Mondal et al. Running Title: Actin-depolymerizing factor in plant … · Mondal et al. 3 47 ABSTRACT 48 The actin cytoskeleton network has an important role in plant cell growth,

Mondal et al.

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Running Title: Actin-depolymerizing factor in plant defense 1

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Author for Correspondence: Jyoti Shah 4

Department of Biological Sciences and BioDiscovery Institute, University of North Texas, 5

Denton, TX 76203, USA 6

Phone: (940) 565-3535; Fax: (940) 565-4136; E-mail: [email protected] 7

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Research Area: Signaling and Response 11

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Plant Physiology Preview. Published on November 13, 2017, as DOI:10.1104/pp.17.01438

Copyright 2017 by the American Society of Plant Biologists

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Arabidopsis ACTIN-DEPOLYMERIZING FACTOR3 is required for controlling aphid 16

feeding from the phloem 17

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Hossain A. Mondal1,3, Joe Louis1,4, Lani Archer1,2, Monika Patel1,2, Vamsi J. Nalam1,5, Sujon 19

Sarowar1, Vishala Sivapalan1, Douglas D. Root1, and Jyoti Shah1,2 20

21 1Department of Biological Sciences and 2BioDiscovery Institute, University of North Texas, 22

Denton, TX 76203, USA 23 3Uttar Banga Krishi Viswavidyalaya, Pundibari, Cooch Behar, India 24

4Department of Entomology and Department of Biochemistry, University of Nebraska, Lincoln, 25

NE 68583, USA 26 5Department of Biology, Indiana University-Purdue University, Fort Wayne, IN 46805, USA 27

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Present address for Sujon Sarowar: Botanical Genetics, Buffalo, NY 14203, USA 29

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One sentence summary: Green peach aphid feeding from the sieve elements is shown to be 31

restricted by ACTIN-DEPOLYMERIZING FACTOR3, thus implicating actin-dependent process 32

in controlling insect feeding from the phloem. 33

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Author contributions: HAM, JL, LA, MP, DDR and JS conceived and designed the study. 35

HAM, JL, LA, MP, VJM, SS and VS conducted the experiments, and analyzed and interpreted 36

the data. DDR contributed to methods development and data analysis. The manuscript was 37

written by HAM, JL and JS with input from all authors. 38

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Funding information: This work was partially supported by a grant from the National Science 40

Foundation (IOS-0919192) to JS. MP was supported by graduate assistantship from the 41

University of North Texas. 42

43

The author responsible for distribution of materials integral to the findings presented in this 44

article in accordance with the Journal policy described in the Instructions for Authors 45

(http://www.plantphysiol.org) is Jyoti Shah ([email protected]). 46



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ABSTRACT 47

The actin cytoskeleton network has an important role in plant cell growth, division and stress 48

response. Actin-depolymerizing factors (ADFs) are a group of actin-binding proteins that 49

contribute to reorganization of the actin network. Here we show that the Arabidopsis thaliana 50

ADF3 is required in the phloem for controlling infestation by Myzus persicae Sülzer, commonly 51

known as the green peach aphid (GPA), which is an important phloem sap-consuming pest of 52

more than fifty plant families. In agreement with a role for the actin-depolymerizing function of 53

ADF3 in defense against the GPA, we show that resistance in adf3 was restored by 54

overexpression of the related ADF4, and the actin cytoskeleton destabilizers, cytochalasin D and 55

latrunculin B. Electrical monitoring of the GPA feeding behavior indicates that the GPA stylets 56

found sieve elements faster when feeding on the adf3 mutant compared to the wild-type (WT) 57

plant. In addition, once they found the sieve elements, the GPA fed for a more prolonged period 58

from sieve elements of adf3 compared to the WT plant. The longer feeding period correlated 59

with an increase in fecundity and population size of the GPA and a parallel reduction in callose 60

deposition in the adf3 mutant. The adf3-conferred susceptibility to GPA was overcome by 61

expression of the ADF3 coding sequence from the phloem-specific SUC2 promoter, thus 62

confirming the importance of ADF3 function in the phloem. We further demonstrate that the 63

ADF3-dependent defense mechanism is linked to the transcriptional upregulation of 64

PHYTOALEXIN-DEFICIENT4, which is an important regulator of defenses against the GPA. 65

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INTRODUCTION 67

In eukaryotes, the actin cytoskeleton, which is composed of filamentous (F)-actin, has a 68

central role in multiple cellular processes, including cell growth, division and differentiation, 69

regulation of polarity, and facilitating cytoplasmic streaming, organelle movement and response 70

to the environment (Staiger, 2000; Hussey et al., 2006; Staiger and Blanchoin, 2006; Pollard and 71

Cooper, 2009; Szymanski and Cosgrove, 2009; Day et al., 2011; Henty-Ridilla et al., 2013). 72

This microfilament network is dynamic and requires continuous reorganization. Remodeling of 73

the actin network involves severing, depolymerization and polymerization of F-actin, which 74

needs to be spatially and temporally regulated. A variety of actin-binding proteins are involved 75

in remodeling of the actin cytoskeleton (Hussey et al., 2006). These include the actin-nucleating 76

and filament stabilizing proteins like the formins and fimbrins, respectively, and the actin-77

depolymerizing factor (ADF) family of proteins (Vidali et al., 2009; Ye et al., 2009; Wu et al., 78

2010). As a result of their ability to sever and depolymerize F-actin and yield products with ends 79

that can serve as sites for new filament initiation, the ADFs, which are small proteins 80

(approximately 17 kDa), increase the dynamics of the actin cytoskeleton and the balance of F-81

actin versus the free globular (G)-actin (Andrianantoandro and Pollard, 2006; Pavlov et al., 82

2007). ADF’s are also involved in shuttling actin into the nucleus (Nebl et al., 1996; Jiang et al., 83

1997), where actin is a component of chromatin remodeling activities that control gene 84

expression (Farrants, 2008; Jockusch et al., 2006). In addition to the cytosol, some ADF’s also 85

localize to the nucleus (Ruzicka et al., 2007). 86

The ADF family in Arabidopsis thaliana consists of eleven expressed genes, which based 87

on their relatedness to each other have been grouped into four subclasses (Feng et al., 2006). 88

These ADF subclasses exhibit novel and differential tissue-specific and developmental 89

expression patterns (Ruzicka et al., 2007). For example, the subclass I genes, which include 90

ADF1, ADF2, ADF3 and ADF4, are constitutively expressed in a variety of vegetative and 91

reproductive tissues, including roots, leaves, flowers, and young seedlings. Amongst the 92

subclass I genes, ADF3 was the most strongly expressed. The subclass II genes can be 93

subdivided into clade IIa (ADF7 and ADF10) and clade IIb (ADF8 and ADF11). ADF7 and 94

ADF10 exhibit high levels of expression in the reproductive tissues, with strongest expression in 95

mature pollen grain. In comparison, expression of the clade IIa genes is relatively poor in roots 96



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and leaves. By contrast, the clade IIb ADF8 and ADF11 genes, show highest expression in 97

epidermal cells that develop into root hairs. ADF5 and ADF9, which belong to subclass III, 98

exhibit strongest expression in the fast growing tissues, including the root meristem and 99

emerging leaves. ADF6, which is the lone member of subclass IV, is expressed in all tissues 100

including pollen. Biochemical characterization of Arabidopsis ADFs indicated that all Class I, II 101

and IV ADFs possess F-actin-severing/depolymerizing activity, with Class I ADFs possessing 102

the strongest activities (Nan et al., 2017). In comparison, the class III ADFs lacked F-actin-103

severing/depolymerizing activity, and instead possessed F-actin bundling activity (Nan et al., 104

2017). 105

ADFs and actin cytoskeleton dynamics are involved in plant response to pathogens. In 106

Arabidopsis, the actin polymerization inhibitor cytochalasin E attenuated non-host resistance 107

against Blumeria graminis f. sp. tritici (Yun et al., 2003). Cytochalasin E also interfered with the 108

targeting of the resistance protein RPW8.2 to the extrahaustorial membrane in Arabidopsis 109

inoculated with the powdery mildew Golovinomyces spp fungi (Wang et al., 2009). Similarly, 110

