1 short title: synthesis of gdp-l-fucose is required for … · 2020. 6. 4. · 1 1 short title:...

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SHORT TITLE: Synthesis of GDP-L-fucose is required for stomatal closure. 1

TITLE: Genetic screen to saturate guard cell signaling network reveals a role of GDP-L-fucose 2

metabolism in stomatal closure. 3

AUTHORS 4

Cezary Waszczak,a Triin Vahisalu,a Dmitry Yarmolinsky,b Maija Sierla,a Olena Zamora,b Marina Leal 5

Gavarrón,a Julia Palorinne,a Ross Carter,c Ashutosh K. Pandey,b Maris Nuhkat,b Melanie Carmody,a 6

Tuomas Puukko,a Nina Sipari,a,d Airi Lamminmäki,a Jörg Durner,e Dieter Ernst,e J. Barbro Winkler,f 7

Lars Paulin,g Petri Auvinen,g Andrew J. Fleming,h Jarkko Salojärvi,a,i Hannes Kollist,b Jaakko 8

Kangasjärvi,a 9

a Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental 10

Sciences, Viikki Plant Science Centre, University of Helsinki, FI-00014, Helsinki, Finland 11 b Institute of Technology, University of Tartu, 50411 Tartu, Estonia 12 c Sainsbury Laboratory, University of Cambridge, Bateman Street, Cambridge, CB2 1LR, UK 13 d Viikki Metabolomics Unit, Faculty of Biological and Environmental Sciences, University of Helsinki, 14

FI-00014 Helsinki, Finland 15 e Institute of Biochemical Plant Pathology, Helmholtz Zentrum München, German Research Center for 16

Environmental Health, 85764 Neuherberg, Germany 17 f Research Unit Environmental Simulation, Institute of Biochemical Plant Pathology, Helmholtz 18

Zentrum München, German Research Center for Environmental Health, 85764 Neuherberg, Germany 19 g Institute of Biotechnology, University of Helsinki, FI-00014 Helsinki, Finland. 20 h Department of Animal and Plant Sciences, University of Sheffield, Western Bank, S10 2TN Sheffield, 21

UK 22 i School of Biological Sciences, Nanyang Technological University, 637551 Singapore, Singapore 23

The author responsible for distribution of materials integral to the findings presented in this article in 24

accordance with the policy described in the Instructions for Authors (www.plantcell.org) is: 25

Jaakko Kangasjärvi ([email protected]), University of Helsinki. 26

Corresponding author email: [email protected] 27

was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. The copyright holder for this preprint (whichthis version posted June 7, 2020. ; https://doi.org/10.1101/2020.06.04.134353doi: bioRxiv preprint

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ABSTRACT 28

Guard cells regulate plant gas exchange by controlling the aperture of stomatal pores. The process 29

of stomatal closure involves a multi-input signaling network that governs the activity of ion channels, 30

which in turn regulate guard cell turgor pressure and volume. Here we describe a forward genetic 31

screen to identify novel components involved in stomatal movements. Through an ozone-sensitivity 32

approach combined with whole-rosette gas exchange analysis, 130 mutants of established stomatal 33

regulators and 76 novel mutants impaired in stomatal closure were identified. One of the novel mutants 34

was mapped to MURUS1 (MUR1), the first enzyme in de novo GDP-L-fucose biosynthesis. Defects in 35

synthesis or import of GDP-L-Fuc into the Golgi apparatus resulted in impaired stomatal closure to 36

multiple stimuli. Stomatal phenotypes observed in mur1 were independent from the canonical guard 37

cell signaling and instead could be related to altered mechanical properties of guard cell walls. 38

Impaired fucosylation of xyloglucan, N-linked glycans and arabinogalactan proteins did not explain the 39

aberrant function of mur1 stomata, however our data suggest that the stomatal phenotypes observed in 40

mur1 can at least partially be attributed to defective dimerization of rhamnogalactouronan-II. In 41

addition to providing the genetic framework for future studies on guard cell signaling, our work 42

emphasizes the impact of fucose metabolism on stomatal movement. 43


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INTRODUCTION 44

Stomata are epidermal pores surrounded by pairs of guard cells that balance the loss of water and 45

uptake of CO2 for photosynthesis. Guard cells respond to multiple environmental factors e.g. light, CO2 46

concentration, drought, low humidity, pathogens and air pollutants such as ozone (O3), to optimize 47

transpiration or prevent the entry of the pathogens into the leaf tissue. Accumulation of reactive oxygen 48

species (ROS) in the apoplast of guard cells and subsequent activation of plasma membrane Ca2+in 49

channels are among the first events associated with execution of stomatal closure (McAinsh et al., 50

1996; Pei et al., 2000; Kwak et al., 2003). 51

Depending on the stimulus, the apoplastic ROS are generated by NADPH oxidases (Kwak et al., 2003; 52

Kadota et al., 2014), apoplastic peroxidases and amine oxidases (Sierla et al., 2016), however, external 53

application of ROS alone is sufficient to initiate the process of stomatal closure (Price, 1990; McAinsh 54

et al., 1996; Kollist et al., 2007). Hydrogen peroxide (H2O2) is the most stable form of ROS (Waszczak 55

et al., 2018). Apoplastic perception of H2O2 involves activation of HYDROGEN 56

PEROXIDE-INDUCED Ca2+ INCREASES1 (HPCA1) leucine-rich repeat receptor kinase which is 57

necessary for activation of Ca2+in channels (Wu et al., 2020). The subsequent rise in cytoplasmic Ca2+ 58

concentration activates multiple Ca2+-dependent protein kinases (Maierhofer et al., 2014; Brandt et al., 59

2015) that together with OPEN STOMATA1 (OST1) kinase (Geiger et al., 2009; Lee et al., 2009) 60

phosphorylate and activate guard cell anion channels SLOW ANION CHANNEL-ASSOCIATED1 61

(SLAC1), QUICK-ACTIVATING ANION CHANNEL1 (QUAC1) and SLAC1 HOMOLOGUE3 62

(SLAH3; for full list of kinases see Sierla et al., (2016)). The activation of guard cell anion channels, 63

accompanied by deactivation of H+-ATPase1 (AHA1; Merlot et al., 2007) leads to membrane 64

depolarization and activation of K+out channels (Hedrich, 2012). The efflux of ions into the apoplast 65

leads to a decrease of osmotic pressure inside the guard cells which provokes an efflux of H2O from the 66

guard cell cytoplasm and vacuole. The consequent drop in guard cell turgor pressure results in closure 67

of stomatal pores (Franks et al., 1998). 68

As evidenced by measurements of guard cell volume and turgor pressure performed in broad bean 69

(Vicia faba), during stomatal opening guard cell turgor pressure rises from as low as 0.3 MPa to 5 MPa 70

which is accompanied by a 30-40 % increase in guard cell volume (Franks et al., 2001). Importantly, 71

the expansion and flexing of guard cells has to overcome the turgor pressure of the subsidiary cells (for 72

a detailed discussion of the role of subsidiary cells see Lawson and Matthews, (2020)). On the other 73

hand, stomatal closure involves a significant decrease in guard cell volume and surface area (Shope et 74


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al., 2003). To allow these volume and pressure changes, the guard cell walls must have a high degree of 75

plasticity, which must be determined by wall structure. While differences between taxa exist (Popper, 76

2008), plant primary cell walls are typically composed of cellulose, hemicelluloses (xyloglucan, xylan, 77

mannan), structural proteins, and pectins such as homogalactouronan (HG), rhamnogalactouronan-I 78

(RG-I) and rhamnogalactouronan-II (RG-II) that determine the cell wall elasticity (Liepman et al., 79

2010). However, the relative content of these components varies between cell types and depends on 80

the developmental stage. Moreover, the spatial distribution of different cell wall components is not 81

uniform and reflects the mechanical needs of the respective cell type. Guard cells are an excellent 82

example of such specialization. In comparison to epidermal cells, the guard cell walls are significantly 83

thicker, devoid of highly methyl-esterified HG and rich in un-esterified HG (Amsbury et al., 2016; 84

Merced and Renzaglia, 2018). The degree of methyl esterification is inversely correlated with the 85

ability for Ca2+-mediated crosslinking, which leads to a more rigid cell wall, as well as susceptibility to 86

degradation by polygalacturonases that leads to cell wall loosening (Levesque-Tremblay et al., 2015). 87

Plants deficient in pectin demethylesterification exhibit defects in stomatal closure (Amsbury et al., 88

2016) which suggests that pectin crosslinking/degradation has a profound effect on the execution of 89

this process. Further, targeted enzymatic digestion of arabinans that typically constitute the side chains 90

of RG-I inhibits stomatal movements, and this effect can be counteracted by subsequent digestion or 91

depolymerization of HG (Jones et al., 2003) . Based on this observation, Jones et al., (2003) proposed 92

a model in which the arabinan side chains of RG-I prevent crosslinking of HG strands which otherwise 93

increases the cell wall rigidity, making it less capable to react to changes in guard cell turgor. Further, 94

as observed already in the first half of the 20th century (see Shtein et al., (2017) for recent visualization) 95

the cellulose microfibrils within the guard cell walls fan out radially from the pore to provide a hoop 96

reinforcement that limits the increase in guard cell radius and promotes guard cell elongation during the 97

stomatal opening (Woolfenden et al., 2017). Moreover, as inferred from the atomic force microscopy 98

(AFM)-based studies of guard cells, the stiffness of guard cell walls is not uniform; the most rigid areas 99

are localized at the poles of guard cells and ventral walls directly surrounding the pore (Carter et al., 100

2017). During stomatal opening the polar fixing prevents the increase in the length of stomatal complex 101

and forces the elongating guard cells to bend, leading to an increase in pore aperture (Carter et al., 102

2017). Taken together, it is clear, that the mechanics of the guard cell wall has a profound role in the 103

execution of stomatal movements (Rui et al., 2018; Woolfenden et al., 2018). 104


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The synthesis of cell wall glycan polymers relies on the availability of nucleotide sugars that constitute 105

the activated precursor forms serving as a donor of sugar moieties (Bar-Peled and O’Neill, 2011). 106

The importance of nucleotide sugar synthesis and transport is exemplified by the requirement of 107

GDP-L-fucose for proper growth and development (Reiter et al., 1993; Reiter et al., 1997; O’Neill et 108

al., 2001; Van Hengel and Roberts, 2002; Rautengarten et al., 2016), resistance to pathogens (Zhang et 109

al., 2019) and freezing tolerance (Panter et al., 2019). The synthesis of GDP-L-Fuc is initiated by 110

GDP-D-mannose 4,6-dehydratases (GMD1) and MURUS1 (MUR1/GMD2) that catalyze the 111

conversion of GDP-D-mannose to GDP-4-keto-6-deoxy-D-mannose (Figure 1; Bonin et al., 1997; 112

Bonin et al., 2003). In leaf tissues, MUR1 is the major GMD isoform and the level of L-Fuc observed 113

in cell walls of mur1 mutants is reduced by approximately 98% as compared to wild type plants (Reiter 114

et al., 1993). In aerial organs GMD1 is expressed only in stipules and pollen grains (Bonin et al., 2003) 115

and plays a minimal role in the synthesis of GDP-L-Fuc in leaf tissue. 116

Plants lacking MUR1 exhibit dwarfism, reduced apical dominance, brittle stems (Reiter et al., 1993), 117

short root phenotype (Van Hengel and Roberts, 2002), altered lignin structure and inflorescence stem 118

development (Voxeur et al., 2017), and sensitivity to freezing (Panter et al., 2019) and pathogens 119

(Zhang et al., 2019). The product of the MUR1-catalyzed reaction, GDP-4-keto-6-deoxy-D-mannose, 120

serves as a substrate for the GDP-4-keto-6-deoxymannose-3,5-epimerase-4-reductases GER1 (Bonin 121

and Reiter, 2000; Nakayama et al., 2003) and GER2 (Rhomberg et al., 2006) that complete the 122

synthesis of GDP-L-Fuc (Figure 1). Another pathway that leads to synthesis of GDP-L-Fuc (the L-123

fucose salvage pathway) involves a single bifunctional enzyme L-FUCOKINASE/GDP-L-FUCOSE 124

PYROPHOSPHORYLASE (FKGP) that converts L-Fuc to GDP-L-Fuc (Figure 1; Kotake et al., 2008). 125

In contrast to mur1, the fkgp mutants exhibit normal growth phenotype and cell wall composition, 126

suggesting that the L-fucose salvage pathway has a minor role in Arabidopsis (Kotake et al., 2008). 127

Following synthesis in the cytoplasm, GDP-L-Fuc is transported into the Golgi lumen by the 128

GDP-FUCOSE TRANSPORTER1 (GFT1; Figure 1; Rautengarten et al., 2016). Knock-out mutants of 129

GFT1 are not viable, and GFT1 knockdown plants exhibit semi-lethal phenotypes similar to those of 130

mur1 mutants (Rautengarten et al., 2016). Within the Golgi lumen, GDP-L-Fuc serves as a substrate for 131

fucosyltransferases (FUT) that fucosylate components of the cell wall (Figure 1). There are 13 132

fucosyltransferases (FUT1 – FUT13) in Arabidopsis (Sarria et al., 2001; Wilson et al., 2001), and 133

according to the current state of knowledge, all are either confirmed or expected to localize to the Golgi 134

membrane with the catalytic domain facing the lumen (Sarria et al., 2001; Strasser, 2016). FUTs add 135


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L-Fuc residues onto molecules such as xyloglucan (FUT1/MUR2)(Perrin et al., 1999; Vanzin et al., 136

2002), arabinogalactan proteins (FUT4, FUT6; Wu et al., 2010; Liang et al., 2013; Tryfona et al., 2014) 137

and N-linked glycans (FUT11/FUCTA, FUT12/FUCTB, FUT13/FUCTC; Leonard et al., 2002; 138

Strasser et al., 2004). Furthermore, L-Fuc is found in RG-I and RG-II, although the FUTs involved in 139

the synthesis of these pectins are not known. In terms of quantity, the majority of L-Fuc is found in 140

RG-I (Anderson et al., 2012). Importantly, the dwarf phenotype of mur1 mutants has been previously 141

attributed to deficiency in boron-dependent dimerization of RG-II (O’Neill et al., 2001). In mur1, RG-142

