1 Running Title: Characterisation of …...2019/04/19 · 6 144 under SDs and LDs. The subsequent...

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Running Title: Characterisation of CENTRORADIALIS in barley 1

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Corresponding Author: Maria von Korff, Heinrich Heine University, c/o Max Planck Institute 4

for Plant Breeding Research, Carl-von-Linné-Weg 10, 50829 Cologne, Germany. Phone 5

+492215062247, email: [email protected] 6

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Research Area: Genes, Development and Evolution 8

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Plant Physiology Preview. Published on April 19, 2019, as DOI:10.1104/pp.18.01454

Copyright 2019 by the American Society of Plant Biologists

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CENTRORADIALIS interacts with FLOWERING LOCUS T-like genes to control spikelet 13

initiation, floret development and grain number 14

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Xiaojing Bi, Wilma van Esse, Mohamed Aman Mulki, Gwendolyn Kirschner, Jinshun Zhong, 16

Rüdiger Simon, Maria von Korff* 17

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Max Planck Institute for Plant Breeding Research, D-50829, Cologne, Germany (M.K., X.B., 19

M.A.M., J.Z.); Institute of Plant Genetics, Heinrich-Heine-University, 40225 Düsseldorf, 20

Germany (M.K., X.B., J.Z. ); Institute for Developmental Genetics, Heinrich-Heine-21

University, 40225 Düsseldorf, Germany (G.K., R.S.), Cluster of Excellence on Plant Sciences 22

“SMART Plants for Tomorrow’s Needs” 40225 Düsseldorf, Germany (G.K., R.S., M.K.); 23

Laboratory of Molecular Biology, Wageningen University and Research, Droevendaalsesteeg 24

1, 6708 PB Wageningen, the Netherlands (W.E.). 25

*Address correspondence: [email protected] 26

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Authors’ Contribution 29

M.K. conceived the original research project. X.B., W.E., M.A.M. and M.K. designed the 30

experiments. X.B. carried out the experiments and analysed the data. G.K. and R.S. conducted 31

the in situ RNA hybridization. J.Z. contributed to the bioinformatic analyses. X. B. and M.K 32

wrote the manuscript, M.K., W.E., R.S. and J.Z. edited the manuscript. 33

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Summary: CENTRORADIALIS interacts with FLOWERING LOCUS T-like genes to 36

control spikelet initiation, floret development and grain number in barley. 37

Funding information 38

This research was funded by the Deutsche Forschungsgemeinschaft (DFG, German Research 39

Foundation) under Germany´s Excellence Strategy – EXC 2048/1 – 390686111, the Priority 40

Programme (SPP1530 Flowering time control- from natural variation to crop improvement) 41

and the Max Planck Society. X. B. received a fellowship from the CSC (Chinese Scholarship 42

Council) 43

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Corresponding Author: Maria von Korff. [email protected] 45

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mailto:[email protected]


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Abstract 47

CENTRORADIALIS (CEN) is a key regulator of flowering time and inflorescence architecture in 48

plants. Natural variation in the barley homolog HvCEN was important for agricultural range expansion 49

of barley cultivation. However, its effects on shoot and spike architecture and consequently yield have 50

not yet been characterised. We evaluated 23 independent hvcen, also termed mat-c, mutants to 51

determine the pleiotropic effects of HvCEN on developmental timing and shoot and spike 52

morphologies of barley under outdoor and controlled conditions. All hvcen mutants flowered early and 53

showed a reduction in spikelet number per spike, tiller number and yield in the outdoor experiments. 54

Mutations in hvcen accelerated spikelet initiation and reduced axillary bud number in a photoperiod 55

independent manner, but promoted floret development only under long days (LDs). The analysis of an 56

hvcen hvft3 double mutant showed that HvCEN interacts with FLOWERING LOCUS T3 (HvFT3) to 57

control spikelet initiation. Furthermore, hvcen hvelf3 (EARLY FLOWERING 3) double mutants with 58

high HvFT1 expression levels under short days (SDs) suggested that HvCEN interacts with HvFT1 to 59

repress floral development. Global transcriptome profiling in developing shoot apices and 60

inflorescences of mutant and wild-type plants revealed that HvCEN controlled transcripts involved in 61

chromatin remodeling activities, cytokinin and cell cycle regulation and cellular respiration under LDs 62

and SDs, while HvCEN affected floral homeotic genes only under LDs. Understanding the stage and 63

organ specific functions of HvCEN and downstream molecular networks will allow manipulating 64

different shoot and spike traits and thereby yield. 65

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Introduction 71

Shoot architecture is a major determinant of grain yield and therefore a primary target for crop 72

improvement. Shoot architecture is largely defined by branching (tillering) patterns, plant height, leaf 73

shape and arrangement, and inflorescence morphologies. These traits are controlled by the combined 74

activities of the shoot apical meristem (SAM) and the axillary meristems (AXMs) (Nakagawa et al., 75

2002; Teichmann and Muhr, 2015). A vegetative SAM gives rise to leaves and AXMs that form in the 76

leaf axils (Turnbull, 2005). As plants transition from vegetative to reproductive growth, the SAM 77

forms an inflorescence, flowers and eventually seeds. Grasses exhibit a striking diversity in their 78

inflorescence architectures that is determined by meristem initiation and determinacy decisions, the 79

acquisition of spikelet meristem identity and the determinacy of the spikelet meristem (Bommert and 80

Whipple 2018). The spikes of barley display a raceme-like branchless shape and consist of triple 81

spikelets produced on two opposite sides along the main axis (rachis). While the barley inflorescence 82

is indeterminate, the barley spikelet is determinate as a defined number of florets is produced, a 83

maximum of a single floret and grain, per spikelet. The triple spikelet meristem (TSM) of barley 84

consists of one central spikelet meristem (CSM) and two lateral spikelet meristems (LSM). Most 85

studies on barley spike architecture have focused on the genetic basis of two-rowed versus six-rowed 86

spikes which is determined by differential development of the lateral spikelets (Komatsuda et al., 87

2007; Koppolu et al., 2013; Ramsay et al., 2011; van Esse et al., 2017; Bull et al., 2017; Youssef et al., 88

2016). However, recent studies have shown that spike architecture is also affected by genetic factors 89

controlling the timing of pre-anthesis development in barley and wheat. The flowering time regulators 90

PHOTOPERIOD1 (PPD-H1) and its downstream target FLOWERING LOCUS T1 (FT1), homolog of 91

Arabidopsis FT, affect the number of spikelets produced on the main inflorescence in barley and 92

wheat, likely by affecting the rate and duration of spikelet initiation (Digel et al., 2015, Boden et al., 93

2015). FT-like genes belong to phosphatidylethanolamine-binding proteins (PEBP), whose homolog in 94

humans was characterised as a Raf kinase inhibitor protein (RKIP), mediating the RAF/MEK/ERK 95

signal transduction pathway (Kobayashi, 1999; Kardailsky, 1999; Bradley et al., 1996, 1997; Yeung et 96

al., 1999; Ohshima et al., 1997). In plants, the PEBP family comprises proteins with antagonistic 97

effects on development as they either promote or inhibit floral transition. FT-like genes generally 98

induce flowering while CENTRORADIALIS (CEN), first described in Antirrhinum, represses the 99

initiation of floral meristems (Digel et al., 2015; Bradley et al., 1996; Danilevskaya et al., 2007; 100

Komiya et al., 2008; Kaneko-Suzuki et al., 2018; Turck et al., 2008). In Arabidopsis and rice, FT is 101

expressed in the leaf vascular tissue and the FT protein is transported through the phloem to the SAM 102

(Corbesier et al., 2007; Tamaki et al., 2007). In rice, the FT homolog HD3A forms a florigen-103

activating complex with 14-3-3 and FLOWERING LOCUS D (FD) homolog OsFD1 to activate the 104

expression of meristem identity genes (Taoka et al., 2011). By contrast, TERMINAL FLOWER1 105

(TFL1) the Arabidopsis homolog of CEN functions as a hypothetical competitor of FT in binding to 106

FD and 14-3-3 proteins in the shoot apex, thereby preventing the induction of flowering (Ahn et al., 107



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2006; Hanano and Goto, 2011; McGarry and Ayre, 2012; Wigge et al., 2005; Abe, 2005; Kaneko-108

Suzuki et al., 2018). An acceleration in flowering time and the formation of a terminal flower were 109

observed in both tfl1 mutants and plants overexpressing FT in Arabidopsis (Bradley et al., 1997; 110

Kardailsky, 1999; Kobayashi, 1999). The function of TFL1 homologs in controlling flowering time 111

and inflorescence architecture are conserved to some extent between grasses and eudicots. For 112

example, the Antirrhinum TFL1 homolog CENTRORADIALIS controls the determinacy of the 113

inflorescence with no effects on flowering time (Bradley et al., 1996), whereas LpTFL1 in ryegrass 114

(Lolium perenne) represses flowering and controls axillary meristem identity (Jensen et al., 2001). 115

RCN1 and RCN2, rice homologs of TFL1, delay flowering and alter the panicle architecture 116

(Nakagawa et al., 2002). Likewise, ectopic expression of the TFL1 homologs ZCN2, ZCN4, ZCN5 in 117

maize leads to late flowering and a bushy tassel with denser spikelets (Danilevskaya et al., 2010). 118

HvCEN, the barley homolog of Antirrhinum CEN and Arabidopsis TFL1, was found to contribute to 119

the expansion of barley cultivation into diverse habitats (Comadran et al., 2012). A natural Pro135Ala 120

substitution in the HvCEN protein has been selected in spring barley cultivars and prolongs vegetative 121

growth, while hvcen mutations lead to early flowering under natural long-day conditions (Comadran et 122

al., 2012). The authors could also show that induced hvcen mutants, originally designated as maturity-123

c (mat-c) mutants, flowered a few days earlier under natural long-day conditions (Druka et al., 2011, 124

Lundqvist, 2014, Comadran et al. 2012). However, the effects of HvCEN on shoot and inflorescence 125

architecture and its interaction with FT-like genes have not been characterised so far. 126

Barley has six different FT-like homologs, of which only HvFT1 and HvFT3 were functionally 127

characterised (Halliwell et al., 2016; Casao et al., 2011; Yan et al., 2006; Schmitz et al., 2000; 128

Hemming et al., 2008; Sasani et al., 2009; Chen and Dubcovsky, 2012). Elevated HvFT1 expression in 129

leaves is correlated with a strong acceleration of floral development, while HvFT3 only induces 130

spikelet initiation without effects on later floret development (Mulki et al., 2018; Digel et al., 2015). In 131

this study, we analysed a large collection of independent hvcen mutants to: 1) identify pleiotropic 132

effects of HvCEN on developmental timing and shoot and spike morphology, 2) determine 133

transcriptional targets of HvCEN in the developing SAM under different photoperiods and 3) 134

investigate the genetic interactions between HvCEN and the FT-like genes HvFT1 and HvFT3. 135

We demonstrate that HvCEN has pleiotropic effects on several shoot traits, as it delays reproductive 136

development and flowering, promotes axillary bud initiation/tillering, and increases the number of 137

spikelet primordia and plant height. Mutations in HvCEN shortened the vegetative phase under both 138

LDs and SDs but accelerated inflorescence development only under LDs. These photoperiod-specific 139

effects of HvCEN were likely dependent on antagonistic interactions with different HvFT-like proteins 140

during development. Global transcriptome analysis in developing shoot apical meristems suggested 141

that spikelet initiation as promoted by mutations in hvcen coincided with a strong reprogramming of 142

transcriptional networks, the induction of cell proliferation and changes in the energy metabolism 143



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under SDs and LDs. The subsequent rapid floral development in the hvcen mutant correlated with the 144

LD specific upregulation of floral homeotic genes. 145

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Results 147

HvCEN controls shoot and spike architecture in barley 148

To characterise the effects of different HvCEN mutations on shoot development, we analysed 149

flowering time, grains per spike, plant height, tiller number and different seed parameters in 23 allelic 150

hvcen mutants in six different spring barley backgrounds under outdoor conditions (Figure 1, Table 151

S1). All the hvcen mutants flowered significantly earlier and produced fewer grains per main spike 152

than the respective wild-type genotypes (Figure 1A, B). In addition, all hvcen mutants, except for mat-153

c.19 and mat-c.913, produced fewer tillers at flowering time compared to the corresponding parental 154

genotypes (Figure 1C). Likewise, plant height of most hvcen mutants, excluding mat-c.19, mat-c.32, 155

mat-c.93 in the Bonus background and all mutants in the Frida background, was reduced compared to 156

the respective wild-type genotypes (Figure 1D). Seed traits, including length, width, area and thousand 157

kernel weight (TKW) were not significantly different between all mutants versus wild-type plants 158

(Figure S1). 159

We calculated the degree of amino acid conservation across taxa for individual mutations. The 160

analysis revealed that all single amino acid substitutions are located in conserved positions within the 161

protein (Table S2). The mutations were positioned in the potential ligand-binding pocket (mat-c.913, 162

mat-c.93, mat-c.32, mat-c.907, mat-c.1115), the 14-3-3 protein interaction site (mat-c.943, mat-c.745) 163

and the external loop (mat-c.400) (Figure S2, Ahn et al. 2006; Ho and Weigel 2014). The external 164

loop has been shown to represent the most substantial differences between FT and TFL1 and is critical 165

for FT versus TFL1 activity in vivo (Ahn et al. 2006). The binding pocket has previously been 166

suggested to play an important role in binding to phosphorylated interacting partners (Ahn et al. 2006; 167

Ho and Weigel 2014) while the 14-3-3 protein interaction site is important for the formation of the 168

flowering activation complex (Taoka et al. 2011). All analysed mutants showed a significant 169

difference in the scored developmental traits, suggesting that these all altered the function of the 170

protein. 171

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To further illustrate the effects of HvCEN on reproductive development, we focused on the 173

development of primary shoots of three selected hvcen mutants (mat-c.907, mat-c.94, and mat-c.943) 174

and cv.Bonus under different photoperiods and compared the timing of spikelet initiation and 175

inflorescence development between the hvcen mutants and the wild-type parent Bonus (Figure 2). The 176

hvcen mutants initiated spikelet primordia (W2.0) significantly earlier under both LDs and SDs 177

compared to the wildtype cv. Bonus (Figure 3A, 3B). However, hvcen mutants exhibited photoperiod-178

specific patterns of growth and floret development. Particularly, under LDs, floret development was 179



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greatly accelerated in hvcen mutants starting from the stamen primordium stage (W3.5) (Figure 3A). 180

Consequently, the hvcen mutant plants flowered (W9.0-10.0) about 18 days earlier than Bonus under 181

LDs (Figure 3A). In contrast, floret development under SDs did not proceed beyond the stages W4.0-182

5.5 in the mutants and the stagesW3.5-W4.5 in Bonus (Figure 2B, 3B). 183

Since hvcen mutants displayed a reduction in grain number per main spike (Figure 1 B), we also 184

evaluated the number of spikelet primordia initiated on the main shoot spike during development. 185

Under LDs, the number of initiated spikelet primordia reached its maximal level at different 186

developmental stages, in the mutant at W4.0 with 24.40± 1.58 spikelet primordia and in the wild type 187

at W5.0 with 40.00 ± 0.76 primordia (Figure 3C). Although the mutants developed fewer spikelet 188

primordia, we did not observe the formation of a terminal spikelet as has been described for the 189

Arabidopsis tfl1 mutant and is typical for the determinate wheat inflorescence (Figure S3). Under SDs, 190

the reduction in the number of spikelet primordia of hvcen mutants was only apparent prior to the 191

lemma primordium stage (W3.0) and no differences were observed between the mutants and wild type 192

after this stage (Figure 3D). Furthermore, hvcen mutants produced fewer and shorter leaves on the 193

main shoot (Figure S4) and exhibited a reduced number of axillary buds under both LDs and SDs 194

(Figure 3E, 3F). No significant differences in leaf width were observed in the mutants compared to 195

wildtype except for mat-c.943 with wider leaves under LDs (Figure S4). 196

Taken together, mutations in the three hvcen (mat-c) mutants accelerated the initiation of spikelet 197

primordia under both LDs and SDs, while floret development of the mutants was only accelerated 198

under LDs. Moreover, hvcen mutants exhibited a reduced number of spikelet primordia on the main 199

shoot apex (MSA). Finally, HvCEN affected the total number of leaves and leaf size on the main culm 200

and tiller number. 201

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HvCEN interacts with HvFT1 and HvFT3 to control reproductive development under LDs and 203

SDs 204

The HvCEN homolog TFL1 acts antagonistically to FT in the meristem and the relative abundance of 205

FT and TFL1 proteins controls floral development and shoot architecture in Arabidopsis and other 206

crop plants (McGarry and Ayre, 2012; Kaneko-Suzuki et al., 2018). Therefore, we tested if and how 207

the effects of HvCEN on development are dependent on the function of FT-like genes in barley. 208

Barley has several FT-like genes (Halliwell et al., 2016), however, only HvFT1 and HvFT3 have been 209

functionally analysed and integrated into flowering pathways. HvFT1 is only transcribed under LDs 210

and its protein promotes spikelet initiation and floret development (Yan et al., 2006, Digel et al. 2015). 211

In contrast, HvFT3 is expressed under SDs and LDs and specifically accelerates the timing of spikelet 212

initiation but has no effects on floret development (Mulki et al., 2018). To test whether the effects of 213

HvCEN on spikelet initiation are dependent on HvFT3, we examined the development of single and 214



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double hvft3 hvcen mutants under SDs. Both genotypes, the hvft3 HvCEN single and the hvft3 hvcen 215

double mutants, did not differ in development and architecture, both showed a delayed development, 216

and an increase in the number of tillers and spikelet primordia (Figure 4A-C). Consequently, only in 217

the presence of a functional HvFT3 did the hvcen allele advances spikelet initiation, and decreases the 218

number of spikelet primordia and axillary buds. We, therefore, inferred that the photoperiod-219

independent effects of HvCEN on spikelet initiation are dependent on HvFT3. However, there was no 220

clear effect of HvCEN on the expression of HvFT3 (Figure S5), suggesting that both interact on the 221

protein level. 222

Next, we examined if the LD-specific effects on floret development could be explained by the 223

interaction of HvCEN with HvFT1 as HvFT1 is only expressed under LDs, but not SDs. For this 224

purpose, we crossed the hvcen mutant with a mutant line carrying a non-functional EARLY 225

FLOWERING 3 (HvELF3) allele. Arabidopsis ELF3 is a circadian clock gene that modulates light 226

signal transduction downstream of phytochromes and mediates the circadian gating of light perception 227

and responses (Hicks et al., 1996; Zagotta et al., 1996; Liu et al., 2001). Barley hvelf3 mutants are 228

characterised by photoperiod independent expression of HvFT1 and early flowering (Faure et al. 2012). 229

We confirmed that expression levels of HvFT1 were comparable between HvELF3 wild-type plants 230

grown under LDs and hvelf3 mutant plants grown under SDs. (Figure S6). MSA development was 231

examined in single and double hvelf3 hvcen mutants under SDs. The plants carrying hvelf3 mutations 232

developed significantly faster when compared to those carrying the wild-type HvELF3 allele 233

irrespective of the HvCEN allele (Figure 4D). More interestingly, variation at HvCEN strongly 234

repressed floral development under SDs in the background of the hvelf3 mutant but had only a minor 235

effect on floral development in HvELF3 wild-type plants. This suggested that under conditions where 236

HvFT1 is expressed, either under LDs or in the hvelf3 mutant background, HvCEN genetically 237

interacted with HvFT1 to modulate floral development. The mutation in hvcen reduced the number of 238

spikelets per MSA and axillary buds in the background of hvelf3 as it did in the HvELF3 background 239

under LDs (Figure 4E, 4F). The genotype and photoperiod specific expression patterns suggested that 240

the photoperiod dependent effects of hvcen on floret development and spikelet number was likely 241

regulated via HvFT1. However, mutations in the clock gene HvELF3 modify the expression of a large 242

number of genes (Faure et al. 2012), we therefore cannot rule out that the hvelf3 specific effect of 243

hvcen might be caused by genes other than HvFT1. 244

Molecular characterisation of HvCEN 245

To further characterise the function of HvCEN, we examined the spatial expression patterns of HvCEN 246

in the main shoot apex and crown tissue of cv. Bonus and cv. Bowman by RNA in-situ hybridization 247

at the spikelet initiation and stamen primordium stages. HvCEN RNA was localized in the axillary 248

meristems and leaf axils, but no signals were detected in the inflorescence meristems (Figure 5A-D, 249

Figure S7). To test if HvCEN was also expressed in the inflorescence but at levels too low for 250

detection by in situ hybridization, we dissected the inflorescence meristems only and tested for 251



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HvCEN expression by qRT-PCR. HvCEN mRNA was expressed in the inflorescence at both spikelet 252

initiation (W2.0) and stamen primordium stages (W3.5) with a higher level at spikelet initiation 253

(Figure 5E). These results indicate that HvCEN is expressed in both the axillary meristems and leaf 254

axils as well as the main inflorescence. 255

To further understand how HvCEN regulates the development of MSA in a photoperiod-dependent 256

and independent manner, we performed genome-wide transcriptome profiling in developing MSA in 257

two allelic hvcen mutants (mat-c.907 and mat-c.943) and the wild type (cv. Bonus). We focused on 258

three developmental stages during which genotypes exhibited phenotypic differences under LDs and 259

SDs (Figure 3, S4), including the vegetative stage (W1.0, MSA enriched tissue), the spikelet initiation 260

stage (W2.0, MSA enriched tissue) and the stamen primordium stage (W3.5, MSA tissue). 261

