions of a 2

Pathogen-responsive MPK3/MPK6 Cascade and Abscisic Acid 3

4 Jianbin Su1,2, Mengmeng Zhang1, Lawrence Zhang3,5, Tiefeng Sun1, Yidong Liu2, Wolfgang 5 Lukowitz4, Juan Xu1,#, and Shuqun Zhang2,1,#6

7 1 Key Laboratory of Plant Physiology and Biochemistry, College of Life Sciences, Zhejiang 8 University, Hangzhou, Zhejiang 310058, China 9 2 Division of Biochemistry, Interdisciplinary Plant Group, and Bond Life Sciences Center, 10 University of Missouri, Columbia, MO 65211, USA 11 3 David H. Hickman High School, Columbia, MO 65203, USA 12 4 Department of Plant Biology, University of Georgia, Athens, GA 30602, USA 13 5 Current address: School of Medicine, University of Missouri-Kansas City, Kansas City, MO 14 64108 15 # Corresponding authors: Juan Xu, [email protected] and Shuqun Zhang, 16

[email protected] 17 18

Short title: MPK3 and MPK6 regulate stomatal immunity 19 One-sentence summary: MPK3/MPK6 cascade and ABA function interdependently to regulate 20 stomatal immunity by modulating the organic acid metabolism and ion channels, respectively. 21

22 The author responsible for distribution of materials integral to the findings presented in this 23 article in accordance with the policy described in the Instructions for Authors (www.plantcell.org) 24 is: Juan Xu, [email protected]. 25

26 ABSTRACT 27 Activation of mitogen-activated protein kinases (MAPKs) is one of the earliest responses after 28 plants sense an invading pathogen. Here we show that MPK3 and MPK6, two Arabidopsis 29 thaliana pathogen-responsive MAPKs, and their upstream MAPK Kinases, MKK4 and MKK5, 30 are essential to both stomatal and apoplastic immunity. Loss of function of MPK3 and MPK6, or 31 their upstream MKK4 and MKK5, abolishes pathogen/microbe-associated molecular pattern- and 32 pathogen-induced stomatal closure. Gain-of-function activation of MPK3/MPK6 induces 33 stomatal closure independently of abscisic acid (ABA) biosynthesis and signaling. In contrast, 34 exogenously applied organic acids such as malate or citrate are able to reverse the stomatal 35 closure induced by MPK3/MPK6 activation. Gene expression analysis and in situ enzyme 36 activity staining revealed that malate metabolism increases in guard cells after activation of 37 MPK3/MPK6 or inoculation of pathogen. In addition, pathogen-induced malate metabolism 38

Plant Cell Advance Publication. Published on March 2, 2017, doi:10.1105/tpc.16.00577

©2017 American Society of Plant Biologists. All Rights Reserved

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requires functional MKK4/MKK5 and MPK3/MPK6. We propose that the pathogen-responsive 39 MPK3/MPK6 cascade and ABA are two essential signaling pathways that control, respectively, 40 the organic acid metabolism and ion channels, two main branches of osmotic regulation in guard 41 cells that function interdependently to control stomatal opening/closure. 42 43 INTRODUCTION 44

For successful pathogenesis, the pathogen must gain entry into the interior of the plant to acquire 45

nutrients. Some pathogenic fungi secrete digestive enzymes or use mechanical force to overcome 46

the impermeable cuticle layer that covers most parts of the plants, yet most other pathogens can 47

only gain entry through wounds or natural openings such as hydathodes or stomatal pores 48

(Grimmer et al., 2012). As a countermeasure, plants can rapidly close their stomata upon 49

perception of pathogens to restrict pathogen entry, a response known as stomatal immunity 50

(Melotto et al., 2006). Once pathogens overcome the surface barrier and enter the interior space 51

of leaves, the perception of pathogen invasion by host cells induces rapid strengthening of their 52

cell walls and secretion of a set of antimicrobial molecules and/or compounds to the apoplastic 53

region to suppress pathogen growth, a process known as apoplastic immunity (Monaghan and 54

Zipfel, 2012; Doehlemann and Hemetsberger, 2013). Plants have surface pattern-recognition 55

receptors (PPRs) to perceive pathogen/microbe-associated molecular patterns (PAMPs), which 56

initiates an array of defense signaling events including activation of mitogen activated protein 57

kinases (MAPKs), Ca2+ influx, reactive oxygen species (ROS) burst, NO production, and 58

activation/inhibition of ion channels (Ausubel, 2005; Chisholm et al., 2006; Jones and Dangl, 59

2006; Boller and Felix, 2009; Bohm et al., 2014). These signaling events function independently 60

or interdependently to form an intricate signaling network to ensure robust stomatal and 61

apoplastic immunity (Melotto et al., 2008; McLachlan et al., 2014; Arnaud and Hwang, 2015). 62

Abscisic acid (ABA) plays an important role in controlling plant water balance by 63

regulating stomatal opening and closure (Hetherington and Woodward, 2003; Kim et al., 2010; 64

Ruszala et al., 2011; Chater et al., 2014). It is also essential to stomatal defense during plant 65

immunity (Melotto et al., 2006; Zeng and He, 2010; Du et al., 2014; Lim et al., 2014). A recent 66

study concluded that ABA and PAMP (flg22) signals converged at SLAC1 and ABA signaling 67

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plays only a minor role in stomatal defense (Montillet et al., 2013). By using a noninvasive 68

nanoinfusion technique, another study demonstrated that ABA- and flg22- induced stomatal 69

closure converge at the level of OST1 and then activate downstream SLAC1 and SLAH3 ion 70

channels (Guzel Deger et al., 2015). ROS bursts play important roles in ABA- and 71

PAMP-induced stomatal closure. However, the mechanism of the PAMP-induced ROS burst is 72

distinct from the ABA-induced ROS burst. Upon perception of PAMPs, the plasma-associated 73

kinase BIK1 directly phosphorylates and so activates the NADPH oxidase RbohD to induce ROS 74

production (Kadota et al., 2014; Li et al., 2014). In contrast, ABA-induced ROS production is 75

dependent on the kinase OST1 (Mustilli et al., 2002), which physically interacts with and 76

phosphorylates RbohD/RbohF (Sirichandra et al., 2009; Acharya et al., 2013). MPK9 and 77

MPK12, two guard cell-specific MAPKs that function downstream of ROS in ABA-mediated 78

stomatal movement (Jammes et al., 2009), were also essential to plant resistance against 79

spray-inoculated Pseudomonas syringae pv tomato DC3000 (Pst) (Jammes et al., 2011). 80

Interestingly, activation of EDS1/PAD4-dependent plant immune responses inhibits 81

ABA-induced stomatal closure at the level of Ca2+ signaling (Kim et al., 2011), suggesting an 82

intricate crosstalk between ABA signaling and plant immune networks. Besides ABA, salicylic 83

acid (SA) biosynthesis and signaling are also required for pathogen/PAMP-induced stomatal 84

closure (Melotto et al., 2006; Zeng and He, 2010; Zheng et al., 2012; Zheng et al., 2015). 85

Plant pathogens also have several sophisticated strategies to interfere with plant stomatal 86

immunity. Coronatine (COR), a phytotoxin produced by some strains of Pseudomonas syringae 87

(Cuppels and Ainsworth, 1995; Brooks et al., 2004), promotes stomatal reopening and inhibits 88

ABA-induced stomatal closure (Melotto et al., 2006). COR hijacks the plant jasmonic acid 89

receptor COI1 to activate several closely related NAC transcription factors, which subsequently 90

prevents the accumulation of SA by inhibiting SA biosynthesis and promoting SA metabolism 91

(Zheng et al., 2012; Du et al., 2014). In addition to COR, pathogen-secreted cytokinin (e.g. 92

kinetin), fusicoccin, type III effectors, and oxalate also function differentially in counteracting 93

plant stomatal immunity (Grimmer et al., 2012). Cytokinin exerts its inhibitory effect on 94

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ABA-induced stomatal closure through enhancing ethylene production (Tanaka et al., 2006), 95

while fusicoccin directly stimulates plasma membrane-localized H+-ATPases (Johansson et al., 96

1993). HopF2 and HopM1, two type III effectors secreted by Pst, were shown to inhibit stomatal 97

immunity through inhibiting the PAMP-induced ROS burst (Hurley et al., 2014; Lozano-Duran 98

et al., 2014). 99

Most interestingly, oxalate, an intermediate of the tricarboxylic acid (TCA) cycle, was 100

shown to function as a virulence factor to counteract stomatal immunity by promoting 101

accumulation of osmotically active molecules and inhibiting ABA-induced stomatal closure 102

(Guimaraes and Stotz, 2004). The finding of oxalate as a virulence factor indicates plant 103

pathogens may counteract stomatal immunity by targeting guard cell organic acid metabolism to 104

enhance pathogenicity. Malate, together with Cl- and NO3-, are the major anions to balance the 105

positively charged K+ during stomatal opening (Kim et al., 2010; Araujo et al., 2011a). As a 106

result, coordinated down-regulation of guard cell organic and inorganic anions through metabolic 107

conversion and transporter-mediated efflux could be an effective strategy to close stomata in 108

response to various stimuli. 109

For a long time, the role of organic solutes was considered to be secondary in both 110

stomatal closure and opening (Blatt, 2016). However, this viewpoint was challenged by three 111

recent reports. One study found that guard cell triacylglycerol breakdown was essential to 112

light-induced stomatal opening (McLachlan et al., 2016). It was proposed that triacylglycerol 113

breakdown provided energy for plasma membrane localized H+-ATPase. By using quantitative 114

analysis of starch turnover and molecular genetic analyses, rapid starch degradation in guard 115

cells was shown to play an important role in stomatal opening and plant fitness (Horrer et al., 116

2016). At the same time, the role of guard cell starch biosynthesis in stomatal closure was also 117

reported (Azoulay-Shemer et al., 2016). Collectively, these recent studies highlight the essential 118

role of metabolism in stomatal movement. These new findings also raise the question of how 119

these metabolic processes are regulated during stomatal movement in response to various 120

stimuli. 121

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Arabidopsis thaliana MPK3, MPK4, and MPK6 are rapidly activated upon plant 122

perception of PAMPs. They play critical roles in multiple plant defense responses (Meng and 123

Zhang, 2013). It was shown that elevated MPK4 activity compromises Arabidopsis resistance to 124

spray-inoculated Pst, but not Pst- or flg22-induced stomatal closure (Berriri et al., 2012). The 125

role of MPK3 and MPK6 in regulating defense-related secondary metabolite biosynthesis has 126

been intensively studied (Ren et al., 2008; Mao et al., 2011; Xu et al., 2016). Here, we show that 127

MPK3 and MPK6, as well as their upstream kinases, MKK4 and MKK5, are essential to 128

Arabidopsis immunity. They are involved in stomatal immunity by regulating guard cell primary 129

organic acid metabolism. Loss of function of both MPK3 and MPK6 resulted in compromised 130

stomatal immunity and high susceptibility phenotype of the double mutant. In contrast, Pst- and 131

flg22-induced stomatal closure was not affected in mpk3 or mpk6 single mutant, which is 132

consistent with their normal resistance against Pst. MPK3/MPK6 activation-induced stomatal 133

closure is independent of ABA biosynthesis and signaling. It is also insensitive to COR, a 134

phytotoxin capable of inhibiting ABA-induced stomatal closure. Our data suggest that MPK3 135

and MPK6 regulate stomatal immunity by controlling the metabolism of organic acids such as 136

malate and citrate, well-known osmolytes in stomatal opening and closure. On the basis of these 137

findings, we propose an interdependent model for MAPK and ABA signaling pathways in 138

stomatal immunity. 139

140 RESULTS 141

Arabidopsis MPK3 and MPK6 have redundant functions in plant immunity 142

MPK3 and MPK6, as well as their orthologs in other plant species, were believed to play 143

positive roles in plant immunity based on their rapid activation by PAMPs and pathogen 144

inoculation (Zhang and Klessig, 2001). They positively regulate various defense responses 145

including phytoalexin induction and defense gene activation (Ren et al., 2008; Mao et al., 2011; 146

Meng and Zhang, 2013; Xu et al., 2016). However, clear genetic evidence to demonstrate a 147

positive role of MPK3 and MPK6 in disease resistance is still lacking. Plants with MPK6, but not 148

MPK3, suppressed by RNAi were reported to be more susceptible to Pst, with about a 1-log 149

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increase in pathogen growth (Menke et al., 2004). Pathogen resistance assays using mpk3 or 150

mpk6 single mutants revealed that mpk3, but not mpk6, plants were more susceptible, with less 151

than 1 log increase in Pst proliferation (Beckers et al., 2009). Our repeated trials revealed no 152

clear susceptible phenotype in mpk3 single mutants, and only a slightly higher bacterial growth 153

in two independent mpk6 mutant alleles (0.01< P<0.05) when spray- or infiltration-inoculated 154

with Pst (Supplemental Figure 1A and C). Also, flg22-induced stomatal closure was normal in 155

mpk3 or two independent mpk6 alleles (Supplemental Figure 1B). These contradictory results are 156

likely a result of functional redundancy between MPK3 and MPK6, and the disease resistance 157

phenotype of the mpk3 or mpk6 single mutant/RNAi line is subtle and subject to influence of 158

experimental conditions. 159

MPK3 and MPK6 play redundant function in many developmental processes (Meng and 160

Zhang, 2013; Xu and Zhang, 2015), including their essential function in embryogenesis. 161

Previously, we generated a partially rescued mpk3 mpk6 double mutant using a steroid-inducible 162

promoter-driven MPK6 (Wang et al., 2007). However, the rescued mpk3 mpk6 double mutant 163

seedlings lack normal pavement cells and cannot survive in soil, and as a result, we could not 164

examine the stomatal behavior and pathogen resistance of such mutant. Recently, using a 165

chemical genetic approach, we generated another conditional mpk3 mpk6 double mutant, named 166

MPK6SR (genotype: mpk3 mpk6 PMPK6:MPK6YG), in which mpk3 mpk6 double mutant was 167

rescued with MPK6YG, a NA-PP1-sensitized version of MPK6. Replacement of Tyr144 with Gly 168

enlarges the ATP binding pockets of MPK6, and makes MPK6YG sensitive to NA-PP1, a PP1 169

analog with a bulkier side chain (Xu et al., 2014). We also generated a similar system using a 170

modified MPK3, MPK3TG, and the rescued plants were named MPK3SR (genotype: mpk3 mpk6 171

PMPK3:MPK3TG). NA-PP1-treated MPK3SR (line #58) and MPK6SR (line #64) plants were more 172

susceptible to spray-inoculated Pst (Figure 1A and B), with close to 3-log increase in pathogen 173

growth. In the absence of NA-PP1 inhibitor, both MPK3SR and MPK6SR plants behaved like 174

wild-type (Col-0) control. In addition, Col-0 plants treated with NA-PP1 did not show increased 175

susceptibility to Pst, demonstrating the specificity of NA-PP1 inhibitor to the 176

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chemical-sensitized MPK3 and MPK6. Similar results were observed in independent MPK3SR 177

and MPK6SR lines (Supplemental Figure 2A). 178

We also tested Pst inoculation by syringe infiltration. As shown in Figure 1C and D, 179

NA-PP1-treated MPK3SR (line #58) and MPK6SR (line #64) plants were more susceptible. 180

Independent MPK3SR and MPK6SR lines also exhibited enhanced susceptibility (Supplemental 181

Figure 3). In comparison to ~3 logs of more pathogen growth when Pst was spray inoculated, 182

infiltration inoculation only resulted in ~1 log of more Pst growth. This result suggests that 183

MPK3 and MPK6 are involved in both stomatal and apoplastic immunity. 184

185

Arabidopsis MPK3 and MPK6 play a major role in stomatal immunity 186

Consistent with our speculation based on two different pathogen inoculation methods, we 187

observed that the clustered stomata are always wide open in the rescued mpk3 mpk6 double 188

mutants (Wang et al., 2007; Xu et al., 2014). These observations prompted us to examine the 189

stomatal behavior in MPK3SR and MPK6SR plants after NA-PP1 treatment. The beauty of the 190

chemical genetically rescued MPK3SR and MPK6SR plants is that we can allow the plants to 191

grow and develop normally in the absence of inhibitor, and then treat the plants with NA-PP1 to 192

specifically block the kinase activity of the modified MPK3 or MPK6, which gives us activity 193

null mutant of MPK3 and MPK6 for functional analyses. We found that stomatal closure in 194

response to flg22 and Pst was impaired in both epidermal peels and intact leaves of MPK3SR and 195

MPK6SR plants treated with NA-PP1 (Figure 1E and F, Supplemental Figure 4), indicating that 196

reduced stomatal immunity contributes to the enhanced susceptibility of NA-PP1-treated 197

MPK3SR and MPK6SR plants, i.e. MPK3 and MPK6 play essential function in plant stomatal 198

immunity. 199

The best way to assess the levels of stomatal immunity is to quantify the number of 200

pathogens that enter into the apoplastic spaces of leaves per stomata in a specific period of time. 201

To achieve this, we developed a pathogen entry assay by utilizing the luxCDABE-tagged Pst 202

strain (Pst-lux) (Fan et al., 2008) together with Photek photon-counting camera system, which 203

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can distinguish more than two orders of magnitude of Pst-lux bacteria (Supplemental Figure 5A). 204

To remove the surface-attached Pst-lux cells, we washed the leaves in 0.02% Silwet L-77 with 205

agitation. To assess the effectiveness, we pressed the washed leaves against Pseudomonas Agar 206

(Difco) plates, and then cultured the plates for 24 hr to generate “leaf prints”. As shown in 207

Supplemental Figure 5B, no Pst-lux was detected after 0.02% Silwet L-77 wash, indicating that 208

the surface Pst-lux cells were removed efficiently, and the Pst-lux we detected in our pathogen 209

entry assay resided inside leaves. We observed that more than 10-fold more Pst-lux bacteria 210

entered into leaf interior of NA-PP1-treated MPK3SR and MPK6SR plants (Figure 2A and B). 211

After being normalized to stomatal density (Figure 2C), the number of Pst-lux entered into the 212

leaves of NA-PP1-treated MPK3SR and MPK6SR plants was >10 times more than those of the 213

control plants (Figure 2D). Compromised stomatal immunity was also observed in independent 214

MPK6SR and MPK3SR lines (Supplemental Figure 2B-E). Based on these data, we conclude that 215

MPK3 and MPK6 have redundant functions in Arabidopsis stomatal immunity. 216

217

MKK4 and MKK5 play redundant roles in both stomatal and apoplastic immunity 218

Our previous study demonstrated that MKK4 and MKK5 are the MAPKKs upstream of 219

MPK3/MPK6 in regulating stomatal development and patterning (Wang et al., 2007). To 220

determine whether these two MAPKKs are also required for stomatal immunity, we generated 221

mkk4 mkk5 double mutants using the recently reported mkk4 and mkk5 single TILLING mutants 222

(Zhao et al., 2014) after the er-105 mutant allele was crossed away. Similar to the 223

NA-PP1-treated MPK3SR and MPK6SR plants, the mkk4 mkk5 double mutant plants were also 224

highly susceptible to spray-inoculated Pst (Figure 3A and B). The mkk4 mkk5 double mutant also 225

exhibited impaired stomatal closure in response to flg22 and Pst (Figure 3C and D) and a higher 226

level of Pst-lux entry (Figure 3E and F). We observed that the stomatal density of mkk4 mkk5 227

double mutant was ~2.5-times higher than that of the wild type (Figure 3G) as a result of 228

clustered stomata (Supplemental Figure 6A and B), which is consistent with our previous report 229

(Wang et al., 2007). However, the clustered stomatal phenotype of this mkk4 mkk5 double 230

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mutant is not as severe as the double MKK4/MKK5 RNAi lines due to the partial knockdown 231

nature of mkk4 in this double mutant (Zhao et al., 2014). After being normalized to the stomatal 232

density, the calculated pathogen entry rate of mkk4 mkk5 double mutant was still significantly 233

higher than that of the wild type (Figure 3H). 234

Based on gain-of-function evidence, it was concluded that MKK4/MKK5 function as 235

redundant MAPKKs upstream of MPK3/MPK6 in plant immunity (Asai et al., 2002; Ren et al., 236

2002; Ren et al., 2008; Tsuda et al., 2013; Guan et al., 2015). With the available mkk4 mkk5 237

double mutant, we were able to gain loss-of-function evidence to support the idea that 238

MKK4/MKK5 are indeed upstream of MPK3/MPK6 in plant immune response. As shown in 239

Figure 3I, flg22-induced MPK3/MPK6 activation was impaired in mkk4 mkk5 double mutant 240

(Figure 3I). There was a compensatory increase in MPK4 activation, similar to the mpk3 mpk6 241

double mutant inoculated with Botrytis cinerea (Ren et al., 2008). Based on these results, we 242

conclude that MKK4/MKK5, similar to their downstream kinases MPK3/MPK6, are essential to 243

stomatal immunity. Furthermore, mkk4 mkk5 double mutants were also more susceptible to 244

infiltration-inoculated Pst (Figure 3J and K). In contrast, no change in pathogen growth was 245

observed in mkk4 and mkk5 single mutants when Pst was either spray- or infiltration-inoculated 246

(Supplemental Figure 7A and B). Furthermore, stomatal closure in response to flg22 treatment 247

and pathogen entry remained the same in mkk4 and mkk5 single mutants (Supplemental Figure 248

7C and D), demonstrating that MKK4 and MKK5 function redundantly in stomatal immunity. It 249

was previously reported that MKK4 or MKK5 overexpressing lines have enhanced resistance 250

against powdery mildew Golovinomyces cichoracearum (Zhao et al., 2014). However, no change 251

in apoplastic or stomatal immunity against Pst was observed in these MKK4 or MKK5 252

overexpressing lines (Supplemental Figure 7A-D), possibly due to differential defense 253

mechanisms against Pst and powdery mildew in Arabidopsis. Taking these results together, we 254

can conclude that the MKK4/MKK5-MPK3/MPK6 module plays essential roles in both stomatal 255

and apoplastic immunity against Pst. In this study, we mainly focus on the role of this signaling 256

module in stomatal immunity. 257

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258

MPK3/MPK6 activation-induced stomatal closure is independent of ABA biosynthesis and 259

signaling 260

Using the conditional gain-of-function GVG-Nt-MEK2DD transgenic Arabidopsis 261

(abbreviated as DD) (Ren et al., 2002), we found that activation of MPK3/MPK6 in DD plants 262

after dexamethasone (DEX) treatment was sufficient to induce stomatal closure (Figure 4A), 263

which supports our conclusion based on loss-of-functional analyses. DEX itself had no effect on 264

stomatal behavior in Col-0 plants (Figure 4A), demonstrating that DEX-induced stomatal closure 265

in DD plants was a result of MPK3/MPK6 activation. To test whether the gain-of-function 266