II L-Fuc residues are replaced by L-galactose (Zablackis et al., 1996) which leads to an approximately 143

50% decrease in RG-II dimer formation (O’Neill et al., 2001) possibly caused by RG-II chain A 144

truncation (Pabst et al., 2013). 145

To understand processes controlling stomatal movements, O3 can be used as an apoplastic ROS donor 146

to stimulate stomatal closure (Kollist et al., 2007; Vahisalu et al., 2010). Ozone enters plants through 147

stomata and subsequently decomposes to various ROS that further provoke active production of ROS 148

by plant cells (Vainonen and Kangasjärvi, 2014). Plants deficient in O3-induced stomatal closure 149

receive high doses of O3 that trigger formation of visible hypersensitive response-like lesions. These 150

lesions are easy to score and allow the identification of stomatal mutants in forward genetic approaches 151

(Overmyer et al., 2000). Previously, such a genetic screen led to the identification of several proteins 152

involved in stomatal closure, i.e., SLOW ANION CHANNEL1 (SLAC1; Vahisalu et al., 2008), the 153


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receptor-like pseudokinase GUARD CELL HYDROGEN PEROXIDE-RESISTANT1 (GHR1) that 154

participates in activation of SLAC1 (Sierla et al., 2018), and HIGH TEMPERATURE1 (HT1) kinase 155

which acts in high CO2-induced guard cell signaling as an inhibitor of SLAC1 activation (Hõrak et al., 156

2016). Here we present an O3 exposure-based forward genetic screen specifically aimed at the 157

identification of novel components involved in stomatal closure. We report the isolation of 76 novel 158

mutants, and 130 new mutants of established stomatal regulators. In particular, we describe the 159

identification of the MUR1 mutant and show that synthesis and import of GDP-L-fucose into the Golgi 160

lumen play an important role in stomatal closure. Our results are consistent with the hypothesis that the 161

stomatal deficiencies observed in mur1 mutants are due to altered mechanical properties of the guard 162

cell walls, and we propose that reduced dimerization of rhamnogalactouronan-II contributes to the 163

impaired stomatal function observed in mur1 mutants. 164


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165


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RESULTS 166

A forward genetic screen identifies novel regulators of guard cell signaling 167

To fill the gaps in known guard cell signaling networks, we have performed forward genetic screen 168

based on an O3-sensitivity (Figure 2). This screen is fully independent from our previous screen 169

(Overmyer et al., 2000). A total of 125,000 seeds of Arabidopsis line pGC1:YC3.6 (Yang et al., 2008), 170

expressing yellow cameleon 3.6 (YC3.6; Nagai et al., 2004), a biosensor probe that allows for imaging 171

of intracellular Ca2+, were treated with ethyl methanesulfonate (EMS). Later, 380,000 M2 plants were 172

exposed to O3. Individual plants displaying visible O3-damage were then subjected to thermal imaging 173

and water loss assay (see Materials and Methods). The progeny of the most prominent mutants 174

(approximately 3,200 lines) was subjected to a secondary screen utilizing the same screening methods. 175

Lines for which at least one of the phenotypes was clearly confirmed (551 lines) were then analyzed 176

with a whole-rosette gas exchange system (Kollist et al., 2007) to investigate stomatal responses to a 177

variety of stimuli inducing stomatal closure i.e. apoplastic ROS (delivered by means of O3-fumigation), 178

5 µM ABA spray, high CO2 and reduced air humidity which is often termed as vapor pressure deficit 179

(VPD; Merilo et al., 2018). Lines demonstrating lack, or impairment of stomatal responses to at least a 180

single stimulus (206 lines) were subjected to targeted sequencing of genomic regions encoding 22 181

well-established stomatal regulators, hereafter referred to as “usual suspects”, to avoid potential 182

re-discoveries (Figure 2). The list of usual suspects included ion channels, ABA biosynthesis enzymes, 183

protein kinases, phosphatases, and other proteins where the corresponding mutant lines are known to 184

exhibit impaired stomatal closure and/or higher stomatal conductance (see Supplementary Table 1 for 185

a full list). Approximately 60% of the tested lines (130) had mutations in coding regions of at least one 186

usual suspect. Most frequently mutations were identified within AHA1, GHR1 (Sierla et al., 2018), 187

MITOGEN-ACTIVATED PROTEIN KINASE12 (MPK12) and MORE AXILLARY BRANCHES2 188

(MAX2) coding sequences (Figure 2). Among the 76 mutants with no mutations in usual suspects, the 189

most frequently observed stomatal phenotypes were impaired responses to elevated CO2 (48 out of 76) 190

and high loss of water from detached leaves (46 out of 76). Notably, the majority of newly identified 191

mutants were affected in stomatal responses to more than one stimulus (Figure 2). With the exception 192

of new alleles of HT1 and AHA1 that will be published elsewhere, all new alleles of the usual suspects 193

are listed in (Supplementary Data Set 1). Due to the high number of mutant lines, we were not able to 194

perform allelism tests for all new alleles of the usual suspects. However, published results (Sierla et al., 195


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2018), as well as currently ongoing experiments, indicate that in most of the lines the observed 196

phenotypes are linked to the mutations in the tested genes. Therefore, only the lines with no mutations 197

in the usual suspects were retained for further analysis. 198

Mapping of a novel mutant impaired in stomatal closure identifies MUR1 199

From the screen described above, mutant T7-9 exhibited higher water loss from detached leaves (water 200

loss) compared to wild type (Figure 3A), and partially impaired stomatal responses to O3, high CO2 201

concentration, ABA, and darkness, while retaining the ability to close stomata in response to high VPD 202

(Supplemental Figure 1). Throughout this paper, the water loss assay, also known as “mass loss of 203

detached leaves, MLD” (Duursma et al., 2019) is used as a simple indicator of stomatal function (if the 204

cuticle permeability is intact) and has proven reliable in our previous work on stomatal signaling 205

(Vahisalu et al., 2008; Hõrak et al., 2016; Sierla et al., 2018). To identify the causative mutation in 206

T7-9, we applied the SHOREmap backcross pipeline (Hartwig et al., 2012). For this, T7-9 was 207


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backcrossed to YC3.6. The resulting BC1F1 plants had a WT-like water loss indicating recessive 208


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inheritance. Approximately 21% of the BC1F2 plants (123 out of 593) exhibited increased water loss, 209

indicating that the trait was determined by a single locus. Nuclear DNA from the 123 BC1F2 plants 210

displaying the mutant phenotype was bulked and subjected to next generation sequencing (NGS). 211

Analysis of the NGS data with marker frequency threshold set to 0.9 resulted in identification of five 212

non-synonymous EMS-specific mutations enriched in BC1F2 plants displaying high water loss 213

(Supplementary Table 2). All polymorphisms localized to the lower arm of chromosome 3 within a 214

17.0 - 19.1 Mb physical interval. Screening of the mutant lines for the five candidate genes revealed 215

that three independent mutant lines: mur1-1, mur1-2 (Reiter et al., 1993; Bonin et al., 1997) and mur1-216

9 (SALK_057153, Supplementary Figure 2), carrying mutations within AT3G51160 (Figure 3B), 217

exhibited highly elevated water loss (Figure 3C). The mutant lines for the remaining candidate genes 218

had WT-like water loss (Supplemental Figure 3). AT3G51160 encodes GDP-mannose-4,6-dehydratase 219

MURUS1 (GMD2, MUR1) which catalyzes the first step in de novo biosynthesis of GDP-L-Fucose 220

(Figure 1; Bonin et al., 1997). Therefore, we investigated the level of GDP-L-Fuc in the T7-9 mutant. 221

Similarly to other mur1 mutants, we were not able to detect this metabolite in T7-9 suggesting the 222

complete loss of MUR1 enzymatic activity (Figure 3D). Finally, an allelism test between T7-9 and a 223

mur1-2 mutant revealed lack of complementation, confirming that the T7-9 MUR1 E175K mutation 224

(hereafter referred to as mur1-10) conferred its high water loss (Figure 3E). 225

MUR1 is involved in stomatal development 226

As multiple phenotypes observed in mur1 mutants have been attributed to abnormal cell wall 227

composition (Reiter et al., 1993; O’Neill et al., 2001; Van Hengel and Roberts, 2002), we first focused 228

on characterizing mur1 stomata and cuticle development. No phenotypes related to cuticle permeability 229

were detected in any of the four mur1 mutants by means of dye-exclusion assay (Supplemental Figure 230

4). To assess stomatal development, we performed scanning electron microscopy-based examination of 231

abaxial epidermis of mur1-1 and mur1-2 cotyledons. In agreement with an earlier report (Zeng et al., 232

2011), no consistent phenotype related to stomatal density was observed in mur1 mutants. The stomatal 233

density in mur1-2 mutant was higher than in mur1-1, which had a similar stomatal density to Col-0 234

(Supplementary Figure 5A). The average size of stomatal pores was moderately increased in the mur1-235

1 mutant, while in mur1-2 there were no significant differences compared to Col-0 (Supplementary 236

Figure 5B). Notably, in both mur1 mutants we observed a larger variability in size of stomatal pores 237

(Supplementary Figure 5C) and the biggest stomatal complexes (2-8% of stomata, Supplementary 238


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Figure 5D) were irregular in shape and often obstructed (Supplementary Figure 5E). Moreover, as 239


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observed before (Zeng et al., 2011; Zhang et al., 2019), a fraction of mur1 stomata exhibited aberrant 240

structure of the outer cuticular ledge. In the most severe cases (3-17% of stomata, Supplementary 241

Figure 5D) the outer stomatal ledges completely sealed the stomatal pores (Supplementary Figure 5F) 242

similarly as in mutants lacking FUSED OUTER CUTICULAR LEDGE1 (FOCL1), a proline-rich 243

protein necessary for formation of outer cuticular ledge (Hunt et al., 2017). Overall, despite the 244

differences in stomata size and morphology, the whole plant steady state stomatal conductance of the 245

mur1 mutants did not differ significantly from that of the wild type plants (Supplementary Figure 1, 246

Supplementary Figure 6, Figure 4I). 247

MUR1 is required for stomatal closure 248

To further characterize the stomatal function in mur1 mutants, we subjected them to a variety of 249

treatments provoking stomatal movements and followed the time-resolved whole-rosette stomatal 250

conductance (Kollist et al., 2007). Gas exchange measurements were performed for mur1-1 and mur1-2 251

mutants (Reiter et al., 1993; Bonin et al., 1997). For the treatments inducing stomatal closure, either 252

ghr1-3 (Sierla et al., 2018) or ost1-3 (Yoshida et al., 2002) were used as non-responsive controls, while 253

in stomata opening assays ht1-2 (Hashimoto et al., 2006) was used. Across all gas-exchange assays, 254

the stomatal responses of mur1 mutants were consistent (Supplementary Figures 6 and 7), therefore, 255

representative results obtained for mur1-1 allele are shown (Figure 4). As observed earlier in T7-9, 256

(Supplementary Figure 1) plants lacking MUR1 were impaired in the rapid transient decrease of 257

stomatal conductance in response to a three-minute O3 pulse (Figure 4A-B, Supplementary Figure 6A), 258

that otherwise induces a rapid decrease in Col-0 plants (Kollist et al., 2007; Vahisalu et al., 2010; Sierla 259

et al., 2018). This suggests that the activity of MUR1 is required for rapid stomatal movements in 260

response to ROS. Further, we observed impaired stomatal closure upon treatment with elevated CO2 261

concentration (800 µl l-1), 5 µM ABA spray or application of darkness during the light period (Figure 262

4C-H, Supplementary Figure 6B-D). Similarly, during diurnal light/dark cycles, the transition to 263

darkness induced a rapid drop in stomatal conductance of Col-0 plants while the mur1 mutants 264

exhibited a much less pronounced response, and maintained higher stomatal conductance during the 265

darkness period (Figure 4I). In contrast to these observations, as seen earlier in T7-9 (Supplementary 266

Figure 1), the stomatal responses of mur1 mutants to high VPD were not significantly different from 267

those of Col-0 plants (Supplementary Figure 6E-F). The stimuli provoking stomatal opening, such as 268

exposure to low CO2 concentration (400 → 100 µl l-1) or increase in light intensity (150 → 500 µmol 269


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m-2 s-1), did not show any differences between mur1 mutants and the wild type (Supplementary Figure 270

7) suggesting that MUR1 activity is not required for stomatal opening. Taken together, our data 271

indicate that MUR1 activity is required for normal stomatal closure in response to O3, high CO2 272

concentration, ABA and darkness, but not for stomatal opening. 273

Import of GDP-L-fucose into the Golgi apparatus is necessary for stomatal function 274

To further study the role of MUR1 in stomatal closure, we investigated stomatal function in mutants 275

impaired in the subsequent steps of the GDP-L-Fuc synthesis and transport (Figure 1). For this, two 276

T-DNA insertion mutants of GER1, (ger1-2, ger1-3) as well as GER2 (ger2-1, ger2-2) were isolated 277

(Supplementary Figure 2B, C) and subjected to a water loss assay. None of the tested ger mutants 278

exhibited elevated water loss, suggesting functional redundancy (Figure 5A). Similarly, the water loss 279

of fkgp mutants (fkgp-1, fkgp-2; Kotake et al., 2008) was similar to that of Col-0 (Figure 5A) indicating 280

that the L-fucose salvage pathway has little impact on stomatal closure. To further explore the possible 281

redundancy between GER1 and GER2, we attempted to generate a ger1 ger2 double mutant. To this 282

end, ger1-2 and ger1-3 were crossed with ger2-1 and ger2-2. However, no double homozygous plants 283

were found in any of the four F2 families. Similarly, the F3 progeny of F2 plants homozygous for ger1 284

but heterozygous for ger2 alleles (ger1-2-/- ger2-1+/-; ger1-2-/- ger2-2+/-; ger1-3-/- ger2-1+/-; ger1-3-/- 285

ger2-2+/-) did not contain double mutants. In every case, the observed segregation of ger2 alleles 286