Transcriptome analysis revealed the expression of 24703 and 25037 transcripts, 62.2% and 63.0% of 262

the total number of annotated transcripts in barley (Mascher et al., 2017), at levels greater than 5 263

counts in at least 2 libraries under LDs and SDs, respectively (Table S10). Principle component 264

analysis on all expressed genes clustered the samples according to the developmental stage under LDs 265

and SDs and separated wild-type and mutant samples at all stages under SDs, but not under LDs 266

(Figure S8). To determine differentially expressed transcripts (DETs), we performed individual 267

pairwise comparisons between each mutant (two allelic mutants) with the wildtype background Bonus 268

at each developmental stage (three stages) within each photoperiod treatment (LDs and SDs), yielding 269

12 sets of DETs. A large number of DETs were identified at the spikelet initiation stage (W2.0), with 270

3177 DETs in mat-c.907, 2876 DETs in mat-c.943 under LDs and 5200 DETs in mat-c.907, 5585 271

DETs in mat-c.943 under SDs. At the vegetative stage (W1.0), only 50 DETs in mat-c.907, 419 DETs 272

in mat-c.943 under LDs and under SDs 14 DETs in mat-c.907, 17 DETs in mat-c.943 were found. At 273

the stamen primordium stage (W3.5), we identified 503 DETs in mat-c.907 and 477 DETs in mat-274

c.943 under LDs and 281 DETs in mat-c.907, 489 DETs in mat-c.943 under SDs (Figure S9). 275

Differences in the number of genes that were affected in the two allelic hvcen mutants may be due to 276

other background mutations in the mutants. Variant calling revealed that 12 and 78 transcripts contain 277

mutations in the mutants mat-c.907 and mat-c.943, respectively, compared to Bonus. Among them, 278

only one transcript (HvCEN) carried different mutations in both mutants (Table S3). In order to 279

minimize possible effects of other background mutations, we focused in the further analyses on 280

transcripts that were differentially regulated in both hvcen mutants, with 33, 1926 and 229 DETs under 281

LDs and 7, 4310 and 117 DETs under SDs at W1.0, W2.0 and W3.5, respectively (Figure S9). Most 282

DETs under both LDs and SDs were observed at W2.0, which corresponded to the developmental 283

stage where the highest expression of HvCEN was observed in the MSA (Figure 6A, 8A, S9). 284

Hierarchical cluster analysis separated the developmental stages on the first principal component (PC) 285

and the photoperiods on the second PC. Only samples harvested at W2.0, were separated for mutants 286

and wild type (Figure 6B, S8C). The majority of genes showed a photoperiod specific regulation. At 287



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W2.0 approximately 73% of the DETs, while at W3.5 92% of the DETs were photoperiod specific 288

(Figure 6A). 289

Taken together, HvCEN had the strongest effect on gene expression at spikelet initiation, specifically 290

under SDs. In addition, the majority of transcripts showed a photoperiod and stage specific regulation 291

in the mutant and the wild-type plants. 292

Transcripts regulated at the spikelet initiation stage correlate with an early transition to 293

reproductive growth in the hvcen mutants 294

Since spikelet initiation was accelerated in the mutants under both photoperiods, we first focused on 295

all transcripts that were regulated at W2.0 independent of the photoperiod. Gene ontology enrichment 296

analysis suggested that HvCEN primarily affected transcripts involved in chromatin modification, 297

ribosome biogenesis, response to cytokinin, cell proliferation and metabolic and biosynthetic 298

processes (Table S4). 299

Among the genes with roles in chromatin modification (selected genes in Table S5), we observed, for 300

instance, the upregulation of two homologs of Arabidopsis MULTICOPY SUPRESSOR OF IRA1 301

(AtMSI1, HORVU5Hr1G084160 (Figure 7A) and HORVU5Hr1G093230). MSI1 is involved in de 302

novo nucleosome assembly during DNA replication, SAM organization and promotes floral transition 303

in Arabidopsis by inducing the expression of CONSTANS (CO) and SOC1 (Bouveret et al., 2006; 304

Steinbach and Hennig, 2014). In addition, the expression of homologs of PROTEIN ARGININE 305

METHYLTRANSFERASES (AtPRMT5, HORVU6Hr1G019540 (Figure 7A), and AtPRMT10, 306

HORVU7Hr1G020620) was increased in the mutants compared to wild type. In Arabidopsis, PRMT 307

genes promote flowering by mediating the epigenetic silencing of FLOWERING LOCUS C (AtFLC) 308

and by affecting pre-mRNA splicing (Deng et al., 2010; Schmitz et al., 2008). A putative target of 309

epigenetic factors and repressor of flowering, a homolog of FLC, HvODDSOC2 310

(HORVU3Hr1G095240), was downregulated in the mutants. In addition, we observed a higher 311

expression of a homolog of ARABIDOPSIS TRITHORAX-RELATED PROTEIN 6 (ATXR6) 312

(HORVU6Hr1G011950, Figure 7A), which encodes a SET-domain protein, a H3K27 313

monomethyltransferases required for chromatin structure and gene silencing (Jacob et al. 2008). 314

Among the upregulated genes were also ribosomal proteins (selected genes in Table S5) with 315

functions in inflorescence development, vascular patterning and adaxial cell fate, such as two 316

homologs of PIGGYBACK 2 (PGY2, HORVU0Hr1G006020, Figure 7B; HORVU3Hr1G001140), and 317

a homolog of PGY3 (HORVU5Hr1G092630). Further, a nucleolar GTPase NUCLEOSTEMIN-LIKE 1 318

(AtNSN1) like transcript (HORVU2Hr1G016650, Figure 7B), which plays a role in the maintenance of 319

inflorescence meristem identity and floral organ development by modulating ribosome biogenesis in 320

Arabidopsis (Wang et al., 2012; Jeon et al., 2015), was upregulated in the mutants. Furthermore, we 321

found DETs with putative roles in cytokinin response and cell cycle regulation (Table S5). For 322

example, we observed the upregulation of a barley 26S proteasome non-ATPase regulatory subunit 8 323



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homolog A (RPN12a, HORVU4Hr1G002140, Figure 7C), which regulates cytokinin responses by 324

upregulating type A ARRs, which are in turn negative regulators of cytokinin signaling (Ryu et al., 325

2009). In addition, barley homologs of type A and B RESPONSE REGULATORS (ARR2, 326

HORVU5Hr1G097560.5; ARR3, HORVU2Hr1G077000, HORVU3Hr1G108540; ARR6, 327

HORVU2Hr1G120490, Figure 7C) that act as regulators in the two-component cytokinin signaling 328

pathway were upregulated. In contrast, barley homologs of histidine kinases and putative cytokinin 329

receptors, HISTIDINE KINASE 3 and HISTIDINE KINASE 4 (AHK3, HORVU3Hr1G094870, Figure 330

7C; AHK4, HORVU6Hr1G077070) were downregulated in the mutants compared to wild-type plants. 331

Alterations in the expression of cytokinin response genes were accompanied by the upregulation in the 332

expression of genes involved in cell division. These included, for example, homologs of CELL 333

DIVISION CONTROL 6 (CDC6, HORVU3Hr1G084800, Figure 7B), PROLIFERATING CELL 334

NUCLEAR ANTIGEN 2 (PCNA2, HORVU6Hr1G088120, HORVU0Hr1G031140) and REPLICON 335

PROTEIN A2 (HORVU6Hr1G094080). A faster transition to a reproductive MSA and induction of 336

cell cycle genes coincided with the differential expression of genes involved in cellular respiration, 337

including glycolysis, pyruvate metabolism and citrate cycle (Table S5). For example, transcripts with 338

roles in glycolysis and carbohydrate metabolism were upregulated in the mutant compared to wild 339

type, i.e. CELL WALL INVERTASE 2 (HORVU2Hr1G073210, Figure 7D), a RAFFINOSE 340

SYNTHASE family protein (HORVU7Hr1G027930), and a TREHALOSE-6-PHOSPHATASE (TPS1, 341

HORVU1Hr1G013450). Further proteins with roles in glycolysis, such as a PYRUVATE KINASE 342

family protein (HORVU3Hr1G039200), three homologs of FRUCTOSE-BISPHOSPHATE 343

ALDOLASE 2 (HORVU3Hr1G002780, Figure 7D; HORVU3Hr1G088540, HORVU3Hr1G088570) 344

and three homologs of ATP-dependent 6-PHOSPHOFRUCTOKINASES (PFK2, 345

HORVU2Hr1G081670; PFK3, HORVU3Hr1G070300; PFK7, HORVU6Hr1G070270, Figure 7D) 346

were upregulated in the mutant versus wild-type plants. The upregulation of genes involved in cellular 347

respiration might be a consequence of changes in the source sink balance and a stronger energy 348

demand required for the accelerated vegetative to reproductive stage transitions in the mutant plants 349

compared to the wild type. Taken together, HvCEN had the strongest molecular effects on the MSA 350

transcriptome at spikelet initiation and these involved genes with functions in chromatin remodeling 351

activities, cytokinin and cell cycle regulation and cellular respiration independent of the photoperiod. 352

HvCEN controls floral homeotic genes under LDs 353

Spikelet initiation was advanced in the mutants under both LDs and SDs, but florets only developed 354

and set seeds under LDs, while the MSAs were aborted under SDs. To gain a better understanding of 355

the photoperiod-dependent effects of HvCEN on floret development we analysed the photoperiod-356

dependent transcripts at the stamen primordium stage (W3.5). The photoperiod specific DETs, 357

including 179 LD-specific and 83 SD-specific DETs, were enriched for transcripts with functions in 358

reproductive processes, response to stimuli and cell communication (Table S6). Among the 179 LD 359

specific DETs at W3.5, 50 DETs were upregulated and 129 DETs downregulated in the mutants 360



12

compared to wild type. Among the transcripts (selected genes in table S7) that were upregulated to a 361

higher extent in the mutants compared to wild type only under LD were HvCEN 362

(HORVU2Hr1G072750, Figure 8A) and flowering promoting genes such as barley homologs of SOC1 363

(HORVU1Hr1G051660, Figure 8A), AGAMOUS-LIKE6 (AGL6, HORVU6Hr1G066140), and 364

FLOWERING PROMOTING FACTOR 1 (FPF1, HORVU2Hr1G007350, Figure 8A). In addition, we 365

observed the LD dependent upregulation of transcription factors that act in a combinatorial manner to 366

achieve floral patterning in Arabidopsis (Coen and Meyerowitz, 1991). These are designated as class 367

A, B, C and E genes and, except for the class A gene AP2, encode members of the MIKC type of 368

MADS-box transcription factors. A class A protein, AP1-like (HvBM8, HORVU2Hr1G063800, 369

FDR<0.05, Figure 8C, table S8) and a class B like gene, PISTILLATA (PI)-like 370

(HORVU1Hr1G063620, FDR<0.05, Figure 8C, table S8), were upregulated in the mutant lines under 371

LDs, but not SDs. In addition, the mutations in HvCEN caused an upregulation of five E class genes, 372

the SEP-like genes (SEP1, HORVU7Hr1G025700, HORVU5Hr1G095710, FDR<0.05, SEP2, 373

HORVU4Hr1G067680, SEP3, HORVU7Hr1G054220, HORVU5Hr1G076400, FDR<0.05) at the 374

stamen primordium stage (Table S8, Figure 8B). An ABERRANT PANICLE ORGANIZATION 375

(APO1)-like transcript (HORVU7Hr1G108970) was downregulated in the mutants compared to wild 376

type. Interestingly, the APO1 protein positively controls spikelet number by suppressing the 377

precocious conversion of inflorescence meristems to spikelet meristems in rice (Ikeda et al. 2007). In 378

addition, barley homologs of a SWEET sucrose transporter (HORVU5Hr1G076770, Figure 8C), of a 379

GLYCOGEN SYNTHASE (HORVU2Hr1G106410) and a TREHALOSE-6-PHOSPHATE 380

PHOSPHATASE (HORVU6Hr1G074960) were upregulated specifically at W3.5 in the mutants 381

compared to the wild type. Likewise, a barley homolog of a SHAGGY-related kinase 382

(HORVU3Hr1G034440) required for the establishment of tissue patterning and cell fate determination 383

and a KNOTTED1-like homeobox gene (HORVU7Hr1G114650) involved in meristem differentiation 384

were upregulated in the mutants at W3.5 under LDs. 385

Under SD, 83 DETs were detected at W3.5, 58 among them were downregulated and 25 upregulated 386

in the mutants compared to wild type. Among the upregulated genes (selected genes in Table S7), we 387

identified a number of stress related genes involved in detoxification such as a stress responsive A/B 388

Barrel Domain protein (HORVU0Hr1G011450, Figure 8D) and a homolog of ACYL-COA-BINDING 389

PROTEIN 6 (AtACBP6, HORVU7Hr1G008320, Figure 8D). Furthermore, we recorded the mutant 390

specific upregulation of transcripts with roles in cellular transport, such as a homolog of CYCLIC 391

NUCLEOTIDE-GATED ION CHANNEL2 (AtCNGC2, HORVU5Hr1G096440, Figure 8D), building 392

nonselective cation channels and of PLASMA MEMBERANE INTRINSIC PROTEIN 3 (AtPIP3, 393

HORVU5Hr1G125600) forming water channels (Figure 8C). Finally, proteins with roles in starch and 394

sugar metabolism, such as a TREHALOSE-6-PHOSPHATE PHOSPHATASE (TPPH, 395

HORVU5Hr1G058300), a SUCROSE SYNTHASE (HORVU7Hr1G033230) and a UDP-GLUCOSE 396



13

4-EPIMERASE (HORVU7Hr1G053260) showed a stronger upregulation in the wild type than the 397

mutants. 398

Taken together, the LD specific expression patterns of floral homeotic genes with putative functions in 399

inflorescence, spikelet and flower development correlated with the differential floret development of 400

mutants and wild type under LDs. Under SDs, the hvcen mutant lines were characterised by the 401

differential regulation of genes implicated in abiotic stress responses, in cellular transport, and 402

carbohydrate metabolism. 403

404

405



14

Discussion 406

In the present work, we analysed induced hvcen mutants to dissect the effects of HvCEN on spike 407

development and plant architecture under different photoperiods and to identify potential molecular 408

targets of HvCEN in the MSA. Mutations in HvCEN accelerated spikelet initiation under LDs and 409

SDs while subsequent floral development required LDs. Therefore, HvCEN showed stage and 410

photoperiod-dependent effects on the development of the MSA. Previous studies have already 411

indicated that spikelet initiation occurs under LDs and SDs, while floral development requires LDs in 412

spring barley genotypes (Digel et al., 2015). Floral development, but not spikelet initiation requires the 413

expression of HvFT1 and TaFT in barley and wheat, respectively (Digel et al., 2015; Pearce et al., 414

2013). Spikelet initiation occurs in the absence of FT1 expression likely promoted by FT3, however, 415

the timing of spikelet initiation is affected by variation in FT1 expression (Digel et al., 2015; Dixon et 416

al., 2018). 417

In Arabidopsis, TFL1 acts antagonistically to the FT protein to repress floral transition and 418

development (Ruiz-García et al., 1997; Jaeger et al., 2013). In addition, rice TFL1-like proteins, RICE 419

CENTRORADIALIS (RCN), compete with the rice FT ortholog Hd3a for the binding activities of the 420

14-3-3 protein, thereby antagonizing the activities of the florigen (Kaneko-Suzuki et al., 2018). 421

Therefore, we also tested if HvCEN genetically interacts with HvFT1 and HvFT3 that affect different 422

stages of pre-anthesis development (Mulki et al., 2018; Halliwell et al., 2016; Casao et al., 2011; Yan 423

et al., 2006). While HvFT1 primarily accelerates floral development under long days (Hemming et al., 424

2008; Sasani et al., 2009), HvFT3 controls spikelet initiation under long and short days (Mulki et al., 425

2018). The double mutant hvcen hvft3 did not differ in the timing of spikelet initiation under SDs from 426

the HvCEN hvft3 line, suggesting that the repressive effect of HvCEN on the timing of spikelet 427

initiation depends on a functional HvFT3 gene. In contrast to HvFT3, HvFT1 is only expressed under 428

LDs and HvFT1 expression is crucial for floral development and flowering (Digel et al. 2015). 429

Consequently, we tested if the LD specific effect of HvCEN on floral development was dependent on 430

HvFT1 expression. For this purpose, we analysed an HvCEN hvelf3 single and hvcen hvelf3 double 431

mutant that both expressed HvFT1 under SDs to similar levels as seen under LDs. The strong delay in 432

inflorescence development and higher number of the spikelets in the HvCEN hvelf3 single mutant as 433

compared to the hvcen hvelf3 double mutant under SDs indicated that the repressive effect of HvCEN 434

on floral development depended on HvFT1 expression in the hvelf3 mutant. In addition, the interaction 435

between HvCEN and FT1 had a strong effect on the determination of spikelet primordia number. 436

However, as many light-dependent transcripts are misregulated in the hvelf3 mutant (Faure et al., 437

2012), we cannot rule out that other genes influenced the observed phenotype. Our results suggested 438

that HvCEN genetically interacted with different FT-like genes in the shoot apical meristem to control 439

different phases of inflorescence development: with HvFT3 to control spikelet initiation; and with 440

HvFT1 to control floral development. The photoperiod specific effects of HvCEN are therefore likely 441

caused by the photoperiod specific expression of its putative competitors HvFT3 and HvFT1. 442



15

443

444

The hvcen mutants were altered in different shoot architecture traits including the number of leaves on 445

the main culm, leaf length, the number of tillers per plant and the number of seeds per spike. The 446

mutant plants developed 1-2 fewer leaves on the main culm under long and short-day conditions, 447

possibly as a consequence of the earlier transition from a vegetative to a reproductive meristem. Since 448

AXMs initiate in the leaf axils a reduction in the number of leaves may have been causal for the 449

reduction in tiller number observed in the mutant lines. In addition, the leaves in the hvcen mutants 450

were shorter indicating that leaf size is controlled by HvCEN-dependent progression of plant 451

development. It was already demonstrated before that flowering time genes may affect leaf size in 452

barley by affecting the duration of leaf growth and consequent variation in leaf cell number (Digel et 453

al. 2016). The earlier termination of leaf growth in the hvcen mutants was matched by an earlier 454

termination of spikelet induction. However, the barley hvcen mutants did not form a terminal spikelet 455

as has been described for the Arabidopsis tfl1 mutant. (Shannon and Ry Meeks-Wagner, 1991). 456

Nevertheless, the mutation in hvcen reduced the period of spikelet primordia initiation and thereby 457

decreased the number of spikelets on the MSA. The wild type reached the maximal number of 458

spikelets at awn primordium stage as observed in previous studies (Kirby and Appleyard, 1987; 459

Alqudah and Schnurbusch, 2014; Riggs and Kirby, 1978; Kernich et al., 1997; Waddington et al., 460

1983), while the hvcen (mat-c) mutants stopped making further spikelets at the pistil primordium stage 461

under inductive LD conditions. It has already been shown before that variation in the developmental 462

timing of the early reproductive phase has strong effects on the number of spikelet primordia on the 463

MSA (Digel et al., 2015; Ejaz and von Korff, 2017; Campoli and von Korff, 2014). For instance, in 464

the presence of a wild-type PPD-H1 allele the development of the MSA is accelerated and this is 465

associated with a reduction in number of spikelet primordia and final grain number per main spike 466

(Digel et al., 2015). In wheat, Alvarez et al. (2016) observed an acceleration of flowering time and an 467

associated reduction in spikelet number in an elf3 mutant. Both, PPD-H1 and HvELF3 likely affect 468

developmental timing and spikelet number by inducing or repressing HvFT1, respectively. Taken 469

together, the pleiotropic phenotypes of the hvcen mutants suggested that changes in the 470

HvFT1/HvCEN ratios may control meristematic activity or its termination in different meristems of 471

the barley shoot as has been proposed for tomato (Lifschitz et al., 2014; Krieger et al., 2010; Park et al., 472

2014; Jiang et al., 2013). However, the molecular basis for these coordinated effects on meristems 473

remain poorly understood. 474

In Arabidopsis, TFL1 mRNA is expressed in young axillary meristems but later becomes restricted to 475

the central inflorescence meristem (Conti and Bradley, 2007; Wigge et al., 2005; Shannon and Ry 476

Meeks-Wagner, 1991; Bradley et al., 1997). By contrast, HvCEN mRNA signals were detected 477

primarily in the AXMs and expression levels were low in the shoot apex despite the strong effect of 478

mutations in HvCEN on developmental timing and spike morphology, i.e. the number of spikelets and 479



16

seeds per spike. Similarly, expression of the HvCEN homologs RCN1-4 in rice was not detected in the 480

SAM, but in the vasculature of the subtending stem and young primordia and leaf blade (Kaneko-481

Suzuki et al., 2018). However, the authors could demonstrate that the RCN protein is transported 482

through the phloem to the inflorescence meristem where it controls phase transition and inflorescence 483

determinacy. Consequently, HvCEN protein might also travel to the inflorescence meristem where it 484

acts on spike development. Accordingly, the largest number of differentially transcribed genes was 485

detected at the spikelet initiation stage when the SAM switches from vegetative to reproductive 486

growth. This acceleration of spikelet initiation in the hvcen mutants under LDs and SDs coincided 487

with the differential regulation of transcripts associated with chromatin modifications. Epigenetic 488

genes putatively upstream of floral regulators included, for example, MSI1, which is part of the 489

evolutionarily conserved Polycomb group (PcG) chromatin-remodeling complex and controls spatial 490

and temporal expression of several homeotic genes that regulate plant development and organ identity 491

(Steinbach and Hennig, 2014; Derkacheva et al., 2013; Chanvivattana, 2004; Hennig et al., 2005). 492

