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1

Running Head Auxin regulates herbivory-induced secondary metabolites 1

Correspondence Matthias Erb University of Bern Institute of Plant Sciences 2

Altenbergrain 21 3013 Bern Switzerland Tel +41 31 631 86 68 matthiaserbipsunibech 3

Research area Signaling and response 4

Plant Physiology Preview Published on August 2 2016 as DOI101104pp1600940

Copyright 2016 by the American Society of Plant Biologists

wwwplantphysiolorgon June 1 2020 - Published by Downloaded from Copyright copy 2016 American Society of Plant Biologists All rights reserved

2

Auxin is rapidly induced by herbivore attack and regulates a subset of systemic 5

jasmonate-dependent secondary metabolites 6

Ricardo AR Machado12 Christelle AM Robert12 Carla CM Arce123 Abigail P Ferrieri1 7

Shuqing Xu1 Guillermo H Jimenez-Aleman1 Ian T Baldwin1 and Matthias Erb12 8

9

1Max Planck Institute for Chemical Ecology Hans-Knoumlll-Str 8 07745 Jena Germany 10

2Institute of Plant Sciences University of Bern Altenbergrain 21 3013 Bern Switzerland 11

3Departamento de Entomologia Universidade Federal de Viccedilosa 36570-000 Viccedilosa MG 12

Brazil 13

14

One sentence summary Herbivory-induced auxin promotes the production of anthocyanins 15

and phenolamides 16

17

18

This work was supported by the Max Planck Society a Humboldt Postdoctoral Research 19

Fellowship (AF) the Brazilian National Council for Research CNPq Grant No 2379292012-20

0 (CA) a Marie Curie Intra European Fellowship Grant No 328935 (SX) a Marie Curie 21

Intra European Fellowship Grant No 273107 (ME) a Swiss National Foundation Fellowship 22

Grant No 140196 (CR) a European Research Council advanced Grant No 293926 (ITB) and 23

Human Frontier Science Program Grant No RGP00022012 (ITB) 24

25

26

Author for correspondence (Phone +41 31 631 8668 E-mail matthiaserbipsunibech)27


3

ABSTRACT 28

Plant responses to herbivore attack are regulated by phytohormonal networks To date the 29

role of the auxin indole-3-acetic acid (IAA) in this context is not well understood We 30

quantified and manipulated the spatiotemporal patterns of IAA accumulation in herbivore-31

attacked Nicotiana attenuata plants to unravel its role in the regulation of plant secondary 32

metabolism We found that IAA is strongly rapidly and specifically induced by herbivore 33

attack IAA is elicited by herbivore oral secretions and fatty acid conjugate elicitors and is 34

accompanied by a rapid transcriptional increase of auxin biosynthetic YUCCA-like genes 35

IAA accumulation starts 30-60 seconds after local induction and peaks within 5 minutes after 36

induction thereby preceding the jasmonate (JA) burst IAA accumulation does not require JA 37

signaling and spreads rapidly from the wound site to systemic tissues Complementation and 38

transport inhibition experiments reveal that IAA is required for the herbivore-specific 39

jasmonate-dependent accumulation of anthocyanins and phenolamides in the stems In 40

contrast IAA does not affect the accumulation of nicotine or 7-hydroxygeranyllinalool 41

diterpene glycosides in the same tissue Taken together our results uncover IAA as a rapid 42

and specific signal that regulates a subset of systemic jasmonate-dependent secondary 43

metabolites in herbivore-attacked plants 44

45


4

INTRODUCTION 46

Plants withstand herbivore attack by specifically recognizing the attacker and mounting 47

appropriate defenses Induced defense responses are activated by hormone-mediated 48

signaling cascades (Erb et al 2012 Wu and Baldwin 2009) and jasmonates (JA) have 49

emerged as key regulators in this context (Geyter et al 2012 Howe and Jander 2008) As a 50

consequence their behavior and mode of action have been studied in great detail (Wasternack 51

and Hause 2013) Similarly other stress-related hormones such as salicylic acid abscisic 52

acid and ethylene have been shown to play important roles in the orchestration of plant 53

defenses against herbivores (Dahl et al 2007 Winz and Baldwin 2001 Thaler and Bostock 54

2004 Zhang et al 2013 Kroes et al 2014) Recent evidence also suggests that hormones 55

which have traditionally been classified as growth regulators participate in induced defense 56

responses Cytokinins for instance modulate wound-induced local and systemic defense 57

responses (Schaumlfer et al 2015) and gibberellins are involved in regulating the plantrsquos 58

investment into growth and defense (Li et al 2015 Hou et al 2010 Yang et al 2012) 59

In contrast to the hormones mentioned above little is known about the role of auxins in 60

induced responses against herbivores Auxins regulate a vast array of plant processes 61

including growth and development as well as responses to light gravity abiotic stress and 62

pathogen attack (Glick 2015 Mano and Nemoto 2012 Yang et al 2014) Several studies 63

suggest that the auxin indole-3-acetic acid (IAA) also regulates gall formation by many 64

herbivores since some gall-forming herbivores contain high levels of IAA (Mapes and 65

Davies 2001b 2001a Tooker and Moraes 2011a Straka et al 2010 Dorchin et al 2009 66

Yamaguchi et al 2012 Tanaka et al 2013) IAA pools and signaling are enhanced in 67

parasitized plant tissue (Yamaguchi et al 2012 Tooker and Moraes 2011b) and direct 68

applications of IAA can result in the formation of gall-resembling structures (Hamner and 69

Kraus 1937 Guiscafrearrillaga 1949 Schaumlller 1968 Bartlett and Connor 2014 Connor et 70

al 2012) In the context of chewing insects however our understanding is more limited 71

(Dafoe et al 2013) IAA levels seem to remain unaltered in Solidago altissima and Triticum 72

aestivum attacked by Heliothis virescens caterpillars (Tooker and Moraes 2011a 2011b) and 73

to be reduced in Helicoverpa zea attacked Zea mays (Schmelz et al 2003) and Manduca 74

sexta-challenged Nicotiana attenuata leaves (Onkokesung et al 2010 Woldemariam et al 75

2012) Moreover mechanical wounding alone can either increase or decrease IAA levels in 76

the leaves (Thornburg and Li 1991 Tanaka and Uritani 1979 Machado et al 2013) A 77

limitation of some of these early studies is that IAA was measured at single time points or 78


5

during the later stages of infestation (Onkokesung et al 2010 Schmelz et al 2003 Tooker 79

and Moraes 2011a 2011b) which may have resulted in an incomplete picture of IAA 80

dynamics under herbivore attack We recently demonstrated in N attenuata that IAA is 81

induced in locally damaged leaves upon simulated M sexta attack (Machado et al 2013) 82

IAA signaling may influence plant responses to herbivore attack by modulating other 83

hormonal pathways and defenses (Erb et al 2012) Exogenous IAA for instance reduces the 84

herbivory-induced accumulation of nicotine and jasmonates (Baldwin et al 1997 Baldwin 85

1989) gene expression of jasmonate-dependent proteinase inhibitors genes (Kernan and 86

Thornburg 1989) and vegetative storage proteins (DeWald et al 1994 Liu et al 2005) 87

Conversely IAA promotes the production of phenolics and flavonoids in root-cell cultures in 88

a dose-dependent manner (Lulu et al 2015 Mahdieh et al 2015) and the auxin homologue 89

24-dichlorophenoxyacetic acid (24-D) acts as a strong inducer of defense responses in rice 90

(Xin et al 2012 Song 2014) 91

In this study we aimed to understand the spatiotemporal patterns of IAA accumulation in 92

herbivore-attacked Nicotiana attenuata plants as well as the role of IAA in regulating the 93

biosynthesis of secondary metabolites In an earlier study we found that IAA accumulates 94

within 1 h following the application of M sexta oral secretions to wounded leaves To 95

understand this pattern in more detail we first evaluated IAA accumulation dynamics in 96

several plant organs in response to real and simulated M sexta attack including the 97

application of a specific herbivore elicitor to wounded leaves at different time points ranging 98

from 15 seconds to 6 h Secondly we analyzed the induction of potential IAA biosynthetic 99

genes Lastly we manipulated IAA accumulation and transport as well as jasmonate 100

signaling to unravel the impact of M sexta-induced IAA on systemic jasmonate-dependent 101

secondary metabolites Our experiments reveal that IAA is a rapid herbivory-induced signal 102

that acts in concert with jasmonates to regulate the systemic induction of plant secondary 103

metabolites104


6

RESULTS 105

Real and simulated M sexta attack induce the accumulation of indole-3-acetic acid 106

(IAA) in the leaves 107

To investigate the behavior of IAA in herbivore-attacked plants we measured IAA 108

concentrations in the leaves of Nicotiana attenuata subjected to either real or simulated M 109

sexta attack (Figure 1A to 1D) We observed a significant increase in IAA levels in response 110

to real M sexta herbivory 3h after infestation This effect could be mimicked by leaf 111

wounding and simultaneous application of either M sexta oral secretions (W+OS) or the fatty 112

acid-amino acid conjugate N-linolenoyl-glutamic acid as a specific herbivore elicitor 113

(W+FAC) (Figure 1A to 1D) Wounding alone led to a delayed and weaker increase in IAA 114

(Figure 1C) The herbivory-induced accumulation of IAA started 30-60 seconds after 115

induction (Figure 1B) and occurred independently of the time of day at which the induction 116

took place (Supplemental Figure 1) Overall IAA concentrations increased 2-3 fold in 117

herbivore induced leaves compared to controls 118

IAA induction gradually spreads through the shoots of attacked plants 119

To explore whether IAA also increases in systemic tissues we induced N attenuata plants 120

and measured IAA concentrations in local treated plant tissues and systemic untreated plant 121

tissues at different time points over a 2 h time period Again we found a rapid increase in 122

IAA levels locally upon simulated M sexta attack (W+OS) which transiently and steadily 123

spread to systemic untreated tissues (Figure 2A to 2F) IAA levels slightly increased in 124

petioles 10 min post treatment in stems 60 min post treatment and in systemic leaves 120 125

min post treatment No significant changes were found in the main and lateral roots (Figure 126

2A to 2F) 127

IAA induction in leaves is conserved across different developmental stages 128

Herbivore-induced jasmonate and ethylene signaling are influenced by plant development 129

(Diezel et al 2011a) To test whether plant development specifically influences M sexta-130

induced IAA levels we induced plants by simulated M sexta attack and measured IAA levels 131

in the leaves of early rosette elongated and flowering plants We found that the herbivore-132

elicited increase in IAA concentration was independent of plant developmental stage (Figure 133

3A to 3C) However the absolute IAA levels and magnitude of induction were strongest in 134

early rosette plants (Figure 3A to 3C) 135


7

YUCCA-like IAA-biosynthesis homologues are rapidly upregulated upon herbivore 136

attack 137

In Arabidopsis thaliana YUCCA-genes encode for flavin monooxygenase-like proteins that 138

convert indole-3-pyruvic acid into IAA a reaction which likely represents the rate-limiting 139

step in IAA biosynthesis (Mashiguchi et al 2011) (Figure 4A) We identified YUCCA-like 140

genes in N attenuata and measured their transcript levels upon herbivore elicitation To 141

achieve this we first searched the sequence of the Arabidopsis thaliana YUCCA2 gene 142

(NCBI accession number NM_1173993) in N attenuata draft genome (Ling et al 2015) and 143

reconstructed the phylogenetic tree of the gene family (Mashiguchi et al 2011) Our analysis 144

revealed that the N attenuata genome contains at least nine YUCCA-like genes that share 145

high similarity with AtYUCCA2 and contain the four conserved amino acid motifs 146

characteristic of this gene family (Supplemental Figure 2) (Expoacutesito-Rodriacuteguez et al 2011 147

Expoacutesito-Rodriacuteguez et al 2007) We designed specific primers and profiled the expression 148

patterns of these genes upon simulated M sexta attack Several YUCCA-like genes were 149

upregulated in response to simulated M sexta attack (Figure 4B to 4I) NaYUCCA-like 1 3 150

5 6 and 9 were upregulated 3 min after the application of M sexta oral secretions and fatty 151

acid-conjugates (Figure 4B to 4H) The upregulation of NaYUCCA-like 1 and 3 was 152

maintained for at least one hour (Figure 4G to 4H) The expression of NaYUCCA-like 2 4 7 153

and 8 was not significantly influenced by simulated M sexta attack (Supplemental Figure 3) 154

