1 : cytokinin, shoot branching and pin accumulation 2 · 79 action of auxin on axillary bud growth...

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Short title: Cytokinin, shoot branching and PIN accumulation 1

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Corresponding author: 3

Ottoline Leyser 4

Sainsbury Laboratory Cambridge University 5

Bateman Street 6

Cambridge 7

CB2 1LR 8

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Email: [email protected] 10

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Plant Physiology Preview. Published on May 1, 2018, as DOI:10.1104/pp.17.01691

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Cytokinin targets auxin transport to promote shoot branching 12

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Tanya Waldie and Ottoline Leyser 14

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Sainsbury Laboratory Cambridge University 16

Bateman Street 17

Cambridge 18

CB2 1LR 19

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One sentence summary: Cytokinin promotes shoot branching in part by promoting 21

plasma membrane accumulation of PIN3, PIN4 and PIN7 auxin exporters in the 22

shoot. 23

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Author contributions: 25

T.W. designed, performed and analysed the results of all the experiments, co-26

conceived the research plans, interpreted the results and wrote the manuscript. 27

O.L. co-conceived the research plans, interpreted the results and wrote the 28

manuscript. 29

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ORCIDs: T.W. 0000-0001-7760-0950, O.L. 0000-0003-2161-3829 31

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Funding information: This work was funded by the European Research Council (N° 33

294514 – EnCoDe) and the Gatsby Charitable Foundation (GAT3272C) 34

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Corresponding author email: [email protected] 36


mailto:[email protected]


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ABSTRACT 37

Cytokinin promotes shoot branching by activating axillary buds, but its mechanism of 38

action in this process is unclear. We have previously shown that a hextuple mutant 39

lacking a clade of type-A Arabidopsis Response Regulators (ARRs) known to act in 40

cytokinin signalling has reduced shoot branching compared to wild type. Since these 41

proteins typically act as negative regulators of cytokinin signalling, this is an 42

unexpected result. To explore this paradox more deeply, we characterised the 43

effects of loss of function of the type-B ARR, ARR1, which positively regulates 44

cytokinin-induced gene expression. The arr1 mutant has increased branching, 45

consistent with a role antagonistic to the type-A ARRs, but in apparent conflict with 46

the known positive role for cytokinin in bud activation. We show that the arr 47

branching phenotypes correlate with increases in stem auxin transport and steady-48

state levels of the auxin export proteins PIN3 and PIN7 on the plasma membrane of 49

xylem-associated cells in the main stem. Cytokinin treatment results in increased 50

accumulation of PIN3, PIN7 and the closely related PIN4 within several hours, and 51

loss of PIN3, PIN4 and PIN7 can partially rescue the arr1 branching phenotype. This 52

suggests there are multiple signalling pathways for cytokinin in bud outgrowth; one of 53

these pathways regulates PIN proteins in shoots, independently of the canonical 54

signalling function of the ARR genes tested here. A hypothesis consistent with the 55

arr shoot phenotypes is that feedback control of biosynthesis leads to altered 56

cytokinin accumulation, driving cytokinin signalling via this pathway. 57

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INTRODUCTION 63

Plant developmental plasticity is exemplified by the diversity in shoot forms seen 64

within a species, which are tuned according to environmental conditions. One 65

process underlying this diversity is differential activation of axillary buds throughout 66

the plant’s life cycle, which results in diverse shoot branching habits. The hormonal 67

signalling network controlling bud activity involves auxin, strigolactone (SL) and 68

cytokinin (CK), all of which have well-defined physiological roles, although the 69

molecular mechanisms through which they control bud outgrowth are not yet entirely 70

clear (reviewed in Domagalska and Leyser 2011; Teichmann and Muhr 2015). It is 71

well-established that apical dominance, the inhibitory effect imposed by an actively 72

growing shoot apex on axillary buds, is mediated at least in part by the synthesis and 73

movement of auxin from young expanding leaves into the basipetal polar auxin 74

transport stream (PATS) in the main stem (Thimann and Skoog 1933; Ljung et al., 75

2001). Auxin in the PATS does not enter axillary buds to exert this repression, and 76

thus acts indirectly (Hall and Hillman 1975; Morris 1977; Booker et al., 2003). There 77

is a substantial body of evidence supporting two parallel mechanisms for the indirect 78

action of auxin on axillary bud growth (reviewed in Domagalska and Leyser 2011). 79

One is that auxin in the main stem regulates the synthesis of second messengers 80

that move up into the buds and regulate their activity. The other is that stem auxin 81

influences the establishment of canalised auxin flow out of buds into the PATS. 82

According to this canalisation-based mechanism, auxin movement begins as a weak 83

flux from the bud – an auxin source – into the main stem PATS – an auxin sink. This 84

flux narrows and strengthens due to positive feedback between auxin flux and the 85

auxin transport machinery (Sachs 1981; Prusinkiewicz et al., 2009). This process 86

results in the formation of specialised cell files that conduct auxin from source to 87



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sink, and this is hypothesised to be required for sustained bud activation. The action 88

of SL and CK in bud activation control can be considered in terms of these two 89

models. 90

91

In dicots, SL is thought to act via both mechanisms. Auxin up-regulates the 92

transcription of SL biosynthetic genes in the stem and SL can move upward into 93

buds, presumably in the transpiration stream (Foo et al., 2001; Bainbridge et al., 94

2005; Foo et al., 2005; Johnson et al., 2006; Hayward et al., 2009). There, SL 95

modulates expression of the TEOSINTE BRANCHED1/CYCLOIDEA/PCNA (TCP) 96

family transcription factor BRANCHED1 (BRC1), an inhibitor of shoot branching 97

(Aguilar-Martínez et al., 2007; Poza-Carrion et al., 2007; Braun et al., 2012; Dun et 98

al., 2012). However, high BRC1 transcript levels are neither necessary nor sufficient 99

for bud inhibition and mutant buds lacking BRC1 can be inhibited by SL (Seale et al., 100

2017). Furthermore, SL addition can promote branching in an auxin transport 101

compromised genetic background, demonstrating that this simple second messenger 102

mechanism cannot be the only mode of action for SL (Shinohara et al., 2013). 103

Consistent with this idea, SL triggers rapid removal of the auxin export protein, PIN1, 104

from the plasma membrane (Crawford et al., 2010; Shinohara et al., 2013; Bennett et 105

al., 2016). This effect is sensitive to inhibitors of clathrin-mediated endocytosis, but 106

not to the translation inhibitor cycloheximide, suggesting a non-transcriptional mode 107

of action of SL on PIN1 endocytosis. In the context of an auxin transport 108

canalisation-based model for bud activation, PIN1 removal can account for the 109

inhibitory effect of SL on shoot branching since it is predicted to make canalisation 110

more difficult to achieve by dampening the feedback between auxin flux and auxin 111

transporter accumulation. Furthermore, when auxin transport is compromised and 112



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auxin fluxes are systemically low, the effect of SL on PIN1 endocytosis is predicted 113

to promote branching, as observed (Shinohara et al., 2013). 114

115

Half a century before the discovery of SL, one of the earliest described roles for CK 116

in plant development was in bud activation (Wickson and Thimann 1958). In 117

Arabidopsis (Arabidopsis thaliana), basally applied CK can overcome the inhibitory 118

effects of apical auxin on bud activity (Chatfield et al., 2000). Furthermore, 119

isopentenyl transferase (ipt) 3,5,7 mutants impaired in CK biosynthesis have 120

reduced CK levels and form fewer branches than wild-type plants (Miyawaki et al., 121

2006; Müller et al., 2015). As for SL, there is good evidence that CK can act as a 122

second messenger for stem auxin. Removal of the shoot apex correlates with 123

increased stem CK levels, and addition of auxin reduces them (Bangerth 1994; 124

Tanaka et al., 2006). In Arabidopsis, CK is perceived at the endoplasmic reticulum 125

by the HISTIDINE KINASE (AHK) receptor kinase family, which initiate a 126

phosphorelay cascade that targets the large, multi-member family of RESPONSE 127

REGULATORs (ARRs) in the nucleus via HISTIDINE PHOSPHOTRANSFER (AHP) 128

proteins (for review see Schaller et al., 2015). The ARRs possess an N-terminal 129

phosphoreceiver domain and comprise two sub-classes based on the presence (in 130

type-Bs) or absence (in type-As) of a DNA-binding domain. Type-B ARRs directly 131

regulate transcription and function as positive regulators of CK signalling, whereas 132

type-A ARRs typically function as negative regulators. It has therefore been widely 133

assumed that CK activates buds by regulating the transcription of relevant genes in 134

the bud, such as BRC1. There is good evidence to support this model since in both 135

pea (Pisum sativum) and Arabidopsis, BRC1 expression is down-regulated by CK 136

(Braun et al., 2012; Dun et al., 2012; Seale et al., 2017). However, CK can promote 137



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bud activation in pea brc1 mutants, demonstrating that CK has BRC1-independent 138

effects on the regulation of bud outgrowth (Braun et al., 2012). Furthermore, the 139

Arabidopsis hextuple type-A arr3,4,5,6,7,15 mutant exhibits reduced bud activation, 140

a phenotype opposite to that predicted, based on the established roles of type-A 141

ARR proteins as negative regulators of transcriptional CK signalling (To et al., 142

2004a; Müller et al., 2015). 143

144

To explore this paradoxical result further, we describe here the analysis of the type-B 145