MPK3/MPK6 activation-induced stomatal closure is dependent on ABA, we crossed DD with 267

mutants of ABA biosynthesis (aba2-1) (Gonzalez-Guzman et al., 2002) and signaling (abi1-11 268

and ost1-3) (Hua et al., 2012; Acharya et al., 2013) as well as downstream components in 269

stomatal closure (slac1-3, and rbohD) (Torres et al., 2002; Negi et al., 2008; Vahisalu et al., 270

2008). F3 double homozygous lines were used for stomatal closure assay. We found that 271

MPK3/MPK6 activation-induced stomatal closure was normal in the absence of ABA 272

biosynthesis or signaling (Figure 4B and C), demonstrating that MPK3/MPK6 activation induces 273

stomatal closure through a pathway independent of ABA biosynthesis and signaling. Moreover, 274

MPK3/MPK6-activation induced stomatal closure is independent of slow anion 275

channel-associated 1 (SLAC1). In addition, ABA treatment could induce stomatal closure in 276

MPK3SR and MPK6SR plants with and without NA-PP1 pretreatment and in the mkk4 mkk5 277

double mutant (Supplemental Figure 8A-C), suggesting again that ABA and MPK3/MPK6 are 278

independent of each other in inducing stomatal closure. Interestingly, MPK3/MPK6-activation 279

induces stomatal closure is more pronounced in aba2-1 and abi1-11 background (Figure 4B). In 280

addition, stomatal closure induced by ABA treatment is more pronounced in NA-PP1-treated 281

MPK3SR and MPK6SR plants and mkk4 mkk5 double mutant (Supplemental Figure 8B and C). It 282

is possible that both MAPK and ABA signaling pathways are essential for stomatal regulation. In 283

the absence of one pathway, there is a compensatory enhancement in the sensitivity of the other 284

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pathway. Alternatively, these two pathways may negatively regulate each other’s activity to fine 285

tune stomatal aperture in response to changing environmental conditions. Future studies are 286

needed to determine the possible link between these two pathways. 287

288

MPK3/MPK6-activation induced stomatal closure is related to malate/citrate metabolism 289

To explore whether MPK3/MPK6 activation-induced stomatal closure is related to 290

metabolic conversion of osmotically active molecules, such as sucrose, malate, and citrate, we 291

examined starch accumulation and malate/citrate metabolism in guard cells after MPK3/MPK6 292

activation. As shown in Supplemental Figure 9, starch accumulation in guard cells was 293

comparable in ethanol (EtOH, solvent control) and DEX-treated DD plants. Interestingly, 294

MPK3/MPK6 activation-induced stomatal closure was strongly impaired in the presence of 295

malate or citrate (Figure 4D). Next, we checked the expression of several genes encoding 296

enzymes in organic acid metabolism. As shown in Figure 4E, MPK3/MPK6 activation increased 297

the expression of genes encoding NAD-isocitrate dehydrogenase 1 (NAD-IDH1), NAD-IDH2, 298

NAD-IDH5, NADP-malate enzyme 2 (NADP-ME2) and NADP-ME3, while the expression of 299

genes encoding NADP-ME4, NAD-malate enzyme 2 (NAD-ME1), and NAD-ME2 were not 300

affected. Associated with the activation of genes encoding enzymes involved in malate 301

metabolism, we detected a decrease in cellular malate contents in whole leaves (Supplemental 302

Figure 10A). 303

The above experiments were done using whole leaves because wounding associated with 304

guard cell protoplast isolation will activate MPK3/MPK6, which will complicated the results. In 305

silico analysis revealed that genes encoding enzymes in malate metabolism and TCA cycle are 306

expressed at comparable levels in guard cells and whole leaves (Supplemental Table 1). To gain 307

more direct evidence, we examined the enzyme activities in guard cells by using in situ enzyme 308

activity staining (Baud and Graham, 2006). The activity of NAD-ME, NADP-ME, NAD-IDH 309

and NADP-IDH all increased upon MPK3/MPK6 activation, among them the increase in 310

NADP-ME and NAD-IDH activities were much higher compared with the others (Figure 4F and 311

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Supplemental Figure 11). These results suggest that MPK3/MPK6 activation-induced stomatal 312

closure might involve metabolic conversion of osmotically active organic acids. 313

314

Pathogen/PAMP-induced NADP-ME and NAD-IDH activities are dependent on 315

MKK4/MKK5-MPK3/MPK6 316

Gain-of-function evidence revealed that MPK3/MPK6 activation could enhance the 317

activities of enzymes involved in malate metabolism in guard cells. We next tested whether 318

pathogen/PAMPs could also increase the gene expression and activity of these enzymes. As 319

shown in Supplemental Figure 12A and B, NADP-ME2 expression was induced after flg22 320

treatment or Pst inoculation. For the enzyme activity staining, we mainly focused on NADP-ME 321

and NAD-IDH because their high induction by MPK3/MPK6 activation. As shown in Figure 5, 322

enzymatic activities of NADP-ME and NAD-IDH were highly induced by flg22 or Pst in 323

wild-type plants, which is again associated with a decrease in cellular malate contents in whole 324

leaves (Supplemental Figure 10B and C). 325

To gain loss-of-function evidence to support the role of MKK4/MKK5-MPK3/MPK6 326

module in regulating malate metabolism, we analyzed the NADP-ME and NAD-IDH activities 327

in mkk4 mkk5 and the conditional mpk3 mpk6 double mutants. To our surprise, the basal 328

activities of NADP-ME and NAD-IDH were substantially higher in untreated mkk4 mkk5 double 329

mutant (Figure 5). After treatment with flg22 or Pst, the activity of NADP-ME and NAD-IDH 330

did not increase; instead, we observed a slight decrease in their activities (Figure 5). Furthermore, 331

we found that the basal activities of NADP-ME was also higher in NA-PP1-treated, but not in 332

DMSO-treated, MPK3SR and MPK6SR plants (Supplemental Figure 13). Similar with that in 333

mkk4 mkk5 double mutant, activity of NADP-ME in NA-PP1-treated MPK3SR and MPK6SR 334

plants failed to increase after Pst inoculation; and instead, we observed a decrease in NADP-ME 335

activity. These results demonstrated that the MKK4/MKK5-MPK3/MPK6 module is required for 336

pathogen/PAMP-induced increase in organic acid metabolism in guard cells as indicated by the 337

13

increase in enzymatic activities, and loss of function of MKK4/MKK5-MPK3/MPK6 signaling 338

also affects the basal level activities of these enzymes. 339

340

Stomatal immunity is impaired in the presence of malate or citrate 341

The results above support a notion that metabolic conversion of osmotically active 342

organic acids plays a role in stomatal immunity. To provide further evidence, we performed 343

pathogen entry assay in leaves treated with malate or citrate. Mannitol was included as a control. 344

There was a large increase in Pst-lux entry in the presence of malate or citrate (Figure 6A and B), 345

which was accompanied by the impairment of Pst-induced stomatal closure (Figure 6C). In 346

contrast, the non-metabolizable mannitol did not have these effects. These results further support 347

the importance of metabolic conversion of osmotically active organic acids in stomatal 348

immunity. 349

A recent study showed that plant-secreted bioactive metabolites including malate and 350

citrate could stimulate the expression of type III effectors, and therefore, enhance the pathogen 351

virulence (Anderson et al., 2014). To test whether the impaired stomatal immunity is a result of 352

enhanced virulence of Pst in the presence of malate or citrate, we first treated Pst-lux with 353

mannitol, malate, or citrate for 1 hr, and then performed pathogen entry assay using these 354

pretreated Pst-lux. As shown in Figure 6D and E, Pst-lux entry into leaves were not affected. 355

356

Differential sensitivity of ABA- and MPK3/MPK6-induced stomatal closure to coronatine 357

The essential function of ABA in plant stomatal defense, as suggested in a number of 358

reports (Melotto et al., 2006; Zeng and He, 2010; Du et al., 2014; Lim et al., 2014), was recently 359

called into question (Montillet et al., 2013). Thus, we re-examined the importance of ABA 360

biosynthesis and signaling in stomatal immunity using the pathogen entry assay. As shown in 361

Figure 7A and B, Pst-lux entry was much higher in aba2-1, ost1-3, slca1-3, and rbohD mutants. 362

In addition, flg22- or Pst-induced stomatal closure was strongly impaired in aba2-1, ost1-3, 363

14

slca1-3, and rbohD (Figure 7C and D), supporting the conclusion that ABA biosynthesis and 364

signaling are essential to stomatal immunity. 365

Coronatine (COR), a phytotoxin produced by Pst, is able to open plant stomata to 366

facilitate the pathogenesis process (Melotto et al., 2006; Zheng et al., 2012). Consistent with 367

previous reports, ABA-induced stomatal closure could be blocked by COR (Figure 8A). 368

However, MPK3/MPK6 activation-induced stomatal closure was insensitive to COR (Figure 8B). 369

In addition, we found that activities of NADP-ME and NAD-IDH were not induced by ABA 370

(Supplemental Figure 14), indicating that ABA-induced stomatal closure is independent of 371

metabolic conversion of osmotically active organic acids. Furthermore, we found that 372

MPK3/MPK6 activation-induced stomatal closure could not be reopened by Pst (Figure 8C and 373

D). These observations further support that ABA and MKK4/MKK5-MPK3/MPK6 represent 374

two different pathways in mediating stomatal closure. 375

376 DISCUSSION 377

In this study, we demonstrate that Arabidopsis MPK3 and MPK6 are essential to plant 378

immunity. They play redundant functions in both stomatal and apoplastic immunity. MKK4 and 379

MKK5 are redundant MAPKKs upstream of MPK3 and MPK6 in the same pathway based on 380

genetic and biochemical analyses. Stomatal aperture measurements and pathogen entry assays 381

revealed that mpk3 mpk6 and mkk4 mkk5 double mutants were unable to close their stomata in 382

response to PAMP treatment or Pst inoculation, which leads to compromised resistance in these 383

mutants (Figure 1-3). It appears that MPK3/MPK6 activation promotes the metabolic conversion 384

or consumption of osmotically active organic acids, therefore, induces stomatal closure during 385

plant defense responses. Furthermore, we found that MPK3/MPK6-induced stomatal closure is 386

independent of ABA biosynthesis and signaling (Figure 4A-C), despite the fact that ABA is 387

essential to stomatal immunity (Figure 7A-D) (Melotto et al., 2006; Zheng et al., 2012; Du et al., 388

2014). We proposed an interdependent model in which the MPK3/MPK6 pathway and the ABA 389

pathway can independently trigger stomatal closure when MPK3/MPK6 are activated in the 390

15

gain-of-function system or when ABA is exogenously applied, but both pathways are required 391

for an effective stomatal immunity upon sensing of pathogen PAMPs (Figure 8E). 392

Previous studies showed that mpk3 and mpk6 single mutants as well as guard cell-specific 393

MPK3 antisense plants were strongly impaired in flg22-, bacterium-, or LPS-induced stomatal 394

closure (Gudesblat et al., 2009; Montillet et al., 2013). It was also reported that ABA-dependent 395

and ABA-independent (but MPK3- or MPK6-dependent) pathways converge on SLAC1 in 396

stomatal immunity (Montillet et al., 2013). Our gain-of-function data showed that MAPK 397

activation-induced stomatal closure is independent of SLAC1 (Figure 4B), but related to organic 398

acids metabolism (Figure 4D-F). More importantly, no clear susceptible phenotype was observed 399

in mpk3 single mutant, and only a slightly higher bacterial growth was observed in mpk6 mutant 400

(0.01< P<0.05) when spray- or infiltration-inoculated with Pst (Supplemental Figure 1A and C), 401

which is not consistent with the reported deficiency in stomatal immunity in the mpk3 and mpk6 402

single mutants. Furthermore, our detailed analysis revealed that mpk3 and mpk6 single mutants 403

exhibited normal flg22-induced stomatal closure (Supplemental Figure 1B). It is possible that the 404

inconsistency is a result of the weak phenotype of the single MAPK mutant/RNAi lines and the 405

variation of assay conditions. In contrast, the double mpk3 mpk6 and mkk4 mkk5 mutant plants 406

showed consistent and strong phenotypes that could be easily observed. 407

In our model, the MKK4/MKK5-MPK3/MPK6 pathway and ABA pathway function 408

interdependently. Instead of converging on SLAC1, they modulate two different facets of 409

osmoregulation in guard cells during pathogen/PAMP-induced stomatal closure. Activation of 410

ABA signaling pathway leads to the activation of OST1 kinase that can directly phosphorylate 411

RbohD and SLAC1, resulting in efflux of inorganic anions (mainly Cl-) through SLAC1 (Negi et 412

al., 2008; Vahisalu et al., 2008; Geiger et al., 2009) and cause subsequent plasma membrane 413

depolarization, activation of quick anion channel (QUAC1) (Sasaki et al., 2010) and outward 414

recertifying K+ channel (Hosy et al., 2003). In contrast, activation of MPK3/MPK6 cascade 415

stimulates metabolic conversion of osmotically active organic acids. Together, these two 416

pathways would confer robust stomatal immunity. Interdependent regulation appears to be a 417

16

common strategy to confer signaling robustness. A recent study demonstrated that SLAC1 418

activity was also interdependently regulated by OST1 and CDPKs, in which phosphorylation of 419

SLAC1 at Ser59 and Ser120 by OST1 and CDPKs, respectively, are both essential for 420

ABA-induced stomatal closure (Brandt et al., 2015). 421

ABA and MPK3/MPK6 pathways also differ in their sensitivity to COR, a phytotoxin. 422

The ABA signaling pathway can be blocked by COR (Figure 8A), as reported (Zheng et al., 423

2012). In contrast, the MPK3/MPK6 activation-mediated stomatal closure is insensitive to COR 424

(Figure 8B). It is worth noting that MPK3/MPK6 activation in the conditional gain-of-function 425

DD system is longer lasting or higher in magnitude than that after flg22 treatment or Pst 426

infection (Figure 3I) (Ren et al., 2002; Guan et al., 2015). Instead, it is similar to MPK3/MPK6 427

activation in plants after necrotrophic fungal infection or during effector-triggered immunity 428

(ETI) in duration and magnitude (Ren et al., 2008; Tsuda et al., 2013), implying a possible role 429

for this stronger and longer lasting MPK3/MPK6 activation in necrotrophic fungal infection- or 430

ETI-related stomatal immunity. It was reported previously that MPK9 and MPK12 function 431

downstream of ROS and modulate SLAC1 activity in guard cell ABA signaling (Jammes et al., 432

2009; Jammes et al., 2011). Based on genetic and biochemical analyses, MKK4 and MKK5 are 433

unable to activate MPK9 and MPK12 in vivo and in vitro (Asai et al., 2002; Lee et al., 2009; 434

Nagy et al., 2015). As a result, MPK9/MPK12 and MPK3/MPK6 are likely to function as two 435

independent MAPK cascades in stomatal movement. In our model, we emphasized the potential 436

role of MPK3/MPK6 in promoting the metabolism of malate/citrate. However, we currently 437

cannot exclude the possibility that the transportations of these organic acids in and out of cells 438

and/or vacuoles, are also regulated in the process. 439

AtABCB14, an ABC transporter, was identified as a plasma membrane-localized malate 440

importer to modulate stomatal movement in response to CO2 (Lee et al., 2008). The plasma 441

membrane-localized malate exporter, aluminum-activated malate transporter 12 (ALMT12), was 442

shown to be involved in ABA- and CO2-induced stomatal closure (Meyer et al., 2010). In 443

addition, γ-aminobutyric acid (GABA) can control the transporter activity of ALMTs (Ramesh et 444

17

al., 2015; Gilliham and Tyerman, 2016). At this stage, the role of ALMTs in pathogen- and/or 445

PAMP-mediated stomatal closure remains to be determined. Recently, starch biosynthesis was 446

implicated in high CO2-induced stomatal closure (Azoulay-Shemer et al., 2016). However, 447

Lugol’s IKI staining indicated no change in starch accumulation after MPK3/MPK6 activation 448

(Supplemental Figure 9), suggesting that starch biosynthesis may not play any role in 449

PAMP/Pathogen-induced stomatal closure. 450

At this point, measuring the change in malate or citrate levels in guard cells in response 451

to MPK3/MPK6 activation during plant immunity remains technically challenging. We 452

attempted to enrich the epidermal fraction by using the blender method (Zhang et al., 2008). In 453

parallel, we also included a group with Cellulase R-10 and Macerozyme R-10 treatment to 454

remove mesophyll cells attached to epidermal strips. In both case, we found malate levels were 455

too low to be reliably detected in both EtOH- and DEX-treated DD samples. This could be a 456

result of strong MPK3/MPK6 activation caused by the wounding stress and cell wall elicitor 457

released during protoplast preparation, which in turn can activate the expression of genes 458

encoding enzymes related to organic acid metabolism. Consistent with this, we observed that leaf 459

petiole wounded by forceps during handling showed higher NADP-ME and NAD-IDH activities 460

(Figure 4F and Supplemental Figure 11, 13-15). (A note: we removed portions of the leaf 461

petioles that had higher activity staining when possible before photography. As a result, not all 462

leaf petioles showed higher activity staining.) Developing tools that will allow in situ single cell 463

detection such as cellular sensors of malate or citrate will be required to achieve this goal. 464

Nonetheless, we found that malate content decreases in whole leaves after MPK3/MPK6 465

activation in DD plants (Supplemental Figure 10A) or Col-0 plants after pathogen/PAMP 466

treatment (Supplemental Figure 10B and C). Furthermore, in situ enzyme activity assays 467

revealed higher levels of increase in NADP-ME and NAD-IDH activities in guard cells than 468

other types of leaf cells (Figure 4F, Figure 5), which should correlate with a more pronounced 469

decrease in malate in guard cells, supporting our working model (Figure 8E). In addition, we 470

found that exogenously added malate/citrate could block stomatal closure triggered by 471

18

MPK3/MPK6 activation or Pst inoculation (Figure 4D, Figure 6C), and compromise stomatal 472

immunity (Figure 6A and B). 473

Due to gene redundancy and the nature of metabolic flexibility, it is very challenging to 474

provide genetic evidence for the role of metabolic conversion of organic acids in stomatal 475

immunity. There are four NADP-ME isoforms and two NAD-ME isoforms in Arabidopsis 476

(Wheeler et al., 2005; Tronconi et al., 2008), and their function can be compensated by each 477

other. Although NADP-ME activity was greatly reduced in nadp-me2 mutant, flg22-induced 478

stomatal closure was not affected (Supplemental Figure 15). By using two independent 479

nadp-me2 alleles, a previous study also showed normal disease resistance against Pst (Wheeler et 480

al., 2005; Li et al., 2013). Double mutant plants lacking both NAD-ME1 and NAD-ME2 also 481

grow normally (Tronconi et al., 2008). Gene redundancy and functional compensation were also 482

reported for NAD-IDH and NADP-IDH (Lemaitre et al., 2007; Mhamdi et al., 2010). 483

Nonetheless, in our gain-of-function MPK3/MPK6 system, all these four enzymes were activated 484

(Figure 4F), and MPK3/MPK6 activation- and Pst inoculation-induced stomatal closure was 485

impaired in the presence of malate or citrate (Figure 4D and Figure 6C). Collectively, these 486

results do support a positive role for metabolic conversion of organic acids in stomatal immunity. 487

MPK3/MPK6 activation results in an increase in the activities of NADP-ME and 488

NAD-IDH in guard cells. Unexpectedly, we also observed higher basal-level activities of 489

NADP-ME and NAD-IDH in the loss-of-function mkk4 mkk5 double mutant (Figure 5). The 490

basal-level activities of NADP-ME in NA-PP1-treated MPK3SR and MPK6SR plants were also 491

higher than that of the control plants treated with DMSO (Supplemental Figure 13A and B), 492

indicating that MKK4/MKK5-MPK3/MPK6 signaling module may also play a role in 493

maintaining the basal levels of these enzymes in plants. It is possible that multiple signaling 494

pathways are involved in controlling the activities of NADP-ME and NAD-IDH, which could be 495

important in maintaining the metabolic homeostasis in plants. Loss of function of the 496

MPK3/MPK6 cascade may result in the overcompensation as a result of the activation of the 497

other signaling pathway. In mammals, both extracellular signal-regulated kinase 2 (ERK2, a 498

19

MAPK) and cyclin-dependent kinase 5 (Cdk5) are involved in maintaining glucose homeostasis 499

in adipose tissues by phosphorylating PPARγ (peroxisome proliferator-activated receptor γ) on 500

Ser273 (Choi et al., 2010). However, when Cdk5 is knocked out, PPARγ phosphorylation on 501

Ser273 residue was found to be much higher. It turns out that, besides PPARγ, Cdk5 can also 502

phosphorylate the MAPKK upstream of ERK2 on a novel site and inhibit ERK2 activation 503

(Banks et al., 2015). The loss of Cdk5, therefore, leads to the hyper-activation of ERK2, 504

resulting in a higher level of phosphorylation of PPARγ on Ser273. It will be interesting to 505

determine the signaling pathways that crosstalk with MKK4/MKK5-MPK3/MPK6 in 506

maintaining the basal organic acid metabolism in guard cells. 507

During light-induced stomatal opening, tobacco guard cells were capable of fixing CO2, 508

operating the TCA cycle, and synthesizing amino acids (Daloso et al., 2015). Furthermore, the 509

levels of TCA cycle intermediates and amino acids are dynamically regulated and respond 510

differently to K+ and sucrose, suggesting that guard cells may take advantage of the flexibility of 511

metabolism to fine tune stomatal aperture during stomatal opening. It was previously shown that 512