IAA accumulation precedes the JA burst 155

To investigate the temporal dynamics of IAA and JA accumulation in M sexta-attacked 156

plants we quantified IAA and JA in plants subjected to simulated M sexta herbivory at 157

different time points We found that IAA peaked more rapidly than jasmonic acid in response 158

to herbivore attack (Figure 5) IAA accumulation commenced within minutes after the onset 159

of the elicitation and reached its maximum five minutes after induction JA accumulated in an 160

equally rapid fashion but peaked significantly later than IAA (Figure 5) 161


8

Jasmonate signaling is not required for the M sexta-induced IAA accumulation 162

Plant responses to attackers are modulated by a complex signaling network consisting of 163

antagonistic neutral and synergistic effects (Erb et al 2012) For example jasmonate 164

signaling antagonizes IAA signaling (Chen et al 2011) To further explore the potential 165

crosstalk between these two phytohormones we measured M sexta-induced IAA in 166

transgenic plants that are impaired to different degrees in jasmonate signaling biosynthesis 167

andor perception (Table 1) We found that the M sexta-triggered accumulation of IAA does 168

not require JA signaling as it was induced in all of the evaluated JA-deficient genotypes 169

(Figure 6 and supplemental Figure 4) 170

M sexta-induced IAA is required for the induction of anthocyanins in the stems 171

To investigate the impact of IAA on plant secondary metabolites we sought to manipulate its 172

perception in planta Our initial attempts to create transgenic dexamethasone (DEX) 173

inducible plants (Schaumlfer et al 2013) harboring a silencing construct for the IAA receptor 174

TIR1 failed either because of promotor methylation in the F2 crosses (Weinhold et al 2013) 175

or because the identified TIR1 homologue was inactive We therefore took advantage of our 176

knowledge on systemic IAA accumulation to devise a series of chemical manipulation 177

experiments First we exogenously applied IAA and MeJA at doses that exceed endogenous 178

levels (Baldwin 1989 Machado et al 2013) Second we inhibited local IAA synthesis with 179

L-kynurenine (L-Kyn) L-kynurenine is a specific inhibitor of tryptophan aminotransferases 180

(TATs) which are key enzymes of the indole-3-pyruvic acid pathway that leads to IAA 181

formation (He et al 2011) Third we inhibited IAA transport at the leaf base and petiole of 182

the induced leaves using 235-triiodobenzoic acid (TIBA) TIBA inhibits auxin polar 183

transport by blocking auxin efflux transporter PIN-FORMED PIN1 cycling (Geldner et al 184

2001) We observed that within hours following M sexta attack N attenuata stems became 185

red (Figure 7D inset) a phenotype that is likely due to anthocyanin accumulation As IAA 186

can regulate the production of anthocyanins in plants (Pasqua et al 2005) we quantitatively 187

and qualitatively evaluated anthocyanin accumulation in the stems following several 188

simulated and real herbivory in combination with IAA manipulation We observed that the 189

levels of anthocyanins in the stems were strongly induced by real M sexta attack an effect 190

that could be mimicked by wounding and applications of M sexta oral secretions (W+OS) 191

but not by wounding alone (W+W) (Figure 7A) Application of IAA or MeJA alone did not 192

trigger anthocyanin accumulation (Figure 7A) By contrast the simultaneous application of 193


9

IAA and MeJA (IAA+MeJA) triggered anthocyanin accumulation (Figure 7A) Chemical 194

inhibition of IAA biosynthesis or transport as well as genetic inhibition of JA biosynthesis led 195

to the complete disappearance of induced anthocyanin accumulation (Figure 7B and 7C) 196

Furthermore we found a positive correlation between anthocyanin contents and red 197

pigmentation in the stems (Figure 7D) 198

IAA specifically potentiates the herbivore-induced accumulation of phenolamides in the 199

stems 200

To investigate the role of IAA in the accumulation of known defensive metabolites in the 201

stems of N attenuata (Onkokesung et al 2012 Heiling et al 2010 Paschold et al 2007) 202

we induced leaves of N attenuata plants by different simulated and real herbivory treatments 203

and complemented them with IAA at doses that exceed endogenous levels (Baldwin 1989 204

Machado et al 2013) The stems of N attenuata are often attacked by herbivores including 205

stem borers (Diezel et al 2011b Lee et al 2016) and are very important for plant fitness 206

(Machado et al 2016) We observed a strong upregulation of defensive secondary 207

metabolites in the stems in response to M sexta attack (Figure 8A to 8D) Petiole 208

pretreatments with IAA dramatically increased the accumulation of caffeoylputrescine and 209

dicaffeoylspermidine in response to real and simulated herbivory as well as MeJA 210

application IAA application alone did not induce the metabolites (Figure 8A and 8B) By 211

contrast nicotine and 7-hydroxygeranyllinalool diterpene glycosides did not respond to IAA 212

petiole pretreatments (Figure 8A to 8D) 213


10

DISCUSSION 214

In this study we show that auxin is a rapidly and specifically induced regulator of defensive 215

secondary metabolites in Nicotiana attenuata Infestation by M sexta caterpillars induced the 216

accumulation of IAA levels in local tissues an effect that could be mimicked by both the 217

applications of M sexta oral secretions and the application of the well-known insect elicitor 218

N-linolenoyl-glutamic acid (Halitschke et al 2003) and to a lesser extent by mechanical 219

wounding These results are in contrast to earlier studies in maize goldenrod and coyote 220

tobacco which found either a slight decrease or no changes in IAA levels in response to 221

herbivore attack (Schmelz et al 2003 Tooker and Moraes 2011a Onkokesung et al 2010 222

Tooker and Moraes 2011b) but are in agreement with our previous study (Machado et al 223

2013) Interestingly in comparison with our previous study we observed differences in both 224

absolute quantities and timing of IAA induction One possible explanation for these 225

differences is that plants were grown using different substrates While sand was used in the 226

previous study potting soil was used in the present paper Given the strong feedback effects 227

of soil bacteria soil nutrients and root growth on IAA signaling (Lambrecht et al 2000 228

Kurepin et al 2015 Tian et al 2008 Sassi et al 2012) it is likely that the growth substrate 229

affected IAA homeostasis and responsiveness in N attenuata On the other hand the absence 230

of IAA induction reported in earlier studies may be due to the fact that late time points were 231

measured (Onkokesung et al 2010 Schmelz et al 2003 Tooker and Moraes 2011a) which 232

may not have captured the rapid and dynamic accumulation of IAA following herbivore 233

attack To further investigate these contradicting results we determined IAA responses in 234

herbivore attacked maize plants (Maag et al submitted) We found that IAA levels increased 235

in an herbivore-specific manner 1-6 h after the onset of the attack Together these 236

experiments suggest that the rapid and transient herbivory-induced accumulation of IAA may 237

be a conserved plant response to insect attack 238

Spatiotemporal IAA profiling revealed that the rapid increase in IAA pools at the site of 239

attack is followed by a weak and transient increase in auxin pools in systemic tissues Similar 240

to what has been observed for other phytohormones (Koo et al 2009 Stitz et al 2011 241

VanDoorn et al 2011) IAA levels increased sequentially in petioles stems and systemic 242

leaves Together with the rapid local induction of YUCCA-like IAA biosynthetic homologues 243

and the absence of IAA dependent systemic defense induction in transport inhibitor treated 244

plants these data suggest that IAA might be synthesized de novo at the site of the attack and 245

then transported across the plant Several studies have demonstrated that auxin is a mobile 246


11

signal in plants (Reed et al 1998 Bhalerao et al 2002 Jin et al 2015 van Noorden et al 247

2006) Based on the IAA accumulation kinetics we estimate that herbivory-induced IAA 248

would need to be transported at a speed of at least 029 cmmin-1 to reach the petioles 5-10 249

minutes after elicitation (based on the fact that IAA accumulates locally 30-60 seconds after 250

elicitation) This value is at least tenfold greater than typical values of polar auxin transport 251

velocities (Kramer et al 2011) but twenty fold slower than wound-induced electrical signals 252

that trigger systemic JA accumulation (Mousavi et al 2013) We propose two hypotheses 253

that may be responsible for the atypical signal propagation speed that we observed First it is 254

possible that IAA is transported to systemic tissues by a combination of both polar and non-255

polar phloem-based transport (Friml 2003) Second rapid secondary signals including 256

electrical potentials may spread through the plant at high speeds and induce de novo IAA 257

biosynthesis in systemic tissues Further experiments with IAA radiotracers (Agtuca et al 258

2014) and transient tissue-specific deactivation of IAA biosynthesis (Koo et al 2009) would 259

help to shed further light on the exact mechanisms responsible for the systemic spread of IAA 260

following herbivore attack 261

Impairing key genes of the jasmonate signaling cascade including mitogen-activated protein 262

kinases jasmonate biosynthesis and jasmonate perception elements did not impair the 263

herbivory-induced accumulation of IAA suggesting that IAA induction does not require JA 264

signaling This observation is consistent with the temporal dynamics of herbivory-induced 265

IAA and JA that we observed IAA accumulation peaks within 5 minutes after the onset of 266

the elicitation while JA starts accumulating in an equally rapid fashion but peaks 267

significantly later than IAA (Figure 5) 268

An important aim of our study was to understand whether IAA is involved in the regulation 269

of induced secondary metabolites in N attenuata Because of the systemic accumulation 270

pattern of IAA and the possibility to block this effect through the local application of 271

transport inhibitors we chose to focus on the induction of stem secondary metabolites The 272

stem of N attenuata is vital for its reproduction and can be attacked by a wide variety of 273

organisms including vertebrates and invertebrate stem borers (Machado et al 2016 Diezel 274

et al 2011b) We observed that real and simulated M sexta attack induced anthocyanin 275

accumulation in the stems an effect that could not be reproduced by MeJA or IAA treatments 276

alone but by the combination of these two hormones Together with the IAA transport and 277

biosynthesis inhibitor treatments and the genetic silencing of JA biosynthesis all of which led 278

to the disappearance of the anthocyanin response these results strongly suggest that IAA is 279


12

required to activate the JA-dependent accumulation of stem anthocyanins In A thaliana 280

anthocyanin production is controlled by the MYB75 transcription factor Production of 281

Anthocyanin Pigment 1 (PAP1) (Shin et al 2015 Borevitz et al 2000) which is 282

transcriptionally upregulated by IAA (Lewis et al 2011) and postranscriptionally repressed 283

by jasmonate-ZIM-Domain (JAZ) proteins (Qi et al 2011) The resulting co-regulation of 284

MYB transcription factors by IAA and JA provides a potential mechanism for the synergistic 285

interaction between JA and IAA observed in our study 286

In a second set of experiments we found that IAA also boosts the production of 287

phenolamides in herbivore-attacked plants Phenolamide accumulation in N attenuata is 288

controlled by the transcription factor MYB8 in a JA-dependent manner (Onkokesung et al 289

2012 Paschold et al 2007) This transcription factor may therefore represent a target for the 290

integration of IAA and JA signaling While IAA strongly potentiated the accumulation of 291

stem phenolamides it had little effect on the accumulation of other JA-dependent secondary 292

metabolites including nicotine and 7-hydroxygeranyllinalool diterpene glycosides (Machado 293

et al 2013 Paschold et al 2007 Jimenez-Aleman et al 2015 Machado et al 2016) This 294

result is consistent with earlier studies showing neutral to negative effects of auxin 295

application on nicotine accumulation in Nicotiana spp (Baldwin 1989 Baldwin et al 1997 296

Shi et al 2006) The direct application of IAA to wounded tissues can even suppress local 297

damage-induced JA accumulation (Dahl and Baldwin 2004 Baldwin et al 1997 Shi et al 298

2006) From these results it is evident that IAA does not simply enhance JA signaling but 299

that it specifically modulates a plantrsquos defensive network Thereby IAA signaling may help 300

plants to mount specific fine-tuned responses to different attackers 301

The ecological function of an upregulation of anthocyanin and phenolamide compounds in 302

the stems upon M sexta attack remains an open question The current literature however 303

provides interesting insights in this context Trichobaris stem weevils prefer to feed and 304

perform better on defenseless jasmonate-deficient plants in a species-specific manner T 305

compacta grows better on nicotine-impaired N attenuata plants while T mucorea is not 306

affected by nicotine but by other yet unknown jasmonate-dependent defenses (Diezel et al 307