RPW8.2 localization was also affected in plants overexpressing ADF6, thus suggesting that the 111

specific targeting of RPW8.2 to the extrahaustorial membrane is dependent on actin cytoskeleton 112

dynamics (Wang et al., 2009). Arabidopsis ADF4 is required for resisting infection by the 113

bacterial pathogen Pseudomonas syringae pv tomato DC3000 expressing the AvrPphB effector 114

protein (Tian et al., 2009). ADF4 function in defense against this pathogen was linked to the 115

transcriptional regulation of the Arabidopsis RPS5 gene, which encodes a resistance protein that 116

facilitates the recognition of the AvrPphB effector protein and the activation of downstream 117

defense signaling (Porter et al., 2012). Thus, it was suggested that ADF4 links actin cytoskeleton 118

dynamics to pathogen perception and defense activation (Porter et al., 2012). ADF4 was also 119

shown to regulate actin dynamics and callose deposition in response to elf26, a microbial elicitor 120

of immunity (Henty-Ridilla et al., 2014). In contrast, knockdown of ADF4 resulted in enhanced 121

resistance against an Arabidopsis adapted strain of the powdery mildew fungus (Inada et al., 122

2016), while knockdown of ADF2 resulted in enhanced resistance to the root-knot nematode 123

Meloidogyne incognita (Clément et al., 2009). Hence, there is a wider involvement of ADFs in 124

plant defense as well as susceptibility to pests. 125



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Aphids (Hemiptera: Aphididae) encompass a large group of insects that consume phloem 126

sap. Nearly 250 species of aphids are considered as pests of plants that limit plant growth and 127

productivity due to removal of nutrients from sieve elements and their ability to alter source-sink 128

patterns, and thus the flow of nutrients to growing parts of the infested plant (Pollard, 1973; 129

Blackman and Eastop, 2000; Goggin, 2007). Furthermore, some aphids also vector viral 130

diseases (Kennedy et al., 1962; Matthews, 1991; Guerrieri and Digilio, 2008). Aphids utilize 131

their mouthparts, which are modified into slender stylets, to feed from the sieve elements. Prior 132

to inserting stylets into the plant tissue, aphids utilize chemosensory hairs on their antennae to 133

assess the plant surface, potentially for gustatory cues (Powell, 2006). Subsequently the stylets 134

briefly penetrate non-vascular cells to sample cell contents for additional gustatory cues. The 135

puncturing of non-vascular cells along the path of the stylet penetration likely results in the 136

activation of host defenses that could potentially interfere with the ability of the stylet to reach a 137

sieve element. Aphids produce two distinct salivary secretions that facilitate infestation (Miles, 138

1999). The proteinaceous gelling saliva, which is released by the stylets on their way to the 139

sieve elements, forms a sheath that facilitates stylet movement through the plant tissue and 140

simultaneously minimizes wound-responses in the plant by quickly sealing off wounds. In 141

comparison, the watery saliva, which is released when the stylets penetrate the sieve elements, 142

contains factors suggested to enable the insect to prevent and maybe reverse phloem occlusion 143

and thus facilitate feeding from the sieve elements (Miles, 1999; Will et al., 2007, 2009). 144

The plant surface provides the first line of defense (e.g. trichomes and glandular 145

secretions) that could deter aphid settling on a plant (Walling, 2008). In addition, during the 146

different stages of stylet penetration into the plant tissue, the insect encounters defenses that 147

deter feeding and adversely impact insect fecundity (Walling, 2008; Louis and Shah, 2013). 148

These include factors that contribute to sieve element occlusion (e.g. callose deposition and 149

phloem protein aggregation) as well as insecticidal factors present in the phloem sap (Pedigo, 150

1999; Smith, 2005; Powell et al., 2006; Goggin, 2007; Walling, 2008). The interaction between 151

Arabidopsis and the green peach aphid (GPA; Myzus persicae Sülzer) has been utilized as a 152

model system to characterize plant genes and mechanism that contribute to defense (Louis et al., 153

2012; Louis and Shah, 2013). GPA is a polyphagous insect that is an important pest of more 154

than 400 plant species belonging to over 50 plant families, and the vector of more than 100 viral 155

diseases (Kennedy et al., 1962; Matthews, 1991; Blackman and Eastop, 2000). The 156



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PHYTOALEXIN-DEFICIENT 4 (PAD4) gene is an important regulator of Arabidopsis defenses 157

that deter GPA feeding and adversely impact GPA fecundity (Pegadaraju et al., 2005, 2007; 158

Louis et al., 2010a). Genetic studies have indicated that the upregulation of PAD4 expression in 159

response to GPA infestation correlated with PAD4’s involvement in deterring GPA feeding from 160

sieve elements. This feeding deterrence was intensified in plants overexpressing PAD4 from the 161

Cauliflower mosaic virus 35S gene promoter (Pegadaraju et al., 2007). In contrast, basal 162

expression of PAD4 was sufficient for the accumulation of an antibiotic activity that adversely 163

impacts insect fecundity (Louis et al., 2010a, 2012). 164

ADF proteins, and profilin, an actin-binding protein that influences actin polymerization, 165

have been identified in phloem exudates from a variety of plants (Schobert et al., 1998, 2000; 166

Kulikova and Puryaseva, 2002; Lin et al., 2009; Rodriguez-Medina, 2011; Fröhlich et al., 2012). 167

Furthermore, microfilament meshwork have been revealed in sieve elements of fava bean (Vicia 168

faba) injected with the actin-binding fluorescent phalloidin (Hafke et al., 2013). Immunolabeling 169

with anti-actin antibodies further confirmed the presence of actin cytoskeleton in the sieve 170

elements (Hafke et al., 2013). The identification of an actin-binding protein in aphid saliva has 171

led to the suggestion that an actin-dependent process contributes to plant defense in the phloem 172

and aphids attempt to curtail these defenses by targeting actin dynamics (Nicholson et al., 2012). 173

We therefore investigated the contribution of ADF genes in the interaction of Arabidopsis with 174

the GPA. We show that an ADF3-dependent mechanism is required for controlling GPA feeding 175

from the sieve elements. We further demonstrate that the PAD4 gene is a critical downstream 176

component of this ADF3-dependent defense mechanism. 177

178

RESULTS 179

ADF3 is Required for Limiting GPA Infestation on Arabidopsis 180

To determine if the ADF genes influence infestation of Arabidopsis by the GPA, 181

Arabidopsis lines that were previously shown to lack or accumulate reduced levels of the ADF1 182

(At3g46010), ADF2 (At3g46000), ADF3 (At5g59880), ADF4 (At5g59890), ADF5 (At2g16700) 183

and ADF9 (At4g34970) transcripts (Clément et al., 2009; Tian et al., 2009) were utilized in no-184



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choice bioassays with the GPA. For the no-choice bioassay, twenty adult apterous asexually 185

reproducing insects were released on each plant and the insect population size (adults + nymphs) 186

determined two days post infestation (dpi). As shown in Figure 1A and 1B, only plants lacking 187

ADF3 function repeatedly showed GPA numbers that were significantly higher than GPA 188

numbers on the wild-type (WT) plant. Compared to the WT plant, the GPA population was 189

significantly larger (P < 0.05) on the adf3-1 (Salk_139265) and adf3-2 (SAIL_501_F01) 190

mutants. adf3-1 and adf3-2 contain T-DNA insertions within the first intron and last exon of 191

ADF3, respectively (Supplemental Fig. S1A), which is associated with reduced accumulation of 192

the ADF3 transcript in these mutant lines compared to the WT (Supplemental Fig. S1B). The 193

increase in GPA population on the adf3-1 mutant correlated with the significantly higher 194

fecundity of GPA on the mutant compared to the WT (Fig. 1C). Compared to an average of 1.2 195

nymphs/day produced by a GPA on the WT plant, 1.9 nymphs/day were produced on the adf3-1 196

mutant. 197

To confirm that loss of ADF3 function is indeed responsible for the better performance of 198

the GPA on the adf3 plants compared to the WT plants, a genomic clone of ADF3 was 199

transformed into the adf3-1 mutant. As shown in Figure 1D, ADF3 expression was restored in 200

two independently-derived gADF3 transgenic lines. In comparison to the adf3-1 mutant, the 201