Sclerotinia sclerotiorum, a phytopathogenic fungus, interferes with the stomatal immunity by 513

secreting oxalate, an intermediate of TCA cycle (Guimaraes and Stotz, 2004). In addition, 514

oxalate could promote stomatal opening and inhibit ABA-induced stomatal closure (Guimaraes 515

and Stotz, 2004). Citrate, another intermediate of the TCA cycle, could also promote stomatal 516

opening and delay dark-induced stomatal closure (Jinno, 1981). It is worth noting that, in these 517

two studies, oxalate or citrate was added to closed stomata. Upon illumination, oxalate or citrate 518

is imported into guard cell and thus increases turgor pressure, which delays ABA- or 519

dark-induced stomatal closure. In another study, it was found that malate or fumarate, both 520

intermediates of TCA cycles, induces stomatal closure when added to opened stomata (Araujo et 521

al., 2011b). These TCA cycle intermediates may affect stomatal movement in two different ways: 522

1) by functioning as osmolytes to open stomata (Lee et al., 2008), and 2) by functioning as 523

signals to close the opened stomata (Araujo et al., 2011b). In a similar case, sucrose, as an 524

osmolyte, accumulates gradually during the day to maintain stomatal in an open state (Lawson, 525

20

2009). However, exogenous application of sucrose to open stomata induces ABA 526

signaling-dependent stomatal closure (Kelly et al., 2013). 527

In order to address the role of malate or citrate in MPK3/MPK6 activation- or 528

Pst-induced stomatal closure, we first allowed the plants to dark-adapt for 1 hr and then treated 529

them with malate or citrate (Figure 4D, Figure 6C). Under such conditions, both organic acids 530

can block the stomatal closure induced by gain-of-function activation of MPK3/MPK6 or Pst 531

inoculation. Together with previous studies, our data here highlight the importance of metabolic 532

shift of organic acids such as malate and citrate in regulating stomatal movement. In plant 533

immune response, the MPK3/MPK6 cascade and ABA are two essential and interdependent 534

signaling pathways that control, respectively, organic acid metabolism and ion channels, two 535

branches of osmotic regulation in guard cells that control stomatal opening/closure. The 536

activation of both pathways would confer a robust response in plant stomatal immunity. 537

538 METHODS 539

Plant materials and growth conditions 540

Arabidopsis thaliana plants were grown in soil under 10 hr day/14 hr night cycle at 22 oC 541

in a CONVIRON walk-in growth chamber equipped with high intensity discharge lights. The 542

light intensity was set to 100 µmol m-2 s-1. The humidity of the growth chamber was set to 75% 543

RH. Three- to four-week-old plants were used for experiments. Col-0 ecotype was used as the 544

wild type. Mutant alleles and transgenic lines of mpk3-1 (SALK_151594), mpk6-2 545

(Salk_073907), mpk6-3 (Salk_127507), DD (GVG-NtMEK2DD), and MPK6SR (mpk3 mpk6 546

PMPK6:MPK6YG, lines # 31 and #58) were reported previously (Ren et al., 2002; Wang et al., 547

2007; Xu et al., 2014). MPK3SR (mpk3 mpk6 PMPK3:MPK3TG, lines #28 and #64) was generated 548

similarly as MPK6SR (Xu et al., 2014). Thr116 residue in the ATP-binding pocket of MPK3 was 549

mutated to Gly (MPK3TG) in a genomic clone in pCambia3300 vector (www.cambia.org). 550

PMPK3:MPK3TG construct was transformed into mpk3 mpk6/+ plants. Single-insertion lines with 551

MPK3TG expressed at a similar level as the endogenous MPK3 were identified by immunoblot 552

analysis using an anti-MPK3 antibody (Sigma, http:// www.sigmaaldrich.com). F3 homozygous 553

21

PMPK3:MPK3TG transgenic plants in the mpk3 mpk6 background (mpk3 mpk6 PMPK3:MPK3TG), 554

also known as inhibitor-sensitized MPK3 variant-rescued plants (MPK3SR), lines #64 was used 555

in this study. 556

Mutant alleles of nadp-me2-1 (Salk_073818), aba2-1 (CS156), ost1-3 (Salk_008068), 557

slac1-3 (Salk_099139) and rbohD (CS9555) were obtained from the Arabidopsis Biological 558

Resource Center. The abi1-11 mutant was kindly provided by Prof. Zhizhong Gong (China 559

Agricultural University). The mkk4 and mkk5 single TILLING mutants (Zhao et al., 2014) were 560

first backcrossed with Col-0 to remove the er-105 mutant allele and then crossed to generate 561

mkk4 mkk5 double mutant. The MKK4 and MKK5 over-expressing lines, MKK4OE #5, 562

MKK4OE #7, MKK5OE #2, and MKK5OE #5, were kindly provided by Prof. Dingzhong Tang 563

(Chinese Academy of Sciences, China). 564

565

Pathogen assay and pathogen entry assay 566

For Pst resistance assay, three- to four-week-old Arabidopsis plants were 567

spray-inoculated with Pst suspension (OD600 = 0.5) in 10 mM MgCl2 with 0.02% Silwet L-77. 568

The 5th and 6th leaves were detached and washed with 0.02% Silwet L-77 before leaf discs were 569

punched out for bacterial growth assays as previously described (Guan et al., 2015). 570

For pathogen entry assays, detached leaves were first illuminated for 2.5 hr to fully open 571

stomata, and then treated with Pst-lux (final OD = 0.5) for 1 hr. The treated leaves were washed 572

by 0.02% Silwet L-77 for 10 seconds with stirring at maximum setting. Pathogen entry was 573

measured by using a photon counting image system (Image 32 system, Photek, United Kingdom) 574

or by direct counting of colony-forming units. 575

576

In situ enzyme activity staining 577

In situ enzyme activity staining was performed as previously described with 578

modifications (Baud and Graham, 2006). Leaves were fixed by vacuum infiltration in fixation 579

solution (2% paraformaldehyde, 2% polyvinylpyrrolidone 40000 and 1 mM dithiothreitol, pH = 580

22

7.0) for 1 hr on ice. After fixation, the leaves were rinsed five times with distilled water and 581

stored in water overnight at 4 oC. For NAD-ME assay, the leaves were incubated in reaction 582

solution [100 mM HEPES-KOH (pH 6.5), 4 mM nicotinamide adenine dinucleotide (NAD+), 0.1 583

mM Acetyl Co enzyme A, 2 mM fructose-1,6-biphosphate (FBP), 2 mM fumarate, 5 mM MnCl2, 584

5 mM malate, 0.8 mM nitroblue tetrazolium dye (NBT), and 0.4 mM phenazine methosulfate 585

(PMS)] for 3 hr at 30 oC. For NAD-IDH assay, the leaves were incubated in reaction solution 586

[100 mM HEPES (pH 7.5), 5 mM MgCl2, 2 mM NAD+, 10 mM D, L-isocitrate, 0.8 mM NBT, 587

and 0.4 mM PMS] for 3 hr at 30 oC. For ABA-treated leaves, a 4-hr incubation at 30

oC was 588

carried out. For NADP-IDH assay, the leaves were incubated in reaction solution [100 mM 589

HEPES (pH 7.5), 5 mM MgCl2, 1 mM NADP+, 10 mM DL-isocitrate, 0.8 mM NBT, and 0.4 590

mM PMS] for 3 hr at 30 oC. For NADP-ME assay, the leaves were incubated in reaction solution 591

[50 mM HEPES-KOH (pH 7.65), 1 mM NADP+, 5 mM malate, 10 mM MgSO4, 0.4 mM PMS, 592

and 0.8 mM NBT] for 1 hr at 30 oC. For ABA treated leaves, a 90-min incubation at 30 oC was 593

carried out. 594

595

Stomatal aperture measurement 596

Epidermal peels or intact leaves from at least 4 independent 3- to 4-week-old plants of 597

various genotypes were used for stomatal closure analysis. Epidermal peels or intact leaves were 598

floated on stomatal opening buffer [10 mM 2-(N-morpholino) ethanesulfonic acid (MES), pH = 599

6.15 and 30 mM KCl] under light for 2.5 hr to fully open the stomata. They were then treated 600

with flg22 (3 µM), Pseudomonas syringae pv tomato DC3000 (OD = 0.1), dexamethasone (DEX) 601

(5 µM), coronatine (COR) (1 µg/ml), ABA (10 µM) for indicated period of time. For MPK6SR 602

and MPK3SR, 1-(1,1-dimethylethyl)-3-(1-naphthalenyl)-1H-pyrazolo[3,4-d]pyrimidin-4-amine 603

(NA-PP1) (2 µM final concentration) or equal volume of DMSO (solvent for NA-PP1 stock) was 604

added at the beginning of illumination. For testing the effect of mannitol, malate, or citrate on 605

Pst-induced stomatal closure, plants were dark adapted for 1 hr to fully close the stomata and 606

then were floated on MES buffer (10 mM MES, pH = 6.15 and 30 mM KCl) containing mannitol 607

23

(60 mM), malate (20 mM) or citrate (20 mM). Stomatal aperture was measured using ImageJ 608

software. 609

610

Starch staining 611

Leaves were floated on MES buffer (10 mM MES pH 6.15, and 30 mM KCl) under light 612

for 2.5 hr to fully open the stomata and then were treated with either ethanol (solvent control) or 613

DEX (5 µM) for 5 hr. The treated leaves were fixed in 80% ethanol and 5% formic acid solution 614

for 10 min at 80 oC. After being wash twice with 80% ethanol for 5 min at 80 oC each, the leaves 615

were stained with Lugol’s IKI solution (5.7 mM iodine and 43.4 mM potassium iodide) for 10 616

min at room temperature (Fritzius et al., 2001). Cell images were taken using an Olympus 617

microscope fitted with a digital camera. 618

619

In silico and quantitative RT-PCR analysis 620

Genes involved in malate metabolism and TCA cycle were retrieved from BioCyc 621

database (Caspi et al., 2016), and their expression levels were extracted from a previous 622

microarray profiling analysis (Bates et al., 2012). Real-time quantitative PCR (qPCR) was 623

performed as previously described (Mao et al., 2011). Total RNA was extracted using TRIzol 624

reagent (Invitrogen). After DNase treatment, 1 µg of total RNA was used for reverse 625

transcription. Real-time qPCR analysis was performed using an ABI 7500 real-time PCR 626

machine (Life Technologies). EF-1α was used as an internal control. The primer pairs used for 627

qPCR are listed in Supplemental Table 2 online. 628

629

Protein extraction and immunoblot analysis 630

Protein was extracted as previously described (Liu and Zhang, 2004). Anti-Flag (Sigma, 631

St. Louis, MO) and anti-pTEpY (Cell Signaling Technology, Danvers, MA) antibodies were 632

used to detect NtMEK2DD protein expression and MPK3/MPK6 activation, respectively. 633

634

24

Spectrophotometric assay of malate levels 635

After treatment, about 0.1 g of leaf tissue was boiled in 1 ml ddH2O for 10 min to release 636

malate. Quantification of malate was performed as described (Peleg et al., 1990) with 637

modifications. Two µl of extract was used in a 100 µl reaction containing 0.6 M glycylglycine 638

pH 10, 0.1 M L-glutamate, 0.5 mM 3-(4,5-Dimethylthiazol-2-yl)-2,5-Diphenyltetrazolium 639

Bromide (MTT), 4 mM NAD+, 0.04 mM PMS, 1 µg glutamate oxaloacetate transaminase (GOT), 640

and 2.5 µg malate dehydrogenase (MDH) for 15 min at 37 oC. The reaction was stopped by the 641

addition of stop solution [20% SDS and 50% dimethyformamide (DMF)], and the absorbance at 642

570 nm was measured. 643

644

Statistical tests 645

For experiments with multiple time points, at least two independent repetitions were 646

performed. For single time-point experiments, at least three independent repetitions were 647

performed. Data from one of the independent repetitions with similar results were shown in the 648

figures. One-way ANOVA Tukey’s test was used for statistical analysis. One and two asterisks 649

above the columns were used to indicate differences that are statistically significant (P < 0.05) 650

and very significant (P < 0.01), respectively. 651

652

Accession Numbers 653

Sequence data from this article can be found in the Arabidopsis Genome Initiative or 654 GenBank/EMBL databases under the following accession numbers: 655

MPK3 (At3g45640), MPK6 (At2g43790), MKK4 (At1g51660), MKK5 (At3g21220), 656

ABA2 (At1g52340), ABI1 (At4g26080), OST1 (At4g33950), SLAC1 (At1g12480), RbohD 657

(At5g47910), NADP-ME2 (At5g11670), NADP-ME3 (At5g25880), NADP-ME4 (At1g79750), 658

NAD-ME1 (At2g13560), NAD-ME2 (At4g00570), NAD-IDH1 (At4g35260), NAD-IDH2 659

(At2g17130), NAD-IDH5 (At5g03290), ACS2 (At1g01480), and EF1α (At5g60390). 660

661

25

Supplemental Data 662

Supplemental Figure 1. Stomatal and apoplastic immunity in mpk3 and mpk6 single mutants. 663

Supplemental Figure 2. Stomatal immunity in independent MPK3SR and MPK6SR lines. 664

Supplemental Figure 3. Apoplastic immunity in independent MPK3SR and MPK6SR lines 665

Supplemental Figure 4. Flg22- or Pst-induced stomatal closure was abolished in leaves of the 666

conditional mpk3 mpk6 double mutants. 667

Supplemental Figure 5. Establishing the pathogen entry assay. 668

Supplemental Figure 6. Stomata patterning in Col-0 and mkk4 mkk5 double mutant. 669

Supplemental Figure 7. Apoplastic and stomatal immunity in MKK4 and MKK5 single mutants 670

and their over-expressing lines. 671

Supplemental Figure 8. Loss of function of MPK3/MPK6 or their upstream MKK4/MKK5 672

leads to hypersensitivity to ABA-induced stomatal closure. 673

Supplemental Figure 9. Normal starch accumulation in guard cells of DD plants after DEX 674

treatment. 675

Supplemental Figure 10. Activation of MPK3/MPK6 in DD plants or Col-0 plants after 676

pathogen/PAMP treatment decreases total malate contents in leaves. 677

Supplemental Figure 11. Validation of the in situ enzyme activity assay. 678

Supplemental Figure 12. Quantitative RT-PCR analysis the expression of malate metabolism 679

related genes in response to flg22 or Pst. 680

Supplemental Figure 13. MPK3 and MPK6 are required for Pst-induced activation of 681

NADP-ME. 682

Supplemental Figure 14. NADP-ME and NAD-IDH activities were not affected by ABA 683

treatment. 684

Supplemental Figure 15. Flg22-induced stomatal closure was not affected in nadp-me2 mutant. 685

Supplemental Table 1. Expression of TCA cycle-related genes in guard cells is comparable 686

with that in intact leaves. 687

Supplemental Table 2. Primers used for reverse transcription quantitative PCR analysis. 688

26

Supplemental Table 3. ANOVA tables for statistical analysis. 689

690

ACKNOWLEDGEMENTS 691

We thank Dr. Zhizhong Gong (China Agricultural University), Dingzhong Tang (Chinese 692

Academy of Sciences), and Arabidopsis Biological Resource Center for mutant seeds. This work 693

was supported by grants from the Natural Science Foundation of China (31670268 and 694

31272029), the 111 Project (Grant B14027), and the National Science Foundation (IOS-0743957 695

and MCB-0950519) to J.X. and S.Z. 696

697

AUTHOR CONTRIBUTIONS 698

J.S., M.Z., T.S., and L.Z. performed experiments and data analysis with substantial inputs699

from W.L., J.X., and S.Z. J.S. and S.Z. designed the experiments. J.S., J.X., and S.Z. wrote the 700

manuscript. 701

702

COMPETING INTERESTS 703

The authors declare no competing financial interest. 704

705

Figure 1. Arabidopsis MPK3 and MPK6 play an essential role in stomatal immunity. (A and B) Loss of function of MPK3 and MPK6 compromises plant resistance against Pst. Col-0, MPK6SR (line #58) and MPK3SR (line #64) plants were pre-treated with DMSO (solvent control) or NA-PP1 (10 mM) for 3 hr, and then spray-inoculated with Pst (OD600 = 0.5). A second spray-treatment of DMSO or NA-PP1 was performed 1.5 days post inoculation (dpi). Photos were taken at 3 dpi (A) and Bacterial growth was measured at 2.5 dpi (B). Values are means ± SD, **P ≤ 0.01, n = 3. (C and D) Loss of function of MPK3 and MPK6 compromises apoplastic immunity. Col-0, MPK3SR and MPK6SR plants were first spray with 10 mM NA-PP1 and then infiltration-inoculated with Pst (OD600 = 0.0005). Photos were taken at 3 dpi (C) and bacterial growth was measured at 0 and 3 dpi (D). Values are means ± SD, **P ≤ 0.01, n = 3. (E and F) Pathogen/PAMP-triggered stomatal closure is dependent on MPK3/MPK6. Epidermal peels of Col-0, MPK6SR (line #58) and MPK3SR (line #64) plants were floated on stomatal opening buffer containing DMSO or NA-PP1 (2 mM) under light for 2.5 hr and then exposed to flg22 for 3 hr (E) or Pst (OD = 0.1) for 1 hr (F) before stomatal aperture measurements. Values are means ± SD, **P ≤ 0.01, n ≥ 30.

Figure 2. Pathogen entry assay revealed a compromised stomatal immunity in mpk3 mpk6 double mutant. (A and B) Detached leaves of Col-0, MPK6SR (line #58) and MPK3SR (line #64) plants were floated on stomatal opening buffer containing DMSO or NA-PP1 (2 mM) under light for 2.5 hr and then exposed to Pst-lux (OD = 0.5) for 1 hr. Leaves were washed with 0.02% Silwet L-77 with stirring. Pst entry was measured by photon counting imaging (A) or direct counting of the bacterial population (B). Values are means ± SD, *P ≤ 0.05 and **P ≤ 0.01, n = 3. (C) Stomatal density was determined by using leaves of the same position from Col-0, MPK6SR (line #58), and MPK3SR (line #64) plants. Values are means ± SD, **P ≤ 0.01, n = 6. (D) Pst entry rates were calculated as Pst number per mm2 divided by stomatal density (number of stomata per mm2) in one hour. Values are means ± SD, *P ≤ 0.05 and **P ≤ 0.01, n = 3.

Figure 3. MKK4 and MKK5 play redundant function in stomatal immunity. (A and B) Loss of function of MKK4 and MKK5 compromises plant resistance against Pst. Col-0 and mkk4 mkk5 double mutant plants were spray-inoculated with Pst (OD = 0.5). Photos were taken at 3 dpi (A) and bacterial growth was measured at 2.5 dpi (B). Values are means ± SD, **P ≤ 0.01, n = 3. (C and D) Pathogen/PAMP-induced stomatal closure is dependent on MKK4/MKK5. Leaves of Col-0 and mkk4 mkk5 plants were floated on stomatal opening buffer under light for 2.5 hr and then exposed to flg22 for 3 hr (C) or Pst (OD = 0.1) for 2 hr (D) before stomatal aperture was measured. Values are means ± SD, **P ≤ 0.01, n ≥ 30. (E and F) Pathogen entry assay revealed a compromised stomatal immunity in mkk4 mkk5 double mutant. Detached leaves of Col-0 and mkk4 mkk5 plants were floated on stomatal opening buffer under light for 2.5 hr and then exposed to Pst-lux (OD = 0.5) for 1 hr. Leaves were washed in 0.02% Silwet L-77 with stirring. Pst entry was measured by photon counting (E) or direct counting of the bacterial population (F). Values are means ± SD, **P ≤ 0.01, n = 3. (G) Increased stomatal density of mkk4 mkk5 double mutant. Values are means ± SD, **P ≤ 0.01, n = 6. (H) Pst entry rate was calculated by dividing Pst number per mm2 by stomatal density. Values are means ± SD, **P ≤ 0.01, n = 3. Note: For measuring stomatal aperture in mkk4 mkk5, only separated stomata were selected for analysis. (I) Loss-of-function of MKK4 and MKK5 compromises flg22-induced MPK3/MPK6 activation. Twelve-day-old Col-0 and mkk4 mkk5 seedlings grown in liquid ½ MS medium were treated with 50 nM flg22 for indicated time. MAPK activation was detected by immunoblot analysis using anti-pTEpY antibody. Equal loading was confirmed by Coomassie Brilliant Blue (CBB) staining. (J and K) Loss of function of MKK4 and MKK5 compromises apoplastic immunity. Col-0 and mkk4 mkk5 plants were infiltration-inoculated with Pst (OD600 = 0.0005). Photos were taken at 3 dpi (J) and bacterial growth was measured at 0 and 3 dpi (K). Values are means ± SD, **P ≤ 0.01, n = 3.

Figure 4. MPK3/MPK6 activation-induced stomatal closure is independent of ABA but related to organic acid metabolism. (A) Gain-of-function activation of MPK3/MPK6 induces stomatal closure.Epidermal peels of DD and Col-0 plants were floated on stomatal opening bufferunder light for 2.5 hr and then treated with DEX (5 mM) or ethanol (EtOH, solventof DEX stock solution). Stomatal aperture was determined at indicated time afterDEX treatment. Values are means ± SD, *P ≤ 0.05 and **P ≤ 0.01, n ≥ 30. (B)MPK3/MPK6 activation is sufficient to triggered stomatal closure in the absenceof ABA biosynthesis or signaling. Detached leaves from DD, DD aba2-1, DDabi1-11, DD ost1-3, DD slac1-3, and DD rbohD plants were floated on stomatalopening buffer under light for 2.5 hr and then treated with EtOH or DEX (5 mM)for 5 hr before stomatal aperture was determined. Values are means ± SD, *P ≤0.05, n ≥ 30. (C) DD induction and MPK3/MPK6 activation in DD, DD aba2-1, DDabi1-11, DD ost1-3, DD slac1-3, and DD rbohD plants. Leaf tissues werecollected at the time of aperture measurement as in (B). Induction of DDexpression and MPK3/MPK6 activation were detected by immunoblot analysisusing anti-Flag and anti-pTEpY antibodies, respectively. Equal loading wasconfirmed by Coomassie Brilliant Blue (CBB) staining. (D) Exogenously addedmalate or citrate can reverse MPK3/MPK6-induced stomatal closure. DD plantswere first dark-adapted for 1 hr. Detached leaves were floated on stomatalopening buffer containing mannitol, malate, or citrate (20 mM) under light for 2.5hr and then treated with EtOH or DEX (5 mM) for 5 hr before stomatal aperturewas measured. Values are means ± SD, **P ≤ 0.01, n ≥ 30. (E) Induction ofgenes encoding enzymes in organic acid metabolism. Total RNA was extractedfrom 3-week-old DD plants treated as in (B). Transcript levels were determined

by quantitative RT-PCR. EF1a was used as an internal control. Values are means ± SD, *P ≤ 0.05 and **P ≤ 0.01, n =3. (F) In situ enzyme activity assays revealed an increase in malate/citrate metabolic enzymes in guard cells after MPK3/MPK6 activation. DD plants were treated as in (B), and the in situ enzyme activity assay was performed as described.