2011b Lee et al 2016) It is therefore possible that the IAA-triggered potentiation of 308

jasmonate-dependent secondary metabolite accumulation in the stems may reduce the 309

performance of stem feeders To disentangle the specific effects that IAA signaling has in this 310

context requires the development of IAA-signaling impaired genotypes and represents an 311

interesting prospect of this study 312


13

In conclusion this study identifies IAA as a rapid and specific signal that regulates a 313

biologically relevant subset of herbivory-induced secondary metabolites Current models on 314

plant defense signaling networks in plant-herbivore interactions can now be expanded to 315

include auxins as potentially important defense hormones 316

METHODS 317

Plant genotypes germination and planting conditions 318

Wild-type N attenuata Torr Ex Watson plants of the 31th inbred generation derived from 319

seeds collected at the Desert Inn Ranch in Utah in 1988 and all genetically engineered plant 320

genotypes were germinated on Gamborgrsquos B5 medium as described (Kruumlgel et al 2002) 321

Nine to ten days later seedlings were transferred to Teku pots (Poumlppelmann GmbH amp Co 322

KG Lohne Germany) for 10-12 days before transferring them into 1 L pots filled with either 323

sand (to facilitate the harvesting of belowground tissues) or soil All plants were grown at 45-324

55 relative humidity and 23-25 degC during days and 19-23 degC during nights under 16 h of 325

light (6am-10pm) Plants planted in soil were watered every day by a flood irrigation system 326

Plants planted in sand were watered twice a day The characteristics of the transgenic plants 327

used in this study are presented in table 1 328

Auxin and jasmonate measurements 329

Phytohormone measurements were conducted as described earlier (Machado et al 2013 330

Machado et al 2015) Briefly plant tissues were harvested flash frozen and stored at -80degC 331

After grinding 100 mg of plant tissue per sample were extracted with 1 mL ethyl acetate 332

formic acid (99505 vv) containing the following phytohormone standards 40ng of 910-333

D2-910-dihydrojasmonic acid (JA) 8 ng of jasmonic acid-[13C6] isoleucine (JA-Ile) and 20 334

ng of D5-indole-3-acetic-acid (IAA) All samples were then vortexed for 10 min and 335

centrifuged at 14000 rpm for 20 min at 4 degC Supernatants were evaporated to dryness in a 336

centrifugal vacuum concentrator (Eppendorf 5301 Eppendorf Hamburg Germany) at room 337

temperature The remaining pellets were resuspended in 50 μL methanol water (7030) and 338

dissolved using an ultrasonic cleaner (Branson 1210 Branson Ultrasonics 339

Danbury Connecticut USA) for 5 min Samples were then analyzed using liquid 340

chromatography (Agilent 1260 Infinity Quaternary LC system Agilent Technologies Santa 341

Clara California USA) coupled to a triple quadrupole mass spectrometer (API 5000 342

LCMSMS Applied Biosystems Foster City California USA) 343


14

IAA levels in herbivore attacked plants 344

IAA levels were determined in local treated leaves of plant subjected to real or simulated M 345

sexta attack Plants were infested by placing 3 first-instar larvae on one fully developed 346

rosette leaf (n=3) Caterpillars were removed and attacked leaves were harvested M sexta 347

attack was simulated by rolling a pattern wheel over the leaves on each side of the midvein 348

Three fully developed rosette leaves were wounded and the resulting wounds were 349

immediately treated with either 15 (vv) water-diluted M sexta oral secretions (W+OS) with 350

pure water (W+W) or with fatty acid-amino acid conjugates (FACs N-linolenoyl-glutamic 351

acid) as described (Xu et al 2015 Machado et al 2013) Intact plants were used as controls 352

(n=5) 353

M sexta-induced auxin levels in different plant tissues 354

Forty-day-old elongating plants were subjected to simulated M sexta attack as described 355

above Five 10 30 60 and 120 min after elicitation treated leaves and their untreated 356

petioles as well as stems systemic leaves (young leaves directly above treated leaves) and 357

main and lateral roots were harvested The same plant tissues were collected from untreated 358

control plants at each time point (n=5) 359

M sexta-induced auxin levels at different developmental stages 360

IAA levels were measured at three developmental stages early rosette (32 days after 361

germination DAG) elongating (39 DAG) and flowering (46 DAG) Tissues were harvested 362

at three time points after elicitation as described above 05 1 and 3h (n=5) 363

Identification and expression profiling of YUCCA-like genes 364

YUCCA genes encode for flavin monooxygenase-like proteins that convert indole-3-pyruvic 365

acid into indole-3-acetic acid (IAA) a catalytic reaction that is currently seen as the limiting 366

step of IAA biosynthesis (Mashiguchi et al 2011) To identify YUCCA-like genes in N 367

attenuata we searched the Arabidopsis thaliana YUCCA2 gene sequence (NCBI accession 368

number NM_1173993) in the N attenuata draft genome (Ling et al 2015) using BLAST (E-369

valuelt1e-10 bit scoregt200) and reconstructed the phylogenetic tree of the gene family We 370

then designed specific primers (Supplemental Table 1) for each gene using Primique 371

(Fredslund and Lange 2007) and profiled gene expression patterns upon simulated M sexta 372

attack by quantitative real-time PCR (qPCR)(n=3) Total RNA was extracted by the TRIZOL 373


15

method followed by DNase-I treatment (Fermentas St Leon-Rot Germany) according to 374

the manufacturerrsquos instructions Five micrograms of total RNA were reverse-transcribed 375

using oligo (dT)18 and the SuperScript-II Reverse Transcriptase kit (Invitrogen) The 376

obtained cDNA was used for gene expression profiling with SYBR Green I following the 377

manufacturerrsquos protocol and the ∆Ct method was used for transcript evaluation The 378

housekeeping gene actin was used as reference Gene expression levels were determined 3 5 379

and 60 minutes after elicitation 380

Characterization of the YUCCA-like gene family 381

The YUCCA-like gene family sequences were aligned by Clustal W (Thompson et al 1994) 382

in BioEdit (Hall 1999) and the occurrence of the already described conserved amino acid 383

motifs characteristic of the flavin monooxygenase gene family was determined (Expoacutesito-384

Rodriacuteguez et al 2011 Expoacutesito-Rodriacuteguez et al 2007) 385

OS-induced auxin and jasmonate kinetics 386

Rosette leaves of wild type plants were subjected to simulated M sexta attack (W+OS) as 387

described and harvested 5 45 and 90 min after elicitation (n=5) Phytohormone 388

measurements were carried out as described 389

M sexta-induced auxin levels in jasmonate and signaling impaired genotypes 390

Three rosette leaves of rosette-stage plant genotypes impaired in salicylic acid-induced and 391

wound-induced mitogen-activated protein kinases (irSIPK irWIPK respectively) jasmonic 392

acid biosynthesis (irGLA irAOS irAOC irOPR3) jasmonic acid-isoleucine biosynthesis 393

(irJAR46) jasmonate perception (irCOI1) and wild type empty vector (EV) were subjected 394

to M sexta simulated attack as described 45 min after elicitation the leaves were harvested 395

and analyzed for IAA jasmonic acid (JA) and jasmonic acid-isoleucine (JA-Ile) (n=5) These 396

transgenic plant genotypes were selected as they are impaired at different layers of the 397

jasmonate signaling cascade early regulatory elements (irSIPK irWIPK) jasmonate 398

biosynthesis (irGLA irAOS irAOC irOPR3) hormone activation (irJAR46) and hormone 399

perception (irCOI1) and their main characteristics are listed in table 1 400

Stem anthocyanin quantifications 401

To determine the role of IAA in M sexta induced stem anthocyanin accumulation we carried 402

out three experiments First we measured anthocyanins in the stem of plants whose rosette 403


16

leaves were either left intact (Control) wounded and treated with water (W+W) wounded 404

and treated with M sexta oral secretions (W+OS) subjected to real continuous M sexta 405

attack (M sexta) treated with the natural auxin IAA (IAA) methyl jasmonic acid (MeJA) or 406

with both IAA and MeJA (IAA+MeJA) dissolved in lanoline paste (n=5) Simulated M sexta 407

attack treatments were carried out as described above Hormonal treatments were carried out 408

as described below In the second experiment we measured stem anthocyanins in plants 409

whose petioles were treated (petiole pretreatment) with the IAA biosynthesis inhibitor L-410

kynurenine (L-Kyn) (He et al 2011) the IAA transport inhibitor 235-triiodobenzoic acid 411

(TIBA) (Hertel et al 1983 Goldsmith 1982 Rubery 1979) or with the natural auxin indole-412

3-acetic acid (IAA) prior to eliciting the plants by simulated M sexta attack (W+OS) (n=12) 413

One hour prior to the simulated M sexta attack treatments approximately 2 microg of L-Kyn 414

TIBA or IAA or 150 microg MeJA dissolved in lanoline paste were applied to the petioles 415

Applied doses were selected according to previous studies (Baldwin 1989 Morris et al 416

1973 Kang et al 2006 He et al 2011) (n=12) In a third experiment we measured changes 417

in stem anthocyanin levels upon simulated M sexta herbivory in jasmonate-deficient irAOC 418

and empty vector (EV) controls (n=10) Simulated and real M sexta attack treatments were 419

carried out as described For all the experiments the stems were harvested five days after 420

treatments and the anthocyanin content of the outer layer (epidermis cortex phloem and 421

cambium) was determined 5 cm above the shoot-root junction as described (Steppuhn et al 422

2010) 423

Stem secondary metabolite quantifications 424

To further explore the regulatory role of IAA in secondary metabolite production we induced 425

the leaves of N attenuata plants using real and simulated M sexta attack treatments Plants 426

were either pretreated with IAA in lanolin paste or with pure lanolin as controls as described 427

above Petiole pretreatments with IAA were carried out one hour prior to induction Five days 428

after induction the stems were harvested and secondary metabolites were measured as 429

described (Gaquerel et al 2010 Ferrieri et al 2015)(n=5) 430

Statistics 431

All data were analyzed by ANOVA using Sigma Plot 120 (Systat Software Inc San Jose 432

CA USA) Normality and equality of variance were verified using ShapirondashWilk and 433

Levenersquos tests respectively HolmndashSidak post hoc tests were used for multiple comparisons 434

Datasets from experiments that did not fulfill the assumptions for ANOVA were natural log- 435


17

root square- or rank-transformed before analysis Correlation between jasmonate and IAA 436

levels and stem coloration index and stem anthocyanin content were evaluated by Pearson 437

product moment test 438

ACKNOWLEDGEMENTS 439

All experimental work of this study was supported by the Max Planck Society We would 440

also like to thank the members of the Department of Molecular Ecology and the glasshouse 441

team of the MPI-CE for their help Special thanks go to Mareike Schirmer and Mareike 442

Schmidt for technical support and to Wenwu Zhou Martin Schaumlfer and Michael Reichelt for 443

their valuable help with the auxin measurements CAMR was supported by a Swiss National 444

Foundation Fellowship (grant no 140196) CCMA by the Brazilian National Council for 445

Research (grant no 2379292012-0) APF by an Alexander von Humboldt Postdoctoral 446

Fellowship SX by a Marie Curie Intra European Fellowship (grant no 328935) ITB by a 447

European Research Council advanced (grant no 293926) and by a Human Frontier Science 448

Program (grant no RGP00022012) and ME by an SNF early post doc fellowship (grant no 449

134930) and a Marie Curie Intra European Fellowship (grant no 273107) 450

AUTHOR CONTRIBUTIONS 451

Designed the research RARM ME ITB Carried out the experimental work RARM 452

CCMA APF CAMR GHJA SX Analyzed data RARM ME ITB Wrote the first draft of 453

the paper RARM ME Revised the paper ME RARM ITB APF CCMA GHJA SX 454

CAMR All authors read and approved the final manuscript 455

456


18

Table 1 Characteristics of the inverted repeat (ir) transgenic lines used in the present study 457

Genotype Gene silencedoverexpressed

Impaired function Phenotype Reference

irSIPK Salicylic acid-induced

mitogen activated protein kinase Early

jasmonate signalling

Reduced levels of jasmonates

Meldau et al 2009

irWIPK Wound-induced

mitogen activated protein kinase

irGLA1 Glycerolipase A1

Jasmonate biosynthesis

Bonaventure et al 2011

irAOS Allene oxide synthase

Kallenbach et al 2012 irAOC Allene oxide cyclase

irOPR3 12-oxo-phytodienoic acid reductase

irJAR46 JA-Ile synthetase Reduced levels of JA-Ile

Wang et al 2008

irCOI1 Coronatine-insensitive 1 JA-Ile perception

Reduced JA-Ile perception

Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465

Figure 1 Indole-3-acetic acid (IAA) is induced specifically and rapidly by real and simulated 466

M sexta attack Average (plusmnSE) IAA levels in leaves that are attacked by M sexta caterpillars 467

(A) treated with M sexta oral secretions (B C) or treated with an herbivore elicitor (D) 468

