gad1 encodes a secreted peptide that regulates grain ...plants contain 46 several signal peptides,...

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RESEARCH ARTICLE 1

GAD1 Encodes a Secreted Peptide That Regulates Grain Number, Grain 2

Length and Awn Development in Rice Domestication 3

4

Jing Jina, Lei Huaa, Zuofeng Zhua, Lubin Tana, Xinhui Zhaoa, Weifeng Zhanga, 5 Fengxia Liua, Yongcai Fua, Hongwei Caia, Xianyou Suna, Ping Gua, Daoxin Xieb, 6 Chuanqing Suna,1 7

8 a State Key Laboratory of Plant Physiology and Biochemistry, National Center for 9 Evaluation of Agricultural Wild Plants (Rice), Beijing Key Laboratory of Crop Genetic 10 Improvement, Department of Plant Genetics and Breeding, China Agricultural 11 University, Beijing, 100193, China 12 b Tsinghua-Peking Center for Life Sciences, School of Life Sciences, Tsinghua 13 University, Beijing 100084, China 14 1Corresponding author: [email protected] 15

16 Short title: GAD1 Regulates Grain and Awn Development 17

18 One-sentence summary: GAD1 encodes a secreted peptide that controls critical 19 transitions of rice domestication such from fewer, longer grains with long awns to 20 more, shorter grains with no or short awns. 21

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

26 ABSTRACT 27 Cultivated rice (Oryza sativa L.) was domesticated from wild rice (Oryza rufipogon 28 Griff.), which typically displays fewer grains per panicle and longer grains than 29 cultivated rice. In addition, wild rice has long awns, whereas cultivated rice has short 30 awns or lacks them altogether. These changes represent critical events in rice 31 domestication. Here, we identified a major gene, GRAIN NUMBER, GRAIN LENGTH 32 AND AWN DEVELOPMENT 1 (GAD1), that regulates those critical changes during 33 rice domestication. GAD1 is located on chromosome 8 and is predicted to encode a 34 small secretary signal peptide belonging to the EPIDERMAL PATTERNING 35 FACTOR-LIKE family. A frame-shift insertion in gad1 destroyed the conserved 36 cysteine residues of the peptide, resulting in a loss of function, and causing the 37 increased number of grains per panicle, shorter grains and awnless phenotype 38 characteristic of cultivated rice. Our findings provide a useful paradigm for revealing 39 functions of peptide signal molecules in plant development and helps elucidate the 40 molecular basis of rice domestication.41

Plant Cell Advance Publication. Published on September 15, 2016, doi:10.1105/tpc.16.00379

©2016 American Society of Plant Biologists. All Rights Reserved

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INTRODUCTION 42

Cell-to-cell communication helps coordinate developmental and environmental 43

responses. Secreted peptides mediate cell-to-cell signaling during 44

development and pattern formation in multicellular organisms. Plants contain 45

several signal peptides, such as systemin, which functions in the wounding 46

response (Pearce et al., 1991), phytosulfokine, which induces the proliferation 47

of single mesophyll cells (Matsubayashi and Sakagami, 1996), S-locus 48

cysteine-rich protein/S-locus protein 11, which determines self-incompatibility 49

(Schopfer et al., 1999), LUREs, which direct pollen-tube elongation (Okuda et 50

al., 2009), and CLAVATA3, which mediates the signaling of cell fate decisions 51

in the Arabidopsis thaliana shoot meristem (Fletcher et al., 1999). 52

EPIDERMAL PATTERING FACTORs (EPFs), such as EPF1, EPF2 and 53

EPF-like 9 (EPFL9)/Stomagen, act as key regulators of stomatal development 54

in Arabidopsis (Engineer et al., 2014; Hara et al., 2007; Hara et al., 2009; Hunt 55

et al., 2009; Sugano et al., 2010). 56

Cultivated rice (Oryza sativa L.), a staple food for more than half of the 57

global population (Khush, 1997), was domesticated from the wild rice (Oryza 58

rufipogon) ~8,000 years ago (Fuller et al., 2010; Zong et al., 2007). Both 59

morphological traits and physiological characteristics changed remarkably 60

during domestication. Thorough understanding of rice domestication 61

processes and molecular mechanisms active in cultivated rice will be 62

beneficial for rice improvement. During the past ten years, the rice evolutionary 63

mechanisms and several domestication-related genes, including Sh4/SHA1 64

(Konishi et al., 2006; Li et al., 2006; Lin et al., 2007), PROG1 (Jin et al., 2008; 65

Tan et al., 2008), Bh4 (Zhu et al., 2011), Rc (Sweeney et al., 2006), OsLG1 66

(Ishii et al., 2013; Zhu et al., 2013), An-1 (Luo et al., 2013) and LABA1/An-2 67

(Gu et al., 2015; Hua et al., 2015) were characterized. 68

Wild rice typically produces a few, long grains per panicle, with long awns 69

(Figure 1A) that are crucial for seed dispersal and deterring granivore 70

predation. However, most current rice cultivars generate many grains per 71

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panicle, shorter grains and show no or only short awns (Figure 1B). These 72

characteristics facilitate rice storage and processing (Hua et al., 2015) and the 73

differences between wild and cultivated rice represent critical events in rice 74

domestication. However, the molecular mechanisms responsible for these 75

changes are still largely unknown. 76

Here, we found that an EPIDERMAL PATTERNING FACTOR-LIKE (EPFL) 77

peptide, encoded by GRAIN NUMBER, GRAIN LENGTH AND AWN 78

DEVELOPMENT 1 (GAD1) regulates grain number, grain length and awn 79

development, all of which were altered during rice domestication. A frame-shift 80

insertion in gad1 destroyed the conserved cysteine residues of the EPFL 81

peptide, causing the increased number of grains per panicle, the shorter grains 82

and the awnless phenotype found in cultivated rice. Our results thus establish 83