Figure 5. MKK4 and MKK5 are required for flg22- or Pst-induced activation of NADP-ME and NAD-IDH. Intact leaves of Col-0 and mkk4 mkk5 plants were floated on stomatal opening buffer under light for 2.5 hr and then exposed to flg22 for 3 hr (A and C) or Pst (OD = 0.1) for 2 hr (B and D). After being fixed in paraformaldehyde solution for 1 hr, and washed overnight, the leaves were subjected to in situ NADP-ME (A and B) or NAD-IDH (C and D) activity staining.

Figure 6. Stomatal immunity is impaired in the presence of malate or citrate. (A and B) Pathogen entry assay demonstrated that treatment with malate or citrate reduces stomatal immunity. Plants were first dark-adapted for 1 hr. Detached leaves were floated on stomatal opening buffer containing mannitol, malate, or citrate (20 mM) under light for 2.5 hr, and then exposed to Pst-lux (OD600 = 0.5) for 1 hr. After washing, Pst entry was measured by photon counting (A) or by direct counting of the bacterial population (B). Values are means ± SD.**P ≤ 0.01, n = 3. (C) Pathogen-induced stomatal closure is impaired by malateor citrate. Plants were first dark-adapted for 1 hr. Detached leaves were floatedon stomatal opening buffer containing mannitol, malate, or citrate under light for2.5 hr and then exposed to Pst-lux (OD600 = 0.1) for 2 hr. Values are means± SD. **P ≤ 0.01, n ≥ 30. (D and E) Compromised stomatal immunity by malateor citrate treatment is not related to induced pathogen virulence. Pst-lux was firstsuspended in stomatal opening buffer containing mannitol, malate, or citrate for 1hr and washed twice with stomatal opening buffer. These pre-treated Pst-luxbacteria were then used for inoculation. Pst entry was measured by photoncounting (D) or by direct counting of the bacterial population (E). Values aremeans ± SD. **P ≤ 0.01, n = 3.

Figure 7. ABA signaling is essential to stomatal immunity. (A and B) Elevated pathogen entry in ABA biosynthesis and signaling mutants. Intact leaves of Col-0, aba2-1, ost1-3, slac1-3, and rbohD plants were floated on stomatal opening buffer under light for 2.5 hr and then exposed to Pst-lux (OD600 = 0.5) for 1 hr. Pst entry was measured by photon counting (A) or by direct counting of the bacterial population (B). Values are means ± SD. **P ≤ 0.01, n = 3. (C and D) Pathogen/PAMP-induced stomatal closure is dependent on ABAbiosynthesis and signaling. Intact leaves of Col-0, aba2-1, ost1-3, slac1-3, andrbohD plants were floated on stomatal opening buffer under light for 2.5 hr andthen exposed to flg22 for 3 hr (C) or Pst (OD600 = 0.1) for 2 hr (D). Values aremeans ± SD, **P ≤ 0.01, n ≥ 30.

Figure 8. MPK3/MPK6 activation-induced stomatal closure is insensitive to coronatine. (A) ABA-induced stomatal closure is sensitive to coronatine. Intact leaves of Col-0 plants were floated on stomatal opening buffer under light for 2.5 hr and thentreated with EtOH (solvent control), ABA (10 mM), or coronatine (COR, 1mg/mL). Stomatal aperture was measured 3 hr later. (B) Gain-of-functionMPK3/MPK6 activation-induced stomatal closure is insensitive to coronatine.Intact leaves of DD plants were floated on stomatal opening buffer under light for2.5 hr and then treated with EtOH (solvent control), DEX (5 mM), or coronatine(COR, 1 mg/mL). Stomatal aperture was measured 5 hr later. (C and D) Gain-of-function activation of MPK3/MPK6 abolishes pathogen-induced stomatalreopening. Intact leaves of DD plants were floated on stomatal opening bufferunder light for 2.5 hr and then treated with EtOH or DEX (5 mM) for 3 hr, whichwas followed by treatment with Pst for another 1 hr (C) or 4 hr (D). In A-D, valuesare means ± SD. *P ≤ 0.05 and **P ≤ 0.01, n ≥ 30. (E) MPK3/MPK6 cascade andABA function interdependently to regulate stomatal immunity. In this model,pathogen-responsive MPK3/MPK6 cascade and ABA are both required forstomatal immunity. Upon perception of pathogen/PAMPs, both pathways areactivated. Activation of MPK3/MPK6 by MKK4/MKK5 increases malate/citratemetabolism, which leads to a decrease in malate/citrate content. ABA signalingpathway activates OST1 that can directly phosphorylated RbohD and SLAC1,and activate SLAC1-mediated Cl- efflux, which leads to plasma depolarizationand subsequent K+ efflux. Different from MPK3/MPK6 cascade, MPK9 andMPK12, two guard cell-specific MAPKs, are involved in SLAC1 activation andstomatal closure in response to ABA and ROS. COR inhibits ABA-, but notMPK3/MPK6 cascade-mediated stomatal closure.

Parsed CitationsAcharya, B.R., Jeon, B.W., Zhang, W., and Assmann, S.M. (2013). Open Stomata 1 (OST1) is limiting in abscisic acid responses ofArabidopsis guard cells. New Phytol. 200: 1049-1063.

Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Anderson, J.C., Wan, Y., Kim, Y.M., Pasa-Tolic, L., Metz, T.O., and Peck, S.C. (2014). Decreased abundance of type III secretionsystem-inducing signals in Arabidopsis mkp1 enhances resistance against Pseudomonas syringae. Proc. Natl. Acad. Sci. USA 111:6846-6851.


Araujo, W.L., Fernie, A.R., and Nunes-Nesi, A. (2011a). Control of stomatal aperture: a renaissance of the old guard. Plant SignalBehav. 6: 1305-1311.


Araujo, W.L., Nunes-Nesi, A., Osorio, S., Usadel, B., Fuentes, D., Nagy, R., Balbo, I., Lehmann, M., Studart-Witkowski, C., Tohge, T.,et al. (2011b). Antisense inhibition of the iron-sulphur subunit of succinate dehydrogenase enhances photosynthesis and growthin tomato via an organic acid-mediated effect on stomatal aperture. Plant Cell 23: 600-627.


Arnaud, D., and Hwang, I. (2015). A sophisticated network of signaling pathways regulates stomatal defenses to bacterialpathogens. Mol. Plant 8: 566-581.


Asai, T., Tena, G., Plotnikova, J., Willmann, M.R., Chiu, W.L., Gomez-Gomez, L., Boller, T., Ausubel, F.M., and Sheen, J. (2002). MAPkinase signalling cascade in Arabidopsis innate immunity. Nature 415: 977-983.


Ausubel, F.M. (2005). Are innate immune signaling pathways in plants and animals conserved? Nat. Immunol. 6: 973-979.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Azoulay-Shemer, T., Bagheri, A., Wang, C., Palomares, A., Stephan, A.B., Kunz, H.H., and Schroeder, J.I. (2016). Starch biosynthesisin guard cells but not in mesophyll cells is involved in CO2-induced stomatal closing. Plant Physiol. 171: 788-798.


Banks, A.S., McAllister, F.E., Camporez, J.P.G., Zushin, P.-J.H., Jurczak, M.J., Laznik-Bogoslavski, D., Shulman, G.I., Gygi, S.P., andSpiegelman, B.M. (2015). An ERK/Cdk5 axis controls the diabetogenic actions of PPAR?. Nature 517: 391-395.


Bates, G.W., Rosenthal, D.M., Sun, J., Chattopadhyay, M., Peffer, E., Yang, J., Ort, D.R., and Jones, A.M. (2012). A comparative studyof the Arabidopsis thaliana guard-cell transcriptome and its modulation by sucrose. PLoS One 7: e49641.


Baud, S., and Graham, I.A. (2006). A spatiotemporal analysis of enzymatic activities associated with carbon metabolism in wild-typeand mutant embryos of Arabidopsis using in situ histochemistry. Plant J. 46: 155-169.


Beckers, G.J., Jaskiewicz, M., Liu, Y., Underwood, W.R., He, S.Y., Zhang, S., and Conrath, U. (2009). Mitogen-activated proteinkinases 3 and 6 are required for full priming of stress responses in Arabidopsis thaliana. Plant Cell 21: 944-953.


Berriri, S., Garcia, A.V., Frei dit Frey, N., Rozhon, W., Pateyron, S., Leonhardt, N., Montillet, J.L., Leung, J., Hirt, H., and Colcombet,J. (2012). Constitutively active mitogen-activated protein kinase versions reveal functions of Arabidopsis MPK4 in pathogendefense signaling. Plant Cell 24: 4281-4293.

Pubmed: Author and Title

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http://scholar.google.com/scholar?as_q=Starch biosynthesis in guard cells but not in mesophyll cells is involved in CO2-induced stomatal closing&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Azoulay-Shemer,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Banks%2C A%2ES%2E%2C McAllister%2C F%2EE%2E%2C Camporez%2C J%2EP%2EG%2E%2C Zushin%2C P%2E%2DJ%2EH%2E%2C Jurczak%2C M%2EJ%2E%2C Laznik%2DBogoslavski%2C D%2E%2C Shulman%2C G%2EI%2E%2C Gygi%2C S%2EP%2E%2C and Spiegelman%2C B%2EM%2E %282015%29%2E An ERK%2FCdk5 axis controls the diabetogenic actions of PPAR%3F%2E Nature 517%3A 391%2D395%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Banks, A.S., McAllister, F.E., Camporez, J.P.G., Zushin, P.-J.H., Jurczak, M.J., Laznik-Bogoslavski, D., Shulman, G.I., Gygi, S.P., and Spiegelman, B.M. (2015). An ERK/Cdk5 axis controls the diabetogenic actions of PPAR?. Nature 517: 391-395.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Banks,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=An ERK/Cdk5 axis controls the diabetogenic actions of PPAR&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=An ERK/Cdk5 axis controls the diabetogenic actions of PPAR&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Banks,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Bates%2C G%2EW%2E%2C Rosenthal%2C D%2EM%2E%2C Sun%2C J%2E%2C Chattopadhyay%2C M%2E%2C Peffer%2C E%2E%2C Yang%2C J%2E%2C Ort%2C D%2ER%2E%2C and Jones%2C A%2EM%2E %282012%29%2E A comparative study of the Arabidopsis thaliana guard%2Dcell transcriptome and its modulation by sucrose%2E PLoS One 7%3A e49641%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Bates, G.W., Rosenthal, D.M., Sun, J., Chattopadhyay, M., Peffer, E., Yang, J., Ort, D.R., and Jones, A.M. (2012). A comparative study of the Arabidopsis thaliana guard-cell transcriptome and its modulation by sucrose. PLoS One 7: e49641.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Bates,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A comparative study of the Arabidopsis thaliana guard-cell transcriptome and its modulation by sucrose&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A comparative study of the Arabidopsis thaliana guard-cell transcriptome and its modulation by sucrose&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bates,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Baud%2C S%2E%2C and Graham%2C I%2EA%2E %282006%29%2E A spatiotemporal analysis of enzymatic activities associated with carbon metabolism in wild%2Dtype and mutant embryos of Arabidopsis using in situ histochemistry%2E Plant J%2E 46%3A 155%2D169%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Baud, S., and Graham, I.A. (2006). A spatiotemporal analysis of enzymatic activities associated with carbon metabolism in wild-type and mutant embryos of Arabidopsis using in situ histochemistry. Plant J. 46: 155-169.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Baud,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A spatiotemporal analysis of enzymatic activities associated with carbon metabolism in wild-type and mutant embryos of Arabidopsis using in situ histochemistry&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A spatiotemporal analysis of enzymatic activities associated with carbon metabolism in wild-type and mutant embryos of Arabidopsis using in situ histochemistry&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Baud,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Beckers%2C G%2EJ%2E%2C Jaskiewicz%2C M%2E%2C Liu%2C Y%2E%2C Underwood%2C W%2ER%2E%2C He%2C S%2EY%2E%2C Zhang%2C S%2E%2C and Conrath%2C U%2E %282009%29%2E Mitogen%2Dactivated protein kinases 3 and 6 are required for full priming of stress responses in Arabidopsis thaliana%2E Plant Cell 21%3A 944%2D953%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Beckers, G.J., Jaskiewicz, M., Liu, Y., Underwood, W.R., He, S.Y., Zhang, S., and Conrath, U. (2009). Mitogen-activated protein kinases 3 and 6 are required for full priming of stress responses in Arabidopsis thaliana. Plant Cell 21: 944-953.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Beckers,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Mitogen-activated protein kinases 3 and 6 are required for full priming of stress responses in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Mitogen-activated protein kinases 3 and 6 are required for full priming of stress responses in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Beckers,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Berriri%2C S%2E%2C Garcia%2C A%2EV%2E%2C Frei dit Frey%2C N%2E%2C Rozhon%2C W%2E%2C Pateyron%2C S%2E%2C Leonhardt%2C N%2E%2C Montillet%2C J%2EL%2E%2C Leung%2C J%2E%2C Hirt%2C H%2E%2C and Colcombet%2C J%2E %282012%29%2E Constitutively active mitogen%2Dactivated protein kinase versions reveal functions of Arabidopsis MPK4 in pathogen defense signaling%2E Plant Cell 24%3A 4281%2D4293%2E&dopt=abstract

CrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Blatt, M.R. (2016). Plant Physiology: Redefining the enigma of metabolism in stomatal movement. Curr. Biol. 26: R107-109.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Bohm, H., Albert, I., Fan, L., Reinhard, A., and Nurnberger, T. (2014). Immune receptor complexes at the plant cell surface. Curr.Opin. Plant Biol. 20: 47-54.


Boller, T., and Felix, G. (2009). A renaissance of elicitors: perception of microbe-associated molecular patterns and danger signalsby pattern-recognition receptors. Annu. Rev. Plant Biol. 60: 379-406.


Brandt, B., Munemasa, S., Wang, C., Nguyen, D., Yong, T., Yang, P.G., Poretsky, E., Belknap, T.F., Waadt, R., Aleman, F., et al.(2015). Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells. Elife 4: e03599.


Brooks, D.M., Hernandez-Guzman, G., Kloek, A.P., Alarcon-Chaidez, F., Sreedharan, A., Rangaswamy, V., Penaloza-Vazquez, A.,Bender, C.L., and Kunkel, B.N. (2004). Identification and characterization of a well-defined series of coronatine biosyntheticmutants of Pseudomonas syringae pv. tomato DC3000. Mol. Plant Microbe Interact. 17: 162-174.


Caspi, R., Billington, R., Ferrer, L., Foerster, H., Fulcher, C.A., Keseler, I.M., Kothari, A., Krummenacker, M., Latendresse, M.,Mueller, L.A., et al. (2016). The MetaCyc database of metabolic pathways and enzymes and the BioCyc collection ofpathway/genome databases. Nucleic Acids Res. 44: D471-480.


Chater, C.C., Oliver, J., Casson, S., and Gray, J.E. (2014). Putting the brakes on: abscisic acid as a central environmental regulatorof stomatal development. New Phytol. 202: 376-391.


Chisholm, S.T., Coaker, G., Day, B., and Staskawicz, B.J. (2006). Host-microbe interactions: shaping the evolution of the plantimmune response. Cell 124: 803-814.


Choi, J.H., Banks, A.S., Estall, J.L., Kajimura, S., Bostrom, P., Laznik, D., Ruas, J.L., Chalmers, M.J., Kamenecka, T.M., Bluher, M., etal. (2010). Anti-diabetic drugs inhibit obesity-linked phosphorylation of PPAR? by Cdk5. Nature 466: 451-456.


Cuppels, D.A., and Ainsworth, T. (1995). Molecular and physiological characterization of Pseudomonas syringae pv. tomato andPseudomonas syringae pv. maculicola strains that produce the phytotoxin coronatine. Appl. Environ. Microbiol. 61: 3530-3536.


Daloso, D.M., Antunes, W.C., Pinheiro, D.P., Waquim, J.P., Araujo, W.L., Loureiro, M.E., Fernie, A.R., and Williams, T.C. (2015).Tobacco guard cells fix CO2 by both Rubisco and PEPcase while sucrose acts as a substrate during light-induced stomatalopening. Plant Cell Environ. 38: 2353-2371.


Doehlemann, G., and Hemetsberger, C. (2013). Apoplastic immunity and its suppression by filamentous plant pathogens. NewPhytol. 198: 1001-1016.


Du, M., Zhai, Q., Deng, L., Li, S., Li, H., Yan, L., Huang, Z., Wang, B., Jiang, H., Huang, T., et al. (2014). Closely related NACtranscription factors of tomato differentially regulate stomatal closure and reopening during pathogen attack. Plant Cell 26: 3167-3184.