GPA population size was significantly lower on these gADF3 transgenic lines and comparable to 202

that on the WT plants (Fig. 1D). Similarly, resistance was restored in adf3-1 plants expressing 203

ADF3 from the heterologous Cauliflower mosaic virus 35S gene promoter. The GPA population 204

size on two independently-derived 35Spro:ADF3 (in adf3-1 background) lines was significantly 205

smaller than that on the adf3-1 mutant, and comparable to that on the WT plant (Fig. 1E and 206

Supplemental Fig. S2). Taken together, these results confirm that ADF3 has a critical role in 207

limiting GPA infestation on Arabidopsis. 208

209

Actin Cytoskeleton Destabilizers Restore Resistance to the adf3 Mutant 210

ADF3 is an actin-binding protein that was recently shown to possess F-actin-211

severing/depolymerizing activity (Nan et al., 2017). F-actin depolymerization assays conducted 212

by us confirm the ability of ADF3 to depolymerize F-actin (Supplementary Fig. S3). To 213



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determine if ADF3-dependent actin reorganization is critical for controlling GPA infestation, the 214

ability of the actin cytoskeleton destabilizers cytochalasin D and latrunculin B to compensate for 215

ADF3 deficiency in the adf3-1 mutant was evaluated. Two adult aphids were caged onto 216

cytochalasin D- and latrunculin B-treated, and as control on dimethyl sulfoxide (DMSO)-treated 217

leaves of adf3-1 and WT plants, and the number of nymphs born per adult aphid monitored two 218

days later. As expected, significantly larger number of nymphs were born on the DMSO-treated 219

adf3-1 compared to the WT plant (Fig. 2A). However, compared to DMSO treatment, 220

significantly fewer nymphs were born on adf3-1 leaves that were treated with cytochalasin D and 221

latrunculin B. The number of nymphs born on the cytochalasin D- and latrunculin B-treated 222

adf3-1 leaves was comparable to that observed on the cytochalasin D- and latrunculin B-treated 223

leaves of WT plants, thus indicating that these actin destabilizers compensate for the lack of 224

ADF3 function in the adf3-1 mutant. Taken together, these results confirm the importance of 225

ADF3’s actin-depolymerizing function in Arabidopsis defense against the GPA. 226

To further test the involvement of ADF3’s actin-depolymerizing function in controlling 227

GPA infestation, we tested if the adf3-1 defect could be compensated by overexpression of 228

another actin depolymerizing factor, ADF4 (Henty et al., 2011; Henty-Ridilla et al., 2014; Nan et 229

al., 2017), which exhibits 83% identity and 91% similarity to ADF3 (Supplemental Fig. S4). A 230

previously described 35Spro:ADF4 construct (Henty-Ridilla et al., 2014), in which the ADF4 231

protein coding sequence is expressed from the 35S promoter, was transformed into the adf3-1 232

mutant. No choice assays were conducted on three independently-derived adf3-1 35Spro:ADF4 233

lines, and as a control on the WT and adf3-1 mutant. As shown in Figure 2B, GPA population 234

size was comparable on the WT and the adf3-1 35Spro:ADF4 plants, and significantly lower than 235

that on the adf3-1 mutant, thus confirming the ability of ADF4 overexpression to overcome the 236

adf3-1 defect in limiting GPA infestation. Collectively, the above results in conjunction with the 237

studies of Nan et al. (2017) lead us to conclude that the impact of ADF3 on controlling GPA 238

population size is linked to ADF3’s function in actin cytoskeleton reorganization. 239

240

ADF3 Expression in the Phloem is Required for Controlling GPA Infestation 241



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ADF3 expression was reported to be the strongest amongst the ADF family of genes in 242

Arabidopsis (Ruzicka et al., 2007). Gene expression data available on the Arabidopsis eFP 243

Browser (http://bbc.botany.utoronto.ca/efp/cgi-bin/efpWeb.cgi) revealed that ADF3 is expressed 244

throughout the development of Arabidopsis in most organs, except pollen. Furthermore, biotic 245

stress, including GPA infestation did not have a pronounced effect on ADF3 expression 246

(http://bbc.botany.utoronto.ca/efp/cgi-bin/efpWeb.cgi). Real-time RT-PCR analysis confirmed 247

that ADF3 expression was not significantly different between uninfested and GPA-infested 248

plants (Fig. 3A). Histochemical analysis for GUS activity in leaves of transgenic ADF3pro:UidA 249

plants in which the bacterial UidA-encoded GUS reporter was expressed from the ADF3 250

promoter, further confirmed that ADF3 promoter activity was comparable between uninfested 251

and GPA-infested leaves (Supplemental Fig. S5). ADF3 promoter activity was strong in the 252

lateral and minor veins compared to the non-vascular tissues (Fig. 3B and Supplemental Fig. S5). 253

Within the vasculature, GUS activity was found to be strong in the phloem (Fig. 3C). 254

To test if ADF3 expression in phloem is important for ADF3’s role in controlling GPA 255

infestation, we tested if the ADF3 coding sequence expressed from the phloem-specific SUC2 256

promoter (Gottwald et al., 2000) was sufficient to restore resistance against the GPA in the adf3-257

1 mutant background. Transgenic adf3-1 plants expressing the ADF3 coding sequence from the 258

35S promoter provided the positive controls for this experiment. As shown in Figure 3D, the 259

SUC2pro:ADF3 chimera complemented the adf3-1 defect. GPA population size was significantly 260

lower on three independently-derived adf3-1 SUC2pro:ADF3 lines compared to the adf3-1 261

mutant. The level of resistance observed in the adf3-1 SUC2pro:ADF3 lines was comparable to 262

that observed in adf3-1 35Spro:ADF3 line. There results confirm an important role for ADF3 in 263

the phloem in controlling GPA infestation on Arabidopsis. 264

265

ADF3 is Required for Limiting GPA Feeding From Sieve Elements 266

The adverse effect of ADF3 on insect fecundity could result from its effect on the 267

accumulation of antibiosis activity, or alternatively due to its impact on the insects feeding 268

behavior. Previous studies have indicated the presence of an antibiotic activity, which is 269

detrimental to GPA fecundity, in leaf petiole exudates that are enriched in phloem sap (Louis et 270



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al., 2010a; Nalam et al., 2012). To determine if ADF3 controls the accumulation of this antibiotic 271

activity, petiole exudates collected from leaves of WT, and the adf3-1 and adf3-2 mutants were 272

added to a synthetic diet and GPAs were reared on the supplemented diet for five days. As 273

shown in Figure 4, in comparison to the GPA population on the synthetic diet lacking petiole 274

exudates, insect population size was significantly smaller (P < 0.05) on diet supplemented with 275

petiole exudates from WT plants. The GPA population size was similarly lower on diet 276

supplemented with petiole exudates collected from the adf3-1 and adf3-2 mutant plants, thus 277

confirming that ADF3 does not significantly influence the accumulation of this antibiotic activity 278

in the phloem sap-enriched petiole exudates. 279

To test if ADF3 adversely influences GPA feeding, GPA feeding behavior was monitored 280

on leaves of the WT and adf3-1 mutant plants with the Electrical Penetration Graph (EPG) 281

technique in which the plant and the insect, with a wire glued to its dorsum, are part of a low-282

voltage circuit (van Helden and Tjallingii, 2000). The different waveform patterns in an EPG 283

are characteristic of the different feeding behavioral activities of the insect. EPG provides 284

information on the time the insect takes to first probe (FP) the plant with its stylets, the time 285

spent by the insect to find and tap into a sieve element for the first time (f-SEP; first sieve 286

element phase), and the sum of time spent during the recording period by the insect in all the 287

SEPs (s-SEP), in the xylem phase (XP) when the stylet is in the xylem and the insect is 288

consuming xylem sap, the pathway phase (PP) when the stylet is inserted into the leaf tissue but 289

is outside the sieve elements and likely sampling other cells, and the non-probing (NP) phase 290

when the stylet is not inserted into the plant tissue. As shown in Figure 5A, comparison of these 291

behavioral activities of the GPA on the WT and the adf3-1 mutant revealed that the insect spent 292

significantly less time (P < 0.05) attaining the f-SEP on the adf3-1 mutant. In addition, the s-293