the function of a peptide signal molecule in plant development and help 84

uncover the genetic basis of rice domestication. 85

86

RESULTS 87

Characterization and Cloning of GAD1 88

We obtained one introgression line (OIL31) from a set of introgression lines 89

derived from a cross between indica variety NA93-11 and an accession of 90

common wild rice (W2014, O. rufipogon). OIL31, which harbored four W2014 91

chromosomal segments, produced fewer grains per panicle and longer grains 92

as well as a high percentage of awned seeds (~80%), with long awns (18 ± 93

1.35 mm) in comparison with NA93-11 (Figure 2). 94

A genetic linkage analysis of 236 F2 individuals derived from a cross 95

between OIL31 and NA93-11 indicated that the grain number per panicle, grain 96

length, and awn phenotype were controlled by a single dominant gene, GAD1. 97

We used the awn phenotype as a representative morphological marker to 98

locate the GAD1 gene between simple sequence repeat markers M10 and 99

RM5485 on the long arm of chromosome 8 (Figure 3A). Using 4,250 F2 100

individuals, we further delimited GAD1 to within an ~6-kb region between the 101

4

markers MX14 and MX16, in which there was only one predicted gene 102

(Os08g0485500) in the Nipponbare genome (The Rice Annotation Project 103

Database) (Figure 3B). A comparison of the Os08g0485500 coding region 104

between OIL31 and NA93-11 revealed one single nucleotide polymorphism 105

(SNP1) and three insertion/deletions (Indel1, Indel2 and Indel3). SNP1 was a 106

synonymous mutation, while Indel1 (a 3-bp deletion in NA93-11), Indel2 (a 107

6-bp insertion in NA93-11) and Indel3 (a 7-bp insertion in NA93-11) led to a 108

single amino-acid residue deletion, a two amino-acid residue insertion, and a 109

frame shift, respectively (Figure 3C; Supplemental Figure 1). 110

To verify the function of Os08g0485500, we generated a construct (pCPL) 111

that contained a ~4.0-kb genomic fragment including the entire Os08g0485500 112

gene from wild rice W2014 (Figure 3D), and introduced the pCPL construct 113

into the domesticated japonica variety Nipponbare. Compared with 114

Nipponbare (controls), 32 independent transgenic lines of the wild rice 115

Os08g0485500 (pCPL) showed fewer grains per panicle, longer grains and 116

long awns (Figures 3E to 3I, 3O and 3Q). These results demonstrated that 117

Os08g0485500 in the O. rufipogon genome was a key gene (GAD1) regulating 118

grain number, grain length and awn development. 119

We further transformed a GAD1 interfering RNA (RNAi) construct (Figure 120

3D) into an introgression line harboring the GAD1 locus. The down-regulation 121

of the GAD1 transcript level in the RNAi transgenic plants led to more grains 122

per panicle, shorter grains, and the short-awns (or awnless) phenotype, 123

compared with control plants (Figures 3J to 3N, 3P and 3R). Thus, GAD1 is the 124

key gene controlling the change in grain number, grain length and awn 125

development that occurred during rice domestication. 126

127

Expression of GAD1 Promotes Grain Elongation and Awn Development 128

Using reverse transcription-quantitative PCR (qRT-PCR), we analyzed the 129

GAD1 expression pattern. GAD1 was preferentially expressed in the 130

developing panicles of OIL31, especially in panicles less of than 0.5 cm in 131

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length. Lower levels of transcript accumulation were detected in roots, leaves, 132

leaf-sheaths and stems. With the development of panicles, GAD1 expression 133

levels dropped dramatically in both NA93-11 and OIL31 (Figure 4A). Using 134

RNA in situ hybridization, we detected GAD1 transcripts in the primary and 135

secondary branch meristem in OIL31 and NA93-11 (Supplemental Figures 2 136

and 3). Because awns are organs at the tips of lemmas, we also compared the 137

GAD1 expression in NA93-11 and OIL31 at the apices of glumes, which are 138

composed of lemmas and paleae. GAD1 expression was higher in OIL31 than 139

in NA93-11 at the glume apices (Figure 4B). 140

To reveal the effects of GAD1, we compared awn development between 141

OIL31 and NA93-11 using scanning electron microscopy. Based on the rice 142

spikelet development (Sp) stages defined by Itoh et al., (2005), we detected no 143

difference between OIL31 and NA93-11 until the Sp6 stage. At this stage, awn 144

primordia appeared at the apices of the lemmas in OIL31 and extended from 145

Sp6 to Sp8. However, during the same stages, no awns formed in NA93-11 146

(Supplemental Figure 4). 147

The detection of GAD1 expression during spikelet development using 148

RNA in situ hybridization consistently showed no differences in the GAD1 149

expression patterns between OIL31 and NA93-11 until the Sp6 stage (Figures 150

4C1 and 4D1). At this stage, the transcripts of GAD1 were more abundant at 151

the apices of lemmas in OIL31 than in NA93-11 (Figures 4C2 and 4D2). At the 152

Sp7 stage and early Sp8 stage (Sp8e), the abundant expression of GAD1 was 153

maintained at the apices of lemmas that formed the awn primordia in OIL31, 154

and GAD1 expression was still low in NA93-11 (Figures 4C3, 4C4 and 4D3, 155

4D4). Thus, the GAD1 expression pattern was consistent with its role in the 156

promotion of awn development. Additionally, at the late Sp8 stage (Sp8I), when 157

spikelet development is almost complete, the GAD1 transcript was weakly 158

detected in lemmas of both OIL31 and NA93-11 (Figures 4C5 and 4D5). 159

Previous studies indicated that cell division plays a crucial role in 160

promoting awn formation and elongation (Hua et al., 2015; Luo et al., 2013). 161

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Therefore, we examined the expression of HISTONE H4, a marker gene of 162

active cell division (Marzluff and Duronio, 2002) using RNA in situ hybridization. 163

The HISTONE H4 expression patterns were similar between OIL31 and 164

NA93-11 until the Sp7 stage (Figures 4E1, 4E2 and 4F1, 4F2). However, at the 165

Sp7 and Sp8 stages, HISTONE H4 expression was higher in the awn 166

primordia of OIL31 than in those of NA93-11 (Figures 4E3 to 4E5 and 4F3 to 167

4F5). Thus, there was more active cell division in the awn primordia of OIL31 168

than of NA93-11. A qRT-PCR analysis of HISTONE H1, another marker gene 169

of active cell division (Luo et al., 2013; Marzluff and Duronio, 2002), 170

consistently showed that the expression of HISTONE H1 was higher in the 171

young panicles of OIL31 than those of NA93-11 (Supplemental Figure 5). 172

An analysis of cell cycle-related gene expression at the apices of glumes 173

showed that 22 genes involved in the G1/S and G2/M phases regulating mitotic 174

division (Li et al., 2011) were more highly expressed at the apices of glumes in 175

