Running Head: Rice TUT1 Regulates Panicle Development · 2015. 8. 4. · 9 144 RESULTS 145 146 Identification and characterization of the rice tut1 mutant 147 To understand the cellular

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Running Head: Rice TUT1 Regulates Panicle Development 3

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Corresponding author: 5

Jiaoteng Bai 6

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Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, 8

China 9

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Tel: (+8610)6480 6585 11

Fax: (+8610)6480 6595 12

E-mail: [email protected] 13

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Research Area: Genes, Development and Evolution 15

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Plant Physiology Preview. Published on August 4, 2015, as DOI:10.1104/pp.15.00229

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Rice TUTOU1 Encodes a SCAR-like Protein that Is Important for Actin 17

Organization and Panicle Development1 18

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Jiaoteng Bai2*, Xudong Zhu2, Qing Wang2, Jian Zhang, Hongqi Chen, Guojun Dong, Lei Zhu, 21

Huakun Zheng, Qingjun Xie, Jinqiang Nian, Fan Chen, Ying Fu, Qian Qian, Jianru Zuo 22

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State Key Laboratory of Plant Genomics and National Plant Gene Research Center, Institute of 24

Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China (J.B., 25

Q.W., J.Z., H.Z., Q.X., J.N. and J.Z.); University of Chinese Academy of Sciences, Beijing 100049, 26

China (J.B., Q.W., H.Z. and Q.X.); State Key Laboratory of Rice Biology, China National Rice 27

Research Institute, Chinese Academy of Agricultural Sciences, Hangzhou 310006, China (X.Z., H.C., 28

G.D. and Q.Q.). State Key Laboratory of Plant Physiology and Biochemistry, Department of Plant 29

Sciences, College of Biological Sciences, China Agricultural University (L.Z. and Y.F.), State Key 30

Laboratory of Molecular Developmental Biology and National Plant Gene Research Center, Institute 31

of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China 32

(F.C.) 33

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One sentence summary: 35

Rice TUTOU1 protein regulates panicle development by modulating actin organization. 36

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Total word count: 9,333 39

Total characters (including space): 59,829 40

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Footnotes 42

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1 This work was supported by grants from the Ministry of Science and Technology of China 44

(2014CB943402), State Key Laboratory of Plant Genomics (SKLPG2011A0210) and National 45

Natural Science Foundation of China (30700500). 46

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*Corresponding author e-mail: [email protected]. 48

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1 The author responsible for the distribution of materials integral to the findings presented in this 51

article in accordance with the policy described in the Instructions for Authors (www.plantphysiol.org) 52

is: Jiaoteng Bai ([email protected]). 53

2 These authors contributed equally to this work. 54

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ABSTRACT 57

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Panicle development, a key event in rice (Oryza sativa) reproduction and a critical determinant of 59

grain yield, forms a branched structure containing multiple spikelets. Genetic and environmental 60

factors can perturb panicle development, causing panicles to degenerate, and producing characteristic 61

whitish, small spikelets with severely reduced fertility and yield; however, little is known about the 62

molecular basis of the formation of degenerating panicles in rice. Here, we report the identification 63

and characterization of the rice panicle degenerative mutant tutou1 (tut1), which shows severe 64

defects in panicle development. The tut1 also shows a pleiotropic phenotype, characterized by short 65

roots, reduced plant height, and abnormal development of anthers and pollen grains. Molecular 66

genetic studies revealed that TUT1 encodes a SCAR/WAVE (suppressor of cAMP receptor/WASP 67

family verprolin-homologous) like protein. We found that TUT1 contains conserved functional 68

domains found in eukaryotic SCAR/WAVE proteins, and was able to activate Arp2/3 to promote 69

actin nucleation and polymerization in vitro. Consistently, tut1 mutants show defects in the 70

arrangement of actin filaments in trichome. These results indicate that TUT1 is a functional 71

SCAR/WAVE protein and plays an important role in panicle development. 72

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Key words: rice panicle development; TUT1; SCAR/WAVE; actin organization 74

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INTRODUCTION 76

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Rice (Oryza sativa L.) yield is closely associated with panicle development, including panicle 78

structure and panicle number. The basic unit of the rice panicle is the spikelet, which consists of 79

multiple florets (each having a lodicule, carpel, and six stamens) enclosed by glumes (Yoshida and 80

Nagato, 2011). During rice growth and development, panicle degeneration may occur under various 81

conditions (Komatsu et al., 2001; Ashikari et al., 2005; Ikeda et al., 2005). The abortion of spikelets 82

usually causes degeneration of floral organs, leading to reduction of rice yield. Spikelet degeneration 83

can occur from the basal or apical end of the panicle, and lead to defective, whitish spikelets, and 84

eventually fewer spikelets per panicle (Li et al., 2009; Cheng et al., 2011). 85

Various genetic and environmental factors cause panicle degeneration, but the underlying 86

mechanisms remain poorly understood (Xing and Zhang, 2010). Several quantitative trait loci (QTLs) 87

related to panicle apical spikelet abortion have been reported. One such QTL, qPAA8, affects the 88

formation of small and whitish anthers and sterile pollen, and an elevated level of reactive oxygen 89

species (ROS) in the aborted spikelets compared with wild type (Cheng et al., 2011). Several reports 90

also indicate that cuticle deficiency can cause degenerated anthers and abnormal pollen grains (Jung 91

et al., 2006; Li et al., 2010). Cuticles, the hydrophobic, waxy structures produced by the epidermal 92

cells in most land plants, play an important role in plant development and in the tolerance to 93

environmental stresses. The constituents of the cuticle include fatty acids, esters, alcohols, aldehydes, 94

ketones, and alkanes (Kunst and Samuels, 2003; Panikashvili et al., 2007). Defects in fatty acid 95

metabolism or transport can also cause anther degeneration. Mutations in rice cytochrome 96

CYP704B2, encoding an enzyme catalyzing the production of Ω-hydroxylated C16 and C18 fatty 97

acids, cause a degenerative phenotype in anther development, associated with defective anther cutin 98

biosynthesis (Li et al., 2010). OsC6 is a fatty acid transporter that affects cutin formation in rice 99

anther, and mutations in OsC6 gene also produce a degenerative anther phenotype (Zhang et al., 100

2010). 101

In plants, a complicated genetic regulatory network controls cytoskeletal organization (Hussey 102

et al., 2006). Actin filaments, one of the major components of the cytoskeleton, are composed of 103



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linear polymers that polymerize from G-actin subunits under the control of regulatory proteins 104

(Elzinga et al., 1973). The Arp2/3 complex forms a nucleation site to initiate new F-actin branches 105

and is essential for actin filament polymerization in animal, yeast, and plant cells (Vartiainen and 106

Machesky, 2004; Yanagisawa et al., 2013). During actin polymerization, SCAR/WAVE (suppressor 107

of cAMP receptor/WASP family verprolin-homologous) proteins activate the Arp2/3 complex. The 108

carboxyl regions of SCAR/WAVE proteins contain a conserved VCA domain that has three 109

subdomains, the verprolin-homology domain (V), the cofilin-homology domain (C) and the acidic 110

domain (A) (Takenawa and Miki, 2001). SCAR/WAVE proteins bind to both actin monomer and the 111

Arp2/3 complex through the VCA domain to promote actin polymerization by activating the Arp2/3 112

complex (Miki and Takenawa, 2003; Frank et al., 2004). 113

Work in plant systems has identified and characterized a number of the components of the 114

highly conserved SCAR-Arp2/3 pathway (Le et al., 2003; Mathur et al., 2003; El-Assal et al., 2004a; 115

Deeks and Hussey, 2005; Harries et al., 2005). SCAR proteins share limited amino acid sequence 116

similarity among various species, but the VCA domain shows high evolutionary conservation in 117

animals and plants (Frank et al., 2004; Basu et al., 2005; Zhang et al., 2005; Zhang et al., 2008). 118

Studies in Arabidopsis and maize showed that the VCA domain of SCARs can activate the Arp2/3 119

complex (Frank et al., 2004; Basu et al., 2005; Uhrig et al., 2007; Zhang et al., 2008). Mutations in 120

the genes encoding SCAR/WAVE complex proteins show various defects in actin arrangement and 121

trichome development (Frank and Smith, 2002; Frank et al., 2003; Mathur et al., 2003; Basu et al., 122

2005; Szymanski, 2005; Zhang et al., 2005; Uhrig et al., 2007; Zhang et al., 2008; Yanagisawa et al. 123

2015). Moreover, mutations in all of the identified Arabidopsis Arp2/3 complex subunit genes also 124

display a similar phenotype, with defects in actin organization and aberrant leaf trichome 125

morphology (Le et al., 2003; Mathur et al., 2003; El-Assall et al., 2004a; Saedler et al., 2004; 126

Kotchoni et al., 2009; Yanagisawa et al. 2015), suggesting that the Arp2/3 and SCAR/WAVE 127

complexes genetically act in a linear pathway. In addition to SCAR proteins, several proteins have 128

been identified in the Arabidopsis SCAR/WAVE complex (Deeks et al., 2004; El-Assal et al., 2004b; 129

Le et al., 2006; Jorgens et al., 2010). Among those, BRK1 is involved in the stabilization of SCAR 130

protein in vivo (Le et al., 2006), and mutations in the BRK genes result in defects of epidermal cell 131