Similarly PRMT-like genes were upregulated in the mutants and these control the epigenetic silencing 493

of the floral repressor FLC and flowering time in Arabidopsis (Niu et al., 2007; Schmitz et al., 2008; 494

Wang et al., 2007; Pei et al., 2007). The upregulation of PRMT-like genes was correlated with a 495

downregulation of a putative repressor of flowering and homolog of FLC, HvODDSOC2, suggesting 496

that its expression might also be controlled epigenetically. Further, we recorded a strong upregulation 497

of Trithorax like proteins that act as H3K27 methyltransferases required for transcriptional repression 498

in Arabidopsis (Jacob et al., 2009). HvCEN dependent regulation of epigenetic regulators in the MSA 499

of barley specifically at spikelet initiation suggested that the transition from vegetative to reproductive 500

meristem requires strong reprogramming of transcriptional networks by epigenetic modifiers. These 501

modifiers and their role for developmental transitions in barley await further functional 502

characterisation. 503

Further, we observed the upregulation of genes involved in cell cycle regulation, cytokinin signalling 504

and response and many ribosomal proteins that are thought to control cellular growth (Bhavsar et al., 505

2010; Naora and Naora, 1999; Schaller et al., 2014). The upregulation of ribosomal proteins in the 506

mutants was specific for the spikelet initiation stage suggesting that the transition from a vegetative to 507

a reproductive meristem requires a strong increase in translational efficiency and de novo protein 508

synthesis. However, several of the ribosomal proteins upregulated in the mutants may have more 509

specific effects on organ development as these are also described as important regulators of leaf and 510

inflorescence development, vascular patterning and phase change, such as RPS13-like and PGY-like 511

genes (Pinon et al., 2008; Ito et al., 2000). Spikelet initiation also coincided with the upregulation of 512

genes involved in carbon metabolism, glycolysis, cellular respiration and tricarboxylic acid cycle. 513

These transcripts may be important to prepare the plant for the subsequent fast inflorescence growth 514

and increased energy demand. Ghiglione et al. (2008) have demonstrated that fast growing wheat 515

inflorescences show strongly reduced soluble carbohydrate levels as compared to slowly developing 516



17

spikes and suggested that these fast growing tissues suffer from carbohydrate starvation. Consequently, 517

the upregulation of cellular respiration genes was possibly important to allow for a higher energy 518

supply to the developing organs in the fast growing mutants. 519

In contrast to spikelet initiation, floral development was photoperiod dependent and the majority of 520

transcripts in the MSA were regulated by HvCEN only under LDs. In particular, floral homeotic genes 521

were primarily regulated under LD conditions by HvCEN and these included homologs of floral 522

patterning transcription factors designated as class A, B, and C genes (for review, see Theißen, 2001). 523

The mutations in HvCEN caused an upregulation of five E class, SEP-like, genes, a class B gene (PI, 524

OsMADS4) and one AP1-like gene (HvBM8) at the stamen primordium stage. In rice, the SEP-like 525

gene OsMADS34 (PANICLE PHYTOMER2) is important for controlling inflorescence and spikelet 526

development (Gao et al., 2010; Kobayashi et al., 2010), while the other SEP-like genes OsMADS1, 527

OsMADS5, OsMADS7, OsMADS8 , and OsMADS34 control the development of different floral 528

organs (Malcomber and Kellogg, 2005; Zahn et al., 2005; Arora et al., 2007). Interestingly, the barley 529

homolog of OSMADS34 was regulated by HvCEN under both photoperiods, while the other SEP-like 530

genes were controlled by HvCEN only under LDs. This suggested that genes important for floral 531

development but not those involved in spikelet initiation were LD dependent. The repression of these 532

genes by HvCEN is not only essential to delay floral development but also to prolong leaf initiation 533

and leaf maturation probably as a strategy to match vegetative and reproductive development of the 534

plant. 535



18

Material and methods 536

Germplasms of hvcen (praematurum-c, mat-c), hvelf3 hvcen and hvft3 hvcen double mutants 537

All mutants and parental lines were obtained from the Nordic Gene Bank (NordGen; 538

http://www.nordgen.org/). The 21 allelic hvcen mutants were originally generated using different 539

mutagens in various barley spring cultivars (wild-type parents): Bonus, Foma, Frida, Kristina and 540

Semira (Matyszczak, 2014; Comadran et al., 2012; Franckowiak and Lundqvist, 2012; Table S1, 541

Figure S2). The different mutants were characterised by distinct changes in the gene body including 542

premature stop codons, changes in splice sites, amino acid replacement, frameshift and whole gene 543

deletions (Comadran et al. 2012). The mutation in mat-c.770 introduced a stop codon, resulting in a 544

truncated protein. Mutations in mat-c.94, mat-c.1111 and mat-c.1114 caused splice site changes. 545

Amino-acid substitution at conserved sites were detected in mat-c.32, mat-c.770, mat-c.907, mat-c.943, 546

mat-c.913, mat-c.93, mat-c.400 and mat-c.1115 (Table S2). The mat-c.1109 genotype carries a 1-bp 547

deletion that resulted in a frame shift. Sanger Sequencing of HvCEN in mat-c.1118 revealed a 12-bp 548

deletion (GATGCAAACAAT) including 2-bp from the end of the 4th exon and 10-bp from the UTR 549

region (10-bp) leading to a larger protein consisting of 229 instead of 173 amino acids (Figure S2). 550

The mat-c.16, mat-c.19, mat-c.1096, mat-c.1107, mat-c.1102, mat-c.1108, mat-c.1120 mutants are 551

putative deletion mutants as HvCEN could not be amplified in these genotypes. In addition, we 552

analysed two backcross derived introgression lines BW508, with an introgression of mat-c.19 and 553

BW507 with an introgression of mat-b.7 in the background of Bowman (Druka et al., 2011). BW507 554

likely contains a large deletion of HvCEN since no amplicons of this gene were found (Matyszczak, 555

2014). The effect of a single amino acid substitution in the point mutants were evaluated by 556

PROVEAN (Protein Variation Effect Analyzer, http://provean.jcvi.org) that computationally predicts 557

the influence of alternations in amino acids on the biological function of the protein (Table S2). The 558

program assesses the functional effects of protein variation using an alignment-based score (PROVEN 559

score) that measures the change in sequence similarity between a query sequence and its variants to a 560

homologous protein sequence (Choi et al. 2012). 561

To determine if HvCEN interacts with HvFT1 and HvFT3 genetically, we produced hvelf3 hvcen 562

(with HvFT1 expression under SDs) and hvft3 hvcen double mutants. The hvelf3 hvcen double mutants 563

were derived from the cross of the hvcen mutant in Bonus (mat-c.907) with the hvelf3 mutant in Bonus 564

(mat-a.8, NGB110008). Three F2 progenies verified as homozygous hvelf3 hvcen double and two as 565

hvelf3 HvCEN single mutants were propagated, respectively. F4 plants from these selected five lines 566

were grown and dissected in a controlled climate chamber under SD conditions (8h/16h, light/dark, 567

20°C/18°C). 568

To obtain hvft3 hvcen double mutants, the hvcen mutant (mat-c.907) in the Bonus background was 569

crossed to an introgressed line carrying a natural mutation in hvft3 in the background of Golden 570

Promise. This introgression line was an F3 progeny derived from crosses between the winter barley 571


http://www.nordgen.org/

http://provean.jcvi.org/


19

Igri and the spring cultivar Golden Promise. Specifically, the introgression line carried a non-572

functional hvft3 allele from Igri and the natural mutation at Ppd-H1, a deletion in the first regulatory 573

intron of VRN1 and a deletion of the VRN2 locus from Golden Promise. This introgression line shows 574

a reduced photoperiod response and does not require vernalization. By genotyping, three F2 lines 575

carrying homozygous hvft3 hvcen double mutations and two hvft3 HvCEN progenies were identified 576

and propagated. 577

Plant growth conditions and phenotyping in outdoor condition and climate chambers 578

We evaluated all mutants and their parental lines (Table S1) under outdoor conditions over two 579

consecutive years for the number of tillers and spikelets per main spike, plant height and flowering 580

time. Flowering time was scored as the time from sowing to heading, when the awns emerge from the 581

flag leaf of the main culm. Tiller number was recorded at heading, while the number of grains per 582

main spike and plant height (soil to base of topmost spike) were measured at full maturity, two weeks 583

before harvest. The numbers of mat-c mutants and wild-type plants grown in each year are indicated in 584

table S1. 585

In parallel, we conducted phenotyping in environment-controlled growth chambers using selected 586

hvcen mutants. Specifically, three hvcen mutants in the Bonus background (mat-c.907, mat-c.94 and 587

mat-c.943) and Bonus were grown in 96-well trays using “Mini Tray” (Einheitserde®) soil. To 588

synchronize germination, the trays were stratified in dark at 4°C for three days followed by growth 589

under LDs (16h, 22°C day; 8h, 18°C night) or SDs (8h, 22°C day; 16h, 18°C night). The 590

developmental stage of the MSA was determined using the Waddington quantitative scale of shoot 591

apex development that is based on the progression of spike initiation and then the most advanced floret 592

primordium and pistil of the inflorescence (Waddington et al., 1983). Five to six individual plants for 593

each focal accession were scored for their MSA development every 3 days and the number of spikelet 594

primordia per main spike, the number of axillary buds and leaf primordia number were scored during 595

the dissection. The axillary buds scored include all the axillary buds, primary, secondary and higher 596

order buds. Each leaf were dissected to see the axillary bud under the leaf sheath. Leaf size and visible 597

leaf number on the main culm were scored in 20-well trays as described by Digel et.al (2016). 598

The hvelf3 hvcen, the hvft3 hvcen double mutants and their control or parental lines were scored for 599

differences in pre-anthesis development, number of spikelet primordia per main spike and number of 600

axillary buds including all the axillary buds, primary, secondary and higher order buds by dissecting 601

the lines every two/four days (hvelf3 hvcen) or weekly (hvft3 hvcen) in 96-well trays under SDs. 602

Statistical analysis 603

We conducted a one-way ANOVA followed by a posthoc Tukey HSD test to test for differences in 604

flowering time, plant height, tiller number and number of seeds per main spike of the mutants 605

compared to their respective parental lines. In addition, we conducted a one-way ANOVA to compare 606



20

HvFT3 expression in HvCEN wild-type and hvcen mutant backgrounds. In the dissection experiments 607

differences between the parental lines, the mutant and double mutant lines were revealed by 608

calculating a polynomial regression model at a 95% confidence interval (Loess smooth line). 609

RNA isolation and sample preparation for RNA sequencing 610

Total mRNA was isolated from plants grown under LDs and SDs. MSA enriched tissues were 611

harvested and pooled for three distinct developmental stages, the vegetative stage (W1.0), the spikelet 612

initiation stage (W2.0) and the anther primordium stage (W3.5) (Waddington et al., 1983). The 613

samples were harvested 2 hours before dark under both LD and SD conditions. To enrich shoot apex 614

specific mRNA, leaves surrounding the MSA were removed manually using a microsurgical stab knife 615

(5-mm blade at 15° [SSC#72-1551]). The enriched MSA tissue was cut from the base of the MSA and 616

still included leaf primordia. At least 10 MSAs were pooled for each of the three biological replicates 617

per time point. All samples harvested for RNA extraction were frozen immediately in liquid nitrogen 618

and stored at -80°C. Total mRNA was extracted using TRIzol reagent (Invitrogen, Carlsbad, CA, USA) 619

and further purified using an RNA easy Micro Kit (Qiagen). The residual DNA was removed using a 620

DNA-free kit (Ambion) and the quality of the RNA was assessed using a bioanalyzer (Agilent 2100 621

Bioanalyzer). The Illumina cDNA libraries were prepared according to the TruSeq RNA sample 622

preparation (version 2; Illumina). A cBot (Illumina) was used for clonal sequence amplification, and 623

generation of sequence clusters. Single-end sequencing was performed using a HiSeq 3000 (Illumina) 624

platform by multiplexing 8 libraries resulting in ~18 million reads per library. The requested single 625

end read length was 100 bp for LD samples and 150bp for SD samples. The initial quality control of 626

the raw reads was performed using the FastQC software (version0.10.1; 627

http://www.bioinformatics.bbsrc.ac.uk/projects/fastqc/). 628

Transcriptome profiling and variant calling 629

The obtained RNA sequencing reads were mapped to a barley High Confidence (HC) transcripts 630

reference (Mascher et al., 2017) using Salmon in quasi-mapping-based mode. When building the 631

quasi-mapping-based index, an auxiliary k-mer hash over k-mers of length 31 was used. U (unstranded 632

single end read) was chosen as library type to quantify the reads of each library. The expected number 633

of reads (NumReads) that have originated from each transcript given the structure of the uniquely 634

mapped and multi-mapped reads and the relative abundance estimates for each transcript and 635

transcripts Per Million (TPM) values were extracted using Salmon (Patro et al., 2015). Transcripts 636

with expression levels greater than 5 NumReads in at least two libraries under LDs or SDs were 637

retained. Tables with expected NumReads (raw counts) and expression levels (normalized counts per 638

million, cpm) are provided in a supplementary table (Table S10). 639

To identify differentially expressed transcripts (DETs), pairwise comparisons, including mat-c.907 vs. 640

Bonus, mat-c.943 vs. Bonus at each stage and photoperiod condition were done using the R 641



21

bioconductor package Limma-vroom with a Benjamin & Hochberg adjustment for multiple testing 642

(false discovery rate, FDR) (Ritchie et al., 2015). The comparisons between the stages and between the 643

photoperiods were not conducted to avoid false positive DETs due to differences in the sample types 644

and potential effects of diurnal gene expression differences between photoperiods. The FDR value of 645

0.01 was used as initial cut-off value for the selection of DETs. In the end, the DETs were extracted 646

per mutant per developmental stage per photoperiod. To minimize the background mutation effects in 647

the two hvcen mutants, we focused on the DETs regulated in both mutants compared with wildtype. 648

To visualize the number of genotype or stage or photoperiod dependent DETs, venn diagrams were 649

drawn using the R package VennDiagram (Chen and Boutros, 2011). The DETs that were observed in 650

both mutants were considered as candidate DETs regulated by HvCEN under each condition. Totally, 651

4527 DETs regulated in the two hvcen mutant at least at one stage compared to wildtype under LDs or 652

SDs were obtained. We then examined the expression patterns using hierarchical cluster analysis 653

(using Pearson correlation coefficients) and principle component analysis (PCA) of the DETs with R. 654

To determine which gene categories were enriched under each treatment, we first produced de novo 655

Gene Ontology (GO) annotations for High Confidence (HC) transcripts using Blast2Go local blast (e 656

value cut-off 1 X 10-5) (Conesa et al., 2005). To assess the effect of HvCEN on biological process, the 657

overrepresentation analysis of particular GO terms was performed based on the Fisher's Exact Test 658

(significant cut-off 0.05) using TopGo package in R (Alexa, 2018). The redundant go categories were 659

removed by first filtering annotated >=5 and allowing similarity as 0.5 (Supek et al., 2011). Then the 660

representative go categories were retained by keeping the enriched transcripts number >= 10 and >=5 661

at W2.0 and W3.5, respectively. 662

Variant calling to verify the HvCEN mutation in mat-c.907 and mat-c.943 and evaluate the number of 663

mutated transcripts in the RNA sequencing reads was done as described in van Esse et al. (2017). 664

Briefly, RNA sequencing reads were mapped to a barley High Confidence (HC) CDS reference 665

(Mascher et al., 2017) using BWA-MEM (version 0.7.15;(Li, 2013)), allowing a mismatch penalty of 666

3. Mapping was evaluated using PicardTools (version 1.1.00; http://picard.sourceforge.net) 667

CollectAlignmentSummaryMetrics and SAMtools (version 1.1.3; (Li et al., 2009)) was used to 668

determine the number of reads mapped with good mapping quality scores (MAPQ > 1). Read 669

duplicate removal and indel realingment were done using PicardTools MarkDuplicates and GATK 670

IndelRealigner (version 3.1-1; (McKenna et al., 2010)), respectively. Variant calling was done with 671

GATK UnifiedGenotyper using 30.0 as a minimum confidence threshold and 10.0 for emitting of 672

called SNPs and 1 for ploidy. Filtered variants with a depth of coverage >= 100, a quality of the 673

assigned genotype >= 98 and a value of Phred-scaled likelihood >= 2000 were taken into 674

consideration. Mutation types include SNP and INDEL. The number of mutations and number of 675

mutated transcripts were summarized in Table S3. 676

Gene expression using qRT-PCR and RNA in situ hybridization 677


https://www.broadinstitute.org/gatk/guide/article?id=4260


22

RNA was isolated from leaf tissues harvested from plants grown under LD and SD conditions. Under 678

LD condition, the second youngest leaf on the main shoot of Bonus (HvCEN HvELF3) and mat-c.907 679

(hvcen HvELF3) at W3.5 was harvested at two hours before dark. Under SD condition, the second 680

youngest leaf of three hvcen hvelf3, two HvCEN hvelf3, mat-a.8 (HvCEN hvelf3 -3), mat-c.907 and 681

Bonus were harvested at seven hours after the beginning of the dark period at the four- to five-leaf 682

stage. Total RNA extraction, first-strand cDNA synthesis and quantitative real-time polymerase chain 683

reaction (qRT-PCR) were performed as described in Campoli et al. (2012). The primer for HvFT1 684

expression is listed in table S9. Two technical replicates were used for each cDNA sample and starting 685

amounts for each data point were calculated based on the titration curve for each target gene and the 686

reference (HvActin) gene using the LightCycler 480 Software (Roche; version 1.5). 687

The expression of HvCEN (HORVU2Hr1G072750.4) was detected using RNA in situ hybridization on 688

SAMs of Waddington stages W2, and W3.5 as described in Kirschner et al. (2018). The probes were 689

prepared using the whole gene sequences followed by carbonate hydrolysis to ca. 200 bp fragments. 690

The anti-sense probe was used for detecting HvCEN expression and the sense probe was used for the 691

negative control. 692

Images were taken using a plan-neofluar 10x objective with a NA of 0.30 using the Zeiss Axioskop 693

light microscope, and image processing, i.e. stitching, was performed with the Stitching Plugin in Fiji 694

(Preibisch et al., 2009, Schindelin et al., 2012). 695

Data availability: Illumina data in the European Short Read Archive: E-MTAB-7807 696

697



23

Figure legends 698

Figure 1. Phenotypes of hvcen (mat-c) mutants trialled under outdoor conditions over two 699

consecutive years. Mutants in six different genetic backgrounds are shown next to their respective 700

parental lines (Table S1). Various traits including A) flowering time, B) grain number per main spike, 701

C) number of tillers at flowering time, and D) plant height were measured over two years. Flowering 702

time was scored as time from sowing to heading when the awn of the main spike emerged from the 703

flag leaf. The line within each box denotes the median. Significant differences between the mutants 704

and their wild-type parents were calculated by a one-way ANOVA using a Tukey HSD as post hoc 705

test, p<0.001 “***”, P<0.01 “**”, P<0.05 “*”, not significant: “ns”. 706

Figure 2. Representative shoot apices of the hvcen mutant and the wildtype scored under A) LD 707

and B) SD conditions. Under SDs, the MSA did not develop beyond the stages W3.5 and W4.5 in the 708

mutant and Bonus, respectively. DAG: days after germination; W: Waddington stage; white bar: 1mm. 709

Figure 3. Phenotyping of three hvcen mutants (mat-c.907, mat-c.94, mat-c.943) and the wildtype 710

Bonus under controlled LD and SD conditions. Development of the main shoot apex (MSA), 711

number of spikelet primordia and of axillary buds at different Waddington stages under LDs (A, C, E) 712

and SDs (B, D, F). The number of spikelet primordia is referring to the number of primordia per main 713

spike. The axillary buds scored include all the axillary buds, primary, secondary and higher order buds. 714

Five or six plants per genotype were dissected at each time point under LDs (16 h light/8 h night) and 715

SDs (8h/16h, light/dark). Statistical differences (P<0.05) were calculated using a polynomial 716

regression model at 95% confidence interval (Loess smooth line) shown in grey-shaded regions. 717

Figure 4. Microscopic phenotypes of double and single hvft3 x hvcen (mat-c.907) and hvelf3 x 718

hvcen (mat-c.907) mutants in Bonus under SDs. Development of the MSA, number of spikelet 719

primordia and number of axillary buds in HvFT3 or hvft3 background (A, B, C) and HvELF3 or hvelf3 720

background (C, D, E) at different Waddington stages under SDs (8h/16h, light/dark). The number of 721

spikelet primordia is referring to the number of primordia per main spike. The axillary buds scored 722

include all the axillary buds, primary, secondary and higher order buds. Three to six plants per 723

genotype were dissected at each time point. Statistical differences (P<0.05) were calculated using a 724

polynomial regression model at 95% confidence interval (Loess smooth line) ) shown in grey-shaded 725

regions. 726

Figure 5. Expression pattern of HvCEN under LDs. A, B) RNA in situ hybridization using the anti-727

sense probe C, D) negative controls using the sense probe of HvCEN in the shoot apex of cv. Bowman 728

at Waddington stages 2.0 and 3.5; E) expression of HvCEN in the pure inflorescence meristem of cv. 729

Bonus. Pure meristems were obtained by carefully removing leaf primordia and stem tissue beneath 730

the inflorescence under the microscope. LDs: long day condition (16h/8h light/dark), W: Waddington 731

stage. 732



24

733

Figure 6. DETs regulated between two hvcen mutants (mat-c.907 and mat-c.943) and wildtype 734

(Bonus) in at least one photoperiod condition (LD/SDs) and co-expression patterns of the DETs. 735