http://search.crossref.org/?page=1&rows=2&q=Berriri, S., Garcia, A.V., Frei dit Frey, N., Rozhon, W., Pateyron, S., Leonhardt, N., Montillet, J.L., Leung, J., Hirt, H., and Colcombet, J. (2012). Constitutively active mitogen-activated protein kinase versions reveal functions of Arabidopsis MPK4 in pathogen defense signaling. Plant Cell 24: 4281-4293.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Berriri,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Constitutively active mitogen-activated protein kinase versions reveal functions of Arabidopsis MPK4 in pathogen defense signaling&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Constitutively active mitogen-activated protein kinase versions reveal functions of Arabidopsis MPK4 in pathogen defense signaling&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Berriri,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Blatt%2C M%2ER%2E %282016%29%2E Plant Physiology%3A Redefining the enigma of metabolism in stomatal movement%2E Curr%2E Biol%2E 26%3A R107%2D109%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Blatt, M.R. (2016). Plant Physiology: Redefining the enigma of metabolism in stomatal movement. Curr. Biol. 26: R107-109.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Blatt,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Plant Physiology: Redefining the enigma of metabolism in stomatal movement&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Plant Physiology: Redefining the enigma of metabolism in stomatal movement&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Blatt,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Bohm%2C H%2E%2C Albert%2C I%2E%2C Fan%2C L%2E%2C Reinhard%2C A%2E%2C and Nurnberger%2C T%2E %282014%29%2E Immune receptor complexes at the plant cell surface%2E Curr%2E Opin%2E Plant Biol%2E 20%3A 47%2D54%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Bohm, H., Albert, I., Fan, L., Reinhard, A., and Nurnberger, T. (2014). Immune receptor complexes at the plant cell surface. Curr. Opin. Plant Biol. 20: 47-54.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Bohm,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Immune receptor complexes at the plant cell surface&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Immune receptor complexes at the plant cell surface&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bohm,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Boller%2C T%2E%2C and Felix%2C G%2E %282009%29%2E A renaissance of elicitors%3A perception of microbe%2Dassociated molecular patterns and danger signals by pattern%2Drecognition receptors%2E Annu%2E Rev%2E Plant Biol%2E 60%3A 379%2D406%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Boller, T., and Felix, G. (2009). A renaissance of elicitors: perception of microbe-associated molecular patterns and danger signals by pattern-recognition receptors. Annu. Rev. Plant Biol. 60: 379-406.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Boller,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A renaissance of elicitors: perception of microbe-associated molecular patterns and danger signals by pattern-recognition receptors&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A renaissance of elicitors: perception of microbe-associated molecular patterns and danger signals by pattern-recognition receptors&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Boller,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Brandt%2C B%2E%2C Munemasa%2C S%2E%2C Wang%2C C%2E%2C Nguyen%2C D%2E%2C Yong%2C T%2E%2C Yang%2C P%2EG%2E%2C Poretsky%2C E%2E%2C Belknap%2C T%2EF%2E%2C Waadt%2C R%2E%2C Aleman%2C F%2E%2C et al%2E %282015%29%2E Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells%2E Elife 4%3A e03599%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Brandt, B., Munemasa, S., Wang, C., Nguyen, D., Yong, T., Yang, P.G., Poretsky, E., Belknap, T.F., Waadt, R., Aleman, F., et al. (2015). Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells. Elife 4: e03599.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Brandt,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Calcium specificity signaling mechanisms in abscisic acid signal transduction in Arabidopsis guard cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Brandt,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Brooks%2C D%2EM%2E%2C Hernandez%2DGuzman%2C G%2E%2C Kloek%2C A%2EP%2E%2C Alarcon%2DChaidez%2C F%2E%2C Sreedharan%2C A%2E%2C Rangaswamy%2C V%2E%2C Penaloza%2DVazquez%2C A%2E%2C Bender%2C C%2EL%2E%2C and Kunkel%2C B%2EN%2E %282004%29%2E Identification and characterization of a well%2Ddefined series of coronatine biosynthetic mutants of Pseudomonas syringae pv%2E tomato DC3000%2E Mol%2E Plant Microbe Interact%2E 17%3A 162%2D174%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Brooks, D.M., Hernandez-Guzman, G., Kloek, A.P., Alarcon-Chaidez, F., Sreedharan, A., Rangaswamy, V., Penaloza-Vazquez, A., Bender, C.L., and Kunkel, B.N. (2004). Identification and characterization of a well-defined series of coronatine biosynthetic mutants of Pseudomonas syringae pv. tomato DC3000. Mol. Plant Microbe Interact. 17: 162-174.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Brooks,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Identification and characterization of a well-defined series of coronatine biosynthetic mutants of Pseudomonas syringae pv&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Identification and characterization of a well-defined series of coronatine biosynthetic mutants of Pseudomonas syringae pv&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Brooks,&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Caspi%2C R%2E%2C Billington%2C R%2E%2C Ferrer%2C L%2E%2C Foerster%2C H%2E%2C Fulcher%2C C%2EA%2E%2C Keseler%2C I%2EM%2E%2C Kothari%2C A%2E%2C Krummenacker%2C M%2E%2C Latendresse%2C M%2E%2C Mueller%2C L%2EA%2E%2C et al%2E %282016%29%2E The MetaCyc database of metabolic pathways and enzymes and the BioCyc collection of pathway%2Fgenome databases%2E Nucleic Acids Res%2E 44%3A D471%2D480%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Caspi, R., Billington, R., Ferrer, L., Foerster, H., Fulcher, C.A., Keseler, I.M., Kothari, A., Krummenacker, M., Latendresse, M., Mueller, L.A., et al. (2016). The MetaCyc database of metabolic pathways and enzymes and the BioCyc collection of pathway/genome databases. Nucleic Acids Res. 44: D471-480.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Caspi,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The MetaCyc database of metabolic pathways and enzymes and the BioCyc collection of pathway/genome databases&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The MetaCyc database of metabolic pathways and enzymes and the BioCyc collection of pathway/genome databases&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Caspi,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Chater%2C C%2EC%2E%2C Oliver%2C J%2E%2C Casson%2C S%2E%2C and Gray%2C J%2EE%2E %282014%29%2E Putting the brakes on%3A abscisic acid as a central environmental regulator of stomatal development%2E New Phytol%2E 202%3A 376%2D391%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Chater, C.C., Oliver, J., Casson, S., and Gray, J.E. (2014). Putting the brakes on: abscisic acid as a central environmental regulator of stomatal development. New Phytol. 202: 376-391.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Chater,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Putting the brakes on: abscisic acid as a central environmental regulator of stomatal development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Putting the brakes on: abscisic acid as a central environmental regulator of stomatal development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chater,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Chisholm%2C S%2ET%2E%2C Coaker%2C G%2E%2C Day%2C B%2E%2C and Staskawicz%2C B%2EJ%2E %282006%29%2E Host%2Dmicrobe interactions%3A shaping the evolution of the plant immune response%2E Cell 124%3A 803%2D814%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Chisholm, S.T., Coaker, G., Day, B., and Staskawicz, B.J. (2006). Host-microbe interactions: shaping the evolution of the plant immune response. Cell 124: 803-814.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Chisholm,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Host-microbe interactions: shaping the evolution of the plant immune response&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Host-microbe interactions: shaping the evolution of the plant immune response&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chisholm,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Choi%2C J%2EH%2E%2C Banks%2C A%2ES%2E%2C Estall%2C J%2EL%2E%2C Kajimura%2C S%2E%2C Bostrom%2C P%2E%2C Laznik%2C D%2E%2C Ruas%2C J%2EL%2E%2C Chalmers%2C M%2EJ%2E%2C Kamenecka%2C T%2EM%2E%2C Bluher%2C M%2E%2C et al%2E %282010%29%2E Anti%2Ddiabetic drugs inhibit obesity%2Dlinked phosphorylation of PPAR%3F by Cdk5%2E Nature 466%3A 451%2D456%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Choi, J.H., Banks, A.S., Estall, J.L., Kajimura, S., Bostrom, P., Laznik, D., Ruas, J.L., Chalmers, M.J., Kamenecka, T.M., Bluher, M., et al. (2010). Anti-diabetic drugs inhibit obesity-linked phosphorylation of PPAR? by Cdk5. Nature 466: 451-456.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Choi,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Anti-diabetic drugs inhibit obesity-linked phosphorylation of PPAR&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Anti-diabetic drugs inhibit obesity-linked phosphorylation of PPAR&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Choi,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Cuppels%2C D%2EA%2E%2C and Ainsworth%2C T%2E %281995%29%2E Molecular and physiological characterization of Pseudomonas syringae pv%2E tomato and Pseudomonas syringae pv%2E maculicola strains that produce the phytotoxin coronatine%2E Appl%2E Environ%2E Microbiol%2E 61%3A 3530%2D3536%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Cuppels, D.A., and Ainsworth, T. (1995). Molecular and physiological characterization of Pseudomonas syringae pv. tomato and Pseudomonas syringae pv. maculicola strains that produce the phytotoxin coronatine. Appl. Environ. Microbiol. 61: 3530-3536.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Cuppels,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Molecular and physiological characterization of Pseudomonas syringae pv&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Molecular and physiological characterization of Pseudomonas syringae pv&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Cuppels,&as_ylo=1995&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Daloso%2C D%2EM%2E%2C Antunes%2C W%2EC%2E%2C Pinheiro%2C D%2EP%2E%2C Waquim%2C J%2EP%2E%2C Araujo%2C W%2EL%2E%2C Loureiro%2C M%2EE%2E%2C Fernie%2C A%2ER%2E%2C and Williams%2C T%2EC%2E %282015%29%2E Tobacco guard cells fix CO2 by both Rubisco and PEPcase while sucrose acts as a substrate during light%2Dinduced stomatal opening%2E Plant Cell Environ%2E 38%3A 2353%2D2371%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Daloso, D.M., Antunes, W.C., Pinheiro, D.P., Waquim, J.P., Araujo, W.L., Loureiro, M.E., Fernie, A.R., and Williams, T.C. (2015). Tobacco guard cells fix CO2 by both Rubisco and PEPcase while sucrose acts as a substrate during light-induced stomatal opening. Plant Cell Environ. 38: 2353-2371.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Daloso,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Tobacco guard cells fix CO2 by both Rubisco and PEPcase while sucrose acts as a substrate during light-induced stomatal opening&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Tobacco guard cells fix CO2 by both Rubisco and PEPcase while sucrose acts as a substrate during light-induced stomatal opening&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Daloso,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Doehlemann%2C G%2E%2C and Hemetsberger%2C C%2E %282013%29%2E Apoplastic immunity and its suppression by filamentous plant pathogens%2E New Phytol%2E 198%3A 1001%2D1016%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Doehlemann, G., and Hemetsberger, C. (2013). Apoplastic immunity and its suppression by filamentous plant pathogens. New Phytol. 198: 1001-1016.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Doehlemann,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Apoplastic immunity and its suppression by filamentous plant pathogens&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Apoplastic immunity and its suppression by filamentous plant pathogens&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Doehlemann,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1


Fan, J., Crooks, C., and Lamb, C. (2008). High-throughput quantitative luminescence assay of the growth in planta of Pseudomonassyringae chromosomally tagged with Photorhabdus luminescens luxCDABE. Plant J. 53: 393-399.


Fritzius, T., Aeschbacher, R., Wiemken, A., and Wingler, A. (2001). Induction of ApL3 expression by trehalose complements thestarch-deficient Arabidopsis mutant adg2-1 lacking ApL1, the large subunit of ADP-glucose pyrophosphorylase. Plant Physiol. 126:883-889.


Geiger, D., Scherzer, S., Mumm, P., Stange, A., Marten, I., Bauer, H., Ache, P., Matschi, S., Liese, A., Al-Rasheid, K.A., et al. (2009).Activity of guard cell anion channel SLAC1 is controlled by drought-stress signaling kinase-phosphatase pair. Proc. Natl. Acad. Sci.USA 106: 21425-21430.


Gilliham, M., and Tyerman, S.D. (2016). Linking metabolism to membrane signaling: The GABA-malate connection. Trends Plant Sci.21: 295-301.


Gonzalez-Guzman, M., Apostolova, N., Belles, J.M., Barrero, J.M., Piqueras, P., Ponce, M.R., Micol, J.L., Serrano, R., andRodriguez, P.L. (2002). The short-chain alcohol dehydrogenase ABA2 catalyzes the conversion of xanthoxin to abscisic aldehyde.Plant Cell 14: 1833-1846.


Grimmer, M.K., John Foulkes, M., and Paveley, N.D. (2012). Foliar pathogenesis and plant water relations: a review. J. Exp. Bot. 63:4321-4331.


Guan, R., Su, J., Meng, X., Li, S., Liu, Y., Xu, J., and Zhang, S. (2015). Multilayered regulation of ethylene induction plays a positiverole in Arabidopsis resistance against Pseudomonas syringae. Plant Physiol. 169: 299-312.


Gudesblat, G.E., Torres, P.S., and Vojnov, A.A. (2009). Xanthomonas campestris overcomes Arabidopsis stomatal innate immunitythrough a DSF cell-to-cell signal-regulated virulence factor. Plant Physiol. 149: 1017-1027.


Guimaraes, R.L., and Stotz, H.U. (2004). Oxalate production by Sclerotinia sclerotiorum deregulates guard cells during infection.Plant Physiol. 136: 3703-3711.


Guzel Deger, A., Scherzer, S., Nuhkat, M., Kedzierska, J., Kollist, H., Brosche, M., Unyayar, S., Boudsocq, M., Hedrich, R., andRoelfsema, M.R. (2015). Guard cell SLAC1-type anion channels mediate flagellin-induced stomatal closure. New Phytol. 208: 162-173.


Hetherington, A.M., and Woodward, F.I. (2003). The role of stomata in sensing and driving environmental change. Nature 424: 901-908.


Horrer, D., Flutsch, S., Pazmino, D., Matthews, J.S., Thalmann, M., Nigro, A., Leonhardt, N., Lawson, T., and Santelia, D. (2016). Bluelight induces a distinct starch degradation pathway in guard cells for stomatal opening. Curr. Biol. 26: 362-370.


http://www.ncbi.nlm.nih.gov/pubmed?term=Du%2C M%2E%2C Zhai%2C Q%2E%2C Deng%2C L%2E%2C Li%2C S%2E%2C Li%2C H%2E%2C Yan%2C L%2E%2C Huang%2C Z%2E%2C Wang%2C B%2E%2C Jiang%2C H%2E%2C Huang%2C T%2E%2C et al%2E %282014%29%2E Closely related NAC transcription factors of tomato differentially regulate stomatal closure and reopening during pathogen attack%2E Plant Cell 26%3A 3167%2D3184%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Du, M., Zhai, Q., Deng, L., Li, S., Li, H., Yan, L., Huang, Z., Wang, B., Jiang, H., Huang, T., et al. (2014). Closely related NAC transcription factors of tomato differentially regulate stomatal closure and reopening during pathogen attack. Plant Cell 26: 3167-3184.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Du,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Closely related NAC transcription factors of tomato differentially regulate stomatal closure and reopening during pathogen attack&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Closely related NAC transcription factors of tomato differentially regulate stomatal closure and reopening during pathogen attack&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Du,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Fan%2C J%2E%2C Crooks%2C C%2E%2C and Lamb%2C C%2E %282008%29%2E High%2Dthroughput quantitative luminescence assay of the growth in planta of Pseudomonas syringae chromosomally tagged with Photorhabdus luminescens luxCDABE%2E Plant J%2E 53%3A 393%2D399%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Fan, J., Crooks, C., and Lamb, C. (2008). High-throughput quantitative luminescence assay of the growth in planta of Pseudomonas syringae chromosomally tagged with Photorhabdus luminescens luxCDABE. Plant J. 53: 393-399.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Fan,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=High-throughput quantitative luminescence assay of the growth in planta of Pseudomonas syringae chromosomally tagged with Photorhabdus luminescens luxCDABE&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=High-throughput quantitative luminescence assay of the growth in planta of Pseudomonas syringae chromosomally tagged with Photorhabdus luminescens luxCDABE&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Fan,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Fritzius%2C T%2E%2C Aeschbacher%2C R%2E%2C Wiemken%2C A%2E%2C and Wingler%2C A%2E %282001%29%2E Induction of ApL3 expression by trehalose complements the starch%2Ddeficient Arabidopsis mutant adg2%2D1 lacking ApL1%2C the large subunit of ADP%2Dglucose pyrophosphorylase%2E Plant Physiol%2E 126%3A 883%2D889%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Fritzius, T., Aeschbacher, R., Wiemken, A., and Wingler, A. (2001). Induction of ApL3 expression by trehalose complements the starch-deficient Arabidopsis mutant adg2-1 lacking ApL1, the large subunit of ADP-glucose pyrophosphorylase. Plant Physiol. 126: 883-889.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Fritzius,&hl=en&c2coff=1


http://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Fritzius,&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Geiger%2C D%2E%2C Scherzer%2C S%2E%2C Mumm%2C P%2E%2C Stange%2C A%2E%2C Marten%2C I%2E%2C Bauer%2C H%2E%2C Ache%2C P%2E%2C Matschi%2C S%2E%2C Liese%2C A%2E%2C Al%2DRasheid%2C K%2EA%2E%2C et al%2E %282009%29%2E Activity of guard cell anion channel SLAC1 is controlled by drought%2Dstress signaling kinase%2Dphosphatase pair%2E Proc%2E Natl%2E Acad%2E Sci%2E USA 106%3A 21425%2D21430%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Geiger, D., Scherzer, S., Mumm, P., Stange, A., Marten, I., Bauer, H., Ache, P., Matschi, S., Liese, A., Al-Rasheid, K.A., et al. (2009). Activity of guard cell anion channel SLAC1 is controlled by drought-stress signaling kinase-phosphatase pair. Proc. Natl. Acad. Sci. USA 106: 21425-21430.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Geiger,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Activity of guard cell anion channel SLAC1 is controlled by drought-stress signaling kinase-phosphatase pair&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Activity of guard cell anion channel SLAC1 is controlled by drought-stress signaling kinase-phosphatase pair&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Geiger,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Gilliham%2C M%2E%2C and Tyerman%2C S%2ED%2E %282016%29%2E Linking metabolism to membrane signaling%3A The GABA%2Dmalate connection%2E Trends Plant Sci%2E 21%3A 295%2D301%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Gilliham, M., and Tyerman, S.D. (2016). Linking metabolism to membrane signaling: The GABA-malate connection. Trends Plant Sci. 21: 295-301.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Gilliham,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Linking metabolism to membrane signaling: The GABA-malate connection&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Linking metabolism to membrane signaling: The GABA-malate connection&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gilliham,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Gonzalez%2DGuzman%2C M%2E%2C Apostolova%2C N%2E%2C Belles%2C J%2EM%2E%2C Barrero%2C J%2EM%2E%2C Piqueras%2C P%2E%2C Ponce%2C M%2ER%2E%2C Micol%2C J%2EL%2E%2C Serrano%2C R%2E%2C and Rodriguez%2C P%2EL%2E %282002%29%2E The short%2Dchain alcohol dehydrogenase ABA2 catalyzes the conversion of xanthoxin to abscisic aldehyde%2E Plant Cell 14%3A 1833%2D1846%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Gonzalez-Guzman, M., Apostolova, N., Belles, J.M., Barrero, J.M., Piqueras, P., Ponce, M.R., Micol, J.L., Serrano, R., and Rodriguez, P.L. (2002). The short-chain alcohol dehydrogenase ABA2 catalyzes the conversion of xanthoxin to abscisic aldehyde. Plant Cell 14: 1833-1846.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Gonzalez-Guzman,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The short-chain alcohol dehydrogenase ABA2 catalyzes the conversion of xanthoxin to abscisic aldehyde&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The short-chain alcohol dehydrogenase ABA2 catalyzes the conversion of xanthoxin to abscisic aldehyde&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gonzalez-Guzman,&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Grimmer%2C M%2EK%2E%2C John Foulkes%2C M%2E%2C and Paveley%2C N%2ED%2E %282012%29%2E Foliar pathogenesis and plant water relations%3A a review%2E J%2E Exp%2E Bot%2E 63%3A 4321%2D4331%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Grimmer, M.K., John Foulkes, M., and Paveley, N.D. (2012). Foliar pathogenesis and plant water relations: a review. J. Exp. Bot. 63: 4321-4331.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Grimmer,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Foliar pathogenesis and plant water relations: a review&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Foliar pathogenesis and plant water relations: a review&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Grimmer,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Guan%2C R%2E%2C Su%2C J%2E%2C Meng%2C X%2E%2C Li%2C S%2E%2C Liu%2C Y%2E%2C Xu%2C J%2E%2C and Zhang%2C S%2E %282015%29%2E Multilayered regulation of ethylene induction plays a positive role in Arabidopsis resistance against Pseudomonas syringae%2E Plant Physiol%2E 169%3A 299%2D312%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Guan, R., Su, J., Meng, X., Li, S., Liu, Y., Xu, J., and Zhang, S. (2015). Multilayered regulation of ethylene induction plays a positive role in Arabidopsis resistance against Pseudomonas syringae. Plant Physiol. 169: 299-312.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Guan,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Multilayered regulation of ethylene induction plays a positive role in Arabidopsis resistance against Pseudomonas syringae&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Multilayered regulation of ethylene induction plays a positive role in Arabidopsis resistance against Pseudomonas syringae&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Guan,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Gudesblat%2C G%2EE%2E%2C Torres%2C P%2ES%2E%2C and Vojnov%2C A%2EA%2E %282009%29%2E Xanthomonas campestris overcomes Arabidopsis stomatal innate immunity through a DSF cell%2Dto%2Dcell signal%2Dregulated virulence factor%2E Plant Physiol%2E 149%3A 1017%2D1027%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Gudesblat, G.E., Torres, P.S., and Vojnov, A.A. (2009). Xanthomonas campestris overcomes Arabidopsis stomatal innate immunity through a DSF cell-to-cell signal-regulated virulence factor. Plant Physiol. 149: 1017-1027.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Gudesblat,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Xanthomonas campestris overcomes Arabidopsis stomatal innate immunity through a DSF cell-to-cell signal-regulated virulence factor&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Xanthomonas campestris overcomes Arabidopsis stomatal innate immunity through a DSF cell-to-cell signal-regulated virulence factor&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gudesblat,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Guimaraes%2C R%2EL%2E%2C and Stotz%2C H%2EU%2E %282004%29%2E Oxalate production by Sclerotinia sclerotiorum deregulates guard cells during infection%2E Plant Physiol%2E 136%3A 3703%2D3711%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Guimaraes, R.L., and Stotz, H.U. (2004). Oxalate production by Sclerotinia sclerotiorum deregulates guard cells during infection. Plant Physiol. 136: 3703-3711.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Guimaraes,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Oxalate production by Sclerotinia sclerotiorum deregulates guard cells during infection&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Oxalate production by Sclerotinia sclerotiorum deregulates guard cells during infection&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Guimaraes,&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Guzel Deger%2C A%2E%2C Scherzer%2C S%2E%2C Nuhkat%2C M%2E%2C Kedzierska%2C J%2E%2C Kollist%2C H%2E%2C Brosche%2C M%2E%2C Unyayar%2C S%2E%2C Boudsocq%2C M%2E%2C Hedrich%2C R%2E%2C and Roelfsema%2C M%2ER%2E %282015%29%2E Guard cell SLAC1%2Dtype anion channels mediate flagellin%2Dinduced stomatal closure%2E New Phytol%2E 208%3A 162%2D173%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Guzel Deger, A., Scherzer, S., Nuhkat, M., Kedzierska, J., Kollist, H., Brosche, M., Unyayar, S., Boudsocq, M., Hedrich, R., and Roelfsema, M.R. (2015). Guard cell SLAC1-type anion channels mediate flagellin-induced stomatal closure. New Phytol. 208: 162-173.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Guzel&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Guard cell SLAC1-type anion channels mediate flagellin-induced stomatal closure&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Guard cell SLAC1-type anion channels mediate flagellin-induced stomatal closure&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Guzel&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Hetherington%2C A%2EM%2E%2C and Woodward%2C F%2EI%2E %282003%29%2E The role of stomata in sensing and driving environmental change%2E Nature 424%3A 901%2D908%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Hetherington, A.M., and Woodward, F.I. (2003). The role of stomata in sensing and driving environmental change. Nature 424: 901-908.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Hetherington,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The role of stomata in sensing and driving environmental change&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The role of stomata in sensing and driving environmental change&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hetherington,&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Horrer%2C D%2E%2C Flutsch%2C S%2E%2C Pazmino%2C D%2E%2C Matthews%2C J%2ES%2E%2C Thalmann%2C M%2E%2C Nigro%2C A%2E%2C Leonhardt%2C N%2E%2C Lawson%2C T%2E%2C and Santelia%2C D%2E %282016%29%2E Blue light induces a distinct starch degradation pathway in guard cells for stomatal opening%2E Curr%2E Biol%2E 26%3A 362%2D370%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Horrer, D., Flutsch, S., Pazmino, D., Matthews, J.S., Thalmann, M., Nigro, A., Leonhardt, N., Lawson, T., and Santelia, D. (2016). Blue light induces a distinct starch degradation pathway in guard cells for stomatal opening. Curr. Biol. 26: 362-370.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Horrer,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Blue light induces a distinct starch degradation pathway in guard cells for stomatal opening&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Blue light induces a distinct starch degradation pathway in guard cells for stomatal opening&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Horrer,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

Hosy, E., Vavasseur, A., Mouline, K., Dreyer, I., Gaymard, F., Poree, F., Boucherez, J., Lebaudy, A., Bouchez, D., Very, A.A., et al.(2003). The Arabidopsis outward K+ channel GORK is involved in regulation of stomatal movements and plant transpiration. Proc.Natl. Acad. Sci. USA 100: 5549-5554.


Hua, D., Wang, C., He, J., Liao, H., Duan, Y., Zhu, Z., Guo, Y., Chen, Z., and Gong, Z. (2012). A plasma membrane receptor kinase,GHR1, mediates abscisic acid- and hydrogen peroxide-regulated stomatal movement in Arabidopsis. Plant Cell 24: 2546-2561.


Hurley, B., Lee, D., Mott, A., Wilton, M., Liu, J., Liu, Y.C., Angers, S., Coaker, G., Guttman, D.S., and Desveaux, D. (2014). ThePseudomonas syringae type III effector HopF2 suppresses Arabidopsis stomatal immunity. PLoS One 9: e114921.