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morphogenesis and cell division in maize (Frank et al., 2003; Frank et al., 2004). However, given the 132

critical role of the Arp2/3 complex and SCAR/WAVE in the organization of the cytoskeleton, it is 133

somewhat unexpected that none of these mutations substantially affects plant growth and 134

development in Arabidopsis and maize. 135

In this study, we report the characterization of TUTOU1 (TUT1), which encodes a functional 136

SCAR/WAVE protein and plays an important role in rice growth and development. The tut1 shows a 137

pleiotropic phenotype with severe defects in panicle development. We show that TUT1 can activate 138

the Arp2/3 complex to promote actin polymerization in vitro, and is involved in modulating actin 139

organization in vivo. We demonstrate that TUT1 plays important roles in growth and development of 140

rice plants. 141

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RESULTS 144

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Identification and characterization of the rice tut1 mutant 146

To understand the cellular and molecular mechanisms that regulate panicle development, we 147

performed a genetic screen for tutou (tut; means bald head in Chinese) mutants that have panicle 148

apical degeneration phenotype from a γ-ray-mutagenized M2 population in the japonica variety 149

Zhonghua11 background. From this screen, we identified several tut mutants and we report here a 150

detailed study of the tut1. The tut1 was backcrossed to wild-type plants (Zhonghua11) three times, 151

and progeny showing the tut1 phenotypes were collected for all studies described below, unless 152

otherwise indicated. 153

During early seedling growth, the aerial part of tut1 seedlings showed a wild-type-like 154

phenotype (Figure 1A). However, the tut1 roots were shorter than wild type roots (Figure 1A and 1B). 155

Compared with wild type plants, the mature tut1 plants were shorter and had fewer tillers (Figure 156

1C-1E). During all growth stages, the tut1 leaves were smaller than wild type leaves, and showed 157

degeneration that initiated from the leaf tips, accompanying by a reduction of the chlorophyll levels 158

(Figure 1F-1H). Young tut1 leaves usually were pale white and then turned green upon maturation 159

(Figure 1F). Consistent with the observation of degenerated leaf tips in tut1, massive cell death was 160

detected by staining with Evans blue along the leaf tips of tut1, but rarely detectable in wild type 161

leaves (Figure 1I). Moreover, tut1 leaves had higher levels of superoxide than that in wild type 162

(Figure 1J), suggesting that degenerative cell death in tut1 leaves was related to the production of 163

ROS. 164

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Degenerated panicles in tut1 166

The tut1 showed a delayed heading phenotype, approximately 15 days later than the wild type plants 167

(Figure 1C). The tut1 panicles were significantly shorter than wild-type panicles, and showed severe 168

degeneration of spikelets in the apical portion of the panicle (Figures 2A). In the tut1 panicles, the 169

degenerating spikelets occurred in the apical parts of all branches, including the primary and 170

secondary branches (Figures 2B and Supplemental Figures 1A, 1D and 1E). Similar to that in leaves, 171



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we also observed significantly increased ROS levels in the degenerated tut1 spikelets compared to 172

that in wild type (Figure 2C and 2D). The tut1 had smaller flower filaments than the wild type, and 173

the tut1 anthers were usually pale white and showed various degrees of degeneration (Figures 2E; 174



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Supplemental Figure 2). 175

To analyze the cellular basis of panicle degeneration in tut1, we examined panicle development 176

of wild type and tut1 by scanning electron microscopy (SEM) over the nine stages (In1 through In9) 177



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of rice panicle development (the panicle development stages were followed as defined by Ikeda et al., 178

2004). In the early stages, the spikelets at the tip of wild type and tut1 panicles showed no significant 179

difference, and spikelets at In7 stage were shown in Figure 2F. At stage In8, the difference between 180

wild type and tut1 became apparent, as the panicle apical spikelet of tut1 was smaller than wild type 181

(Figure 2G). At stages In8-In9, the tut1 spikelets were misshapen and disorganized compared with 182

wild type (Figure 2H). A closer examination revealed that the surface of wild type spikelets was well 183

organized and smooth, whereas the surface of the tut1 spikelet contained shrunken and disorganized 184

cell files (Figure 2I). During later developmental stages, the degenerative phenotype of tut1 panicles 185

became more severe, resulting in the formation of small, aborted spikelets that lacked recognizable 186

cell files on the surface of the tut1 spikelets (Figures 2J and 2K). These results suggested that TUT1 187

plays an important role in panicle development, and is likely involved in cell expansion. 188

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Defective anther and pollen development in tut1 190

Anthers in tut1 also showed a degenerative phenotype, as mutant anthers lacked organized cell files 191

on the epidermis and abortive or shrunk pollen grains (Figures 2E, 3A through 3C). We analyzed 192

pollen development in tut1 in comparison with that of wild type by light microscopy. The developing 193

tut1 pollen showed no abnormalities at the microspore mother cell stage (Figure 3D). However, after 194

entering the vacuolated pollen stage, the tut1 microspores became misshapen and disorganized 195

(Figure 3E), and eventually aborted (Figures 3F through 3H). Notably, the degeneration of the 196

epidermal cell layer usually occurred after the mature pollen stage in wild type, whereas the 197

premature degeneration of the epidermal cells in tut1 anthers was observed at the vacuolated pollen 198

stage (Figure 3E). Moreover, the tut1 pollen sac also had an irregular shape compared with wild type 199

at the first pollen mitosis stage (Figures 3F and 3G). At the mature pollen stage, tut1 had 200

significantly fewer mature pollen grains, compared with wild type (Figure 3H). Due to these defects, 201

tut1 showed poor fertility and produced smaller seeds (Figure 3I and Supplemental Figures 1B and 202

1C). 203

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Defective cuticle formation in tut1 205



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The epidermis of wild type anthers was coated with epicuticular wax crystals, but these wax crystals 206

were substantially fewer and disorganized on the surface of the tut1 anthers (Figures 3A and 3B). On 207

the tut1 anthers, the formation of wax cuticle was reduced in the basal panicle and completely absent 208



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in the apical panicle (Figures 4A-4C). Consistent with this, transmission electron microscopy (TEM) 209

revealed that tut1 cuticle had significantly fewer wax structures compared with wild type (Figure 4D). 210

Similarly, the tut1 mutation also caused abnormalities in pollen exine development (Figure 4E) and 211



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formation of cuticle on the surface of leaves and leaf stomata (Figure 4F; Supplemental Figures 3A 212

and 3B). These results suggested that TUT1 is involved in cuticle formation, which may explain the 213

defective development of anthers and pollen in tut1. 214

We reasoned that the defective development of wax cuticles might be related to abnormal 215

metabolism of fatty acids. To test this possibility, we analyzed the fatty acid levels in heading 216

panicles by gas chromatography. We found that the level of palmitic acid (C16:0), eicosenoic acid 217

(C20:1), arachidonic acid (C20:4), and docosanoic acid (C22:0) were reduced in tut1 panicles 218

compared with wild type (Figure 4G), suggesting that altered fatty acid metabolism may be related to 219

the tut1 phenotype. 220

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Molecular cloning of TUT1 222

In a cross between wild type (Zhonghua11) and tut1, all F1 progenies displayed a wild-type 223

phenotype. In the self-pollinated F2 population, the wild-type and tut1 mutant plants segregated in a 224

ratio of approximately 3:1 (wild type: tut1 = 203: 54; χ2 = 2.17, P < 0.05), indicating that tut1 is a 225

recessive mutation in a single nuclear gene. To genetically map TUT1, we generated an F2 226

population (~10,000 plants) derived from the cross between tut1 and Zhenshan97 (an indica cultivar). 227

TUT1 was roughly mapped on the short arm of Chromosome 1, between the simple sequence repeat 228

(SSR) markers RM151 and RM272 (Figure 5A). Using 2,600 F2- and F3-derived tut1 segregants, we 229

mapped TUT1 to an interval of 196 kb (kilo base pairs) between the SNP (single nucleotide 230

polymorphism) markers P18 and P22 (Figure 5A). This region contained 28 annotated genes and 231

DNA sequencing analysis of tut1 revealed a C-to-G transition at nucleotide 1,556 of the coding 232

sequence of LOC_OS01g11040. The mutation occurred in exon 6, which converted a Ser into a 233

premature stop codon (TCA to TGA), resulting in the formation of a truncated polypeptide lacking 234

815 amino acid residues of the carboxyl region (Figures 5A and 5B). Reverse transcription 235

quantitative PCR (RT-qPCR) revealed that the transcript levels of the putative TUT1 gene were 236

slightly lower in tut1 than in wild type (Figure 5E). Also, the tut1 mutation occurred in the highly 237

conserved envelope glycoprotein C (EGC) domain and also caused the loss of the VCA domain at 238

the carboxyl region (Figure 5B), which is functionally important for the SCAR/WAVE proteins 239



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(Machesky et al., 1999; Basu et al., 2005; Zhang et al., 2008). Therefore, tut1 is likely a null mutant 240

allele. 241

To verify the identity of the putative TUT1 gene, we conducted a complementation assay. To 242



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this end, we transformed a full length wild-type cDNA fragment of LOC_Os01g11040 under the 243

control of the maize ubiquitin (Ubi) promoter into the TUT1/tut1 heterozygous plants. The transgene 244

fully rescued the tut1 phenotype, including the degenerated panicle phenotype (Figures 5C and 5D). 245