(ger2+/+ : ger2+/- : ger2-/-) within the F3 families was 1 : 1 : 0 (Supplementary Table 3). We therefore 287

inspected both pollen viability and embryo development in F2 ger1-/- ger2+/- plants but did not find any 288

abnormalities. Hence, the inability to generate ger1 ger2 mutants might be related to defects in 289

fertilization with ger1- ger2- pollen. Thus, we focused on characterization of the GDP-L-Fuc 290

transporter GFT1. 291

Loss-of-function mutants of GFT1 are not viable, for that reason we utilized hairpin RNAi knockdown 292

plants (hpGFT1; (Rautengarten et al., 2016). A total of 66 independent hpGFT1 T1 plants were 293

selected and transplanted to soil. As described before, the hpGFT1 plants exhibited varying growth 294

phenotypes, i.e., reduced projected rosette area, short petioles, and wavy leaves (Supplementary Figure 295

8A) that correlated with the residual GFT1 transcript level and cell wall L-Fuc content (Rautengarten et 296

al., 2016). For each of the T1 hpGFT1 plant that survived in soil (64 plants) we measured the projected 297

rosette area and loss of water from detached leaves, as well as the GFT1 transcript level (58 plants). 298

The hpGFT1 T1 plants exhibited varying water loss (Figure 5B) and residual GFT1 transcript level 299


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(Supplementary Figure 8B), and displayed a clear negative correlation between these two traits (Figure 300


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5C). Similarly, the projected rosette area was also negatively correlated with the water loss 301

(Supplementary figure 8C). The majority of hpGFT1 plants with GFT1 transcript level lower than 10% 302

of that observed in the empty vector control (EVC) displayed water loss comparable to that of mur1 303

mutants (Figure 5C). Together, our data indicate that the import of GDP-L-Fuc into the Golgi lumen is 304

important for stomatal closure. 305

Lack of MUR1 affects mechanical properties of guard cell walls. 306

To investigate the genetic interactions between MUR1 and proteins regulating canonical stomata 307

closure pathways we crossed mur1-1 and mur1-2 mutants to slac1-4 (Vahisalu et al., 2008), aba2-11 308

(González-Guzmán et al., 2002), ost2-2D (Merlot et al., 2007), ost1-3 (Yoshida et al., 2002) and ghr1-3 309

(Sierla et al., 2018) and assayed the stomatal function of the double mutants via water loss assay. In 310

every double mutant an additive effect of combining two mutations (Figure 6A) indicated that the 311

phenotypes observed in mur1 plants were independent from the canonical guard cell signaling 312

pathways. Because of the additive effects (Figure 6A), the previously documented role of MUR1 in cell 313

wall development (Reiter et al., 1993; Reiter et al., 1997; O’Neill et al., 2001), as well as the general 314

deficiency in responses to stomata-closing stimuli observed in mur1 mutants (Figure 4, Supplementary 315

Figure 1, Supplementary Figure 6), we investigated the mechanical properties of mur1 guard cell walls 316

with atomic force microscopy (Carter et al., 2017). The patterning of the apparent modulus (Ea) in the 317

stomatal complexes of mur1 mutants was comparable to that of the control lines. However, comparison 318

of the absolute Ea values derived from the AFM scans indicated that mur1 mutants had significantly 319

stiffer subsidiary cells and this difference was even more pronounced when values obtained for guard 320

cell walls were compared (Figure 6B). Thus, taken together, our data indicate that the stomatal 321

phenotypes observed in mur1 mutants are most likely related to the altered mechanical properties of the 322

stomatal complexes. 323

mur1 stomatal phenotypes are not linked to xyloglucan structure. 324

Xyloglucan has been previously linked to stomatal movements (Rui and Anderson, 2016). To 325

investigate whether the phenotypes observed in mur1 mutants might be related to the lack of 326

xyloglucan fucosylation, we analyzed the stomatal function in mutants lacking FUT1/MUR2, the major 327

xyloglucan fucosyltransferase (Figure 1; Vanzin et al., 2002), and KATAMARI1/MURUS3 (KAM1, 328

MUR3) a xyloglucan galactosyltransferase (Madson et al., 2003). The cell wall fucose content in mur2-329

1, mur3-1 and mur3-2 mutants is reduced by approximately 50 % (Reiter et al., 1997). Unlike MUR3 330


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EMS mutants (mur3-1, mur3-2) the T-DNA insertion mutants mur3-3 and mur3-7 exhibit a dwarf 331


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rosette phenotype (Tamura et al., 2005; Kong et al., 2015) which can be rescued by removing the 332


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activity of XYLOSYLTRANSFERASE1 (XXT1) and XXT2 leading to plants devoid of xyloglucan 333

(Kong et al., 2015). Therefore, we assessed the stomatal function of mur2-1, all four mur3 mutants, as 334

well as a mur3-3 xxt1 xxt2 triple mutant (Kong et al., 2015) and an xxt1 xxt2 double mutant (Cavalier et 335

al., 2008). The water loss of mur2-1 was not significantly different from that observed for Col-0 336

control, indicating that the lack of xyloglucan fucosylation does not affect the stomatal closure process 337

(Figure 7A). All mur3 mutants exhibited a moderately elevated water loss, however, much lower than 338

that of mur1 mutants. The water loss of xyloglucan-deficient xxt1 xxt2 double mutant was similar to 339

that of Col-0 (Figure 7A). Therefore, we concluded that the stomatal phenotypes observed in mur1 340

mutants were not linked to defects in XyG fucosylation. 341

mur1 stomatal phenotypes are related to pectin structure 342

To investigate whether the stomatal phenotypes of mur1 were related to the dimerization of RG-II, we 343

grew mur1 mutants in soil supplemented with 1 mM borate. Such treatments were previously shown to 344

compensate for the deficiency in RG-II crosslinking (O’Neill et al., 2001). We observed a significant 345

decrease of water loss of mur1 mutants grown in the presence of 1 mM borate as compared to the 346

control conditions (Figure 7B). A similar trend was observed in Col-0, slac1-4 and ghr1-3 mutants, 347

albeit to a much lower extent. To validate this finding, we utilized plants lacking boron transporter 348

REQUIRES HIGH BORON1 (BOR1) required for boron xylem loading (Noguchi et al., 1997; Takano 349

et al., 2002) as previously the impaired uptake of boron was demonstrated to affect the dimerization of 350

RG-II (Miwa et al., 2013; Panter et al., 2019). Lack of BOR1 leads to impaired expansion of rosette 351

leaves which can be rescued by increase of B concentration in the growth medium (Noguchi et al., 352

1997). We found that a soil–grown bor1-3 mutant (Kasai et al., 2011) exhibited high water loss which 353

could be reverted by supplementing the soil with 50 µM borate, while lower concentrations (10 µM 354

and 20 µM) had no effect (Supplementary Figure 9). Thus, the data supported the hypothesis that the 355

phenotypes observed in mur1 plants were related to a deficiency in RG-II crosslinking. 356

To further investigate the role of RG-II in stomatal closure, we analyzed the stomatal function of 357

mutants lacking other enzymes involved in RG-II biosynthesis. The structure of RG-II is highly 358

complex and in Arabidopsis only few enzymes involved in its synthesis have been identified 359

(Funakawa and Miwa, 2015). Thus far, apart from enzymes involved in synthesis or transport of GDP-360

L-Fuc, the following proteins have been shown to play a role in RG-II synthesis: 361

RHAMNOGALACTURONAN XYLOSYLTRANSFERASE1 (RGXT1), RGXT2 (Egelund et al., 362


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2006), RGXT3 (Egelund et al., 2008), RGXT4/MGP4 (Liu et al., 2011) catalyzing the transfer of D-363

xylose onto L-fucose, KDO-8-P SYNTHASE1 (AtKDSA1) and AtKDSA2, required for synthesis of 3-364

deoxy-d-manno-octulosonate (KDO; Matsuura et al., 2003; Delmas et al., 2008); and GOLGI GDP‐L‐365

GALACTOSE TRANSPORTER1 (GGLT1) required for import of GDP-L-galactose into the Golgi 366

apparatus (Sechet et al., 2018). Deficiency in RGXT4 or KDSA activity leads to defective pollen tube 367

formation, preventing the generation of mutant lines (Delmas et al., 2008; Liu et al., 2011). Therefore, 368

we investigated the water loss of plants deficient in RGXT1 (rgxt1-1), RGXT2 (rgxt2-1; Egelund et al., 369

2006), KDSA1 (AtkdsA1-S), KDSA2 (AtkdsA2-S; Delmas et al., 2008) and three GGLT1 RNAi 370

knockdown lines (Sechet et al., 2018); but no elevated water loss was observed in any of the tested 371

lines (Supplementary Figure 10). It should be noted that, under our growth conditions, none of the 372

tested mutants phenocopied the mur1 rosette phenotype, which suggests functional redundancy 373

between the members of the respective gene families (Delmas et al., 2008) or different structural 374

consequences for the cell wall caused by the alterations of RG-II composition in the respective mutants. 375

In addition, we investigated whether mur1 stomatal phenotypes might be related to lack of fucosylation 376

of other cell wall components. For this, we measured water loss of mutants deficient in FUT4 and 377

FUT6 (fut4 fut6; Tryfona et al., 2014), FUT11 and FUT12 (fucta fuctb; Strasser et al., 2004), FUT13 378

(fuctc-1; Rips et al., 2017) as well as other fucosyltransferases with yet unidentified targets for which 379

the mutant lines were available: fut2-1, fut3-1, fut8-1, fut8-2, fut10-1 and fut10-2 (Supplementary 380

Figure 11). We did not detect FUT9 transcript in 2-week-old seedlings, and at the time of the analysis 381

no mutant lines for FUT5 and FUT7 were available, therefore FUT5, FUT7 and FUT9 were not 382

included in these experiments. The water loss of fut4 fut6, fucta fuctb and fuctc-1 mutants was 383

comparable to that of Col-0 (Figure 7C), suggesting that mur1 stomatal deficiencies are not related to 384

lack of fucosylation of arabinogalactan proteins or N-linked glycans. Similarly, mutants of the 385

remaining fucosyltransferases did not show elevated water loss, which likely reflects a high degree of 386

functional redundancy within the FUT family. 387

388


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DISCUSSION 389

Forward genetic screen identifies new components of guard cell signaling network 390

Stomatal movements are coordinated by multiple signaling pathways that converge to regulate the 391

activity of guard cell tonoplast and plasma membrane ion channels. Over the last three decades, many 392

components of guard cell signaling networks have been identified (Hedrich, 2012; Sierla et al., 2016; 393

Ehonen et al., 2019; Lawson and Matthews, 2020), however multiple key components await 394

characterization. Here we describe a forward genetic screen that aims to saturate the gene network 395

controlling stomatal closure. Our results indicate that the current gene network is far from complete as 396

we identified 76 mutants affected in stomatal closure (Figure 2) that do not represent mutations in any 397

of the 22 well-established stomatal regulators. The majority of the novel mutants were impaired in 398

stomatal closure to several stimuli, however mutants deficient in responses to a single stimulus e.g. 399

apoplastic ROS generated by O3 exposure, or CO2 were also identified (Figure 2). We expect that 400

future characterization of the newly identified mutants will provide insight into global, and stimuli-401

specific regulation of stomatal closure. As expected, apart from identifying novel mutants, our 402

screening strategy yielded 130 new mutants of genes encoding known stomatal regulators 403

(Supplementary Data Set 1). Mutations in AHA1, GHR1, MPK12 and MAX2 (Figure 2) were the most 404

prevalent, which might emphasize the significance of these genes in stomatal closure or the importance 405

of intact protein sequence for the whole-protein function. New alleles of the usual suspects described 406

here can act as a useful resource for detailed studies of their molecular function. Recently we used 10 407

novel alleles of GHR1 to dissect the role of specific mutations on its function and stability (Sierla et al., 408

2018). Currently, we are investigating the newly identified mutants of AHA1 and HT1 to get a deeper 409

insight into their mode of action. An unexpected finding was the identification of multiple novel alleles 410

of MAX2 (Supplementary Data Set 1). MAX2 is mostly recognized for its role in strigolactone 411

signaling (Waters et al., 2017) and has been previously implicated in guard cell functions (Ha et al., 412

2014; Piisilä et al., 2015) that are likely independent from SLs signaling (Kalliola et al., 2020). 413

However, our data suggests that the role of MAX2 in stomatal closure might be more central than 414

previously realized. In conclusion, our work provides genetic resources that enable further studies of 415

established stomatal regulators. Furthermore, mapping of newly identified mutants will contribute to 416

saturation of the genetic landscape of stomatal regulation. 417

Synthesis and import of GDP-L-fucose into the Golgi is necessary for stomatal closure 418


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At the organellar/cellular level, the flux of water through the guard cell tonoplast and plasma 419

membranes results in changes in volume and surface area of guard cells (Franks et al., 2001; Shope et 420

al., 2003; Meckel et al., 2007) and their central vacuoles (Diekmann et al., 1993; Gao et al., 2005). 421

Stomatal opening involves an increase in guard cell turgor (Franks et al., 2001), volume, and surface 422

area that in turn lead to an increase in guard cell length (Meckel et al., 2007). Contrary to stomatal 423

opening, stomatal closure involves guard cell shrinking. However, it is not entirely understood how the 424

repetitive, and fast changes in guard cell wall surface area are executed during stomatal movements. 425

According to current understanding, this process relies on the flexibility of the cell wall matrix which is 426

determined during guard cell morphogenesis (Rui et al., 2018), and is presumably promoted by the 427

turgor pressure of the subsidiary cells. Data available thus far point towards the key role of the pectin 428

network in controlling the cell wall flexibility that enables stomatal movements, and results presented 429

in this study support this hypothesis. The first mutant characterized from the screen described here 430

corresponds to MUR1 - an enzyme catalyzing the first step in de novo GDP-L-Fuc synthesis pathway 431

(Bonin et al., 1997). Plants lacking GDP-L-Fuc exhibited high loss of water from detached leaves and 432

impaired responses to apoplastic ROS, ABA, darkness and high CO2 concentration (Figure 4, 433

Supplementary Figures 1 and 6). Elevated water loss was also observed in plants impaired in import of 434

GDP-L-Fuc into Golgi apparatus (Figure 5B-C). Therefore, we conclude that not just the synthesis, but 435

also the import of GDP-L-Fuc into the Golgi lumen is necessary for stomatal closure. Recently, mur1 436

mutants were also shown to lack stomatal responses to treatment with Pseudomonas syringae (Pst 437