A) Number of DETs regulated in both mutants (mat-c.907 and mat-c.943) compared to WT (Bonus) 736

under LDs and/or SDs. The red circles display DETs detected under LDs and blue circles show DETs 737

detected under SDs. 738

B) Heatmap of co-expression clusters for 4527 DETs. Colors represent log2-fold changes (log2-FC) in 739

expression levels relative to the mean transcript abundance across the tested conditions, i.e., apex 740

(enriched) samples of mat-c.907, mat-c.943 and Bonus grown under LD and SD conditions and 741

harvested at different developmental stages (W1.0, W2.0, W3.5). LDs: long day condition (16h/8h, 742

light/dark), SDs: short day condition (8h/16h, light/dark), W: Waddington stage, DETs: differentially 743

expressed transcripts. Transcripts with FDR<0.01 were considered as DETs. 744

Figure 7. Expression profiles of selected DETs between the hvcen mutants (mat-c.907, mat-c.943) 745

and wildtype (Bonus) at W2.0 related to A) chromatin modification, regulation of development and 746

organ initiation; B) ribosomal proteins, leaf development, leaf patterning; C) regulation of hormone 747

level and hormone response D) cellular respiration, sink strength, carbohydrate metabolism, glycolytic 748

processes under both LDs and SDs. LDs and SDs are shown by white and grey colors. LDs: long day 749

condition (16h/8h, light/dark), SDs: short day condition (8h/16h, light/dark), white: light period (16h), 750

grey: dark period (8h), W: Waddington stage, cpm: normalized counts per million, DETs: differential 751

expressed transcripts, error bars: standard deviation. Transcripts with FDR<0.01 were considered as 752

DETs. 753

Figure 8. Expression profiles of selected DETs between the hvcen mutants (mat-c.907, mat-c.943) 754

and wildtype (Bonus) at W3.5 under both LDs and SDs. LDs and SDs are shown by white and grey 755

colors. LDs: long day conditions (16h/8h, light/dark), SDs: short day conditions (8h/16h, light/dark), 756

white: light period (16h), gray: dark period (8h), W: Waddington stage, cpm: normalized counts per 757

million, DETs: differential expressed transcripts, error bars: standard deviation. 758

759

Supplemental Material 760

Figure S1. Seed parameters of hvcen (mat-c) mutants trialled under outdoor condition. A) Seed 761

length, B) seed width, C) seed area and D) thousand kernel weight (TKW) in field conditions in one 762

year, 5 plants per genotype and 5 representative spikes per plant were analysed. The line within each 763

box denotes the median. Significant differences between the mutants and their wild-type parents were 764

calculated by a one-way ANOVA using a Tukey HSD as post hoc test, p<0.001 “***”, P<0.01 “**”, 765

P<0.05 “*”, not significant: “ns”. 766



25

Figure S2. The mutation sites of the 14 mat-c mutants on the schematic structure of the HvCEN gene. 767

The positions of the 11 mutant alleles, including mat-c.32, mat-c.400, mat-c.770, mat-c.907, mat-c.913, 768

mat-c.93, mat-c.94, mat-c.943, mat-c.1109, mat-c.1114 and mat-c.1115, were obtained from 769

Comadran et al. (2012). The positions of mat-c.745 and mat-c.1111 and were described in Matyszczak, 770

2014. Sequencing of HvCEN in mat-c.1118 identified a 12-bp deletion which translated into a protein 771

containing 229 instead of 173 amino acids. Stop indicates introduction of a stop codon; 1-bp and 12-772

bp deletions are represented by “:”; splice site means a single nucleotide polymorphism affecting an 773

intron-exon splice junction. 774

Figure S3. Determinate spike with a terminal spikelet in wheat (Chinese Spring) and indeterminate 775

spike without terminal spikelet in barley (mat-c.907 and Bonus). 776

Figure S4. Leaf length and number of leaves on main culm of hvcen mutants (mat-c.907, mat-777

c.94, mat-c.943) and Bonus under LDs and SDs. A-B) Width of leaves on the main culm of hvcen 778

mutants (mat-c.907, mat-c.94, mat-c.943) and Bonus under LDs and SDs; C) number of visible leaves 779

on the main culm at different Waddington stages of hvcen mutants (mat-c.907, mat-c.943) and Bonus 780

under LDs and SDs. LDs (16h/8h light/dark), SDs (8h/16h, light/dark). The line within each box 781

denotes the median. 782

Figure S5. HvFT3 expression in leaves of Bonus, mat-c.907 and mat-c.943 at W2.0 and W3.5 under 783

SDs. Letters indicate significant differences (p≤ 0.05), determined with a one-way ANOVA using a 784

Tukey HSD as post hoc test. Significant differences between the lines are calculated for each 785

developmental stage separately. W: Waddington stage, SDs: (8h/16h, light/dark). The line within each 786

box denotes the median. 787

Figure S6. HvFT1 expression in leaves. hvelf3 hvcen-1,2,3 and hvelf3 HvCEN-1,2 are progenies of 788

lines selected from the cross (mat-c.907 x mat-a.8); hvelf3 HvCEN-3 is mat-a.8; HvELF3 hvcen is 789

mat-c.907; HvELF3 HvCEN is Bonus. LD: long day (16h/8h, light/dark), SD: (8h/16h, dark/light). The 790

line within each box denotes the median. 791

Figure S7. Expression pattern of HvCEN in cv. Bonus under LDs. A, B) RNA in situ hybridization 792

using anti-sense probe; C, D) negative controls using sense probe of HvCEN in the shoot apex of cv. 793

Bonus at Waddington stages 2.0 and 3.5. LD: long day (16h/8h, light/dark). 794

Figure S8. Principal component analysis (PCA) of normalized expression profiles for all 795

expressed genes under LDs and SDs. A) The first two PCs account for ∼40% of the overall variation 796

under LDs, B) the first two PCs account for 31% of the overall variation under SDs. C) PCA of the 797

4528 DETs in both mutants (mat-c.907 and mat-c.943) compared to Bonus under LDs or/and SDs, 798

FDR>0.01. LD: long day (16h/8h, light/dark), SD: short day (8h/16h, dark/light), W: Waddington 799

stage. 800



26

Figure S9. A) DETs in each mutant (mat-c.907 or mat-c.943) compared with wildtype (Bonus) under 801

each photoperiod (LDs, SDs) at W1.0, W2.0 and W3.5; B) DETs (FDR<0.01) were found in both 802

mutants at W1.0, W2.0 and W3.5 under both LDs and SDs. LD: long day (16h/8h, light/dark), SD: 803

short day (8h/16h, light/dark), W: Waddington stage. 804

Figure S1. Seed parameters of hvcen (mat-c) mutants trialled under outdoor condition. 805

Figure S2. The mutation sites of the 14 mat-c mutants on the schematic structure of the HvCEN gene. 806

Figure S3. Determinate spike with a terminal spikelet in wheat and indeterminate spike without 807

terminal spikelet in barley. 808

Figure S4. Leaf length and number of leaves on main culm of hvcen mutants (mat-c.907, mat-c.94, 809

mat-c.943) and Bonus under LDs and SDs. 810

Figure S5. HvFT3 expression in leaves of Bonus, mat-c.907 and mat-c.943 at W2.0 and W3.5 under 811

SDs. 812

Figure S6. HvFT1 expression in leaves. 813

Figure S7. Expression pattern of HvCEN in cv. Bonus under LDs. 814

Figure S8. Principal component analysis (PCA) of normalized expression profiles for all expressed 815

genes under LDs and SDs. 816

Figure S9. DETs in each mutant compared with wildtype under each photoperiod 817

Table S1. mat-c (hvcen) mutants carrying different type of mutations in HvCEN 818

Table S2. Effects of amino acid substitutions in the mat-c (hvcen) mutants 819

Table S3. Number of transcripts mutated in the two mutants compared to wild type 820

Table S4. Go enrichment analysis of photoperiod independent DETs at W2.0 821

Table S5. Selected* transcripts differentially regulated in the mutant MSA at W2.0 under LDs and 822

SDs (FDR<0.01) 823

Table S6. Go enrichment analysis of photoperiodic specific DETs (FDR<0.01) at W3.5 824

Table S7. Selected photoperiodic independent/dependent DETs at W3.5 (FDR<0.01) 825

Table S8. Selected* DETs involved in floral development and identity at W3.5 at less stringent cut-off 826

(FDR<0.05) 827

Table S9. Primers used in qRT-PCR and genotyping 828

Table S10. Normalized expression value (cpm), log2FC, FDR and annotation of all expressed 829

transcripts 830

831



27

832

Acknowledgments 833

We cordially thank Kerstin Luxa, Caren Dawidson, Thea Rütjes, Andrea Lossow for excellent 834

technical assistance and Artem Pankin for uploading the raw sequencing data to the European 835

Nucleotide Archive. 836

Competing Interests 837

The authors do not have any financial, personal or professional interests that have influenced 838

this present paper. 839

840



28

841

References 842

Abe, M. (2005). FD, a bZIP Protein Mediating Signals from the Floral Pathway Integrator FT at the Shoot Apex. 843 Science. 309: 1052–1056. 844

Ahn, J.H., Miller, D., Winter, V.J., Banfield, M.J., Lee, J.H., Yoo, S.Y., Henz, S.R., Brady, R.L., and 845 Weigel, D. (2006). A divergent external loop confers antagonistic activity on floral regulators FT and 846 TFL1. EMBO J. 25: 605–14. 847

Alexa, A., Rahnenführer, J., and Lengauer, T. (2006). Improved scoring of functional groups from gene 848 expression data by decorrelating GO graph structure. Bioinformatics, 22, pp.1600-1607. 849

Alqudah, A.M. and Schnurbusch, T. (2014). Awn primordium to tipping is the most decisive developmental 850 phase for spikelet survival in barley. Funct. Plant Biol. 41: 424–436. 851

Bhavsar, R.B., Makley, L.N., and Tsonis, P.A. (2010). The other lives of ribosomal proteins. Hum Genomics 4: 852 327–344. 853

Boden, S.A., Cavanagh, C., Cullis, B.R., Ramm, K., Greenwood, J., Jean Finnegan, E., Trevaskis, B., and 854 Swain, S.M. (2015). Ppd-1 is a key regulator of inflorescence architecture and paired spikelet development 855 in wheat. Nat. Plants 1: 1–6. 856

Bommert, P., and Whipple, C. (2018). Grass inflorescence architecture and meristem determinacy. Semin. Cell 857 Dev. Biol. 79: 37–47. 858

Bouveret, R., Schönrock, N., Gruissem, W., and Hennig, L. (2006). Regulation of flowering time by 859 Arabidopsis MSI1. Development 133: 1693-1702 860

Bradley, D., Carpenter, R., Copsey, L., Vincent, C., Rothstein, S., and Coen, E. (1996). Control of 861 inflorescence architecture in Antirrhinum. Nature 379: 791–797. 862

Bradley, D., Ratcliffe, O., Vincent, C., Carpenter, R., and Coen, E. (1997). Inflorescence Commitment and 863 Architecture in Arabidopsis. Science 275: 80–83. 864

Bull, H. et al. (2017). Barley SIX-ROWED SPIKE3 encodes a putative Jumonji C-type H3K9me2/me3 865 demethylase that represses lateral spikelet fertility. Nat. Commun. 8: 936. 866

Campoli, C. and von Korff, M. (2014). Genetic control of reproductive development in temperate cereals. In 867 Advances in botanical research (Elsevier), pp. 131–158. 868

Casao, M.C., Karsai, I., Igartua, E., Gracia, M.P., Veisz, O., and Casas, A.M. (2011). Adaptation of barley 869 to mild winters: A role for PPDH2. BMC Plant Biol. 11. 164. 870

Chanvivattana, Y., Bishopp, A., Schubert, D., Stock, C., Moon, Y. H., Sung, Z. R., and Goodrich, J. (2004). 871 Interaction of Polycomb-group proteins controlling flowering in Arabidopsis. Development, 131: 5263-872 5276. 873

Chen, A. and Dubcovsky, J. (2012). Wheat TILLING mutants show that the vernalization gene VRN1 down-874 regulates the flowering repressor VRN2 in leaves but is not essential for flowering. PLoS Genet. 8: 875 e1003134. 876

Coen, E.S. and Meyerowitz, E.M. (1991). The war of the whorls: genetic interactions controlling flower 877 development. Nature 353: 31–37. 878

Comadran, J. et al. (2012). Natural variation in a homolog of Antirrhinum CENTRORADIALIS contributed to 879 spring growth habit and environmental adaptation in cultivated barley. Nat. Genet. 44: 1388–1392. 880

Conesa, A., Götz, S., García-Gómez, J.M., Terol, J., Talón, M., and Robles, M. (2005). Blast2GO: A 881 universal tool for annotation, visualization and analysis in functional genomics research. Bioinformatics 21: 882 3674–3676. 883

Conti, L. and Bradley, D. (2007). TERMINAL FLOWER1 Is a Mobile Signal Controlling Arabidopsis 884 Architecture. Plant Cell 19: 767–778. 885

Corbesier, L., Vincent, C., Jang, S., Fornara, F., Fan, Q., Searle, I., Giakountis, A., Farrona, S., Gissot, L., 886



29

Turnbull, C., and Coupland, G. (2007). FT Protein Movement Contributes to Long-Distance Signaling 887 in Floral Induction of Arabidopsis. Science 316: 1030–1033. 888

Danilevskaya, O.N., Meng, X., and Ananiev, E. V (2010). Concerted modification of flowering time and 889 inflorescence architecture by ectopic expression of TFL1-like genes in maize. Plant Physiol. 153: 238–251. 890

Danilevskaya, O.N., Meng, X., Hou, Z., Ananiev, E. V., and Simmons, C.R. (2007). A Genomic and 891 Expression Compendium of the Expanded PEBP Gene Family from Maize. Plant Physiol. 146: 250–264. 892

Deng, X., Gu, L., Liu, C., Lu, T., Lu, F., Lu, Z., Cui, P., Pei, Y., Wang, B., Hu, S., and Cao, X. (2010). 893 Arginine methylation mediated by the Arabidopsis homolog of PRMT5 is essential for proper pre-mRNA 894 splicing. Proc. Natl. Acad. Sci. 107: 19114–19119. 895

Derkacheva, M., Steinbach, Y., Wildhaber, T., Mozgová, I., Mahrez, W., Nanni, P., Bischof, S., Gruissem, 896 W., and Hennig, L. (2013). Arabidopsis MSI1 connects LHP1 to PRC2 complexes. EMBO J. 32: 2073–897 2085. 898

Digel, B., Pankin, A., and von Korff, M. (2015). Global Transcriptome Profiling of Developing Leaf and Shoot 899 Apices Reveals Distinct Genetic and Environmental Control of Floral Transition and Inflorescence 900 Development in Barley. Plant Cell 27: 2318-2334 901

Digel, B., Tavakol, E., Verderio, G., Tondelli, A., Xu, X., Cattivelli, L., Rossini, L. and von Korff, M. 902 (2016). Photoperiod1 (Ppd-H1) controls leaf size. Plant physiology 172: 405-415. 903

Dixon, L.E., Farré, A., Finnegan, E.J., Orford, S., Griffiths, S. and Boden, S.A. (2018). Developmental 904 responses of bread wheat to changes in ambient temperature following deletion of a locus that includes 905 FLOWERING LOCUS T1. Plant, cell & environment 41: 1715-1725. 906

Ejaz, M. and von Korff, M. (2017). The Genetic Control of Reproductive Development under High Ambient 907 Temperature. Plant Physiol. 173: 294–306. 908

van Esse, G.W., Walla, A., Finke, A., Koornneef, M., Pecinka, A., and von Korff, M. (2017). Six-Rowed 909 Spike3 (VRS3) Is a Histone Demethylase That Controls Lateral Spikelet Development in Barley. Plant 910 Physiol. 174: 2397–2408. 911

Faure, S., Turner, A.S., Gruszka, D., Christodoulou, V., Davis, S.J., von Korff, M., and Laurie, D.A. 912 (2012). Mutation at the circadian clock gene EARLY MATURITY 8 adapts domesticated barley 913 (Hordeum vulgare) to short growing seasons. Proc. Natl. Acad. Sci. 109: 8328–8333. 914

Franckowiak, J.D. and Lundqvist, U. (2012). Descriptions of Barley Genetic Stocks for 2012. Barley Genet. 915 Newsl. 42: 36–173. 916

Gao, X., Liang, W., Yin, C., Ji, S., Wang, H., Su, X., Guo, C., Kong, H., Xue, H., and Zhang, D. (2010). The 917 SEPALLATA-Like Gene OsMADS34 Is Required for Rice Inflorescence and Spikelet Development. Plant 918 Physiol. 153: 728–740. 919

Halliwell, J., Borrill, P., Gordon, A., Kowalczyk, R., Pagano, M.L., Saccomanno, B., Bentley, A.R., Uauy, 920 C., and Cockram, J. (2016). Systematic Investigation of FLOWERING LOCUS T-Like Poaceae Gene 921 Families Identifies the Short-Day Expressed Flowering Pathway Gene, TaFT3 in Wheat (Triticum 922 aestivum L.). Front. Plant Sci. 7: 1–15. 923

Hanano, S. and Goto, K. (2011). Arabidopsis TERMINAL FLOWER1 Is Involved in the Regulation of 924 Flowering Time and Inflorescence Development through Transcriptional Repression. Plant Cell 23: 3172–925 3184. 926

Hemming, M.N., Peacock, W.J., Dennis, E.S., and Trevaskis, B. (2008). Low-Temperature and Daylength 927 Cues Are Integrated to Regulate FLOWERING LOCUS T in Barley. Plant Physiol. 147: 355–366. 928

Hennig, L., Bouveret, R., and Gruissem, W. (2005). MSI1-like proteins: An escort service for chromatin 929 assembly and remodeling complexes. Trends Cell Biol. 15: 295–302. 930

Hicks, K.A., Millar, A.J., Carre, I.A., Somers, D.E., Straume, M., Meeks-Wagner, R., and Kay, S.A. (1996). 931 Conditional circadian dysfuncion of the Arabidopsisearly-flowering 3 mutant. Science 274: 790–792. 932

Ito, T., Kim, G.T., and Shinozaki, K. (2000). Disruption of an Arabidopsis cytoplasmic ribosomal protein S13-933 homologous gene by transposon-mediated mutagenesis causes aberrant growth and development. Plant J. 934 22: 257–264. 935



30

Jacob, Y., Feng, S., LeBlanc, C.A., Bernatavichute, Y. V., Stroud, H., Cokus, S., Johnson, L.M., Pellegrini, 936 M., Jacobsen, S.E., and Michaels, S.D. (2009). ATXR5 and ATXR6 are H3K27 monomethyltransferases 937 required for chromatin structure and gene silencing. Nat. Struct. Mol. Biol. 16: 763–768. 938

Jaeger, K.E., Pullen, N., Lamzin, S., Morris, R.J., and Wigge, P.A. (2013). Interlocking Feedback Loops 939 Govern the Dynamic Behavior of the Floral Transition in Arabidopsis. Plant Cell 25: 820–833. 940

Jensen, C.S., Salchert, K., and Nielsen, K.K. (2001). A TERMINAL FLOWER1-like gene from perennial 941 ryegrass involved in floral transition and axillary meristem identity. Plant Physiol. 125: 1517–1528. 942

Jeon, Y., Park, Y.J., Cho, H.K., Jung, H.J., Ahn, T.K., Kang, H., and Pai, H.S. (2015). The nucleolar 943 GTPase nucleostemin-like 1 plays a role in plant growth and senescence by modulating ribosome 944 biogenesis. J. Exp. Bot. 66: 6297–6310. 945

Jiang, K., Liberatore, K.L., Park, S.J., Alvarez, J.P., and Lippman, Z.B. (2013). Tomato Yield Heterosis Is 946 Triggered by a Dosage Sensitivity of the Florigen Pathway That Fine-Tunes Shoot Architecture. PLoS 947 Genet. 9: e1004043 948

Kaneko-Suzuki, M., Kurihara-Ishikawa, R., Okushita-Terakawa, C., Kojima, C., Nagano-Fujiwara, M., 949 Ohki, I., Tsuji, H., Shimamoto, K., and Taoka, K.I. (2018). TFL1-Like Proteins in Rice Antagonize 950 Rice FT-Like Protein in Inflorescence Development by Competition for Complex Formation with 14-3-3 951 and FD. Plant Cell Physiol. 59: 458–468. 952

Kardailsky, I. (1999). Activation Tagging of the Floral Inducer FT. Science 286: 1962–1965. 953

Kernich, G.C., Halloran, G.M., and Flood, R.G. (1997). Variation in duration of pre-anthesis phases of 954 development in barley (Hordeum vulgare). Aust. J. Agric. Res. 48: 59–66. 955

Kirby, E.J.M. and Appleyard, M. (1987). Development and structure of the wheat plant. In: Wheat breeding 956 (Springer), pp. 287–311. 957

Kobayashi, K., Maekawa, M., Miyao, A., Hirochika, H., and Kyozuka, J. (2010). PANICLE PHYTOMER2 958 (PAP2), encoding a SEPALLATA subfamily MADS-box protein, positively controls spikelet meristem 959 identity in rice. Plant Cell Physiol. 51: 47–57. 960

Kobayashi, Y. (1999). A Pair of Related Genes with Antagonistic Roles in Mediating Flowering Signals. 961 Science 286: 1960–1962. 962

Komatsuda, T. et al. (2007). Six-rowed barley originated from a mutation in a homeodomain-leucine zipper I-963 class homeobox gene. Proc. Natl. Acad. Sci. 104: 1424–1429. 964

Komiya, R., Ikegami, A., Tamaki, S., Yokoi, S., and Shimamoto, K. (2008). Hd3a and RFT1 are essential for 965 flowering in rice. Development 135: 767–774. 966