SEP was significantly longer (P < 0.05) when the insects were on the adf3-1 mutant compared to 294

the WT, thus indicating that the insects spent significantly more time feeding from the sieve 295

elements of the adf3-1 mutant than the WT plant. A corresponding reduction in the time spent 296

by GPA in PP was observed on the adf3-1 mutant compared to the WT plant, thus confirming 297

that the insect encounters fewer obstacles that allow it to find sieve elements faster and feed for 298

longer periods on the adf3-1 mutant. In contrast to f-SEP, s-SEP and PP, the insect took 299

comparable amount of time to reach the FP, and spent comparable time in the NP and XP on the 300



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WT and adf3-1 mutant. These results suggest that ADF3 and/or an ADF3-dependent 301

mechanism(s) interferes with the ability of the GPA to find and feed from the sieve elements. 302

During the SEP, the watery saliva injected by the insect into the sieve elements may 303

inhibit phloem-sealing mechanisms (Powell et al., 2006; Tjallingii, 2006; Will and van Bel, 304

2006). This watery saliva injection phase, the E1 phase, is followed by the E2 phloem sap-305

ingestion phase. A prolonged E1 and/or reduced E2 is suggestive of a lowered ability of the 306

insect to suppress rapid phloem-sealing mechanisms (Garzo et al., 2002). To determine if insects 307

feeding from sieve elements of the adf3 mutant encounter less resistance that results in a shorter 308

duration of watery saliva release into the sieve elements, the length of the E1 waveform in the 309

first SEP was compared between insects released on the WT and the adf3-1 mutant. As shown in 310

Figure 5B, the length of the E1 phase was comparable between insects placed on the WT and 311

adf3-1 mutant, thus indicating that ADF3 does not influence the length of the period of watery 312

saliva release by GPA into Arabidopsis sieve elements. However, the E2 phase was significantly 313

shorter (P < 0.05) on the WT compared to the adf3-1, thus confirming that ADF3 and/or an 314

ADF3-dependent mechanism in the WT plant limits ingestion of phloem sap by the GPA. 315

316

ADF3 Controls Callose Deposition in GPA-infested Plants 317

Callose deposition is one of the processes involved in phloem occlusion (Will and van 318

Bel, 2006; Kempema et al., 2007; Hao et al., 2008). Similarly, callose deposited outside cells that 319

are in the path of the stylets trying to reach the sieve element, could interfere with the ability of 320

the stylets to reach sieve elements. Considering that ADF3 is required for limiting GPA feeding, 321

and callose synthesis is subject to the participation of the actin cytoskeleton (Cai et al., 2010), we 322

tested the impact of ADF3 on callose deposition in GPA-infested plants. As shown in Figure 6, 323

in response to GPA infestation a significant increase in the number of callose spots was observed 324

in WT plants. In comparison to the WT, the number of callose deposits were lower in the GPA-325

infested leaves of adf3-1 mutant. The number of callose spots in the GPA-infested adf3-1 mutant 326

was comparable to that observed in the GPA-infested pad4-1 mutant (Fig. 6), which like adf3-1, 327

is also defective in controlling GPA feeding from the sieve elements (Pegadaraju et al., 2007; 328



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Louis et al., 2012), thus further relating callose deposition with the ability of the plant to control 329

GPA feeding. 330

331

An ADF3-dependent Mechanism Controls PAD4 Expression in GPA-infested Plants 332

In the nucleus, actin is a component of the chromatin remodeling complexes that function 333

to regulate gene expression (Jockusch et al., 2006; Farrants, 2008). Furthermore, some ADFs are 334

involved in shuttling actin into the nucleus and thus in regulating gene expression (Nebl et al., 335

1996; Jiang et al., 1997). We noted that the upregulation of PAD4 expression, which is observed 336

in the GPA-infested leaves of the WT, was attenuated in the adf3-1 mutant. As shown in Figure 337

7A, in comparison to the significant (P<0.05) increase in PAD4 expression observed in GPA-338

infested compared to uninfested WT plants, in the adf3-1 mutant, PAD4 transcript levels did not 339

exhibit a significant increase in response to GPA infestation. These results suggest that an 340

ADF3-dependent mechanism regulates PAD4 transcript accumulation in GPA-infested WT 341

plants. 342

To further define the relationship between ADF3 and PAD4 in Arabidopsis defense 343

against the GPA, adf3-1 35Spro:PAD4 and pad4-1 35Spro:ADF3 plants in which PAD4 and 344

ADF3, respectively, are expressed from the 35S promoter were generated. adf3-1 35Spro:ADF3 345

plants provided the controls. As expected, ADF3 expression from the 35S promoter restored 346

basal resistance against GPA in the adf3-1 35Spro:ADF3 plants (Fig. 7B and Supplemental Fig. 347

S6). By comparison, in two independently-derived transgenic lines, ADF3 expression from the 348

35S promoter was unable to restore resistance in the absence of a functional PAD4 gene in the 349

pad4-1 35Spro ADF3 plants (Fig. 7B and Supplemental Fig. S6), thus suggesting a critical role 350

for PAD4 in ADF3-dependent defense against the GPA. Moreover, in no-choice assays 351

conducted on plants of two independent adf3-1 35Spro PAD4 lines, GPA population size was 352

comparable to that on the WT plants, but smaller than on the adf3-1 mutant (Fig. 7B and 353

Supplemental Fig. S6), thus indicating that overexpression of PAD4 from the heterologous 35S 354

promoter is sufficient to restore significant level of resistance in the absence of ADF3 function. 355

Collectively, these results reaffirm the importance of ADF3-dependent regulation of PAD4 356

expression in Arabidopsis defense against the GPA. 357



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358

DISCUSSION 359

The actin cytoskeleton and actin-binding proteins are present in the phloem (Schobert et 360

al., 1998, 2000; Kulikova and Puryaseva, 2002; Lin et al., 2009; Rodriguez-Medina, 2011; 361

Fröhlich et al., 2012; Hafke et al., 2013). However, their biological function in the phloem is 362

poorly understood. We provide multiple lines of evidence indicating an important role for ADF3 363

and actin reorganization in Arabidopsis defense against the GPA, particularly in controlling GPA 364

feeding from the sieve elements. (i) Compared to the WT plant, GPA population was larger on 365

adf3 mutant plants. (ii) The adf3 deficiency in controlling GPA infestation was compensated by 366

the actin cytoskeleton destabilizers cytochalasin D and latrunculin B, and by overexpression of 367

actin depolymerizing factor ADF4. (iii) ADF3 was expressed in the phloem and expression of 368

ADF3 from the phloem-specific SUC2 promoter was sufficient to restore resistance against the 369

GPA in the adf3-1 mutant. (iv) In agreement with it functioning in the phloem, ADF3 was 370

required for limiting GPA feeding from the sieve elements. Our results further demonstrate that 371

ADF3’s involvement in controlling GPA infestation is in part mediated via its influence on 372

PAD4 expression and hence defense signaling. 373

EPG analysis indicated that besides limiting GPA feeding from sieve elements, an ADF3-374

dependent mechanism interferes with the ability of the insect stylets to reach the sieve elements. 375

Thus, we propose that an ADF3-dependent process also impacts events occurring prior to the 376

first penetration of a sieve element by the aphid stylets. On their way to finding a sieve element, 377

the stylets penetrate and sample contents of non-vascular cells (Pollard, 1973; Powell et al., 378

2006). These punctures of plant cells by the stylets could stimulate plant defenses. Salivary 379

components that are released into the plant tissue are also known to stimulate host defenses 380

(DeVos and Jander, 2009; Bos et al., 2010). For example, callose deposition increases in 381

response to GPA infestation (Elzinga et al., 2014) as well as application of GPA-derived elicitors 382

(Prince et al., 2014). Callose has been implicated in the control of infestation by insects that feed 383

from the phloem (Will and van Bel, 2006; Kempema et al., 2007; Hao et al., 2008). Callose 384

deposited in the sieve elements interferes with the ability to the aphid to feed from the sieve 385

elements. In addition, callose deposited outside the sieve elements could interfere with the ability 386