DC3118) and salicylic acid (Zhang et al., 2019). The stomatal defects observed in mur1 appear 438

independent from canonical guard cell signaling pathways (Figure 6A) and instead stem from defects in 439

cellular metabolism. 440

The Golgi apparatus (GA) serves as a hub for synthesis of pectic polysaccharides which are later 441

exported to the apoplastic space (Caffall and Mohnen, 2009). Accordingly, the fucosylation of cell wall 442

polysaccharides, AGPs and N-linked glycans is thought to occur in the GA (Figure 1; Chou et al., 443

2015; Strasser, 2016) and the majority of fucose is incorporated into the cell wall via the GA-derived 444

vesicles (Anderson et al., 2012). We found that mutants of fucosyltransferases responsible for synthesis 445

of AGPs, N-linked glycans and xyloglucan did not exhibit elevated water loss (Figure 7 A,C), therefore 446

we excluded the possibility that phenotypes observed in mur1 might be linked to impaired fucosylation 447

of these cell wall components. The partial reversion of the mur1 water loss phenotype by borate (Figure 448

7B), and high water loss observed in bor1-3 mutant (Supplementary Figure 9), suggests that the 449


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phenotype is linked either directly, or indirectly, to the structure and dimerization of RG-II. Earlier it 450

was found that mur1 mutants, even when grown under high borate conditions, had 74-78 % of RG-II in 451

dimer form while nearly all RG-II (95%) was dimerized in Col-0 (O’Neill et al., 2001). Therefore, 452

partial restoration of mur1 stomatal function might be related to incomplete RG-II dimerization even 453

after borate supplementation. 454

A complementary explanation for the observed phenotype might be related to shortening of the RG-II 455

chain A observed in mur1 plants (Pabst et al., 2013). As RG-II and HG share the same backbone and 456

are covalently cross-linked (Caffall and Mohnen, 2009; Harholt et al., 2010) the shortening of the RG-457

II chain A might have similar consequences as digestion of RG-I arabinan side chains (as observed 458

previously by Jones et al., (2003)). According to this scenario, the lack of GDP-L-Fuc would promote 459

crosslinking of HG and render the guard cell walls stiffer and, thus, less responsive to changes in turgor 460

pressure. This hypothesis is supported by our AFM data as we found that mur1 mutants have 461

significantly stiffer epidermal cell walls than wild type plants (Figure 6B). It is noteworthy, that 462

mutants with highly decreased HG content, quasimodo1 (qua1; Bouton et al., 2002) and qua2 (Mouille 463

et al., 2007) exhibit morphological phenotypes related to loss of cell adhesion which are not a direct 464

consequence of decreased HG content (Verger et al., 2016). In addition to impaired development, both 465

mutants exhibit high loss of water from detached leaves (Bouton et al., 2002; Krupková et al., 2007) 466

however, it is yet unknown whether this phenotype is related to impaired stomatal closure. 467

One argument against the hypothesis of increased HG crosslinking comes from the observation of the 468

stomatal opening process in mur1 mutants. In contrast to stimuli inducing stomatal closure, mur1 469

mutants responded properly to stomatal opening cues such as increase in light intensity and decrease in 470

CO2 concentration (Supplementary Figure 7), while Jones et al., (2003) observed impaired stomatal 471

opening after the arabinanase treatment. However, a similar phenotype to that reported here, i.e. 472

impaired closure and normal opening, was observed in plants lacking POLYGALACTURONASE 473

INVOLVED IN EXPANSION3 (PGX3; Rui et al., 2017). Polygalacturonases constitute a large group 474

of pectin-hydrolyzing enzymes (Yang et al., 2018). PGX3 is expressed in expanding tissues and guard 475

cells where it controls the abundance and molecular mass of HG. PGX3 deficiency led to increased 476

crosslinking of HG and impaired stomatal responses to ABA and darkness while the light- or 477

fusicoccin-induced stomatal opening was not affected (Rui et al., 2017). On the basis of this phenotype 478


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Rui et al., (2017) proposed a model in which the loosening of pectin structure is necessary for stomatal 479

closure, which is supported by the results of our study. 480

The normal stomatal opening observed in mur1 and pgx3 mutants might be explained by the relatively 481

low severity of phenotypes observed in these two mutants. Jones et al., (2003) reported that treatment 482

with arabinanase completely inhibited stomatal closure, while slower and “stepwise” stomatal closure 483

was observed in mur1 and pgx3 mutants, respectively. The digestion of arabinan via exogenous enzyme 484

treatment in epidermal strips likely affects the structure of the guard cell pectin network to a much 485

greater extent than that observed in the intact leaves of mur1 and pgx3 mutants, leading to a stomatal 486

opening phenotype. 487

Because of the observed changes in mechanical properties of guard cell walls, and the independence of 488

mur1 stomatal phenotypes from canonical guard cell signaling pathways, we hypothesize that the 489

impaired function of mur1 stomata is linked to a relatively stiffer epidermis. Since stomatal movements 490

occur in the context of a mechanical environment determined by the subsidiary cells it is possible that 491

the increased stiffness of the subsidiary cells also contributes to the mur1 phenotypes. 492

Alternative hypotheses and future challenges 493

Since mur1 mutants exhibit a radical reduction in availability of GDP-L-Fuc, additional factors may 494

contribute to impaired function of mur1 stomata. For example, a fraction of stomatal pores of mur1 495

plants were sealed with outer cuticular ledges (Supplementary Figure 5). A similar but more severe 496

phenotype, which was also associated with impaired stomatal closure in response to ABA, has been 497

observed in the focl1 mutant (Hunt et al., 2017). The precise molecular function of FOCL1 is not 498

known, however, it has been proposed to control the mechanical properties of guard cell walls and the 499

formation of the cuticle-cell wall bond (Hunt et al., 2017). Thus, it is tempting to speculate that, while 500

being independent from guard cell signaling, impaired mur1 stomatal responses might be linked to 501

those observed in focl1. 502

As the majority of cell wall L-fucose is localized in RG-I (Anderson et al., 2012), the lack of 503

fucosylation of this cell wall component might also contribute to phenotypes observed in mur1. To our 504

knowledge, the identity of fucosyltransferases responsible for incorporation of L-Fuc into RG-I and 505

RG-II remains unknown. Despite screening of multiple fucosyltransferase mutants we were not able to 506

reproduce the mur1 phenotype (Figure 7C), which probably reflects functional redundancy within the 507


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FUT family. In support of this hypothesis, the coding sequences of multiple FUTs are highly similar 508

and physically localized in groups (Sarria et al., 2001) which suggests that closely related FUTs arose 509

during gene duplication and, thus, might function redundantly. Therefore, due to the current lack of 510

genetic tools, we were unable to precisely pinpoint the cell wall components responsible for mur1 511

stomatal phenotypes. Analysis of high-order mutant lines, or the application of a CRISPR-512

Cas/amiRNA strategy, might be suitable for identification of fucosyltransferases that fucosylate RG-I 513

and RG-II. 514

Depletion of GDP-L-Fuc likely results also in defects in protein o-fucosylation. According to 515

phylogenetic analysis of plant and metazoan o-fucosyltransferases, Arabidopsis POFUTs constitute a 516

multigene family with nearly 40 members (Smith et al., 2018). Among them SPINDLY was shown to 517

fucosylate and activate DELLA proteins (Zentella et al., 2017) and according to (Zhang et al., 2019), 518

part of the pathogen susceptibility observed in mur1 mutants can be explained by lack of 519

SPINDLY-mediated protein o-fucosylation. However, we were not able to detect stomata-related 520

phenotypes in spy mutant lines. Another two putative o-fucosyltransferases FRIABLE1 (FRB1) and 521

ESMERALDA (ESMD) were shown to influence cell adhesion (Neumetzler et al., 2012; Verger et al., 522

2016). Similarly to qua1 and qua2, plants lacking FRB1 exhibit loss of cell adhesion (Neumetzler et 523

al., 2012). Strikingly, the introduction of the esmd mutation into qua1, qua2 and frb1 backgrounds 524

restored their cell adhesion defects with no apparent changes in cell wall composition, implying the 525

existence of a signaling pathway controlling cell adhesion (Verger et al., 2016). More research efforts 526

are needed to investigate whether the above-discussed mutants are impaired in stomatal function. 527

In summary, our study reveals that the synthesis of GDP-L-fucose is necessary for stomatal closure and 528

highlights the key role of fucose metabolism for stomatal closure. The interpretation of our data in the 529

context of earlier observations/hypotheses (Jones et al., 2003; Amsbury et al., 2016; Rui et al., 2017) 530

leads us to conclude that the impaired stomatal closure observed in mur1 is linked to increased stiffness 531

of guard cell walls which likely stems from enhanced crosslinking of HG caused by impaired structure 532

of RG-II. While the information obtained from mutant lines reported here provides valuable indications 533

as to which cell wall components are necessary for the execution of stomatal movements, precisely 534

how wall structure is modified/reorganized upon perception of stomatal opening/closing stimuli to 535

accommodate volume/pressure changes awaits future investigation. 536


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MATERIALS & METHODS 537

Plant material and growth conditions 538

All Arabidopsis thaliana mutants used in this study were in the Col-0 genetic background. The 539

following lines were obtained from Nottingham Arabidopsis Stock Center: mur1-1, mur1-2 (Reiter et 540

al., 1993; Bonin et al., 1997); mur1-9 (SALK_057153); ger1-2 (WiscDsLox_425G11), ger1-3 541

(GK_296H02), ger2-1 (GK_113G05), ger2-2 (SALK_091781); mur2-1 (Reiter et al., 1997; Vanzin et 542

al., 2002); ghr1-3 (GK_760C07; Sierla et al., 2018); slac1-4 (SALK_137265; Vahisalu et al., 2008); 543

ost1-3 (SALK_008068; Yoshida et al., 2002); mur3-7 (SALK_127057; Tamura et al., 2005); AtkdsA1-544

S (SALK_024867), AtkdsA2-S (SALK_066700; Delmas et al., 2008); rgxt1-1 (SALK_073748), rgxt2-1 545

(SALK_023883; Egelund et al., 2006), bor1-3 (SALK_037312; Kasai et al., 2011),fut2-1 546

(GK_320C07), fut3-1 (SALK_045666), fut8-1 (SALK_010965), fut8-2 (WiscDsLox_449F06), fut10-1 547

(WiscDsLox_432A01), fut10-2 (SALK_020408), fuctc-1 (SALK_067444; Rips et al., 2017). 548

pGC1:YC3.6 line (Yang et al., 2008) was donated by Julian Schroeder. FKGP mutants fkgp-1 549

(SALK_012400), fkgp-2 (SALK_053913; Kotake et al., 2008) were donated by Toshihisa Kotake; 550

mur3-1, mur3-2 (Reiter et al., 1997), mur3-3 (SALK_141953; Tamura et al., 2005), xxt1 551

(SAIL_785_E02) xxt2 (SALK_101308) double mutant (Cavalier et al., 2008), and xxt1 xxt2 mur3-3 552

triple mutant (Kong et al., 2015) were donated by Malcolm A. O’Neill; ost2-2D mutant (Merlot et al., 553

2007) was donated by Jeffrey Leung; aba2-11 mutant (González-Guzmán et al., 2002) was donated by 554

Pedro L. Rodríguez; fucTA (SALK_087481) fucTB (SALK_063355) double mutant (Strasser et al., 555

2004) seeds were obtained from Richard Strasser; fut4 (SAIL_284_B05) fut6 (SALK_078357) double 556

mutant seeds (Tryfona et al., 2014) were obtained from Paul Dupree; snrk2.236 triple mutant seeds 557

(Fujii and Zhu, 2009; Cui et al., 2016) were donated by Hiroaki Fujii and Kirk Overmyer; ht1-2 mutant 558

(Hashimoto et al., 2006) was donated by Koh Iba. All mutant lines were genotyped with gene specific 559

primers (Supplementary Data Set 2). GGLT1 RNAi lines (Sechet et al., 2018), together with the 560

corresponding empty-vector control, were donated by Julien Sechet and Jenny C. Mortimer; T1 seeds 561

of hpGFT1 and empty-vector control as well as E.coli and Agrobacterium tumefaciens strains carrying 562

the hpGFT1 construct in pART27 (Rautengarten et al., 2016) were obtained from Joshua L. 563

Heazlewood. To generate hpGFT1 lines, Col-0 plants were transformed via the floral-dip method with 564

Agrobacterium tumefaciens strain AGL1 carrying the hpGFT1 construct in pART27 vector 565

(Rautengarten et al., 2016). T1 transformants were selected on half-strength Murashige & Skoog 566


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medium (Duchefa Biochemie) supplemented with 50 µg ml-1 kanamycin, in a controlled growth 567

chambers (MLR-350, Sanyo) under 12 h light (130-160 µmol m-2 s-1)/12 h dark cycle, 22°C/18°C 568

(day/night). After two weeks plants were transplanted to soil and grown for additional three weeks as 569

described below. The same selection procedure has been applied to select for empty-vector control 570

lines. Col-0, mur1-1, mur1-2, slac1-4 and ghr1-3 lines were grown in parallel, except the selection 571

antibiotic was not included in the growth medium. The double mutants of mur1-1 and mur1-2 with 572

ost1-3, aba2-11, ghr1-3, slac1-4 and ost2-2D were generated by crossing mur1-1 and mur1-2 (pollen 573

acceptors) with the respective pollen donor. Double mutants were identified in F2 generation by 574

genotyping with gene-specific primers (Supplementary Data Set 2). An attempt to generate ger1 ger2 575

double mutants was performed by crossing ger1-2 and ger1-3 (pollen acceptors) with ger2-1 and ger2-576

2 (pollen donors). Later, F2 populations were genotyped with gene-specific primers (Supplementary 577

Data Set 2). F2 plants homozygous for ger1 but heterozygous in ger2 locus were allowed to self-578

pollinate and the F3 plants were genotyped as before to determine the segregation of ger2 alleles. 579

Unless specified otherwise, seeds were suspended in 0.1% agarose solution, vernalized in the dark for 580

two days at 4°C and sown on a 1:1 mixture of peat and vermiculite. Plants were grown in controlled 581

growth rooms under 12 h light (200 µmol m-2 s-1)/12 h dark cycle, 22°C/18°C (day/night), 60%/70% 582

relative humidity. 583

Mutagenesis and screening procedure 584

Mutagenesis of Arabidopsis pGC1:YC3.6 (Yang et al., 2008) seeds was performed as described before 585