Moreover, the transgenic plants carrying an RNA interference (RNAi) transgene had a reduced 246

expression level of LOC_Os01g11040 (Supplemental Figure 4A), and showed tut1-like phenotypes, 247

including reduced plant height, degenerated panicles, cuticle defects, blunt trichomes, and increased 248

ROS levels (Figures 5F-5H, Supplemental Figures 4B-4F). These results demonstrate that 249

LOC_Os01g11040 represents the TUT1 gene. 250

251

Analysis of TUT1 expression 252

To investigate the expression pattern of TUT1 in rice, we used RT-qPCR to measure the TUT1 253

mRNA levels in various organs or tissues of wild type plants, including root, node, stem, immature 254

leaf, mature leaf, immature panicle, and mature panicle (Figure 6A). Consistent with the tut1 255

phenotype, the level of TUT1 mRNA was highest in the panicles, followed by leaves. We also 256

detected TUT1 expression in roots, stems, and nodes, but at lower levels compared to that in panicles 257

and leaves (Figure 6A). 258

To analyze the spatial expression pattern of TUT1, we generated transgenic plants harboring a 259

TUT1:GUS (β-glucuronidase) reporter gene. Five independent TUT1:GUS transgenic lines were 260

analyzed and gave similar results. We found GUS activity in mature glume, seed germ, root, 261

coleoptile, node, and leaf (Figure 6B-6E), consistent with the tut1 phenotype of short roots, small 262

seeds, dwarf plants, and degenerating leaves. Close examination of TUT1:GUS expression in the 263

panicle revealed TUT1 expression in glumes, anthers, and mature pollen grains (Figure 6F-6I). In the 264

anther, TUT1:GUS expression mainly occurred in the epidermis and endothecium, with a lower level 265

in the tapetum and microspore (Figure 6H). TUT1:GUS was also highly expressed in mature pollen 266

grains (Figure 6I). These results suggest that the expression pattern of TUT1 is consistent with its 267

important role in panicle development, especially in anther development. 268

269

TUT1 encodes a functional SCAR/WAVE protein 270



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Sequence alignment suggested that TUT1 encodes a SCAR/WAVE-like protein (Supplemental Figure 271

5A and 5B). The classic SCAR/WAVE family proteins contain a SCAR/WAVE homolog domain 272

(SHD), a short stretch of basic amino acids (B) in the N-terminal region, and a VCA domain in the 273



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C-terminal region (Figure 7A). A sequence comparison indicated that TUT1 contains all functionally 274

important domains found in eukaryotic SCAR/WAVE proteins (Figure 7A). TUT1 exists as a single 275

copy gene in the rice genome. In a previous study, TUT1 was also named OsSCAR1 (Zhang et al., 276



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2008). Three additional rice genes encode proteins containing the SCAR homolog domain, annotated 277

as OsSCAR2, OsSCAR3, and OsSCAR4, of which OsSCAR2 lacks the putative VCA domain (Figure 278

7A and Supplemental Figure 5B). TUT1/OsSCAR1 shared the highest sequence similarity with 279

SCAR-like proteins from Brachypodium distachyon (68% identity), Sorghum bicolor (63% identity), 280

Setaria italica (62% identity) and Aegilops tauschii (62% identity), followed by those from Sorghum 281

bicolor and maize (Zea mays), Arabidopsis, and other plant species (Figure 7B, Supplemental Figure 282

5A and 5B). Arabidopsis has one SCAR-like protein (AtSCAR5/SCARL) and four SCAR proteins 283

(AtSCAR1-4) (Zhang et al., 2008), of which AtSCAR1 and AtSCAR3 show higher sequence 284

similarity to TUT1 than do AtSCAR2 and AtSCAR4 (Figure 7B and Supplemental Figure 6A and 285

6B). 286

Mutations in the SCAR/WAVE complex genes of Arabidopsis (atscar2) and maize (brk1) cause 287

defective trichome development (Frank and Smith, 2002; Frank et al., 2003; Basu et al., 2005; Zhang 288

et al., 2005). In the tut1 mutant, development of trichomes was also impaired, characterized by 289

blunted tips of the trichomes (Figures 7C and 7D), suggesting that TUT1 plays an evolutionarily 290

conserved role in trichome development. Moreover, similar to that in the maize brk mutants (Frank et 291

al., 2003), the cell wall at the junction between two epidermal cell layers was affected in tut1. The 292

tut1 cells were more rounded compared with wild type (Supplemental Figure 7A and 7B). 293

The VCA domain of SCAR/WAVE proteins acts to activate the Arp2/3 complex, and 294

consequently increases the actin filament nucleation and branching activities of the complex (Stradal 295

et al., 2004). Sequence alignment analysis showed that the carboxyl-terminal region of TUT1 protein 296

contains a conserved WH2 region and acidic region (Supplemental Figure 5B). In an actin 297

polymerization assay, the VCA domain of TUT1 significantly increased actin filament assembly in 298

the presence of bovine Arp2/3 (Supplemental Figure 8), indicating that the TUT1-VCA domain is 299

capable of activating Arp2/3. Taken together, these results indicate that TUT1 is a functional 300

SCAR/WAVE protein. 301

302

Impaired actin filament organization in tut1 303

The data presented above indicate that TUT1 can activate Arp2/3. The SCAR/WAVE-activated 304



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Arp2/3 complex is involved in the regulation of the formation of loosely aligned actin bundles at the 305

earliest stages of trichome branch elongation (El-Assall et al., 2004b). To assess the possible 306

regulatory role of TUT1, we examined the organization of F-actin in the trichomes of wild-type and 307



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tut1 leaves by Alexa Fluor 488-phalloidin staining. In the trichomes of wild-type leaves, the F-actin 308

filaments were well organized along the longitudinal axis (Figure 8A). However, in tut1 leaves, we 309

observed significantly fragmented F-actin filaments (Figure 8B), consistent with the altered shape of 310

these cells in tut1 (see Figures 7C and 7D). Statistical analysis showed that the mean length of actin 311

filaments in tut1 trichomes was significantly shorter compared with wild type (Figure 8C). The 312

number of actin filaments increased in tut1 trichomes (Figure 8D). Our results suggest that TUT1 is a 313

SCAR/WAVE protein and is involved in F-actin organization in rice. 314

315

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DISCUSSION 317

318

In this study, we functionally characterized rice TUT1 and presented multiple lines of evidence 319

showing that TUT1 encodes a functional SCAR/WAVE protein that plays an important role in 320

regulating F-actin organization. We also demonstrate that TUT1 functioning in multiple aspects of 321

rice growth and development, especially in panicle development. 322

Our present work shows that TUT1, a SCAR/WAVE-like protein in rice, can activate the Arp2/3 323

complex to modulate actin polymerization. The SCAR/WAVE complex plays an important role in 324

regulating actin filament arrangement by activating the Arp2/3 complex to modify actin filament 325

organization (Blanchoin et al., 2000; Frank et al., 2004; Beltzner and Pollard, 2008). Mutations in 326

key components of the SCAR/WAVE-Arp2/3 complex result in various defects in yeast and animals 327

(Winter et al., 1999; Di Nardo et al., 2005). In many cases, these mutations cause embryonic arrest 328

and zygotic lethality by impairing various cellular activities (Severson et al., 2002; Sawa et al., 2003; 329

Vartiainen and Machesky, 2004). However, in Arabidopsis and maize, mutations in the SCAR/WAVE 330

complex genes cause relatively weak phenotypes, affecting cell morphogenesis and development of 331

the trichome and the hypocotyl (Frank and Smith, 2002; Uhrig et al., 2007). Notably, the rice tut1 332

mutant shows a pleiotropic phenotype during its entire life cycle, including shorter roots, reduced 333

plant height, degenerating leaves, and degenerating panicles, suggesting that TUT1 functions as a 334

housekeeping gene during rice growth and development. A recent study reveals that EARLY 335

SENESCENCE 1 (ES1), identical to TUT1, plays an important role in regulating water loss (Rao et 336

al., 2015), suggesting that TUT1/ES1 is involved in various developmental and physiological 337

processes. 338

The most severe defects in tut1 occur in panicles, consistent with the observation that the panicle 339

has the highest level of TUT1 transcripts among the examined organs and tissues. In the panicle, we 340

detected strong TUT1-GUS expression in glumes and anthers but not in the pistil. Consistent with the 341

expression pattern of TUT1, whereas development of glumes, anthers, and pollen grains is severely 342

affected in tut1, resulting in partial male sterility, the pistils and carpels remain relatively normal. 343

Since TUT1 encodes a SCAR/WAVE-like protein, we presume that TUT1 is involved in panicle 344



24

development by regulating F-actin organization. The biochemical analysis shows that the activation 345

of TUT1-VCA domain is capable of activating the Arp2/3 complex in vitro. We noticed that the 346

TUT1-VCA polypeptide shows a lower activity compared with SCAR/WAVE proteins of 347

Arabidopsis and maize. We speculate that the reduced activity might be caused by the assay 348

condition, which may not be optimized for the rice TUT1-VCA polypeptide, or by improper folding 349

of TUT1-VCA recombinant polypeptide. 350

Panicle and spikelet development is a controlled by multiple genes that function in different 351

cellular processes, such as cuticle formation and cell differentiation (Komatsu et al., 2001; Ikeda et 352

al., 2004; Zheng et al., 2005; Panikashvili et al., 2009). Cuticle plays an important role in rice flower 353

morphogenesis, anther and pollen development, pathogen resistance, and salt tolerance (Bird et al., 354