(n=5) Different letters indicate significant differences between treatments (P lt 005) 469

Control intact plants W+W wounded and water-treated plants W+OS wounded and M 470

sexta oral secretion-treated plants W+FACs wounded and fatty acid-amino acid conjugate-471

treated plants 472


19

Figure 2 Herbivory induces IAA both locally and systemically Average (plusmnSE) IAA levels 473

following simulated M sexta attack in local treated leaves (A) and in untreated petioles (B) 474

stem (C) systemic leaves (D) main root (E) and lateral roots (F) (n=5) Asterisks indicate 475

significant differences between treatments within plant tissues and time points ( P lt 005 476

P lt 0001) Control intact plants W+OS wounded and M sexta oral secretion-treated 477

plants 478

Figure 3 IAA induction in leaves occurs across different developmental stages Average 479

(plusmnSE) IAA levels in local treated leaves following simulated M sexta attack at the early 480

rosette (A) elongated (B) and flowering stage (C) (n=5) Different letters indicate significant 481

differences between treatments within developmental stages and time points (P lt 005) 482


sexta oral secretion-treated plants 484

Figure 4 YUCCA-like genes are upregulated in response to simulated M sexta herbivory 485

(A) Schematic representation of YUCCA-mediated conversion of indole-3-pyruvic acid into 486

IAA Average (plusmnSE) transcript abundance relative to control of YUCCA-like 3 (B) YUCCA-487

like 5 (C) YUCCA-like 6 (D) and YUCCA-like 9 (E) in treated leaves three minutes after 488

elicitation and YUCCA-like 1 (F) and YUCCA-like 3 (G) 5 and 60 min following simulated 489

M sexta attack (n=3) Different letters indicate significant differences between treatments (P 490

lt 005) Control intact plants W+W wounded and water-treated plants W+OS wounded 491

and M sexta oral secretion-treated plants W+FACs wounded and fatty acid-amino acid 492

conjugate-treated plants 493

Figure 5 Manduca sexta-induced IAA peaks earlier than jasmonic acid (JA) Left Y-axis 494

average (plusmnSE) leaf IAA levels in response to M sexta attack Right Y-axis average (plusmnSE) 495

leaf JA levels in response to M sexta attack Closed squares IAA levels upon W+OS 496

treatments closed triangles IAA levels in control untreated plants Grey squares JA levels 497

upon W+OS treatments grey triangles jasmonic acid (JA) levels in control untreated plants 498

(n=5) Different letters indicate significant differences between treatments for individual 499

metabolites (P lt 005) IAA Time P = 0015 treatment P lt 0001 Time treatment P = 500

0638 JA Time P lt 0001 treatment P lt 0001 Time treatment P lt 0001) Control 501

intact plants W+OS wounded and M sexta oral secretion-treated plants 502

Figure 6 Jasmonate signaling is not required for the M sexta-induced accumulation of IAA 503

(A) Average (plusmnSE) IAA levels in local treated leaves of wild type plants (empty vector EV) 504


20

and plant genotypes impaired in early JA signaling jasmonate biosynthesis andor JA-Ile 505

perception 45 minutes after elicitation (n=5) Different letters indicate significant differences 506

between treatments within each genotype (P lt 005) C control intact plants W wounded 507

and water-treated plants OS wounded and M sexta oral secretions-treated plants 508

Figure 7 Manduca sexta-induced IAA and JA act synergistically to trigger anthocyanin 509

accumulation in the stems (A) Average (plusmnSE) stem anthocyanin content 5 days following 510

either simulated or continuous M sexta attack exogenous application of methyl jasmonate 511

(MeJA) andor IAA (n=5) (B) Average (plusmnSE) stem anthocyanin content 5 days following 512

simulated M sexta attack and petiole-pretreatments with either IAA the IAA biosynthesis 513

inhibitor L-kynurenine (L-Kyn) or the IAA transport inhibitor TIBA (235-triiodobenzoic 514

acid) (n=12) (C) Average (plusmnSE) stem anthocyanin contents following simulated M sexta 515

attack of wild type and JA-impaired irAOC plants (n=10) (D) Correlation between stem 516

anthocyanin content and stem coloration Inset Photograph of the red stem phenotype 517

Asterisks indicate significant differences between treatments and control (A) between 518

simulated herbivory treatments within petiole pretreatments (B) and between treatments 519

within genotypes (C) ( P lt 005 P lt 001 P lt 0001) The correlation between 520

stem coloration index and stem anthocyanin content was evaluated by a Pearson product 521

moment test Leaf treatments Control intact plants W+W wounded and water-treated 522

plants W+OS wounded and M sexta oral secretion-treated plants M sexta plants subjected 523

to actual M sexta attack IAA rosette leaves treated with indole-3-acetic acid MeJA rosette 524

leaves treated with methyl jasmonic acid IAA+MeJA rosette leaves treated with IAA and 525

MeJA Petiole pretreatments Petioles treated with either pure lanoline paste (Lanoline) L-526

kynurenine (L-Kyn) 235-triiodobenzoic acid (TIBA) or indole-3-acetic acid (IAA) 527

dissolved in lanoline 1h prior to leaf treatments 528

Figure 8 IAA specifically potentiates the herbivore-induced systemic production of 529

phenolamides Average (plusmnSE) caffeoylputrescine (A) dicaffeoylspermidine (B) nicotine (C) 530

and diterpene glycoside (D) levels in the stems 5 days following simulated or real M sexta 531

attack and petiole pretreatments with IAA (n=5) Asterisks indicate significant differences 532

between petiole pretreatments within simulated M sexta attack treatments ( P lt 005 P 533

lt 001 P lt 0001) Petiole pretreatments Control petioles treated with pure lanoline 534

paste 1h prior to leaf treatments IAA petioles treated with IAA dissolved in lanoline 1h prior 535

to leaf treatments Leaf treatments Control intact plants W+W wounded and water-treated 536



21

to actual M sexta attack MeJA rosette leaves treated with methyl jasmonic acid dissolved in 538

lanoline paste 539

SUPPLEMENTAL DATA 540

Supplemental Figure 1 IAA is induced locally in response to simulated M sexta herbivory 541

independently of time of day 542

Supplemental Figure 2 The N attenuata genome contains nine YUCCA-like genes 543

Supplemental Figure 3 Gene expression patterns of YUCCA-like genes upon simulated M 544

sexta attack 545

Supplemental Figure 4 Jasmonate signaling is not required for the M sexta-induced 546

accumulation of IAA 547

Supplemental Table 1 Sequence of primers used for quantitative PCR analysis 548

549


22

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root growth in Zea mays Journal of plant growth regulation 33 (2) 328ndash339 554

Baldwin IT (1989) Mechanism of damage-induced alkaloid production in wild tobacco 555

Journal of Chemical Ecology 15 (5) 1661ndash1680 556

Baldwin IT Zhang Z-P Diab N Ohnmeiss TE McCloud ES Lynds GY and 557

Schmelz EA (1997) Quantification correlations and manipulations of wound-induced 558

changes in jasmonic acid and nicotine in Nicotiana sylvestris Planta 201 (4) 397ndash404 559

Bartlett L and Connor EF (2014) Exogenous phytohormones and the induction of plant 560

galls by insects Arthropod-Plant Interactions 8 (4) 339ndash348 561

Bhalerao RP Ekloumlf J Ljung K Marchant A Bennett M and Sandberg G 562

(2002) Shoot‐derived auxin is essential for early lateral root emergence in Arabidopsis 563

seedlings The Plant Journal 29 (3) 325ndash332 564

Bonaventure G Schuck S and Baldwin IT (2011) Revealing complexity and 565

specificity in the activation of lipase‐mediated oxylipin biosynthesis a specific role of the 566

Nicotiana attenuata GLA1 lipase in the activation of jasmonic acid biosynthesis in leaves 567

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Borevitz JO Xia Y Blount J Dixon RA and Lamb C (2000) Activation tagging 569

identifies a conserved MYB regulator of phenylpropanoid biosynthesis The Plant Cell 12 570

(12) 2383ndash2393 571

Chen Q Sun J Zhai Q Zhou W Qi L Xu L Wang B Chen R Jiang H and 572

Qi J (2011) The basic helix-loop-helix transcription factor MYC2 directly represses 573

PLETHORA expression during jasmonate-mediated modulation of the root stem cell 574

niche in Arabidopsis The Plant Cell 23 (9) 3335ndash3352 575

Connor EF Bartlett L OrsquoToole S Byrd S Biskar K and Orozco J (2012) The 576

mechanism of gall induction makes galls red Arthropod-Plant Interactions 6 (4) 489ndash577

495 578

Dafoe NJ Thomas JD Shirk PD Legaspi ME Vaughan MM Huffaker A 579

Teal PE and Schmelz EA (2013) European corn borer (Ostrinia nubilalis) induced 580

responses enhance susceptibility in maize PloS one 8 (9) 581


23

Dahl CC von and Baldwin IT (2004) Methyl jasmonate and cis‐jasmone do not dispose 582

of the herbivore‐induced jasmonate burst in Nicotiana attenuata Physiologia Plantarum 583

120 (3) 474ndash481 584

Dahl CC von Winz RA Halitschke R Kuumlhnemann F Gase K and Baldwin IT 585

(2007) Tuning the herbivore‐induced ethylene burst the role of transcript accumulation 586

and ethylene perception in Nicotiana attenuata The Plant Journal 51 (2) 293ndash307 587

DeWald DB Sadka A and Mullet JE (1994) Sucrose modulation of soybean Vsp 588

gene expression is inhibited by auxin Plant Physiology 104 (2) 439ndash444 589

Diezel C Allmann S and Baldwin IT (2011a) Mechanisms of optimal defense patterns 590

in Nicotiana attenuata Flowering attenuates herbivory‐elicited ethylene and jasmonate 591

Signaling Journal of integrative plant biology 53 (12) 971ndash983 592

Diezel C Kessler D and Baldwin IT (2011b) Pithy protection Nicotiana attenuatarsquos 593

jasmonic acid-mediated defenses are required to resist stem-boring weevil larvae Plant 594

Physiology 155 (4) 1936ndash1946 595

Dorchin N Hoffmann JH Stirk WA NOVAacuteK O Strnad M and van Staden J 596

(2009) Sexually dimorphic gall structures correspond to differential phytohormone 597

contents in male and female wasp larvae Physiological Entomology 34 (4) 359ndash369 598

Erb M Meldau S and Howe GA (2012) Role of phytohormones in insect-specific 599

plant reactions Trends in plant science 17 (5) 250ndash259 600

Expoacutesito-Rodriacuteguez M Borges AA Borges-Peacuterez A Hernaacutendez M and Peacuterez JA 601

(2007) Cloning and biochemical characterization of ToFZY a tomato gene encoding a 602

flavin monooxygenase involved in a tryptophan-dependent auxin biosynthesis pathway 603

Journal of plant growth regulation 26 (4) 329ndash340 604

Expoacutesito-Rodriacuteguez M Borges AA Borges-Peacuterez A and Peacuterez JA (2011) Gene 605

structure and spatiotemporal expression profile of tomato genes encoding YUCCA-like 606

flavin monooxygenases the ToFZY gene family Plant Physiology and Biochemistry 49 607

(7) 782ndash791 608

Ferrieri AP Arce C Machado RAR Meza‐Canales ID Lima E Baldwin IT 609

and Erb M (2015) A Nicotiana attenuata cell wall invertase inhibitor (NaCWII) 610

reduces growth and increases secondary metabolite biosynthesis in herbivore‐attacked 611

plants New Phytologist 612

Fredslund J and Lange M (2007) Primique automatic design of specific PCR primers 613

for each sequence in a family BMC bioinformatics 8 (1) 369 614


24

Friml J (2003) Auxin transportmdashshaping the plant Current opinion in plant biology 6 (1) 615

7ndash12 616

Gaquerel E Heiling S Schoumlttner M Zurek G and Baldwin IT (2010) 617

Development and validation of a liquid chromatographyminus electrospray ionizationminus time-618

of-flight mass spectrometry method for induced changes in Nicotiana attenuata leaves 619

during simulated herbivory Journal of Agricultural and Food Chemistry 58 (17) 9418ndash620

9427 621

Geldner N Friml J Stierhof Y-D Juumlrgens G and Palme K (2001) Auxin transport 622

inhibitors block PIN1 cycling and vesicle trafficking Nature 413 (6854) 425ndash428 623

Geyter N de Gholami A Goormachtig S and Goossens A (2012) Transcriptional 624

machineries in jasmonate-elicited plant secondary metabolism Trends in plant science 17 625