(Kim et al., 2006). In the primary screen, M2 plants were grown for two weeks and exposed to 275-350 586

nl l-1 O3 for 6 h. Presence of O3-induced lesions was assessed visually, and sensitive plants were 587

rescued. One to two weeks later, rosettes were imaged with an Optris PI450 thermal imager (Optris 588

GmbH) and subjected to water loss assay as described below. M3 seeds were collected from the M2 589

plants that exhibited the most pronounced phenotypes i.e. severe O3 sensitivity, high loss of water 590

and/or low leaf temperature. Later, to confirm the inheritance of the observed phenotypes, the M3 591

plants were exposed to 350 nl l-1 O3 for 6 h, and subjected to water loss assay. Approximately half of 592

both, the primary and the secondary screen, was performed at the ExpoSCREEN facility (HMGU, 593

Munich) and the remaining half at the plant growth facilities of the University of Helsinki. Lines for 594

which at least one phenotype was reproduced, were selected for whole-rosette gas exchange-based 595

phenotyping of stomatal function. Gas exchange measurements were performed on 2 - 4 plants per 596


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mutant line and in total 551 lines were analyzed for stomatal responses to O3 pulse (547 lines), elevated 597

CO2 (550 lines), darkness (106 lines), VPD (498 lines) and ABA spray (482 lines). Stomatal 598

conductance of 505 lines was tested. 599

Lines with defective stomatal responses to at least one stimulus were subjected to sequencing of “usual 600

suspects” either by NGS-based sequencing of PCR amplicons obtained with the use of gene-specific 601

primers (Supplementary Data Set 2) and Phusion DNA polymerase (ThermoFisher Scientific; ROUND 602

1 to 6), or whole-genome sequencing (ROUND 7 and 8). In the PCR-based approach, for each line, 603

amplicons were pooled and subjected to NGS with the use of GS FLX+ system (Roche) or MiSeq 604

system (Illumina) at DNA Sequencing and Genomics Laboratory, Institute of Biotechnology, 605

University of Helsinki. Sequencing with the use of GS FLX+ system was performed exactly as 606

described before (Sierla et al., 2018). For MiSeq-based sequencing, pooled amplicons were sheared 607

(Bioruptor NGS, Diagenode) into approximately 500 bp fragments, end-repaired, A-tailed and ligated 608

to truncated TruSeq adapters (Illumina). After purification a PCR reaction was performed to introduce 609

full-length P5 adapter and indexed P7 adapter sequences. After PCR, all products were pooled, purified 610

with AMPure XP (Agencourt, Beckman Coulter), size selected to 600-800 bp (as described above), and 611

analyzed on a Fragment Analyzer (Advanced Analytical Technologies Inc., Ankeny, IA, USA). Paired-612

end sequencing were performed on a MiSeq using a 600 cycle kit v3 (Illumina). The obtained 613

sequences were trimmed using Cutadapt (Martin, 2011) and assembled with SPAdes (Bankevich et al., 614

2012). In the whole genome sequencing approach (ROUND 7 and 8, performed at Institute for 615

Molecular Medicine Finland, FIMM) leaf samples of approximately 10 plants per line were used for 616

nuclear DNA isolation. Nextera Flex library preparation was performed using 50-100 ng of dsDNA 617

according to Nextera DNA Flex Library Prep Reference Guide (Illumina, San Diego, CA, USA) with 618

the following modifications: all reactions were performed in half of the normal volume and library 619

normalization was done according to the concentration measured on LabChip GX Touch HT 620

(PerkinElmer, USA). Later, 500-850 bp fragments were size selected from the pool using BluePippin 621

(Sage Science, USA). In ROUND 7, sequencing was performed with Illumina NovaSeq 6000 system 622

using S4 flow cell with lane divider (Illumina, San Diego, CA, USA). Read length for the paired-end 623

run was 2x151 bp. In ROUND 8, sequencing was performed with the Illumina NovaSeq 6000 system 624

using S1 flow cell with lane divider (Illumina, San Diego, CA, USA). Read length for the paired-end 625

run was 2 x 101 bp. Following the data acquisition, reads were subjected to demultiplexing with 626

Illumina bcl2fastq2.20 software. 627


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30

Quality control of the reads was carried out using FastQC software 628

(https://www.bioinformatics.babraham.ac.uk/projects/fastqc/), and the adapter removal and read 629

trimming were performed with Trimmomatic v0.36 (Bolger et al., 2014). After removal of adaptor 630

sequences, all bases were removed from the beginning and the end of the reads if their Phred quality 631

score was below 20. Additionally, reads with mean Phred score <15 in a sliding window of 3 bases 632

were clipped, and the reads with read length < 35 bp after trimming were removed. The trimmed reads 633

were aligned to the TAIR10 version of the Arabidopsis genome using Bowtie 2 v2.2.9 (Langmead and 634

Salzberg, 2012). Next, the sequence alignment data was converted to binary alignment file and sorted 635

using Picard v2.5.0 (http://broadinstitute.github.io/picard/). The same toolbox was used for removing 636

the read PCR duplicates and for adding read group information. Next, Genome Analysis Toolkit 637

(GATK, https://gatk.broadinstitute.org) by Broad Institute was used to obtain genomic VCF files using 638

HaplotypeCaller (Poplin et al., 2017), for combining the gVCF files, and finally for the joint calling of 639

variants using GenotypeGVCFs. The called SNPs were annotated using snpEff v4.3t (Cingolani et al., 640

2012) and Arabidopsis gene annotation downloaded from TAIR on May 1st 2018. The SNP calls in the 641

list of usual suspects were then collected for manual inspection. 642

Water loss measurement 643

Two middle age leaves of 3.5-week-old soil-grown plants were cut and dried abaxial side up at room 644

temperature for 2 h, unless specified otherwise. Weight of leaves was determined before and after 645

drying, and water loss was calculated as percentage of initial fresh weight loss. 646

Quantification of GDP-L-fucose 647

Approximately 160 mg of frozen plant material was ground to powder and extracted first with 1 ml of 648

80% (v/v) methanol and then with 0.5 ml of 100% methanol. During each extraction, the homogenate 649

was vortexed at 4 °C for 2 h, then centrifuged at 21,500 g for 5 min at 4 °C. The supernatants were 650

combined and evaporated to dryness in vacuum (MiVac Duo concentrator, GeneVac Ltd, Ipswich, UK) 651

and reconstituted in 50 µl of 50% (v/v) methanol. The GDP-L-fucose was quantified with UPLC-652

6500+ QTRAP/MS system (Sciex, CA) in negative (ESI-) multiple reaction monitoring (MRM) mode. 653

Two MRM transitions for GDP-L-fucose (C16H25O15N5P2, molecular weight 589.34 g mol-1) were 654

used: 588.0 → 442.0 for quantitative, and 588.0 → 424.0 for qualitative purposes. The 655

chromatographic separation was performed in Waters BEH Amide column (100 mm × 2.1 mm, ø1.7 656

µm) at a flow rate of 0.4 ml min-1. Column compartment temperature of UPLC system (Sciex, CA) was 657


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set to 35 °C while the samples were kept at 10 °C. Initial chromatographic conditions were 15% buffer 658

A (10 mM ammonium formate, pH 9.0) and 85% buffer B (acetonitrile). The initial conditions were 659

held for 1 min and followed by a linear gradient to 60 % (A) in 6 min, then returned to initial 660

conditions in 1 min and left to stabilize for 2 min. Analysis has been performed with the following 661

parameters: declustering potential of -85, entrance potential of -10, collision cell exit potential of -36 662

and 40, respectively. The ion spray voltage was set at -4,500V, source temperature (TEM) at 425 °C, 663

collision gas (CAD) was set to medium, curtain gas to 20, and source gas 1 (GS1) and 2 (GS2) were 664

both set to 10. All data were acquired with Analyst 1.5.1 software (Sciex, CA) and GDP-L-fucose was 665

quantified with MultiQuant 2.1 software (Sciex, CA) by utilizing standard calibration curve of GDP-L-666

fucose standard (Sigma Aldrich). 667

Gas exchange analysis 668

The gas-exchange experiments were performed as described before (Sierla et al., 2018). Briefly, plants 669

were grown in 4:3 peat : vermiculite mix under 12 h day (23°C) / 12 h night (18°C) regime at 70% 670

relative humidity and 100-150 μmol m-2 s-1 light intensity. The stomatal function of 3 - 4 week-old 671

plants was analyzed with the use of a custom-built gas-exchange device (Kollist et al., 2007). First, 672

plants were inserted into the device and pre-incubated for about 1 h to stabilize the stomatal 673

conductance. The experimental conditions in the chambers were as follows: ambient CO2 (~400 µL L-674 1), 150 µmol m-2 s-1 light, ~70% relative air humidity, 24oC. Later, stimuli triggering stomatal 675

movements were applied and changes in stomatal conductance were monitored in time. To induce 676

stomatal closure O3 pulse (450 nl l-1 during 3 min), elevated CO2 (from 400 µl l-1 to 800 µl l-1), 677

darkness, increase in VPD (reduction in relative air humidity (from 70% to ~30-35%)) or 5 µM ABA 678

spray were applied. Similarly, reduction of CO2 concentration (from 400 µl l-1 to 100 µl l-1) or increase 679

of light intensity (from 150 μmol m-2 s-1 to 500 μmol m-2 s-1) were used to trigger stomatal opening. In 680

order to study diurnal light/darkness responses, stomatal conductance was recorded for approx. 40 h 681

under conditions that mimicked the growth day/night regime. 682


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Mapping by next-generation sequencing 683

Leaf tissue samples from the BC1F2 plants exhibiting the mutant phenotype were harvested in bulk and 684

used for preparation of nuclear DNA. The nuclear DNA-enriched sample was sheared to 400 – 600 bp 685

fragments (Bioruptor NGS), end-repaired, A-tailed and ligated to truncated TruSeq adapters (Illumina). 686

Full-length P5 and indexed P7 adapter sequences were introduced by PCR. The library was purified 687

and size selected to 500-700 bp fragments as described above. Paired-end sequencing (150 – 150 bp) of 688

the obtained library was done on a NextSeq 500 sequencer (Illumina) to an approximately 50-fold 689

genome coverage at DNA Sequencing and Genomics Laboratory, Institute of Biotechnology, 690

University of Helsinki. Further, data were analyzed with SHORE v0.9.0 pipeline (Ossowski et al., 691

2008) implementing GenomeMapper v0.4.4s read alignment tool (Schneeberger et al., 2009) according 692

to the procedure described earlier (Sun and Schneeberger, 2015). In parallel, the same procedure was 693

performed for line YC3.6. Next, SHOREmap v3.2 (Sun and Schneeberger, 2015) was used to identify, 694

prioritize and annotate mutations enriched in plants displaying mutant phenotype. This final step 695

involved subtraction of all innate YC3.6 polymorphisms. 696

Nuclear DNA enrichment 697

For nuclear DNA enrichment, plant material was ground in liquid nitrogen and 1 g of tissue powder 698

was homogenized with 15 ml of HBM buffer (25 mM Tris-HCl pH 7.5, 440 mM sucrose, 10 mM 699

MgCl2, 0.1% Triton X-100, 10mM β-mercaptoethanol, 2 mM spermine). The homogenate was filtered 700

through Miracloth and centrifuged for 10 min at 1,075g, 4ºC. Pellet was resuspended in 1 ml of NIB 701

buffer [20 mM Tris-HCl pH 7.5, 250 mM sucrose, 5 mM MgCl2, 5 mM KCl, 0.1% (v/v) Triton X-100, 702

10 mM β-mercaptoethanol] and applied to a 15/50% (v/v) Percoll gradient in NIB buffer. Samples were 703

centrifuged for 10 min at 3000g, 4ºC and pelleted nuclei were resuspended in 0.5 ml NIB buffer and 704

centrifuged again. Pellets were mixed with 750 µL of DNA extraction buffer [2 % (w/v) 705

cetyltrimethylammonium bromide, 100 mM Tris-HCl pH 8.0, 20 mM EDTA pH 8.0, 1.4 M NaCl, 1 % 706

(w/v) polyvinylpyrrolidone 40] and incubated for 30 min at 60ºC. Next, samples were extracted with 707

chloroform : isoamyl alcohol (24:1, v/v) and centrifuged for 10 min at 7,000g, 4ºC. The water-soluble 708

phase was collected, treated with RNase A (Merck), and DNA was precipitated with isopropanol 709

at -20ºC. Samples were centrifuged for 6 min at 13,000g, 4 ºC, the pellet was washed twice in 70% 710

(v/v) ethanol, air-dried, and resuspended in water. DNA concentration was measured with Qubit 711

fluorometer (Thermo Fisher Scientific) according to manufacturer’s instructions. 712


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33

Expression Analysis by qPCR 713

For gene expression analysis of T-DNA insertion mutants, plants were grown vertically on half-714

strength Murashige & Skoog medium (Duchefa Biochemie) in controlled growth chambers (model 715

MLR-350, Sanyo) under 12 h light (130-160 µmol m-2 s-1)/12 h dark cycle, 22°C/18°C (day/night). For 716

every biological replicate (3 in total), approximately 10 whole two-week-old plants were pooled, frozen 717

and ground in liquid nitrogen. Total RNA was extracted using a Genejet plant RNA isolation kit 718

(Thermo Fisher Scientific). For analysis of GFT1 transcript level, whole rosettes of T1 hpGFT1 plants 719

(minus two middle-aged leaves that were used for water loss assay) were frozen in liquid nitrogen and 720

ground with a mortar and pestle. Total RNA was extracted as described above. Two micrograms of 721

RNA were treated with DNaseI and reverse-transcribed using oligo-dT(20) priming with Maxima 722

Reverse Transcriptase (RT) and Ribolock RNase inhibitor (Thermo Fisher Scientific) in a 31.5 µl 723

volume. The reactions were diluted to the final volume of 100 µl, 1µl of which was used as template 724

for PCR with 5x HOT FIREPol EvaGreen qPCR Mix Plus (no ROX; Solis Biodyne) with gene-specific 725

primers (Supplemental Table 5). The PCR was performed on the CFX384 Real-Time System (Bio-726