OIL31 than in NA93-11 (Supplemental Figure 6). Thus, cell division was more 176

active at the apices of glumes in OIL31 than in NA93-11, indicating that GAD1 177

promoted continuous cell division at the apices of glumes, which likely led to 178

the promotion of grain elongation and awn development in wild rice. 179

180

Cytokinin is Decreased in the Introgression Line OIL31 181

The plant hormone cytokinin is one of the factors controlling shoot apical 182

meristem activity, which is a central determinant of grain number. Previous 183

studies revealed that down-regulation of CKX2 or DST causes cytokinin 184

accumulation and increases grain number in rice (Ashikari et al., 2013; Li et al., 185

2013; Zhang and Yuan, 2014). Our RNA in situ hybridization and real-time 186

PCR analysis showed that expression of CKX2 and DST were increased in the 187

introgression line OIL31 compared to cultivar NA93-11 (Figures 4G and 4H; 188

Supplemental Figure 7A). Consistent with the high expression of CKX2 and 189

DST, the cytokinin (iP and tZ) contents were substantially decreased in the 190

inflorescence meristem of OIL31, and expression of the cytokinin-response 191

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marker genes RR1 and RR2 (Jain et al., 2006) was significantly decreased in 192

the young panicles of OIL31 (Figures 4I - 4K; Supplemental Figure 7B). 193

Together with our observations of higher GAD1 expression and fewer grains 194

per panicle in OIL31 (Figures 2F and 4A) and previous genetic evidence 195

(Ashikari et al., 2013; Li et al., 2013), our results suggest a hypothesis that 196

GAD1 activates DST and CKX2 to reduce the cytokinin level and cause less 197

grain number per panicle in wild rice. 198

199

GAD1 Encodes a Predicted Small Secreted Cysteine-rich Peptide with a 200

Conserved Cysteine Residue 201

The GAD1 cDNA in wild rice W2014 encodes a 127-amino-acid protein that is 202

a homolog of a small secreted cysteine-rich EPF/EPFL peptide in Arabidopsis 203

thaliana (Engineer et al., 2014; Hara et al., 2007; Hara et al., 2009; Hunt et al., 204

2009; Sugano et al., 2010; Takata et al., 2013) (Supplemental Figures 8 and 9 205

and Supplemental Data set 1). Similar to the Arabidopsis EPF/EPFL peptides, 206

the GAD1 prepropeptide contained an N-terminal secretory signal peptide and 207

a mature peptide with a conserved region of six cysteine residues 208

(Supplemental Figures 10 and 11; Supplemental Table 1). The six conserved 209

cysteine residues are essential for proper formation of intramolecular disulfide 210

bonds that are critical for peptide function (Abrash and Bergmann, 2010; 211

Jewaria et al., 2013; Katsir et al., 2011; Lee et al., 2012; Ohki et al., 2011; 212

Takata et al., 2013). A comparison of the predicted mature peptides between 213

wild rice W2014 and cultivar NA93-11 indicated that the 7-bp (Indel3) insertion 214

in gad1 caused a frame-shift mutation that led to the loss of two (5th and 6th) 215

essential cysteine residues and, accordingly, likely prevented the formation of 216

the proper mature peptide, resulting in the loss of the gad1 function in NA93-11 217

(Figure 5A). 218

To determine the amount of GAD1 variation among wild and cultivated rice, 219

we sequenced the GAD1 coding region in 53 accessions of wild rice 220

originating from 12 countries and 203 accessions of cultivated rice from 15 221

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countries. We found 10 kinds of GAD1/gad1 haplotypes in wild and cultivated 222

rice (Supplemental Figure 12). The GAD1 gene of most wild rice contained a 223

9-bp insertion or was similar to W2014, and encoded a mature protein 224

containing six cysteine residues (6C). This gene was associated with the 225

long-awn phenotype, suggesting that GAD1 with six cysteine residues is 226

functional. In the cultivated rice, there were 180 varieties that, like NA93-11, 227

had frame shifts that destroyed the classic 5th and 6th cysteine residues in 228

GAD1. Among those, 137 varieties contained a 7-bp insertion in the Indel3 of 229

gad1 to generate a dysfunctional mutated mature peptide containing seven 230

cysteine residues (7C), while the remaining 43 varieties contained a 5-bp 231

insertion in Indel3 to encode the dysfunctional mutated protein with four 232

cysteine residues (4C). Among the 180 varieties with the dysfunctional gad1 233

peptide, 161 (~90%) of the varieties showed the awnless phenotype (Figures 234

5B and 5C; Supplemental Figure 12), while 19 varieties displayed an awn 235

phenotype, which may be caused by other awn-regulatory gene(s). 236

237

Neutrality Test and Positive Selection on gad1 in O. sativa 238

The nucleotide diversity of gad1 in cultivated rice was remarkably lower than 239

that of GAD1 in wild rice. Tajima’s D test suggested that gad1 in cultivated rice 240

deviated significantly from the neutral expectation (P < 0.01), while GAD1 in 241

wild rice did not deviate from the neutral expectation (P > 0.10) (Supplemental 242

Table 2). We further examined the nucleotide diversity of 16 loci in the 243

GAD1-containing 1.9-Mb genomic region, and identified an interval of ~900-kb 244

surrounding the GAD1 locus that exhibited a significantly decreased level of 245

nucleotide diversity in cultivated rice relative to wild rice (Figure 6). Our results 246

are consistent with recent whole-genome investigations (Huang et al., 2012; 247

Xu et al., 2012). These data indicated that the gad1 gene regulating grain 248

number, grain length and awn development in O. sativa was strongly selected 249

by humans during rice domestication. 250

To further investigate the selection of GAD1 during rice domestication, we 251

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used 159 accessions of cultivated rice from 15 countries to compare the 252

appearance of GAD1 with two recently identified genes involved in awn 253

development, An-1, encoding a basic helix-loop-helix transcription factor (Luo 254

et al., 2013), and LABA1/An-2, encoding a cytokinin-activating enzyme (Gu et 255

al., 2015; Hua et al., 2015). We found that the proportions of cultivars 256

harboring the GAD1, LABA1 and An-1 alleles were 14%, 30% and 30% 257

respectively (Supplemental Figure 13), suggesting that the gad1 selection 258

might be stronger than that for either laba1 or an-1 during rice domestication. 259