ARR mutant, arr1. ARR1 binds to the promoters of CK up-regulated genes, including 146

those induced during CK-triggered bud activation. We therefore hypothesised the 147

arr1 mutant should show reduced and CK-resistant shoot branching. However, our 148

results demonstrate arr1 has increased and CK-responsive shoot branching, 149

suggesting an alternative mechanism for CK-mediated shoot branching control. We 150

provide evidence that the mechanism involves CK-mediated accumulation of the 151

PIN3, PIN4 and PIN7 auxin transporters. 152

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RESULTS 155

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Type-A and type-B arr mutations confer opposite shoot branching and auxin 157

transport phenotypes 158

Previously, we found that the hextuple type-A arr3,4,5,6,7,15 mutant has reduced 159

shoot branching relative to wild type (Müller et al., 2015). Since type-B ARR family 160

members are known to act antagonistically to type-A ARRs in other CK responses, 161

we investigated shoot branching in the arr1 loss-of-function mutant. ARR1 was 162

selected because a group of CK-up-regulated genes in buds possess an ARR1 163

response element in their promoters (Müller et al., 2015). In accordance with action 164

antagonistic to type-A ARRs, the type-B arr1 single mutant has increased branching 165

compared to wild-type controls, forming a mean of 6.8 rosette branches compared to 166

5.5 in wild type when decapitated (Fig. 1A; p < 0.001), and a mean of 6.8 branches 167

compared to 5.7 in wild type when intact (Fig. 1B-E; p < 0.001). 168

169

Buds on isolated nodal stem segments from arr3,4,5,6,7,15 plants treated with apical 170

auxin were previously shown to be resistant to the effects of basal CK and slightly 171

resistant to basal SL (Müller et al., 2015). The same isolated nodal assay system 172

was used to assess bud hormone responses in arr1 (Chatfield et al., 2000; Crawford 173

et al., 2010). Briefly, small nodal stem segments, each bearing an inactive cauline 174

bud, were excised from bolting inflorescences. The apical end of the stem was 175

embedded in an agar block supplemented with synthetic auxin (1-napthalene acetic 176

acid; NAA) or a control solution. The basal end of the stem was embedded in an 177

agar block supplemented with CK (6-benzylaminopurine; BA), SL (rac-GR24) or a 178

control solution (hereafter referred to as apical NAA, basal BA and basal GR24 179



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treatments). The kinetics of outgrowth in mock-treated arr1 buds was similar to wild 180

type (Fig. 2A & B). For both genotypes, apical NAA delayed bud activation by 181

approximately three days. The response of arr1 to BA was also similar to wild type, 182

where basal BA alone had little effect compared to mock treatment, but basal BA 183

could overcome the inhibitory effect of apical NAA and activate buds. As previously 184

reported, basal GR24 prolonged the inhibitory effect of apical NAA in wild type, but 185

arr1 buds were resistant to basal GR24 under these conditions (Fig. 2C & D). To 186

assess whether this altered GR24 response extended beyond isolated nodes, whole 187

plants were grown under axenic conditions on ATS media supplemented with GR24, 188

as per Crawford et al. (2010). Branching in wild-type plants was significantly inhibited 189

by 1 µM GR24, whereas 5 µM was required for significant branch inhibition in arr1 190

plants (Fig. S1). 191

192

The altered GR24 responses of arr1 single and arr3,4,5,6,7,15 hextuple mutants 193

raise the possibility that arr mutations may affect auxin transport processes in the 194

shoot. We therefore compared bulk auxin transport in wild type, type-A and type-B 195

arr mutant stem segments, as previously described (Bennett et al., 2016). Compared 196

to wild type, arr1 transported 40% more auxin (p < 0.01) and arr3,4,5,6,7,15 197

transported 23% less (p < 0.01) over the 6-hour assay period (Fig. 3A). To determine 198

whether this effect was associated with CK signalling, we also assessed the ipt3 CK 199

biosynthesis mutant, which we have previously shown to have a reduced shoot 200

branching phenotype (Müller et al., 2015). Similar to arr3,4,5,6,7,15, the ipt3 mutant 201

also exhibited reduced auxin transport (Fig. S2). Thus, the degree of shoot 202

branching positively correlates with bulk auxin transport in the stems of these mutant 203

genotypes. 204



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To test this correlation further, we assessed the shoot branching response of arr1 206

and arr3,4,5,6,7,15 plants to the auxin transport inhibitor 1-naphthylphthalamic acid 207

(NPA). Plants were grown under axenic conditions on ATS media supplemented with 208

NPA, as per Bennett et al. (2006). Significant reductions in branching were observed 209

in arr1 plants treated with 0.1 μM NPA (p < 0.05) and 1.0 μM NPA (p < 0.001) (Fig. 210

3B), consistent with a causative link between auxin transport defects and shoot 211

branching in the arr1 mutant, similar to the results obtained for SL deficient mutants 212

(Bennett et al., 2006). In contrast, treatment with 0.1 μM NPA significantly increased 213

branching in arr3,4,5,6,7,15 (p < 0.01) (Fig. 3C). This is consistent with the well-214

established promotive effect of very low auxin transport on branching (Ruegger et 215

al., 1997; Geldner et al., 2003). Taken together, the results suggest mutations in 216

type-A and type-B ARR family members perturb auxin transport in the shoot, and this 217

contributes to their effects on shoot branching. 218

219

CK signalling targets PIN proteins in the stem 220

Several PIN proteins contribute to stem auxin transport in Arabidopsis. PIN1 is an 221

important component of the classical PATS and is expressed in a highly polar 222

manner in xylem parenchyma and cambium cells in the stem vasculature. PIN3, 223

PIN4 and PIN7 contribute to PATS and to a less-polar route, termed Connective 224

Auxin Transport (CAT), which connects surrounding tissues and organs, including 225

axillary buds, to the PATS (Bennett et al., 2016). To determine whether arr mutant 226

shoots have alterations in PIN accumulation we analysed the steady-state transcript 227

levels of PIN1, PIN3, PIN4 and PIN7 in arr1 and arr3,4,5,6,7,15. PIN transcript levels 228

were analysed in upper inflorescence internodes of wild-type, arr1 and 229



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arr3,4,5,6,7,15 plants using RT-qPCR (Fig. S3). Apart from a 5-fold increase in PIN7 230

transcripts in arr3,4,5,6,7,15 compared to wild type (p < 0.01), no significant changes 231

were observed, suggesting the arr mutant auxin transport phenotypes do not 232

correlate with alterations in PIN transcript abundance in inflorescence stems. 233

234

In the absence of correlative changes in PIN transcription, we assessed PIN protein 235

localisation and abundance using established PIN-GFP reporter lines crossed into 236

the type-A and B arr mutant backgrounds. PIN-GFP accumulation patterns were 237

analysed in longitudinal or transverse sections of basal inflorescence internodes (the 238

inflorescence internode located directly above the rosette and farthest from the shoot 239

apex) and imaged using confocal microscopy. In arr3,4,5,6,7,15 mutants, the amount 240

of PIN1-GFP on the basal plasma membrane (the rootward-facing membrane 241

farthest from the shoot apex) of xylem parenchyma cells was unchanged compared 242

to wild type (Fig. 4A-C). In contrast, the amount of PIN3-GFP on the basal plasma 243

membrane of xylem parenchyma cells was reduced by approximately 25% in 244

arr3,4,5,6,7,15 compared to wild type (p < 0.001) (Fig. 4D-F). As PIN7-GFP typically 245

shows a broad, cross-stem pattern of accumulation in young wild-type internodes, 246

and PIN7-GFP was not detectable on the basal plasma membrane of xylem 247

parenchyma cells in arr3,4,5,6,7,15, the PIN7-GFP signal was quantified within 248

vascular bundles in inflorescence stem transverse sections (Fig. 4G-I). Like PIN3-249

GFP, PIN7-GFP was also significantly decreased in arr3,4,5,6,7,15 compared to 250

wild-type (p < 0.001). Analysis of the ipt3 CK synthesis mutant revealed similar 251

patterns of PIN1-GFP and PIN7-GFP accumulation as the arr3,4,5,6,7,15 mutant, 252

with wild-type levels of PIN1-GFP present on the basal plasma membrane of xylem 253



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parenchyma cells and decreased levels of PIN7-GFP within vascular bundles (p < 254

0.001) (Fig. S4). 255

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Wild-type levels of basal plasma membrane-localised PIN1-GFP were also observed 257

in the arr1 mutant (Fig. 5A-C). However, in contrast to arr3,4,5,6,7,15, PIN3-GFP 258

was increased approximately 20% in arr1 inflorescence stems compared to wild type 259

(p < 0.05) (Fig. 5D-F). In transverse sections of young basal arr1 inflorescence 260

internodes (8-10 cm high), PIN7-GFP was restricted to the outer xylem parenchyma 261

and cambium of vascular bundles, whereas wild-type plants exhibited a brighter and 262

broader accumulation pattern that decreased at later stages, as previously reported 263

(Bennett et al., 2016) (Fig. S5). In contrast, the accumulation of PIN7-GFP 264

associated with vascular bundles appeared to be slightly higher in arr1 than wild type 265

at later stages of inflorescence development (18-20 cm and 28-30 cm high) (Fig. 266

S5C-F). In accordance with this observation, the amount of PIN7-GFP present on 267

the basal plasma membranes of xylem parenchyma cells in longitudinal sections of 268

inflorescence stems was increased approximately 25% (p < 0.01) in arr1 compared 269

to wild-type (Fig. 5G-I). 270

271

These data suggest that the auxin transport phenotypes of the arr3,4,5,6,7,15 and 272

arr1 mutants are due at least in part to differential accumulation of PINs belonging to 273

the PIN3, 4, 7 clade in the stem. To assess whether this is a direct effect of CK, we 274

assessed the response of these PIN proteins to BA treatment. Inflorescence stem 275

segments from wild-type plants were held between two agar blocks supplemented 276

with NAA in the upper block and BA or a control solution in the lower block. After 4 h 277

treatment, a longitudinal section was made in the basal half of the stem segment, at 278