Rad) with the following cycle conditions: 95°C 10 min, 60 cycles with 95°C 30 s, 60°C 10 s, 72°C 30 727

s, and ending with melting curve analysis. Col-0 cDNA dilution series was used to determine primer 728

amplification efficiencies. Three reference genes were used for data normalization: YELLOW-LEAF-729

SPECIFIC GENE8 (YLS8, AT5G08290), TAP42 INTERACTING PROTEIN OF 41 KDA (TIP41, 730

AT4G34270), and MONENSIN SENSITIVITY1 (MON1, AT2G28390; Czechowski et al., 2005). Data 731

were analyzed with qBase 3.0 (Biogazelle) and the transcript levels were related to those observed in 732

respective control lines. 733

Expression analysis by RT-PCR 734

For gene expression analysis of fut mutants plants were grown, and the cDNA was produced, as 735

described above. PCR was performed with the use of FirePol polymerase (Solis Biodyne) and 1 µl of 736

cDNA was used as a template. PP2AA3 (At1g13320) transcript was used for reference. All primers are 737

listed in Supplementary Data Set 2. The cycling conditions were as follows 95°C 3 min, 40 cycles with 738

95°C 30 s, 56°C 10 s, 72°C 90 s ending with 5 min final elongation at 72°C. PCR products were 739

analyzed by agarose gel electrophoresis using ethidium bromide staining for imaging. 740


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Stomatal morphology and cuticle permeability 741

For scanning electron microscopy, cotyledons of 3-week-old soil-grown plants (1 cotyledon per plant, 742

4-6 plants per line per biological replicate, 3 biological replicates) were harvested and fixed for 16 h in 743

a fixing buffer (2% paraformaldehyde, 2.5% glutaraldehyde, 0.1 M sodium cacodylate pH 7.4). 744

Subsequently, samples were incubated for 1 h at room temperature in 0.1 M sodium cacodylate buffer 745

pH 7.3 containing 1% (w/v) OsO4 and washed twice in distilled water. Later samples were dehydrated 746

by 3 x 10 min washing in 50% , 70%, 96% and 100% (v/v) ethanol, subjected to critical point drying 747

(Leica EM CPD300, Leica Microsystems GmbH), mounted on the aluminum stubs and coated with 5 748

nm layer of platinum (Quorum Q150TS, Quorum Technologies Ltd). Serial images (1,800x 749

magnification, 10 % overlap) were taken with Quanta FEG 250 (Thermo Fisher Scientific) scanning 750

electron microscope at the Electron Microscopy Unit, Institute of Biotechnology, University of 751

Helsinki. Images were stitched in Fiji (Schindelin et al., 2012) implementing MIST plugin (Chalfoun et 752

al., 2017). For each cotyledon from 1 to 2.5 mm2 area was analyzed, containing 100 - 350 stomatal 753

complexes. Morphology of every stomata within the region of interest was assessed visually and 754

assigned into one of the four categories: normal appearance, not determined, sealed (Supplementary 755

Figure 5F), and obstructed (Supplementary Figure 5E). The size of stomatal pores was determined by 756

measuring the length of the stomatal pore defined by the boarders of the outer cuticular ledge. Cuticle 757

permeability assay was performed on middle-age leaves of 3-week-old soil-grown plants as described 758

earlier (Cui et al., 2016) except 0.01% Tween 20 has been added to the staining solution. Mutants 759

aba2-11 (González-Guzmán et al., 2002) and snrk2.236 (Fujii and Zhu, 2009) were used as controls 760

with high cuticle permeability (Cui et al., 2016). 761

Projected rosette area 762

Rosettes were photographed with Nikon D5100 camera equipped with AF-S Micro Nikkor 40 mm 763

1:2.8G objective (Nikon). The projected rosette area was determined by image analysis with ImageJ 764

software (Schneider et al., 2012) equipped with Measure Rosette Area Tool 765

(http://dev.mri.cnrs.fr/projects/imagej-macros/wiki/Measure_Rosette_Area_Tool). 766

767


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Atomic force microscopy 768

For AFM experiments, seeds were stratified for 7 days at 4°C, then grown in a 3:1 compost:perlite mix. 769

Growth conditions were as follows: light intensity 170 μmol m-2 s-1, 12h day (21°C) / 12 h (17°C) 770

night, 60% humidity. Leaves from approx. 21-day-old plants were analyzed as described before (Carter 771

et al., 2017). 772

773


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SUPPLEMENTARY DATA 774

Supplementary Figure 1. Characterization of T7-9 stomatal phenotypes. 775

Supplementary Figure 2. Q-PCR analysis of mur1-9, ger1 and ger2 mutants. 776

Supplementary Figure 3. Water loss-based screen of T-DNA insertion mutants of T7-9 candidate genes. 777

Supplementary Figure 4. Cuticle permeability of mur1 mutants. 778

Supplementary Figure 5. Scanning electron microscopy-based phenotyping of stomatal morphology in 779

mur1 mutants. 780

Supplementary Figure 6. Stomatal responses of mur1 mutants to stomata-closing stimuli. 781

Supplementary Figure 7. Response of mur1 mutants to stomata-opening stimuli. 782

Supplementary Figure 8. Analysis of hpGFT1 T1 plants. 783

Supplementary Figure 9. The influence of soil borate concentration on water loss of bor1-3 mutant. 784

Supplementary Figure 10. Water loss of mutants of enzymes/transporters involved in synthesis of 785

RG-II. 786

Supplementary Figure 11. Characterization of fut T-DNA insertion mutants. 787

Supplementary Table 1. Genes included in candidate gene sequencing. 788

Supplementary Table 2. High-frequency single nucleotide polymorphisms identified in T7-9 BC1F2 789

mapping population 790

Supplementary Table 3. Genotypes of F3 plants obtained from ger1-/- ger2+/- F2 plants. 791

Supplementary Data Set 1. Mutations identified during candidate gene sequencing. 792

Supplementary Data Set 2. List of primers 793


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ACKNOWLEDGEMENTS 794

We thank Julian Schroeder, Toshihisa Kotake, Malcolm A. O’Neill, Jeffrey Leung, Pedro L. 795

Rodríguez, Jenny C. Mortimer, Julien Sechet, Richard Strasser, Hiroaki Fujii, Kirk Overmyer, Koh Iba 796

and Paul Dupree for providing seeds of multiple mutants used in this study. We thank Joshua 797

Heazlewood and Berit Ebert for providing Agrobacterium tumefaciens strain carrying the hpGFT1 798

construct, T1 seeds of hpGFT1 and corresponding EVC. Anna Huusari, Konrad Łosiński, Jelena 799

Odintsova and Samuli Lundström are acknowledged for excellent technical help. Leena Grönholm and 800

Valtteri Lehtonen are acknowledged for maintaining the plant growth facilities. Personnel of the DNA 801

sequencing and genomics laboratory is acknowledged for the technical assistance with NGS. Mervi 802

Lindman is acknowledged for assistance with electron microscopy. We gratefully acknowledge Hans 803

Lang for an excellent technical support during the ozone fumigation experiments at the EUS. We thank 804

Andreas Albert and the technical staff of EUS for outstanding support during the experiment. Thanks to 805

Mikael Brosché and Kirk Overmyer for helpful comments and discussions throughout the duration of 806

the project. This work was supported by the University of Helsinki, Academy of Finland Centre of 807

Excellence program (2014-2019), Academy of Finland post-doctoral fellowships (Decision 294580, 808

C.W.; 266793, T.V.), University of Helsinki 3-year research grant (C.W., M.L.G., J.P.), EDUFI 809

fellowship (M.L.G.), The Ella and Georg Ehrnrooth Foundation (M.S.), Finnish Cultural Foundation 810

(M.S), Dora Plus Programme (Contract No. 36.9-6.1/1372, O.Z.), Estonian Research Council 811

(PRG433, H.K.; PUT311, PRG719, D.Y.; Mobilitas Pluss postdoctoral researcher grant (Contract no. 812

MOBJD291), A.K.P), European Regional Development Fund (Center of Excellence in Molecular Cell 813

Engineering CEMCE, H.K.). The ozone exposure experiments performed at the HMGU were 814

supported by Transnational Access program of the European Plant Phenotyping Network (EPPN, grant 815

no. 284443) funded by the FP7 Research Infrastructures Programme of the European Union. 816

AUTHOR CONTRIBUTIONS 817

J.K., conceived the project; C.W., T.V., D.Y., M.S., M.L.G., R.C., N.S., A.J.F., H.K., J.K., designed 818

experiments, C.W., T.V., D.Y., M.S., O.Z., M.L.G., J.P., R.C., A.K.P., M.N., M.C., T.P., N.S., A.L., 819

L.P., P.A., A.J.F., J.S., performed experiments and analyzed the data, J.D., D.E., J.B.W., provided 820

technological solutions for large-scale O3 exposures; C.W., T.V., D.Y., M.S., A.J.F., H.K. and J.K. 821

wrote the manuscript with comments from all co-authors. 822


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38

FIGURE LEGENDS 823

Figure 1. Synthesis and metabolism of GDP-L-fucose in Arabidopsis thaliana. 824

Figure 2. Ozone sensitivity-based forward genetic screen to identify novel regulators of stomatal 825

closure. 826

(A) The amount of mutant lines of known stomatal regulators identified during candidate gene 827

sequencing. 828

(B) Scheme of the screening procedure. 829

(C) Whole-rosette gas exchange analysis of novel mutants impaired in stomatal closure. Numbers 830

indicate the amount of mutants impaired in stomatal closure to indicated stimuli. 831

Figure 3. Mapping of T7-9 mutant. 832

(A) Leaf fresh weight loss of T7-9 and control lines (YC3.6, Col-0, slac1-4 and ghr1-3) recorded after 833

2h. Data bars represent means ± SD (n = 12 plants). 834

(B) Positions of mutations in mur1 mutants used in this study. 835

(C) Leaf fresh weight loss of T7-9, independent mur1 mutants (mur1-1, mur1-2, mur1-9) and control 836

lines (YC3.6, Col-0, slac1-4 and ghr1-3) recorded after 2h. Data bars represent means ± SD (n = 12 837

plants). 838

(A, C) Asterisks denote statistical differences (*** p < 0.001) to respective control lines (Col-0 or 839

YC3.6) according to one-way ANOVA followed by Sidak’s post-hoc test. 840

(D) GDP-L-fucose content in T7-9, mur1 mutants and respective control lines measured by UPLC-MS. 841

Data bars represent means ± SD (n = 4-5 plants); n.d., not detected. 842

(E) Leaf fresh weight loss of T7-9, mur1-2, F1 T7-9 x mur1-2, F1 YC3.6 x mur1-2 and control lines 843

recorded after 2h. Data bars represent means ± SD (n = 9-12 plants). Asterisks denote statistical 844

differences (* p < 0.05; ** p < 0.01; *** p < 0.001) to Col-0 according to one-way ANOVA followed 845

by Dunnett’s post-hoc test. 846

(A, C, E) Experiments were repeated three times with similar results. Results of the representative 847

experiments are shown. 848


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39

Figure 4. Characterization of mur1 stomatal phenotypes. 849

(A-H) Stomatal responses of mur1 mutants to stomata-closing stimuli. The changes in stomatal 850

conductance are shown in relative and absolute values calculated from the data presented in 851

Supplementary Figure 6. 852

(A, C, E, G) Time course of relative stomatal conductance (normalized to the last time point before the 853

treatment) of 3- to 4-week-old mur1-1, Col-0, and ghr1-3 plants in response to (A) O3 pulse, (C) 854

elevated CO2, (E) ABA spray and (G) darkness. The indicated treatments were applied at t = 0 and 855

whole-rosette stomatal conductance was recorded. Data points represent means ± SEM; n = 7-10 (A), 856

10-12 (C), 9-11 (E), 6-11 (G) plants analyzed in two (A, E, G) or three (C) independent experiments. 857

(B, D, F, H) Changes in stomatal conductance of Col-0, mur1-1, mur1-2 and ghr1-3 in response to (B) 858

O3 pulse, (D) elevated CO2, (F) ABA spray and (H) darkness. Values were calculated by subtracting 859

the initial stomatal conductance at t = 0 (D, F, H) or t = -1 (B) from the stomatal conductance at (B) t 860

= 7 min, (D, F, H) t = 40 min. Data bars represent means ± SEM; n = 7-10 (B), 10-12 (D), 9-11 (F), 6-861

11 (H) plants. Asterisks denote statistical differences to Col-0 (* p < 0.05; ** p < 0.01; *** p < 0.001) 862

according to one-way ANOVA followed by Dunnett’s post-hoc test. 863

(I) Diurnal changes in whole-rosette stomatal conductance of mur1-1, mur1-2 and Col-0 plants. Data 864

points represent means ± SEM; n = 8 plants analyzed in 3 experiments. Each experiment was started at 865

the same time of the day and measurements were recorded for 45 h. 866

867


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40

Figure 5. Stomatal function in mutants affected in GDP-L-fucose metabolism. 868

(A) Leaf fresh weight loss of mutants of genes encoding enzymes involved in synthesis of GDP-L-869

fucose, and control lines (Col-0, mur1-1, mur1-2, slac1-4 and ghr1-3), recorded after 2h. Data bars 870

represent means ± SD (n = 13-16 plants). Asterisks denote statistical differences (*** p < 0.001) to 871

Col-0 according to one-way ANOVA followed by Dunnett’s post-hoc test. Experiment was repeated 872

three times with similar results. Results of the representative experiment are shown. 873

(B) Leaf fresh weight loss of 64 independent hpGFT1 T1 plants, and control lines (Col-0, EVC – 874

empty-vector control, mur1-1, mur1-2, slac1-4 and ghr1-3), recorded after 2h. Data points represented 875

values obtained for separate plants. Data bars represent means ± SD (for control lines n = 11-12 plants). 876

(C) Correlation between residual GFT1 transcript level and fresh weight loss observed in 58 877

independent hpGFT1 T1 plants. Each dot represents values obtained for an independent hpGFT1 T1 878

plant. 879

Figure 6. Stomatal phenotypes observed in mur1 mutants are independent from canonical guard 880

cell signaling. 881

(A) Leaf fresh weight loss of double mutants obtained after crossing slac1-4, ghr1-3, aba2-11, ost1-3 882

and ost2-2D with mur1-1 and mur1-2 recorded after 1h. Data bars represent means ± SD (n = 13-16 883

plants). Asterisks denote statistical differences (*** p < 0.001) to respective single mutant lines 884