(6) 349ndash359 626

Glick BR (2015) Beneficial Plant-bacterial Interactions (Springer) 627

Goldsmith MHM (1982) A saturable site responsible for polar transport of indole-3-628

acetic acid in sections of maize coleoptiles Planta 155 (1) 68ndash75 629

Guiscafrearrillaga J (1949) Formation of galls in stems and leaves of sugar cane in 630

response to injections of growth-regulating substances Phytopathology 39 (6) 489ndash493 631

Halitschke R Gase K Hui D Schmidt DD and Baldwin IT (2003) Molecular 632

interactions between the specialist herbivore Manduca sexta (Lepidoptera Sphingidae) 633

and its natural host Nicotiana attenuata VI Microarray analysis reveals that most 634

herbivore-specific transcriptional changes are mediated by fatty acid-amino acid 635

conjugates Plant Physiology 131 (4) 1894ndash1902 636

Hall TA (1999) BioEdit a user-friendly biological sequence alignment editor and analysis 637

program for Windows 9598NT Nucleic acids symposium series (41) 95-98 638

Hamner KC and Kraus EJ (1937) Histological reactions of bean plants to growth 639

promoting substances Botanical Gazette 735ndash807 640

He W Brumos J Li H Ji Y Ke M Gong X Zeng Q Li W Zhang X and An 641

F (2011) A small-molecule screen identifies L-kynurenine as a competitive inhibitor of 642

TAA1TAR activity in ethylene-directed auxin biosynthesis and root growth in 643

Arabidopsis The Plant Cell 23 (11) 3944ndash3960 644

Heiling S Schuman MC Schoettner M Mukerjee P Berger B Schneider B 645

Jassbi AR and Baldwin IT (2010) Jasmonate and ppHsystemin regulate key 646

malonylation steps in the biosynthesis of 17-hydroxygeranyllinalool diterpene glycosides 647


25

an abundant and effective direct defense against herbivores in Nicotiana attenuata The 648

Plant Cell 22 (1) 273ndash292 649

Hertel R Lomax TL and Briggs WR (1983) Auxin transport in membrane vesicles 650

from Cucurbita pepo L Planta 157 (3) 193ndash201 651

Hou X Lee LYC Xia K Yan Y and Yu H (2010) DELLAs modulate jasmonate 652

signaling via competitive binding to JAZs Developmental cell 19 (6) 884ndash894 653

Howe GA and Jander G (2008) Plant immunity to insect herbivores Annu Rev Plant 654

Biol 59 41ndash66 655

Jimenez-Aleman GH Machado RAR Goumlrls H Baldwin IT and Boland W 656

(2015) Synthesis structural characterization and biological activity of two diastereomeric 657

JA-Ile macrolactones Organic amp biomolecular chemistry 13 (21) 5885ndash5893 658

Jin X Zimmermann J Polle A and Fischer U (2015) Auxin is a long-range signal 659

that acts independently of ethylene signaling on leaf abscission in Populus Frontiers in 660

plant science 6 634 661

Kallenbach M Bonaventure G Gilardoni PA Wissgott A and Baldwin IT 662

(2012) Empoasca leafhoppers attack wild tobacco plants in a jasmonate-dependent 663

manner and identify jasmonate mutants in natural populations Proceedings of the 664

National Academy of Sciences 109 (24) E1548-E1557 665

Kang J-H Wang L Giri A and Baldwin IT (2006) Silencing threonine deaminase 666

and JAR4 in Nicotiana attenuata impairs jasmonic acidndashisoleucinendashmediated defenses 667

against Manduca sexta The Plant Cell 18 (11) 3303ndash3320 668

Kernan A and Thornburg RW (1989) Auxin levels regulate the expression of a 669

wound-inducible proteinase inhibitor II-chloramphenicol acetyl transferase gene fusion in 670

vitro and in vivo Plant Physiology 91 (1) 73ndash78 671

Koo AJK Gao X Daniel Jones A and Howe GA (2009) A rapid wound signal 672

activates the systemic synthesis of bioactive jasmonates in Arabidopsis The Plant Journal 673

59 (6) 974ndash986 674

Kramer EM Rutschow HL and Mabie SS (2011) AuxV a database of auxin 675

transport velocities Trends in plant science 16 (9) 461ndash463 676

Kroes A van Loon JJA and Dicke M (2014) Density-dependent interference of 677

aphids with caterpillar-induced defenses in Arabidopsis Involvement of phytohormones 678

and transcription factors Plant and cell physiology pcu150 679


26

Kruumlgel T Lim M Gase K Halitschke R and Baldwin IT (2002) Agrobacterium-680

mediated transformation of Nicotiana attenuata a model ecological expression system 681

Chemoecology 12 (4) 177ndash183 682

Kurepin LV Park JM Lazarovits G and Bernards MA (2015) Burkholderia 683

phytofirmans-induced shoot and root growth promotion is associated with endogenous 684

changes in plant growth hormone levels Plant Growth Regulation 75 (1) 199ndash207 685

Lambrecht M Okon Y Broek AV and Vanderleyden J (2000) Indole-3-acetic 686

acid a reciprocal signalling molecule in bacteriandashplant interactions evolution 54 59 687

Lee G Joo Y Diezel C Lee EJ Baldwin IT and Kim S (2016) Trichobaris 688

weevils distinguish amongst toxic host plants by sensing volatiles that do not affect larval 689

performance Molecular ecology 690

Lewis DR Ramirez MV Miller ND Vallabhaneni P Ray WK Helm RF 691

Winkel BSJ and Muday GK (2011) Auxin and ethylene induce flavonol 692

accumulation through distinct transcriptional networks Plant Physiology 156 (1) 144ndash693

164 694

Li R Zhang J Li J Zhou G Wang Q Bian W Erb M and Lou Y (2015) 695

Prioritizing plant defence over growth through WRKY regulation facilitates infestation by 696

non-target herbivores Elife 4 e04805 697

Ling Z Zhou W Baldwin IT and Xu S (2015) Insect herbivory elicits genome‐wide 698

alternative splicing responses in Nicotiana attenuata The Plant Journal 84 (1) 228ndash243 699

Liu Y Ahn J-E Datta S Salzman RA Moon J Huyghues-Despointes B 700

Pittendrigh B Murdock LL Koiwa H and Zhu-Salzman K (2005) Arabidopsis 701

vegetative storage protein is an anti-insect acid phosphatase Plant Physiology 139 (3) 702

1545ndash1556 703

Lulu T Park S-Y Ibrahim R and Paek K-Y (2015) Production of biomass and 704

bioactive compounds from adventitious roots by optimization of culturing conditions of 705

Eurycoma longifolia in balloon-type bubble bioreactor system Journal of bioscience and 706

bioengineering 119 (6) 712ndash717 707

Machado RAR Arce C Ferrieri AP Baldwin IT and Erb M (2015) Jasmonate‐708

dependent depletion of soluble sugars compromises plant resistance to Manduca sexta 709

New Phytologist 207 (1) 91ndash105 710


27

Machado RAR Ferrieri AP Am Robert C Glauser G Kallenbach M Baldwin 711

IT and Erb M (2013) Leaf‐herbivore attack reduces carbon reserves and regrowth 712

from the roots via jasmonate and auxin signaling New Phytologist 200 (4) 1234ndash1246 713

Machado RAR McClure M Herveacute M Baldwin IT and Erb M (2016) Benefits 714

of jasmonate-dependent defenses against vertebrate herbivores in nature Elife 5 e13720 715

Mahdieh M Noori M and Hoseinkhani S (2015) Studies of in vitro Adventitious Root 716

Induction and Flavonoid Profiles in Rumex crispus Advances in Life Sciences 5 (3) 53ndash717

57 718

Mano Y and Nemoto K (2012) The pathway of auxin biosynthesis in plants Journal of 719

Experimental Botany ers091 720

Mapes CC and Davies PJ (2001a) Cytokinins in the ball gall of Solidago altissima and 721

in the gall forming larvae of Eurosta solidaginis New Phytologist 151 (1) 203ndash212 722

Mapes CC and Davies PJ (2001b) Indole‐3‐acetic acid and ball gall development on 723

Solidago altissima New Phytologist 151 (1) 195ndash202 724

Mashiguchi K Tanaka K Sakai T Sugawara S Kawaide H Natsume M 725

Hanada A Yaeno T Shirasu K and Yao H (2011) The main auxin biosynthesis 726

pathway in Arabidopsis Proceedings of the National Academy of Sciences 108 (45) 727

18512ndash18517 728

Meldau S Wu J and Baldwin IT (2009) Silencing two herbivory‐activated MAP 729

kinases SIPK and WIPK does not increase Nicotiana attenuatas susceptibility to 730

herbivores in the glasshouse and in nature New Phytologist 181 (1) 161ndash173 731

Morris DA Kadir GO and Barry AJ (1973) Auxin transport in intact pea seedlings 732

(Pisum sativum L) the inhibition of transport by 2 3 5-triiodobenzoic acid Planta 110 733

(2) 173ndash182 734

Mousavi SAR Chauvin A Pascaud F Kellenberger S and Farmer EE (2013) 735

GLUTAMATE RECEPTOR-LIKE genes mediate leaf-to-leaf wound signalling Nature 736

500 (7463) 422ndash426 737

Onkokesung N Gaacutelis I Dahl CC von Matsuoka K Saluz H-P and Baldwin IT 738

(2010) Jasmonic acid and ethylene modulate local responses to wounding and simulated 739

herbivory in Nicotiana attenuata leaves Plant Physiology 153 (2) 785ndash798 740

Onkokesung N Gaquerel E Kotkar H Kaur H Baldwin IT and Galis I (2012) 741

MYB8 controls inducible phenolamide levels by activating three novel 742


28

hydroxycinnamoyl-coenzyme A polyamine transferases in Nicotiana attenuata Plant 743


Paschold A Halitschke R and Baldwin IT (2007) Co (i)‐ordinating defenses 745

NaCOI1 mediates herbivore‐induced resistance in Nicotiana attenuata and reveals the 746

role of herbivore movement in avoiding defenses The Plant Journal 51 (1) 79ndash91 747

Pasqua G Monacelli B Mulinacci N Rinaldi S Giaccherini C Innocenti M and 748

Vinceri FF (2005) The effect of growth regulators and sucrose on anthocyanin 749

production in Camptotheca acuminata cell cultures Plant Physiology and Biochemistry 750

43 (3) 293ndash298 751

Qi T Song S Ren Q Wu D Huang H Chen Y Fan M Peng W Ren C and 752

Xie D (2011) The Jasmonate-ZIM-domain proteins interact with the WD-753

RepeatbHLHMYB complexes to regulate Jasmonate-mediated anthocyanin 754

accumulation and trichome initiation in Arabidopsis thaliana The Plant Cell 23 (5) 755

1795ndash1814 756

Reed RC Brady SR and Muday GK (1998) Inhibition of auxin movement from the 757

shoot into the root inhibits lateral root development in Arabidopsis Plant Physiology 118 758

(4) 1369ndash1378 759

Rubery PH (1979) The effects of 2 4-dinitrophenol and chemical modifying reagents on 760

auxin transport by suspension-cultured crown gall cells Planta 144 (2) 173ndash178 761

Sassi M Lu Y Zhang Y Wang J Dhonukshe P Blilou I Dai M Li J Gong 762

X and Jaillais Y (2012) COP1 mediates the coordination of root and shoot growth by 763

light through modulation of PIN1-and PIN2-dependent auxin transport in Arabidopsis 764

Development 139 (18) 3402ndash3412 765

Schaumlfer M Bruumltting C Gase K Reichelt M Baldwin I and Meldau S (2013) 766

lsquoReal timersquogenetic manipulation a new tool for ecological field studies The Plant Journal 767

76 (3) 506ndash518 768

Schaumlfer M Meza‐Canales ID Bruumltting C Baldwin IT and Meldau S (2015) 769

Cytokinin concentrations and CHASE‐DOMAIN CONTAINING HIS KINASE 2 770

(NaCHK2)‐and NaCHK3‐mediated perception modulate herbivory‐induced defense 771

signaling and defenses in Nicotiana attenuata New Phytologist 772

Schaumlller G (1968) Biochemische Analyse des Aphidenspeichels und seine Bedeutung fuumlr 773

die Gallenbildung Zool Jb Physiol 74 54ndash87 774


29

Schmelz EA Engelberth J Alborn HT ODonnell P Sammons M Toshima H 775

and Tumlinson JH (2003) Simultaneous analysis of phytohormones phytotoxins and 776

volatile organic compounds in plants Proceedings of the National Academy of Sciences 777