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or near the site of BA treatment, and imaged using confocal microscopy. The amount 279

of PIN1-GFP on the basal plasma membrane was unchanged in mock versus BA-280

treated stems (Fig. 6A-C). BA treatment increased the amount of PIN3-GFP on the 281

basal plasma membrane by ~30% (p < 0.01) (Fig. 6D-F), PIN4-GFP by ~20% (p < 282

0.001) (Fig. 6G-I) and PIN7-GFP by ~15% (p < 0.01) (Fig. 6J-L). No changes in the 283

steady-state transcript levels of PIN1, PIN3, PIN4 or PIN7 were found in the basal 5 284

mm of equivalently treated wild-type inflorescence internodes after 4 h BA treatment 285

(Fig. S6). When tracking individual PIN7:PIN7-GFP-expressing xylem parenchyma 286

cells in a longitudinal section for 2 h, the amount of PIN7-GFP on the basal plasma 287

membrane of NAA-treated internodes generally decreased over time, but cells 288

treated with a combination of NAA and BA retained more PIN7-GFP signal 289

compared to NAA alone (p < 0.01) (Fig. S7). 290

291

Together, the results demonstrate CK affects plasma membrane levels of PIN3, 292

PIN4 and PIN7 proteins in inflorescence stems, and this correlates with changes in 293

auxin transport. The effect of CK on these PINs may be post-transcriptional and the 294

response has some specificity, since PIN1 is unaffected. 295

296

Nitrate phenocopies the effect of CK on PIN proteins in stems 297

Nitrate supply promotes shoot branching in Arabidopsis but the mechanisms 298

underlying this response are complex, with multiple modes of action likely (de Jong 299

et al., 2014). The response of branching to nitrate is unlikely to involve PIN1, since 300

steady-state PIN1-GFP levels in the stem remain the same under different nitrate 301

regimes (de Jong et al., 2014). It is well-established that CK can act as a nitrate 302

signal (Sakakibara et al., 2006) and consistent with this, we previously showed that 303



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higher order ipt3,5,7 and arr3,4,5,6,7,15 mutants form similar numbers of branches 304

under high and low nitrate conditions, suggesting CK contributes to the ability of 305

Arabidopsis plants to modulate branching in response to nitrate supply (Müller et al., 306

2015). The CK-responsiveness of PIN3, PIN4 and PIN7 suggests a possible route 307

for nitrate-mediated regulation of branching via CK. To assess whether changes in 308

nitrate status might be reflected in PIN3, PIN4 and PIN7 accumulation, PIN3:PIN3-309

GFP, PIN4:PIN4-GFP and PIN7:PIN7-GFP plants were grown under nitrate sufficient 310

and insufficient conditions and the amount of PIN-GFP signal on the basal plasma 311

membrane of xylem parenchyma cells quantified in basal inflorescence internodes. 312

As for exogenous CK supply, nitrate sufficient conditions were associated with 313

increased levels of PIN3-GFP, PIN4-GFP and PIN7-GFP on the basal plasma 314

membrane (Fig. 7). Under low N conditions, PIN3-GFP was reduced to ~75% of the 315

levels observed in high N plants (p < 0.001) (Fig. 7A), while PIN4-GFP and PIN7-316

GFP were reduced to approximately 85% of that in high N plants (p < 0.001) (Fig. 7B 317

and C). These results are consistent with the idea that nitrate modulates shoot 318

branching at least in part via CK effects on PIN3, PIN4 and PIN7 abundance; 319

although, it is also possible that nitrate supply modules PIN3,4,7 accumulation via a 320

CK-independent mechanism. 321

322

PIN3, PIN4 and PIN7 contribute to shoot branching control in arr1 323

To assess the contribution of changes in PIN3, PIN4 and PIN7 to CK-mediated 324

branching control, we generated the arr1 pin3 pin4 pin7 quadruple mutant (hereafter 325

referred to as arr1 pin3,4,7) and analysed its branching phenotype. As the 326

differences in branch numbers between wild type, arr1 and pin3,4,7 are typically 327

small, plants were grown under short days for 4 weeks initially then shifted to long 328



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days for 5 weeks in order to maximise branch numbers, providing sensitised 329

conditions to assess any differences. At terminal flowering, an intermediate number 330

of branches were formed in the arr1 pin3,4,7 quadruple mutant (9.9) compared to the 331

arr1 single mutant (12.1) and the pin3,4,7 triple mutant (8.4) (Fig. 8A-F). This 332

reduction in branching of arr1 in the pin3,4,7 mutant background is consistent with 333

the hypothesis that CK-regulated changes in PIN3, PIN4 and PIN7 contribute to the 334

increased shoot branching phenotype of arr1. The pin3,4,7 mutant exhibits twisted 335

rosette leaves and an increased cauline branch angle phenotype (Bennett et al., 336

2016) and in both these aspects the arr1 pin3,4,7 quadruple mutant appeared similar 337

to the pin3,4,7 triple mutant (Fig. 8A-E and G). 338

339

To assess the responses of pin3,4,7 buds to exogenous CK treatment, we used the 340

same experimental set-up described for arr1, in which buds on isolated nodal stem 341

segments were treated with apical NAA and basal BA supplied through the stem. 342

Overall, the pin3,4,7 triple mutant responded similarly to wild type, activating in mock 343

and basal BA treatments and remaining inhibited for several days in the presence of 344

apical NAA (Fig. 9A and B). As for wild type, basal BA could override the effect of 345

apical NAA in pin3,4,7 mutants, however, mutant buds activated significantly faster 346

than wild type during the first four days (Fig. 9C). Together, the results suggest PIN3, 347

PIN4 and PIN7 contribute to CK-mediated bud activation, but the phenotypic effects 348

of pin3,4,7 mutation suggest that CK also functions via other mechanisms. 349

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DISCUSSION 351

Perturbing CK levels through exogenous application or through manipulation of 352

endogenous levels provides clear evidence that CK can promote bud activation 353

(Wickson and Thimann 1958; Faiss et al., 1997; Chatfield et al., 2000; Müller et al., 354

2015). Elucidating the role of the known CK signalling pathway in bud activation has 355

been less straightforward due to the large gene families involved and consequent 356

functional redundancy. Previously, we showed that the arr3,4,5,6,7,15 mutant has 357

reduced branching and its buds are CK resistant – an unexpected result since type-A 358

ARRs are generally considered to be negative regulators of CK signalling (Müller et 359

al., 2015). Since the type-A ARRs are themselves transcriptionally induced by CK as 360

part of a negative feedback loop, loss-of-function mutations in these genes could 361

have complex phenotypes with respect to CK response. However, our demonstration 362

that the arr1 mutant has a branching phenotype opposite to arr3,4,5,6,7,15 suggests 363

that the type-A and type-B ARRs do indeed function antagonistically in bud 364

activation, leaving the paradox unresolved. 365

366

ARR1 and ARR3,4,5,6,7,15-independent CK signalling and shoot branching 367

ARR1 is a well-characterised positive regulator of CK signalling and regulates the 368

expression of CK-responsive genes (Sakai et al., 2001). Throughout our analyses of 369

shoot branching and associated auxin transport phenotypes, arr1 phenocopied CK 370

treatment, whereas arr3,4,5,6,7,15 phenocopied CK depletion, as in ipt3 (Figs. 1, 3-371

6, Figs. S2 and S4). This raises the interesting prospect that the arr mutant 372

phenotypes are due to feedback regulation of the pool of active CKs via synthesis or 373

degradation, such that reduced CK signalling in arr1 leads to an increase in the pool 374

of active CKs, while increased CK signalling in arr3,4,5,6,7,15 reduces the pool of 375



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active CKs. Such feedback between hormone signalling and hormone levels is well-376

established for most plant hormones, including CK. For example, CK application 377

induces expression of genes encoding CYTOKININ OXIDASE (CKX) degrading 378

enzymes (Kiba et al., 2002; Rashotte et al., 2003), and this has also been observed 379

in CK-treated buds (Müller et al., 2015). Consistent with this feedback, the triple ahk2 380

ahk3 cre1 CK receptor mutant has elevated levels of N6-(Δ2-isopentenyl)-adenine 381

precursor and trans-zeatin-type CKs in young plants (Riefler et al., 2006). This 382

feedback has the potential to account for the “high CK” increased branching 383

phenotype of the arr1 mutant and the “low CK” reduced branching phenotype of the 384

arr3,4,5,6,7,15 mutant (Fig. 10). 385

386

Importantly, in addition to shoot branching, the phenotypic correlations we observe in 387

arr mutants extend to auxin transport properties. CK treatment, nitrate supply, and 388

loss of ARR1 result in over-accumulation of PIN3, PIN4 and PIN7, whereas CK-389

depleting conditions such as ipt3 mutation or nitrate starvation, and loss of the 390

ARR3, 4, 5, 6, 7, 15 clade result in PIN3, PIN4, and PIN7 depletion. If CK 391

homeostasis is indeed altered in the arr mutants, it implies that CK can signal 392

independently of the ARR genes analysed here to promote PIN3, PIN4 and PIN7 393

accumulation (Fig. 10). One possibility is that CK can signal entirely independently of 394

ARRs via a non-canonical pathway or simply independently of ARR1 and ARR3, 4, 395

5, 6, 7, 15, for example through a specialised type-B family member (or members) 396

via the canonical pathway. Further work will be required to understand precisely 397

which CK signalling components target PIN proteins in shoots. No differences were 398

observed in PIN1 accumulation, suggesting specific auxin transporters are targeted 399

by this signalling route. 400



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401

The lack of change in PIN gene expression levels in type-A and type-B arr mutants 402

and in wild-type plants in response to CK addition suggests PIN transcription is not a 403

direct target for CK signalling in bud activation, by whatever pathway, and suggests 404

that the effect of CK on PINs is post-transcriptional (Figs. S3 and S6). This could be 405

a protein-level effect on PINs, similar to that established for SLs and PIN1, or it could 406

be a transcriptional effect on PIN regulators. For example, PINOID-dependent (PID) 407

phosphorylation can modulate the apical-basal polarity of PINs (Friml et al., 2004) 408

and PID expression is reduced in shoots within 2 h of CK treatment in Arabidopsis 409

seedlings, consistent with CK promoting basal localisation of PINs (Brenner and 410