2007; Qin et al., 2011; Qin et al., 2013). However, our understanding of cuticle function in panicle 355

development remains limited. Considering that the panicle apical degeneration phenotype of tut1 is 356

closely associated with cuticle disorganization and deficiency, it is reasonable to presume that the 357

formation of the cuticle affects panicle development in rice. The complex regulation of cuticle 358

formation includes vesicle trafficking, fatty acid metabolism, and transport processes (Yeats and 359

Rose, 2013). Vesicle trafficking that can be regulated by F-actin is important for wax secretion from 360

epidermal cells in Arabidopsis (McFarlane et al., 2014). Consistently, we found that fatty acid 361

metabolism is impaired in tut1, suggesting that abnormal fatty acid metabolism may cause the 362

deficient cuticle formation with affecting F-actin regulated vesicle trafficking in tut1. A recent study 363

revealed that Arp2/3 is involved in modulating the cell wall thickness gradient from tip to base of the 364

trichome branch by regulating F-actin meshwork formation (Yanagisawa et al., 2015). Consistently, 365

we also observed the abnormal arrangement of F-actin in the tut1 mutant, suggesting that the 366

Arp2/3-SCAR/WAVE complex plays a conserved role in leaf trichome development in plants. 367

Compared with mutants in Arabidopsis SCAR/WAVE proteins, the rice tut1 mutants show multiple 368

phenotypes, especially in panicle development. Therefore, the diverse functions of SCAR/WAVE 369

family proteins need to be studied in different varieties of plants, to provide a comprehensive 370

understanding of SCAR/WAVE functions. 371

Additionally, one of the important reasons caused rice yield reduction is defective panicle 372



25

development (Xing and Zhang, 2010). Understanding the mechanism of panicle development should 373

be helpful to mitigate the crop yield reduction caused by defective panicle development. However, 374

agricultural applications to avoid panicle degeneration remain relatively limited and additional 375

research will be required to identify the molecular mechanisms that can be modified to confer fully 376

developed panicles in rice and to understand the complicated regulation of panicle apical spikelet 377

development, including the effects of temperature, nutrition, and genetic background (Cheng et al., 378

2011). The identification and characterization of genes that function in rice panicle development and 379

production, such as TUT1, provide the first steps toward genetic manipulation to engineer or breed 380

high-production rice. 381

382

383



26

MATERIALS AND METHODS 384

385

Plant Materials and Growth Conditions 386

The tut1 was screened from γ-ray-induced M2 population in the japonica cultivar (Oryza sativa L. 387

subsp.) Zhonghua11. Plants were grown in the experimental field of Beijing (39° 54′ 20″ N 116° 23′ 388

29″ E) from May to January under long-day conditions and in Lingshui (18° 30′ 00″ N 110° 01′ 20″ 389

E) from January to May under short-day conditions. When grown under the laboratory condition, 390

seeds of wild type and tut1 were germinated in darkness at 30°C for 3 days and then transferred to a 391

SANYO culture chamber with 16/8 hrs photoperiod at 30°C. 392

393

Histochemical Analysis 394

For light microscopy, samples were fixed at 4°C for 12 hrs in 3.7% (v/v) glutaraldehyde in 0.025 M 395

phosphate buffer (pH 6.8) and dehydrated with ethanol. The samples were embedded overnight at 396

room temperature in embedding medium (Leica Historesin) and then sectioned at 4 μm thickness by 397

using a Leica RM2265 rotary microtome. The sections were then stained with 0.05% (w/v) toluidine 398

blue for 1 min and observed with a light microscope (OLYMPUS BX51). To measure chlorophyll 399

contents, chlorophyll was extracted from fresh leaves of wild type and tut1 plants and analyzed by 400

using a spectrophotometer (BECKMAN COULTER DU800) as described (Lichtenthaler, 1987). For 401

cell death staining, samples were treated with 1% (w/v) Evans blue for 6 hrs and then treating with 402

ethanol for observation. For ROS staining, samples were treated with 0.05% (w/v) tetranitroblue 403

tetrazolium chloride (NBT) at 28°C overnight and then treating with ethanol for observation. 404

405

Gene Expression Analysis 406

For RT-qPCR analysis, total RNA was extracted from root, node, stem, immature leaves, mature 407

leaves, immature panicle, and mature panicle using the RNAprep pure Plant RNA Purification Kit 408

(Tiangen Biotech). Total RNA (2 μg) was reverse transcribed into cDNA and then used as template 409

for RT-qPCR analysis with specific primers (TUT1_ RT-qPCR _F and TUT1_ RT-qPCR _R; see 410

Supplemental Table 1). The RT-qPCR was performed using the UltraSYBR mixture (CWBIO) 411



27

according to the manufacturer’s instructions, and the reactions were run in a CFX96TM real-time 412

PCR system (Bio-Rad). The rice GAPDH gene was used as an internal control with 40 PCR cycles 413

using primers GAPDH_F and GAPDH_R (see Supplemental Table 1). 414

415

TUT1:GUS Construct and Histological GUS Assay 416

The 785-bp fragment immediately upstream of the start codon of TUT1 was amplified using the 417

gene-specific primers GUS_F and GUS_R and inserted in front of the GUS gene in pCAMBIA1300 418

with PstI and BamHI (see Supplemental Table 1). The final construct containing the TUT1:GUS 419

reporter gene was introduced into the wild-type background plant by Agrobacterium 420

tumefaciens-mediated transformation(Hiei et al., 1994). Transgenic plants were selected on 421

hygromycin medium and T1 transgenic plants were used for analysis of GUS activity. 422

GUS activity was detected using a standard protocol (Jefferson et al., 1987). Transgenic plant 423

tissues were incubated in X-gluc buffer overnight at 37°C. The samples were cleared of chlorophyll 424

by dehydration with ethanol. The stained samples were photographed using a Cannon digital camera 425

and stereoscope (OLYMPUS SZX12). The stained stamen was sliced using resin sections (Leica 426

Historesin) and analyzed by light microscopy (OLYMPUS BX51). 427

428

Scanning and Transmission Electron Microscopy 429

For scanning electron microscopy, samples were fixed overnight at 4°C with 2.5% (v/v) 430

glutaraldehyde in 0.1 M phosphate buffer (pH 7.4). The samples were dehydrated in a graded ethanol 431

series and substitution with a graded isoamyl acetate series. The samples were critical-point dried, 432

sputter coated with gold, and observed using a HITACHI S-3000N scanning electron microscope at 433

10 kV. For transmission electron microscopy, samples were fixed overnight at 4°C with 2.5% (v/v) 434

glutaraldehyde in 0.1 M phosphate buffer (pH 7.4). After three washes with phosphate buffer, 435

samples were fixed overnight in 2% (v/v) OsO4 in phosphate buffer. After being stained and 436

dehydrated in a graded ethanol series, the samples were submerged in LR White resin and 437

polymerized for 2 days. Ultrathin cross-sections were prepared using a Leica EM UC6 438

ultra-microtome. Ultrathin sections were observed under a Tecnai Spirit (120 kV) transmission 439



28

electron microscope. 440

441

Map-Based Cloning of TUT1 442

The tut1 was crossed with Zhenshan 97 (indica) to generate the F2 mapping population. TUT1 was 443

mapped to a 196-kb region on Chromosome 1 using simple sequence repeat (SSR) markers and SNP 444

(single nucleotide polymorphism) markers (see Supplemental Table 1). Exons of candidate genes in 445

this region were amplified from tut1 and wild-type plants. The sequences were compared using 446

DNASTAR software and the NCBI website. The full-length ORF of TUT1 was amplified using 447

reverse-transcribed DNA as template. The primers used were TUT1_OE_F and TUT1_OE_R (see 448

Supplemental Table 1). The full-length ORF was then cloned into the SK vector using the SalI and 449

SmaI restriction sites and then cloned into the binary vector pTCK303 using the KpnI and SpeI 450

restriction sites (see Supplemental Table 1) (Wang et al., 2004). Agrobacterium tumefaciens–451

mediated transformation was used to transform the resulting construct, pTCK303-TUT1, into a 452

compound callus pool containing wild-type, heterozygous, and homozygous callus generated from 453

TUT1/tut1 heterozygous seeds. More than 2 independent transgenic lines were produced that 454

complemented the tut1 phenotypes. A specific DNA fragment of TUT1 was amplified from the 455

Zhonghua11 cDNA using the PCR primers RNAi_F (KpnI and SpeI) and RNAi_R (BamHI and SacI) 456

(see Supplemental Table 1). The PCR products were then digested with SpeI/SacI and KpnI/BamHI, 457

respectively. The two digested fragments were sequentially cloned into the vector pTCK303, which 458

contains a maize (Zea mays) ubiquitin promoter, to make the final pTCK303:TUT1 RNAi plasmid as 459

previously described (Wang et al., 2004). This final construct was introduced into the wild type 460

background plant by Agrobacterium tumefaciens-mediated transformation (Hiei et al., 1994). 461