100 (18) 10552ndash10557 778

Shi Q Li C and Zhang F (2006) Nicotine synthesis in Nicotiana tabacum L induced 779

by mechanical wounding is regulated by auxin Journal of Experimental Botany 57 (11) 780

2899ndash2907 781

Shin DH Cho M Choi MG Das PK Lee S-K Choi S-B and Park Y-I 782

(2015) Identification of genes that may regulate the expression of the transcription factor 783

production of anthocyanin pigment 1 (PAP1)MYB75 involved in Arabidopsis 784

anthocyanin biosynthesis Plant cell reports 34 (5) 805ndash815 785

Song Y (2014) Insight into the mode of action of 2 4‐dichlorophenoxyacetic acid (2 4‐D) 786

as an herbicide Journal of integrative plant biology 56 (2) 106ndash113 787

Steppuhn A Gaquerel E and Baldwin IT (2010) The two α-dox genes of Nicotiana 788

attenuata overlapping but distinct functions in development and stress responses BMC 789

plant biology 10 (1) 171 790

Stitz M Gase K Baldwin IT and Gaquerel E (2011) Ectopic expression of AtJMT 791

in Nicotiana attenuata creating a metabolic sink has tissue-specific consequences for the 792

jasmonate metabolic network and silences downstream gene expression Plant Physiology 793

157 (1) 341ndash354 794

Straka JR Hayward AR and Emery RN (2010) Gall-inducing Pachypsylla celtidis 795

(Psyllidae) infiltrate hackberry trees with high concentrations of phytohormones Journal 796

of Plant Interactions 5 (3) 197ndash203 797

Tanaka Y Okada K Asami T and Suzuki Y (2013) Phytohormones and willow gall 798

induction by a gall-inducing sawfly Biosci Biotechnol Biochem 77 1942ndash1948 799

Tanaka Y and Uritani I (1979) Polar transport and content of indole-3-acetic acid in 800

wounded sweet potato root tissues Plant and cell physiology 20 (6) 1087ndash1095 801

Thaler JS and Bostock RM (2004) Interactions between abscisic-acid-mediated 802

responses and plant resistance to pathogens and insects Ecology 85 (1) 48ndash58 803

Thompson JD Higgins DG and Gibson TJ (1994) CLUSTAL W improving the 804

sensitivity of progressive multiple sequence alignment through sequence weighting 805

position-specific gap penalties and weight matrix choice Nucleic acids research 22 (22) 806

4673ndash4680 807


30

Thornburg RW and Li X (1991) Wounding Nicotiana tabacum leaves causes a decline 808

in endogenous indole-3-acetic acid Plant Physiology 96 (3) 802ndash805 809

Tian Q Chen F Liu J Zhang F and Mi G (2008) Inhibition of maize root growth 810

by high nitrate supply is correlated with reduced IAA levels in roots Journal of plant 811

physiology 165 (9) 942ndash951 812

Tooker JF and Moraes CM de (2011a) Feeding by a gall-inducing caterpillar species 813

alters levels of indole-3-acetic and abscisic acid in Solidago altissima (Asteraceae) stems 814

Arthropod-Plant Interactions 5 (2) 115ndash124 815

Tooker JF and Moraes CM de (2011b) Feeding by Hessian fly (Mayetiola destructor 816

[Say]) larvae on wheat increases levels of fatty acids and indole-3-acetic acid but not 817

hormones involved in plant-defense signaling Journal of plant growth regulation 30 (2) 818

158ndash165 819

van Noorden GE Ross JJ Reid JB Rolfe BG and Mathesius U (2006) 820

Defective long-distance auxin transport regulation in the Medicago truncatula super 821

numeric nodules mutant Plant Physiology 140 (4) 1494ndash1506 822

VanDoorn A Bonaventure G Schmidt DD and Baldwin IT (2011) Regulation of 823

jasmonate metabolism and activation of systemic signaling in Solanum nigrum COI1 and 824

JAR4 play overlapping yet distinct roles New Phytologist 190 (3) 640ndash652 825

Wang L Allmann S Wu J and Baldwin IT (2008) Comparisons of 826

LIPOXYGENASE3-and JASMONATE-RESISTANT46-silenced plants reveal that 827

jasmonic acid and jasmonic acid-amino acid conjugates play different roles in herbivore 828

resistance of Nicotiana attenuata Plant Physiology 146 (3) 904ndash915 829

Wasternack C and Hause B (2013) Jasmonates biosynthesis perception signal 830

transduction and action in plant stress response growth and development An update to 831

the 2007 review in Annals of Botany Annals of Botany 111 (6) 1021ndash1058 832

Weinhold A Kallenbach M and Baldwin IT (2013) Progressive 35S promoter 833

methylation increases rapidly during vegetative development in transgenic Nicotiana 834

attenuata plants BMC plant biology 13 (1) 99 835

Winz RA and Baldwin IT (2001) Molecular interactions between the specialist 836

herbivore Manduca sexta (Lepidoptera Sphingidae) and its natural host Nicotiana 837

attenuata IV Insect-induced ethylene reduces jasmonate-induced nicotine accumulation 838

by regulating putrescine N-methyltransferase transcripts Plant Physiology 125 (4) 2189ndash839

2202 840


31

Woldemariam MG Onkokesung N Baldwin IT and Galis I (2012) Jasmonoyl‐l‐841

isoleucine hydrolase 1 (JIH1) regulates jasmonoyl‐l‐isoleucine levels and attenuates plant 842

defenses against herbivores The Plant Journal 72 (5) 758ndash767 843

Wu J and Baldwin IT (2009) Herbivory‐induced signalling in plants perception and 844

action Plant cell amp environment 32 (9) 1161ndash1174 845

Xin Z Yu Z Erb M Turlings TCJ Wang B Qi J Liu S and Lou Y (2012) 846

The broad‐leaf herbicide 2 4‐dichlorophenoxyacetic acid turns rice into a living trap for a 847

major insect pest and a parasitic wasp New Phytologist 194 (2) 498ndash510 848

Xu S Zhou W Pottinger S and Baldwin IT (2015) Herbivore associated elicitor-849

induced defences are highly specific among closely related Nicotiana species BMC plant 850

biology (1) 2 851

Yamaguchi H Tanaka H Hasegawa M Tokuda M Asami T and Suzuki Y 852

(2012) Phytohormones and willow gall induction by a gall‐inducing sawfly New 853

Phytologist 196 (2) 586ndash595 854

Yang D-L Yao J Mei C-S Tong X-H Zeng L-J Li Q Xiao L-T Sun T-p 855

Li J and Deng X-W (2012) Plant hormone jasmonate prioritizes defense over growth 856

by interfering with gibberellin signaling cascade Proceedings of the National Academy of 857

Sciences 109 (19) E1192-E1200 858

Yang F Song Y Yang H Liu Z Zhu G and Yang Y (2014) An auxin‐responsive 859

endogenous peptide regulates root development in Arabidopsis Journal of integrative 860

plant biology 56 (7) 635ndash647 861

Zhang P-J Li W-D Huang F Zhang J-M Xu F-C and Lu Y-B (2013) 862

Feeding by whiteflies suppresses downstream jasmonic acid signaling by eliciting 863

salicylic acid signaling Journal of Chemical Ecology 39 (5) 612ndash619 864


0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min

Treatment P lt 0001Time P = 0570TT P = 0782

IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001

Time after M sextafeeding start (h)

a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC

Figure 1 Indole-3-acetic acid (IAA) is induced specifically and rapidly by real and simulated M sextaattack Average (plusmnSE) IAA levels in leaves that are attacked by M sexta caterpillars (A) treated with Msexta oral secretions (B C) or treated with an herbivore elicitor (D) (n=5) Different letters indicatesignificant differences between treatments (P lt 005) Control intact plants W+W wounded and water-treated plants W+OS wounded and M sexta oral secretion-treated plants W+FACs wounded and fattyacid-amino acid conjugate-treated plants

0

1

2

3

4

Control 15 30 60 180

aa a

bb

Time after W+OS-induction (s)IA

A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D

Time after treatment Time after treatment


020406080

100

0 30 60 90 120

Stem

Time P = 0764Treatment P = 0558TT P = 0093

IAA

(ng

gFW

)

Time after treatment (min)

0

35

70

0 30 60 90 120

Main root

Time P = 0232Treatment P = 0486TT P = 0146 0

10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves

Time P = 0131Treatment P lt 0001TT P = 0085

ControlW+OS

A B

C D

E F

Figure 2 Herbivory induces IAA both locally and systemically Average (plusmnSE) IAA levels followingsimulated M sexta attack in local treated leaves (A) and in untreated petioles (B) stem (C) systemicleaves (D) main root (E) and lateral roots (F) (n=5) Asterisks indicate significant differences betweentreatments within plant tissues and time points ( P lt 005 P lt 0001) Control intact plantsW+OS wounded and M sexta oral secretion-treated plants

10 10

10

1010

10


Time after treatment (h)

01020304050

05 1 3

W+W

Time P lt 0001Treatment P lt 0001TT P = 0036

a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3

Time P = 0002Treatment P lt 0001TT P lt 0001

b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C

Figure 3 IAA induction in leaves occurs across different developmental stages Average (plusmnSE) IAAlevels in local treated leaves following simulated M sexta attack at the early rosette (A) elongated (B)and flowering stage (C) (n=5) Different letters indicate significant differences between treatments withindevelopmental stages and time points (P lt 005) Control intact plants W+W wounded and water-treated plants W+OS wounded and M sexta oral secretion-treated plants


0

5

10

0123

0

2

4

Control 5 60Time after W+OS treatment (min)

YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge

YUCCA-mediated oxidative decarboxylation

Indole-3-pyruvic acid Indole-3-acetic acid

A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2

YUCCA-like 6P = 0001 b

a

b

a

D

P lt 0001

Figure 4 YUCCA-like genes are upregulated in response to simulated M sexta herbivory (A)Schematic representation of YUCCA-mediated conversion of indole-3-pyruvic acid into IAA Average(plusmnSE) transcript abundance relative to control of YUCCA-like 3 (B) YUCCA-like 5 (C) YUCCA-like 6(D) and YUCCA-like 9 (E) in treated leaves three minutes after elicitation and YUCCA-like 1 (F) andYUCCA-like 3 (G) 5 and 60 min following simulated M sexta attack (n=3) Different letters indicatesignificant differences between treatments (P lt 005) Control intact plants W+W wounded and water-treated plants W+OS wounded and M sexta oral secretion-treated plants W+FACs wounded and fattyacid-amino acid conjugate-treated plants

012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001

Time after W+OS treatment (min)


IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control


Figure 5 Manduca sexta-induced IAA peaks earlier than jasmonic acid (JA) Left Y-axis average (plusmnSE)leaf IAA levels in response to M sexta attack Right Y-axis average (plusmnSE) leaf JA levels in response toM sexta attack Closed squares IAA levels upon W+OS treatments closed triangles IAA levels incontrol untreated plants Grey squares JA levels upon W+OS treatments grey triangles jasmonic acid(JA) levels in control untreated plants (n=5) Different letters indicate significant differences betweentreatments for individual metabolites (P lt 005) IAA Time P = 0015 treatment P lt 0001 Timetreatment P = 0638 JA Time P lt 0001 treatment P lt 0001 Time treatment P lt 0001) Controlintact plants W+OS wounded and M sexta oral secretion-treated plants

5


0

1

2

3

4

C W OS C W OS C W OS C W OS C W OS C W OS C W OS C W OS C W OS

EV irSIPK irWIPK irGLA irAOS irAOC irOPR3 irJAR46 irCOI1

a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)

Genotype P lt 0001Treatment P lt 0001GT P = 0113

Early JA-signaling JA-biosynthesis JA-Ile-perception

Wild type

Impaired in

JA-Ile-biosynthesis

Figure 6 Jasmonate signaling is not required for the M sexta-induced accumulation of IAA (A)Average (plusmnSE) IAA levels in local treated leaves of wild type plants (empty vector EV) and plantgenotypes impaired in early JA signaling jasmonate biosynthesis andor JA-Ile perception 45 minutesafter elicitation (n=5) Different letters indicate significant differences between treatments within eachgenotype (P lt 005) C control intact plants W wounded and water-treated plants OS wounded andM sexta oral secretions-treated plants