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15

of the stylet to reach sieve elements to further limit insect feeding. The GPA infestation-387

associated increase in callose deposition was attenuated the adf3-1 mutant, suggesting an 388

important role for ADF3 in this process. The actin cytoskeleton is involved in the localization of 389

callose synthases to the cell membrane, presumably due to the involvement of the actin 390

cytoskeleton in vesicular trafficking (Cai et al., 2010). Whether ADF3 is similarly involved in 391

the localization of callose synthase is not known. However, since the GPA infestation-associated 392

increase in callose deposition was also attenuated in the pad4 mutant, we suggest that the impact 393

of ADF3 on callose deposition in response to GPA infestation is likely exerted via ADF3’s 394

engagement of the PAD4 defense signaling pathway. The contribution of PAD4 in promoting 395

callose deposition is further supported by a recent study, which showed that the endophytic 396

Bacillus velezensis-induced resistance against the GPA was accompanied by an increase in 397

callose deposition, which was dependent on PAD4 (Rashid et al., 2017). 398

Actin is a component of chromatin remodeling activities that control gene expression in 399

the nucleus (Jockusch et al., 2006; Farrants, 2008), and ADF’s are involved in shuttling actin 400

into the nucleus (Nebl et al., 1996; Jiang et al., 1997) and in regulating gene expression (Burgos-401

Rivera et al., 2008; Porter et al., 2012). Amongst the subclass I ADFs in Arabidopsis, ADF4’s 402

involvement in effector-triggered immunity is linked to downstream activation of gene 403

expression (Porter et al., 2012). Similarly, our results indicate that ADF3’s involvement in 404

defense against the GPA is linked to the upregulation of PAD4. However, although PAD4 is 405

required for ADF3-conferrred resistance to the GPA, our results also indicate that ADF3 and 406

PAD4 have additional functions in Arabidopsis defense against the GPA that are independent of 407

each other. For example, unlike PAD4, an ADF3-dependent mechanism hinders with the ability 408

of the insect to find sieve elements. In contrast, unlike ADF3, a PAD4-dependent mechanism is 409

required for the accumulation of an antibiosis activity in the vascular sap (Pegadaraju et al., 410

2005, 2007; Louis et al., 2010a). Previous studies have shown that while basal expression of 411

PAD4 contributes to the antibiosis activity, the upregulation of PAD4 expression is associated 412

with the feeding deterrence function of the PAD4-dependent pathway (Pegadaraju et al., 2007; 413

Louis et al., 2010a, 2010b). These conclusions are in agreement with the observations described 414

here that ADF3, which is required for the upregulation of PAD4 expression in response to GPA 415

infestation, but not for the basal expression of PAD4, is required for controlling GPA feeding, 416

but not for the accumulation of the antibiotic activity. It is likely that different cell types are 417



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16

involved in exerting these effects. For example, the contribution of ADF3 and PAD4 to feeding 418

deterrence is exerted at the level of the sieve elements, while ADF3’s involvement in hindering 419

with the ability of the GPA to find sieve elements is exerted outside the sieve elements. 420

Similarly, PAD4’s contribution to antibiosis is likely exerted in tissues where these antibiotic 421

factors are produced. 422

ADF3 (At5g59880) and ADF4 (At5g59890), which are located in tandem on 423

chromosome 5, are both subclass I ADFs that share high level of sequence identity. Both possess 424

actin-severing/depolymerizing activity (Henty-Ridilla et al., 2014; Nan et al., 2017). However, 425

while ADF3, not ADF4, is essential for defense against the GPA, ADF4, but not ADF3, is 426

required for AvrPphB -triggered immunity against the bacterial pathogen P. syringae (Tian et al., 427

2009). The ability of ADF4 expressed from the strong and ubiquitously expressed 35S promoter 428

to limit GPA population in the adf3-1 mutant background indicates that ADF3 and ADF4 have 429

overlapping biochemical functions. We therefore conclude that the unique biological functions of 430

ADF3 in defense against the GPA is likely determined by differences in the spatial expression 431

pattern and/or overall level of expression of ADF3, compared to ADF4, and likely other subclass 432

I ADF genes. 433

434

CONCLUSION 435

Our results demonstrate that an ADF3-dependent mechanism hinders with the ability of 436

the GPA to find and feed from sieve elements. Considering that ADF3 is required for 437

upregulating PAD4 expression and callose deposition in GPA-infested leaves, we postulate that 438

an ADF3-dependent mechanism is involved in signaling associated with Arabidopsis defense 439

against the GPA. 440

441

MATERIAL AND METHODS 442

Plant and Insect Materials 443



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17

All plants were cultivated at 22°C under a 14 h light (100 μE m2 s-1) and10 h dark regime 444

on a peat-based soil mix (Fafard #2, fafard.com). The Arabidopsis adf1 (Salk_144459) (Tian et 445

al., 2009), adf2 (RNAi line; Clément et al., 2009), adf3-1 (Salk_139265C) (Tian et al., 2009), 446

adf3-2 (SAIL_501_F01), adf4 (Garlic_823_A11.b.1b.Lb3Fa) (Tian et al., 2009), adf5 447

(Salk_030145C), and adf9 (Salk_056064) (Tian et al., 2009) lines were used in this study. 448

Silencing of ADF2 in the ADF2 RNAi line was induced by treating plants with 0.5% ethanol 449

solution (Clément et al., 2009). 450

A GPA (Kansas State University, Museum of Entomological and Prairie Arthropod 451

Research, voucher specimen 194) colony was reared at 22°C under a 14 h light (100 μE m2 s-1) 452

and10 h dark regime on a 1:1 mixture of commercially available radish (Raphanus sativus) 453

(Early Scarlet Globe) and mustard (Brassica juncea) (Florida Broadleaf). 454

455

No-choice Tests 456

No-choice assays were conducted as previously described (Pegadaraju et al., 2005; Louis 457

et al., 2010a). Unless stated otherwise, 20 adult asexually reproducing apterous (wingless) 458

aphids were placed on each 4-week-old plant. Two days later, the number of insects on each 459

plant was counted. To monitor aphid fecundity, one young nymph (3-4 day old) was placed on 460

each plant and the total number of progeny born over a 10-day period was recorded. 461

462

Monitoring Aphid Feeding Behavior 463

The electrical penetration graph (EPG) technique (van Helden and Tjallingii, 2000; 464

Walker, 2000) was used to simultaneously monitor the feeding behavior of GPA on 4-week-old 465

plants of two different genotypes, as previously described (Pegadaraju et al., 2007). An eight-466

channel GIGA-8 direct current amplifier (http://www.epgsystems.eu/systems.htm) was used for 467

simultaneous recordings of eight individual aphids. The waveform recordings were analyzed 468

using the EPG analysis software PROBE 3.4 (provided by W.F. Tjallingii, Wageningen 469



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18

University, The Netherlands). The analysis of the first sieve element phase was divided into the 470

length of the first salivation E1 and the time of active ingestion phase E2. 471

472

Petiole Exudate Collection and Artificial Diet Feeding Assays 473

Phloem sap-enriched petiole exudates were collected from Arabidopsis leaves as 474

previously described (Chaturvedi et al., 2008). After concentration, equal volumes of petiole 475

exudates were added to a synthetic diet (Mittler et al., 1965) contained in a feeding chamber 476

(Louis et al., 2010b). Three adult aphids were released on each feeding chamber and the total 477

number of GPA in each feeding chamber was determined 5 days later. 478

479

Actin Cytoskeleton Destabilizer Treatment 480

Two leaves of each 4-week-old plant were infiltrated with 2 μM of Cytochalasin D or 481

Latruculin B solubilized in 0.5% dimethylsulfoxide (DMSO), followed by release of two adult 482

GPA on each leaf. Each leaf was caged in a perforated 2 ml microfuge tube to prevent escape of 483

insects. Total number of nymphs on each leaf were determined two days later. Plants treated 484

with 0.5% DMSO provided the negative control. A minimum of twelve leaves of each genotype 485

were used for each treatment. 486

487

PCR Analysis for Mutant Screening 488

All primers used in this study to confirm the knock-down of individual ADF genes in the 489

adf mutants and the ADF2 RNAi knock-down lines are listed in Supplemental Table S1. PCR 490

with gene-specific primers was performed under the following conditions: 94°C for 5 min, 491

followed by 36 cycles of 94°C for 30 sec, 56°C for 30 sec and 72°C for 1 min, with a final 492

extension of 72°C for 5 min. PCR for genotyping the adf mutants utilized the T-DNA left border 493

primer and a gene-specific primer. The PCR conditions included a 5 min denaturation at 94°C, 494



Mondal et al.