Koppolu, R. et al. (2013). Six-rowed spike4 (Vrs4) controls spikelet determinacy and row-type in barley. Proc. 967 Natl. Acad. Sci. 110: 13198–13203. 968

Krieger, U., Lippman, Z.B., and Zamir, D. (2010). The flowering gene SINGLE FLOWER TRUSS drives 969 heterosis for yield in tomato. Nat. Genet. 42: 459–463. 970

Lifschitz, E., Ayre, B.G., and Eshed, Y. (2014). Florigen and anti-florigen - a systemic mechanism for 971 coordinating growth and termination in flowering plants. Front. Plant Sci. 5: 1–14. 972

Liu, X.L., Covington, M.F., Fankhauser, C., Chory, J., and D. Ry Wagner (2001). ELF3 Encodes a 973 Circadian Clock-Regulated Nuclear Protein That Functions in an Arabidopsis PHYB Signal Transduction 974 Pathway. Plant Cell Online 13: 1293–1304. 975

Mascher, M. et al. (2017). A chromosome conformation capture ordered sequence of the barley genome. Nature 976 544: 427–433. 977

Matyszczak, I. (2014). Characterization of early maturity barley mutants praematurum-a, -b and –c. PhD. Thesis 978 Aarhus University, Department of Molecular Biology and Genetics, Faculty of Science and Technology, 979 Denmark. 980

McGarry, R.C. and Ayre, B.G. (2012). Manipulating plant architecture with members of the CETS gene 981 family. Plant Sci. 188–189: 71–81. 982

Mulki, M.A., Bi, Xi., and von Korff, M. (2018). FLOWERING LOCUS T3 controls spikelet initiation but not 983



31

floral development. Plant Physiol.: pp.00236. 984

Nakagawa M., Shimamoto K., and Kyozuka J. (2002). Overexpression of RCN1and RCN2, rice TERMINAL 985 FLOWER 1/CENTRORADIALIS homologs, confers delay of phase transition and altered panicle 986 morphology in rice. Plant J 29: 743–750 987

Naora, H. and Naora, H. (1999). Involvement of ribosomal proteins in regulating cell growth and apoptosis: 988 Translational modulation or recruitment for extraribosomal activity? Immunol. Cell Biol. 77: 197–205. 989

Niu, L., Lu, F., Pei, Y., Liu, C., and Cao, X. (2007). Regulation of flowering time by the protein arginine 990 methyltransferase AtPRMT10. EMBO Rep. 8: 1190–1195. 991

Park, S.J., Jiang, K., Tal, L., Yichie, Y., Gar, O., Zamir, D., Eshed, Y., and Lippman, Z.B. (2014). 992 Optimization of crop productivity in tomato using induced mutations in the florigen pathway. Nat. Genet. 993 46: 1337–1342. 994

Pearce, S., Vanzetti, L.S., and Dubcovsky, J. (2013). Exogenous Gibberellins Induce Wheat Spike 995 Development under Short Days Only in the Presence of VERNALIZATION1. Plant Physiol. 163: 1433–996 1445. 997

Pei, Y., Niu, L., Lu, F., Liu, C., Zhai, J., Kong, X., and Cao, X. (2007). Mutations in the Type II Protein 998 Arginine Methyltransferase AtPRMT5 Result in Pleiotropic Developmental Defects in Arabidopsis. Plant 999 Physiol. 144: 1913–1923. 1000

Pinon, V., Etchells, J.P., Rossignol, P., Collier, S.A., Arroyo, J.M., Martienssen, R.A., and Byrne, M.E. 1001 (2008). Three PIGGYBACK genes that specifically influence leaf patterning encode ribosomal proteins. 1002 Development 135: 1315–1324. 1003

Ramsay, L., Comadran, J., Druka, A., Marshall, D.F., Thomas, W.T., Macaulay, M., MacKenzie, K., 1004 Simpson, C., Fuller, J., Bonar, N. and Hayes, P.M. (2011). INTERMEDIUM-C, a modifier of lateral 1005 spikelet fertility in barley, is an ortholog of the maize domestication gene TEOSINTE BRANCHED 1. 1006 Nature genetics, 43: 169-173. 1007

Riggs, T.J. and Kirby, E.J.M. (1978). Developmental consequences of two-row and six-row ear type in spring 1008 barley: 1. Genetical analysis and comparison of mature plant characters. J. Agric. Sci. 91: 199–205. 1009

Ruiz-García, L., Madueno, F., Wilkinson, M., Haughn, G., Salinas, J., and Martínez-Zapater, J.M. (1997). 1010 Different Roles of Flowering-Time Genes in the Activation of Floral lnitiation Genes in Arabidopsis. Plant 1011 Cell 9: 1921–1934. 1012

Ryu, M.Y., Cho, S.K., and Kim, W.T. (2009). RNAi suppression of RPN12a decreases the expression of type-1013 A ARRs, negative regulators of cytokinin signaling pathway, in Arabidopsis. Mol. Cells 28: 375–382. 1014

Sasani, S., Hemming, M.N., Oliver, S.N., Greenup, A., Tavakkol-Afshari, R., Mahfoozi, S., Poustini, K., 1015 Sharifi, H.R., Dennis, E.S., Peacock, W.J., and Trevaskis, B. (2009). The influence of vernalization and 1016 daylength on expression of flowering-time genes in the shoot apex and leaves of barley (Hordeum vulgare). 1017 J. Exp. Bot. 60: 2169–2178. 1018

Schaller, G.E., Street, I.H., and Kieber, J.J. (2014). Cytokinin and the cell cycle. Curr. Opin. Plant Biol. 21: 1019 7–15. 1020

Schmitz, J., Franzen, R., Ngyuen, T.H., Garcia-Maroto, F., Pozzi, C., Salamini, F., and Rohde, W. (2000). 1021 Cloning, mapping and expression analysis of barley MADS-box genes. Plant Mol. Biol. 42: 899–913. 1022

Schmitz, R.J., Sung, S., and Amasino, R.M. (2008). Histone arginine methylation is required for vernalization-1023 induced epigenetic silencing of FLC in winter-annual Arabidopsis thaliana. Proc. Natl. Acad. Sci. U. S. A. 1024 105: 411–416. 1025

Shannon, S. and Ry Meeks-Wagner, O. (1991). A Mutation in the Arabidopsis TFL1 Gene Affects Inf 1026 lorescence Meristem Development. Plant Cell 3: 877–892. 1027

Steinbach, Y. and Hennig, L. (2014). Arabidopsis MSI1 functions in photoperiodic flowering time control. 1028 Front. Plant Sci. 5: 77. 1029

Supek, F., Bošnjak, M., Škunca, N., and Šmuc, T. (2011). Revigo summarizes and visualizes long lists of 1030 gene ontology terms. PLoS One 6: e21800 1031



32

Tamaki, S., Matsuo, S., Wong, H.L., Yokoi, S., and Shimamoto, K. (2007). Hd3a Protein Is a Mobile 1032 Flowering Signal in Rice. Science 316: 1033–1036. 1033

Taoka, K. et al. (2011). 14-3-3 proteins act as intracellular receptors for rice Hd3a florigen. Nature 476: 332–1034 335. 1035

Teichmann, T. and Muhr, M. (2015). Shaping plant architecture. Front. Plant Sci. 6: 1–18. 1036

Theißen, G. (2001). Development of floral organ identity: Stories from the MADS house. Curr. Opin. Plant Biol. 1037 4: 75–85. 1038

Turck, F., Fornara, F., and Coupland, G. (2008). Regulation and Identity of Florigen: FLOWERING LOCUS 1039 T Moves Center Stage. Annu. Rev. Plant Biol. 59: 573–594. 1040

Turnbull, C.G.N. (2005). Plant Architecture and Its Manipulation (Blackwell). 1041

Waddington, S.R., Cartwright, P.M., and Wall, P.C. (1983). A Quantitative Scale of Spike Initial and Pistil 1042 Development in Barley and Wheat. Ann. Bot. 51: 119–130. 1043

Wang, X., Gingrich, D.K., Deng, Y., and Hong, Z. (2012). A nucleostemin-like GTPase required for normal 1044 apical and floral meristem development in Arabidopsis. Mol. Biol. Cell 23: 1446–56. 1045

Wang, X., Zhang, Y., Ma, Q., Zhang, Z., Xue, Y., Bao, S., and Chong, K. (2007). SKB1-mediated symmetric 1046 dimethylation of histone H4R3 controls flowering time in Arabidopsis. EMBO J. 26: 1934–1941. 1047

Wigge, P.A., Kim, M.C., Jaeger, K.E., Busch, W., Schmid, M., Lohmann, J.U., and Weigel, D. (2005). 1048 Integration of Spatial and Temporal Information During Floral Induction in Arabidopsis. Science 309: 1049 1056–1059. 1050

Yan, L., Fu, D., Li, C., Blechl, A., Tranquilli, G., Bonafede, M., Sanchez, A., Valarik, M., Yasuda, S., and 1051 Dubcovsky, J. (2006). The wheat and barley vernalization gene VRN3 is an orthologue of FT. Proc. Natl. 1052 Acad. Sci. 103: 19581–19586. 1053

Yeung, K., Seitz, T., Li, S., Janosch, P., McFerran, B., Kaiser, C., Fee, F., Katsanakis, K.D., Rose, D.W., 1054 Mischak, H., Sedivy, J.M., and Kolch, W. (1999). Suppression of Raf-1 kinase activity and MAP kinase 1055 signalling by RKIP. Nature 401: 173–177. 1056

Youssef, H.M. et al. (2016). VRS2 regulates hormone-mediated inflorescence patterning in barley. Nat. Genet. 1057 49: 157–161. 1058

Zagotta, M.T., Hicks, K.A., Jacobs, C.I., Young, J.C., Hangarter, R.P., and Meeks-Wagner, D.R. (1996). 1059 The Arabidopsis ELF3 gene regulates vegetative photomorphogenesis and the photoperiodic induction of 1060 flowering. Plant J. 10: 691–702. 1061

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D

Figure 1. Phenotypes of hvcen (mat-c) mutants trialled under outdoor conditions over two

consecutive years. Mutants in six different genetic backgrounds are shown next to their respective

parental lines (Table S1). Various traits including A) flowering time, B) grain number per main spike,

C) number of tillers at flowering time, and D) plant height were measured over two years. Flowering

time was scored as time from sowing to heading when the awn of the main spike emerged from the flag

leaf. The line within each box denotes the median. Significant differences between the mutants and their

wild-type parents were calculated by a one-way ANOVA using a Tukey HSD as post hoc test, p<0.001

“***”, P<0.01 “**”, P<0.05 “*”, not significant: “ns”.



BonusLDs

c.907LDs

W1.0 W1.5 W3.5 W5.5 W6.5 W9.5

W1.0 W2.0 W5.5 W9.5

c.907SDs

W1.0 W1.5 W2.0 W2.25 W3.5 W4.5

DAG 6 12 24 39 42 54

BonusSDs

W1.0 W1.5 W1.75 W2.0 W2.5 W3.5

DAG 9 15 21 27 33 45

A

B



c.907

c.94

c.943

Bonus

Figure 3. Phenotyping of three hvcen mutants (mat-c.907, mat-c.94, mat-c.943) and the wildtype

Bonus under controlled LD and SD conditions. Development of the main shoot apex (MSA), number

of spikelet primordia and of axillary buds at different Waddington stages under LDs (A, C, E) and SDs

(B, D, F). The number of spikelet primordia is referring to the number of primordia per main spike. The

axillary buds scored include all the axillary buds, primary, secondary and higher order buds. Five or six

plants per genotype were dissected at each time point under LDs (16 h light/8 h night) and SDs (8h/16h,

light/dark). Statistical differences (P<0.05) were calculated using a polynomial regression model at 95%

confidence interval (Loess smooth line) shown in grey-shaded regions. www.plantphysiol.orgon August 16, 2020 - Published by Downloaded from

Copyright © 2019 American Society of Plant Biologists. All rights reserved.


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geno●●

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hvelf3 hvcen

hvelf3 HvCEN

HvELF3 hvcen

HvELF3 HvCEN

D

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2 3 4 5 6 7 8 9 10Waddington stage

Num

ber o

f spi

kele

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ordi

a

n●

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geno●●

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hvelf3 hvcen

hvelf3 HvCEN

HvELF3 hvcen

HvELF3 HvCEN

E

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1020304050607080

1 2 3 4 5 6 7 8 9 10Waddington stage

Num

ber o

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llary

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n●

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hvelf3 hvcen

hvelf3 HvCEN

HvELF3 hvcen

HvELF3 HvCEN

F

Figure 4. Microscopic phenotypes of double and single hvft3 x hvcen(mat-c.907) and hvelf3 x hvcen

(mat-c.907) mutants in Bonus under SDs. Development of the MSA, number of spikelet primordia and

number of axillary buds in HvFT3 or hvft3 background (A, B, C) and HvELF3 or hvelf3 background (C,

D, E) at different Waddington stages under SDs (8h/16h, light/dark). The number of spikelet primordia

is referring to the number of primordia per main spike. The axillary buds scored include all the axillary

buds, primary, secondary and higher order buds. Three to six plants per genotype were dissected at each

time point. Statistical differences (P<0.05) were calculated using a polynomial regression model at 95%

confidence interval (Loess smooth line) shown in grey-shaded regions.



Figure 5. Expression pattern of HvCEN under LDs. A, B) RNA in situ hybridization using the anti-

sense probe C, D) negative controls using the sense probe of HvCEN in the shoot apex of cv. Bowman

at Waddington stages 2.0 and 3.5; E) expression of HvCEN in the pure inflorescence meristem of cv.

Bonus. Pure meristems were obtained by carefully removing leaf primordia and stem tissue beneath the

inflorescence under the microscope. LDs: long day condition (16h/8h light/dark), W: Waddington stage.



W 1.0

mutants vs Bonus(2+4)

W 2.0


W 3.5




2173330

675446

3

15421002

1

50129

0

8131

25580


7210

110

220

A

SD_c.907_W3.5SD_c.943_W3.5SD_Bonus_W3.5LD_Bonus_W2.0SD_Bonus_W2.0LD_c.907_W3.5LD_c.943_W3.5LD_Bonus_W3.5LD_c.943_W2.0LD_c.907_W2.0LD_c.943_W1.0LD_c.907_W1.0LD_Bonus_W1.0SD_c.907_W1.0SD_c.943_W1.0SD_Bonus_W1.0SD_c.943_W2.0SD_c.907_W2.0

−3 −2 −1 0 1 2 3

I II III IV

B

Cluster No.

Log2-FCLDs SDs

Figure 6. DETs regulated between two hvcen mutants (mat-c.907 and mat-c.943) and wildtype

(Bonus) in at least one photoperiod condition (LD/SDs) and co-expression patterns of the DETs.

A) Number of DETs regulated in both mutants (mat-c.907 and mat-c.943) compared to WT (Bonus)

under LDs and/or SDs. The red circles display DETs detected under LDs and blue circles show DETs

detected under SDs.

B) Heatmap of co-expression clusters for 4527 DETs. Colors represent log2-fold changes (log2-FC) in

expression levels relative to the mean transcript abundance across the tested conditions, i.e., apex

(enriched) samples of mat-c.907, mat-c.943 and Bonus grown under LD and SD conditions and

harvested at different developmental stages (W1.0, W2.0, W3.5). LDs: long day condition (16h/8h,

light/dark), SDs: short day condition (8h/16h, light/dark), W: Waddington stage, DETs: differentially

expressed transcripts. Transcripts with FDR<0.01 were considered as DETs.



cpm

W1.0 W2.0 W3.5

020

4060

8010

0 MSI HORVU5Hr1G084160

cpm

W1.0 W2.0 W3.5

020

4060

8010

0

cpm

W1.0 W2.0 W3.5

050

100

150

200

PRMT5 HORVU6Hr1G019540

cpm

W1.0 W2.0 W3.5

050

100

150

200

cpm

W1.0 W2.0 W3.5

010

2030

4050

60

ATXR6 HORVU6Hr1G011950

cpm

W1.0 W2.0 W3.5

010

2030

4050

60

cpm

W1.0 W2.0 W3.5

020

4060

80

PGY2 HORVU0Hr1G006020

cpm

W1.0 W2.0 W3.5

020

4060

80

cpm

W1.0 W2.0 W3.5

010

2030

4050

60

CDC6 HORVU3Hr1G084800

cpm

W1.0 W2.0 W3.5

010

2030

4050

60

cpm

W1.0 W2.0 W3.5

050

100

PRN12a HORVU4Hr1G002140

cpm

W1.0 W2.0 W3.5

050

100

cpm

W1.0 W2.0 W3.5

020

4060

80

ARR6 HORVU2Hr1G120490

cpm

W1.0 W2.0 W3.5

020

4060

80

cpm

W1.0 W2.0 W3.5

050

100

150

AHK3 HORVU3Hr1G094870

cpm

W1.0 W2.0 W3.5

050

100

150

cpm

W1.0 W2.0 W3.5

050

100

150

200

CWINV2 HORVU2Hr1G073210

cpm

W1.0 W2.0 W3.5

050

100

150

200

cpm

W1.0 W2.0 W3.5

050

100

150

Fructose-bisphosphate aldolase2 HORVU3Hr1G002780

cpm

W1.0 W2.0 W3.5

050

100

150

cpm

W1.0 W2.0 W3.5

010

020

030

040

050

0

PFK7 HORVU6Hr1G070270

cpm

W1.0 W2.0 W3.5

010

020

030

040

050

0

cpm

W1.0 W2.0 W3.5

050

100

200

300

NSN1 HORVU2Hr1G016650

cpm

W1.0 W2.0 W3.5

050

100

200

300

A

D

C

B

Bonus c.907 c.943

Figure 7. Expression profiles of selected DETs between the hvcen mutants (mat-c.907, mat-c.943)

and wildtype (Bonus) at W2.0 related to A) chromatin modification, regulation of development and

organ initiation; B) ribosomal proteins, leaf development, leaf patterning; C) regulation of hormone

level and hormone response D) cellular respiration, sink strength, carbohydrate metabolism, glycolytic

processes under both LDs and SDs. LDs and SDs are shown by white and grey colors. LDs: long day

condition (16h/8h, light/dark), SDs: short day condition (8h/16h, light/dark), white: light period (16h),

grey: dark period (8h), W: Waddington stage, cpm: normalized counts per million, DETs: differential

expressed transcripts, error bars: standard deviation. Transcripts with FDR<0.01 were considered as

DETs.



cpm

W1.0 W2.0 W3.5

050

100

HvCEN HORVU2Hr1G072750

cpm

W1.0 W2.0 W3.5

050

100

cpm

W1.0 W2.0 W3.5

05

1015

2025

SOC1 HORVU1Hr1G051660

cpm

W1.0 W2.0 W3.5

05

1015

2025

cpm

W1.0 W2.0 W3.5

05

1015

20

FPF1 HORVU2Hr1G007350

cpm

W1.0 W2.0 W3.5

05

1015

20

cpm

W1.0 W2.0 W3.5

05

1015

2025

SEP2 HORVU4Hr1G067680

cpm

W1.0 W2.0 W3.5

05

1015

2025

cpm

W1.0 W2.0 W3.5

05

1015


cpm

W1.0 W2.0 W3.5

05

1015

cpm

W1.0 W2.0 W3.5

01

23

45

6

SWEET HORVU5Hr1G076770

cpm

W1.0 W2.0 W3.5

01

23

45

6

cpm

W1.0 W2.0 W3.5

010

2030

40


cpm

W1.0 W2.0 W3.5

010

2030

40

cpm

W1.0 W2.0 W3.5

05

1015

20

PI HORVU1Hr1G063620

cpm

W1.0 W2.0 W3.5

05

1015

20

cpm

W1.0 W2.0 W3.5

05

1015

20

CNGC2 HORVU5Hr1G096440

cpm

W1.0 W2.0 W3.5

05

1015

20

cpm

W1.0 W2.0 W3.5

050

100

150

200

AB Barrel Domain protein HORVU0Hr1G011450

cpm

W1.0 W2.0 W3.5

050

100

150

200

cpm

W1.0 W2.0 W3.5

020

4060

8010

0

ACBP6 HORVU7Hr1G008320

cpm

W1.0 W2.0 W3.5

020

4060

8010

0

cpm

W1.0 W2.0 W3.5

020

4060

80

HvBM8 HORVU2Hr1G063800

cpm

W1.0 W2.0 W3.5

020

4060

80

A

D

C

B

Bonus c.907 c.943

Figure 8. Expression profiles of selected DETs between the hvcen mutants (mat-c.907, mat-c.943)

and wildtype (Bonus) at W3.5 under both LDs and SDs. LDs and SDs are shown by white and grey

colors. LDs: long day conditions (16h/8h, light/dark), SDs: short day conditions (8h/16h, light/dark),

white: light period (16h), gray: dark period (8h), W: Waddington stage, cpm: normalized counts per

million, DETs: differential expressed transcripts, error bars: standard deviation.



Parsed CitationsAbe, M. (2005). FD, a bZIP Protein Mediating Signals from the Floral Pathway Integrator FT at the Shoot Apex. Science. 309: 1052–1056.

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

Ahn, J.H., Miller, D., Winter, V.J., Banfield, M.J., Lee, J.H., Yoo, S.Y., Henz, S.R., Brady, R.L., and Weigel, D. (2006). A divergent externalloop confers antagonistic activity on floral regulators FT and TFL1. EMBO J. 25: 605–14.


Alexa, A., Rahnenführer, J., and Lengauer, T. (2006). Improved scoring of functional groups from gene expression data bydecorrelating GO graph structure. Bioinformatics, 22, pp.1600-1607.


Alqudah, A.M. and Schnurbusch, T. (2014). Awn primordium to tipping is the most decisive developmental phase for spikelet survival inbarley. Funct. Plant Biol. 41: 424–436.