Schmulling 2012). Such a post-transcriptional effect on PINs acting via the canonical 411

CK signalling pathway has been established in roots. In the octuple 412

arr3,4,5,6,7,8,9,15 mutant, lower levels of GFP-tagged translational fusions of PIN1, 413

3 and 4 are observed in root tips, whereas PIN7 is reduced in the stele but increased 414

in the root cap, and these changes do not correlate with PIN transcript abundances 415

(Zhang et al., 2011). In contrast to our results, this effect mirrors CK treatment, as 416

expected if the type-A ARRs are acting as negative regulators of CK signalling. 417

418

The role of the auxin transport system in the regulation of shoot branching by 419

CK 420

Our results show a strong correlation between CK, basal membrane abundance of 421

PIN3, PIN4 and PIN7, bulk auxin transport and shoot branching across all our 422

experiments. Reducing auxin transport with the pharmacological inhibitor NPA has 423

opposite effects on branching in arr1 and arr3,4,5,6,7,15 (Fig. 3), suggesting a 424

causal link between auxin transport perturbation and the arr mutant shoot branching 425



19

phenotypes. The pin3,4,7 triple mutant partially suppresses the increased branching 426

of arr1 mutants, consistent with the idea that this phenotype is caused in part by 427

over-accumulation of these PINs (Bennett et al., 2016) (Fig. 8). One explanation for 428

this relationship is that CK-mediated increases in PIN3, PIN4 and PIN7 accumulation 429

increase the initial flow of auxin between the bud and the PATS, thereby supporting 430

the establishment of canalised auxin transport between the bud and the stem and 431

increasing the ease with which buds can activate. Consistent with this idea, polarised 432

PIN1 protein can be observed at the bud-stem junction in pea within 24 h of CK 433

treatment (Kalousek et al., 2010) and CK applied to axillary buds can promote the 434

export of auxin out of the buds (Li and Bangerth 2003). This hypothesis is also 435

consistent with the resistance of arr1 buds to the inhibitory effects of the SL 436

analogue GR24 (Fig. 2). SL acts in part by dampening the positive feedback in auxin 437

transport canalisation between the bud and the stems. Increased PIN3, PIN4 and 438

PIN7-mediated bud-stem auxin flux could counteract SL-mediated PIN1 removal by 439

promoting additional flux-correlated PIN1 allocation. According to this model, it is the 440

role of PIN3, PIN4 and PIN7 in cross-stem auxin flux, rather than basipetal transport 441

down the stem that is important in CK-mediated bud activation. This is consistent 442

with our previous analyses demonstrating that PIN3, PIN4 and PIN7 play an 443

important role in the ability of consecutive buds on opposite sides of the stem to 444

inhibit one another’s outgrowth, while having only limited impact on the total level of 445

shoot branching in intact plants. 446

447

Importantly, the pin3,4,7 mutant only partially suppresses the arr1 shoot branching 448

phenotype and pin3,4,7 buds respond strongly to basal BA supply in isolated nodal 449

assays, even activating slightly earlier than wild-type buds when treated with a 450



20

combination of apical NAA and basal BA. These results demonstrate that CK can 451

activate buds through PIN3-, PIN4- and PIN7-independent mechanisms. The 452

mechanism(s) underlying faster pin3,4,7 bud activation in response to apical NAA 453

and basal BA is not known. In Arabidopsis, the ability of strigolactone to inhibit bud 454

activity appears to depend jointly on its ability to promote accumulation of transcripts 455

of the BRC1 gene and to trigger endocytosis of PIN1, reducing the ability of buds to 456

establish canalised auxin export into the stem. Our data for CK support a similar dual 457

activity for CK, reducing BRC1 transcript abundance and increasing PIN3,4,7 458

accumulation at the plasma membrane (Seale et al., 2017) (Fig. 6). One highly 459

speculative possibility is that the combination of low BRC1 expression and reduced 460

peripheral stem auxin in pin3,4,7 mutants might allow for rapid early expansion of the 461

bud. 462

463

Other targets for CK signalling in shoot branching 464

Our data suggest CK promotes shoot branching in part by driving plasma membrane 465

accumulation of PIN3, PIN4 and PIN7. This would operate in parallel with the 466

established ability of CK to regulate transcription in buds, for example by down-467

regulating expression of the BRC1 bud regulatory gene (Fig. 10) (Aguilar-Martínez et 468

al., 2007; Poza-Carrion et al., 2007; Braun et al., 2012; Dun et al., 2012; Seale et al., 469

2017). The parallel operation of these two mechanisms makes interpretation of the 470

arr mutant phenotypes complicated. For example, in the arr1 mutant, impaired CK-471

induced changes in transcription should lead to reduced shoot branching. Consistent 472

with this, the arr1 mutant has reduced steady-state type-A ARR gene expression, as 473

expected if there is reduced CK signalling via the canonical pathway (Fig. S8). At the 474

same time, it is possible that this results in over-accumulation of CK due to impaired 475



21

feedback regulation on CK levels. This CK could signal in an ARR1-independent 476

manner to promote PIN3, PIN4 and PIN7 accumulation, thereby promoting shoot 477

branching. Thus, it is likely that the arr1 shoot branching phenotype is a compromise 478

between these opposing effects. Added to this, there is likely to be some redundancy 479

in the type-B ARR family and it is possible that different family members are 480

differentially important in regulating feedback on CK synthesis versus modulation of 481

transcription of bud-regulating genes. Given all these considerations, there are many 482

alternative ways to interpret our result that arr1 mutant buds inhibited by apical NAA 483

supply can be activated by basal BA similar to wild type, whereas arr3,4,5,6,7,15 484

buds cannot (Müller et al., 2015). 485

486

CONCLUSIONS 487

Several studies have proposed a causal link between CK and increased auxin 488

transport as one mechanism for CK-mediated bud outgrowth (Davies et al., 1966; 489

Chatfield et al., 2000; Li and Bangerth 2003). Our results support this hypothesis, 490

and in particular we demonstrate that CK drives accumulation of PIN3, PIN4 and 491

PIN7 on the plasma membrane, and this contributes to the branching phenotype 492

observed in the arr1 mutant. Interestingly, the phenotypes of type-A and type-B arr 493

CK signalling mutants are the opposite of those expected given their respective 494

negative and positive roles in CK-mediated transcriptional control in roots. This 495

suggests strong specialisation within the ARR gene families or an ARR-independent 496

mechanism for CK signalling in the control of shoot branching. 497

498

499



22

500



23

MATERIALS & METHODS 501

502

Plant lines 503

Col-0 was used as wild type for all experiments. The arr3,4,5,6,7,15 line was 504

published previously (Müller et al., 2015). The homozygous arr1-4 (SALK_042196) 505

T-DNA insertion line was obtained from the Nottingham Arabidopsis Stock Centre 506

and identified using ARR1-4 LP (5’-GAT CAA ACC CAT TCA ATG TCG-3’), ARR1-4 507

RP (5’-GAG ATG GCA TTG TCT CTG CTC-3’) and LBb1.3 as per the SALK T-DNA 508

website (www.signal.salk.edu/tdnaprimers.2.html) (Alonso et al., 2003). The 509

PIN1:PIN1-GFP (Benková et al., 2003), PIN3:PIN3-GFP (Žádníková et al., 2010), 510

PIN4:PIN4-GFP (Bennett et al., 2016) and PIN7:PIN7-GFP (Blilou et al., 2005) 511

reporter lines have been described previously. All PIN:PIN-GFP reporter lines on the 512

arr3,4,5,6,7,15 mutant background were generated by crossing arr3,4,5,6,7,15 to 513

each of the arr3,4,5,6,7,8,9,15 PIN-GFP lines described previously (Zhang et al., 514

2011) and screened for the presence of wild-type ARR8 (F 5’-CAA ATG GCT GTT 515

AAA ACC CAC CAA TA-3’ and R 5’-CCA TTG TTA GTG TGC TAT CAC CTG AGT 516

G-3’) and ARR9 genes (F 5’-CAG ACT CTT TAT TTC TCT TCC TC-3’ and R 5’-CCC 517

ACA TAC AAC ATC ATC ATC ATA TTC C-3’). For arr1, the four Col-0 PIN:PIN-GFP 518

lines were crossed to arr1-4. Segregants were screened in the F2 and F3 519

generations for GFP and the correct genotype using the above gene-specific primers 520

and ARR3, -4, -5, -6, -7 and -15 primers published previously (To et al., 2004b; 521

Zhang et al., 2011). 522

523

Growth conditions 524


http://www.signal.salk.edu/tdnaprimers.2.html


24

Seeds were stratified for 3-5 days at 4°C. Soil-grown plants were sown onto F2 soil 525

treated with Intercept 70WG (both Levington, http://www.scottsprofessional.co.uk) or 526

Exemptor (ICL, http://icl-sf.com/uk-en) in P24 or P40 trays in controlled environment 527

rooms under long-day (16 h light/ 8 h dark) or short-day (8 h light/16 h dark) 528

photoperiods. For plants grown under sterile conditions, seeds were vapour-529

sterilised with 100 mL 10% (w/v) chlorine bleach and 3 mL 10.2 M HCl for 4 h and 530

sown onto solidified ATS medium (containing 0.8% agar) (Wilson et al., 1990). Soil 531

and sterile-grown plants were subject to an average light intensity of 170 or 100 μmol 532

m-2 sec-1, respectively, and an average temperature range of 17–21°C. 533

534

Branch counts 535

Primary rosette and cauline branches ≥1 cm were counted on intact plants at or near 536

terminal flowering. Decapitation assays were performed as per Greb et al. (2003). 537