19

followed by 36 cycles of 94°C for 30 sec, 56°C for 30 sec and 72°C for 1 min, with a final 495

extension of 72°C for 5 min. 496

497

Gene expression 498

RNA extraction from leaves was performed as previously described (Pegadaraju et al., 499

2005). Each sample included a minimum of two leaves. A minimum of three samples were 500

analyzed for each treatment. For Real-time RT-PCR, gene expression levels were normalized to 501

that of EF1α, while for RT-PCR ACT8 was used as a control. Gene-specific primers used for 502

Real-time RT-PCR and RT-PCR are listed in Supplemental Table S1. 503

504

Expression and Purification of Recombinant ADF3 Proteins 505

The primers 5’-CGCCCCATATGGCTAATGCAGCATCAGGAATGGCAGTCC-3’ and 506

5’- AAGCTTCTCGAGTCAATTGGCTCGGCTTTTGAAAAC -3’ were used to amplify the 507

ADF3 coding sequence, using cDNA prepared from mRNA harvested from Arabidopsis Col-0 as 508

the template. The resultant product was digested with NdeI and XhoI, which cut within the two 509

primer regions. The digested product was ligated between the NdeI and XhoI sites of pET28a 510

vector. The resultant pET28a-ADF3 plasmid in which a 6X-His tag was incorporated at the N-511

terminal end of ADF3 was transformed into E. coli strain BL21-DE3. Expression of the 512

recombinant ADF3 protein was induced by the addition of Isopropyl-β-D-thiogalactopyranoside 513

(IPTG) prior to bacterial cell harvest. The recombinant, 6X His-tagged protein was purified over 514

an affinity Ni-NTA column (QIAexpress, Qiagen, http://www.qiagen.com). The purified ADF3 515

protein was centrifuged at 20,000g for 30 min at 4°C before use in the F-actin co-sedimentation 516

assays. The GST-ADF3 and GST-ADF4 clones used in the actin depolymerization assays 517

encode recombinant proteins that contain GST fused to the N-terminal end of ADF3 and ADF4. 518

These were a gift of Brad Day. The recombinant proteins were purified over a GST-affinity 519

column before use in actin depolymerization assays. 520

521



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20

Transgenic ADF3 and ADF4 Expressing Plants 522

The ADF3 genomic fragment (gADF3) was amplified using the primers 5’-523

TAAATGAATTTTTTTTACGGGA-3’ and 5’-TCAATTGGCTCGGCTTTTGA-3’, and the 524

resultant product cloned into the pCR8/GW/TOPO® vector (Life technologies; 525

www.lifetechnologies.com). Gateway LR clonase® (Life technologies; 526

www.lifetechnologies.com) was used to mobilize the cloned fragment into the binary vector 527

pMDC107 (Curtis and Grossniklaus, 2003) to yield pMDC107-gADF3. To generate a construct 528

for in planta expression of the ADF3 coding sequence from the Cauliflower mosaic virus 35S 529

gene promoter, the pET28a-ADF3 plasmid (see above) was used as a template with the primer 530

5’-ATGGGCAGCAGCCATCATC-3’, which is derived from the pET28a vector and primer 5’-531

TCAATTGGCTCGGCTTTTGA-3’ to amplify the ADF3 coding region. The resultant PCR 532

product was cloned into pCR8/GW/TOPO® from where it was mobilized into the destination 533

vector pMDC32 (Curtis and Grossniklaus, 2003) to yield the pMDC32-35Spro:ADF3 plasmid. 534

The pMDC107-gADF3 and pMDC32-35Spro:ADF3 plasmids were individually 535

electroporated into Agrobacterium tumefaciens strain GV3101, which was subsequently used for 536

transforming the adf3-1 plants by the floral-dip method (Zhang et al., 2006) to generate 537

transgenic plants in which the adf3-1 mutant phenotype was complemented by expression of the 538

gADF3 and 35Spro:ADF3. Transgenic plants were selected on ½ strength MS agar plates 539

containing hygromycin (25 μg ml-1). The 35Spro:PAD4 construct has been previously described 540

(Xing and Chen, 2006). 541

The 35Spro:ADF4 construct (Henty-Ridilla et al., 2014), in which the ADF4 coding 542

sequence containing a N-terminal T7 epitope tag is expressed from the 35S promoter was used for 543

generating the adf3-1 35Spro:ADF4 plants by transforming adf3-1 plants with the recombinant 544

construct. Transformed plants were screened based on their resistance to kanamycin (50 μg ml-1) 545

and the presence of the recombinant construct and its expression confirmed by PCR and RT-PCR, 546

respectively. To verify the construct in the adf3-1 35S:ADF4 transgenic lines, a forward primer 547

5’-GGTGGTCAACAAATGGGT- 3’ that is specific to the region containing the T7 tag and a 548

reverse primer 5’-TTAGTTGACGCGGCTTTTC- 3’, which is specific to the ADF4 coding 549

sequence were utilized. The same primers were also utilized to monitor expression of the 550



Mondal et al.

21

recombinant constructed by RT-PCR. The PCR conditions were as follows: 94°C for 5 min, 551

followed by 30 cycles of 94°C for 30 sec, 56°C for 30 sec and 72°C for 30 sec, followed by a 552

final extension step of 72°C for 5 min. 553

To generate a SUC2pro:ADF3 construct, the ADF3 coding sequence with a 6X-His tag at 554

the N-terminus was amplified from plants containing the gADF3 construct which contains a His 555

tag on the 5' end. The primers 5’- ATGGGCAGCAGCCATCATC -3’ and 5’-TCAATTG 556

GCTCGGCTTTTGA-3’ were used to amplify the ADF3 coding sequence and the resultant PCR 557

product was cloned into pCR8/GW/TOPO®. The pCR8 product was then mobilized into a 558

pMDC85 destination vector (Curtis and Grossniklaus, 2003) which, had been modified by the 559

removal of the 2x 35S promoter and replaced by the SUC2 promoter (Elzinga et al., 2014). The 560

resultant construct, pMDC85-SUC2pro:ADF3 was transformed into adf3-1 using Agrobacterium 561

tumefaciens via the floral-dip method (Zhang et al., 2006). Transformed plants were screened 562

based on resistance to hygromycin (20 μg ml-1) and presence of the recombinant construct and its 563

expression confirmed by PCR and RT-PCR, respectively. A forward primer 5’-564

AGCCATCATCATCATCATCAC-3’ designed to the 6X-HIS tag at the 5’ end of the ADF3 565

coding sequence and a reverse primer 5’-TCAATTG GCTCGGCTTTTG-3’ that is specific to 566

the ADF3 coding sequence were utilized in PCR reactions to confirm the presence of the 567

SUC2pro:ADF3 construct. These same primers were also utilized to monitor expression of the 568

recombinant construct by RT-PCR. The PCR conditions were as follows: 94°C for 5 min, 569

followed by 30 cycles of 94°C for 30 sec, 57°C for 30 sec and 72°C for 45 sec, followed by a 570

final extension step of 72°C for 5 min. 571

572

GUS Reporter Constructs and Histochemical Analysis 573

A 2035 bp DNA fragment upstream of the transcriptional start site of ADF3 was 574

amplified with Platinum® Taq polymerase (Invitrogen) using Arabidopsis Col-0 genomic DNA 575

as template and the primers 5’-TAAATGAATTTTTTTTACGGGA-3’ and 5’- 576

GGTTGAATCAAAGCTAGTCTCA-3’. The PCR product was cloned into pCR8/GW/TOPO 577

from where it was mobilized into the pMDC132 vector (Curtis and Grossniklaus, 2003), which 578

contains the coding sequence of the bacterial UidA gene, which encodes the GUS protein 579



Mondal et al.