260

GAD1 Might Have Additive Effects with An-1 and LABA1/An-2 261

Two key genes, An-1 and LABA1/An-2, that affect awn development during 262

rice domestication have been characterized previously. An-1 encodes a basic 263

helix-loop-helix protein and regulates awn development (Luo et al., 2013). 264

LABA1/An-2 encodes a cytokinin-activating enzyme and controls awn barb 265

formation and awn elongation (Gu et al., 2015; Hua et al., 2015). To determine 266

the relationships among An-1, LABA1/An-2 and GAD1, we generated a set of 267

near-isogenic lines and pyramiding lines. The awn length of the pyramiding 268

line harboring GAD1 and An-1 (6.27 ± 0.12 cm) was much longer than those of 269

near-isogenic lines containing only GAD1 (1.89 ± 0.16 cm) or An-1 (4.6 ± 0.26 270

cm). Additionally, the awn length of a pyramiding line harboring GAD1 and 271

LABA1/An-2 (2.68 ± 0.16 cm) was longer than that of a near-isogenic line 272

containing LABA1/An-2 (no awn) and GAD1 (Supplemental Figure 14). Thus, 273

GAD1 might have an additive effect with An-1 and LABA1/An-2. 274

275

DISCUSSION 276

In this paper, we found that the GAD1-encoded EPFL peptide regulates grain 277

number, grain length and awn development in rice. A frame-shift insertion in 278

coding region of the GAD1 gene was responsible for the transition from a few, 279

long grains per panicle, with long awns to many grains per panicle, shorter 280

grains and show no or only short awns during rice domestication. 281

10

Crop domestication is the genetic modification of a wild species to 282

generate a new form to meet human needs through cultivation (Doebley et al., 283

2006). Wild rice shows prostrate growth, severe shattering, spread panicles 284

and long awns, while cultivated rice exhibits erect growth, no seed-shattering 285

and compact panicles, which may be more amenable to cultivation and allow 286

harvest of more seeds (Jin et al., 2008; Ishii et al., 2013; Konishi et al., 2006; Li 287

et al., 2006; Lin et al., 2007; Tan et al., 2008; Zhu et al., 2013). Recently, 288

several genes controlling rice domestication-related traits have been 289

characterized. These genes encode transcription factors or enzymes (Gu et al., 290

2015; Hua et al., 2015; Ishii et al., 2013; Jin et al., 2008; Konishi et al., 2006; Li 291

et al., 2006; Lin et al., 2007; Luo et al., 2013; Sweeney et al., 2006; Tan et al., 292

2008; Zhu et al., 2011; Zhu et al., 2013). By contrast, our study revealed that 293

key rice domestication-related traits, including awn development, and 294

yield-related traits were regulated by a predicted small secretory signal peptide 295

GAD1, related to the Arabidopsis EPFL family. Our findings not only provide 296

evidence revealing novel functions of peptide signal molecules during plant 297

development, but also offer intriguing insights on the role of peptide signal 298

molecules in rice domestication. 299

300

The EPF/EPFL Peptides Have Multiple Biological Functions 301

Multiple biological functions of EPF/EPFL family members have been 302

characterized in plants. In Arabidopsis, EPF1, EPF2 and STOMAGEN/EPFL9 303

regulate stomatal development (Engineer et al., 2014; Hara et al., 2007; Hara 304

et al., 2009; Hunt et al., 2009; Sugano et al., 2010); EPFL4 and 305

EPFL6/CHALLAN(CHAL), two EPFL family members, promote inflorescence 306

growth (Uchida et al., 2012); EPFL5 promotes fertility and fruit growth (Abrash 307

et al., 2011). In Brassica napus, EFFL6 regulates filament elongation and may 308

also play affect organ morphology and floral organ specification (Huang et al., 309

2014). In this study, we verified that the EPFL peptide encoded by GAD1 310

regulates grain number, grain length and awn development in rice. 311

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EPF1, EPF2 and STOMAGEN/EPFL9 regulate plant development by 312

binding ERECTA family members (Lee et al., 2012; Lee et al., 2015). Two 313

genes (LOC_Os06g03970 and LOC_Os06g10230) in the rice genome show 314

high homology to Arabidopsis ERECTA. These genes therefore appear to be 315

good initial candidates to encode proteins that serve as GAD1 receptors. 316

317

GAD1 Has Pleiotropic Effects on Awn and Grain Development 318

Previous studies showed that some key genes controlling the domestication 319

processes of crops have pleiotropic effects. For example, the selection of 320

prog1 led to erect growth, greater grain number and higher grain yields in 321

cultivated rice (Tan et al., 2008). LG1 controls rice ligule development and the 322

transition of panicle architecture in cultivars during rice domestication (Ishii et 323

al., 2013; Lee et al. 2007; Zhu et al., 2013). Q is a major gene involved in 324

wheat domestication-related traits, such as free-threshing character, glume 325

shape and tenacity, and rachis fragility (Simons et al., 2006). In our study, we 326

found that GAD1 also had pleiotropic effects – on grain length, grain number 327

and awn development – during rice domestication. These findings indicate that 328

targeting a gene with pleiotropic effects might be an efficient strategy for crop 329

domestication. 330

331

Crop domestication represents the genetic modification of a wild species to 332

create a new form to meet human needs. Our findings provided insights into 333

the puzzles surrounding rice domestication and revealed a previously 334

unrecognized function of peptide signal molecules in plant development. 335

12

METHODS 336

337

Plant Materials and Growth Conditions 338

For the fine mapping, expression pattern analysis, and panicle development 339

analysis, we used introgression line OIL31 and indica cultivar NA93-11, which 340

is a sibling of 93-11 with no awn. OIL31 was derived from a cross between 341

W2014 (common wild rice from India) and NA93-11. An F2 segregation 342

population, containing 4,250 individuals, derived from a backcross between 343

OIL31 and NA93-11 was used for the fine mapping of GAD1. The introgression 344

line TIL189 was derived from a cross between the indica cultivar Teqing and 345

common wild rice W2014. The japonica variety Nipponbare and the 346

introgression line TIL189 were used as transgenic recipients. We generated 347

two pyramiding lines (An1/GAD1-NIL and GAD1/LABA1-NIL) that harbored 348

GAD1 and An-1 (Luo et al., 2013), and GAD1 and LABA1/An-2 (Gu et al., 349

2015; Hua et al., 2015), respectively, in the NA93-11 genetic background. The 350

