nucleotide metabolism in plants - plant physiology1 1 author for contact: claus-peter witte 1,...

1

Author for contact: Claus-Peter Witte 1, Leibniz Universität Hannover, Department of 1

Molecular Nutrition and Biochemistry of Plants, Herrenhäuser Str. 2, 30419 Hannover, 2

Germany 3

Update: Nucleotide Metabolism in Plants 4

Claus-Peter Wittea,2,3 and Marco Herdea 5

a Leibniz Universität Hannover, Department of Molecular Nutrition and Biochemistry of 6

Plants, Herrenhäuser Str. 2, 30419 Hannover, Germany 7

2 Author for contact 8 3 Senior author 9

ORCID ID: 0000-0002-3617-7807 (C.-P.W.) 10

ORCID ID: 0000-0003-2804-0613 (M.H.) 11

One-sentence summary: Nucleotide metabolism is an essential function in plants. 12

13

Author contributions: C.-P.W. conceived the study, C.-P.W. and M.H. wrote the article 14

Funding: The authors acknowledge funding from the Deutsche Forschungsgemeinschaft 15

(WI3411/4-1 to C.-P.W. and HE 5949/3-1 to M.H.) and the Bundesministerium für Bildung 16

und Forschung (Nutzpflanzen der Zukunft - 031B0540). 17

E-mail address of the author for contact: [email protected] 18

19

Nucleotide metabolism in plants 20

Nucleotides are essential for life. It is easy to validate this statement – one just needs to 21

recall that nucleotides are the building blocks of DNA and RNA, and that many molecules 22

which are central for metabolism, for example ATP, NADH, Co-A and UDP-glucose, are 23

nucleotides or contain nucleotide moieties. Generally, a nucleotide is defined as a 24

phosphorylated ribose or deoxyribose linked to a nitrogen-containing heterocyclic group 25

called the nucleobase via a glycosidic bond (Figure 1). Because of the phosphate groups, 26

nucleotides are negatively charged, whereas at neutral pH nucleosides and nucleobases are 27

uncharged. The exception is xanthine, which is partially charged as a free base (pKa = 7.4) 28

but completely charged at the base in xanthosine (pKa = 5.5) or the corresponding 29

nucleotides (Figure 1, Sigel et al., 2009). 30

Many excellent reviews focus on general (Wagner and Backer, 1992; Zrenner et al., 2006; 31

Zrenner and Ashihara, 2011; Stasolla et al., 2003; Moffatt and Ashihara, 2002) or particular 32

Plant Physiology Preview. Published on October 22, 2019, as DOI:10.1104/pp.19.00955

Copyright 2019 by the American Society of Plant Biologists

www.plantphysiol.orgon April 24, 2020 - Published by Downloaded from Copyright © 2019 American Society of Plant Biologists. All rights reserved.

mailto:[email protected]

http://www.plantphysiol.org

2

aspects of (Smith and Atkins, 2002; Ashihara et al., 2018; Kafer et al., 2004) plant nucleotide 33

metabolism. The aim of this review is to provide an update on how nucleotide metabolism is 34

hardwired, mostly focusing on the cellular level, because our understanding of the 35

organization at the tissue and organ level remains very limited. The presented models are 36

mostly based on results from Arabidopsis thaliana. These will often be valid for most plants, 37

but certainly there will be species-dependent variations. We also cover extracellular 38

nucleotide metabolism and review the evidence for overlap between cytokinin metabolism 39

and central nucleotide metabolism. Figure 2 shows a general overview of plant nucleotide 40

metabolism. 41

DE NOVO SYNTHESIS 42

Purine de novo synthesis 43

Plants possess the metabolic pathways for the de novo synthesis of purine nucleotides 44

generating AMP as well as pyrimidine nucleotides yielding UMP. During de novo 45

biosynthesis, nucleotides are newly synthesized from the general metabolites activated 46

ribose (5-phosphoribosyl-1-pyrophosphate, PRPP), glutamine, aspartate, and bicarbonate as 47

well as specifically for the purine nucleotides, glycine and formyl tetrahydrofolate (Figure 2). 48

There is strong evidence that AMP biosynthesis occurs entirely in the plastids, because the 49

11 enzymes (catalyzing 12 reactions; Smith and Atkins, 2002) required for AMP biosynthesis 50

in Arabidopsis all have an N-terminal organelle targeting peptide, and C-terminal yellow 51

fluorescent protein (YFP)-fusion proteins of several of these enzymes were observed 52

exclusively in the plastids when they were transiently expressed in Nicotiana benthamiana 53

in our laboratory (N. Medina Escobar and C.-P. Witte, unpublished data) (Figure 3A). In rice 54