22

(Jefferson et al., 1987). Transgenic plants containing the ADF3pro:UidA construct were 580

generated in the accession Col-0 background. Histochemical analysis of GUS activity was 581

performed using X-gluc (5-bromo-4-chloro-3-indolyl-β-D-glucopyranosiduronic acid) as the 582

substrate. 583

584

Callose Staining 585

Callose staining and quantification of callose deposits was conducted using a modification 610

of a previously described protocol (Ton and Mauch-Mani, 2004). Briefly, leaf samples were 611

cleared overnight in 96% ethanol, followed by a 30 min incubation in sodium phosphate buffer 612

(0.07 M, pH 9). Leaves were then incubated for 60 min in a solution containing 0.005% aniline 613

blue in sodium phosphate buffer (0.07 M, pH 9), followed by the addition of calcofluor to a final 614

concentration of 0.005%. The tissues were immediately observed under a Nikon e600 615

epifluorescence microscope equipped with a X-cite 120 Fluor System, UV (DAPI) filter and a 616

digital SPOT color camera. Digital images were used to quantify callose spots, which were 617

expressed as number of callose spots per mm2 of leaf tissue. 618

619

Actin Depolymerization Assays 620

Rabbit skeletal muscle G-actin was prepared by the method of Spudich and Watt (1971) 621

and flash frozen in liquid nitrogen in 0.5 mL aliquots for storage at -70°C until use. After rapid 622

thawing, the G-actin was chromatographed on a superdex 75 gel filtration column immediately 623

prior to measurements. Concentrations of G-actin were determined using an extinction 624

coefficient of 0.63 mg/ml/cm at 290 nm. A portion of the actin was labeled with pyrene 625

iodoacetamide by the method of Cooper et al. (1983) and stored by the same method as the 626

unlabeled G-actin. 627

Measurements of pyrene-labeled actin were performed with an Aminco-Bowman II 628

luminescence spectrometer using methods similar to those previously described (Xu and Root 629

2000). Briefly, 15% pyrene-labeled actin was polymerized by the addition of 0.1 M KCl and 2 630



Mondal et al.

23

mM MgCl2 yielding a 20-fold enhancement of pyrene fluorescence intensity. Actin 631

concentrations were tested in the ranges of 3 to 7.1 μM for separate experiments. The 632

polymerized actin was titrated at 25°C with a range of concentrations of recombinant GST-633

tagged ADF3 and ADF4 or buffer as a control, and changes in fluorescence intensity measured. 634

Control experiments indicated a reproducibility of ±10% error or better. 635

636

Statistical Analysis 637

When comparing two treatments or genotypes, 2-tail t-test was used to determine if the 638

mean values were significantly different from each other (P<0.05). When simultaneously 639

comparing multiple genotypes and/or treatments to each other, analysis of variance (ANOVA) 640

performed following the General Linear Model GLM was used followed by Tukey’s multiple 641

comparison test to identify mean values that were significantly different from each other 642

(P<0.05) (Minitab v15; www.minitab.com). When comparing multiple experimental groups to a 643

single control group, Dunnett’s multiple comparison test was used to compare means to identify 644

values that were significantly different (P<0.05) from the control group. All data conform to the 645

assumptions of ANOVA and no transformations were necessary. For the EPG analysis, the 646

nonparametric Kruskal-Wallis test (Minitab v15; www.minitab.com) was used to analyze the 647

mean time spent by aphids on various activities. 648

649

Arabidopsis Gene Accession Numbers 650

ADF1 (At3g46010); ADF2 (At3g46000); ADF3 (At5g59880); ADF4 (At5g59890); ADF5 651

(At2g16700); ADF9 (At4g34970); ACT8 (At1g49240); EF1α (At5g60390); PAD4 (At3g52430). 652

653

ACKNOWLEDGEMENTS 654

We thank the Arabidopsis Biological Resource Center for Arabidopsis seeds, Brad Day 655

for adf1, adf3-1, adf4, aadf5 and adf9 mutant lines, the GST-ADF3, GST-ADF4 and 35S:ADF4 656



Mondal et al.

24

clones, Janice de Almeida Engler for the ADF2-RNAi line, Zhixiang Chen for the 35Spro:PAD4 657

construct, and Jane Parker and Bart Feys for discussion and making available unpublished 658

results. 659

660

661

SUPPLEMENTAL DATA 662 The following materials are available in the online version of this article. 663

Supplemental Table S1. Primers used for monitoring gene expression. 664

Supplemental Figure S1. Arabidopsis lines carrying T-DNA insertions at the ADF3 locus. 665

Supplemental Figure S2. ADF3 expression from the 35S promoter restores resistance against 666

the GPA in the adf3-1 mutant. 667

Supplemental Figure S3. ADF3 exhibits actin depolymerizing activity. 668

Supplemental Figure S4. Alignment of ADF3 and ADF4 669

Supplemental Figure S5. Histochemical staining for GUS activity in uninfested (-GPA) and 670

GPA-infested (+GPA) leaves of a ADF3pro:UidA plant. 671

Supplemental Figure S6. Expression of PAD4 from the constitutively expressed 35S promoter 672

restores resistance against GPA in plants lacking ADF3 function. 673

674

675



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25

FIGURE LEGENDS 676

677

Figure 1. Arabidopsis ADF3 gene is required for limiting GPA infestation. A, GPA population 678

size on Arabidopsis wild-type (WT) accession Col-0 and adf1 (a1), adf2 (a2), adf3-1 (a3-1), 679

adf4 (a4), adf5 (a5) and adf9 (a9) mutants (n=10) two days post infestation. B, GPA population 680

size on WT accession Col-0 and adf3-1 (a3-1) and adf3-2 (a3-2) mutant plants (n=10) two days 681

post infestation. C, One nymph was released on each WT and a3-1 plant and the number of 682

progeny nymphs produced by each insect over a 10 day period determined (n=6). D, GPA 683

population size on WT accession Col-0, a3-1 mutant, and two independently-derived gADF3 684

plants in which a genomic clone of ADF3 was transformed into the a3-1 mutant (n= 6). Top 685

panel shows RT-PCR analysis of ADF3 and as a control ACT8 expression in uninfested plants of 686

indicated genotypes. E, GPA population size on WT, adf3-1 and a transgenic 35Spro:ADF3 line 687

#1 in which the ADF3 coding sequence was expressed from the heterologous 35S promoter 688

(n=6). See Supplemental Fig. S2 for results from an independently derived 35Spro:ADF3 line. 689

In A, B, D and E, twenty adult apterous aphids were released on each plant and the population 690

size (adults + nymphs) on each plant determined two days later. All values are mean + SE for 691

each genotype. In A and B, asterisks above bars denote values that are significantly different 692

from the WT (P < 0.05; Dunnett test). In C, an asterisk denotes significant difference from the 693

WT (P < 0.05; t-test). In D and E, different letters above bars denote values that are significantly 694

different from each other (P < 0.05; Tukey test). 695

696

697

Figure 2. Actin cytoskeleton destabilizers restore resistance to GPA in the adf3 mutant. 698

A, Effect of the actin destabilizers cytochalasin D (CytD, left panel) and latrunculin B (LatB, 699

right panel) on insect numbers. Number of nymphs on leaves of WT and adf3-1 (a3-1) mutant 700

treated with 2 μM CytD or LatB, and as control with DMSO (0.5%), which was used as a solvent 701

for CytD and LatB. The number of nymphs per adult was determined two days after release of 702

two adult aphids on each leaf (n=12). B, Complementation of adf3-1-conferred susceptibility to 703

GPA by constitutive expression of ADF4 from the 35S promoter. Upper panel: RT-PCR analysis 704

of ADF4 expression from the 35S promoter in three independent a3-1 35SPpro:ADF4 lines, and 705

as control in WT and a3-1 plants. EF1α expression provided the control for RT-PCR. Lower 706



Mondal et al.