(slac1-4, ghr1-3, aba2-11, ost1-3 and ost2-2D) according to one-way ANOVA followed by Sidak’s 885

post-hoc test. Experiment was repeated three times with similar results. Results of the representative 886

experiment are shown. 887

(B) Average apparent Young’s modulus (Ea) values derived from AFM scans of subsidiary cells (SC) 888

and guard cells (GC) of control (Col-0, YC3.6) and mur1 plants. Bars represent means ± SD (n = 2-3 889

plants, 2 stomata per plant). Data analyzed with two-way repeated measures ANOVA with “genotype” 890

and “cell type” as factors, followed by Tukey’s post-hoc test. Asterisks denote statistical differences 891

(* p < 0.05, ** p < 0.01, *** p < 0.001) to respective control lines (Col-0, YC3.6) observed within a 892

cell type. 893

894


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41

Figure 7. Screen for fucose-containing cell wall components affecting stomatal function. 895

(A) Leaf fresh-weight loss of mutants affected in fucosylation and synthesis of xyloglucan, recorded 896

after 2h. Values obtained for Col-0, mur1-1, mur1-2, slac1-4 and ghr1-3 are provided for reference. 897

Data bars represent means ± SD (n = 15-16 plants). 898

(B) The effect of borate supplementation on the leaf fresh-weight loss of mur1 mutants and control 899

lines (Col-0, ghr1-3 and slac1-4). Bars represent means ± SD (n = 14-16 plants). Data analyzed with 900

two-way ANOVA with “genotype” and “borate concentration” as factors, followed by Sidak’s post-hoc 901

test. Asterisks denote statistical significances (** p < 0.01, *** p < 0.001) of treatment effect within 902

each genotype. Experiment was repeated five times with similar results. Results of the representative 903

experiment are shown. 904

(C) Leaf fresh-weight loss of mutants deficient in fucosyltransferases. Values obtained for Col-0, 905

mur1-1, mur1-2, slac1-4 and ghr1-3 are provided for reference. 906

(A, C) Experiments were repeated at least 3 times with similar results. Results of the representative 907

experiments are shown. Asterisks denote statistical differences (* p < 0.05, ** p < 0.01, *** p < 0.001) 908

to Col-0 according to one-way ANOVA followed by Dunnett’s post-hoc test. 909

910


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Supplementary Figure 1. Characterization of T7-9 stomatal phenotypes. 911

Time course of whole-plant stomatal conductance of T7-9, YC3.6 and ghr1-3 (A-D) or ost1-3 (E) in 912

response to (A) O3 pulse, (B) elevated CO2, (C) ABA spray, (D) darkness and (E) drop in relative air 913

humidity (VPD increase from 1.01 ± 0.01 kPa to 2.04 ± 0.03 kPa (mean ± SE)). The indicated 914

treatments were applied at t = 0 and whole-rosette stomatal conductance of 3- to 4-week-old plants was 915

recorded. Data points represent means ± SEM; n = 8-9 (A), 8-10 (B), 9-10 (C), 6-8 (D), 3-12 (E) plants 916

analyzed in 2 independent experiments. 917

Supplementary Figure 2. Q-PCR analysis of mur1-9, ger1 and ger2 mutants. 918

(A) Relative MUR1 transcript level in Col-0 and mur1-9 plants. 919

(B) Relative GER1 transcript level in Col-0, ger1-2 and ger1-3 plants. 920

(C) Relative GER2 transcript level in Col-0, ger2-1 and ger2-2 plants. 921

(A-C) Experiments performed on whole two-week-old plants grown in vitro on ½ x MS medium under 922

12 h light (120 – 160 µmol m-2 s-1)/12 h dark cycle at 22°C/18°C (day/night) temperature. Data bars 923

represent means of three biological replicates ± SD. Asterisks indicate significant differences (*** p < 924

0.001) to Col-0 according to (A) Student's t-test, (B, C) one-way ANOVA followed by Dunnett’s post-925

hoc test; n.d., not detected. 926

Supplementary Figure 3. Water loss-based screen of T-DNA insertion mutants of T7-9 candidate 927

genes. 928

Fresh weight loss of T7-9, control lines (YC3.6, Col-0, slac1-4, ghr1-3) and T-DNA insertion mutants 929

of T7-9 candidate genes recorded after 2h. Data bars represent means ± SD (n = 12 plants). Asterisks 930

denote statistical differences (*** p < 0.001) to respective control lines (Col-0 or YC3.6) according to 931

one-way ANOVA followed by Sidak’s post-hoc test. This figure is an integral part of Figure 3C, values 932

recorded for T7-9 and control lines are provided for reference. Experiment was performed three times 933

with similar results. 934

935


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Supplementary Figure 4. Cuticle permeability of mur1 mutants. 936

Representative photos of middle-aged leaves of 3.5 weeks-old mur1 mutants, respective control lines 937

and cuticule-deficient mutants (aba2-11, snrk2.236) subjected to dye exclusion assay. Leaves were 938

stained with 5 µL drops of toluidine blue solution (0.05 % toluidine blue, 0.01% Tween 20) for 2h and 939

rinsed with water. Dark blue staining indicates high cuticule permeability. Experiment was performed 940

three times with similar results. Scale bar, 10 mm. 941

Supplementary Figure 5. Scanning electron microscopy-based phenotyping of stomatal 942

morphology in mur1 mutants. 943

(A) Stomatal density, (B) mean stomatal pore length and (C) distribution of stomatal pore length in 944

individual plants on abaxial side of 3-week-old cotyledons of Col-0, mur1-1 and mur1-2. (A, B) Data 945

bars represent means ± SD (n = 4-6 cotyledons, 1 cotyledon per plant). Asterisks denote statistical 946

differences (** p < 0.01, *** p < 0.001) to Col-0 according to one-way ANOVA followed Dunnett’s 947

post-hoc test. 948

(D) Frequency of abnormal stomata in Col-0, mur1-1 and mur1-2. Bar represent frequencies obtained 949

for separate plants. 950

(A-D) Experiments were performed three times with similar results. 951

(E, F) Representative images of (E) irregular/obstructed and (F) sealed stomata observed on abaxial 952

side of 3-week-old cotyledons of mur1 mutants. 953

Supplementary Figure 6. Stomatal responses of mur1 mutants to stomata-closing stimuli. 954

(A-E) Whole-rosette stomatal conductance of 3- to 4-week-old mur1-1, mur1-2, Col-0 and ghr1-3 955

(A-D) or ost1-3 (E) plants in response to (A) O3 pulse, (B) elevated CO2, (C) ABA spray, (D) darkness, 956

(E) drop in relative humidity (VPD increase from 1.01 ± 0.01 kPa to 2.04 ± 0.03 kPa (mean ± SE)). 957

The indicated treatments were applied at t = 0. Data points represent means ± SEM; n = 7-10 (A), 10-958

12 (B), 9-11 (C), 6-11 (D), 3-12 (E) plants analyzed in 2 (A, C, D, E) or 3 (B) independent 959

experiments. 960

(F) Changes in low humidity-induced stomatal conductance in Col-0, mur1-1, mur1-2 and ost1-3. 961

Values were calculated by subtracting the stomatal conductance at t = 0 from the stomatal conductance 962

at t = 16 min based on data presented in (E). Data bars represent means ± SEM; n = 3-12 plants. 963


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44

Asterisks denote statistical differences to Col-0 (** p < 0.01) according to one-way ANOVA followed 964

by Dunnett’s post-hoc test. 965

Supplementary Figure 7. Response of mur1 mutants to stomata-opening stimuli. 966

(A, C) Time course of stomatal conductance of mur1-1, Col-0 and ht1-2 plants in response to (A) low 967

CO2 concentration, (C) increase in light intensity. The indicated treatments were applied at t = 0 and 968

whole-rosette stomatal conductance of 3- to 4-week-old plants was recorded. Data points represent 969

means ± SEM; n = 8-10 plants analyzed in 2 independent experiments. 970

(B) Change in stomatal conductance 104 min after decrease in CO2 concentration, calculated based on 971

the data presented in (A). 972

(D) Change in stomatal conductance 56 min after increase of light intensity, calculated based on the 973

data presented in (C). 974

(B, D) Values were calculated by subtracting the initial stomatal conductance recorded at t = 0 from the 975

stomatal conductance obtained at (B) t = 104 min, (D) t = 56 min. Data bars represent means ± SEM; 976

n = 8-10 plants. Asterisks denote statistical differences to Col-0 (*** p < 0.001) according to one-way 977

ANOVA followed by Dunnett’s post-hoc test. 978

Supplementary Figure 8. Analysis of hpGFT1 T1 plants. 979

(A) Rosette morphology of selected hpGFT1 T1 plants, empty vector control (EVC), Col-0 and mur1 980

mutants. Scale bar = 20 mm. 981

(B) Variability in GFT1 transcript level observed in 58 independent hpGFT1 T1 plants. Data points 982

represent values obtained for separate plants. Data bars represent means ± SD; Col-0 n = 3 plants, EVC 983

n = 10 plants. 984

(C) The relationship between the fresh weight loss in 2h and projected rosette area observed in 64 985

independent hpGFT1 T1 plants. Each data point represents values obtained for a single hpGFT1 T1 986

plant. 987

Supplementary Figure 9. The influence of soil borate concentration on leaf fresh-weight loss 988

of bor1-3 mutant. 989


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45

Data bars represent means ± SD (n = 10-16 plants per genotype per condition). Experiment was 990

performed three times with similar results. 991

992


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46

Supplementary Figure 10. Water loss of mutants of enzymes/transporters involved in synthesis 993

of RG-II. 994

Data bars represent means ± SD (n = 13-16 plants). Asterisks denote statistical differences (*** p < 995

0.001) to respective control lines (Col-0 or EVC-empty vector control) according to one-way ANOVA 996

followed by Sidak’s post-hoc test. Experiment was performed three times with similar results. 997

Supplementary Figure 11. Characterization of fut T-DNA insertion mutants. 998

(A-D) Results of RT-PCR obtained for indicated fut mutants with primers specific for (A) FUT2, (B) 999

FUT3, (C) FUT8 and (D) FUT10. PP2AA3 was used as a reference gene and within each panel the 1000

same cDNA sample was used for amplification with all primer sets. The experiment was performed 1001

with three biological replicates and with similar results. Representative results are shown. 1002


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Supplementary Figure 1. Characterization of T7-9 stomatal phenotypes.

Time course of whole-plant stomatal conductance of T7-9, YC3.6 and ghr1-3 (A-D) or ost1-3 (E) in

response to (A) O3 pulse, (B) elevated CO2, (C) ABA spray, (D) darkness and (E) drop in relative air

humidity (VPD increase from 1.01 ± 0.01 kPa to 2.04 ± 0.03 kPa (mean ± SE)). The indicated treatments

were applied at t = 0 and whole-rosette stomatal conductance of 3- to 4-week-old plants was recorded.

Data points represent means ± SEM; n = 8-9 (A), 8-10 (B), 9-10 (C), 6-8 (D), 3-12 (E) plants analyzed

in 2 independent experiments.


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Supplementary Figure 2. Q-PCR analysis of mur1-9, ger1 and ger2 mutants.

(A) Relative MUR1 transcript level in Col-0 and mur1-9 plants.

(B) Relative GER1 transcript level in Col-0, ger1-2 and ger1-3 plants.

(C) Relative GER2 transcript level in Col-0, ger2-1 and ger2-2 plants.

(A-C) Experiments performed on whole two-week-old plants grown in vitro on ½ x MS medium under

12 h light (120 – 160 µmol m-2 s-1)/12 h dark cycle at 22°C/18°C (day/night) temperature. Data bars

represent means of three biological replicates ± SD. Asterisks indicate significant differences (*** p <

0.001) to Col-0 according to (A) Student's t-test, (B, C) one-way ANOVA followed by Dunnett’s

post-hoc test; n.d., not detected.


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Supplementary Figure 3. Water loss-based screen of T-DNA insertion mutants of T7-9 candidate

genes.

Fresh weight loss of T7-9, control lines (YC3.6, Col-0, slac1-4, ghr1-3) and T-DNA insertion mutants

of T7-9 candidate genes recorded after 2h. Data bars represent means ± SD (n = 12 plants). Asterisks

denote statistical differences (*** p < 0.001) to respective control lines (Col-0 or YC3.6) according to

one-way ANOVA followed by Sidak’s post-hoc test. This figure is an integral part of Figure 3C, values

recorded for T7-9 and control lines are provided for reference. Experiment was performed three times

with similar results.


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Supplementary Figure 4. Cuticle permeability of mur1 mutants.

Representative photos of middle-aged leaves of 3.5 weeks-old mur1 mutants, respective control lines and

cuticule-deficient mutants (aba2-11, snrk2.236) subjected to dye exclusion assay. Leaves were stained

with 5 µL drops of toluidine blue solution (0.05 % toluidine blue, 0.01% Tween 20) for 2h and rinsed

with water. Dark blue staining indicates high cuticule permeability. Experiment was performed three

times with similar results. Scale bar, 10 mm.


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Supplementary Figure 5. Scanning electron microscopy-based phenotyping of stomatal

morphology in mur1 mutants.

(A) Stomatal density, (B) mean stomatal pore length and (C) distribution of stomatal pore length in

individual plants on abaxial side of 3-week-old cotyledons of Col-0, mur1-1 and mur1-2. (A, B) Data


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6

bars represent means ± SD (n = 4-6 cotyledons, 1 cotyledon per plant). Asterisks denote statistical

differences (** p < 0.01, *** p < 0.001) to Col-0 according to one-way ANOVA followed Dunnett’s

post-hoc test.

(D) Frequency of abnormal stomata in Col-0, mur1-1 and mur1-2. Bar represent frequencies obtained for

separate plants.

(A-D) Experiments were performed three times with similar results.

(E, F) Representative images of (E) irregular/obstructed and (F) sealed stomata observed on abaxial side

of 3-week-old cotyledons of mur1 mutants.