462

Fatty Acid Content Analysis 463

For fatty acid analysis, 50 mg of tissue was ground in liquid nitrogen and then added to 1.5 mL 464

extraction buffer (2.5% sulfuric acid in methanol, v/v) and incubated at 80°C for 2 hrs. After the 465

sample cooled to room temperature, 2 mL 0.9% (w/v) NaCl and 1 mL hexyl hydride were added to 466

the extraction buffer and the mixture was vortexed. The mixture was centrifuged at 4,000 rpm for 10 467



29

min at room temperature and the supernatant was transferred to a new microfuge tube then dried to 468

powder with a vacuum pump. The powder was dissolved in 50 μL isoamyl acetate for analysis. 469

470

Fluorescence Microscopy of Actin Filaments 471

F-actin was stained using the previously described method (Olyslaegers and Verbelen, 1998; Zhang 472

et al., 2011). Mature rice leaves were incubated in PEM buffer (100 mM PIPES, 10 mM EGTA, 0.3 473

M mannitol, 5 mM MgSO4, pH 6.9) and 0.66 mM Alexa Fluor 488-phalloidin [Life Technologies]. 474

After overnight incubation at 4°C, samples were observed in 50% glycerol with a confocal 475

microscope (OLYMPUS FLUOVIEW FV1000). 476

477

Accession Numbers 478

Sequence data from this article can be found in the GenBank/EMBL databases and the Gramene 479

website under the following accession numbers: TUT1/OsSCAR1 (NP_001172227.1), OsSCAR2 480

(NP_001049845), OsSCAR3 (Q84TX2), OsSCAR4 (BAC75554), AtSCAR1 (NP_001031474), 481

AtSCAR2 (NP_181378), AtSCAR3 (NP_174212), AtSCAR4 (NP_195793), AtSCAR5 (NP_193595), 482

ZmSCAR1 (AFW80370), ZmSCAR2 (DAA44917), ZmSCAR3 (GRMZM2G475139), ZmSCAR4 483

(ACN28010), ZmSCAR5 (DAA52161), ZmSCAR6 (AFW88660), AetSCAR (EMT14604), 484

BdSCAR (XP_003565325), SiSCAR (XP_004968593), SbSCAR (Sb03g002220), CsSCAR 485

(XP_004149068), GmSCAR (XP_006585168), SlSCAR (XP_004233023), StSCAR 486

(XP_006362414), HsWAVE1 (AAD33052), MmWAVE1 (AAL75574), CeWVE-1 (NP_493028), 487

DmSCAR (NP_609477), and DdSCAR1 (AAD29083). 488

489

Supplemental Data 490

The following materials are available. 491

Supplemental Figure 1. Quantitation of agronomic traits in wild type and tut1. 492

Supplemental Figure 2. Quantitation of anther degeneration in wild type and tut1. 493

Supplemental Figure 3. SEM and TEM examination of cuticle in wild type and tut1 leaves. 494

Supplemental Figure 4. The phenotypes of TUT1 RNAi transgenic lines. 495



30

Supplemental Figure 5. Amino acid sequence alignment of TUT1 and other WAVE/SCAR family 496

proteins. 497

Supplemental Figure 6. Phylogenetic tree of SHD of SCAR/WAVE proteins from Arabidopsis and 498

rice. 499

Supplemental Figure 7. Analysis of mature leaves of wild type and tut1 by light microscopy. 500

Supplemental Figure 8. Activation analysis of TUT1-VCA domain in vitro. 501

Table S1. Primers used in this study. 502

503

AUTHORS’ CONTRIBUTION 504

J.B., X.Z., F.C., Y.F., Q.Q. and J. Zuo designed the experiment and analyzed the data. J.B. and Q.W. 505

performed most of the experiments with assistance of J. Zhang, H.Q., G.D., L.Z., H.K., Q.X. and J.N. 506

J.B. and J. Zuo wrote the manuscript. 507

508

ACKNOWLEDGEMENTS 509

510

We thank Dr. Kang Chong (Institute of Botany, Chinese Academy of Sciences) for providing the 511

pTCK303 vector and Shufeng Sun (Institute of Biophysics, Chinese Academy of Sciences) for 512

technical assistance in transmission electron microscopy. 513

514

Figure Legends 515

516

Figure 1. Phenotypes of tut1. 517

A, Seedling phenotype of wild type (Zhonghua11, left 3 seedlings) and tut1 (right 3 seedlings). Seeds 518

were germinated at 30°C for 3 days and then transferred to a culture chamber with 16/8 hr 519

photoperiod at 30°C. Bar = 1 cm. B, Statistical analysis of root length in wild type and tut1 as shown 520

in (A). The data presented are mean values of two independent experiments (n > 45). Error bars 521

indicate SD; *** indicates P < 0.01 (Student’s t -test). C, Phenotypes of wild type (left) and tut1 522

(right) mutants at the grain-filling stage. Bar = 5 cm. D and E, Statistical analysis of tiller number 523

and plant height at maturity of wild type and tut1. The data presented are mean values (n > 50). Error 524



31

bars indicate SD; *** indicates P < 0.01 (Student’s t -test). F, Leaves of wild type (leaf 4 leaves) and 525

tut1 (right 4 leaves) at different growth stages. Bar = 5 cm. G, Leaves of wild type (left) and tut1 526

(right) at 10 days after leafing. Bar = 2 cm. H, Analysis of chlorophyll content of wild type and tut1 527

leaves at different times after leafing. Blue line indicates wild type, red line indicates tut1. Leaves at 528

4, 8, 12, 16, and 20 days after leafing were used in the analysis. Fresh leaves were used for 529

measurement of chlorophyll contents. Two technical repeats were made in the test. Error bars 530

indicate SD. I, Cell death analysis of wild type (left) and tut1 (right) seedling leaves by using Evans 531

blue staining. Bar = 0.5 cm. J, ROS observation of wild-type (left) and tut1 (right) mature leaves 532

using NBT staining. Bar = 2 cm. 533

534

Figure 2. Morphology of panicle and spikelet in tut1. 535

A, Panicle rachis of wild-type (left) and tut1 (right) plants at the grain-filling stage. Bar = 2 cm. B, 536

The expanding panicle branch of wild-type (left) and tut1 (right) plants at the grain-filling stage. The 537

part of the tut1 panicle shown in the white box was enlarged. The arrows show the degenerated 538

branches and spikelets in the tut1. Bars = 2 cm. C, ROS analysis of wild type (bottom) and tut1 (top) 539

by NBT staining. Bar = 2 mm. D, ROS analysis of spikelets showing different degrees of 540

degeneration in tut1 at the heading stage, by NBT staining. Bar = 2 mm. E, Comparison of wild type 541

(left one) and tut1 spikelets (right three) after removing the lemma and palea at heading stage. Right 542

three spikelets, named Type 1, Type 2 and Type 3, show varying degrees of developmental defects in 543

tut1. Bar = 1 mm. F to K, SEM observation of spikelet at the tip of the panicle at stage In7 (F), stage 544

In8, arrows indicate the apical spikelets in wild type and the tut1 (G), stage In8-In9 (H), the white 545

square frame region shown in (H) is enlarged in (I), stage In9 (J), the white square frame region 546

shown in (J) is enlarged in (K). Bars = 100 μm (F and K), 400 μm (G), 200 μm (H), 10 μm (I), and 547

500 μm (K). 548

549

Figure 3. Scanning electron microscopy and semithin section analysis of wild-type and tut1 anther 550

morphology. 551

A to C, SEM observation of mature anthers of wild type and the tut1 (A), enlarged view of the anther 552

surface of wild type and the tut1 (B), mature pollen grain of wild type and the tut1 (C). Bars = 200 553



32

μm (A), 20 μm (B), and 10 μm (C). D to H, The cross sections were stained with Toluidine blue. Ep, 554

epidermis; En, endothecium; ML, middle layer; Ms, microsporocyte; Msp, microspore; MP, mature 555

pollen; T, tapetum. Cross-section of single locule at the microspore mother cell stage, Ep, En, ML, T, 556

and Ms are indicated (D); vacuolated pollen stage, Ep, En, T, and Msp are indicated (E); Pollen first 557

mitosis stage, Ep, En, T, and Msp are indicated (F); Pollen second mitosis stage, Ep and Mp are 558

indicated (G); Mature pollen stage, Ep, and Mp are indicated (H). The arrows show the degenerated 559

epidermal cell. I, Phenotype of mature seeds without glumes of wild type (top) and tut1 (bottom). 560

Bar = 5 mm. 561

562

Figure 4. Cuticle formation and fatty acid metabolism in wild type and tut1. 563

A to C, SEM observation of mature anther cutin of wild type (A) and the tut1 at middle panicle (B) 564

and apical panicle (C). tut1 anthers showing different degrees of degeneration are shown in (B) and 565

(C). Bars = 10 μm. D, TEM observation of wild type and tut1 mature anthers. Arrowhead indicated 566

cuticle. Bar = 5 μm. E and F, SEM observation of the surface of mature pollen grains (E) and the 567

cuticle of mature leaves (F) of wild type and the tut1. Bars = 2 μm (E) and 5 μm (F). G, Analysis of 568

fatty acid contents in wild type and tut1, using gas chromatography. Fresh panicles at heading stage 569

of wild type and tut1 were used in the measurement. C17 was added as an internal standard. Seven 570

types of fatty acid, C16:0, C18:1, C18:2, C18:3, C20:1, C20:4 and C22:0 were measured and three 571

biological repeats were conducted. Error bars indicate SD. ** indicates P < 0.05, *** indicates P < 572