introgression and pyramiding lines were generated as described previously 351

(Tan et al., 2007). The plant materials were growth in the field at the 352

Experimental Stations of China Agriculture University, Beijing and Hainan, 353

China. The wild rice and cultivated rice accessions used in the haplotype 354

analysis, neutrality test and selective sweep analysis are listed in 355

Supplemental Data sets 2, 3 and 4, respectively. 356

357

Primers 358

The primers used in this study are listed in Supplemental Table 3. 359

360

Generation of Constructs and Transformations 361

The functional complementation construct (pCPL), which harbored the ~4.0-kb 362

W2014 genomic fragment covering the entire GAD1 gene, was inserted into 363

the binary vector pCAMBIA1300 (AF234296）. The GAD1 DNA fragments were 364

PCR-amplified by 1300-3 (Supplemental Table 3) and cloned into the 365

13

pCAMBIA1300 vector KpnⅠ and SalⅠ site. The RNAi construct contained 366

an inverted repeat harboring the 350-bp GAD1 cDNA fragment in the vector 367

pTCK303. The two plasmid constructs were introduced to Agrobacterium 368

tumefaciens strain EHA105, and then transformed into the japonica variety 369

Nipponbare and the introgression line TIL189. 370

371

Phenotypic Evaluation 372

We used 20 plants to measure the grain number of main panicles, grain length, 373

awn length and the percentage of awned seeds in the introgression line OIL31 374

and NA93-11. The average awn length was measured on the apical spikelet of 375

each primary branch on the main stem panicle. The percentages of awned 376

seeds were determined from awned spikelets on the main stem panicle, with 377

the awns > 1 mm on mature grains considered to represent an awned 378

phenotype. Two independent complemented (with pCPL) and RNAi T2 379

homozygous lines were used for the measurements. The main panicles of 10 380

plants were measured for each line. Ten Nipponbare and TIL189 plants 381

harboring empty vectors were used as the controls. 382

383

5′ - and 3′ - RACE and Quantitative RT-PCR 384

We extracted total RNA using TRIzol reagent (Invitrogen Life Technologies) 385

and purified RNA with a purification kit (TIANGEN). Total RNA was reverse 386

transcribed with an oligo (dT20) primer using SuperScript III Reverse 387

Transcriptase (Life Technologies). 5′ and 3′ RACE were conducted with 5′- and 388

3′-Full RACE Kits (TaKaRa) following the manufacturer’s instructions. 389

Quantitative RT-PCR was performed on a CFX96 real-time system (Bio-Rad). 390

Diluted cDNA was amplified using the SsoFast Evagreen Supermix (Bio-Rad). 391

The transcript levels were normalized to endogenous ACTIN transcripts 392

amplified with primers ActinF and ActinR. Each set of experiments was 393

repeated three times, and the relative quantification method (2-ΔΔCT) was used 394

to evaluate quantitative variation (Livak and Schmittgen, 2001). 395

14

396

Scanning Electron Microscopy 397

We fixed the young panicles in 2.5% glutaraldehyde-phosphate buffer saline 398

fixative solution, dehydrated the samples using an ethanol series, and dried 399

them with a carbon dioxide critical-point dryer. The dried panicles were gold 400

plated and observed using a Hitachi S-2460 scanning electron microscope at 401

15 kV. Scanning electron microscopy was performed as described previously 402

(Hua et al., 2015). 403

404

RNA in situ Hybridization 405

RNA in situ hybridization was performed as described previously (De Block M 406

and Debrouwer D, 1993) with minor modification. The young panicles of all 407

stages from OIL31 and NA93-11 were fixed in 4% (w/v) paraformaldehyde, 408

subjected to a dehydration series and infiltration, and embedded in paraplast 409

(Paraplast Plus; Sigma–Aldrich). The tissues were sliced into 8–10 μm 410

sections with a microtome (Leica RM2145). A ~300-bp fragment of GAD1 411

cDNA was amplified and used as the template to generate sense and 412

antisense RNA probes that were labeled by digoxigenin using a DIG Northern 413

Starter Kit (Catalog No. 2039672; Roche) following the manufacturer’s 414

instructions. Five independent experiments have been performed for RNA in 415

situ Hybridization. 416

417

Sequencing and Data Analysis 418

Multiple sequences were aligned with ClustalX (Thompson et al., 1997). The 419

program DnaSP, version 5.0, was used to calculate nucleotide diversity and 420

Tajima’s D test (Rozas et al., 2003). The phylogenetic tree was constructed by 421

the MEGA 5 program The evolutionary history was inferred using the 422

Neighbor-Joining method. The percentage of replicate trees in which the 423

associated taxa clustered together in the bootstrap test (1000 replicates) are 424

shown next to the branches (Tamura et al., 2011). 425

15

426

Signal Peptide Analysis 427

The cleavage sites and a signal peptide/non-signal peptide in the GAD1 428

protein were predicted using SignalP4.1 429

(http://www.cbs.dtu.dk/services/SignalP) (Petersen et al., 2011). 430

431

Accession Numbers 432

Sequence data from this article can be found in the GenBank/EMBL databases 433

under the following accession numbers: GAD1 cDNA from W2014 (XXX) and 434

gad1 cDNA from NA93-11 (XXX) 435

436

Supplemental Data 437

Supplemental Figure 1. GAD1 sequence comparison between OIL31 and 438

NA93-11. 439

Supplemental Figure 2. RNA in situ hybridization of GAD1 in inflorescence 440

meristem. 441

Supplemental Figure 3. RNA in situ hybridization of the GAD1 sense probe at 442

different developmental stages in OIL31. 443

Supplemental Figure 4. Scanning electron microscopy images of spikelets at 444

different developmental stages. 445

Supplemental Figure 5. qRT-PCR analysis of HISTONE H1 in young panicles 446

of NA93-11 and OIL31. 447

Supplemental Figure 6. Transcript level comparison of 22 cell 448

cycle-associated genes between NA93-11 and OIL31. 449

Supplemental Figure 7. Comparison of CKX2, DST and RR transcript levels 450

between NA93-11 and OIL31. 451

Supplemental Figure 8. Full-length cDNA of GAD1 and its predicted amino 452

acid sequence. 453

Supplemental Figure 9. Phylogenetic tree of EPF/EPFL family genes from 454

rice and Arabidopsis. 455

16

Supplemental Figure 10. Protein structure and maturation process 456

predictions of GAD1. 457

Supplemental Figure 11. Analysis of the consensus sequence from the GAD1 458

protein. 459

Supplemental Figure 12. Haplotypes of GAD1 in diverse wild and cultivated 460

rice accessions. 461

Supplemental Figure 13. Distribution of GAD1/gad1, LABA1/laba1 and 462

An-1/an-1 in 159 varieties of cultivated rice. 463

Supplemental Figure 14. Phenotypes of near-isogenic and pyramiding lines 464

containing GAD1, An-1 and LABA1. 465

Supplemental Table 1. The signal peptide and cleavage site predictions in the 466