538

Hormone and NPA treatments 539

For testing bud responses to NAA, BA and GR24, plants were grown under sterile 540

conditions and one-node assays performed as previously described (Chatfield et al., 541

2000; Müller et al., 2015). For treating whole plants with NPA or GR24, plants were 542

grown for 6 weeks under sterile conditions on solidified ATS in glass jars as 543

described previously (Bennett et al., 2006; Crawford et al., 2010). For NAA and BA 544

treatment of stems, 2 cm segments from the basal inflorescence internode were 545

collected from soil-grown plants. Similar to the one-node assay system, segments 546

were placed vertically into split plates prepared with 1 μM NAA in the upper apical 547

portion and either mock (DMSO) or 1 μM BA in the lower basal portion, or the apical 548

portion. Plates were prepared by cutting a 1 cm trough through the middle of square 549


http://www.scottsprofessional.co.uk/

http://icl-sf.com/uk-en


25

10 cm plates containing 50 mL solidified ATS without sucrose. Hormone or mock 550

solutions were pipetted into the upper or lower half (25 μL of 1000X stocks). Plates 551

were placed under standard light conditions for sterile-grown plants. 552

553

Reverse transcription quantitative PCR 554

For analysis of PIN1, PIN3, PIN4 and PIN7 gene expression levels in the arr1 and 555

arr3,4,5,6,7,15 mutants, seeds were sown in glass jars on solidified ATS as 556

described above for whole-plant NPA and GR24 treatments. Plants were grown for 6 557

weeks and the uppermost inflorescence internodes (i.e. the inflorescence stem 558

between the uppermost two cauline nodes) harvested onto liquid nitrogen. Three 559

biological replicates each containing 10-15 internodes were collected. For NAA and 560

BA treatments, 2 cm segments from the basal inflorescence internodes were 561

collected from 6 week old soil-grown plants and placed vertically into plates 562

containing apical NAA with or without basal BA, prepared as above. Treatments 563

were left for 4 h and the basal 5 mm of the 2 cm segments were harvested into three 564

biological replicates of 6-7 segments each on liquid nitrogen. RNA extractions, cDNA 565

synthesis and quantification of transcript levels were carried out as described 566

previously (Müller et al., 2015) using 650 or 500 ng total RNA for the arr or BA 567

response analyses, respectively. Sequences of primers used are as follows: PIN1 F 568

5’-CAG TCT TGG GTT GTT CAT GGC-3’, PIN1 R 5’-ATC TCA TAG CCG CCG CAA 569

AA-3’, PIN3 F 5’-CCA TGG CGG TTA GGT TCC TT-3’, PIN3 R 5’-ATG CGG CCT 570

GAA CTA TAG CG-3’, PIN4 F 5’-AAT GCT AGA GGT GGT GGT GAT G-3’, PIN4 R 571

5’-TAG CTC CGC CGT GGA ATT AG-3’, PIN7 F 5’-GGT GAA AAC AAA GCT GGT 572

CCG-3’ and PIN7 R 5’-CCG AAG CTT GTG TAG TCC GT-3’. For Fig. S8, ARR1 F 573

5’-TAC GAA GTA ACG AAA TGC AAC AGA-3’, ARR1 R 5’-GAA ACC GTC CAT 574



26

GTC AGG CA-3’, and previously published primer sequences for ARR4, 5, 6, 7 and 575

15 were used (Müller et al., 2015). 576

577

Microscopy 578

For PIN1:PIN1-GFP and PIN3:PIN3-GFP, which exhibit relatively stable levels of 579

expression in the stem throughout development, basal inflorescence internodes from 580

~6 week old soil-grown plants were collected for imaging. For PIN4:PIN4-GFP and 581

PIN7:PIN7-GFP, which exhibit a reduction in stem expression during development, 582

inflorescence internodes were taken from 4–5-week-old soil-grown plants when the 583

inflorescence was approximately 5-15 cm high. Between treatments, individual 584

plants were stage-matched so that the same internodes from two plants with 585

inflorescence heights within 1 cm of each other were compared. Internodes were 586

hand-sectioned longitudinally through vascular bundles using a razor blade under a 587

dissecting microscope, or transversely in 2 mm segments, then secured to a petri 588

dish with micropore tape or embedded in solidified ATS and covered in water. 589

Confocal microscopy was performed using a Zeiss LSM700 imaging system with 590

20X or 10X water immersion lenses. For longitudinal sections, xylem parenchyma 591

tissues were located by focussing on the helical pattern of xylem cell wall lignin 592

under transmitted light. Excitation was performed using 488 nm (3-6% laser power) 593

and 639 nm (2% laser power) lasers. Images were acquired using SP 555 and 594

LP640 emission filters for GFP and chloroplast autofluorescence, respectively. The 595

same detection settings were used within an experiment. For longitudinal sections, 596

GFP fluorescence intensities were quantified by manually tracing around the basal 597

plasma membrane of a non-saturated cell using Zeiss ZEN 2012 software. At least 598

five cells each from 8-10 individual plants were analysed per treatment and repeated 599



27

at least once. For transverse sections, Z-stacks were acquired using a 3-5 μm step 600

size. Maximum projections generated using ImageJ software and GFP fluorescence 601

(as arbitrary units per pixel) quantified by manually tracing around vascular bundles. 602

Five vascular bundles were analysed from at least five individual plants and repeated 603

at least once. For representative images, projections were made using LSM Image 604

Browser software version 4.2.0.121 and processed in Adobe Photoshop. 605

606

Auxin transport assays 607

Auxin transport assays were performed as per (Bennett et al., 2016). 608

609

Branch angle measurements 610

Inflorescence stems bearing two branches from mature plants were trimmed, laid flat 611

and photographed. The angle between the point of emergence and the inflorescence 612

stem was measured using ImageJ software. 613

614

Statistical analyses 615

Statistical analyses were performed using SPSS Statistics version 22. 616

ACCESSION NUMBERS 617

Sequence data form this article can be found on The Arabidopsis Information 618

Resource (TAIR; www.arabidopsis.org ) under accession numbers AT3G16857 619

(ARR1), AT1G59940 (ARR3), AT1G10470 (ARR4), AT3G48100 (ARR5), 620

AT5G62920 (ARR6), AT1G19050 (ARR7), AT1G74890 (ARR15), AT3G63110 621

(IPT3), AT1G73590 (PIN1), AT1G70940 (PIN3), AT2G01420 (PIN4) and 622

AT1G23080 (PIN7). 623



28

624

SUPPLEMENTAL FIGURES 625

Figure S1. GR24 dose response of wild-type and arr1. 626

Figure S2. ipt3 has reduced stem auxin transport. 627

Figure S3. PIN gene expression in wild-type, arr1 and arr3,4,5,6,7,15 mutants. 628

Figure S4. PIN7-GFP is reduced in ipt3 inflorescence stems. 629

Figure S5. PIN7-GFP expression in arr1 inflorescence stems over time. 630

Figure S6. PIN gene expression is unchanged in inflorescence stems after 4 h BA 631

treatment. 632

Figure S7. BA promotes accumulation of PIN7:PIN7-GFP on the basal plasma 633

membrane of xylem parenchyma cells within 2 h. 634

Figure S8. Expression of ARR genes in arr mutants. 635

636

ACKNOWLEDGEMENTS 637

We thank Joseph Kieber for kindly supplying the arr3,4,5,6,7,8,9,15 PIN-GFP 638

reporter lines, Ruth Stephens for excellent technical assistance and Tom Bennett 639

and Martin van Rongen for helpful advice and discussions. 640

641

642



29

FIGURE LEGENDS 643

644

Figure 1. An ARR1 mutation confers increased shoot branching. A) Rosette 645

branches formed 10 days after primary shoot decapitation in plants grown in short 646

days for 4 weeks and long days for ~3 weeks, as per Greb et al. (2006) (n = 38-40). 647

B) Total branches (rosette and cauline) formed in Col-0 and arr1-4 plants grown in 648

long days for ~6 weeks (n = 30-31). C-E) Shoot branching phenotypes of Col-0 and 649

arr1-4 plants grown in long days for ~7 weeks. A close-up view of the rosettes in C 650

are shown in D (Col-0) and E (arr1-4). Bars indicate S.E.M., statistical comparisons 651

shown are between mutant and wild type (Mann-Whitney test, ** = p < 0.01, *** = < 652

0.001). 653

654

Figure 2. arr1 buds have wild-type responses to auxin and cytokinin but are resistant 655

to strigolactone. Wild type (A and C) or arr1 (B and D) isolated nodal segments 656

bearing one bud were treated for 8 days with (A and B) mock, 1 μM NAA (apical) 1 657

μM BA (basal) or combined 1 μM NAA (apical) and 1 μM BA (basal) (n = 18-20 per 658

treatment), or (C and D) 0.5 μM NAA (apical) or combined 0.5 μM NAA (apical) and 659

5 μM GR24 (n = 18-20). Bars indicate S.E.M. Statistical comparisons shown were 660

made between NAA and NAA + GR24 treated buds at each time point (Student’s t-661

test, * = p < 0.05, ** = p < 0.01). 662

663

Figure 3. Type-A and type-B arr mutant branching phenotypes are associated with 664

altered auxin transport. A) Amount of apically applied 14C-IAA (CPM; counts per 665

minute) transported to the basal end of inflorescence stem internodes from ~6 week 666

old wild-type, arr1 and arr3,4,5,6,7,15 plants in 6 hours (n = 24-32). Different letters 667