0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS

Lanolin L-Kyn TIBA IAA Petiole pretreatment

Leaf treatment P lt 0001Petiole pretreatment P lt 0001LTPPT P lt 0001

Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC

Genotype P lt 0001Treatment P lt 0001GT P lt 0001

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta

Figure 7 Manduca sexta-induced IAA and JA act synergistically to trigger anthocyanin accumulation in thestems (A) Average (plusmnSE) stem anthocyanin content 5 days following either simulated or continuous M sextaattack exogenous application of methyl jasmonate (MeJA) andor IAA (n=5) (B) Average (plusmnSE) stemanthocyanin content 5 days following simulated M sexta attack and petiole-pretreatments with either IAA theIAA biosynthesis inhibitor L-kynurenine (L-Kyn) or the IAA transport inhibitor TIBA (235-triiodobenzoic acid)(n=12) (C) Average (plusmnSE) stem anthocyanin contents following simulated M sexta attack of wild type and JA-impaired irAOC plants (n=10) (D) Correlation between stem anthocyanin content and stem coloration InsetPhotograph of the red stem phenotype Asterisks indicate significant differences between treatments and control(A) between simulated herbivory treatments within petiole pretreatments (B) and between treatments withingenotypes (C) ( P lt 005 P lt 001 P lt 0001) The correlation between stem coloration index andstem anthocyanin content was evaluated by a Pearson product moment test Leaf treatments Control intactplants W+W wounded and water-treated plants W+OS wounded and M sexta oral secretion-treated plants Msexta plants subjected to actual M sexta attack IAA rosette leaves treated with indole-3-acetic acid MeJArosette leaves treated with methyl jasmonic acid IAA+MeJA rosette leaves treated with IAA and MeJA Petiolepretreatments Petioles treated with either pure lanoline paste (Lanoline) L-kynurenine (L-Kyn) 235-triiodobenzoic acid (TIBA) or indole-3-acetic acid (IAA) dissolved in lanoline 1h prior to leaf treatments


0

15

30

45

60

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90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Control W+W W+OS M sexta MeJA

0

100

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Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine

Dicaffeoylspermidine

Ste

m c

onte

nt

Leaf treatment P lt 0001Petiole pretreatment P lt 0001LTPPT P = 0004

Nicotine

DTGsP

eak

area

103

gFW

Leaf treatment P lt 0001Petiole pretreatment P = 0800LTPPT P = 0968

0

03

06

09

12

15

18

mg

gFW

Leaf treatment P lt 0001Petiole pretreatment P = 0431 LTPPT P = 0888

ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300

360 Leaf treatment P lt 0001Petiole pretreatment P lt 0001LTPPT P lt 0001

microgg

FWmicrog

gFW

Petiole pretreatment

Leaf treatment

A C

B D

Figure 8 IAA specifically potentiates the herbivore-induced systemic production of phenolamides Average (plusmnSE) caffeoylputrescine (A) dicaffeoylspermidine (B) nicotine (C) and diterpene glycoside (D) levels in the stems 5 days following simulated or real M sexta attack and petiole pretreatments with IAA (n=5) Asterisks indicate significant differences between petiole pretreatments within simulated M sexta attack treatments ( P lt 005 P lt 001 P lt 0001) Petiole pretreatments Control petioles treated with pure lanoline paste 1h prior to leaf treatments IAA petioles treated with IAA dissolved in lanoline 1h prior to leaf treatments Leaf treatments Control intact plants W+W wounded and water-treated plants W+OS wounded and M sexta oral secretion-treated plants M sexta plants subjected to actual M sexta attack MeJA rosette leaves treated with methyl jasmonic acid dissolved in lanoline paste


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Bartlett L and Connor EF (2014) Exogenous phytohormones and the induction of plant galls by insects Arthropod-PlantInteractions 8 (4) 339-348


Bhalerao RP Ekloumlf J Ljung K Marchant A Bennett M and Sandberg G (2002) Shoot-derived auxin is essential for earlylateral root emergence in Arabidopsis seedlings The Plant Journal 29 (3) 325-332


Bonaventure G Schuck S and Baldwin IT (2011) Revealing complexity and specificity in the activation of lipase-mediatedoxylipin biosynthesis a specific role of the Nicotiana attenuata GLA1 lipase in the activation of jasmonic acid biosynthesis inleaves and roots Plant cell amp environment 34 (9) 1507-1520


Borevitz JO Xia Y Blount J Dixon RA and Lamb C (2000) Activation tagging identifies a conserved MYB regulator ofphenylpropanoid biosynthesis The Plant Cell 12 (12) 2383-2393


Chen Q Sun J Zhai Q Zhou W Qi L Xu L Wang B Chen R Jiang H and Qi J (2011) The basic helix-loop-helixtranscription factor MYC2 directly represses PLETHORA expression during jasmonate-mediated modulation of the root stem cellniche in Arabidopsis The Plant Cell 23 (9) 3335-3352


Connor EF Bartlett L OToole S Byrd S Biskar K and Orozco J (2012) The mechanism of gall induction makes galls redArthropod-Plant Interactions 6 (4) 489-495


Dafoe NJ Thomas JD Shirk PD Legaspi ME Vaughan MM Huffaker A Teal PE and Schmelz EA (2013) Europeancorn borer (Ostrinia nubilalis) induced responses enhance susceptibility in maize PloS one 8 (9)


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Dahl CC von Winz RA Halitschke R Kuumlhnemann F Gase K and Baldwin IT (2007) Tuning the herbivore-inducedethylene burst the role of transcript accumulation and ethylene perception in Nicotiana attenuata The Plant Journal 51 (2) 293-307



DeWald DB Sadka A and Mullet JE (1994) Sucrose modulation of soybean Vsp gene expression is inhibited by auxin PlantPhysiology 104 (2) 439-444


Diezel C Allmann S and Baldwin IT (2011a) Mechanisms of optimal defense patterns in Nicotiana attenuata Floweringattenuates herbivory-elicited ethylene and jasmonate Signaling Journal of integrative plant biology 53 (12) 971-983


Diezel C Kessler D and Baldwin IT (2011b) Pithy protection Nicotiana attenuatas jasmonic acid-mediated defenses arerequired to resist stem-boring weevil larvae Plant Physiology 155 (4) 1936-1946


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Erb M Meldau S and Howe GA (2012) Role of phytohormones in insect-specific plant reactions Trends in plant science 17(5) 250-259


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Ferrieri AP Arce C Machado RAR Meza-Canales ID Lima E Baldwin IT and Erb M (2015) A Nicotiana attenuata cellwall invertase inhibitor (NaCWII) reduces growth and increases secondary metabolite biosynthesis in herbivore-attacked plantsNew Phytologist


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Halitschke R Gase K Hui D Schmidt DD and Baldwin IT (2003) Molecular interactions between the specialist herbivoreManduca sexta (Lepidoptera Sphingidae) and its natural host Nicotiana attenuata VI Microarray analysis reveals that mostherbivore-specific transcriptional changes are mediated by fatty acid-amino acid conjugates Plant Physiology 131 (4) 1894-1902


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He W Brumos J Li H Ji Y Ke M Gong X Zeng Q Li W Zhang X and An F (2011) A small-molecule screen identifies L-kynurenine as a competitive inhibitor of TAA1TAR activity in ethylene-directed auxin biosynthesis and root growth in ArabidopsisThe Plant Cell 23 (11) 3944-3960


Heiling S Schuman MC Schoettner M Mukerjee P Berger B Schneider B Jassbi AR and Baldwin IT (2010)Jasmonate and ppHsystemin regulate key malonylation steps in the biosynthesis of 17-hydroxygeranyllinalool diterpeneglycosides an abundant and effective direct defense against herbivores in Nicotiana attenuata The Plant Cell 22 (1) 273-292


Hertel R Lomax TL and Briggs WR (1983) Auxin transport in membrane vesicles from Cucurbita pepo L Planta 157 (3) 193-201


Hou X Lee LYC Xia K Yan Y and Yu H (2010) DELLAs modulate jasmonate signaling via competitive binding to JAZsDevelopmental cell 19 (6) 884-894


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Jin X Zimmermann J Polle A and Fischer U (2015) Auxin is a long-range signal that acts independently of ethylene signalingon leaf abscission in Populus Frontiers in plant science 6 634


Kallenbach M Bonaventure G Gilardoni PA Wissgott A and Baldwin IT (2012) Empoasca leafhoppers attack wild tobaccoplants in a jasmonate-dependent manner and identify jasmonate mutants in natural populations Proceedings of the NationalAcademy of Sciences 109 (24) E1548-E1557

Pubmed Author and Title wwwplantphysiolorgon June 1 2020 - Published by Downloaded from Copyright copy 2016 American Society of Plant Biologists All rights reserved

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

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Kernan A and Thornburg RW (1989) Auxin levels regulate the expression of a wound-inducible proteinase inhibitor II-chloramphenicol acetyl transferase gene fusion in vitro and in vivo Plant Physiology 91 (1) 73-78


Koo AJK Gao X Daniel Jones A and Howe GA (2009) A rapid wound signal activates the systemic synthesis of bioactivejasmonates in Arabidopsis The Plant Journal 59 (6) 974-986


Kramer EM Rutschow HL and Mabie SS (2011) AuxV a database of auxin transport velocities Trends in plant science 16 (9)461-463


Kroes A van Loon JJA and Dicke M (2014) Density-dependent interference of aphids with caterpillar-induced defenses inArabidopsis Involvement of phytohormones and transcription factors Plant and cell physiology pcu150


Kruumlgel T Lim M Gase K Halitschke R and Baldwin IT (2002) Agrobacterium-mediated transformation of Nicotianaattenuata a model ecological expression system Chemoecology 12 (4) 177-183


Kurepin LV Park JM Lazarovits G and Bernards MA (2015) Burkholderia phytofirmans-induced shoot and root growthpromotion is associated with endogenous changes in plant growth hormone levels Plant Growth Regulation 75 (1) 199-207


Lambrecht M Okon Y Broek AV and Vanderleyden J (2000) Indole-3-acetic acid a reciprocal signalling molecule in bacteria-plant interactions evolution 54 59


Lee G Joo Y Diezel C Lee EJ Baldwin IT and Kim S (2016) Trichobaris weevils distinguish amongst toxic host plants bysensing volatiles that do not affect larval performance Molecular ecology


Lewis DR Ramirez MV Miller ND Vallabhaneni P Ray WK Helm RF Winkel BSJ and Muday GK (2011) Auxin andethylene induce flavonol accumulation through distinct transcriptional networks Plant Physiology 156 (1) 144-164


Li R Zhang J Li J Zhou G Wang Q Bian W Erb M and Lou Y (2015) Prioritizing plant defence over growth throughWRKY regulation facilitates infestation by non-target herbivores Elife 4 e04805


Ling Z Zhou W Baldwin IT and Xu S (2015) Insect herbivory elicits genome-wide alternative splicing responses in Nicotianaattenuata The Plant Journal 84 (1) 228-243


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Machado RAR Arce C Ferrieri AP Baldwin IT and Erb M (2015) Jasmonate-dependent depletion of soluble sugarscompromises plant resistance to Manduca sexta New Phytologist 207 (1) 91-105


Machado RAR Ferrieri AP Am Robert C Glauser G Kallenbach M Baldwin IT and Erb M (2013) Leaf-herbivore attackreduces carbon reserves and regrowth from the roots via jasmonate and auxin signaling New Phytologist 200 (4) 1234-1246


Machado RAR McClure M Herveacute M Baldwin IT and Erb M (2016) Benefits of jasmonate-dependent defenses againstvertebrate herbivores in nature Elife 5 e13720


Mahdieh M Noori M and Hoseinkhani S (2015) Studies of in vitro Adventitious Root Induction and Flavonoid Profiles inRumex crispus Advances in Life Sciences 5 (3) 53-57


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Mousavi SAR Chauvin A Pascaud F Kellenberger S and Farmer EE (2013) GLUTAMATE RECEPTOR-LIKE genes mediateleaf-to-leaf wound signalling Nature 500 (7463) 422-426


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Onkokesung N Gaquerel E Kotkar H Kaur H Baldwin IT and Galis I (2012) MYB8 controls inducible phenolamide levels wwwplantphysiolorgon June 1 2020 - Published by Downloaded from

Copyright copy 2016 American Society of Plant Biologists All rights reserved

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Paschold A Halitschke R and Baldwin IT (2007) Co (i)-ordinating defenses NaCOI1 mediates herbivore-induced resistance inNicotiana attenuata and reveals the role of herbivore movement in avoiding defenses The Plant Journal 51 (1) 79-91


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Qi T Song S Ren Q Wu D Huang H Chen Y Fan M Peng W Ren C and Xie D (2011) The Jasmonate-ZIM-domainproteins interact with the WD-RepeatbHLHMYB complexes to regulate Jasmonate-mediated anthocyanin accumulation andtrichome initiation in Arabidopsis thaliana The Plant Cell 23 (5) 1795-1814