Bhavsar, R.B., Makley, L.N., and Tsonis, P.A. (2010). The other lives of ribosomal proteins. Hum Genomics 4: 327–344.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Boden, S.A., Cavanagh, C., Cullis, B.R., Ramm, K., Greenwood, J., Jean Finnegan, E., Trevaskis, B., and Swain, S.M. (2015). Ppd-1 is akey regulator of inflorescence architecture and paired spikelet development in wheat. Nat. Plants 1: 1–6.


Bommert, P., and Whipple, C. (2018). Grass inflorescence architecture and meristem determinacy. Semin. Cell Dev. Biol. 79: 37–47.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Bouveret, R., Schönrock, N., Gruissem, W., and Hennig, L. (2006). Regulation of flowering time by Arabidopsis MSI1. Development 133:1693-1702


Bradley, D., Carpenter, R., Copsey, L., Vincent, C., Rothstein, S., and Coen, E. (1996). Control of inflorescence architecture inAntirrhinum. Nature 379: 791–797.


Bradley, D., Ratcliffe, O., Vincent, C., Carpenter, R., and Coen, E. (1997). Inflorescence Commitment and Architecture in Arabidopsis.Science 275: 80–83.


Bull, H. et al. (2017). Barley SIX-ROWED SPIKE3 encodes a putative Jumonji C-type H3K9me2/me3 demethylase that represses lateralspikelet fertility. Nat. Commun. 8: 936.


Campoli, C. and von Korff, M. (2014). Genetic control of reproductive development in temperate cereals. In Advances in botanicalresearch (Elsevier), pp. 131–158.


Casao, M.C., Karsai, I., Igartua, E., Gracia, M.P., Veisz, O., and Casas, A.M. (2011). Adaptation of barley to mild winters: A role for PPDH2.BMC Plant Biol. 11. 164.


Chanvivattana, Y., Bishopp, A., Schubert, D., Stock, C., Moon, Y. H., Sung, Z. R., and Goodrich, J. (2004). Interaction of Polycomb-groupproteins controlling flowering in Arabidopsis. Development, 131: 5263-5276.


Chen, A. and Dubcovsky, J. (2012). Wheat TILLING mutants show that the vernalization gene VRN1 down-regulates the floweringrepressor VRN2 in leaves but is not essential for flowering. PLoS Genet. 8: e1003134.



https://www.ncbi.nlm.nih.gov/pubmed?term=Abe%2C M%2E %282005%29%2E FD%2C a bZIP Protein Mediating Signals from the Floral Pathway Integrator FT at the Shoot Apex%2E Science%2E 309%3A 1052%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=FD, a bZIP Protein Mediating Signals from the Floral Pathway Integrator FT at the Shoot Apex&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

https://scholar.google.com/scholar?as_q=FD, a bZIP Protein Mediating Signals from the Floral Pathway Integrator FT at the Shoot Apex&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Abe,&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ahn%2C J%2EH%2E%2C Miller%2C D%2E%2C Winter%2C V%2EJ%2E%2C Banfield%2C M%2EJ%2E%2C Lee%2C J%2EH%2E%2C Yoo%2C S%2EY%2E%2C Henz%2C S%2ER%2E%2C Brady%2C R%2EL%2E%2C and Weigel%2C D%2E %282006%29%2E A divergent external loop confers antagonistic activity on floral regulators FT and TFL1%2E EMBO J%2E 25%3A 605%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A divergent external loop confers antagonistic activity on floral regulators FT and TFL1&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

https://scholar.google.com/scholar?as_q=A divergent external loop confers antagonistic activity on floral regulators FT and TFL1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ahn,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Alexa%2C A%2E%2C Rahnenf�hrer%2C J%2E%2C and Lengauer%2C T%2E %282006%29%2E Improved scoring of functional groups from gene expression data by decorrelating GO graph structure%2E Bioinformatics%2C 22%2C pp&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Improved scoring of functional groups from gene expression data by decorrelating GO graph structure&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

https://scholar.google.com/scholar?as_q=Improved scoring of functional groups from gene expression data by decorrelating GO graph structure&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Alexa,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Alqudah%2C A%2EM%2E and Schnurbusch%2C T%2E %282014%29%2E Awn primordium to tipping is the most decisive developmental phase for spikelet survival in barley%2E Funct%2E Plant Biol%2E 41%3A 424%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Awn primordium to tipping is the most decisive developmental phase for spikelet survival in barley&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

https://scholar.google.com/scholar?as_q=Awn primordium to tipping is the most decisive developmental phase for spikelet survival in barley&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Alqudah,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bhavsar%2C R%2EB%2E%2C Makley%2C L%2EN%2E%2C and Tsonis%2C P%2EA%2E %282010%29%2E The other lives of ribosomal proteins%2E Hum Genomics 4%3A 327%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The other lives of ribosomal 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

https://scholar.google.com/scholar?as_q=The other lives of ribosomal proteins&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bhavsar,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Boden%2C S%2EA%2E%2C Cavanagh%2C C%2E%2C Cullis%2C B%2ER%2E%2C Ramm%2C K%2E%2C Greenwood%2C J%2E%2C Jean Finnegan%2C E%2E%2C Trevaskis%2C B%2E%2C and Swain%2C S%2EM%2E %282015%29%2E Ppd%2D1 is a key regulator of inflorescence architecture and paired spikelet development in wheat%2E Nat%2E Plants 1%3A 1%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Ppd-1 is a key regulator of inflorescence architecture and paired spikelet development in wheat&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

https://scholar.google.com/scholar?as_q=Ppd-1 is a key regulator of inflorescence architecture and paired spikelet development in wheat&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Boden,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bommert%2C P%2E%2C and Whipple%2C C%2E %282018%29%2E Grass inflorescence architecture and meristem determinacy%2E Semin%2E Cell Dev%2E Biol%2E 79%3A 37%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Grass inflorescence architecture and meristem determinacy&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

https://scholar.google.com/scholar?as_q=Grass inflorescence architecture and meristem determinacy&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bommert,&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bouveret%2C R%2E%2C Sch�nrock%2C N%2E%2C Gruissem%2C W%2E%2C and Hennig%2C L%2E %282006%29%2E Regulation of flowering time by Arabidopsis MSI1%2E Development&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Regulation of flowering time by Arabidopsis MSI1&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

https://scholar.google.com/scholar?as_q=Regulation of flowering time by Arabidopsis MSI1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bouveret,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bradley%2C D%2E%2C Carpenter%2C R%2E%2C Copsey%2C L%2E%2C Vincent%2C C%2E%2C Rothstein%2C S%2E%2C and Coen%2C E%2E %281996%29%2E Control of inflorescence architecture in Antirrhinum%2E Nature 379%3A 791%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Control of inflorescence architecture in Antirrhinum&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

https://scholar.google.com/scholar?as_q=Control of inflorescence architecture in Antirrhinum&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bradley,&as_ylo=1996&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bradley%2C D%2E%2C Ratcliffe%2C O%2E%2C Vincent%2C C%2E%2C Carpenter%2C R%2E%2C and Coen%2C E%2E %281997%29%2E Inflorescence Commitment and Architecture in Arabidopsis%2E Science 275%3A 80%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Inflorescence Commitment and Architecture 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

https://scholar.google.com/scholar?as_q=Inflorescence Commitment and Architecture in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bradley,&as_ylo=1997&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bull%2C H%2E et al%2E %282017%29%2E Barley SIX%2DROWED SPIKE3 encodes a putative Jumonji C%2Dtype H3K9me2%2Fme3 demethylase that represses lateral spikelet fertility%2E Nat%2E Commun&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Barley SIX-ROWED SPIKE3 encodes a putative Jumonji C-type H3K9me2/me3 demethylase that represses lateral spikelet fertility&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

https://scholar.google.com/scholar?as_q=Barley SIX-ROWED SPIKE3 encodes a putative Jumonji C-type H3K9me2/me3 demethylase that represses lateral spikelet fertility&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bull,&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Campoli%2C C%2E and von Korff%2C M%2E %282014%29%2E Genetic control of reproductive development in temperate cereals%2E In Advances in botanical research %28Elsevier%29%2C pp%2E 131%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Genetic control of reproductive development in temperate cereals.&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

https://scholar.google.com/scholar?as_q=Genetic control of reproductive development in temperate cereals.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Campoli,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Casao%2C M%2EC%2E%2C Karsai%2C I%2E%2C Igartua%2C E%2E%2C Gracia%2C M%2EP%2E%2C Veisz%2C O%2E%2C and Casas%2C A%2EM%2E %282011%29%2E Adaptation of barley to mild winters%3A A role for PPDH2%2E BMC Plant Biol&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Adaptation of barley to mild winters: A role for PPDH2.&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

https://scholar.google.com/scholar?as_q=Adaptation of barley to mild winters: A role for PPDH2.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Casao,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chanvivattana%2C Y%2E%2C Bishopp%2C A%2E%2C Schubert%2C D%2E%2C Stock%2C C%2E%2C Moon%2C Y%2E H%2E%2C Sung%2C Z%2E R%2E%2C and Goodrich%2C J%2E %282004%29%2E Interaction of Polycomb%2Dgroup proteins controlling flowering in Arabidopsis%2E Development&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Interaction of Polycomb-group proteins controlling flowering 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

https://scholar.google.com/scholar?as_q=Interaction of Polycomb-group proteins controlling flowering in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chanvivattana,&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chen%2C A%2E and Dubcovsky%2C J%2E %282012%29%2E Wheat TILLING mutants show that the vernalization gene VRN1 down%2Dregulates the flowering repressor VRN2 in leaves but is not essential for flowering%2E PLoS Genet%2E 8%3A e&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Wheat TILLING mutants show that the vernalization gene VRN1 down-regulates the flowering repressor VRN2 in leaves but is not essential for flowering&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

https://scholar.google.com/scholar?as_q=Wheat TILLING mutants show that the vernalization gene VRN1 down-regulates the flowering repressor VRN2 in leaves but is not essential for flowering&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chen,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1


Coen, E.S. and Meyerowitz, E.M. (1991). The war of the whorls: genetic interactions controlling flower development. Nature 353: 31–37.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Comadran, J. et al. (2012). Natural variation in a homolog of Antirrhinum CENTRORADIALIS contributed to spring growth habit andenvironmental adaptation in cultivated barley. Nat. Genet. 44: 1388–1392.


Conesa, A., Götz, S., García-Gómez, J.M., Terol, J., Talón, M., and Robles, M. (2005). Blast2GO: A universal tool for annotation,visualization and analysis in functional genomics research. Bioinformatics 21: 3674–3676.


Conti, L. and Bradley, D. (2007). TERMINAL FLOWER1 Is a Mobile Signal Controlling Arabidopsis Architecture. Plant Cell 19: 767–778.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Corbesier, L., Vincent, C., Jang, S., Fornara, F., Fan, Q., Searle, I., Giakountis, A., Farrona, S., Gissot, L., Turnbull, C., and Coupland, G.(2007). FT Protein Movement Contributes to Long-Distance Signaling in Floral Induction of Arabidopsis. Science 316: 1030–1033.


Danilevskaya, O.N., Meng, X., and Ananiev, E. V (2010). Concerted modification of flowering time and inflorescence architecture byectopic expression of TFL1-like genes in maize. Plant Physiol. 153: 238–251.


Danilevskaya, O.N., Meng, X., Hou, Z., Ananiev, E. V., and Simmons, C.R. (2007). A Genomic and Expression Compendium of theExpanded PEBP Gene Family from Maize. Plant Physiol. 146: 250–264.


Deng, X., Gu, L., Liu, C., Lu, T., Lu, F., Lu, Z., Cui, P., Pei, Y., Wang, B., Hu, S., and Cao, X. (2010). Arginine methylation mediated by theArabidopsis homolog of PRMT5 is essential for proper pre-mRNA splicing. Proc. Natl. Acad. Sci. 107: 19114–19119.


Derkacheva, M., Steinbach, Y., Wildhaber, T., Mozgová, I., Mahrez, W., Nanni, P., Bischof, S., Gruissem, W., and Hennig, L. (2013).Arabidopsis MSI1 connects LHP1 to PRC2 complexes. EMBO J. 32: 2073–2085.


Digel, B., Pankin, A., and von Korff, M. (2015). Global Transcriptome Profiling of Developing Leaf and Shoot Apices Reveals DistinctGenetic and Environmental Control of Floral Transition and Inflorescence Development in Barley. Plant Cell 27: 2318-2334


Digel, B., Tavakol, E., Verderio, G., Tondelli, A., Xu, X., Cattivelli, L., Rossini, L. and von Korff, M. (2016). Photoperiod1 (Ppd-H1)controls leaf size. Plant physiology 172: 405-415.


Dixon, L.E., Farré, A., Finnegan, E.J., Orford, S., Griffiths, S. and Boden, S.A. (2018). Developmental responses of bread wheat tochanges in ambient temperature following deletion of a locus that includes FLOWERING LOCUS T1. Plant, cell & environment 41:1715-1725.


Ejaz, M. and von Korff, M. (2017). The Genetic Control of Reproductive Development under High Ambient Temperature. Plant Physiol.173: 294–306.


van Esse, G.W., Walla, A., Finke, A., Koornneef, M., Pecinka, A., and von Korff, M. (2017). Six-Rowed Spike3 (VRS3) Is a HistoneDemethylase That Controls Lateral Spikelet Development in Barley. Plant Physiol. 174: 2397–2408.


Faure, S., Turner, A.S., Gruszka, D., Christodoulou, V., Davis, S.J., von Korff, M., and Laurie, D.A. (2012). Mutation at the circadian clockgene EARLY MATURITY 8 adapts domesticated barley (Hordeum vulgare) to short growing seasons. Proc. Natl. Acad. Sci. 109: 8328–8333.



https://www.ncbi.nlm.nih.gov/pubmed?term=Coen%2C E%2ES%2E and Meyerowitz%2C E%2EM%2E %281991%29%2E The war of the whorls%3A genetic interactions controlling flower development%2E Nature 353%3A 31%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The war of the whorls: genetic interactions controlling flower 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

https://scholar.google.com/scholar?as_q=The war of the whorls: genetic interactions controlling flower development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Coen,&as_ylo=1991&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Comadran%2C J%2E et al%2E %282012%29%2E Natural variation in a homolog of Antirrhinum CENTRORADIALIS contributed to spring growth habit and environmental adaptation in cultivated barley%2E Nat%2E Genet%2E 44%3A 1388%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Natural variation in a homolog of Antirrhinum CENTRORADIALIS contributed to spring growth habit and environmental adaptation in cultivated barley&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

https://scholar.google.com/scholar?as_q=Natural variation in a homolog of Antirrhinum CENTRORADIALIS contributed to spring growth habit and environmental adaptation in cultivated barley&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Comadran,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Conesa%2C A%2E%2C G�tz%2C S%2E%2C Garc�a%2DG�mez%2C J%2EM%2E%2C Terol%2C J%2E%2C Tal�n%2C M%2E%2C and Robles%2C M%2E %282005%29%2E Blast2GO%3A A universal tool for annotation%2C visualization and analysis in functional genomics research%2E Bioinformatics 21%3A 3674%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Blast2GO: A universal tool for annotation, visualization and analysis in functional genomics research&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

https://scholar.google.com/scholar?as_q=Blast2GO: A universal tool for annotation, visualization and analysis in functional genomics research&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Conesa,&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Conti%2C L%2E and Bradley%2C D%2E %282007%29%2E TERMINAL FLOWER1 Is a Mobile Signal Controlling Arabidopsis Architecture%2E Plant Cell 19%3A 767%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=TERMINAL FLOWER1 Is a Mobile Signal Controlling Arabidopsis Architecture&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

https://scholar.google.com/scholar?as_q=TERMINAL FLOWER1 Is a Mobile Signal Controlling Arabidopsis Architecture&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Conti,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Corbesier%2C L%2E%2C Vincent%2C C%2E%2C Jang%2C S%2E%2C Fornara%2C F%2E%2C Fan%2C Q%2E%2C Searle%2C I%2E%2C Giakountis%2C A%2E%2C Farrona%2C S%2E%2C Gissot%2C L%2E%2C Turnbull%2C C%2E%2C and Coupland%2C G%2E %282007%29%2E FT Protein Movement Contributes to Long%2DDistance Signaling in Floral Induction of Arabidopsis%2E Science 316%3A 1030%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=FT Protein Movement Contributes to Long-Distance Signaling in Floral Induction of 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

https://scholar.google.com/scholar?as_q=FT Protein Movement Contributes to Long-Distance Signaling in Floral Induction of Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Corbesier,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Danilevskaya%2C O%2EN%2E%2C Meng%2C X%2E%2C and Ananiev%2C E%2E V %282010%29%2E Concerted modification of flowering time and inflorescence architecture by ectopic expression of TFL1%2Dlike genes in maize%2E Plant Physiol%2E 153%3A 238%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Concerted modification of flowering time and inflorescence architecture by ectopic expression of TFL1-like genes in maize&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

https://scholar.google.com/scholar?as_q=Concerted modification of flowering time and inflorescence architecture by ectopic expression of TFL1-like genes in maize&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Danilevskaya,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Danilevskaya%2C O%2EN%2E%2C Meng%2C X%2E%2C Hou%2C Z%2E%2C Ananiev%2C E%2E V%2E%2C and Simmons%2C C%2ER%2E %282007%29%2E A Genomic and Expression Compendium of the Expanded PEBP Gene Family from Maize%2E Plant Physiol%2E 146%3A 250%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A Genomic and Expression Compendium of the Expanded PEBP Gene Family from Maize&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

https://scholar.google.com/scholar?as_q=A Genomic and Expression Compendium of the Expanded PEBP Gene Family from Maize&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Danilevskaya,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Deng%2C X%2E%2C Gu%2C L%2E%2C Liu%2C C%2E%2C Lu%2C T%2E%2C Lu%2C F%2E%2C Lu%2C Z%2E%2C Cui%2C P%2E%2C Pei%2C Y%2E%2C Wang%2C B%2E%2C Hu%2C S%2E%2C and Cao%2C X%2E %282010%29%2E Arginine methylation mediated by the Arabidopsis homolog of PRMT5 is essential for proper pre%2DmRNA splicing%2E Proc%2E Natl%2E Acad%2E Sci%2E 107%3A 19114%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Arginine methylation mediated by the Arabidopsis homolog of PRMT5 is essential for proper pre-mRNA splicing&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

https://scholar.google.com/scholar?as_q=Arginine methylation mediated by the Arabidopsis homolog of PRMT5 is essential for proper pre-mRNA splicing&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Deng,&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Derkacheva%2C M%2E%2C Steinbach%2C Y%2E%2C Wildhaber%2C T%2E%2C Mozgov�%2C I%2E%2C Mahrez%2C W%2E%2C Nanni%2C P%2E%2C Bischof%2C S%2E%2C Gruissem%2C W%2E%2C and Hennig%2C L%2E %282013%29%2E Arabidopsis MSI1 connects LHP1 to PRC2 complexes%2E EMBO J%2E 32%3A 2073%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Arabidopsis MSI1 connects LHP1 to PRC2 complexes&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

https://scholar.google.com/scholar?as_q=Arabidopsis MSI1 connects LHP1 to PRC2 complexes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Derkacheva,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Digel%2C B%2E%2C Pankin%2C A%2E%2C and von Korff%2C M%2E %282015%29%2E Global Transcriptome Profiling of Developing Leaf and Shoot Apices Reveals Distinct Genetic and Environmental Control of Floral Transition and Inflorescence Development in Barley%2E Plant Cell&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Global Transcriptome Profiling of Developing Leaf and Shoot Apices Reveals Distinct Genetic and Environmental Control of Floral Transition and Inflorescence Development in Barley&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

https://scholar.google.com/scholar?as_q=Global Transcriptome Profiling of Developing Leaf and Shoot Apices Reveals Distinct Genetic and Environmental Control of Floral Transition and Inflorescence Development in Barley&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Digel,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Digel%2C B%2E%2C Tavakol%2C E%2E%2C Verderio%2C G%2E%2C Tondelli%2C A%2E%2C Xu%2C X%2E%2C Cattivelli%2C L%2E%2C Rossini%2C L%2E and von Korff%2C M%2E %282016%29%2E Photoperiod1 %28Ppd%2DH1%29 controls leaf size%2E Plant physiology&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Photoperiod1 (Ppd-H1) controls leaf 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

https://scholar.google.com/scholar?as_q=Photoperiod1 (Ppd-H1) controls leaf size&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Digel,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Dixon%2C L%2EE%2E%2C Farr�%2C A%2E%2C Finnegan%2C E%2EJ%2E%2C Orford%2C S%2E%2C Griffiths%2C S%2E and Boden%2C S%2EA%2E %282018%29%2E Developmental responses of bread wheat to changes in ambient temperature following deletion of a locus that includes FLOWERING LOCUS T1%2E Plant%2C cell %26 environment&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Developmental responses of bread wheat to changes in ambient temperature following deletion of a locus that includes FLOWERING LOCUS T1&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

https://scholar.google.com/scholar?as_q=Developmental responses of bread wheat to changes in ambient temperature following deletion of a locus that includes FLOWERING LOCUS T1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Dixon,&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ejaz%2C M%2E and von Korff%2C M%2E %282017%29%2E The Genetic Control of Reproductive Development under High Ambient Temperature%2E Plant Physiol%2E 173%3A 294%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The Genetic Control of Reproductive Development under High Ambient Temperature&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

https://scholar.google.com/scholar?as_q=The Genetic Control of Reproductive Development under High Ambient Temperature&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ejaz,&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=van Esse%2C G%2EW%2E%2C Walla%2C A%2E%2C Finke%2C A%2E%2C Koornneef%2C M%2E%2C Pecinka%2C A%2E%2C and von Korff%2C M%2E %282017%29%2E Six%2DRowed Spike3 %28VRS3%29 Is a Histone Demethylase That Controls Lateral Spikelet Development in Barley%2E Plant Physiol%2E 174%3A 2397%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Six-Rowed Spike3 (VRS3) Is a Histone Demethylase That Controls Lateral Spikelet Development in Barley&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

https://scholar.google.com/scholar?as_q=Six-Rowed Spike3 (VRS3) Is a Histone Demethylase That Controls Lateral Spikelet Development in Barley&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=van&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Faure%2C S%2E%2C Turner%2C A%2ES%2E%2C Gruszka%2C D%2E%2C Christodoulou%2C V%2E%2C Davis%2C S%2EJ%2E%2C von Korff%2C M%2E%2C and Laurie%2C D%2EA%2E %282012%29%2E Mutation at the circadian clock gene EARLY MATURITY 8 adapts domesticated barley %28Hordeum vulgare%29 to short growing seasons%2E Proc%2E Natl%2E Acad%2E Sci%2E 109%3A 8328%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Mutation at the circadian clock gene EARLY MATURITY 8 adapts domesticated barley (Hordeum vulgare) to short growing seasons&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

https://scholar.google.com/scholar?as_q=Mutation at the circadian clock gene EARLY MATURITY 8 adapts domesticated barley (Hordeum vulgare) to short growing seasons&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Faure,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1


Franckowiak, J.D. and Lundqvist, U. (2012). Descriptions of Barley Genetic Stocks for 2012. Barley Genet. Newsl. 42: 36–173.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Gao, X., Liang, W., Yin, C., Ji, S., Wang, H., Su, X., Guo, C., Kong, H., Xue, H., and Zhang, D. (2010). The SEPALLATA-Like GeneOsMADS34 Is Required for Rice Inflorescence and Spikelet Development. Plant Physiol. 153: 728–740.