30

denote significant difference (ANOVA, Tukey-b post-hoc test, p < 0.05). B) NPA dose 668

response of wild-type and arr1 plants grown for 6 weeks on different concentrations 669

of NPA under sterile conditions (n = 18-42). C) NPA dose response of wild-type and 670

arr3,4,5,6,7,15 plants grown for 6 weeks on different concentrations of NPA under 671

sterile conditions (n = 13-24). For A-C bars indicate S.E.M. For B and C, letters 672

denote significant difference within a genotype (Kruskal-Wallis H test, p < 0.05). 673

674

Figure 4. PIN3-GFP and PIN7-GFP accumulation is attenuated in arr3,4,5,6,7,15 675

inflorescence stems. Representative accumulation patterns of PIN1-GFP (A and B), 676

PIN3-GFP (D and E) and PIN7-GFP (G and H) in basal inflorescence internodes of 677

Col-0 (A, D and G) and arr3,4,5,6,7,15 (B, E and H) plants, imaged at ~5-6 weeks of 678

age using confocal microscopy. For confocal images, green shows PIN-GFP signal 679

and magenta shows chloroplast autofluorescence. Scale bars represent 10 μm (A, B, 680

D and E) or 200 μm (G and H). For PIN1:PIN1-GFP and PIN3:PIN3-GFP, plants 681

were sectioned longitudinally and the amount of GFP signal on the basal plasma 682

membrane was quantified in the xylem parenchyma using at least five cells each 683

from eight independent plants (C and F) (n = 40). For PIN7:PIN7-GFP, plants were 684

sectioned transversely (~2 mm thickness) and the amount of GFP signal was 685

quantified in at least five vascular bundles from five independent plants with 686

inflorescence stems stage-matched at 24-26 cm in height (I) (n = 25). Bars indicate 687

S.E.M., statistical comparisons shown were made between mutant and wild type 688

(Student’s t-test, p < 0.001). 689

690

Figure 5. PIN3-GFP and PIN7-GFP accumulation is elevated in arr1 inflorescence 691

stems. Representative accumulation patterns of PIN1-GFP (A and B), PIN3-GFP (D 692



31

and E) and PIN7-GFP (G and H) in basal inflorescence internodes of 4–7-week-old 693

Col-0 (A, D and G) and arr1 (B, E and H) plants, sectioned longitudinally and imaged 694

using confocal microscopy. Green shows PIN-GFP signal and magenta shows 695

chloroplast autofluorescence. Scale bars represent 10 μm. The amount of GFP 696

signal on the basal plasma membrane was quantified in the xylem parenchyma 697

using at least five cells each from eight independent plants (C, F and I) (n = 40). For 698

PIN1:PIN1-GFP and PIN3:PIN3-GFP, plants were analysed at 6-7 weeks of age. For 699

PIN7:PIN7-GFP, plants were stage-matched by comparing plants with inflorescence 700

stems 18-20 cm in height. Bars indicate S.E.M., statistical comparisons shown were 701

made between mutant and wild type (Student’s t-test, * = p < 0.05, ** = p < 0.01). 702

703

Figure 6. Cytokinin promotes the accumulation of PIN3-GFP, PIN4-GFP and PIN7-704

GFP on the basal plasma membrane of xylem parenchyma cells in inflorescence 705

stems. Vertically held basal inflorescence internode segments (~2 cm) from 706

PIN1:PIN1-GFP (A-C), PIN3:PIN3-GFP (D-F), PIN4:PIN4-GFP (G-I) and PIN7:PIN7-707

GFP (J-L) plants (all Col-0 background), were treated apically with 1 μM NAA and 708

basally with either 0.1% DMSO control (“Mock”) or 1 μM BA (“+ BA”). After 4 hours, 709

stem segments were sectioned longitudinally at their basal ends and imaged using 710

confocal microscopy. The amount of GFP signal on the basal plasma membrane of 711

xylem parenchyma cells was quantified in five cells each from at least seven 712

independent plants for PIN1:PIN1-GFP (A), ten independent plants for PIN3:PIN3-713

GFP (D) and PIN4:PIN4-GFP (G), or eleven independent plants for PIN7:PIN7-GFP 714

(J). Bars indicate S.E.M., statistical comparisons shown were made between mock 715

and BA treated stems (Student’s t-test, * = p < 0.05, ** = p < 0.01). Representative 716

confocal images used for GFP quantifications are shown for mock (B, E, H and K) 717



32

and BA (C, F, I and L) treated stems. Green shows PIN-GFP signal and magenta 718

shows chloroplast autofluorescence. PIN1:PIN1-GFP and PIN3:PIN3-GFP plants 719

were analysed at 5-6 weeks of age. PIN4:PIN4-GFP and PIN7:PIN7-GFP plants 720

were analysed at 4-5 weeks of age and stage-matched across treatments according 721

to inflorescence height. Scale bars represent 10 μm. 722

723

Figure 7. Growth on low nitrate (N) conditions reduces PIN-GFP accumulation. 724

PIN3:PIN3-GFP (A), PIN4:PIN4-GFP (B) and PIN7:PIN7-GFP (C) plants (all Col-0 725

background) were grown on a sand and terragreen mix supplemented with ATS 726

medium containing 9.0 mM NO3 (high N) or 1.8 mM NO3 (low N). Basal inflorescence 727

internodes were sectioned longitudinally and imaged at 4-6 weeks of age using 728

confocal microscopy. PIN7:PIN7-GFP plants were stage matched based on 729

inflorescence heights across N treatments. The amount of GFP signal on the basal 730

plasma membrane was quantified in the xylem parenchyma using five cells each 731

from eight independent plants for (C) (n = 40). Bars indicate S.E.M., statistical 732

comparisons shown were made between high and low N treated plants (Student’s t-733

test, p < 0.001). 734

735

Figure 8. Interactions between arr1 and pin3,4,7 in shoot branching control. A-E) 736

Shoot phenotypes of ~6 week old Col-0, arr1, pin3,4,7 and arr1 pin3,4,7 plants 737

grown under long days. A close-up view of the rosettes in A are shown in B (Col-0), 738

C (arr1), D (pin3,4,7) and E (arr1 pin3,4,7). F) Total number of branches formed at 739

terminal flowering on ~9 week old wild-type, arr1, pin3,4,7 and arr1 pin3,4,7 plants 740

grown under short days for 4 weeks then long days for 5 weeks (n = 16-47). Bars 741

indicate S.E.M. Letters denote significant difference between genotypes (Kruskal-742



33

Wallis H test, p < 0.05). G) Angle between emergence point of branches on the 743

primary inflorescence in wild-type, arr1, pin3,4,7 and arr1 pin3,4,7 plants at ~6 744

weeks of age. Two cauline branches were analysed per plant from at least 13 745

independent plants (n = 26-46). Bars indicate S.E.M. Letters denote significant 746

difference (ANOVA, Tukey-b post-hoc test, p < 0.05). 747

748

Figure 9. pin3,4,7 exhibits altered bud activation in response to BA. Bud length of 749

wild type (Col-0) (A) and pin3,4,7 (B) isolated nodal segments treated for 8 days with 750

mock, 1 μM NAA (apical), 1 μM BA (basal) or combined 1 μM NAA (apical) and 1 μM 751

BA (basal) (n = 19-20 per treatment). (C) Close-up of days 0-4 for wild-type and 752

pin3,4,7 buds treated with combined apical NAA and basal BA. Bars indicate S.E.M. 753

Asterisks denote statistically significant difference between treatments (Student’s t-754

test, * = p < 0.05, *** = p < 0.001). 755

756

Figure 10. Proposed model for CK signalling in bud outgrowth regulation. Canonical 757

CK signalling via the type-B ARR1 results in feedback-mediated down-regulation of 758

CK levels as part of signal perception. Members of the type-A ARR family are 759

transcriptionally induced by ARR1 and/or other type-B family members, and dampen 760

type-B-mediated signalling, including the feedback-mediated down-regulation of CK 761

levels. Basal plasma-membrane accumulation of PIN3, PIN4 and PIN7 in xylem 762

parenchyma cells in the main stem is enhanced by CK by an unknown mechanism. 763

In parallel, expression of other bud regulatory genes such as BRC1 may be targeted 764

by the canonical CK signalling pathway. 765

766

LITERATURE CITED 767



34

768

Aguilar-Martínez, J.A., Poza-Carrión, C. and Cubas, P. (2007) Arabidopsis 769

BRANCHED1 acts as an integrator of branching signals within axillary buds. 770

Plant Cell, 19, 458-472. 771

Alonso, J.M., Stepanova, A.N., Leisse, T.J., Kim, C.J., Chen, H., Shinn, P., 772

Stevenson, D.K., Zimmerman, J., Barajas, P., Cheuk, R., Gadrinab, C., Heller, 773

C., Jeske, A., Koesema, E., Meyers, C.C., Parker, H., Prednis, L., Ansari, Y., 774

Choy, N., Deen, H., Geralt, M., Hazari, N., Hom, E., Karnes, M., Mulholland, 775

C., Ndubaku, R., Schmidt, I., Guzman, P., Aguilar-Henonin, L., Schmid, M., 776

Weigel, D., Carter, D.E., Marchand, T., Risseeuw, E., Brogden, D., Zeko, A., 777

Crosby, W.L., Berry, C.C. and Ecker, J.R. (2003) Genome-wide insertional 778

mutagenesis of Arabidopsis thaliana. Science, 301, 653-657. 779

Bainbridge, K., Sorefan, K., Ward, S. and Leyser, O. (2005) Hormonally controlled 780

expression of the Arabidopsis MAX4 shoot branching regulatory gene. Plant 781

J., 44, 569-580. 782

Bangerth, F. (1994) Response of cytokinin concentration in the xylem exudate of 783

bean (Phaseolus vulgaris L.) plants to decapitation and auxin treatment, and 784

relationship to apical dominance. Planta, 194, 4. 785

Benková, E., Michniewicz, M., Sauer, M., Teichmann, T., Seifertová, D., Jürgens, G. 786

and Friml, J. (2003) Local, efflux-dependent auxin gradients as a common 787

module for plant organ formation. Cell, 115, 591-602. 788

Bennett, T., Sieberer, T., Willett, B., Booker, J., Luschnig, C. and Leyser, O. (2006) 789