Jammes, F., Yang, X., Xiao, S., and Kwak, J.M. (2011). Two Arabidopsis guard cell-preferential MAPK genes, MPK9 and MPK12,function in biotic stress response. Plant Signal Behav. 6: 1875-1877.


Jammes, F., Song, C., Shin, D., Munemasa, S., Takeda, K., Gu, D., Cho, D., Lee, S., Giordo, R., Sritubtim, S., et al. (2009). MAPkinases MPK9 and MPK12 are preferentially expressed in guard cells and positively regulate ROS-mediated ABA signaling. Proc.Natl. Acad. Sci. USA 106: 20520-20525.


Jinno, N.F., M. (1981). Effects of calcium-complexing agents, sodium oxalate and soduim citrate on stomatal opening and closing ofCommelina communis L. Science Bulletin of the Faculty of Education, Nagasaki University 32: 83-86.


Johansson, F., Sommarin, M., and Larsson, C. (1993). Fusicoccin activates the plasma membrane H+-ATPase by a mechanisminvolving the C-terminal inhibitory domain. Plant Cell 5: 321-327.


Jones, J.D., and Dangl, J.L. (2006). The plant immune system. Nature 444: 323-329.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Kadota, Y., Sklenar, J., Derbyshire, P., Stransfeld, L., Asai, S., Ntoukakis, V., Jones, J.D., Shirasu, K., Menke, F., Jones, A., et al.(2014). Direct regulation of the NADPH oxidase RBOHD by the PRR-associated kinase BIK1 during plant immunity. Mol. Cell 54: 43-55.


Kelly, G., Moshelion, M., David-Schwartz, R., Halperin, O., Wallach, R., Attia, Z., Belausov, E., and Granot, D. (2013). Hexokinasemediates stomatal closure. Plant J. 75: 977-988.


Kim, T.H., Bohmer, M., Hu, H., Nishimura, N., and Schroeder, J.I. (2010). Guard cell signal transduction network: advances inunderstanding abscisic acid, CO2, and Ca2+ signaling. Annu. Rev. Plant Biol. 61: 561-591.


Kim, T.H., Hauser, F., Ha, T., Xue, S., Bohmer, M., Nishimura, N., Munemasa, S., Hubbard, K., Peine, N., Lee, B.H., et al. (2011).Chemical genetics reveals negative regulation of abscisic acid signaling by a plant immune response pathway. Curr. Biol. 21: 990-997.


Lawson, T. (2009). Guard cell photosynthesis and stomatal function. New Phytol. 181: 13-34.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

http://www.ncbi.nlm.nih.gov/pubmed?term=Hosy%2C E%2E%2C Vavasseur%2C A%2E%2C Mouline%2C K%2E%2C Dreyer%2C I%2E%2C Gaymard%2C F%2E%2C Poree%2C F%2E%2C Boucherez%2C J%2E%2C Lebaudy%2C A%2E%2C Bouchez%2C D%2E%2C Very%2C A%2EA%2E%2C et al%2E %282003%29%2E The Arabidopsis outward K%2B channel GORK is involved in regulation of stomatal movements and plant transpiration%2E Proc%2E Natl%2E Acad%2E Sci%2E USA 100%3A 5549%2D5554%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Hosy, E., Vavasseur, A., Mouline, K., Dreyer, I., Gaymard, F., Poree, F., Boucherez, J., Lebaudy, A., Bouchez, D., Very, A.A., et al. (2003). The Arabidopsis outward K+ channel GORK is involved in regulation of stomatal movements and plant transpiration. Proc. Natl. Acad. Sci. USA 100: 5549-5554.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Hosy,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The Arabidopsis outward K+ channel GORK is involved in regulation of stomatal movements and plant transpiration&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The Arabidopsis outward K+ channel GORK is involved in regulation of stomatal movements and plant transpiration&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hosy,&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Hua%2C D%2E%2C Wang%2C C%2E%2C He%2C J%2E%2C Liao%2C H%2E%2C Duan%2C Y%2E%2C Zhu%2C Z%2E%2C Guo%2C Y%2E%2C Chen%2C Z%2E%2C and Gong%2C Z%2E %282012%29%2E A plasma membrane receptor kinase%2C GHR1%2C mediates abscisic acid%2D and hydrogen peroxide%2Dregulated stomatal movement in Arabidopsis%2E Plant Cell 24%3A 2546%2D2561%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Hua, D., Wang, C., He, J., Liao, H., Duan, Y., Zhu, Z., Guo, Y., Chen, Z., and Gong, Z. (2012). A plasma membrane receptor kinase, GHR1, mediates abscisic acid- and hydrogen peroxide-regulated stomatal movement in Arabidopsis. Plant Cell 24: 2546-2561.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Hua,&hl=en&c2coff=1


http://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hua,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Hurley%2C B%2E%2C Lee%2C D%2E%2C Mott%2C A%2E%2C Wilton%2C M%2E%2C Liu%2C J%2E%2C Liu%2C Y%2EC%2E%2C Angers%2C S%2E%2C Coaker%2C G%2E%2C Guttman%2C D%2ES%2E%2C and Desveaux%2C D%2E %282014%29%2E The Pseudomonas syringae type III effector HopF2 suppresses Arabidopsis stomatal immunity%2E PLoS One 9%3A e114921%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Hurley, B., Lee, D., Mott, A., Wilton, M., Liu, J., Liu, Y.C., Angers, S., Coaker, G., Guttman, D.S., and Desveaux, D. (2014). The Pseudomonas syringae type III effector HopF2 suppresses Arabidopsis stomatal immunity. PLoS One 9: e114921.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Hurley,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The Pseudomonas syringae type III effector HopF2 suppresses Arabidopsis stomatal immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The Pseudomonas syringae type III effector HopF2 suppresses Arabidopsis stomatal immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hurley,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Jammes%2C F%2E%2C Yang%2C X%2E%2C Xiao%2C S%2E%2C and Kwak%2C J%2EM%2E %282011%29%2E Two Arabidopsis guard cell%2Dpreferential MAPK genes%2C MPK9 and MPK12%2C function in biotic stress response%2E Plant Signal Behav%2E 6%3A 1875%2D1877%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Jammes, F., Yang, X., Xiao, S., and Kwak, J.M. (2011). Two Arabidopsis guard cell-preferential MAPK genes, MPK9 and MPK12, function in biotic stress response. Plant Signal Behav. 6: 1875-1877.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Jammes,&hl=en&c2coff=1


http://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jammes,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Jammes%2C F%2E%2C Song%2C C%2E%2C Shin%2C D%2E%2C Munemasa%2C S%2E%2C Takeda%2C K%2E%2C Gu%2C D%2E%2C Cho%2C D%2E%2C Lee%2C S%2E%2C Giordo%2C R%2E%2C Sritubtim%2C S%2E%2C et al%2E %282009%29%2E MAP kinases MPK9 and MPK12 are preferentially expressed in guard cells and positively regulate ROS%2Dmediated ABA signaling%2E Proc%2E Natl%2E Acad%2E Sci%2E USA 106%3A 20520%2D20525%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Jammes, F., Song, C., Shin, D., Munemasa, S., Takeda, K., Gu, D., Cho, D., Lee, S., Giordo, R., Sritubtim, S., et al. (2009). MAP kinases MPK9 and MPK12 are preferentially expressed in guard cells and positively regulate ROS-mediated ABA signaling. Proc. Natl. Acad. Sci. USA 106: 20520-20525.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Jammes,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=MAP kinases MPK9 and MPK12 are preferentially expressed in guard cells and positively regulate ROS-mediated ABA signaling&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=MAP kinases MPK9 and MPK12 are preferentially expressed in guard cells and positively regulate ROS-mediated ABA signaling&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jammes,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Jinno%2C N%2EF%2E%2C M%2E %281981%29%2E Effects of calcium%2Dcomplexing agents%2C sodium oxalate and soduim citrate on stomatal opening and closing of Commelina communis L%2E Science Bulletin of the Faculty of Education%2C Nagasaki University 32%3A 83%2D86%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Jinno, N.F., M. (1981). Effects of calcium-complexing agents, sodium oxalate and soduim citrate on stomatal opening and closing of Commelina communis L. Science Bulletin of the Faculty of Education, Nagasaki University 32: 83-86.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Jinno,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Effects of calcium-complexing agents, sodium oxalate and soduim citrate on stomatal opening and closing of Commelina communis L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Effects of calcium-complexing agents, sodium oxalate and soduim citrate on stomatal opening and closing of Commelina communis L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jinno,&as_ylo=1981&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Johansson%2C F%2E%2C Sommarin%2C M%2E%2C and Larsson%2C C%2E %281993%29%2E Fusicoccin activates the plasma membrane H%2B%2DATPase by a mechanism involving the C%2Dterminal inhibitory domain%2E Plant Cell 5%3A 321%2D327%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Johansson, F., Sommarin, M., and Larsson, C. (1993). Fusicoccin activates the plasma membrane H+-ATPase by a mechanism involving the C-terminal inhibitory domain. Plant Cell 5: 321-327.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Johansson,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Fusicoccin activates the plasma membrane H+-ATPase by a mechanism involving the C-terminal inhibitory domain&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Fusicoccin activates the plasma membrane H+-ATPase by a mechanism involving the C-terminal inhibitory domain&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Johansson,&as_ylo=1993&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Jones%2C J%2ED%2E%2C and Dangl%2C J%2EL%2E %282006%29%2E The plant immune system%2E Nature 444%3A 323%2D329%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Jones, J.D., and Dangl, J.L. (2006). The plant immune system. Nature 444: 323-329.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Jones,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The plant immune system&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The plant immune system&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jones,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Kadota%2C Y%2E%2C Sklenar%2C J%2E%2C Derbyshire%2C P%2E%2C Stransfeld%2C L%2E%2C Asai%2C S%2E%2C Ntoukakis%2C V%2E%2C Jones%2C J%2ED%2E%2C Shirasu%2C K%2E%2C Menke%2C F%2E%2C Jones%2C A%2E%2C et al%2E %282014%29%2E Direct regulation of the NADPH oxidase RBOHD by the PRR%2Dassociated kinase BIK1 during plant immunity%2E Mol%2E Cell 54%3A 43%2D55%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Kadota, Y., Sklenar, J., Derbyshire, P., Stransfeld, L., Asai, S., Ntoukakis, V., Jones, J.D., Shirasu, K., Menke, F., Jones, A., et al. (2014). Direct regulation of the NADPH oxidase RBOHD by the PRR-associated kinase BIK1 during plant immunity. Mol. Cell 54: 43-55.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Kadota,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Direct regulation of the NADPH oxidase RBOHD by the PRR-associated kinase BIK1 during plant immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Direct regulation of the NADPH oxidase RBOHD by the PRR-associated kinase BIK1 during plant immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kadota,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Kelly%2C G%2E%2C Moshelion%2C M%2E%2C David%2DSchwartz%2C R%2E%2C Halperin%2C O%2E%2C Wallach%2C R%2E%2C Attia%2C Z%2E%2C Belausov%2C E%2E%2C and Granot%2C D%2E %282013%29%2E Hexokinase mediates stomatal closure%2E Plant J%2E 75%3A 977%2D988%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Kelly, G., Moshelion, M., David-Schwartz, R., Halperin, O., Wallach, R., Attia, Z., Belausov, E., and Granot, D. (2013). Hexokinase mediates stomatal closure. Plant J. 75: 977-988.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Kelly,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Hexokinase mediates stomatal closure&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Hexokinase mediates stomatal closure&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kelly,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Kim%2C T%2EH%2E%2C Bohmer%2C M%2E%2C Hu%2C H%2E%2C Nishimura%2C N%2E%2C and Schroeder%2C J%2EI%2E %282010%29%2E Guard cell signal transduction network%3A advances in understanding abscisic acid%2C CO2%2C and Ca2%2B signaling%2E Annu%2E Rev%2E Plant Biol%2E 61%3A 561%2D591%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Kim, T.H., Bohmer, M., Hu, H., Nishimura, N., and Schroeder, J.I. (2010). Guard cell signal transduction network: advances in understanding abscisic acid, CO2, and Ca2+ signaling. Annu. Rev. Plant Biol. 61: 561-591.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Kim,&hl=en&c2coff=1


http://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kim,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Kim%2C T%2EH%2E%2C Hauser%2C F%2E%2C Ha%2C T%2E%2C Xue%2C S%2E%2C Bohmer%2C M%2E%2C Nishimura%2C N%2E%2C Munemasa%2C S%2E%2C Hubbard%2C K%2E%2C Peine%2C N%2E%2C Lee%2C B%2EH%2E%2C et al%2E %282011%29%2E Chemical genetics reveals negative regulation of abscisic acid signaling by a plant immune response pathway%2E Curr%2E Biol%2E 21%3A 990%2D997%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Kim, T.H., Hauser, F., Ha, T., Xue, S., Bohmer, M., Nishimura, N., Munemasa, S., Hubbard, K., Peine, N., Lee, B.H., et al. (2011). Chemical genetics reveals negative regulation of abscisic acid signaling by a plant immune response pathway. Curr. Biol. 21: 990-997.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Kim,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Chemical genetics reveals negative regulation of abscisic acid signaling by a plant immune response pathway&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Chemical genetics reveals negative regulation of abscisic acid signaling by a plant immune response pathway&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kim,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Lawson%2C T%2E %282009%29%2E Guard cell photosynthesis and stomatal function%2E New Phytol%2E 181%3A 13%2D34%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lawson, T. (2009). Guard cell photosynthesis and stomatal function. New Phytol. 181: 13-34.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Lawson,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Guard cell photosynthesis and stomatal function&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Guard cell photosynthesis and stomatal function&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lawson,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

Lee, J.S., Wang, S., Sritubtim, S., Chen, J.G., and Ellis, B.E. (2009). Arabidopsis mitogen-activated protein kinase MPK12 interactswith the MAPK phosphatase IBR5 and regulates auxin signaling. Plant J. 57: 975-985.


Lee, M., Choi, Y., Burla, B., Kim, Y.Y., Jeon, B., Maeshima, M., Yoo, J.Y., Martinoia, E., and Lee, Y. (2008). The ABC transporterAtABCB14 is a malate importer and modulates stomatal response to CO2. Nat. Cell Biol. 10: 1217-1223.


Lemaitre, T., Urbanczyk-Wochniak, E., Flesch, V., Bismuth, E., Fernie, A.R., and Hodges, M. (2007). NAD-dependent isocitratedehydrogenase mutants of Arabidopsis suggest the enzyme is not limiting for nitrogen assimilation. Plant Physiol. 144: 1546-1558.


Li, L., Li, M., Yu, L., Zhou, Z., Liang, X., Liu, Z., Cai, G., Gao, L., Zhang, X., Wang, Y., et al. (2014). The FLS2-associated kinase BIK1directly phosphorylates the NADPH oxidase RbohD to control plant immunity. Cell Host Microbe 15: 329-338.


Li, S., Mhamdi, A., Clement, C., Jolivet, Y., and Noctor, G. (2013). Analysis of knockout mutants suggests that Arabidopsis NADP-MALIC ENZYME2 does not play an essential role in responses to oxidative stress of intracellular or extracellular origin. J. Exp.Bot. 64: 3605-3614.


Lim, C.W., Luan, S., and Lee, S.C. (2014). A prominent role for RCAR3-mediated ABA signaling in response to Pseudomonassyringae pv. tomato DC3000 infection in Arabidopsis. Plant Cell Physiol. 55: 1691-1703.


Liu, Y., and Zhang, S. (2004). Phosphorylation of 1-aminocyclopropane-1-carboxylic acid synthase by MPK6, a stress-responsivemitogen-activated protein kinase, induces ethylene biosynthesis in Arabidopsis. Plant Cell 16: 3386-3399.


Lozano-Duran, R., Bourdais, G., He, S.Y., and Robatzek, S. (2014). The bacterial effector HopM1 suppresses PAMP-triggeredoxidative burst and stomatal immunity. New Phytol. 202: 259-269.


Mao, G., Meng, X., Liu, Y., Zheng, Z., Chen, Z., and Zhang, S. (2011). Phosphorylation of a WRKY transcription factor by twopathogen-responsive MAPKs drives phytoalexin biosynthesis in Arabidopsis. Plant Cell 23: 1639-1653.


McLachlan, D.H., Kopischke, M., and Robatzek, S. (2014). Gate control: guard cell regulation by microbial stress. New Phytol. 203:1049-1063.


McLachlan, D.H., Lan, J., Geilfus, C.M., Dodd, A.N., Larson, T., Baker, A., Horak, H., Kollist, H., He, Z., Graham, I., et al. (2016). Thebreakdown of stored triacylglycerols is required during light-induced stomatal opening. Curr. Biol. 26: 707-712.


Melotto, M., Underwood, W., and He, S.Y. (2008). Role of stomata in plant innate immunity and foliar bacterial diseases. Annu. Rev.Phytopathol. 46: 101-122.


Melotto, M., Underwood, W., Koczan, J., Nomura, K., and He, S.Y. (2006). Plant stomata function in innate immunity against bacterialinvasion. Cell 126: 969-980.


Meng, X., and Zhang, S. (2013). MAPK cascades in plant disease resistance signaling. Annu. Rev. Phytopathol. 51: 245-266.

http://www.ncbi.nlm.nih.gov/pubmed?term=Lee%2C J%2ES%2E%2C Wang%2C S%2E%2C Sritubtim%2C S%2E%2C Chen%2C J%2EG%2E%2C and Ellis%2C B%2EE%2E %282009%29%2E Arabidopsis mitogen%2Dactivated protein kinase MPK12 interacts with the MAPK phosphatase IBR5 and regulates auxin signaling%2E Plant J%2E 57%3A 975%2D985%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lee, J.S., Wang, S., Sritubtim, S., Chen, J.G., and Ellis, B.E. (2009). Arabidopsis mitogen-activated protein kinase MPK12 interacts with the MAPK phosphatase IBR5 and regulates auxin signaling. Plant J. 57: 975-985.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Lee,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis mitogen-activated protein kinase MPK12 interacts with the MAPK phosphatase IBR5 and regulates auxin signaling&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis mitogen-activated protein kinase MPK12 interacts with the MAPK phosphatase IBR5 and regulates auxin signaling&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lee,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Lee%2C M%2E%2C Choi%2C Y%2E%2C Burla%2C B%2E%2C Kim%2C Y%2EY%2E%2C Jeon%2C B%2E%2C Maeshima%2C M%2E%2C Yoo%2C J%2EY%2E%2C Martinoia%2C E%2E%2C and Lee%2C Y%2E %282008%29%2E The ABC transporter AtABCB14 is a malate importer and modulates stomatal response to CO2%2E Nat%2E Cell Biol%2E 10%3A 1217%2D1223%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lee, M., Choi, Y., Burla, B., Kim, Y.Y., Jeon, B., Maeshima, M., Yoo, J.Y., Martinoia, E., and Lee, Y. (2008). The ABC transporter AtABCB14 is a malate importer and modulates stomatal response to CO2. Nat. Cell Biol. 10: 1217-1223.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Lee,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The ABC transporter AtABCB14 is a malate importer and modulates stomatal response to CO2&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The ABC transporter AtABCB14 is a malate importer and modulates stomatal response to CO2&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lee,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Lemaitre%2C T%2E%2C Urbanczyk%2DWochniak%2C E%2E%2C Flesch%2C V%2E%2C Bismuth%2C E%2E%2C Fernie%2C A%2ER%2E%2C and Hodges%2C M%2E %282007%29%2E NAD%2Ddependent isocitrate dehydrogenase mutants of Arabidopsis suggest the enzyme is not limiting for nitrogen assimilation%2E Plant Physiol%2E 144%3A 1546%2D1558%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lemaitre, T., Urbanczyk-Wochniak, E., Flesch, V., Bismuth, E., Fernie, A.R., and Hodges, M. (2007). NAD-dependent isocitrate dehydrogenase mutants of Arabidopsis suggest the enzyme is not limiting for nitrogen assimilation. Plant Physiol. 144: 1546-1558.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Lemaitre,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=NAD-dependent isocitrate dehydrogenase mutants of Arabidopsis suggest the enzyme is not limiting for nitrogen assimilation&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=NAD-dependent isocitrate dehydrogenase mutants of Arabidopsis suggest the enzyme is not limiting for nitrogen assimilation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lemaitre,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Li%2C L%2E%2C Li%2C M%2E%2C Yu%2C L%2E%2C Zhou%2C Z%2E%2C Liang%2C X%2E%2C Liu%2C Z%2E%2C Cai%2C G%2E%2C Gao%2C L%2E%2C Zhang%2C X%2E%2C Wang%2C Y%2E%2C et al%2E %282014%29%2E The FLS2%2Dassociated kinase BIK1 directly phosphorylates the NADPH oxidase RbohD to control plant immunity%2E Cell Host Microbe 15%3A 329%2D338%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Li, L., Li, M., Yu, L., Zhou, Z., Liang, X., Liu, Z., Cai, G., Gao, L., Zhang, X., Wang, Y., et al. (2014). The FLS2-associated kinase BIK1 directly phosphorylates the NADPH oxidase RbohD to control plant immunity. Cell Host Microbe 15: 329-338.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Li,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The FLS2-associated kinase BIK1 directly phosphorylates the NADPH oxidase RbohD to control plant immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The FLS2-associated kinase BIK1 directly phosphorylates the NADPH oxidase RbohD to control plant immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Li,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Li%2C S%2E%2C Mhamdi%2C A%2E%2C Clement%2C C%2E%2C Jolivet%2C Y%2E%2C and Noctor%2C G%2E %282013%29%2E Analysis of knockout mutants suggests that Arabidopsis NADP%2DMALIC ENZYME2 does not play an essential role in responses to oxidative stress of intracellular or extracellular origin%2E J%2E Exp%2E Bot%2E 64%3A 3605%2D3614%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Li, S., Mhamdi, A., Clement, C., Jolivet, Y., and Noctor, G. (2013). Analysis of knockout mutants suggests that Arabidopsis NADP-MALIC ENZYME2 does not play an essential role in responses to oxidative stress of intracellular or extracellular origin. J. Exp. Bot. 64: 3605-3614.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Li,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Analysis of knockout mutants suggests that Arabidopsis NADP-MALIC ENZYME2 does not play an essential role in responses to oxidative stress of intracellular or extracellular origin&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Analysis of knockout mutants suggests that Arabidopsis NADP-MALIC ENZYME2 does not play an essential role in responses to oxidative stress of intracellular or extracellular origin&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Li,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Lim%2C C%2EW%2E%2C Luan%2C S%2E%2C and Lee%2C S%2EC%2E %282014%29%2E A prominent role for RCAR3%2Dmediated ABA signaling in response to Pseudomonas syringae pv%2E tomato DC3000 infection in Arabidopsis%2E Plant Cell Physiol%2E 55%3A 1691%2D1703%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lim, C.W., Luan, S., and Lee, S.C. (2014). A prominent role for RCAR3-mediated ABA signaling in response to Pseudomonas syringae pv. tomato DC3000 infection in Arabidopsis. Plant Cell Physiol. 55: 1691-1703.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Lim,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A prominent role for RCAR3-mediated ABA signaling in response to Pseudomonas syringae pv&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A prominent role for RCAR3-mediated ABA signaling in response to Pseudomonas syringae pv&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lim,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Liu%2C Y%2E%2C and Zhang%2C S%2E %282004%29%2E Phosphorylation of 1%2Daminocyclopropane%2D1%2Dcarboxylic acid synthase by MPK6%2C a stress%2Dresponsive mitogen%2Dactivated protein kinase%2C induces ethylene biosynthesis in Arabidopsis%2E Plant Cell 16%3A 3386%2D3399%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Liu, Y., and Zhang, S. (2004). Phosphorylation of 1-aminocyclopropane-1-carboxylic acid synthase by MPK6, a stress-responsive mitogen-activated protein kinase, induces ethylene biosynthesis in Arabidopsis. Plant Cell 16: 3386-3399.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Liu,&hl=en&c2coff=1