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Supplementary Figure 6. Stomatal responses of mur1 mutants to stomata-closing stimuli.

(A-E) Whole-rosette stomatal conductance of 3- to 4-week-old mur1-1, mur1-2, Col-0 and ghr1-3 (A-D)

or ost1-3 (E) plants in response to (A) O3 pulse, (B) elevated CO2, (C) ABA spray, (D) darkness, (E)

drop in relative humidity (VPD increase from 1.01 ± 0.01 kPa to 2.04 ± 0.03 kPa (mean ± SE)). The


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indicated treatments were applied at t = 0. Data points represent means ± SEM; n = 7-10 (A), 10-12 (B),

9-11 (C), 6-11 (D), 3-12 (E) plants analyzed in 2 (A, C, D, E) or 3 (B) independent experiments.

(F) Changes in low humidity-induced stomatal conductance in Col-0, mur1-1, mur1-2 and ost1-3. Values

were calculated by subtracting the stomatal conductance at t = 0 from the stomatal conductance at t = 16

min based on data presented in (E). Data bars represent means ± SEM; n = 3-12 plants. Asterisks denote

statistical differences to Col-0 (** p < 0.01) according to one-way ANOVA followed by Dunnett’s post-

hoc test.


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Supplementary Figure 7. Response of mur1 mutants to stomata-opening stimuli.

(A, C) Time course of stomatal conductance of mur1-1, Col-0 and ht1-2 plants in response to (A) low

CO2 concentration, (C) increase in light intensity. The indicated treatments were applied at t = 0 and

whole-rosette stomatal conductance of 3- to 4-week-old plants was recorded. Data points represent means

± SEM; n = 8-10 plants analyzed in 2 independent experiments.

(B) Change in stomatal conductance 104 min after decrease in CO2 concentration, calculated based on

the data presented in (A).

(D) Change in stomatal conductance 56 min after increase of light intensity, calculated based on the data

presented in (C).

(B, D) Values were calculated by subtracting the initial stomatal conductance recorded at t = 0 from the

stomatal conductance obtained at (B) t = 104 min, (D) t = 56 min. Data bars represent means ± SEM;

n = 8-10 plants. Asterisks denote statistical differences to Col-0 (*** p < 0.001) according to one-way

ANOVA followed by Dunnett’s post-hoc test.


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Supplementary Figure 8. Analysis of hpGFT1 T1 plants.

(A) Rosette morphology of selected hpGFT1 T1 plants, empty vector control (EVC), Col-0 and mur1

mutants. Scale bar = 20 mm.


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(B) Variability in GFT1 transcript level observed in 58 independent hpGFT1 T1 plants. Data points

represent values obtained for separate plants. Data bars represent means ± SD; Col-0 n = 3 plants, EVC

n = 10 plants.

(C) The relationship between the fresh weight loss in 2h and projected rosette area observed in 64

independent hpGFT1 T1 plants. Each data point represents values obtained for a single hpGFT1 T1 plant.


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Supplementary Figure 9. The influence of soil borate concentration on leaf fresh-weight loss

of bor1-3 mutant.

Data bars represent means ± SD (n = 10-16 plants per genotype per condition). Experiment was

performed three times with similar results.


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Supplementary Figure 10. Water loss of mutants of enzymes/transporters involved in synthesis

of RG-II.

Data bars represent means ± SD (n = 13-16 plants). Asterisks denote statistical differences (*** p <

0.001) to respective control lines (Col-0 or EVC-empty vector control) according to one-way ANOVA

followed by Sidak’s post-hoc test. Experiment was performed three times with similar results.


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Supplementary Figure 11. Characterization of fut T-DNA insertion mutants.

(A-D) Results of RT-PCR obtained for indicated fut mutants with primers specific for (A) FUT2, (B)

FUT3, (C) FUT8 and (D) FUT10. PP2AA3 was used as a reference gene and within each panel the same

cDNA sample was used for amplification with all primer sets. The experiment was performed with three

biological replicates and with similar results. Representative results are shown.


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Supplementary Table 1. Genes included in candidate gene sequencing.

Functional group Protein name Abbreviation AGI code References

Ion channels SLOW ANION CHANNEL-ASSOCIATED1 SLAC1 AT1G12480 (Vahisalu et al., 2008)

(Negi et al., 2008)

QUICK-ACTIVATING ANION CHANNEL1 QUAC1 AT4G17970 (Sasaki et al., 2010)

(Meyer et al., 2010)

LRR receptor-like

pseudokinase

GUARD CELL HYDROGEN PEROXIDE-

RESISTANT1

GHR1 AT4G20940 (Hua et al., 2012)

(Sierla et al., 2018)

ABA synthesis ABA DEFICIENT1 ABA1 AT5G67030 (Koornneef et al., 1982)

(Marin et al., 1996)

ABA DEFICIENT2 ABA2 AT1G52340 (Léon-Kloosterziel et al., 1996)

(González-Guzmán et al., 2002)

ABA DEFICIENT3 ABA3 AT1G16540 (Léon-Kloosterziel et al., 1996)

(Xiong et al., 2001)

(Bittner et al., 2001)

ABA DEFICIENT4 ABA4 AT1G67080 (North et al., 2007)

ABSCISIC ALDEHYDE OXIDASE3 AAO3 AT2G27150 (Seo et al., 2000)


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NINE-CIS-EPOXYCAROTENOID DIOXYGENASE3 NCED3 AT3G14440 (Iuchi et al., 2001)

Proton pump H+-ATPASE 1 AHA1 AT2G18960 (Merlot et al., 2007)

Kinases OPEN STOMATA1 OST1 AT4G33950 (Mustilli et al., 2002)

(Yoshida et al., 2002)

HIGH LEAF TEMPERATURE1 HT1 AT1G62400 (Hashimoto et al., 2006)

(Hashimoto-Sugimoto et al.,

2016)

(Hõrak et al., 2016)

MITOGEN-ACTIVATED PROTEIN KINASE12 MPK12 AT2G46070 (Des Marais et al., 2014)

(Jakobson et al., 2016)

(Hõrak et al., 2016)

PP2Cs ABA INSENSITIVE1 ABI1 AT4G26080 (Meyer et al., 1994)

(Leung et al., 1994)

ABA INSENSITIVE2 ABI2 AT5G57050 (Leung et al., 1997)

HYPERSENSITIVE TO ABA1 HAB1 AT1G72770 (Saez et al., 2004)

HIGHLY ABA-INDUCED PP2C GENE1 HAI1 AT5G59220 (Antoni et al., 2012)

HIGHLY ABA-INDUCED PP2C GENE2 HAI2 AT1G07430 (Lim et al., 2012)


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PROTEIN PHOSPHATASE 2CA PP2CA AT3G11410 (Kuhn et al., 2006)

ABA transport ATP-BINDING CASSETTE G22 ABCG22 AT5G06530 (Kuromori et al., 2011)

Strigolactone-

related

MORE AXILLARY BRANCHES2 MAX2 AT2G42620 (Piisilä et al., 2015)

(Bu et al., 2014)

(Ha et al., 2014)

DWARF14 D14 AT3G03990 (Li et al., 2017)

(Zhang et al., 2018)

(Kalliola et al., 2020)

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21

Supplementary Table 2. High-frequency single nucleotide polymorphisms identified in T7-9 BC1F2 mapping population. C

hro

moso

me

Muta

tion

Fre

quen

cy

AG

I co

de

Ref

eren

ce a

a

Muta

nt

aa

Annota

tion

Muta

nt

lines

Wat

er l

oss

3 C 19007831 T 0.95 At3g51160 E K GDP-D-MANNOSE-4,6-DEHYDRATASE2, GMD2,

MUR1, MURUS 1

mur1-1 high

mur1-2 high

SALK_057153 high

3 C 17388552 T 0.93 At3g47220 R C ATPLC9, PHOSPHATIDYLINOSITOL-SPECIFIC

PHOSPHOLIPASE C9, PLC9

SALK_025949 WT-like

SALK_021982 WT-like

3 C 17021471 T 0.91 At3g46330 G D MATERNAL EFFECT EMBRYO ARREST39, MEE39 GK_277G01

WT-like

SALK_108641 WT-like

SALK_065070 WT-like

3 C 18009773 T 0.91 At3g48590 E K ATHAP5A, HAP5A, NF-YC1, NUCLEAR FACTOR Y,

SUBUNIT C1

SALK_086334

WT-like

3 C 18556988 T 0.91 At3g50050 R H Eukaryotic aspartyl protease family protein SALK_038145 WT-like


https://doi.org/10.1101/2020.06.04.134353

22

Supplementary Table 3. Genotypes of F3 plants obtained from ger1/ ger2/ F2 plants.

* not determined; ** not germinated

F2 plant F3 segregation

ger1-2/ ger2-1+/+ ger1-2/ ger2-1+/ ger1-2/ ger2-1/ n.d.* n.g.** Total

ger1-2/ ger2-1+/ plant 7/3 22 23 - - 3 48

ger1-2/ ger2-1+/ plant 9/5 24 22 - 1 1 48

ger1-2/ ger2-2+/+ ger1-2/ ger2-2+/ ger1-2/ ger2-2/

ger1-2/ ger2-2+/ plant 26/2 19 20 - - 1 40

ger1-2/ ger2-2+/ plant 33/3 22 18 - - - 40


ger1-3/ ger2-1+/ plant 28/2 17 23 - - - 40

ger1-3/ ger2-1+/ plant 31/8 28 11 - 1 - 40


ger1-3/ ger2-2+/ plant 37/5 19 21 - - - 40

ger1-3/ ger2-2+/ plant 38/6 19 16 - 1 4 40


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https://www.ncbi.nlm.nih.gov/pubmed?term=Kong Y%2C Pe�a MJ%2C Renna L%2C Avci U%2C Pattathil S%2C Tuomivaara ST%2C Li X%2C Reiter WD%2C Brandizzi F%2C Hahn MG%2C et al %282015%29 Galactose%2Ddepleted xyloglucan is dysfunctional and leads to dwarfism in Arabidopsis%2E Plant Physiol 167%3A 1296%26%23x&dopt=abstract

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https://scholar.google.com/scholar?as_q=Galactose-depleted xyloglucan is dysfunctional and leads to dwarfism 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

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https://www.ncbi.nlm.nih.gov/pubmed?term=Kotake T%2C Hojo S%2C Tajima N%2C Matsuoka K%2C Koyama T%2C Tsumuraya Y %282008%29 A bifunctional enzyme with L%2Dfucokinase and GDP%2DL%2Dfucose pyrophosphorylase activities salvages free L%2Dfucose in Arabidopsis%2E J Biol Chem 283%3A 8125%26%23x&dopt=abstract

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https://scholar.google.com/scholar?as_q=A bifunctional enzyme with L-fucokinase and GDP-L-fucose pyrophosphorylase activities salvages free L-fucose 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 bifunctional enzyme with L-fucokinase and GDP-L-fucose pyrophosphorylase activities salvages free L-fucose in Arabidopsis.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kotake&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Krupkov� E%2C Immerzeel P%2C Pauly M%2C Schm�lling T %282007%29 The TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co%2Dordinated plant development%2E Plant J 50%3A 735%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co-ordinated plant 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 TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co-ordinated plant development.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Krupková&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kwak JM%2C Mori IC%2C Pei Z%2DM%2C Leonhardt N%2C Torres MA%2C Dangl JL%2C Bloom RE%2C Bodde S%2C Jones JDG%2C Schroeder JI %282003%29 NADPH oxidase AtrbohD and AtrbohF genes function in ROS�dependent ABA signaling in Arabidopsis%2E EMBO J 22%3A 2623%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=NADPH oxidase AtrbohD and AtrbohF genes function in ROS�dependent ABA signaling 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=NADPH oxidase AtrbohD and AtrbohF genes function in ROS�dependent ABA signaling in Arabidopsis.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kwak&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Langmead B%2C Salzberg SL %282012%29 Fast gapped%2Dread alignment with Bowtie 2%2E Nat Methods 9%3A 357%26%23x&dopt=abstract

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https://scholar.google.com/scholar?as_q=Fast gapped-read alignment with Bowtie 2.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Langmead&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

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https://scholar.google.com/scholar?as_q=Guard cell metabolism and stomatal function&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=Guard cell metabolism and stomatal function&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lawson&as_ylo=2020&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Lee SC%2C Lan W%2C Buchanan BB%2C Luan S %282009%29 A protein kinase%2Dphosphatase pair interacts with an ion channel to regulate ABA signaling in plant guard cells%2E PNAS 106%3A 21419%26%23x&dopt=abstract

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https://scholar.google.com/scholar?as_q=A protein kinase-phosphatase pair interacts with an ion channel to regulate ABA signaling in plant guard cells.&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=A protein kinase-phosphatase pair interacts with an ion channel to regulate ABA signaling in plant guard cells.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lee&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Leonard R%2C Costa G%2C Darrambide E%2C Lhernould S%2C Fleurat%2DLessard P%2C Carlue M%2C Gomord V%2C Faye L%2C Maftah A %282002%29 The presence of Lewis a epitopes in Arabidopsis thaliana glycoconjugates depends on an active α4%2Dfucosyltransferase gene%2E Glycobiology 12%3A 299%26%23x&dopt=abstract

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https://scholar.google.com/scholar?as_q=The presence of Lewis a epitopes in Arabidopsis thaliana glycoconjugates depends on an active �?±4-fucosyltransferase gene.&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 presence of Lewis a epitopes in Arabidopsis thaliana glycoconjugates depends on an active �?±4-fucosyltransferase gene.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Leonard&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Levesque%2DTremblay G%2C Pelloux J%2C Braybrook SA%2C M�ller K %282015%29 Tuning of pectin methylesterification%3A consequences for cell wall biomechanics and development%2E Planta&dopt=abstract

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https://www.ncbi.nlm.nih.gov/pubmed?term=Merilo E%2C Yarmolinsky D%2C Jalakas P%2C Parik H%2C Tulva I%2C Rasulov B%2C Kilk K%2C Kollist H %282018%29 Stomatal VPD response%3A There is more to the story than ABA%2E Plant Physiol 176%3A 851%26%23x&dopt=abstract

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https://doi.org/10.1101/2020.06.04.134353

1 short title: synthesis of gdp-l-fucose is required for … · 2020. 6. 4. · 1 1 short title:...

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