The Arabidopsis MAX pathway controls shoot branching by regulating auxin 790

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35

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Blilou, I., Xu, J., Wildwater, M., Willemsen, V., Paponov, I., Friml, J., Heidstra, R., 795

Aida, M., Palme, K. and Scheres, B. (2005) The PIN auxin efflux facilitator 796

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Booker, J., Chatfield, S. and Leyser, O. (2003) Auxin acts in xylem-associated or 799

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shoot branching. Plant Physiol., 158, 225-238. 805

Brenner, W.G. and Schmulling, T. (2012) Transcript profiling of cytokinin action in 806

Arabidopsis roots and shoots discovers largely similar but also organ-specific 807

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Chatfield, S.P., Stirnberg, P., Forde, B.G. and Leyser, O. (2000) The hormonal 809

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Crawford, S., Shinohara, N., Sieberer, T., Williamson, L., George, G., Hepworth, J., 811

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competition between shoot branches by dampening auxin transport. 813

Development, 137, 2905-2913. 814

Davies, C.R., Seth, A.K. and Wareing, P.F. (1966) Auxin and kinetin interaction in 815

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Plant Physiol, 166, 384-395. 820

Domagalska, M.A. and Leyser, O. (2011) Signal integration in the control of shoot 821

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1

Col-0 arr1-4

C

Col-0 arr1-4

Rose

tte b

ranc

hes

0

2

4

6

8

10

***

A

Col-0 arr1-4

Tota

l bra

nche

s

0

2

4

6

8

10

B

***

D E

Figure 1. An ARR1 mutation confers increased shoot branching. A) Rosette

branches formed 10 days after primary shoot decapitation in plants grown in short

days for 4 weeks and long days for ~3 weeks, as per Greb et al. (2006) (n = 38-40).



2

B) Total branches (rosette and cauline) formed in Col-0 and arr1-4 plants grown in

long days for ~6 weeks (n = 30-31). C-E) Shoot branching phenotypes of Col-0 and

arr1-4 plants grown in long days for ~7 weeks. A close-up view of the rosettes in C

are shown in D (Col-0) and E (arr1-4). Bars indicate S.E.M., statistical comparisons

shown are between mutant and wild-type (Mann-Whitney test, ** = p < 0.01, *** = <

0.001).



1

A

Days

0 1 2 3 4 5 6 7 8

Me

an

bu

d le

ng

th (

mm

)

0

5

10

15

20

25

30

Mock

BA

NAA

NAA + BA

Col-0

Days

0 1 2 3 4 5 6 7 8

Me

an

bu

d le

ng

th (

mm

)

0

5

10

15

20

NAA + mock

NAA + GR24C

Col-0

*

*

*

*

Days

0 1 2 3 4 5 6 7 8

Me

an

bu

d le

ng

th (

mm

)

0

5

10

15

20

NAA + mock

NAA+GR24 D

arr1-4** **

Days

0 1 2 3 4 5 6 7 8

Me

an

bu

d le

ng

th (

mm

)

0

5

10

15

20

25

30

Mock

BA

NAA

NAA + BA

B

arr1-4

A

Figure 2. arr1 buds have wild-type responses to auxin and cytokinin but are resistant

to strigolactone. Wild-type (A and C) or arr1 (B and D) isolated nodal segments

bearing one bud were treated for 8 days with (A and B) mock, 1 μM NAA (apical) 1

μM BA (basal) or combined 1 μM NAA (apical) and 1 μM BA (basal) (n = 18-20 per

treatment), or (C and D) 0.5 μM NAA (apical) or combined 0.5 μM NAA (apical) and

5 μM GR24 (n = 18-20). Bars indicate S.E.M. Statistical comparisons shown were

made between NAA and NAA + GR24 treated buds at each time point (Student’s t-

test, * = p < 0.05, ** = p < 0.01).



1

Col-0 arr1-4

Tota

l bra

nche

s

0

1

2

3

4

5

0 µM NPA 0.01 µM NPA 0.1 µM NPA 1.0 µM NPA

Col-0 arr1-4 arr3,4,5, 6,7,15

Aux

in tr

ansp

orte

d (C

PM)

0

20

40

60

80

100A

a

b

c

B

ABA A

B

a abbc

c

Col-0 arr3,4,5,6,7,15

Tota

l bra

nche

s

0

1

2

3

4

5

0 µM NPA 0.001 µM NPA 0.01 µM NPA 0.1 µM NPA

C

B

A

a

bcab

cA

AB

Figure 3. Type-A and type-B arr mutant branching phenotypes are associated with

altered auxin transport. A) Amount of apically applied 14C-IAA (counts per minute)

transported to the basal end of inflorescence stem internodes from ~6 week old wild-

type, arr1 and arr3,4,5,6,7,15 plants in 6 hours (n = 24-32). Different letters denote

significant difference (ANOVA, Tukey-b post-hoc test, p < 0.05). B) NPA dose



2

response of wild-type and arr1 plants grown for 6 weeks on different concentrations

of NPA under sterile conditions (n = 18-42). C) NPA dose response of wild-type and

arr3,4,5,6,7,15 plants grown for 6 weeks on different concentrations of NPA under

sterile conditions (n = 13-24). For A-C bars indicate S.E.M. For B and C, letters

denote significant difference within a genotype (Kruskal-Wallis H test, p < 0.05).



1

Col-0

A PIN1

Col-0 arr3,4,5, 6,7,15

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120PIN1CB

arr3,4,5,6,7,15

PIN1

PIN3D

Col-0

PIN3

arr3,4,5,6,7,15

E

Col-0 arr3,4,5, 6,7,15

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

10

20

30

40

50

60PIN3F

***

Col-0 arr3,4,5, 6,7,15

Arb

itrar

y un

its/p

ixel

/vas

cula

r bun

dle

0

10

20

30

40

I

***

PIN7G

Col-0

PIN7 H

arr3,4,5,6,7,15

PIN7

Figure 4. PIN3:PIN3-GFP and PIN7:PIN7-GFP expression is attenuated in

arr3,4,5,6,7,15 inflorescence stems. Representative expression patterns of

PIN1:PIN1-GFP (A and B), PIN3:PIN3-GFP (D and E) and PIN7:PIN7-GFP (G and

H) in basal inflorescence internodes of Col-0 (A, D and G) and arr3,4,5,6,7,15 (B, E

and H) plants, imaged at ~5-6 weeks of age using confocal microscopy. For confocal

images, green shows PIN-GFP signal and magenta shows chloroplast



2

autofluorescence. Scale bars represent 10 μm (A, B, D and E) or 200 μm (G and H).

For PIN1:PIN1-GFP and PIN3:PIN3-GFP, plants were sectioned longitudinally and

the amount of GFP signal on the basal plasma membrane was quantified in the

xylem parenchyma using at least five cells each from eight independent plants (C

and F) (n = 40). For PIN7:PIN7-GFP, plants were sectioned transversely (~2 mm

thickness) and the amount of GFP signal was quantified in at least five vascular

bundles from five independent plants with inflorescence stems stage-matched at 24-

26 cm in height (I) (n = 25). The outline of a vascular bundle is shown by a white

wedge. Bars indicate S.E.M., statistical comparisons shown were made between

mutant and wild-type (Student’s t-test, p < 0.001).



1

Col-0

A PIN1 B

arr1-4

PIN1

Col-0 arr1-4

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

PIN1C

PIN3D

Col-0

PIN3

arr1-4

E

Col-0 arr1-4Re

lativ

e flu

ores

cenc

e (a

rbitr

ary

units

)0

20

40

60

80

PIN3F

*

G

Col-0

PIN7 H

arr1-4

PIN7

Col-0 arr1-4

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

I

**

PIN7

Figure 5. PIN3:PIN3-GFP and PIN7:PIN7-GFP expression is elevated in arr1

inflorescence stems. Representative expression patterns of PIN1:PIN1-GFP (A and

B), PIN3:PIN3-GFP (D and E) and PIN7:PIN7-GFP (G and H) in basal inflorescence

internodes of 4-7 week old Col-0 (A, D and G) and arr1 (B, E and H) plants,

sectioned longitudinally and imaged between using confocal microscopy. Green

shows PIN-GFP signal and magenta shows chloroplast autofluorescence. Scale bars

represent 10 μm. The amount of GFP signal on the basal plasma membrane was



2

quantified in the xylem parenchyma using at least five cells each from eight

independent plants (C, F and I) (n = 40). For PIN1:PIN1-GFP and PIN3:PIN3-GFP,

plants were analysed at 6-7 weeks of age. For PIN7:PIN7-GFP, plants were stage-

matched by comparing plants with inflorescence stems 18-20 cm in height. Bars

indicate S.E.M., statistical comparisons shown were made between mutant and wild-

type (Student’s t-test, * = p < 0.05, ** = p < 0.01).