http://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Liu,&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Lozano%2DDuran%2C R%2E%2C Bourdais%2C G%2E%2C He%2C S%2EY%2E%2C and Robatzek%2C S%2E %282014%29%2E The bacterial effector HopM1 suppresses PAMP%2Dtriggered oxidative burst and stomatal immunity%2E New Phytol%2E 202%3A 259%2D269%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lozano-Duran, R., Bourdais, G., He, S.Y., and Robatzek, S. (2014). The bacterial effector HopM1 suppresses PAMP-triggered oxidative burst and stomatal immunity. New Phytol. 202: 259-269.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Lozano-Duran,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The bacterial effector HopM1 suppresses PAMP-triggered oxidative burst and stomatal immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The bacterial effector HopM1 suppresses PAMP-triggered oxidative burst and stomatal immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lozano-Duran,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Mao%2C G%2E%2C Meng%2C X%2E%2C Liu%2C Y%2E%2C Zheng%2C Z%2E%2C Chen%2C Z%2E%2C and Zhang%2C S%2E %282011%29%2E Phosphorylation of a WRKY transcription factor by two pathogen%2Dresponsive MAPKs drives phytoalexin biosynthesis in Arabidopsis%2E Plant Cell 23%3A 1639%2D1653%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Mao, G., Meng, X., Liu, Y., Zheng, Z., Chen, Z., and Zhang, S. (2011). Phosphorylation of a WRKY transcription factor by two pathogen-responsive MAPKs drives phytoalexin biosynthesis in Arabidopsis. Plant Cell 23: 1639-1653.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Mao,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Phosphorylation of a WRKY transcription factor by two pathogen-responsive MAPKs drives phytoalexin biosynthesis in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Phosphorylation of a WRKY transcription factor by two pathogen-responsive MAPKs drives phytoalexin biosynthesis in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Mao,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=McLachlan%2C D%2EH%2E%2C Kopischke%2C M%2E%2C and Robatzek%2C S%2E %282014%29%2E Gate control%3A guard cell regulation by microbial stress%2E New Phytol%2E 203%3A 1049%2D1063%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=McLachlan, D.H., Kopischke, M., and Robatzek, S. (2014). Gate control: guard cell regulation by microbial stress. New Phytol. 203: 1049-1063.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=McLachlan,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Gate control: guard cell regulation by microbial stress&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Gate control: guard cell regulation by microbial stress&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=McLachlan,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=McLachlan%2C D%2EH%2E%2C Lan%2C J%2E%2C Geilfus%2C C%2EM%2E%2C Dodd%2C A%2EN%2E%2C Larson%2C T%2E%2C Baker%2C A%2E%2C Horak%2C H%2E%2C Kollist%2C H%2E%2C He%2C Z%2E%2C Graham%2C I%2E%2C et al%2E %282016%29%2E The breakdown of stored triacylglycerols is required during light%2Dinduced stomatal opening%2E Curr%2E Biol%2E 26%3A 707%2D712%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=McLachlan, D.H., Lan, J., Geilfus, C.M., Dodd, A.N., Larson, T., Baker, A., Horak, H., Kollist, H., He, Z., Graham, I., et al. (2016). The breakdown of stored triacylglycerols is required during light-induced stomatal opening. Curr. Biol. 26: 707-712.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=McLachlan,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=The breakdown of stored triacylglycerols is required during light-induced stomatal opening&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The breakdown of stored triacylglycerols is required during light-induced stomatal opening&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=McLachlan,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Melotto%2C M%2E%2C Underwood%2C W%2E%2C and He%2C S%2EY%2E %282008%29%2E Role of stomata in plant innate immunity and foliar bacterial diseases%2E Annu%2E Rev%2E Phytopathol%2E 46%3A 101%2D122%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Melotto, M., Underwood, W., and He, S.Y. (2008). Role of stomata in plant innate immunity and foliar bacterial diseases. Annu. Rev. Phytopathol. 46: 101-122.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Melotto,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Role of stomata in plant innate immunity and foliar bacterial diseases&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Role of stomata in plant innate immunity and foliar bacterial diseases&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Melotto,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Melotto%2C M%2E%2C Underwood%2C W%2E%2C Koczan%2C J%2E%2C Nomura%2C K%2E%2C and He%2C S%2EY%2E %282006%29%2E Plant stomata function in innate immunity against bacterial invasion%2E Cell 126%3A 969%2D980%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Melotto, M., Underwood, W., Koczan, J., Nomura, K., and He, S.Y. (2006). Plant stomata function in innate immunity against bacterial invasion. Cell 126: 969-980.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Melotto,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Plant stomata function in innate immunity against bacterial invasion&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Plant stomata function in innate immunity against bacterial invasion&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Melotto,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1


Menke, F.L., van Pelt, J.A., Pieterse, C.M., and Klessig, D.F. (2004). Silencing of the mitogen-activated protein kinase MPK6compromises disease resistance in Arabidopsis. Plant Cell 16: 897-907.


Meyer, S., Mumm, P., Imes, D., Endler, A., Weder, B., Al-Rasheid, K.A., Geiger, D., Marten, I., Martinoia, E., and Hedrich, R. (2010).AtALMT12 represents an R-type anion channel required for stomatal movement in Arabidopsis guard cells. Plant J. 63: 1054-1062.


Mhamdi, A., Mauve, C., Gouia, H., Saindrenan, P., Hodges, M., and Noctor, G. (2010). Cytosolic NADP-dependent isocitratedehydrogenase contributes to redox homeostasis and the regulation of pathogen responses in Arabidopsis leaves. Plant CellEnviron. 33: 1112-1123.


Monaghan, J., and Zipfel, C. (2012). Plant pattern recognition receptor complexes at the plasma membrane. Curr. Opin. Plant Biol.15: 349-357.


Montillet, J.L., Leonhardt, N., Mondy, S., Tranchimand, S., Rumeau, D., Boudsocq, M., Garcia, A.V., Douki, T., Bigeard, J., Lauriere,C., et al. (2013). An abscisic acid-independent oxylipin pathway controls stomatal closure and immune defense in Arabidopsis.PLoS Biol. 11: e1001513.


Mustilli, A.C., Merlot, S., Vavasseur, A., Fenzi, F., and Giraudat, J. (2002). Arabidopsis OST1 protein kinase mediates the regulationof stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production. Plant Cell 14: 3089-3099.


Nagy, S.K., Darula, Z., Kallai, B.M., Bogre, L., Banhegyi, G., Medzihradszky, K.F., Horvath, G.V., and Meszaros, T. (2015). Activationof AtMPK9 through autophosphorylation that makes it independent of the canonical MAPK cascades. Biochem J. 467: 167-175.


Negi, J., Matsuda, O., Nagasawa, T., Oba, Y., Takahashi, H., Kawai-Yamada, M., Uchimiya, H., Hashimoto, M., and Iba, K. (2008). CO2regulator SLAC1 and its homologues are essential for anion homeostasis in plant cells. Nature 452: 483-486.


Peleg, Y., Rokem, J.S., and Goldberg, I. (1990). A simple plate-assay for the screening of L-malic acid producing microorganisms.FEMS Microbiol. Lett. 55: 233-236.


Ramesh, S.A., Tyerman, S.D., Xu, B., Bose, J., Kaur, S., Conn, V., Domingos, P., Ullah, S., Wege, S., Shabala, S., et al. (2015). GABAsignalling modulates plant growth by directly regulating the activity of plant-specific anion transporters. Nat. Commun. 6: 7879.


Ren, D., Yang, H., and Zhang, S. (2002). Cell death mediated by MAPK is associated with hydrogen peroxide production inArabidopsis. J. Biol. Chem. 277: 559-565.


Ren, D., Liu, Y., Yang, K.Y., Han, L., Mao, G., Glazebrook, J., and Zhang, S. (2008). A fungal-responsive MAPK cascade regulatesphytoalexin biosynthesis in Arabidopsis. Proc. Natl. Acad. Sci. USA 105: 5638-5643.


Ruszala, E.M., Beerling, D.J., Franks, P.J., Chater, C., Casson, S.A., Gray, J.E., and Hetherington, A.M. (2011). Land plants acquiredactive stomatal control early in their evolutionary history. Curr. Biol. 21: 1030-1035.

http://www.ncbi.nlm.nih.gov/pubmed?term=Meng%2C X%2E%2C and Zhang%2C S%2E %282013%29%2E MAPK cascades in plant disease resistance signaling%2E Annu%2E Rev%2E Phytopathol%2E 51%3A 245%2D266%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Meng, X., and Zhang, S. (2013). MAPK cascades in plant disease resistance signaling. Annu. Rev. Phytopathol. 51: 245-266.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Meng,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=MAPK cascades in plant disease resistance signaling&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=MAPK cascades in plant disease resistance signaling&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Meng,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Menke%2C F%2EL%2E%2C van Pelt%2C J%2EA%2E%2C Pieterse%2C C%2EM%2E%2C and Klessig%2C D%2EF%2E %282004%29%2E Silencing of the mitogen%2Dactivated protein kinase MPK6 compromises disease resistance in Arabidopsis%2E Plant Cell 16%3A 897%2D907%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Menke, F.L., van Pelt, J.A., Pieterse, C.M., and Klessig, D.F. (2004). Silencing of the mitogen-activated protein kinase MPK6 compromises disease resistance in Arabidopsis. Plant Cell 16: 897-907.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Menke,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Silencing of the mitogen-activated protein kinase MPK6 compromises disease resistance in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Silencing of the mitogen-activated protein kinase MPK6 compromises disease resistance in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Menke,&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Meyer%2C S%2E%2C Mumm%2C P%2E%2C Imes%2C D%2E%2C Endler%2C A%2E%2C Weder%2C B%2E%2C Al%2DRasheid%2C K%2EA%2E%2C Geiger%2C D%2E%2C Marten%2C I%2E%2C Martinoia%2C E%2E%2C and Hedrich%2C R%2E %282010%29%2E AtALMT12 represents an R%2Dtype anion channel required for stomatal movement in Arabidopsis guard cells%2E Plant J%2E 63%3A 1054%2D1062%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Meyer, S., Mumm, P., Imes, D., Endler, A., Weder, B., Al-Rasheid, K.A., Geiger, D., Marten, I., Martinoia, E., and Hedrich, R. (2010). AtALMT12 represents an R-type anion channel required for stomatal movement in Arabidopsis guard cells. Plant J. 63: 1054-1062.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Meyer,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=AtALMT12 represents an R-type anion channel required for stomatal movement in Arabidopsis guard cells&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=AtALMT12 represents an R-type anion channel required for stomatal movement in Arabidopsis guard cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Meyer,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Mhamdi%2C A%2E%2C Mauve%2C C%2E%2C Gouia%2C H%2E%2C Saindrenan%2C P%2E%2C Hodges%2C M%2E%2C and Noctor%2C G%2E %282010%29%2E Cytosolic NADP%2Ddependent isocitrate dehydrogenase contributes to redox homeostasis and the regulation of pathogen responses in Arabidopsis leaves%2E Plant Cell Environ%2E 33%3A 1112%2D1123%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Mhamdi, A., Mauve, C., Gouia, H., Saindrenan, P., Hodges, M., and Noctor, G. (2010). Cytosolic NADP-dependent isocitrate dehydrogenase contributes to redox homeostasis and the regulation of pathogen responses in Arabidopsis leaves. Plant Cell Environ. 33: 1112-1123.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Mhamdi,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Cytosolic NADP-dependent isocitrate dehydrogenase contributes to redox homeostasis and the regulation of pathogen responses in Arabidopsis leaves&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Cytosolic NADP-dependent isocitrate dehydrogenase contributes to redox homeostasis and the regulation of pathogen responses in Arabidopsis leaves&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Mhamdi,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Monaghan%2C J%2E%2C and Zipfel%2C C%2E %282012%29%2E Plant pattern recognition receptor complexes at the plasma membrane%2E Curr%2E Opin%2E Plant Biol%2E 15%3A 349%2D357%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Monaghan, J., and Zipfel, C. (2012). Plant pattern recognition receptor complexes at the plasma membrane. Curr. Opin. Plant Biol. 15: 349-357.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Monaghan,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Plant pattern recognition receptor complexes at the plasma membrane&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Plant pattern recognition receptor complexes at the plasma membrane&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Monaghan,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Montillet%2C J%2EL%2E%2C Leonhardt%2C N%2E%2C Mondy%2C S%2E%2C Tranchimand%2C S%2E%2C Rumeau%2C D%2E%2C Boudsocq%2C M%2E%2C Garcia%2C A%2EV%2E%2C Douki%2C T%2E%2C Bigeard%2C J%2E%2C Lauriere%2C C%2E%2C et al%2E %282013%29%2E An abscisic acid%2Dindependent oxylipin pathway controls stomatal closure and immune defense in Arabidopsis%2E PLoS Biol%2E 11%3A e1001513%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Montillet, J.L., Leonhardt, N., Mondy, S., Tranchimand, S., Rumeau, D., Boudsocq, M., Garcia, A.V., Douki, T., Bigeard, J., Lauriere, C., et al. (2013). An abscisic acid-independent oxylipin pathway controls stomatal closure and immune defense in Arabidopsis. PLoS Biol. 11: e1001513.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Montillet,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=An abscisic acid-independent oxylipin pathway controls stomatal closure and immune defense in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=An abscisic acid-independent oxylipin pathway controls stomatal closure and immune defense in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Montillet,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Mustilli%2C A%2EC%2E%2C Merlot%2C S%2E%2C Vavasseur%2C A%2E%2C Fenzi%2C F%2E%2C and Giraudat%2C J%2E %282002%29%2E Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production%2E Plant Cell 14%3A 3089%2D3099%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Mustilli, A.C., Merlot, S., Vavasseur, A., Fenzi, F., and Giraudat, J. (2002). Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production. Plant Cell 14: 3089-3099.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Mustilli,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Mustilli,&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Nagy%2C S%2EK%2E%2C Darula%2C Z%2E%2C Kallai%2C B%2EM%2E%2C Bogre%2C L%2E%2C Banhegyi%2C G%2E%2C Medzihradszky%2C K%2EF%2E%2C Horvath%2C G%2EV%2E%2C and Meszaros%2C T%2E %282015%29%2E Activation of AtMPK9 through autophosphorylation that makes it independent of the canonical MAPK cascades%2E Biochem J%2E 467%3A 167%2D175%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Nagy, S.K., Darula, Z., Kallai, B.M., Bogre, L., Banhegyi, G., Medzihradszky, K.F., Horvath, G.V., and Meszaros, T. (2015). Activation of AtMPK9 through autophosphorylation that makes it independent of the canonical MAPK cascades. Biochem J. 467: 167-175.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Nagy,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Activation of AtMPK9 through autophosphorylation that makes it independent of the canonical MAPK cascades&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Activation of AtMPK9 through autophosphorylation that makes it independent of the canonical MAPK cascades&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Nagy,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Negi%2C J%2E%2C Matsuda%2C O%2E%2C Nagasawa%2C T%2E%2C Oba%2C Y%2E%2C Takahashi%2C H%2E%2C Kawai%2DYamada%2C M%2E%2C Uchimiya%2C H%2E%2C Hashimoto%2C M%2E%2C and Iba%2C K%2E %282008%29%2E CO2 regulator SLAC1 and its homologues are essential for anion homeostasis in plant cells%2E Nature 452%3A 483%2D486%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Negi, J., Matsuda, O., Nagasawa, T., Oba, Y., Takahashi, H., Kawai-Yamada, M., Uchimiya, H., Hashimoto, M., and Iba, K. (2008). CO2 regulator SLAC1 and its homologues are essential for anion homeostasis in plant cells. Nature 452: 483-486.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Negi,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=CO2 regulator SLAC1 and its homologues are essential for anion homeostasis in plant cells&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=CO2 regulator SLAC1 and its homologues are essential for anion homeostasis in plant cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Negi,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Peleg%2C Y%2E%2C Rokem%2C J%2ES%2E%2C and Goldberg%2C I%2E %281990%29%2E A simple plate%2Dassay for the screening of L%2Dmalic acid producing microorganisms%2E FEMS Microbiol%2E Lett%2E 55%3A 233%2D236%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Peleg, Y., Rokem, J.S., and Goldberg, I. (1990). A simple plate-assay for the screening of L-malic acid producing microorganisms. FEMS Microbiol. Lett. 55: 233-236.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Peleg,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A simple plate-assay for the screening of L-malic acid producing microorganisms&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A simple plate-assay for the screening of L-malic acid producing microorganisms&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Peleg,&as_ylo=1990&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Ramesh%2C S%2EA%2E%2C Tyerman%2C S%2ED%2E%2C Xu%2C B%2E%2C Bose%2C J%2E%2C Kaur%2C S%2E%2C Conn%2C V%2E%2C Domingos%2C P%2E%2C Ullah%2C S%2E%2C Wege%2C S%2E%2C Shabala%2C S%2E%2C et al%2E %282015%29%2E GABA signalling modulates plant growth by directly regulating the activity of plant%2Dspecific anion transporters%2E Nat%2E Commun%2E 6%3A 7879%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Ramesh, S.A., Tyerman, S.D., Xu, B., Bose, J., Kaur, S., Conn, V., Domingos, P., Ullah, S., Wege, S., Shabala, S., et al. (2015). GABA signalling modulates plant growth by directly regulating the activity of plant-specific anion transporters. Nat. Commun. 6: 7879.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Ramesh,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=GABA signalling modulates plant growth by directly regulating the activity of plant-specific anion transporters&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=GABA signalling modulates plant growth by directly regulating the activity of plant-specific anion transporters&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ramesh,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Ren%2C D%2E%2C Yang%2C H%2E%2C and Zhang%2C S%2E %282002%29%2E Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis%2E J%2E Biol%2E Chem%2E 277%3A 559%2D565%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Ren, D., Yang, H., and Zhang, S. (2002). Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis. J. Biol. Chem. 277: 559-565.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Ren,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Cell death mediated by MAPK is associated with hydrogen peroxide production in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ren,&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Ren%2C D%2E%2C Liu%2C Y%2E%2C Yang%2C K%2EY%2E%2C Han%2C L%2E%2C Mao%2C G%2E%2C Glazebrook%2C J%2E%2C and Zhang%2C S%2E %282008%29%2E A fungal%2Dresponsive MAPK cascade regulates phytoalexin biosynthesis in Arabidopsis%2E Proc%2E Natl%2E Acad%2E Sci%2E USA 105%3A 5638%2D5643%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Ren, D., Liu, Y., Yang, K.Y., Han, L., Mao, G., Glazebrook, J., and Zhang, S. (2008). A fungal-responsive MAPK cascade regulates phytoalexin biosynthesis in Arabidopsis. Proc. Natl. Acad. Sci. USA 105: 5638-5643.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Ren,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A fungal-responsive MAPK cascade regulates phytoalexin biosynthesis in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A fungal-responsive MAPK cascade regulates phytoalexin biosynthesis in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ren,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1


Sasaki, T., Mori, I.C., Furuichi, T., Munemasa, S., Toyooka, K., Matsuoka, K., Murata, Y., and Yamamoto, Y. (2010). Closing plantstomata requires a homolog of an aluminum-activated malate transporter. Plant Cell Physiol. 51: 354-365.


Sirichandra, C., Gu, D., Hu, H.C., Davanture, M., Lee, S., Djaoui, M., Valot, B., Zivy, M., Leung, J., Merlot, S., et al. (2009).Phosphorylation of the Arabidopsis AtrbohF NADPH oxidase by OST1 protein kinase. FEBS Lett. 583: 2982-2986.


Tanaka, Y., Sano, T., Tamaoki, M., Nakajima, N., Kondo, N., and Hasezawa, S. (2006). Cytokinin and auxin inhibit abscisic acid-induced stomatal closure by enhancing ethylene production in Arabidopsis. J. Exp. Bot. 57: 2259-2266.


Torres, M.A., Dangl, J.L., and Jones, J.D. (2002). Arabidopsis gp91phox homologues AtrbohD and AtrbohF are required foraccumulation of reactive oxygen intermediates in the plant defense response. Proc. Natl. Acad. Sci. USA 99: 517-522.


Tronconi, M.A., Fahnenstich, H., Gerrard Weehler, M.C., Andreo, C.S., Flugge, U.I., Drincovich, M.F., and Maurino, V.G. (2008).Arabidopsis NAD-malic enzyme functions as a homodimer and heterodimer and has a major impact on nocturnal metabolism. PlantPhysiol. 146: 1540-1552.