GAD1 protein. 467

Supplemental Table 2. GAD1 nucleotide diversity and Tajima’s D test. 468

Supplemental Table 3. Primers used in this study. 469

Supplemental Data set 1. Text file of the alignment used for the phylogenetic 470

analysis in Supplemental Figure 9. 471

Supplemental Data set 2a. Wild rice accessions used in the haplotype and 472

nucleotide diversity analyses. 473

Supplemental Data set 2b. Rice cultivars used in the haplotype and 474

nucleotide diversity analyses. 475

Supplemental Data set 3a. Wild rice accessions used in the selective sweep 476

analysis. 477

Supplemental Data set 3b. Rice cultivars used in the selective sweep 478

analysis. 479

Supplemental Data set 4. Rice cultivars used to analyze the distribution of 480

GAD1/gad1 and two other awn-related genes. 481

482

ACKNOWLEDGEMENTS 483

We thank the International Rice Research Institute, Chinese Rice Research 484

Institute, Institute of Crop Sciences of Chinese Academy of Agricultural 485

17

Sciences, Guangxi Academy of Agricultural Sciences and Guangdong 486

Academy of Agricultural Sciences for providing the wild and cultivated rice 487

samples. This research was supported by the National Key R & D Program for 488

Crop Breeding (2016YFD0100301) and National Natural Science Foundation 489

of China (Grant 91535301). 490

491

AUTHOR CONTRIBUTIONS 492

C.S. designed and supervised this study. J.J. conducted the characterization493

of the introgression line, map-based cloning, genetic transformations, gene 494

expression analysis, and the evolutionary analysis. X.Z. constructed the 495

introgression lines. X.S. and Y.F. maintained plant materials and performed the 496

field management. L.H., W.Z., L.T., F.L., Z.Z., H.C. and P.G. collected the rice 497

germplasm and phenotypic data. C.S., J.J. and D.X. analyzed the data and 498

wrote the article.499

18

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668

669

24

Figure Legends 670

Figure 1. Phenotypes of wild and cultivated rice. 671

(A) O. rufipogon. A panicle of wild rice is shown in the lower left corner, and the 672

grains of wild rice are illustrated in the lower right corner. Bar = 10 mm 673

(B) O. sativa. A panicle of cultivated rice is shown in the lower left corner, and 674

the grains of cultivated rice are illustrated in the lower right corner. Bar = 10 675

mm. 676

677

Figure 2. Phenotypes of NA93-11 and OIL31. 678

(A) Panicle comparison between NA93-11 and OIL31. Bar = 50 mm. 679

(B) Grains of NA93-11 and OIL31. Bar = 10 mm. 680

(C) Genotype of OIL31. The white regions indicate the NA93-11 genetic 681

background. The blue regions indicate wild rice introgressions. The red ellipse 682

indicates the GAD1 locus. 683

(D) and (E) Grain length comparison between NA93-11 and OIL31. Bar = 10 684

mm. 685

(F) Comparison of grain number per panicle between NA93-11 and OIL31. 686

Sample sizes were n = 20. The statistical significance was set at P < 0.05 687

based on a two-tailed Student’s t test. Error bars represent the SD. 688

689

Figure 3. Map-based cloning of GAD1. 690

(A) The target gene was mapped between the markers M10 and RM5485 on 691

the long arm of chromosome 8. The numbers underneath the bars indicate the 692

number of recombinants between GAD1 and the molecular markers. 693

(B) GAD1 was fine mapped to a ~6-kb region between the markers MX14 and 694

MX16. The numbers underneath the bars indicate the numbers of 695

recombinants between GAD1 and molecular markers. Os08g0485500 was the 696

candidate gene. The arrows indicate gene directions. 697

(C) Gene structure of GAD1, and the different sites between OIL31 and 698

NA93-11. The black bars represent 5′ upstream and 3′ downstream regions 699

25

and introns; rectangles represent exons; and triangles represent insertions. 700

There were three Indels (Indel1, Indel2 and Indel3) and an SNP (SNP1) 701

between OIL31 and NA93-11. 702

(D) The constructs used in the GAD1 functional investigation. pCPL represents 703

the ~4.0-kb W2014 genomic fragment covering the entire GAD1 gene region 704

and the 2.5-kb 5′ upstream and 350-bp 3′ downsteam sequences used for the 705

complementation test. pRNAi denotes the RNAi construct that harbors a 706

350-bp specific region of the GAD1 mRNA. UBI is a maize Ubiquitin promoter. 707

Green areas represent the 5′ and 3′ untranslated regions; red areas represent 708

gene coding regions. 709

(E) Panicle comparison between Nipponbare (Nip) and the transgenic line 710

(CPL-1). Bar = 50 mm. 711

(F) Grain comparison between Nip and the transgenic line (CPL-1). Bar = 10 712

mm. 713

(G) Awn length comparison between Nip and two independent transgenic lines, 714

CPL-1 and CPL-2. 715

(H) The comparison of percentages of awned seeds between Nip and two 716

independent transgenic lines, CPL-1 and CPL-2. 717

(I) and (O) Grain length comparison between Nip and transgenic lines. Bar = 718

10 mm. 719

(J) Panicle comparison between a control plant (TIL189) and a transgenic line 720

(RNAi-1). Bar = 50 mm. 721

(K) Grain comparison between TIL189 and a transgenic line (RNAi-1). Bar = 722

10 mm. 723

(L) Awn length comparison between TIL189 and two independent transgenic 724

lines, RNAi-1 and RNAi-2. 725

(M) The comparison of percentages of awned seeds between TIL189 and two 726

independent transgenic lines, RNAi-1 and RNAi-2. 727

(N) and (P) Grain length comparison between TIL189 and RNAi transgenic 728

lines. Bar = 10 mm. 729

26

(Q) Grain number per panicle comparison between Nip and two independent 730

transgenic lines, CPL-1 and CPL-2. 731

(R) Grain number per panicle comparison between ‘TIL189’ and two 732

independent transgenic lines, RNAi-1 and RNAi-2. 733

Sample sizes were n = 10. The statistical significance was set at P < 0.05 734

based on a two-tailed Student’s t test. Error bars represent the SD. 735

736

Figure 4. The expression patterns and effects of GAD1 on spikelet 737

development. 738

(A) Comparison of GAD1 expression patterns between NA93-11 and OIL31. 739

(B) Comparison of GAD1 expression levels of at the apices of glumes between 740

NA93-11 and OIL31. 741

(C1) to (C5) RNA in situ hybridization of GAD1 during spikelet development in 742