0.01 (Student’s t -test). 573

574

Figure 5. Map-based cloning of TUT1. 575

A, TUT1 maps on chromosome 1. TUT1 was roughly mapped to a 440 kb interval between the SSR 576

markers RM151 and RM272. The number of recombinants between the molecular markers and 577

TUT1 are indicated. TUT1 was fine-mapped to an interval of about 196 kb between the SNP markers 578

P18 and P22. This region contained 28 predicted genes. A one-nucleotide transversion mutation was 579

found in a gene annotated as a SCAR-like protein 2 (LOC_Os01g11040). The mutation site at 580

nucleotide 1556 (C to G) in the sixth exon is indicated by the asterisk. B, Conserved domain 581

organization of TUT1, showing the SCAR/WAVE homolog domain (SHD), Basic domain (B), 582



33

envelope glycoprotein C (EGC) domain, pantetheine attachment site (PPAS), and VCA domain as 583

black rectangles. The mutation alters the 519th codon (serine) to a termination codon (shown as a 584

white dot in the EGC domain). The asterisk indicates the termination codon. C, Phenotypes of wild 585

type (WT), complemented transgenic line (Com), and tut1 (tut1) at heading stage. Bar = 10 cm. D, 586

Mature panicle phenotypes of wild type (WT), complemented line (Com) and tut1 (tut1). Bar = 2 cm. 587

E, TUT1 expression analysis of wild type and tut1 by reverse transcription quantitative PCR 588

(RT-qPCR). F, Morphology of wild type, tut1, and RNAi transgenic lines (R1 and R2) at heading 589

stage. Bar = 20 cm. G, Panicles of wild type, tut1 and RNAi transgenic lines (R1 and R2) at 590

flowering stage. Bar = 3 cm. H, Anther cuticle of wild type, tut1 and representative complemented 591

line (Com). Bars = 20 cm (F), 3 cm (G), and 5 μm (H). 592

593

Figure 6. Spatial expression pattern of TUT1. 594

A, TUT1 mRNA levels was measured by RT-qPCR. Total RNA was extracted from the root (R), node 595

(N), stem (S), immature leaf (IL), mature leaf (ML), immature panicle (IP), and mature panicle (MP) 596

of the wild type. B to I, Expression of TUT1, as revealed by promoter-GUS fusion analysis in 597

transgenic plants. GUS activity was observed in the mature glume, seed, and seed germ (B), root, 598

coleoptile and seedling (C), basal node of shoot (D), leaf (E), glume at heading stage (F), anther 599

filaments, and anthers. The arrow indicates the pistil (G), GUS staining of a cross-section of the 600

anther, with Ep, En, T, and Msp indicated (H) and mature pollen (I). Bars = 2 mm (B, C, D, and E), 1 601

mm (F and G), and 30 μm (H and I). 602

603

Figure 7. TUT1 belongs to the SCAR/WAVE family. 604

A, Domain structure of SCAR/WAVE proteins. The rice genome encodes four SCAR proteins, 605

TUT1/OsSCAR1, OsSCAR2, OsSCAR3, and OsSCAR4. All four have SHD and basic domains, but 606

OsSCAR2 lacks the VCA domain. Percentages above each domain indicate the amino acid sequence 607

identities of the SHD and VCA domains in each protein, compared with TUT1. The domains are 608

color-coded: SHD (blue), basic domain (red), VCA domain (green), and the black line represents the 609

amino acid sequence. B, A Phylogenetic tree of TUT1 and its homologous proteins in different 610

species. The phylogenetic tree was constructed using DNAMAN software with 1000 bootstrapping 611



34

trials. Bootstrapping values over 50% and the scale bar are shown in the phylogenetic tree. The tree 612

was constructed using the distance method with maximum likelihood. TUT1 is shown in red. 613

Accession numbers for each sequence are shown. Species are as indicated: Aegilops tauschii (Aet), 614

Arabidopsis thaliana (At), Brachypodium distachyon (Bd), Caenorhabditis elegans (Ce), Cucumis 615

sativus (Cs), Dictyostelium discoideum (Dd), Drosophila melanogaster (Dm), Glycine max (Gm), 616

Homo sapiens (Hs), Mus musculus (Mm), Oryza sativa (Os), Setaria italic (Si), Solanum 617

lycopersicum (Sl), Solanum tuberosum (St), Sorghum bicolor (Sb), and Zea mays (Zm). C and D, 618

Trichome phenotype of wild type and tut1 shown by SEM. Mature trichomes of wild type and tut1 619

are shown in (C). Enlarged view of wild type trichome and tut1 trichome (D) from regions indicated 620

by white squares in (C). Bars = 100 μm (C) and 10 μm (D). 621

622

Figure 8. F-actin organization in wild type and tut1. 623

A and B, F-actin organization in wild type (A) and tut1 (B) mutant leaf trichomes visualized by 624

staining with Alexa Fluor 488-phalloidin. The arrows show the continuous and discontinuous actin 625

filaments in wild type and tut1, respectively. Bars = 20 μm. C, Analysis of average filament length in 626

wild type and tut1. Arrow indicates long actin filaments. Arrowhead indicates fragmented F-actin. 627

*** indicates P < 0.01 (Student’s t -test). D, Analysis of number of filaments in trichomes of wild 628

type and tut1. *** indicates P < 0.01 (Student’s t -test). Over 20 trichomes were used in these 629

analyses. 630



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http://www.ncbi.nlm.nih.gov/pubmed?term=%28Science%5BTitle%5D%29 AND 309%5BVolume%5D AND 741%5BPagination%5D AND Ashikari M%2C Sakakibara H%2C Lin S%2C Yamamoto T%2C Takashi T%2C Nishimura A%2C Angeles ER%2C Qian Q%2C Kitano H%2C Matsuoka M%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Ashikari M, Sakakibara H, Lin S, Yamamoto T, Takashi T, Nishimura A, Angeles ER, Qian Q, Kitano H, Matsuoka M (2005) Cytokinin oxidase regulates rice grain production. Science 309: 741-745

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

http://scholar.google.com/scholar?as_q=Cytokinin oxidase regulates rice grain production&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant Cell%5BTitle%5D%29 AND 17%5BVolume%5D AND 502%5BPagination%5D AND Basu D%2C Le J%2C El%2DEssal Sel%2DD%2C Huang S%2C Zhang C%2C Mallery EL%2C Koliantz G%2C Staiger CJ%2C Szymanski DB%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Basu D, Le J, El-Essal Sel-D, Huang S, Zhang C, Mallery EL, Koliantz G, Staiger CJ, Szymanski DB (2005) DISTORTED3/SCAR2 is a putative Arabidopsis WAVE complex subunit that activates the Arp2/3 complex and is required for epidermal morphogenesis. Plant Cell 17: 502-524

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

http://scholar.google.com/scholar?as_q=DISTORTED3/SCAR2 is a putative Arabidopsis WAVE complex subunit that activates the Arp2/3 complex and is required for epidermal morphogenesis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28J Biol Chem%5BTitle%5D%29 AND 283%5BVolume%5D AND 7135%5BPagination%5D AND Beltzner CC%2C Pollard TD%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Beltzner CC, Pollard TD (2008) Pathway of actin filament branch formation by Arp2/3 complex. J Biol Chem 283: 7135-7144

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http://scholar.google.com/scholar?as_q=Pathway of actin filament branch formation by Arp2/3 complex&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Beltzner&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

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Frank MJ, Cartwright HN, Smith LG (2003) Three Brick genes have distinct functions in a common pathway promoting polarizedcell division and cell morphogenesis in the maize leaf epidermis. Development 130: 753-762


Frank MJ, Smith LG (2002) A small, novel protein highly conserved in plants and animals promotes the polarized growth anddivision of maize leaf epidermal cells. Curr Biol 12: 849-853


Harries PA, Pan A, Quatrano RS (2005) Actin-related protein2/3 complex component ARPC1 is required for proper cellmorphogenesis and polarized cell growth in Physcomitrella patens. Plant Cell 17: 2327-2339


Hiei Y, Ohta S, Komari T, Kumashiro T (1994) Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium andsequence analysis of the boundaries of the T-DNA. Plant J 6: 271-282


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http://scholar.google.com/scholar?as_q=A small, novel protein highly conserved in plants and animals promotes the polarized growth and division of maize leaf epidermal cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Frank&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant Cell%5BTitle%5D%29 AND 17%5BVolume%5D AND 2327%5BPagination%5D AND Harries PA%2C Pan A%2C Quatrano RS%5BAuthor%5D&dopt=abstract

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

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http://scholar.google.com/scholar?as_q=Actin-related protein2/3 complex component ARPC1 is required for proper cell morphogenesis and polarized cell growth in Physcomitrella patens&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Harries&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28 mediated by Agrobacterium and sequence analysis of the boundaries of the T%2DDNA%2E Plant J%5BTitle%5D%29 AND 6%5BVolume%5D AND 271%5BPagination%5D AND Hiei Y%2C Ohta S%2C Komari T%2C Kumashiro T%5BAuthor%5D&dopt=abstract

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28Annu Rev Plant Biol%5BTitle%5D%29 AND 57%5BVolume%5D AND 109%5BPagination%5D AND Hussey PJ%2C Ketelaar T%2C Deeks MJ%5BAuthor%5D&dopt=abstract