Tsuda, K., Mine, A., Bethke, G., Igarashi, D., Botanga, C.J., Tsuda, Y., Glazebrook, J., Sato, M., and Katagiri, F. (2013). Dualregulation of gene expression mediated by extended MAPK activation and salicylic acid contributes to robust innate immunity inArabidopsis thaliana. PLoS Genet. 9: e1004015.


Vahisalu, T., Kollist, H., Wang, Y.F., Nishimura, N., Chan, W.Y., Valerio, G., Lamminmaki, A., Brosche, M., Moldau, H., Desikan, R., etal. (2008). SLAC1 is required for plant guard cell S-type anion channel function in stomatal signalling. Nature 452: 487-491.


Wang, H., Ngwenyama, N., Liu, Y., Walker, J.C., and Zhang, S. (2007). Stomatal development and patterning are regulated byenvironmentally responsive mitogen-activated protein kinases in Arabidopsis. Plant Cell 19: 63-73.


Wheeler, M.C., Tronconi, M.A., Drincovich, M.F., Andreo, C.S., Flugge, U.I., and Maurino, V.G. (2005). A comprehensive analysis ofthe NADP-malic enzyme gene family of Arabidopsis. Plant Physiol. 139: 39-51.


Xu, J., and Zhang, S. (2015). Mitogen-activated protein kinase cascades in signaling plant growth and development. Trends PlantSci. 20: 56-64.


Xu, J., Xie, J., Yan, C., Zou, X., Ren, D., and Zhang, S. (2014). A chemical genetic approach demonstrates that MPK3/MPK6activation and NADPH oxidase-mediated oxidative burst are two independent signaling events in plant immunity. Plant J. 77: 222-234.


Xu, J., Meng, J., Meng, X., Zhao, Y., Liu, J., Sun, T., Liu, Y., Wang, Q., and Zhang, S. (2016). Pathogen-responsive MPK3 and MPK6reprogram the biosynthesis of indole glucosinolates and their derivatives in Arabidopsis immunity. Plant Cell 28: 1144-1162.


Zeng, W., and He, S.Y. (2010). A prominent role of the flagellin receptor FLAGELLIN-SENSING2 in mediating stomatal response to

http://www.ncbi.nlm.nih.gov/pubmed?term=Ruszala%2C E%2EM%2E%2C Beerling%2C D%2EJ%2E%2C Franks%2C P%2EJ%2E%2C Chater%2C C%2E%2C Casson%2C S%2EA%2E%2C Gray%2C J%2EE%2E%2C and Hetherington%2C A%2EM%2E %282011%29%2E Land plants acquired active stomatal control early in their evolutionary history%2E Curr%2E Biol%2E 21%3A 1030%2D1035%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Ruszala, E.M., Beerling, D.J., Franks, P.J., Chater, C., Casson, S.A., Gray, J.E., and Hetherington, A.M. (2011). Land plants acquired active stomatal control early in their evolutionary history. Curr. Biol. 21: 1030-1035.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Ruszala,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Land plants acquired active stomatal control early in their evolutionary history&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Land plants acquired active stomatal control early in their evolutionary history&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ruszala,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Sasaki%2C T%2E%2C Mori%2C I%2EC%2E%2C Furuichi%2C T%2E%2C Munemasa%2C S%2E%2C Toyooka%2C K%2E%2C Matsuoka%2C K%2E%2C Murata%2C Y%2E%2C and Yamamoto%2C Y%2E %282010%29%2E Closing plant stomata requires a homolog of an aluminum%2Dactivated malate transporter%2E Plant Cell Physiol%2E 51%3A 354%2D365%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Sasaki, T., Mori, I.C., Furuichi, T., Munemasa, S., Toyooka, K., Matsuoka, K., Murata, Y., and Yamamoto, Y. (2010). Closing plant stomata requires a homolog of an aluminum-activated malate transporter. Plant Cell Physiol. 51: 354-365.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Sasaki,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Closing plant stomata requires a homolog of an aluminum-activated malate transporter&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Closing plant stomata requires a homolog of an aluminum-activated malate transporter&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sasaki,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Sirichandra%2C C%2E%2C Gu%2C D%2E%2C Hu%2C H%2EC%2E%2C Davanture%2C M%2E%2C Lee%2C S%2E%2C Djaoui%2C M%2E%2C Valot%2C B%2E%2C Zivy%2C M%2E%2C Leung%2C J%2E%2C Merlot%2C S%2E%2C et al%2E %282009%29%2E Phosphorylation of the Arabidopsis AtrbohF NADPH oxidase by OST1 protein kinase%2E FEBS Lett%2E 583%3A 2982%2D2986%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Sirichandra, C., Gu, D., Hu, H.C., Davanture, M., Lee, S., Djaoui, M., Valot, B., Zivy, M., Leung, J., Merlot, S., et al. (2009). Phosphorylation of the Arabidopsis AtrbohF NADPH oxidase by OST1 protein kinase. FEBS Lett. 583: 2982-2986.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Sirichandra,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Phosphorylation of the Arabidopsis AtrbohF NADPH oxidase by OST1 protein kinase&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Phosphorylation of the Arabidopsis AtrbohF NADPH oxidase by OST1 protein kinase&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sirichandra,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Tanaka%2C Y%2E%2C Sano%2C T%2E%2C Tamaoki%2C M%2E%2C Nakajima%2C N%2E%2C Kondo%2C N%2E%2C and Hasezawa%2C S%2E %282006%29%2E Cytokinin and auxin inhibit abscisic acid%2Dinduced stomatal closure by enhancing ethylene production in Arabidopsis%2E J%2E Exp%2E Bot%2E 57%3A 2259%2D2266%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Tanaka, Y., Sano, T., Tamaoki, M., Nakajima, N., Kondo, N., and Hasezawa, S. (2006). Cytokinin and auxin inhibit abscisic acid-induced stomatal closure by enhancing ethylene production in Arabidopsis. J. Exp. Bot. 57: 2259-2266.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Tanaka,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Cytokinin and auxin inhibit abscisic acid-induced stomatal closure by enhancing ethylene production in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Cytokinin and auxin inhibit abscisic acid-induced stomatal closure by enhancing ethylene production in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tanaka,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Torres%2C M%2EA%2E%2C Dangl%2C J%2EL%2E%2C and Jones%2C J%2ED%2E %282002%29%2E Arabidopsis gp91phox homologues AtrbohD and AtrbohF are required for accumulation of reactive oxygen intermediates in the plant defense response%2E Proc%2E Natl%2E Acad%2E Sci%2E USA 99%3A 517%2D522%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Torres, M.A., Dangl, J.L., and Jones, J.D. (2002). Arabidopsis gp91phox homologues AtrbohD and AtrbohF are required for accumulation of reactive oxygen intermediates in the plant defense response. Proc. Natl. Acad. Sci. USA 99: 517-522.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Torres,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis gp91phox homologues AtrbohD and AtrbohF are required for accumulation of reactive oxygen intermediates in the plant defense response&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis gp91phox homologues AtrbohD and AtrbohF are required for accumulation of reactive oxygen intermediates in the plant defense response&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Torres,&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Tronconi%2C M%2EA%2E%2C Fahnenstich%2C H%2E%2C Gerrard Weehler%2C M%2EC%2E%2C Andreo%2C C%2ES%2E%2C Flugge%2C U%2EI%2E%2C Drincovich%2C M%2EF%2E%2C and Maurino%2C V%2EG%2E %282008%29%2E Arabidopsis NAD%2Dmalic enzyme functions as a homodimer and heterodimer and has a major impact on nocturnal metabolism%2E Plant Physiol%2E 146%3A 1540%2D1552%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Tronconi, M.A., Fahnenstich, H., Gerrard Weehler, M.C., Andreo, C.S., Flugge, U.I., Drincovich, M.F., and Maurino, V.G. (2008). Arabidopsis NAD-malic enzyme functions as a homodimer and heterodimer and has a major impact on nocturnal metabolism. Plant Physiol. 146: 1540-1552.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Tronconi,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis NAD-malic enzyme functions as a homodimer and heterodimer and has a major impact on nocturnal metabolism&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis NAD-malic enzyme functions as a homodimer and heterodimer and has a major impact on nocturnal metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tronconi,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Tsuda%2C K%2E%2C Mine%2C A%2E%2C Bethke%2C G%2E%2C Igarashi%2C D%2E%2C Botanga%2C C%2EJ%2E%2C Tsuda%2C Y%2E%2C Glazebrook%2C J%2E%2C Sato%2C M%2E%2C and Katagiri%2C F%2E %282013%29%2E Dual regulation of gene expression mediated by extended MAPK activation and salicylic acid contributes to robust innate immunity in Arabidopsis thaliana%2E PLoS Genet%2E 9%3A e1004015%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Tsuda, K., Mine, A., Bethke, G., Igarashi, D., Botanga, C.J., Tsuda, Y., Glazebrook, J., Sato, M., and Katagiri, F. (2013). Dual regulation of gene expression mediated by extended MAPK activation and salicylic acid contributes to robust innate immunity in Arabidopsis thaliana. PLoS Genet. 9: e1004015.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Tsuda,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Dual regulation of gene expression mediated by extended MAPK activation and salicylic acid contributes to robust innate immunity in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Dual regulation of gene expression mediated by extended MAPK activation and salicylic acid contributes to robust innate immunity in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tsuda,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Vahisalu%2C T%2E%2C Kollist%2C H%2E%2C Wang%2C Y%2EF%2E%2C Nishimura%2C N%2E%2C Chan%2C W%2EY%2E%2C Valerio%2C G%2E%2C Lamminmaki%2C A%2E%2C Brosche%2C M%2E%2C Moldau%2C H%2E%2C Desikan%2C R%2E%2C et al%2E %282008%29%2E SLAC1 is required for plant guard cell S%2Dtype anion channel function in stomatal signalling%2E Nature 452%3A 487%2D491%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Vahisalu, T., Kollist, H., Wang, Y.F., Nishimura, N., Chan, W.Y., Valerio, G., Lamminmaki, A., Brosche, M., Moldau, H., Desikan, R., et al. (2008). SLAC1 is required for plant guard cell S-type anion channel function in stomatal signalling. Nature 452: 487-491.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Vahisalu,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=SLAC1 is required for plant guard cell S-type anion channel function in stomatal signalling&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=SLAC1 is required for plant guard cell S-type anion channel function in stomatal signalling&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Vahisalu,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Wang%2C H%2E%2C Ngwenyama%2C N%2E%2C Liu%2C Y%2E%2C Walker%2C J%2EC%2E%2C and Zhang%2C S%2E %282007%29%2E Stomatal development and patterning are regulated by environmentally responsive mitogen%2Dactivated protein kinases in Arabidopsis%2E Plant Cell 19%3A 63%2D73%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Wang, H., Ngwenyama, N., Liu, Y., Walker, J.C., and Zhang, S. (2007). Stomatal development and patterning are regulated by environmentally responsive mitogen-activated protein kinases in Arabidopsis. Plant Cell 19: 63-73.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Wang,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Stomatal development and patterning are regulated by environmentally responsive mitogen-activated protein kinases in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Stomatal development and patterning are regulated by environmentally responsive mitogen-activated protein kinases in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wang,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Wheeler%2C M%2EC%2E%2C Tronconi%2C M%2EA%2E%2C Drincovich%2C M%2EF%2E%2C Andreo%2C C%2ES%2E%2C Flugge%2C U%2EI%2E%2C and Maurino%2C V%2EG%2E %282005%29%2E A comprehensive analysis of the NADP%2Dmalic enzyme gene family of Arabidopsis%2E Plant Physiol%2E 139%3A 39%2D51%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Wheeler, M.C., Tronconi, M.A., Drincovich, M.F., Andreo, C.S., Flugge, U.I., and Maurino, V.G. (2005). A comprehensive analysis of the NADP-malic enzyme gene family of Arabidopsis. Plant Physiol. 139: 39-51.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Wheeler,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A comprehensive analysis of the NADP-malic enzyme gene family of Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A comprehensive analysis of the NADP-malic enzyme gene family of Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wheeler,&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Xu%2C J%2E%2C and Zhang%2C S%2E %282015%29%2E Mitogen%2Dactivated protein kinase cascades in signaling plant growth and development%2E Trends Plant Sci%2E 20%3A 56%2D64%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Xu, J., and Zhang, S. (2015). Mitogen-activated protein kinase cascades in signaling plant growth and development. Trends Plant Sci. 20: 56-64.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Xu,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Mitogen-activated protein kinase cascades in signaling plant growth and development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Mitogen-activated protein kinase cascades in signaling plant growth and development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Xu,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Xu%2C J%2E%2C Xie%2C J%2E%2C Yan%2C C%2E%2C Zou%2C X%2E%2C Ren%2C D%2E%2C and Zhang%2C S%2E %282014%29%2E A chemical genetic approach demonstrates that MPK3%2FMPK6 activation and NADPH oxidase%2Dmediated oxidative burst are two independent signaling events in plant immunity%2E Plant J%2E 77%3A 222%2D234%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Xu, J., Xie, J., Yan, C., Zou, X., Ren, D., and Zhang, S. (2014). A chemical genetic approach demonstrates that MPK3/MPK6 activation and NADPH oxidase-mediated oxidative burst are two independent signaling events in plant immunity. Plant J. 77: 222-234.


http://scholar.google.com/scholar?as_q=A chemical genetic approach demonstrates that MPK3/MPK6 activation and NADPH oxidase-mediated oxidative burst are two independent signaling events in plant immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A chemical genetic approach demonstrates that MPK3/MPK6 activation and NADPH oxidase-mediated oxidative burst are two independent signaling events in plant immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Xu,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Xu%2C J%2E%2C Meng%2C J%2E%2C Meng%2C X%2E%2C Zhao%2C Y%2E%2C Liu%2C J%2E%2C Sun%2C T%2E%2C Liu%2C Y%2E%2C Wang%2C Q%2E%2C and Zhang%2C S%2E %282016%29%2E Pathogen%2Dresponsive MPK3 and MPK6 reprogram the biosynthesis of indole glucosinolates and their derivatives in Arabidopsis immunity%2E Plant Cell 28%3A 1144%2D1162%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Xu, J., Meng, J., Meng, X., Zhao, Y., Liu, J., Sun, T., Liu, Y., Wang, Q., and Zhang, S. (2016). Pathogen-responsive MPK3 and MPK6 reprogram the biosynthesis of indole glucosinolates and their derivatives in Arabidopsis immunity. Plant Cell 28: 1144-1162.


http://scholar.google.com/scholar?as_q=Pathogen-responsive MPK3 and MPK6 reprogram the biosynthesis of indole glucosinolates and their derivatives in Arabidopsis immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Pathogen-responsive MPK3 and MPK6 reprogram the biosynthesis of indole glucosinolates and their derivatives in Arabidopsis immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Xu,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

Pseudomonas syringae pv tomato DC3000 in Arabidopsis. Plant Physiol. 153: 1188-1198.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Zhang, S., and Klessig, D.F. (2001). MAPK cascades in plant defense signaling. Trends Plant Sci. 6: 520-527.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Zhang, W., Nilson, S.E., and Assmann, S.M. (2008). Isolation and whole-cell patch clamping of Arabidopsis guard cell protoplasts.CSH Protoc. 2008: pdb prot5014.


Zhao, C., Nie, H., Shen, Q., Zhang, S., Lukowitz, W., and Tang, D. (2014). EDR1 physically interacts with MKK4/MKK5 and negativelyregulates a MAP kinase cascade to modulate plant innate immunity. PLoS Genet. 10: e1004389.


Zheng, X.Y., Zhou, M., Yoo, H., Pruneda-Paz, J.L., Spivey, N.W., Kay, S.A., and Dong, X. (2015). Spatial and temporal regulation ofbiosynthesis of the plant immune signal salicylic acid. Proc. Natl. Acad. Sci. USA 112: 9166-9173.


Zheng, X.Y., Spivey, N.W., Zeng, W., Liu, P.P., Fu, Z.Q., Klessig, D.F., He, S.Y., and Dong, X. (2012). Coronatine promotesPseudomonas syringae virulence in plants by activating a signaling cascade that inhibits salicylic acid accumulation. Cell HostMicrobe 11: 587-596.


http://www.ncbi.nlm.nih.gov/pubmed?term=Zeng%2C W%2E%2C and He%2C S%2EY%2E %282010%29%2E A prominent role of the flagellin receptor FLAGELLIN%2DSENSING2 in mediating stomatal response to Pseudomonas syringae pv tomato DC3000 in Arabidopsis%2E Plant Physiol%2E 153%3A 1188%2D1198%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Zeng, W., and He, S.Y. (2010). A prominent role of the flagellin receptor FLAGELLIN-SENSING2 in mediating stomatal response to Pseudomonas syringae pv tomato DC3000 in Arabidopsis. Plant Physiol. 153: 1188-1198.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Zeng,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=A prominent role of the flagellin receptor FLAGELLIN-SENSING2 in mediating stomatal response to Pseudomonas syringae pv tomato DC3000 in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A prominent role of the flagellin receptor FLAGELLIN-SENSING2 in mediating stomatal response to Pseudomonas syringae pv tomato DC3000 in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zeng,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Zhang%2C S%2E%2C and Klessig%2C D%2EF%2E %282001%29%2E MAPK cascades in plant defense signaling%2E Trends Plant Sci%2E 6%3A 520%2D527%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Zhang, S., and Klessig, D.F. (2001). MAPK cascades in plant defense signaling. Trends Plant Sci. 6: 520-527.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Zhang,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=MAPK cascades in plant defense signaling&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=MAPK cascades in plant defense signaling&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhang,&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Zhang%2C W%2E%2C Nilson%2C S%2EE%2E%2C and Assmann%2C S%2EM%2E %282008%29%2E Isolation and whole%2Dcell patch clamping of Arabidopsis guard cell protoplasts%2E CSH Protoc%2E 2008%3A pdb prot5014%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Zhang, W., Nilson, S.E., and Assmann, S.M. (2008). Isolation and whole-cell patch clamping of Arabidopsis guard cell protoplasts. CSH Protoc. 2008: pdb prot5014.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Zhang,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Isolation and whole-cell patch clamping of Arabidopsis guard cell protoplasts&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Isolation and whole-cell patch clamping of Arabidopsis guard cell protoplasts&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhang,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Zhao%2C C%2E%2C Nie%2C H%2E%2C Shen%2C Q%2E%2C Zhang%2C S%2E%2C Lukowitz%2C W%2E%2C and Tang%2C D%2E %282014%29%2E EDR1 physically interacts with MKK4%2FMKK5 and negatively regulates a MAP kinase cascade to modulate plant innate immunity%2E PLoS Genet%2E 10%3A e1004389%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Zhao, C., Nie, H., Shen, Q., Zhang, S., Lukowitz, W., and Tang, D. (2014). EDR1 physically interacts with MKK4/MKK5 and negatively regulates a MAP kinase cascade to modulate plant innate immunity. PLoS Genet. 10: e1004389.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Zhao,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=EDR1 physically interacts with MKK4/MKK5 and negatively regulates a MAP kinase cascade to modulate plant innate immunity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=EDR1 physically interacts with MKK4/MKK5 and negatively regulates a MAP kinase cascade to modulate plant innate immunity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhao,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Zheng%2C X%2EY%2E%2C Zhou%2C M%2E%2C Yoo%2C H%2E%2C Pruneda%2DPaz%2C J%2EL%2E%2C Spivey%2C N%2EW%2E%2C Kay%2C S%2EA%2E%2C and Dong%2C X%2E %282015%29%2E Spatial and temporal regulation of biosynthesis of the plant immune signal salicylic acid%2E Proc%2E Natl%2E Acad%2E Sci%2E USA 112%3A 9166%2D9173%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Zheng, X.Y., Zhou, M., Yoo, H., Pruneda-Paz, J.L., Spivey, N.W., Kay, S.A., and Dong, X. (2015). Spatial and temporal regulation of biosynthesis of the plant immune signal salicylic acid. Proc. Natl. Acad. Sci. USA 112: 9166-9173.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Zheng,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Spatial and temporal regulation of biosynthesis of the plant immune signal salicylic acid&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Spatial and temporal regulation of biosynthesis of the plant immune signal salicylic acid&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zheng,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=Zheng%2C X%2EY%2E%2C Spivey%2C N%2EW%2E%2C Zeng%2C W%2E%2C Liu%2C P%2EP%2E%2C Fu%2C Z%2EQ%2E%2C Klessig%2C D%2EF%2E%2C He%2C S%2EY%2E%2C and Dong%2C X%2E %282012%29%2E Coronatine promotes Pseudomonas syringae virulence in plants by activating a signaling cascade that inhibits salicylic acid accumulation%2E Cell Host Microbe 11%3A 587%2D596%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Zheng, X.Y., Spivey, N.W., Zeng, W., Liu, P.P., Fu, Z.Q., Klessig, D.F., He, S.Y., and Dong, X. (2012). Coronatine promotes Pseudomonas syringae virulence in plants by activating a signaling cascade that inhibits salicylic acid accumulation. Cell Host Microbe 11: 587-596.

http://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Zheng,&hl=en&c2coff=1

http://scholar.google.com/scholar?as_q=Coronatine promotes Pseudomonas syringae virulence in plants by activating a signaling cascade that inhibits salicylic acid accumulation&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Coronatine promotes Pseudomonas syringae virulence in plants by activating a signaling cascade that inhibits salicylic acid accumulation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zheng,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

DOI 10.1105/tpc.16.00577; originally published online March 2, 2017;Plant Cell

and Shuqun ZhangJianbin Su, Mengmeng Zhang, Lawrence Zhang, Tiefeng Sun, Yidong Liu, Wolfgang Lukowitz, Juan Xu

MPK3/MPK6 Cascade and Abscisic AcidRegulation of Stomatal Immunity by Interdependent Functions of a Pathogen-responsive

This information is current as of April 16, 2020

Supplemental Data /content/suppl/2017/03/31/tpc.16.00577.DC2.html /content/suppl/2017/03/02/tpc.16.00577.DC1.html

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