NA93-11. Bar = 100 μm. 743

(D1) to (D5) RNA in situ hybridization of GAD1 during spikelet development in 744

OIL31. Bar = 100 μm. 745

(E1) to (E5) RNA in situ hybridization of HISTONE H4 during spikelet 746

development in NA93-11. Bar = 100 μm. 747

(F1) to (F5) RNA in situ hybridization of HISTONE H4 during spikelet 748

development in OIL31. Bar = 100 μm. 749

(G) RNA in situ hybridization of CKX2 in a young panicle of NA93-11. Bar = 750

100 μm. 751

(H) RNA in situ hybridization of CKX2 in a young panicle of OIL31. Bar = 100 752

μm. 753

(I) RNA in situ hybridization of RR1 in a young panicle of NA93-11. Bar = 100 754

μm. 755

(J) RNA in situ hybridization of RR1 in a young panicle of OIL31. Bar = 100 μm. 756

(K) Comparison of cytokinin concentrations in 5–10 mm-long young panicles 757

between NA93-11 and OIL31. 758

Red arrows (and red dashed lines in E3 to E5 and F3 to F5) indicate the tips of 759

27

lemmas. The statistical significance was set at P < 0.05 based on a two-tailed 760

Student’s t test. Error bars represent the SD. In (A), (B) and (K), the data 761

represent the average of three independent biological replicates. 762

763

Figure 5. GAD1 protein variants in wild and cultivated rice. 764

(A) Differences between GAD1 in W2014 and gad1 in NA93-11. The green 765

rectangles represent the signal peptide. Reddish rectangles represent the 766

pro-peptide. Purple rectangles represent the mature peptide. Red vertical lines 767

represent cysteine residues. Blue rectangles represent the frame-shift region. 768

(B) The distribution of GAD1 protein variants in wild rice. 769

(C) The distribution of gad1 protein variants in cultivated rice. 770

771

Figure 6. Nucleotide diversity across the GAD1 genomic region on 772

chromosome 8. 773

(A) The 16 sampled loci (including GAD1) located in the ~1.9-Mb genomic 774

region around GAD1 gene on chromosome 8. 775

(B) Nucleotide diversity in cultivated rice and wild rice at the 16 sampled loci. 776

(C) Ratio of nucleotide diversity in O. sativa relative to that in O. rufipogon, 777

including the ~900-kb selective sweep region surrounding gad1 in cultivated 778

rice. 779

780

O. rufipogon O. sativa

A B

Figure 1. Phenotypes of wild rice and cultivated rice.

(A) O. rufipogon. A panicle of wild rice is shown in the lower left corner, and the

grains of wild rice are illustrated in the lower right corner. Bar = 10 mm.

(B) O. sativa. A panicle of cultivated rice is shown in the lower left corner, and the

grains of cultivated rice are illustrated in the lower right corner. Bar = 10 mm.

Figure 2. Phenotypes of NA93-11 and OIL31.

(A) Panicle comparison between NA93-11 and OIL31. Bar = 50 mm.

(B) Grains of NA93-11 and OIL31. Bar = 10 mm.

(C) Genotype of OIL31. The white regions indicate the NA93-11 genetic background.

The blue regions indicate wild rice introgressions. The red ellipse indicates the GAD1

locus.

(D) and (E) Grain length comparison between NA93-11 and OIL31. Bar = 10 mm.

(F) Comparison of grain number per panicle between NA93-11 and OIL31.

Sample sizes were n = 20. The statistical significance was set at P < 0.05 based on a

two-tailed Student’s t test. Error bars represent the SD.

GAD1

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R=23 9 4 11

MX9M62 MX10M67 MX16

2 6 61033

GAD1MX14

2

5 kbMarker

R=

MX12MX15M3

47 2 2

OIL31

NA93-11

Indel1 Indel3SNP1Indel2

ATG TGA

ATG TGA--- GCCTCC G CGCCGCC

awn

awnless5’

5’ 3’

3’

**

Figure 3. Map-based cloning of GAD1.

(A) The target gene was mapped between the markers M10 and RM5485 on the

long arm of chromosome 8. The numbers underneath the bars indicate the number

of recombinants between GAD1 and the molecular markers.

(B) GAD1 was fine mapped to a ~6-kb region between the markers MX14 and

MX16. The numbers underneath the bars indicate the numbers of recombinants

between GAD1 and molecular markers. Os08g0485500 was the candidate gene.

The arrows indicate gene directions.

(C) Gene structure of GAD1, and the different sites between OIL31 and NA93-11.

The black bars represent 5′ upstream and 3′ downstream regions and introns;

rectangles represent exons; and triangles represent insertions. There were three

Indels (Indel1, Indel2 and Indel3) and an SNP (SNP1) between OIL31 and NA93-

11.

(D) The constructs used in the GAD1 functional investigation. pCPL represents the

~4.0-kb W2014 genomic fragment covering the entire GAD1 gene region and the

2.5-kb 5′ upstream and 350-bp 3′ downsteam sequences used for the

complementation test. pRNAi denotes the RNAi construct that harbors a 350-bp

specific region of the GAD1 mRNA. UBI is a maize Ubiquitin promoter. Green

areas represent the 5′ and 3′ untranslated regions; red areas represent gene

coding regions.

(E) Panicle comparison between Nipponbare (Nip) and the transgenic line (CPL-1).

Bar = 50 mm.

(F) Grain comparison between Nip and the transgenic line (CPL-1). Bar = 10 mm.

(G) Awn length comparison between Nip and two independent transgenic lines,

CPL-1 and CPL-2.

(H) The comparison of percentages of awned seeds between Nip and two

independent transgenic lines, CPL-1 and CPL-2.

(I) and (O) Grain length comparison between Nip and transgenic lines. Bar = 10

mm.

(J) Panicle comparison between a control plant (TIL189) and a transgenic line

(RNAi-1). Bar = 50 mm.

(K) Grain comparison between TIL189 and a transgenic line (RNAi-1). Bar = 10

mm.

(L) Awn length comparison between TIL189 and two independent transgenic lines,

RNAi-1 and RNAi-2.

(M) The comparison of percentages of awned seeds between TIL189 and two

independent transgenic lines, RNAi-1 and RNAi-2.

(N) and (P) Grain length comparison between TIL189 and RNAi transgenic lines.

Bar = 10 mm.

(Q) Grain number per panicle comparison between Nip and two independent

transgenic lines, CPL-1 and CPL-2.

(R) Grain number per panicle comparison between ‘TIL189’ and two independent

transgenic lines, RNAi-1 and RNAi-2.