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http://scholar.google.com/scholar?as_q=Control of the actin cytoskeleton in plant cell growth&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28Dev Biol%5BTitle%5D%29 AND 282%5BVolume%5D AND 349%5BPagination%5D AND Ikeda K%2C Nagasawa N%2C Nagato Y%5BAuthor%5D&dopt=abstract

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28Embo J%5BTitle%5D%29 AND 6%5BVolume%5D AND 3901%5BPagination%5D AND Jefferson RA%2C Kavanagh TA%2C Bevan MW%5BAuthor%5D&dopt=abstract

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http://scholar.google.com/scholar?as_q=GUS fusions: �-glucuronidase as a sensitive and versatile gene fusion marker in higher plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jefferson&as_ylo=1987&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant J%5BTitle%5D%29 AND 62%5BVolume%5D AND 925%5BPagination%5D AND J�rgens CI%2C Gr�newald N%2C H�lskamp M%2C Uhrig JF%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=J�rgens CI, Gr�newald N, H�lskamp M, Uhrig JF (2010) A role for ABIL3 in plant cell morphogenesis. Plant J 62: 925-935

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

http://scholar.google.com/scholar?as_q=A role for ABIL3 in plant cell morphogenesis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=A role for ABIL3 in plant cell morphogenesis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=J�rgens&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant Cell%5BTitle%5D%29 AND 18%5BVolume%5D AND 3015%5BPagination%5D AND Jung KH%2C Han MJ%2C Lee DY%2C Lee YS%2C Schreiber L%2C Franke R%2C Faust A%2C Yephremov A%2C Saedler H%2C Kim YW%2C Hwang I%2C An G%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Jung KH, Han MJ, Lee DY, Lee YS, Schreiber L, Franke R, Faust A, Yephremov A, Saedler H, Kim YW, Hwang I, An G (2006) Wax-deficient anther1 is involved in cuticle and wax production in rice anther walls and is required for pollen development. Plant Cell 18: 3015-3032

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http://scholar.google.com/scholar?as_q=Wax-deficient anther1 is involved in cuticle and wax production in rice anther walls and is required for pollen development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Wax-deficient anther1 is involved in cuticle and wax production in rice anther walls and is required for pollen development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jung&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Dev Biol%5BTitle%5D%29 AND 231%5BVolume%5D AND 364%5BPagination%5D AND Komatsu M%2C Maekawa M%2C Shimamoto K%2C Kyozuka J%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Komatsu M, Maekawa M, Shimamoto K, Kyozuka J (2001) The LAX1 and FRIZZY PANICLE 2 genes determine the inflorescence architecture of rice by controlling rachis-branch and spikelet development. Dev Biol 231: 364-373

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

http://scholar.google.com/scholar?as_q=The LAX1 and FRIZZY PANICLE 2 genes determine the inflorescence architecture of rice by controlling rachis-branch and spikelet development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The LAX1 and FRIZZY PANICLE 2 genes determine the inflorescence architecture of rice by controlling rachis-branch and spikelet development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Komatsu&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant Physiol%5BTitle%5D%29 AND 151%5BVolume%5D AND 2095%5BPagination%5D AND Kotchoni SO%2C Zakharova T%2C Mallery EL%2C Le J%2C El%2DAssal Sel%2DD%2C Szymanski DB%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Kotchoni SO, Zakharova T, Mallery EL, Le J, El-Assal Sel-D, Szymanski DB (2009) The association of the Arabidopsis actin-related protein2/3 complex with cell membranes is linked to its assembly status but not its activation. Plant Physiol 151: 2095-2109

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http://scholar.google.com/scholar?as_q=The association of the Arabidopsis actin-related protein2/3 complex with cell membranes is linked to its assembly status but not its activation&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=The association of the Arabidopsis actin-related protein2/3 complex with cell membranes is linked to its assembly status but not its activation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kotchoni&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Prog Lipid Res%5BTitle%5D%29 AND 42%5BVolume%5D AND 51%5BPagination%5D AND Kunst L%2C Samuels AL%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Kunst L, Samuels AL (2003) Biosynthesis and secretion of plant cuticular wax. Prog Lipid Res 42: 51-80

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

http://scholar.google.com/scholar?as_q=Biosynthesis and secretion of plant cuticular wax&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Biosynthesis and secretion of plant cuticular wax&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kunst&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Curr Biol%5BTitle%5D%29 AND 13%5BVolume%5D AND 1341%5BPagination%5D AND Le J%2C El%2DAssal Sel%2DD%2C Basu D%2C Saad ME%2C Szymanski DB%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Le J, El-Assal Sel-D, Basu D, Saad ME, Szymanski DB (2003) Requirements for Arabidopsis ATARP2 and ATARP3 during epidermal development. Curr Biol 13: 1341-1347

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http://scholar.google.com/scholar?as_q=Requirements for Arabidopsis ATARP2 and ATARP3 during epidermal development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Requirements for Arabidopsis ATARP2 and ATARP3 during epidermal development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Le&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1


Le J, Mallery EL, Zhang C, Brankle S, Szymanski DB (2006) Arabidopsis BRICK1/HSPC300 is an essential WAVE-complex subunitthat selectively stabilizes the Arp2/3 activator SCAR2. Curr Biol 16: 895-901


Li H, Pinot F, Sauveplane V, Werck-Reichhart D, Diehl P, Schreiber L, Franke R, Zhang P, Chen L, Gao Y, Liang W, Zhang D (2010)Cytochrome P450 family member CYP704B2 catalyzes the ?-hydroxylation of fatty acids and is required for anther cutinbiosynthesis and pollen exine formation in rice. Plant Cell 22: 173-190


Li S, Qian Q, Fu Z, Zeng D, Meng X, Kyozuka J, Maekawa M, Zhu X, Zhang J, Li J, Wang Y (2009) Short panicle1 encodes a putativePTR family transporter and determines rice panicle size. Plant J 58: 592-605


Lichtenthaler HK (1987) Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes. Methods Enzymol 148: 350-382Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Machesky LM, Atkinson SJ, Ampe C, Vandekerckhove J, Pollard TD (1994) Purification of a cortical complex containing twounconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose. J Cell Biol 127: 107-115


Machesky LM, Mullins RD, Higgs HN, Kaiser DA, Blanchoin L, May RC, Hall ME, Pollard TD (1999) Scar, a WASp-related protein,activates nucleation of actin filaments by the Arp2/3 complex. Proc Natl Acad Sci USA 96: 3739-3744


Mathur J, Mathur N, Kernebeck B, Hülskamp M (2003) Mutations in actin-related proteins 2 and 3 affect cell shape development inArabidopsis. Plant Cell 15: 1632-1645


McFarlane HE, Watanabe Y, Yang W, Huang Y, Ohlrogge J, Samuels AL (2014) Golgi- and trans-Golgi network-mediated vesicletrafficking is required for wax secretion from epidermal cells. Plant Physiol 164: 1250-1260


Miki H, Takenawa T (2003) Regulation of actin dynamics by WASP family proteins. J Biochem 134: 309-313Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Miller KG, Field CM, Alberts BM (1989) Actin-binding proteins from Drosophila embryos: a complex network of interacting proteinsdetected by F-actin affinity chromatography. J Cell Biol 109: 2963-2975


Olyslaegers G, Verbelen JP (1998) Improved staining of F-actin and co-localization of mitochondria in plant cells. J Microsc 192: 73-77


Panikashvili D, Savaldi-Goldstein S, Mandel T, Yifhar T, Franke RB, Höfer R, Schreiber L, Chory J, Aharoni A (2007) TheArabidopsis DESPERADO/AtWBC11 transporter is required for cutin and wax secretion. Plant Physiol 145: 1345-1360


Panikashvili D, Shi JX, Schreiber L, Aharoni A (2009) The Arabidopsis DCR encoding a soluble BAHD acyltransferase is requiredfor cutin polyester formation and seed hydration properties. Plant Physiol 151: 1773-1789


Paunola E, Mattila PK, Lappalainen P (2002) WH2 domain: a small, versatile adapter for actin monomers. FEBS Lett 513: 92-97Pubmed: Author and TitleCrossRef: Author and TitleGoogle Scholar: Author Only Title Only Author and Title https://plantphysiol.orgDownloaded on March 14, 2021. - Published by

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28Curr Biol%5BTitle%5D%29 AND 16%5BVolume%5D AND 895%5BPagination%5D AND Le J%2C Mallery EL%2C Zhang C%2C Brankle S%2C Szymanski DB%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Le J, Mallery EL, Zhang C, Brankle S, Szymanski DB (2006) Arabidopsis BRICK1/HSPC300 is an essential WAVE-complex subunit that selectively stabilizes the Arp2/3 activator SCAR2. Curr Biol 16: 895-901