Halliwell, J., Borrill, P., Gordon, A., Kowalczyk, R., Pagano, M.L., Saccomanno, B., Bentley, A.R., Uauy, C., and Cockram, J. (2016).Systematic Investigation of FLOWERING LOCUS T-Like Poaceae Gene Families Identifies the Short-Day Expressed Flowering PathwayGene, TaFT3 in Wheat (Triticum aestivum L.). Front. Plant Sci. 7: 1–15.


Hanano, S. and Goto, K. (2011). Arabidopsis TERMINAL FLOWER1 Is Involved in the Regulation of Flowering Time and InflorescenceDevelopment through Transcriptional Repression. Plant Cell 23: 3172–3184.


Hemming, M.N., Peacock, W.J., Dennis, E.S., and Trevaskis, B. (2008). Low-Temperature and Daylength Cues Are Integrated toRegulate FLOWERING LOCUS T in Barley. Plant Physiol. 147: 355–366.


Hennig, L., Bouveret, R., and Gruissem, W. (2005). MSI1-like proteins: An escort service for chromatin assembly and remodelingcomplexes. Trends Cell Biol. 15: 295–302.


Hicks, K.A., Millar, A.J., Carre, I.A., Somers, D.E., Straume, M., Meeks-Wagner, R., and Kay, S.A. (1996). Conditional circadian dysfuncionof the Arabidopsisearly-flowering 3 mutant. Science 274: 790–792.


Ito, T., Kim, G.T., and Shinozaki, K. (2000). Disruption of an Arabidopsis cytoplasmic ribosomal protein S13-homologous gene bytransposon-mediated mutagenesis causes aberrant growth and development. Plant J. 22: 257–264.


Jacob, Y., Feng, S., LeBlanc, C.A., Bernatavichute, Y. V., Stroud, H., Cokus, S., Johnson, L.M., Pellegrini, M., Jacobsen, S.E., andMichaels, S.D. (2009). ATXR5 and ATXR6 are H3K27 monomethyltransferases required for chromatin structure and gene silencing. Nat.Struct. Mol. Biol. 16: 763–768.


Jaeger, K.E., Pullen, N., Lamzin, S., Morris, R.J., and Wigge, P.A. (2013). Interlocking Feedback Loops Govern the Dynamic Behavior ofthe Floral Transition in Arabidopsis. Plant Cell 25: 820–833.


Jensen, C.S., Salchert, K., and Nielsen, K.K. (2001). A TERMINAL FLOWER1-like gene from perennial ryegrass involved in floraltransition and axillary meristem identity. Plant Physiol. 125: 1517–1528.


Jeon, Y., Park, Y.J., Cho, H.K., Jung, H.J., Ahn, T.K., Kang, H., and Pai, H.S. (2015). The nucleolar GTPase nucleostemin-like 1 plays arole in plant growth and senescence by modulating ribosome biogenesis. J. Exp. Bot. 66: 6297–6310.


Jiang, K., Liberatore, K.L., Park, S.J., Alvarez, J.P., and Lippman, Z.B. (2013). Tomato Yield Heterosis Is Triggered by a DosageSensitivity of the Florigen Pathway That Fine-Tunes Shoot Architecture. PLoS Genet. 9: e1004043


Kaneko-Suzuki, M., Kurihara-Ishikawa, R., Okushita-Terakawa, C., Kojima, C., Nagano-Fujiwara, M., Ohki, I., Tsuji, H., Shimamoto, K.,and Taoka, K.I. (2018). TFL1-Like Proteins in Rice Antagonize Rice FT-Like Protein in Inflorescence Development by Competition forComplex Formation with 14-3-3 and FD. Plant Cell Physiol. 59: 458–468.


Kardailsky, I. (1999). Activation Tagging of the Floral Inducer FT. Science 286: 1962–1965.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title www.plantphysiol.orgon August 16, 2020 - Published by Downloaded from


https://www.ncbi.nlm.nih.gov/pubmed?term=Franckowiak%2C J%2ED%2E and Lundqvist%2C U%2E %282012%29%2E Descriptions of Barley Genetic Stocks for 2012%2E Barley Genet%2E Newsl%2E 42%3A 36%26%23x&dopt=abstract

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

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https://www.ncbi.nlm.nih.gov/pubmed?term=Gao%2C X%2E%2C Liang%2C W%2E%2C Yin%2C C%2E%2C Ji%2C S%2E%2C Wang%2C H%2E%2C Su%2C X%2E%2C Guo%2C C%2E%2C Kong%2C H%2E%2C Xue%2C H%2E%2C and Zhang%2C D%2E %282010%29%2E The SEPALLATA%2DLike Gene OsMADS34 Is Required for Rice Inflorescence and Spikelet Development%2E Plant Physiol%2E 153%3A 728%26%23x&dopt=abstract

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https://www.ncbi.nlm.nih.gov/pubmed?term=Halliwell%2C J%2E%2C Borrill%2C P%2E%2C Gordon%2C A%2E%2C Kowalczyk%2C R%2E%2C Pagano%2C M%2EL%2E%2C Saccomanno%2C B%2E%2C Bentley%2C A%2ER%2E%2C Uauy%2C C%2E%2C and Cockram%2C J%2E %282016%29%2E Systematic Investigation of FLOWERING LOCUS T%2DLike Poaceae Gene Families Identifies the Short%2DDay Expressed Flowering Pathway Gene%2C TaFT3 in Wheat %28Triticum aestivum L%2E%29%2E Front%2E Plant Sci%2E 7%3A 1%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Systematic Investigation of FLOWERING LOCUS T-Like Poaceae Gene Families Identifies the Short-Day Expressed Flowering Pathway Gene, TaFT3 in Wheat (Triticum aestivum L&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

https://scholar.google.com/scholar?as_q=Systematic Investigation of FLOWERING LOCUS T-Like Poaceae Gene Families Identifies the Short-Day Expressed Flowering Pathway Gene, TaFT3 in Wheat (Triticum aestivum L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Halliwell,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hanano%2C S%2E and Goto%2C K%2E %282011%29%2E Arabidopsis TERMINAL FLOWER1 Is Involved in the Regulation of Flowering Time and Inflorescence Development through Transcriptional Repression%2E Plant Cell 23%3A 3172%26%23x&dopt=abstract

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

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https://scholar.google.com/scholar?as_q=Arabidopsis TERMINAL FLOWER1 Is Involved in the Regulation of Flowering Time and Inflorescence Development through Transcriptional Repression&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hanano,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hemming%2C M%2EN%2E%2C Peacock%2C W%2EJ%2E%2C Dennis%2C E%2ES%2E%2C and Trevaskis%2C B%2E %282008%29%2E Low%2DTemperature and Daylength Cues Are Integrated to Regulate FLOWERING LOCUS T in Barley%2E Plant Physiol%2E 147%3A 355%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Low-Temperature and Daylength Cues Are Integrated to Regulate FLOWERING LOCUS T in Barley&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

https://scholar.google.com/scholar?as_q=Low-Temperature and Daylength Cues Are Integrated to Regulate FLOWERING LOCUS T in Barley&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hemming,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hennig%2C L%2E%2C Bouveret%2C R%2E%2C and Gruissem%2C W%2E %282005%29%2E MSI1%2Dlike proteins%3A An escort service for chromatin assembly and remodeling complexes%2E Trends Cell Biol%2E 15%3A 295%26%23x&dopt=abstract

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

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https://scholar.google.com/scholar?as_q=MSI1-like proteins: An escort service for chromatin assembly and remodeling complexes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hennig,&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hicks%2C K%2EA%2E%2C Millar%2C A%2EJ%2E%2C Carre%2C I%2EA%2E%2C Somers%2C D%2EE%2E%2C Straume%2C M%2E%2C Meeks%2DWagner%2C R%2E%2C and Kay%2C S%2EA%2E %281996%29%2E Conditional circadian dysfuncion of the Arabidopsisearly%2Dflowering 3 mutant%2E Science 274%3A 790%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Conditional circadian dysfuncion of the Arabidopsisearly-flowering 3 mutant&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

https://scholar.google.com/scholar?as_q=Conditional circadian dysfuncion of the Arabidopsisearly-flowering 3 mutant&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hicks,&as_ylo=1996&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ito%2C T%2E%2C Kim%2C G%2ET%2E%2C and Shinozaki%2C K%2E %282000%29%2E Disruption of an Arabidopsis cytoplasmic ribosomal protein S13%2Dhomologous gene by transposon%2Dmediated mutagenesis causes aberrant growth and development%2E Plant J%2E 22%3A 257%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Disruption of an Arabidopsis cytoplasmic ribosomal protein S13-homologous gene by transposon-mediated mutagenesis causes aberrant growth and 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

https://scholar.google.com/scholar?as_q=Disruption of an Arabidopsis cytoplasmic ribosomal protein S13-homologous gene by transposon-mediated mutagenesis causes aberrant growth and development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ito,&as_ylo=2000&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jacob%2C Y%2E%2C Feng%2C S%2E%2C LeBlanc%2C C%2EA%2E%2C Bernatavichute%2C Y%2E V%2E%2C Stroud%2C H%2E%2C Cokus%2C S%2E%2C Johnson%2C L%2EM%2E%2C Pellegrini%2C M%2E%2C Jacobsen%2C S%2EE%2E%2C and Michaels%2C S%2ED%2E %282009%29%2E ATXR5 and ATXR6 are H3K27 monomethyltransferases required for chromatin structure and gene silencing%2E Nat%2E Struct%2E Mol%2E Biol%2E 16%3A 763%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=ATXR5 and ATXR6 are H3K27 monomethyltransferases required for chromatin structure and gene silencing&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

https://scholar.google.com/scholar?as_q=ATXR5 and ATXR6 are H3K27 monomethyltransferases required for chromatin structure and gene silencing&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jacob,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jaeger%2C K%2EE%2E%2C Pullen%2C N%2E%2C Lamzin%2C S%2E%2C Morris%2C R%2EJ%2E%2C and Wigge%2C P%2EA%2E %282013%29%2E Interlocking Feedback Loops Govern the Dynamic Behavior of the Floral Transition in Arabidopsis%2E Plant Cell 25%3A 820%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Interlocking Feedback Loops Govern the Dynamic Behavior of the Floral Transition 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

https://scholar.google.com/scholar?as_q=Interlocking Feedback Loops Govern the Dynamic Behavior of the Floral Transition in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jaeger,&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jensen%2C C%2ES%2E%2C Salchert%2C K%2E%2C and Nielsen%2C K%2EK%2E %282001%29%2E A TERMINAL FLOWER1%2Dlike gene from perennial ryegrass involved in floral transition and axillary meristem identity%2E Plant Physiol%2E 125%3A 1517%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A TERMINAL FLOWER1-like gene from perennial ryegrass involved in floral transition and axillary meristem identity&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

https://scholar.google.com/scholar?as_q=A TERMINAL FLOWER1-like gene from perennial ryegrass involved in floral transition and axillary meristem identity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jensen,&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jeon%2C Y%2E%2C Park%2C Y%2EJ%2E%2C Cho%2C H%2EK%2E%2C Jung%2C H%2EJ%2E%2C Ahn%2C T%2EK%2E%2C Kang%2C H%2E%2C and Pai%2C H%2ES%2E %282015%29%2E The nucleolar GTPase nucleostemin%2Dlike 1 plays a role in plant growth and senescence by modulating ribosome biogenesis%2E J%2E Exp%2E Bot%2E 66%3A 6297%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The nucleolar GTPase nucleostemin-like 1 plays a role in plant growth and senescence by modulating ribosome biogenesis&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

https://scholar.google.com/scholar?as_q=The nucleolar GTPase nucleostemin-like 1 plays a role in plant growth and senescence by modulating ribosome biogenesis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jeon,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jiang%2C K%2E%2C Liberatore%2C K%2EL%2E%2C Park%2C S%2EJ%2E%2C Alvarez%2C J%2EP%2E%2C and Lippman%2C Z%2EB%2E %282013%29%2E Tomato Yield Heterosis Is Triggered by a Dosage Sensitivity of the Florigen Pathway That Fine%2DTunes Shoot Architecture%2E PLoS Genet%2E 9%3A e&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Tomato Yield Heterosis Is Triggered by a Dosage Sensitivity of the Florigen Pathway That Fine-Tunes Shoot Architecture.&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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https://www.ncbi.nlm.nih.gov/pubmed?term=Kaneko%2DSuzuki%2C M%2E%2C Kurihara%2DIshikawa%2C R%2E%2C Okushita%2DTerakawa%2C C%2E%2C Kojima%2C C%2E%2C Nagano%2DFujiwara%2C M%2E%2C Ohki%2C I%2E%2C Tsuji%2C H%2E%2C Shimamoto%2C K%2E%2C and Taoka%2C K%2EI%2E %282018%29%2E TFL1%2DLike Proteins in Rice Antagonize Rice FT%2DLike Protein in Inflorescence Development by Competition for Complex Formation with 14%2D3%2D3 and FD%2E Plant Cell Physiol%2E 59%3A 458%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Kaneko-Suzuki,&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=TFL1-Like Proteins in Rice Antagonize Rice FT-Like Protein in Inflorescence Development by Competition for Complex Formation with 14-3-3 and FD&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

https://scholar.google.com/scholar?as_q=TFL1-Like Proteins in Rice Antagonize Rice FT-Like Protein in Inflorescence Development by Competition for Complex Formation with 14-3-3 and FD&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kaneko-Suzuki,&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kardailsky%2C I%2E %281999%29%2E Activation Tagging of the Floral Inducer FT%2E Science 286%3A 1962%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Activation Tagging of the Floral Inducer FT&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

https://scholar.google.com/scholar?as_q=Activation Tagging of the Floral Inducer FT&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kardailsky,&as_ylo=1999&as_allsubj=all&hl=en&c2coff=1


Google Scholar: Author Only Title Only Author and Title

Kernich, G.C., Halloran, G.M., and Flood, R.G. (1997). Variation in duration of pre-anthesis phases of development in barley (Hordeumvulgare). Aust. J. Agric. Res. 48: 59–66.


Kirby, E.J.M. and Appleyard, M. (1987). Development and structure of the wheat plant. In: Wheat breeding (Springer), pp. 287–311.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Kobayashi, K., Maekawa, M., Miyao, A., Hirochika, H., and Kyozuka, J. (2010). PANICLE PHYTOMER2 (PAP2), encoding a SEPALLATAsubfamily MADS-box protein, positively controls spikelet meristem identity in rice. Plant Cell Physiol. 51: 47–57.


Kobayashi, Y. (1999). A Pair of Related Genes with Antagonistic Roles in Mediating Flowering Signals. Science 286: 1960–1962.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Komatsuda, T. et al. (2007). Six-rowed barley originated from a mutation in a homeodomain-leucine zipper I-class homeobox gene.Proc. Natl. Acad. Sci. 104: 1424–1429.


Komiya, R., Ikegami, A., Tamaki, S., Yokoi, S., and Shimamoto, K. (2008). Hd3a and RFT1 are essential for flowering in rice. Development135: 767–774.


Koppolu, R. et al. (2013). Six-rowed spike4 (Vrs4) controls spikelet determinacy and row-type in barley. Proc. Natl. Acad. Sci. 110:13198–13203.


Krieger, U., Lippman, Z.B., and Zamir, D. (2010). The flowering gene SINGLE FLOWER TRUSS drives heterosis for yield in tomato. Nat.Genet. 42: 459–463.


Lifschitz, E., Ayre, B.G., and Eshed, Y. (2014). Florigen and anti-florigen - a systemic mechanism for coordinating growth andtermination in flowering plants. Front. Plant Sci. 5: 1–14.


Liu, X.L., Covington, M.F., Fankhauser, C., Chory, J., and D. Ry Wagner (2001). ELF3 Encodes a Circadian Clock-Regulated NuclearProtein That Functions in an Arabidopsis PHYB Signal Transduction Pathway. Plant Cell Online 13: 1293–1304.


Mascher, M. et al. (2017). A chromosome conformation capture ordered sequence of the barley genome. Nature 544: 427–433.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Matyszczak, I. (2014). Characterization of early maturity barley mutants praematurum-a, -b and –c. PhD. Thesis Aarhus University,Department of Molecular Biology and Genetics, Faculty of Science and Technology, Denmark.


McGarry, R.C. and Ayre, B.G. (2012). Manipulating plant architecture with members of the CETS gene family. Plant Sci. 188–189: 71–81.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Mulki, M.A., Bi, Xi., and von Korff, M. (2018). FLOWERING LOCUS T3 controls spikelet initiation but not floral development. PlantPhysiol.: pp.00236.


Nakagawa M., Shimamoto K., and Kyozuka J. (2002). Overexpression of RCN1and RCN2, rice TERMINAL FLOWER 1/CENTRORADIALIShomologs, confers delay of phase transition and altered panicle morphology in rice. Plant J 29: 743–750


Naora, H. and Naora, H. (1999). Involvement of ribosomal proteins in regulating cell growth and apoptosis: Translational modulation orrecruitment for extraribosomal activity? Immunol. Cell Biol. 77: 197–205.

Pubmed: Author and Title www.plantphysiol.orgon August 16, 2020 - Published by Downloaded from Copyright © 2019 American Society of Plant Biologists. All rights reserved.

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Niu, L., Lu, F., Pei, Y., Liu, C., and Cao, X. (2007). Regulation of flowering time by the protein arginine methyltransferase AtPRMT10.EMBO Rep. 8: 1190–1195.


Park, S.J., Jiang, K., Tal, L., Yichie, Y., Gar, O., Zamir, D., Eshed, Y., and Lippman, Z.B. (2014). Optimization of crop productivity intomato using induced mutations in the florigen pathway. Nat. Genet. 46: 1337–1342.


Pearce, S., Vanzetti, L.S., and Dubcovsky, J. (2013). Exogenous Gibberellins Induce Wheat Spike Development under Short Days Onlyin the Presence of VERNALIZATION1. Plant Physiol. 163: 1433–1445.


Pei, Y., Niu, L., Lu, F., Liu, C., Zhai, J., Kong, X., and Cao, X. (2007). Mutations in the Type II Protein Arginine MethyltransferaseAtPRMT5 Result in Pleiotropic Developmental Defects in Arabidopsis. Plant Physiol. 144: 1913–1923.


Pinon, V., Etchells, J.P., Rossignol, P., Collier, S.A., Arroyo, J.M., Martienssen, R.A., and Byrne, M.E. (2008). Three PIGGYBACK genesthat specifically influence leaf patterning encode ribosomal proteins. Development 135: 1315–1324.


Ramsay, L., Comadran, J., Druka, A., Marshall, D.F., Thomas, W.T., Macaulay, M., MacKenzie, K., Simpson, C., Fuller, J., Bonar, N. andHayes, P.M. (2011). INTERMEDIUM-C, a modifier of lateral spikelet fertility in barley, is an ortholog of the maize domestication geneTEOSINTE BRANCHED 1. Nature genetics, 43: 169-173.


Riggs, T.J. and Kirby, E.J.M. (1978). Developmental consequences of two-row and six-row ear type in spring barley: 1. Geneticalanalysis and comparison of mature plant characters. J. Agric. Sci. 91: 199–205.


Ruiz-García, L., Madueno, F., Wilkinson, M., Haughn, G., Salinas, J., and Martínez-Zapater, J.M. (1997). Different Roles of Flowering-Time Genes in the Activation of Floral lnitiation Genes in Arabidopsis. Plant Cell 9: 1921–1934.


Ryu, M.Y., Cho, S.K., and Kim, W.T. (2009). RNAi suppression of RPN12a decreases the expression of type-A ARRs, negative regulatorsof cytokinin signaling pathway, in Arabidopsis. Mol. Cells 28: 375–382.