Sample sizes were n = 10. The statistical significance was set at P < 0.05 based on

a two-tailed Student’s t test. Error bars represent the SD.

0

200

400

600

800

1000

1200

Rela

tive e

xp

ressio

n l

evel

NA93-11

OIL31

A

**

** **

** **

0.0

0.1

0.1

0.2

0.2

0.3

0.3

0.4

0.4

tZ iP

CK

s c

on

cen

trati

on

(n

g/g

)

NA93-11

OIL31

K

*

*

CKX2

OIL31

CKX2

NA93-11

RR1

NA93-11

RR1

OIL31

G H

I J

B

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

4.0

NA93-11 OIL31

Rela

tive e

xp

ressio

n l

evel

**

HIS

TO

NE

H4

OIL

31

HIS

TO

NE

H4

NA

93-1

1G

AD

1

OIL

31

GA

D1

NA

93-1

1

C1 C2 C3 C4 C5

D1 D2 D3 D4 D5

E1 E2 E3 E4 E5

F1 F2 F3 F4 F5

Figure 4. The expression patterns and effects of GAD1 on spikelet development.

(A) Comparison of GAD1 expression patterns between NA93-11 and OIL31.

(B) Comparison of GAD1 expression levels of at the apices of glumes between

NA93-11 and OIL31.

(C1) to (C5) RNA in situ hybridization of GAD1 during spikelet development in NA93-

11. Bar = 100 μm.

(D1) to (D5) RNA in situ hybridization of GAD1 during spikelet development in OIL31.

Bar = 100 μm.

(E1) to (E5) RNA in situ hybridization of HISTONE H4 during spikelet development in

NA93-11. Bar = 100 μm.

(F1) to (F5) RNA in situ hybridization of HISTONE H4 during spikelet development in

OIL31. Bar = 100 μm.

(G) RNA in situ hybridization of CKX2 in a young panicle of NA93-11. Bar = 100 μm.

(H) RNA in situ hybridization of CKX2 in a young panicle of OIL31. Bar = 100 μm.

(I) RNA in situ hybridization of RR1 in a young panicle of NA93-11. Bar = 100 μm.

(J) RNA in situ hybridization of RR1 in a young panicle of OIL31. Bar = 100 μm.

(K) Comparison of cytokinin concentrations in 5–10 mm-long young panicles

between NA93-11 and OIL31.

Red arrows (and red dashed lines in E3 to E5 and F3 to F5) indicate the tips of

lemmas. The statistical significance was set at P < 0.05 based on a two-tailed

Student’s t test. Error bars represent the SD. In (A), (B) and (K), the data represent

the average of three independent biological replicates.

B

A

Awnless

AwnAwnless

Awn

AwnlessAwn

Cultivated rice

(n=137)(n=23)

(n=43)

Cysteine number =7

Cysteine number =6

Cysteine number =4

(n=203)

Cysteine number =7

Cysteine number =6

Cysteine number =4

(n=53)

0

10

20

30

40

50

60

Cysteine number=4

Cysteine number=6

Cysteine number=7

Wild rice

InDel1 InDel3SNP1InDel2

W2014

NA93-11

Signal peptide Pro-peptide Mature peptide

A - - E

E frame shift

C1 C2 C3 C4 C5 C6

C5’ C6’ C7

C

C1 C2 C3 C4

- AS

Nu

mb

er

of

acce

ssio

ns

Figure 5. GAD1 protein variants in wild and cultivated rice.

(A) Differences between GAD1 in W2014 and gad1 in NA93-11. The green rectangles

represent the signal peptide. Reddish rectangles represent the pro-peptide. Purple

rectangles represent the mature peptide. Red vertical lines represent cysteine residues.

Blue rectangles represent the frame-shift region.

(B) The distribution of GAD1 protein variants in wild rice.

(C) The distribution of gad1 protein variants in cultivated rice.

Physical position (Mb)

0

0.02

π

0

0.5

1

1.5

πra

tio

（O

. sati

va/

O.

rufi

po

go

n)

23.4 23.5 23.6 23.7 23.8 23.9 24.0 24.1 24.2 24.3 24.4 24.5 24.6 24.7 24.8 24.9

.

site 1

. . . . . . .. . . . . . . .

2 3 4 5 6 7 8 9 10 11 1312 1514GAD1

Very low diversity in O. sativa

A

O. sativa

O. rufipogon

B

C

Figure 6. Nucleotide diversity across the GAD1 genomic region on chromosome 8.

(A) The 16 sampled loci (including GAD1) located in the ~1.9-Mb genomic region

around GAD1 gene on chromosome 8.

(B) Nucleotide diversity in cultivated rice and wild rice at the 16 sampled loci.

(C) Ratio of nucleotide diversity in O. sativa relative to that in O. rufipogon, including

the ~900-kb selective sweep region surrounding gad1 in cultivated rice.

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http://www.ncbi.nlm.nih.gov/pubmed?term=Abrash%2C E%2EB%2E%2C and Bergmann%2C D%2EC%2E %282010%29%2E Regional specification of stomatal production by the putative ligand CHALLAH%2E Development 137%3A 447%2D455%2E&dopt=abstract

http://search.crossref.org/?page=1&rows=2&q=Abrash, E.B., and Bergmann, D.C. (2010). Regional specification of stomatal production by the putative ligand CHALLAH. Development 137: 447-455.

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

http://scholar.google.com/scholar?as_q=Regional specification of stomatal production by the putative ligand CHALLAH&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://search.crossref.org/?page=1&rows=2&q=Abrash, E.B., Dacies K.A., and Bergmann, D.C. (2011). Generation of signaling specificity in Arabidopsis by spatially restricted buffering of ligand-receptor interactions. Plant Cell 23: 2864-2879.

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

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DOI 10.1105/tpc.16.00379; originally published online September 15, 2016;Plant Cell

Hongwei Cai, Xianyou Sun, Ping Gu, Daoxin Xie and Chuanqing SunJing Jin, Lei Hua, Zuofeng Zhu, Lubin Tan, Xinhui Zhao, Weifeng Zhang, Fengxia Liu, Yongcai Fu,

Development in Rice DomesticationGAD1 Encodes a Secreted Peptide That Regulates Grain Number, Grain Length and Awn

This information is current as of May 29, 2020

Supplemental Data /content/suppl/2016/09/13/tpc.16.00379.DC1.html

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gad1 encodes a secreted peptide that regulates grain ...plants contain 46 several signal peptides,...

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