1

PIN3-GFP

Mock + BA

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

**

D PIN3-GFP

Mock

E F PIN3-GFP

+ BA

PIN1-GFP

Mock

BPIN1-GFP

Mock + BA

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

A PIN1-GFPC

+ BA

PIN4-GFP

Mock + BARela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

***

G PIN4-GFPH

Mock

PIN4-GFPI

+ BA

PIN7-GFP

Mock + BARela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

**

J PIN7-GFPK

Mock + BA

PIN7-GFPL

Figure 6. Cytokinin promotes the accumulation of PIN3:PIN3-GFP, PIN4:PIN4-GFP

and PIN7:PIN7-GFP on the basal plasma membrane of xylem parenchyma cells in



2

inflorescence stems. Vertically held basal inflorescence internode segments (~2 cm)

from PIN1:PIN1-GFP (A-C), PIN3:PIN3-GFP (D-F), PIN4:PIN4-GFP (G-I) and

PIN7:PIN7-GFP (J-L) plants (all Col-0 background), were treated apically with 1 μM

NAA and basally with either 0.1 % DMSO control (“Mock”) or 1 μM BA (“+ BA”). After

4 hours, stem segments were sectioned longitudinally at their basal ends and

imaged using confocal microscopy. The amount of GFP signal on the basal plasma

membrane of xylem parenchyma cells was quantified in five cells each from at least

seven independent plants for PIN1:PIN1-GFP (A), ten independent plants for

PIN3:PIN3-GFP (D) and PIN4:PIN4-GFP (G), or eleven independent plants for

PIN7:PIN7-GFP (J). Bars indicate S.E.M., statistical comparisons shown were made

between mock and BA treated stems (Student’s t-test, * = p < 0.05, ** = p < 0.01).

Representative confocal images used for GFP quantifications are shown for mock

(B, E, H and K) and BA (C, F, I and L) treated stems. Green shows PIN-GFP signal

and magenta shows chloroplast autofluorescence. PIN1:PIN1-GFP and

PIN3:PIN3GFP plants were analysed at 5-6 weeks of age. PIN4:PIN4-GFP and

PIN7:PIN7-GFP plants were analysed at 4-5 weeks of age and stage-matched

across treatments according to infloresence height. Scale bars represent 10 μm.



1

PIN7-GFP

High N Low N

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

PIN4-GFP

High N Low N

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

PIN3-GFP

High N Low N

Rela

tive

fluor

esce

nce

(arb

itrar

y un

its)

0

20

40

60

80

100

120

A

***

B***

C***

Figure 7. Growth on low nitrate (N) conditions reduces PIN-GFP accumulation.

PIN3:PIN3-GFP (A), PIN4:PIN4-GFP (B) and PIN7:PIN7-GFP (C) plants (all Col-0

background) were grown on a sand and terragreen mix supplemented with ATS

medium containing 9.0 mM NO3 (high N) or 1.8 mM NO3 (low N). Basal inflorescence

internodes were sectioned longitudinally and imaged at 4-6 weeks of age using

confocal microscopy. PIN7:PIN7-GFP plants were stage matched based on

inflorescence heights across N treatments. The amount of GFP signal on the basal



2

plasma membrane was quantified in the xylem parenchyma using five cells each

from eight independent plants for (C) (n = 40). Bars indicate S.E.M., statistical

comparisons shown were made between high and low N treated plants (Student’s t-

test, p < 0.001).



1

Col-0 arr1 pin3,4,7 arr1pin3,4,7

Bran

ch a

ngle

(deg

rees

)

0

20

40

60

80

100

G

a a

b b

Col-0 arr1 pin3,4,7 arr1pin3,4,7

Tota

l bra

nche

s

0

2

4

6

8

10

12

14 F

a

b

ac

Col-0 arr1 pin3,4,7 arr1 pin3,4,7

A B C

D E

Figure 8. Interactions between arr1 and pin3,4,7 in shoot branching control. A-E)

Shoot phenotypes of ~6 week old Col-0, arr1, pin3,4,7 and arr1 pin3,4,7 plants

grown under long days. A close-up view of the rosettes in A are shown in B (Col-0),

C (arr1), D (pin3,4,7) and E (arr1 pin3,4,7). F) Total number of branches formed at

terminal flowering on ~9 week old wild-type, arr1, pin3,4,7 and arr1 pin3,4,7 plants

grown under short days for 4 weeks then long days for 5 weeks (n = 16-47). Bars

indicate S.E.M. Letters denote significant difference between genotypes (Kruskal-

Wallis H test, p < 0.05). G) Angle between emergence point of branches on the

primary inflorescence in wild-type, arr1, pin3,4,7 and arr1 pin3,4,7 plants at ~6

weeks of age. Two cauline branches were analysed per plant from at least 13

independent plants (n = 26-46). Bars indicate S.E.M. Letters denote significant

difference (ANOVA, Tukey-b post-hoc test, p < 0.05).



1

0 1 2 3 4 5 6 7 8

Me

an

bu

d le

ng

th (

mm

)

0

5

10

15

20

25

30

35

Mock

NAA

BA

NAA + BA

A

Col-0

0 1 2 3 4 5 6 7 8

Me

an

bu

d le

ng

th (

mm

)

0

5

10

15

20

25

30

35

Mock

NAA

BA

NAA + BA

B

pin3,4,7

Days

0 1 2 3 4

Me

an

bu

d le

ng

th (

mm

)

0

1

2

3

4

5

6

Col-0

pin3,4,7

N.S.

N.S.

***

*

*

C

Figure 9. pin3,4,7 exhibits altered bud activation in response to BA. Bud length of

wild-type (Col-0) (A) and pin3,4,7 (B) isolated nodal segments treated for 8 days with

mock, 1 μM NAA (apical), 1 μM BA (basal) or combined 1 μM NAA (apical) and 1 μM

BA (basal) (n = 19-20 per treatment). (C) Close-up of days 0-4 for wild-type and

pin3,4,7 buds treated with combined apical NAA and basal BA. Bars indicate S.E.M.



2

Asterisks denote statistically significant difference between treatments (Student’s t-

test, * = p < 0.05, *** = p < 0.001).



1

Figure 10. Proposed model for CK signalling in bud outgrowth regulation. Canonical

CK signalling via the type-B ARR1 results in feedback down-regulation of CK levels

as part of signal perception. Members of the type-A ARR family are transcriptionally

induced by ARR1 and/or other type-B family members, and dampen type-B-

mediated signalling, including the feedback down-regulation of CK levels. Basal

plasma-membrane accumulation of PIN3, PIN4 and PIN7 in xylem parenchyma cells

in the main stem is enhanced by CK by an unknown mechanism. In parallel,

expression of other bud regulatory genes such as BRC1 may be targeted by the

canonical CK signalling pathway.



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Bainbridge, K., Sorefan, K., Ward, S. and Leyser, O. (2005) Hormonally controlled expression of the Arabidopsis MAX4 shoot branchingregulatory gene. Plant J., 44, 569-580.


Bangerth, F. (1994) Response of cytokinin concentration in the xylem exudate of bean (Phaseolus vulgaris L.) plants to decapitationand auxin treatment, and relationship to apical dominance. Planta, 194, 4.


Benková, E., Michniewicz, M., Sauer, M., Teichmann, T., Seifertová, D., Jürgens, G. and Friml, J. (2003) Local, efflux-dependent auxingradients as a common module for plant organ formation. Cell, 115, 591-602.


Bennett, T., Sieberer, T., Willett, B., Booker, J., Luschnig, C. and Leyser, O. (2006) The Arabidopsis MAX pathway controls shootbranching by regulating auxin transport. Curr. Biol., 16, 553-563.


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Friml, J., Yang, X., Michniewicz, M., Weijers, D., Quint, A., Tietz, O., Benjamins, R., Ouwerkerk, P.B., Ljung, K., Sandberg, G., Hooykaas,P.J., Palme, K. and Offringa, R. (2004) A PINOID-dependent binary switch in apical-basal PIN polar targeting directs auxin efflux.Science, 306, 862-865.


Geldner, N., Anders, N., Wolters, H., Keicher, J., Kornberger, W., Muller, P., Delbarre, A., Ueda, T., Nakano, A. and Jurgens, G. (2003)The Arabidopsis GNOM ARF-GEF mediates endosomal recycling, auxin transport, and auxin-dependent plant growth. Cell, 112, 219-230.


Greb, T., Clarenz, O., Schäfer, E., Müller, D., Herrero, R., Schmitz, G. and Theres, K. (2003) Molecular analysis of the LATERALSUPPRESSOR gene in Arabidopsis reveals a conserved control mechanism for axillary meristem formation. Genes Dev., 17, 1175-1187.


Hall, S.M. and Hillman, J.R. (1975) Correlative inhibition of lateral bud growth in Phaseolus vulgaris L.: timing of bud growth followingdecapitation. Planta, 123, 137-143.

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Wickson, M. and Thimann, K.V. (1958) The antagonism of auxin and kinetin in apical dominance. Physiol. Plantarum, 11, 62-74.Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

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Žádníková, P., Petrášek, J., Marhavý, P., Raz, V., Vandenbussche, F., Ding, Z., Schwarzerová, K., Morita, M.T., Tasaka, M., Hejátko, J.,Van Der Straeten, D., Friml, J. and Benková, E. (2010) Role of PIN-mediated auxin efflux in apical hook development of Arabidopsisthaliana. Development, 137, 607-617.


Zhang, W., To, J.P., Cheng, C.Y., Schaller, G.E. and Kieber, J.J. (2011) Type-A response regulators are required for proper root apicalmeristem function through post-transcriptional regulation of PIN auxin efflux carriers. Plant J., 68, 1-10.



http://www.ncbi.nlm.nih.gov/pubmed?term=Wickson%2C M%2E and Thimann%2C K%2EV%2E %281958%29 The antagonism of auxin and kinetin in apical dominance%2E Physiol%2E Plantarum%2C 11%2C 62%2D74%2E&dopt=abstract

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http://www.ncbi.nlm.nih.gov/pubmed?term=Zhang%2C W%2E%2C To%2C J%2EP%2E%2C Cheng%2C C%2EY%2E%2C Schaller%2C G%2EE%2E and Kieber%2C J%2EJ%2E %282011%29 Type%2DA response regulators are required for proper root apical meristem function through post%2Dtranscriptional regulation of PIN auxin efflux carriers%2E Plant J%2E%2C 68%2C 1%2D10%2E&dopt=abstract

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1 : cytokinin, shoot branching and pin accumulation 2 · 79 action of auxin on axillary bud growth...

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