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

http://scholar.google.com/scholar?as_q=Arabidopsis BRICK1/HSPC300 is an essential WAVE-complex subunit that selectively stabilizes the Arp2/3 activator SCAR2&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Arabidopsis BRICK1/HSPC300 is an essential WAVE-complex subunit that selectively stabilizes the Arp2/3 activator SCAR2&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Le&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28hydroxylation of fatty acids and is required for anther cutin biosynthesis and pollen exine formation in rice%2E Plant Cell%5BTitle%5D%29 AND 22%5BVolume%5D AND 173%5BPagination%5D AND Li H%2C Pinot F%2C Sauveplane V%2C Werck%2DReichhart D%2C Diehl P%2C Schreiber L%2C Franke R%2C Zhang P%2C Chen L%2C Gao Y%2C Liang W%2C Zhang D%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Li H, Pinot F, Sauveplane V, Werck-Reichhart D, Diehl P, Schreiber L, Franke R, Zhang P, Chen L, Gao Y, Liang W, Zhang D (2010) Cytochrome P450 family member CYP704B2 catalyzes the ?-hydroxylation of fatty acids and is required for anther cutin biosynthesis and pollen exine formation in rice. Plant Cell 22: 173-190

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

http://scholar.google.com/scholar?as_q=Cytochrome P450 family member CYP704B2 catalyzes the&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Cytochrome P450 family member CYP704B2 catalyzes the&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Li&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant J%5BTitle%5D%29 AND 58%5BVolume%5D AND 592%5BPagination%5D AND Li S%2C Qian Q%2C Fu Z%2C Zeng D%2C Meng X%2C Kyozuka J%2C Maekawa M%2C Zhu X%2C Zhang J%2C Li J%2C Wang Y%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Li S, Qian Q, Fu Z, Zeng D, Meng X, Kyozuka J, Maekawa M, Zhu X, Zhang J, Li J, Wang Y (2009) Short panicle1 encodes a putative PTR family transporter and determines rice panicle size. Plant J 58: 592-605

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

http://scholar.google.com/scholar?as_q=Short panicle1 encodes a putative PTR family transporter and determines rice panicle size&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Short panicle1 encodes a putative PTR family transporter and determines rice panicle size&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Li&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Methods Enzymol%5BTitle%5D%29 AND 148%5BVolume%5D AND 350%5BPagination%5D AND Lichtenthaler HK%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Lichtenthaler HK (1987) Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes. Methods Enzymol 148: 350-382

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

http://scholar.google.com/scholar?as_q=Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lichtenthaler&as_ylo=1987&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28J Cell Biol%5BTitle%5D%29 AND 127%5BVolume%5D AND 107%5BPagination%5D AND Machesky LM%2C Atkinson SJ%2C Ampe C%2C Vandekerckhove J%2C Pollard TD%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Machesky LM, Atkinson SJ, Ampe C, Vandekerckhove J, Pollard TD (1994) Purification of a cortical complex containing two unconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose. J Cell Biol 127: 107-115

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

http://scholar.google.com/scholar?as_q=Purification of a cortical complex containing two unconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Purification of a cortical complex containing two unconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Machesky&as_ylo=1994&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Proc Natl Acad Sci USA%5BTitle%5D%29 AND 96%5BVolume%5D AND 3739%5BPagination%5D AND Machesky LM%2C Mullins RD%2C Higgs HN%2C Kaiser DA%2C Blanchoin L%2C May RC%2C Hall ME%2C Pollard TD%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Machesky LM, Mullins RD, Higgs HN, Kaiser DA, Blanchoin L, May RC, Hall ME, Pollard TD (1999) Scar, a WASp-related protein, activates nucleation of actin filaments by the Arp2/3 complex. Proc Natl Acad Sci USA 96: 3739-3744

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

http://scholar.google.com/scholar?as_q=Scar, a WASp-related protein, activates nucleation of actin filaments by the Arp2/3 complex&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Scar, a WASp-related protein, activates nucleation of actin filaments by the Arp2/3 complex&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Machesky&as_ylo=1999&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant Cell%5BTitle%5D%29 AND 15%5BVolume%5D AND 1632%5BPagination%5D AND Mathur J%2C Mathur N%2C Kernebeck B%2C H�lskamp M%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Mathur J, Mathur N, Kernebeck B, H�lskamp M (2003) Mutations in actin-related proteins 2 and 3 affect cell shape development in Arabidopsis. Plant Cell 15: 1632-1645

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

http://scholar.google.com/scholar?as_q=Mutations in actin-related proteins 2 and 3 affect cell shape development in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Mutations in actin-related proteins 2 and 3 affect cell shape development in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Mathur&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Plant Physiol%5BTitle%5D%29 AND 164%5BVolume%5D AND 1250%5BPagination%5D AND McFarlane HE%2C Watanabe Y%2C Yang W%2C Huang Y%2C Ohlrogge J%2C Samuels AL%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=McFarlane HE, Watanabe Y, Yang W, Huang Y, Ohlrogge J, Samuels AL (2014) Golgi- and trans-Golgi network-mediated vesicle trafficking is required for wax secretion from epidermal cells. Plant Physiol 164: 1250-1260

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

http://scholar.google.com/scholar?as_q=Golgi- and trans-Golgi network-mediated vesicle trafficking is required for wax secretion from epidermal cells&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Golgi- and trans-Golgi network-mediated vesicle trafficking is required for wax secretion from epidermal cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=McFarlane&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28J Biochem%5BTitle%5D%29 AND 134%5BVolume%5D AND 309%5BPagination%5D AND Miki H%2C Takenawa T%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Miki H, Takenawa T (2003) Regulation of actin dynamics by WASP family proteins. J Biochem 134: 309-313

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

http://scholar.google.com/scholar?as_q=Regulation of actin dynamics by WASP family proteins&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Regulation of actin dynamics by WASP family proteins&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Miki&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28J Cell Biol%5BTitle%5D%29 AND 109%5BVolume%5D AND 2963%5BPagination%5D AND Miller KG%2C Field CM%2C Alberts BM%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Miller KG, Field CM, Alberts BM (1989) Actin-binding proteins from Drosophila embryos: a complex network of interacting proteins detected by F-actin affinity chromatography. J Cell Biol 109: 2963-2975

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

http://scholar.google.com/scholar?as_q=Actin-binding proteins from Drosophila embryos: a complex network of interacting proteins detected by F-actin affinity chromatography&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

http://scholar.google.com/scholar?as_q=Actin-binding proteins from Drosophila embryos: a complex network of interacting proteins detected by F-actin affinity chromatography&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Miller&as_ylo=1989&as_allsubj=all&hl=en&c2coff=1

http://www.ncbi.nlm.nih.gov/pubmed?term=%28J Microsc%5BTitle%5D%29 AND 192%5BVolume%5D AND 73%5BPagination%5D AND Olyslaegers G%2C Verbelen JP%5BAuthor%5D&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Olyslaegers G, Verbelen JP (1998) Improved staining of F-actin and co-localization of mitochondria in plant cells. J Microsc 192: 73-77

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http://scholar.google.com/scholar?as_q=Improved staining of F-actin and co-localization of mitochondria in plant cells&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

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http://scholar.google.com/scholar?as_q=The Arabidopsis DESPERADO/AtWBC11 transporter is required for cutin and wax secretion&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Panikashvili&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

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Stradal TE, Rottner K, Disanza A, Confalonieri S, Innocenti M, Scita G (2004) Regulation of actin dynamics by WASP and WAVEfamily proteins. Trends Cell Biol 14: 303-311


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Wang Z, Chen C, Xu YY, Jiang RX, Han Y, Xu ZH, Chong K (2004) A practical vector for efficient knockdown of gene expression inrice (Oryza sativa L.). Plant Mol Biol Rep 22: 409-417


Winter DC, Choe EY, Li R (1999) Genetic dissection of the budding yeast Arp2/3 complex: A comparison of the in vivo andstructural roles of individual subunits. Proc Natl Acad Sci USA 96: 7288-7293


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Copyright (c) 2020 American Society of Plant Biologists. All rights reserved.

http://www.ncbi.nlm.nih.gov/pubmed?term=%28Mol Plant%5BTitle%5D%29 AND 4%5BVolume%5D AND 985%5BPagination%5D AND Qin BX%2C Tang D%2C Huang J%2C Li M%2C Wu XR%2C Lu LL%2C Wang KJ%2C Yu HX%2C Chen JM%2C Gu MH%2C Cheng ZK%5BAuthor%5D&dopt=abstract

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http://scholar.google.com/scholar?as_q=ABCG15 encodes an ABC transporter protein, and is essential for post-meiotic anther and pollen exine development in rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

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

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http://www.ncbi.nlm.nih.gov/pubmed?term=%28elegans Arp2%2F3 complex in cell migration during ventral enclosure%2E J Cell Sci%5BTitle%5D%29 AND 116%5BVolume%5D AND 1505%5BPagination%5D AND Sawa M%2C Suetsugu S%2C Sugimoto A%2C Miki H%2C Yamamoto M%2C Takenawa T%5BAuthor%5D&dopt=abstract

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Zhang Z, Zhang Y, Tan H, Wang Y, Li G, Liang W, Yuan Z, Hu J, Ren H, Zhang D (2011) RICE MORPHOLOGY DETERMINANTencodes the type II formin FH5 and regulates rice morphogenesis. Plant Cell 23: 681-700


Zheng H, Rowland O, Kunst L (2005) Disruptions of the Arabidopsis Enoyl-CoA reductase gene reveal an essential role for very-long-chain fatty acid synthesis in cell expansion during plant morphogenesis. Plant Cell 17: 1467-1481



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Documents

Running Head: Rice TUT1 Regulates Panicle Development · 2015. 8. 4. · 9 144 RESULTS 145 146 Identification and characterization of the rice tut1 mutant 147 To understand the cellular