26

panel: No choice assay comparing aphid numbers two days post release of 20 adult aphids per 707

leaf on the WT, a3-1 and three independently-derived a3-1 35Spro:ADF4 transgenic lines. 708

Values are mean + SE of a minimum of six replicates for each genotype. Different letters above 709

bars denote values that are significantly different from each other (P < 0.05; Tukey test). 710

711

Figure 3. ADF3 function in the phloem is required for controlling GPA infestation. A, Real-712

time RT-PCR analysis of ADF3 expression relative to that of EF1α in uninfested (-GPA) and 713

GPA-infested (+GPA) leaves of the Arabidopsis accession Col-0 at the indicated hours post 714

infestation (hpi). Values are mean + SE (n = 3). B, Histochemical staining for GUS activity 715

(blue color) in a GPA-infested leaf of a ADF3pro:UidA plant in which UidA expression is driven 716

from the ADF3 promoter. Right panel shows a close-up showing strong GUS activity in the 717

veins. C, Histochemical staining in a section through a vein showing strong GUS activity in the 718

phloem (P). X, xylem. D, ADF3 expression in the phloem is sufficient for controlling GPA 719

infestation. Upper panel: RT-PCR analysis of ADF3 expression from the SUC2 promoter in 720

three independent a3-1 SUC2Pro:ADF3 lines, and as control in wild-type (WT) accession Col-0, 721

a3-1 and a3-1 35Spro:ADF3 plants. Primers specific for the SUC2 promoter-driven ADF3 722

construct were used. EF1α expression provided the control for RT-PCR. Lower panel: GPA 723

population size on Arabidopsis plants of indicated genotypes. Twenty adult apterous aphids 724

were released on each plant and the population size (adults + nymphs) on each plant determined 725

two days later. Values are mean + SE of a minimum of six replicates for each genotype. 726

Different letters above bars denote values that are significantly different from each other (P < 727

0.05; Tukey test). 728

729

730

Figure 4. ADF3 is not required for accumulation of antibiosis activity in petiole exudates. 731

Artificial diet assay showing a comparison of GPA numbers on diet containing petiole exudates 732

(Pet-ex) collected from leaves of wild-type (WT), adf3-1 (a3-1) and adf3-2 (a3-2) plants. 733

Control was an artificial diet supplemented with the buffer used to collect petiole exudates. 734

Values are mean + SE of a minimum of seven replicates for each treatment. Different letters 735

above bars denote values that are significantly different from each other (P < 0.05; Tukey test). 736



Mondal et al.

27

Figure 5. ADF3 is required for controlling GPA feeding on Arabidopsis leaves. A, Electrical 737

penetration graph (EPG) comparison of time spent by GPA in various activities on the WT and 738

adf3-1 (a3-1) mutant plants in an 8 h period. FP: time taken for first probe; f-SEP: time taken to 739

reach first sieve element phase; PP: time spent in pathway phase during the 8 h period; NP: total 740

time spent non-probing during the 8 h period; s-SEP: total time spent in sieve element phase 741

during the 8 h period; XP: total time spent in the xylem phase during the 8 h period. B, EPG 742

comparison of time spent by GPA in the E1 (salivation) and E2 (ingestion) phases during the 743

first SEP. In A and B, values are the mean + SE (n=10). Asterisks indicate significant 744

differences (P < 0.05; Kruskal-Wallis test) in an individual feeding parameter between insect 745

feeding on the a3-1 compared to the WT plant. 746

747

Figure 6. ADF3 is required for promoting callose deposition in response to GPA infestation. 748

Upper panel: Callose deposits (light blue spots) in leaves of wild-type (WT) plants of the 749

Arabidopsis accession Col-0 and the adf3-1 (a3-1) mutant, 24 hpi with the GPA (+GPA) or as 750

control in uninfested (-GPA) plants. Bar = 200 μM. Lower panel: Mean number of callose spots 751

+ SE per mm2 of leaf tissue in GPA-infested and uninfested leaves (n=5). Different letters above 752

bars denote values that are significantly different from each other (P < 0.05; Tukey test). 753

754

Figure 7. ADF3 is required for promoting PAD4 expression in response to GPA infestation. A, 755

qRT-PCR analysis of PAD4 expression relative to EF1a expression at 24 hpi in GPA-infested 756

and uninfested WT and adf3-1 (a3-1) mutant plants. Values are mean + SE (n=3). Asterisks 757

indicate values that are significantly different from the corresponding mock-treatment (P < 0.05; 758

t-test). B, GPA population size on WT, a3-1, a3-1 35Spro:ADF3 line #1, a3-1 35Spro:PAD4 line 759

#1, pad4-1, and pad4-1 3Spro:ADF3 line #1. See Supplemental Fig. S6 for data for additional 760

independently-derived transgenic lines. The no-choice assay was conducted by releasing twenty 761

adult apterous aphids on each plant and the population size (adults + nymphs) on each plant 762

determined two days later. Values are mean + SE (n=6) for each genotype. Different letters 763

above bars denote values that are significantly different from each other (P < 0.05; Tukey test). 764

765



Mondal et al.

28

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969



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Parsed CitationsAndrianantoandro E, Pollard TD (2006) Mechanism of actin filament turnover by severing and nucleation at different concentrations ofADF/cofilin. Mol Cell 24: 13-23

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

Blackman RL, Eastop VF (2000) Aphids on the World’s Crops: An Identification and Information Guide, Ed 2. John Wiley, Chichester, UKPubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Bos JIB, Prince, D, Pitino M, Maffei ME, Win J, Hogenhout SA (2010) A functional genomics approach identifies candidate effectorsfrom the aphid species Myzus persicae (Green Peach Aphid). PLoS Genet 6: e1001216


Burgos-Rivera B, Ruzicka DR, Deal RB, McKinney EC, King-Feid L, Meagher RB (2008) ACTIN DEPOLYMERIZING FACTOR9 controlsdevelopment and gene expression in Arabidopsis. Plant Mol Biol 68: 619-632


Cai G, Faleri C, Del Casino C, Emons AMC, Cresti M (2010) Distribution of callose synthase, cellulose synthase, and sucrose synthasein tobacco pollen tube is controlled in dissimilar ways by actin filaments and microtubules. Plant Physiol 155: 1169-1190


Chaturvedi R, Krothapalli K, Makandar R, Nandi A, Sparks AA, Roth MR, Welti R, Shah J (2008) Plastid �-3-fatty acid desaturasedependent accumulation of a systemic acquired resistance inducing activity in petiole exudates of Arabidopsis thaliana is independentof jasmonic acid. Plant J 54:106-117


Clément M, Ketelaar T, Rodiuc N, Banora MY, Smertenko A, Engler G, Abad P, Hussey PJ, de Almeida Engler J (2009) Actindepolymerization factor2-meidated actin dynamics are essential for root-knot nematode infection of Arabidopsis. Plant Cell 21: 2963-2979


Cooper JA, Walker SB, Pollard TD (1983) Pyrene actin: documentation of the validity of a sensitive assay for actin polymerization. JMuscle Res Cell Motil 4: 253-262


Curtis MD, Grossniklaus U (2003) A gateway cloning vector set for high-throughput functional analysis of genes in planta. Plant Physiol133: 462-469


Day B, Henty JL, Porter KJ, Staiger CJ (2011) The pathogen-actin connection: A platform for defense signaling in plants. Annu RevPhytopathol 49: 483-506.


De Vos M, Jander G (2009) Myzus persicae (green peach aphid) salivary components induce defence responses in Arabidopsisthaliana. Plant Cell Environ. 32: 1548-1560


Elzinga DA, De Vos M, Jander G (2014) Suppression of plant defenses by a Myzus persicae (green peach aphid) salivary effectorprotein. Molecular Plant-Microbe Interactions 27: 747-756



http://www.ncbi.nlm.nih.gov/pubmed?term=Andrianantoandro E%2C Pollard TD %282006%29 Mechanism of actin filament turnover by severing and nucleation at different concentrations of ADF%2Fcofilin%2E Mol Cell 24%3A 13%2D23&dopt=abstract

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http://www.ncbi.nlm.nih.gov/pubmed?term=Walling LL %282008%29 Avoiding effective defenses%3A strategies employed by phloem%2Dfeeding insects%2E Plant Physiol 146%3A 859%2D866&dopt=abstract

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Yun BW, Atkinson HA, Gaborit C, Greenland A, Read ND, Pallas JA, Loake GJ (2003) Loss of actin cytoskeletal function and EDS1activity, in combination, severely compromises non-host resistance in Arabidopsis against wheat powdery mildew. Plant J 34: 768-777


Zhang X, Henriques R, Lin SS, Niu QW, Chua NH (2006) Agrobacterium-mediated transformation of Arabidopsis thaliana using the floraldip method. Nature Protocols 1: 641-646



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