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Rubery PH (1979) The effects of 2 4-dinitrophenol and chemical modifying reagents on auxin transport by suspension-culturedcrown gall cells Planta 144 (2) 173-178


Sassi M Lu Y Zhang Y Wang J Dhonukshe P Blilou I Dai M Li J Gong X and Jaillais Y (2012) COP1 mediates thecoordination of root and shoot growth by light through modulation of PIN1-and PIN2-dependent auxin transport in ArabidopsisDevelopment 139 (18) 3402-3412


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Schaumlfer M Meza-Canales ID Bruumltting C Baldwin IT and Meldau S (2015) Cytokinin concentrations and CHASE-DOMAINCONTAINING HIS KINASE 2 (NaCHK2)-and NaCHK3-mediated perception modulate herbivory-induced defense signaling anddefenses in Nicotiana attenuata New Phytologist


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Shi Q Li C and Zhang F (2006) Nicotine synthesis in Nicotiana tabacum L induced by mechanical wounding is regulated byauxin Journal of Experimental Botany 57 (11) 2899-2907


Shin DH Cho M Choi MG Das PK Lee S-K Choi S-B and Park Y-I (2015) Identification of genes that may regulate theexpression of the transcription factor production of anthocyanin pigment 1 (PAP1)MYB75 involved in Arabidopsis anthocyaninbiosynthesis Plant cell reports 34 (5) 805-815 wwwplantphysiolorgon June 1 2020 - Published by Downloaded from



Song Y (2014) Insight into the mode of action of 2 4-dichlorophenoxyacetic acid (2 4-D) as an herbicide Journal of integrativeplant biology 56 (2) 106-113


Steppuhn A Gaquerel E and Baldwin IT (2010) The two a-dox genes of Nicotiana attenuata overlapping but distinct functionsin development and stress responses BMC plant biology 10 (1) 171


Stitz M Gase K Baldwin IT and Gaquerel E (2011) Ectopic expression of AtJMT in Nicotiana attenuata creating a metabolicsink has tissue-specific consequences for the jasmonate metabolic network and silences downstream gene expression PlantPhysiology 157 (1) 341-354


Straka JR Hayward AR and Emery RN (2010) Gall-inducing Pachypsylla celtidis (Psyllidae) infiltrate hackberry trees with highconcentrations of phytohormones Journal of Plant Interactions 5 (3) 197-203


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Thaler JS and Bostock RM (2004) Interactions between abscisic-acid-mediated responses and plant resistance to pathogensand insects Ecology 85 (1) 48-58


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Thornburg RW and Li X (1991) Wounding Nicotiana tabacum leaves causes a decline in endogenous indole-3-acetic acid PlantPhysiology 96 (3) 802-805


Tian Q Chen F Liu J Zhang F and Mi G (2008) Inhibition of maize root growth by high nitrate supply is correlated withreduced IAA levels in roots Journal of plant physiology 165 (9) 942-951


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Tooker JF and Moraes CM de (2011b) Feeding by Hessian fly (Mayetiola destructor [Say]) larvae on wheat increases levels offatty acids and indole-3-acetic acid but not hormones involved in plant-defense signaling Journal of plant growth regulation 30 (2)158-165


van Noorden GE Ross JJ Reid JB Rolfe BG and Mathesius U (2006) Defective long-distance auxin transport regulation wwwplantphysiolorgon June 1 2020 - Published by Downloaded from


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Woldemariam MG Onkokesung N Baldwin IT and Galis I (2012) Jasmonoyl-l-isoleucine hydrolase 1 (JIH1) regulatesjasmonoyl-l-isoleucine levels and attenuates plant defenses against herbivores The Plant Journal 72 (5) 758-767


Wu J and Baldwin IT (2009) Herbivory-induced signalling in plants perception and action Plant cell amp environment 32 (9)1161-1174


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Zhang P-J Li W-D Huang F Zhang J-M Xu F-C and Lu Y-B (2013) Feeding by whiteflies suppresses downstreamjasmonic acid signaling by eliciting salicylic acid signaling Journal of Chemical Ecology 39 (5) 612-619



Foliennummer 1

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Figure 1

Figure 2

Figure 3

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Figure 5

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Parsed Citations

2

Auxin is rapidly induced by herbivore attack and regulates a subset of systemic 5

jasmonate-dependent secondary metabolites 6

Ricardo AR Machado12 Christelle AM Robert12 Carla CM Arce123 Abigail P Ferrieri1 7

Shuqing Xu1 Guillermo H Jimenez-Aleman1 Ian T Baldwin1 and Matthias Erb12 8

9

1Max Planck Institute for Chemical Ecology Hans-Knoumlll-Str 8 07745 Jena Germany 10

2Institute of Plant Sciences University of Bern Altenbergrain 21 3013 Bern Switzerland 11

3Departamento de Entomologia Universidade Federal de Viccedilosa 36570-000 Viccedilosa MG 12

Brazil 13

14

One sentence summary Herbivory-induced auxin promotes the production of anthocyanins 15

and phenolamides 16

17

18

This work was supported by the Max Planck Society a Humboldt Postdoctoral Research 19

Fellowship (AF) the Brazilian National Council for Research CNPq Grant No 2379292012-20

0 (CA) a Marie Curie Intra European Fellowship Grant No 328935 (SX) a Marie Curie 21

Intra European Fellowship Grant No 273107 (ME) a Swiss National Foundation Fellowship 22

Grant No 140196 (CR) a European Research Council advanced Grant No 293926 (ITB) and 23

Human Frontier Science Program Grant No RGP00022012 (ITB) 24

25

26

Author for correspondence (Phone +41 31 631 8668 E-mail matthiaserbipsunibech)27


3

ABSTRACT 28

















45


4

INTRODUCTION 46


































5













(Xin et al 2012 Song 2014) 91













metabolites104


6

RESULTS 105






















2A to 2F) 127










7


attack 137


























8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








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2

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Con

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W+W

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3 min 7 min


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a

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3 min

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a

bP lt 0001


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Main root


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Elongated

0

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ab

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P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

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a

ba

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5


0

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2

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a a

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a a

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a a

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a a

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ba a

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a a

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a a

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Wild type

Impaired in

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0

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nt (micro

mol

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Stem color

plt0001

M sexta

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Ste

m a

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nt

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Ste

m a

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nt

(microm

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A B

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Control M sexta



0

15

30

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60

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Con

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IAA

Con

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IAA

Con

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IAA

Con

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A C

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Foliennummer 1

Foliennummer 1

Foliennummer 1

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Foliennummer 1

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Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

3

ABSTRACT 28

















45


4

INTRODUCTION 46


































5













(Xin et al 2012 Song 2014) 91













metabolites104


6

RESULTS 105






















2A to 2F) 127










7


attack 137


























8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

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AC

W+F

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IAA

(ng

gFW

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IAA

(ng

gFW

)

D



020406080

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Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

4

INTRODUCTION 46


































5













(Xin et al 2012 Song 2014) 91













metabolites104


6

RESULTS 105






















2A to 2F) 127










7


attack 137


























8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

5













(Xin et al 2012 Song 2014) 91













metabolites104


6

RESULTS 105






















2A to 2F) 127










7


attack 137


























8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

6

RESULTS 105






















2A to 2F) 127










7


attack 137


























8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

7


attack 137


























8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

8


































9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

9







stems 200















10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

10

DISCUSSION 214


































11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

11



































12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

12



































13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

13





METHODS 317




























14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

14










(n=5) 353






















15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

15
































16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

16




















2010) 423








Statistics 431






17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

17





















456


18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

18








Meldau et al 2009










Wang et al 2008



Paschold et al 2007

458

TABLE LEGENDS 463


FIGURE LEGENDS 465







treated plants 472


19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

19






plants 478




























20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

20



































21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

21


lanoline paste 539






sexta attack 545




549


22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

22

REFERENCES 550





















(12) 2383ndash2393 571







495 578





23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

23



120 (3) 474ndash481 584
























(7) 782ndash791 608








24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

24


7ndash12 616





9427 621





(6) 349ndash359 626























25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

25



























59 (6) 974ndash986 674







26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

26















164 694









1545ndash1556 703









27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

27








57 718










18512ndash18517 728






(2) 173ndash182 734



500 (7463) 422ndash426 737







28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

28









43 (3) 293ndash298 751





1795ndash1814 756



(4) 1369ndash1378 759









76 (3) 506ndash518 768








29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

29




100 (18) 10552ndash10557 778



2899ndash2907 781













157 (1) 341ndash354 794













4673ndash4680 807


30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

30












158ndash165 819





















2202 840


31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

31











biology (1) 2 851







Sciences 109 (19) E1192-E1200 858








0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

0

2

4

6

8

Control 3 6

0

1

2

3

Con

trol

W+W

W+F

AC

s

Con

trol

W+W

W+F

AC

s

3 min 7 min


IAA

(ng

gFW

)

a

a

b

a

b

c

0

1

2

3

4

C

ontro

l

W

+W

W

+OS

3 min

a

a

bP lt 0001


a

b

bP lt 0015

A B

C

W+F

AC

W+F

AC


0

1

2

3

4

Control 15 30 60 180

aa a

bb


A (n

ggF

W)

P lt 0001

IAA

(ng

gFW

)

IAA

(ng

gFW

)

D



020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

020406080

100

0 30 60 90 120

Stem


IAA

(ng

gFW

)


0

35

70

0 30 60 90 120

Main root


10

20

30

0 30 60 90 120

Lateral roots


01234

0 30 60 90 120

Petioles


012345

0 30 60 90 120

Systemic leaves


05

101520

0 30 60 90 120

Local leaves


ControlW+OS

A B

C D

E F


10 10

10

1010

10



01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations


01020304050

05 1 3

W+W


a

b

c

a

b

a

b

W+OS

Control ab

Early rosette

0

5

10

05 1 3


b

a

c

b

a

c

Elongated

0

2

4

6

05 1 3


a

b

a

a

ab

b

Flowering

IAA

(ng

gFW

)

A B C



0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

0

5

10

0123

0

2

4


YUCCA-like 3

YUCCA-like 9

a

YUCCA-like 3

a

b b

a a

b bP lt 0001

P lt 0001

a

b

c

Fold

cha

nge



A

B

005

115

YUCCA-like 5

a ab b

P lt 0001

C

E

G

0

1

2


a

b

a

D

P lt 0001


012345

Control 5 60

YUCCA-like 1

a

b

c

F

P lt 0001



IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

IAA

(ng

gFW

) JA (nggFW)

0

400

800

1200

1600

0

10

20

30

40

0 45 90

IAA Control

a

ba

b

A

b

a

A

B BJA Control



5


0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

0

1

2

3

4



a a

b

a a

b

a a

b

a a

b

a

c

ba a

b

a a

b

a a

b

a a

b

IAA

(ng

gFW

)



Wild type

Impaired in

JA-Ile-biosynthesis



0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

0

1

2

3

4

5

6

-1 0 1 2 3 4 5

Ste

m a

ntho

cyan

in c

onte

nt (micro

mol

gFW

)

Stem color

plt0001

M sexta

W+OSIAA+MeJA

W+WMeJAControl

IAA

0

1

2

3

4

5

Con

trol

W+W

W+O

SM

sex

taIA

AM

eJA

IAA+

MeJ

A

P lt 0001

0

4

8

12

16

Con

trol

W+O

SC

ontro

lW

+OS

Con

trol

W+O

SC

ontro

lW

+OS



Leaf treatment

0

1

2

3

Con

trol

W+O

SC

ontro

lW

+OS

EV irAOC


Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

Ste

m a

ntho

cyan

in c

onte

nt

(microm

olg

FW)

A B

C D

Control M sexta



0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

0

15

30

45

60

75

90

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


0

100

200

300

400

500

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA

Con

trol

IAA


Caffeoylputrescine


Ste

m c

onte

nt


Nicotine

DTGsP

eak

area

103

gFW


0

03

06

09

12

15

18

mg

gFW


ns

ns

ns

ns

ns

nsns

ns

ns

ns

0

60

120

180

240

300


microgg

FWmicrog

gFW


Leaf treatment

A C

B D

























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations























































































































































































































Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Foliennummer 1

Parsed Citations

Article File

Figure 1

Figure 2

Figure 3

Figure 4

Figure 5

Figure 6

Figure 7

Figure 8

Parsed Citations

Download - 1 Running Head: Auxin regulates herbivory-induced secondary metabolites - Plant … · Induced defense responses are activated by hormone-mediated 49 signaling cascades (Erb et

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