Sasani, S., Hemming, M.N., Oliver, S.N., Greenup, A., Tavakkol-Afshari, R., Mahfoozi, S., Poustini, K., Sharifi, H.R., Dennis, E.S.,Peacock, W.J., and Trevaskis, B. (2009). The influence of vernalization and daylength on expression of flowering-time genes in theshoot apex and leaves of barley (Hordeum vulgare). J. Exp. Bot. 60: 2169–2178.


Schaller, G.E., Street, I.H., and Kieber, J.J. (2014). Cytokinin and the cell cycle. Curr. Opin. Plant Biol. 21: 7–15.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Schmitz, J., Franzen, R., Ngyuen, T.H., Garcia-Maroto, F., Pozzi, C., Salamini, F., and Rohde, W. (2000). Cloning, mapping andexpression analysis of barley MADS-box genes. Plant Mol. Biol. 42: 899–913.


Schmitz, R.J., Sung, S., and Amasino, R.M. (2008). Histone arginine methylation is required for vernalization-induced epigeneticsilencing of FLC in winter-annual Arabidopsis thaliana. Proc. Natl. Acad. Sci. U. S. A. 105: 411–416.


Shannon, S. and Ry Meeks-Wagner, O. (1991). A Mutation in the Arabidopsis TFL1 Gene Affects Inf lorescence Meristem Development.Plant Cell 3: 877–892.


Steinbach, Y. and Hennig, L. (2014). Arabidopsis MSI1 functions in photoperiodic flowering time control. Front. Plant Sci. 5: 77.Pubmed: Author and Title www.plantphysiol.orgon August 16, 2020 - Published by Downloaded from


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

https://scholar.google.com/scholar?as_q=Involvement of ribosomal proteins in regulating cell growth and apoptosis: Translational modulation or recruitment for extraribosomal activity&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

https://scholar.google.com/scholar?as_q=Involvement of ribosomal proteins in regulating cell growth and apoptosis: Translational modulation or recruitment for extraribosomal activity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Naora,&as_ylo=1999&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Niu%2C L%2E%2C Lu%2C F%2E%2C Pei%2C Y%2E%2C Liu%2C C%2E%2C and Cao%2C X%2E %282007%29%2E Regulation of flowering time by the protein arginine methyltransferase AtPRMT10%2E EMBO Rep%2E 8%3A 1190%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Regulation of flowering time by the protein arginine methyltransferase AtPRMT10&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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https://www.ncbi.nlm.nih.gov/pubmed?term=Park%2C S%2EJ%2E%2C Jiang%2C K%2E%2C Tal%2C L%2E%2C Yichie%2C Y%2E%2C Gar%2C O%2E%2C Zamir%2C D%2E%2C Eshed%2C Y%2E%2C and Lippman%2C Z%2EB%2E %282014%29%2E Optimization of crop productivity in tomato using induced mutations in the florigen pathway%2E Nat%2E Genet%2E 46%3A 1337%26%23x&dopt=abstract

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

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https://www.ncbi.nlm.nih.gov/pubmed?term=Pearce%2C S%2E%2C Vanzetti%2C L%2ES%2E%2C and Dubcovsky%2C J%2E %282013%29%2E Exogenous Gibberellins Induce Wheat Spike Development under Short Days Only in the Presence of VERNALIZATION1%2E Plant Physiol%2E 163%3A 1433%26%23x&dopt=abstract

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https://www.ncbi.nlm.nih.gov/pubmed?term=Pei%2C Y%2E%2C Niu%2C L%2E%2C Lu%2C F%2E%2C Liu%2C C%2E%2C Zhai%2C J%2E%2C Kong%2C X%2E%2C and Cao%2C X%2E %282007%29%2E Mutations in the Type II Protein Arginine Methyltransferase AtPRMT5 Result in Pleiotropic Developmental Defects in Arabidopsis%2E Plant Physiol%2E 144%3A 1913%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Mutations in the Type II Protein Arginine Methyltransferase AtPRMT5 Result in Pleiotropic Developmental Defects 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

https://scholar.google.com/scholar?as_q=Mutations in the Type II Protein Arginine Methyltransferase AtPRMT5 Result in Pleiotropic Developmental Defects in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Pei,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Pinon%2C V%2E%2C Etchells%2C J%2EP%2E%2C Rossignol%2C P%2E%2C Collier%2C S%2EA%2E%2C Arroyo%2C J%2EM%2E%2C Martienssen%2C R%2EA%2E%2C and Byrne%2C M%2EE%2E %282008%29%2E Three PIGGYBACK genes that specifically influence leaf patterning encode ribosomal proteins%2E Development 135%3A 1315%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Three PIGGYBACK genes that specifically influence leaf patterning encode ribosomal 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

https://scholar.google.com/scholar?as_q=Three PIGGYBACK genes that specifically influence leaf patterning encode ribosomal proteins&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Pinon,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ramsay%2C L%2E%2C Comadran%2C J%2E%2C Druka%2C A%2E%2C Marshall%2C D%2EF%2E%2C Thomas%2C W%2ET%2E%2C Macaulay%2C M%2E%2C MacKenzie%2C K%2E%2C Simpson%2C C%2E%2C Fuller%2C J%2E%2C Bonar%2C N%2E and Hayes%2C P%2EM%2E %282011%29%2E INTERMEDIUM%2DC%2C a modifier of lateral spikelet fertility in barley%2C is an ortholog of the maize domestication gene TEOSINTE BRANCHED 1%2E Nature genetics&dopt=abstract

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

https://scholar.google.com/scholar?as_q=INTERMEDIUM-C, a modifier of lateral spikelet fertility in barley, is an ortholog of the maize domestication gene TEOSINTE BRANCHED 1&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

https://scholar.google.com/scholar?as_q=INTERMEDIUM-C, a modifier of lateral spikelet fertility in barley, is an ortholog of the maize domestication gene TEOSINTE BRANCHED 1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ramsay,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Riggs%2C T%2EJ%2E and Kirby%2C E%2EJ%2EM%2E %281978%29%2E Developmental consequences of two%2Drow and six%2Drow ear type in spring barley%3A 1%2E Genetical analysis and comparison of mature plant characters%2E J%2E Agric%2E Sci%2E 91%3A 199%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Developmental consequences of two-row and six-row ear type in spring barley: 1&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

https://scholar.google.com/scholar?as_q=Developmental consequences of two-row and six-row ear type in spring barley: 1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Riggs,&as_ylo=1978&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ruiz%2DGarc�a%2C L%2E%2C Madueno%2C F%2E%2C Wilkinson%2C M%2E%2C Haughn%2C G%2E%2C Salinas%2C J%2E%2C and Mart�nez%2DZapater%2C J%2EM%2E %281997%29%2E Different Roles of Flowering%2DTime Genes in the Activation of Floral lnitiation Genes in Arabidopsis%2E Plant Cell 9%3A 1921%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Ruiz-García,&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=Different Roles of Flowering-Time Genes in the Activation of Floral lnitiation Genes 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

https://scholar.google.com/scholar?as_q=Different Roles of Flowering-Time Genes in the Activation of Floral lnitiation Genes in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ruiz-García,&as_ylo=1997&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ryu%2C M%2EY%2E%2C Cho%2C S%2EK%2E%2C and Kim%2C W%2ET%2E %282009%29%2E RNAi suppression of RPN12a decreases the expression of type%2DA ARRs%2C negative regulators of cytokinin signaling pathway%2C in Arabidopsis%2E Mol%2E Cells 28%3A 375%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=RNAi suppression of RPN12a decreases the expression of type-A ARRs, negative regulators of cytokinin signaling pathway, 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

https://scholar.google.com/scholar?as_q=RNAi suppression of RPN12a decreases the expression of type-A ARRs, negative regulators of cytokinin signaling pathway, in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ryu,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sasani%2C S%2E%2C Hemming%2C M%2EN%2E%2C Oliver%2C S%2EN%2E%2C Greenup%2C A%2E%2C Tavakkol%2DAfshari%2C R%2E%2C Mahfoozi%2C S%2E%2C Poustini%2C K%2E%2C Sharifi%2C H%2ER%2E%2C Dennis%2C E%2ES%2E%2C Peacock%2C W%2EJ%2E%2C and Trevaskis%2C B%2E %282009%29%2E The influence of vernalization and daylength on expression of flowering%2Dtime genes in the shoot apex and leaves of barley %28Hordeum vulgare%29%2E J%2E Exp%2E Bot%2E 60%3A 2169%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The influence of vernalization and daylength on expression of flowering-time genes in the shoot apex and leaves of barley (Hordeum vulgare)&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

https://scholar.google.com/scholar?as_q=The influence of vernalization and daylength on expression of flowering-time genes in the shoot apex and leaves of barley (Hordeum vulgare)&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sasani,&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schaller%2C G%2EE%2E%2C Street%2C I%2EH%2E%2C and Kieber%2C J%2EJ%2E %282014%29%2E Cytokinin and the cell cycle%2E Curr%2E Opin%2E Plant Biol%2E 21%3A 7%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Cytokinin and the cell cycle&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

https://scholar.google.com/scholar?as_q=Cytokinin and the cell cycle&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schaller,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schmitz%2C J%2E%2C Franzen%2C R%2E%2C Ngyuen%2C T%2EH%2E%2C Garcia%2DMaroto%2C F%2E%2C Pozzi%2C C%2E%2C Salamini%2C F%2E%2C and Rohde%2C W%2E %282000%29%2E Cloning%2C mapping and expression analysis of barley MADS%2Dbox genes%2E Plant Mol%2E Biol%2E 42%3A 899%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Cloning, mapping and expression analysis of barley MADS-box genes&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

https://scholar.google.com/scholar?as_q=Cloning, mapping and expression analysis of barley MADS-box genes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schmitz,&as_ylo=2000&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schmitz%2C R%2EJ%2E%2C Sung%2C S%2E%2C and Amasino%2C R%2EM%2E %282008%29%2E Histone arginine methylation is required for vernalization%2Dinduced epigenetic silencing of FLC in winter%2Dannual Arabidopsis thaliana%2E Proc%2E Natl%2E Acad%2E Sci%2E U%2E S%2E A%2E 105%3A 411%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Histone arginine methylation is required for vernalization-induced epigenetic silencing of FLC in winter-annual Arabidopsis thaliana&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

https://scholar.google.com/scholar?as_q=Histone arginine methylation is required for vernalization-induced epigenetic silencing of FLC in winter-annual Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schmitz,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Shannon%2C S%2E and Ry Meeks%2DWagner%2C O%2E %281991%29%2E A Mutation in the Arabidopsis TFL1 Gene Affects Inf lorescence Meristem Development%2E Plant Cell 3%3A 877%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A Mutation in the Arabidopsis TFL1 Gene Affects Inf lorescence Meristem 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

https://scholar.google.com/scholar?as_q=A Mutation in the Arabidopsis TFL1 Gene Affects Inf lorescence Meristem Development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Shannon,&as_ylo=1991&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Steinbach%2C Y%2E and Hennig%2C L%2E %282014%29%2E Arabidopsis MSI1 functions in photoperiodic flowering time control%2E Front%2E Plant Sci&dopt=abstract



Supek, F., Bošnjak, M., Škunca, N., and Šmuc, T. (2011). Revigo summarizes and visualizes long lists of gene ontology terms. PLoSOne 6: e21800


Tamaki, S., Matsuo, S., Wong, H.L., Yokoi, S., and Shimamoto, K. (2007). Hd3a Protein Is a Mobile Flowering Signal in Rice. Science 316:1033–1036.


Taoka, K. et al. (2011). 14-3-3 proteins act as intracellular receptors for rice Hd3a florigen. Nature 476: 332–335.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Teichmann, T. and Muhr, M. (2015). Shaping plant architecture. Front. Plant Sci. 6: 1–18.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Theißen, G. (2001). Development of floral organ identity: Stories from the MADS house. Curr. Opin. Plant Biol. 4: 75–85.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Turck, F., Fornara, F., and Coupland, G. (2008). Regulation and Identity of Florigen: FLOWERING LOCUS T Moves Center Stage. Annu.Rev. Plant Biol. 59: 573–594.


Turnbull, C.G.N. (2005). Plant Architecture and Its Manipulation (Blackwell).

Waddington, S.R., Cartwright, P.M., and Wall, P.C. (1983). A Quantitative Scale of Spike Initial and Pistil Development in Barley andWheat. Ann. Bot. 51: 119–130.


Wang, X., Gingrich, D.K., Deng, Y., and Hong, Z. (2012). A nucleostemin-like GTPase required for normal apical and floral meristemdevelopment in Arabidopsis. Mol. Biol. Cell 23: 1446–56.


Wang, X., Zhang, Y., Ma, Q., Zhang, Z., Xue, Y., Bao, S., and Chong, K. (2007). SKB1-mediated symmetric dimethylation of histone H4R3controls flowering time in Arabidopsis. EMBO J. 26: 1934–1941.


Wigge, P.A., Kim, M.C., Jaeger, K.E., Busch, W., Schmid, M., Lohmann, J.U., and Weigel, D. (2005). Integration of Spatial and TemporalInformation During Floral Induction in Arabidopsis. Science 309: 1056–1059.


Yan, L., Fu, D., Li, C., Blechl, A., Tranquilli, G., Bonafede, M., Sanchez, A., Valarik, M., Yasuda, S., and Dubcovsky, J. (2006). The wheatand barley vernalization gene VRN3 is an orthologue of FT. Proc. Natl. Acad. Sci. 103: 19581–19586.


Yeung, K., Seitz, T., Li, S., Janosch, P., McFerran, B., Kaiser, C., Fee, F., Katsanakis, K.D., Rose, D.W., Mischak, H., Sedivy, J.M., andKolch, W. (1999). Suppression of Raf-1 kinase activity and MAP kinase signalling by RKIP. Nature 401: 173–177.


Youssef, H.M. et al. (2016). VRS2 regulates hormone-mediated inflorescence patterning in barley. Nat. Genet. 49: 157–161.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Zagotta, M.T., Hicks, K.A., Jacobs, C.I., Young, J.C., Hangarter, R.P., and Meeks-Wagner, D.R. (1996). The Arabidopsis ELF3 generegulates vegetative photomorphogenesis and the photoperiodic induction of flowering. Plant J. 10: 691–702.



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

https://scholar.google.com/scholar?as_q=Arabidopsis MSI1 functions in photoperiodic flowering time control&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

https://scholar.google.com/scholar?as_q=Arabidopsis MSI1 functions in photoperiodic flowering time control&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Steinbach,&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Supek%2C F%2E%2C Bošnjak%2C M%2E%2C Škunca%2C N%2E%2C and Šmuc%2C T%2E %282011%29%2E Revigo summarizes and visualizes long lists of gene ontology terms%2E PLoS One 6%3A e&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Revigo summarizes and visualizes long lists of gene ontology terms.&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

https://scholar.google.com/scholar?as_q=Revigo summarizes and visualizes long lists of gene ontology terms.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Supek,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Tamaki%2C S%2E%2C Matsuo%2C S%2E%2C Wong%2C H%2EL%2E%2C Yokoi%2C S%2E%2C and Shimamoto%2C K%2E %282007%29%2E Hd3a Protein Is a Mobile Flowering Signal in Rice%2E Science 316%3A 1033%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Hd3a Protein Is a Mobile Flowering Signal 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

https://scholar.google.com/scholar?as_q=Hd3a Protein Is a Mobile Flowering Signal in Rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tamaki,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Taoka%2C K%2E et al%2E %282011%29%2E 14%2D3%2D3 proteins act as intracellular receptors for rice Hd3a florigen%2E Nature 476%3A 332%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=14-3-3 proteins act as intracellular receptors for rice Hd3a florigen&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

https://scholar.google.com/scholar?as_q=14-3-3 proteins act as intracellular receptors for rice Hd3a florigen&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Taoka,&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Teichmann%2C T%2E and Muhr%2C M%2E %282015%29%2E Shaping plant architecture%2E Front%2E Plant Sci%2E 6%3A 1%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Shaping plant architecture&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

https://scholar.google.com/scholar?as_q=Shaping plant architecture&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Teichmann,&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Thei�en%2C G%2E %282001%29%2E Development of floral organ identity%3A Stories from the MADS house%2E Curr%2E Opin%2E Plant Biol%2E 4%3A 75%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Development of floral organ identity: Stories from the MADS house&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

https://scholar.google.com/scholar?as_q=Development of floral organ identity: Stories from the MADS house&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Thei�?en,&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Turck%2C F%2E%2C Fornara%2C F%2E%2C and Coupland%2C G%2E %282008%29%2E Regulation and Identity of Florigen%3A FLOWERING LOCUS T Moves Center Stage%2E Annu%2E Rev%2E Plant Biol%2E 59%3A 573%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Regulation and Identity of Florigen: FLOWERING LOCUS T Moves Center Stage&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

https://scholar.google.com/scholar?as_q=Regulation and Identity of Florigen: FLOWERING LOCUS T Moves Center Stage&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Turck,&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Waddington%2C S%2ER%2E%2C Cartwright%2C P%2EM%2E%2C and Wall%2C P%2EC%2E %281983%29%2E A Quantitative Scale of Spike Initial and Pistil Development in Barley and Wheat%2E Ann%2E Bot%2E 51%3A 119%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A Quantitative Scale of Spike Initial and Pistil Development in Barley and Wheat&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

https://scholar.google.com/scholar?as_q=A Quantitative Scale of Spike Initial and Pistil Development in Barley and Wheat&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Waddington,&as_ylo=1983&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wang%2C X%2E%2C Gingrich%2C D%2EK%2E%2C Deng%2C Y%2E%2C and Hong%2C Z%2E %282012%29%2E A nucleostemin%2Dlike GTPase required for normal apical and floral meristem development in Arabidopsis%2E Mol%2E Biol%2E Cell 23%3A 1446%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A nucleostemin-like GTPase required for normal apical and floral meristem 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

https://scholar.google.com/scholar?as_q=A nucleostemin-like GTPase required for normal apical and floral meristem development in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wang,&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wang%2C X%2E%2C Zhang%2C Y%2E%2C Ma%2C Q%2E%2C Zhang%2C Z%2E%2C Xue%2C Y%2E%2C Bao%2C S%2E%2C and Chong%2C K%2E %282007%29%2E SKB1%2Dmediated symmetric dimethylation of histone H4R3 controls flowering time in Arabidopsis%2E EMBO J%2E 26%3A 1934%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=SKB1-mediated symmetric dimethylation of histone H4R3 controls flowering time 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

https://scholar.google.com/scholar?as_q=SKB1-mediated symmetric dimethylation of histone H4R3 controls flowering time in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wang,&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wigge%2C P%2EA%2E%2C Kim%2C M%2EC%2E%2C Jaeger%2C K%2EE%2E%2C Busch%2C W%2E%2C Schmid%2C M%2E%2C Lohmann%2C J%2EU%2E%2C and Weigel%2C D%2E %282005%29%2E Integration of Spatial and Temporal Information During Floral Induction in Arabidopsis%2E Science 309%3A 1056%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Integration of Spatial and Temporal Information During Floral Induction 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

https://scholar.google.com/scholar?as_q=Integration of Spatial and Temporal Information During Floral Induction in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wigge,&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Yan%2C L%2E%2C Fu%2C D%2E%2C Li%2C C%2E%2C Blechl%2C A%2E%2C Tranquilli%2C G%2E%2C Bonafede%2C M%2E%2C Sanchez%2C A%2E%2C Valarik%2C M%2E%2C Yasuda%2C S%2E%2C and Dubcovsky%2C J%2E %282006%29%2E The wheat and barley vernalization gene VRN3 is an orthologue of FT%2E Proc%2E Natl%2E Acad%2E Sci%2E 103%3A 19581%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The wheat and barley vernalization gene VRN3 is an orthologue of FT&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

https://scholar.google.com/scholar?as_q=The wheat and barley vernalization gene VRN3 is an orthologue of FT&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Yan,&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Yeung%2C K%2E%2C Seitz%2C T%2E%2C Li%2C S%2E%2C Janosch%2C P%2E%2C McFerran%2C B%2E%2C Kaiser%2C C%2E%2C Fee%2C F%2E%2C Katsanakis%2C K%2ED%2E%2C Rose%2C D%2EW%2E%2C Mischak%2C H%2E%2C Sedivy%2C J%2EM%2E%2C and Kolch%2C W%2E %281999%29%2E Suppression of Raf%2D1 kinase activity and MAP kinase signalling by RKIP%2E Nature 401%3A 173%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Suppression of Raf-1 kinase activity and MAP kinase signalling by RKIP&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

https://scholar.google.com/scholar?as_q=Suppression of Raf-1 kinase activity and MAP kinase signalling by RKIP&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Yeung,&as_ylo=1999&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Youssef%2C H%2EM%2E et al%2E %282016%29%2E VRS2 regulates hormone%2Dmediated inflorescence patterning in barley%2E Nat%2E Genet%2E 49%3A 157%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=VRS2 regulates hormone-mediated inflorescence patterning in barley&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

https://scholar.google.com/scholar?as_q=VRS2 regulates hormone-mediated inflorescence patterning in barley&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Youssef,&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zagotta%2C M%2ET%2E%2C Hicks%2C K%2EA%2E%2C Jacobs%2C C%2EI%2E%2C Young%2C J%2EC%2E%2C Hangarter%2C R%2EP%2E%2C and Meeks%2DWagner%2C D%2ER%2E %281996%29%2E The Arabidopsis ELF3 gene regulates vegetative photomorphogenesis and the photoperiodic induction of flowering%2E Plant J%2E 10%3A 691%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The Arabidopsis ELF3 gene regulates vegetative photomorphogenesis and the photoperiodic induction of flowering&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

https://scholar.google.com/scholar?as_q=The Arabidopsis ELF3 gene regulates vegetative photomorphogenesis and the photoperiodic induction of flowering&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zagotta,&as_ylo=1996&as_allsubj=all&hl=en&c2coff=1


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