(Oryza sativa), the pathway also seems to reside in plastids (Zhang et al., 2018). However, it 55

has been reported that in nodules of the tropical legume cowpea (Vigna unguiculata), 56

purine biosynthesis is targeted to plastids and mitochondria (Atkins et al., 1997; Smith and 57

Atkins, 2002). It may be worthwhile to reconfirm this special localization in nodules using 58

fluorescent tagged proteins. 59

AMP is exported from the plastids by the adenine nucleotide uniporter brittle1 (BT1, Figure 60

3A, number 1), which can also transport ADP and ATP (Hu et al., 2017; Kirchberger et al., 61

2008; Leroch et al., 2005). Interestingly, BT1 from Arabidopsis and maize (Zea mays) was 62

reported to be dual localized to the chloroplast and mitochondria (Bahaji et al., 2011b) and 63



3

a bt1 mutant with a severe dwarf phenotype could be complemented with an N-terminally 64

truncated version of BT1 that is exclusively located in the mitochondria and not in the 65

plastids (Bahaji et al., 2011a). This either indicates that there are certain tissues or 66

developmental stages where purine nucleotide biosynthesis occurs mainly in mitochondria, 67

or that BT1 has an alternative and essential function in this organelle. In the latter case, BT1 68

might still also be involved in exporting adenylates from plastids, but the abrogation of the 69

plastidic variant would not cause the strong dwarf phenotype, indicating that de novo 70

synthesized AMP has an alternative, BT1 independent, way to leave the plastids. 71

GMP biosynthesis requires inosine monophosphate (IMP, Figure 3A), which can either be 72

derived from AMP deamination in the cytosolic compartment catalyzed by AMP deaminase 73

(AMPD, Figure 3A, #2) or from direct export of IMP from the plastids, because IMP is 74

generated there en route to AMP (Figure 3A). Mutation of AMPD is zygote lethal (Xu et al., 75

2005) and coformycin, an AMPD inhibitor, is a potent herbicide after its phosphorylation in 76

vivo (Dancer et al., 1997). These phenotypic effects might be caused by hampered GMP 77

biosynthesis, suggesting that AMPD could be required for this process. Consistent with this, 78

AMPD is strongly activated by ATP (Han et al., 2006) and this regulation might balance 79

cellular ATP and GTP concentrations. However, it has also been reported that AMPD 80

inhibition might be detrimental by severely altering the cellular energy charge and that the 81

GTP pool is not altered upon AMPD inhibition (Sabina et al., 2007), implying that GMP 82

biosynthesis is independent of AMPD and that there is an alternative IMP supply from the 83

plastids. The activity of AMPD likely resides in the cytosol, but the protein has an N-terminal 84

transmembrane domain and is clearly attached to a membrane (Han et al., 2006). 85

The following enzymatic reactions for GMP biosynthesis, (i) the oxidation of IMP to 86

xanthosine monophosphate (XMP, Figure 1) by IMP dehydrogenase (IMPDH, Figure 3A, #3), 87

and (ii) the amination of XMP to GMP by GMP synthetase (GMPS, Figure 3A, #4) probably 88

take place in the cytosol. Both enzymes have no apparent subcellular targeting peptide and 89

were detected in the cytosolic proteome of Arabidopsis (Ito et al., 2011). Also, IMPDH from 90

cowpea nodules was associated with the cytosolic fraction (Shelp and Atkins, 1983). GMP is 91

a quite strong competitive inhibitor of IMPDH (Atkins et al., 1985) maybe resulting in 92

feedback regulation in vivo. 93



4

Pyrimidine de novo synthesis 94

The first dedicated reaction of UMP biosynthesis (Figure 3B) is catalyzed by asparate 95

transcarbamoylase (ATCase, Figure 3B, #5). The enzyme is located in the plastids. The next 96

enzyme, dihydroorotatase (DHOase, Figure 3B, #6) was associated to plastids in cell 97

fractionation studies (Doremus and Jagendorf, 1985 and references on the SUBA web 98

server, Hooper et al., 2017) but was located in the cytosol when transiently overexpressed 99

in Arabidopsis protoplasts as a green fluorescent protein (GFP)-tagged fusion protein (Witz 100

et al., 2012). Maybe DHOase is associated with the chloroplast membrane via a protein-101

protein interaction, and the interaction partner is overwhelmed by strong overexpression of 102

DHOase. The following enzyme, dihydroorotate dehydrogenase (DHODH, Figure 3B, #7), is 103

associated with mitochondria (Witz et al., 2012; Doremus and Jagendorf, 1985) and likely 104

located on the outer surface of the inner mitochondrial membrane as observed for the 105

mammalian orthologs (Ullrich et al., 2002). UMP synthase (UMPS, Figure 3B, #8), the final 106

enzyme, was associated with the plastids and the cytosol, as shown by cell fractionation of 107

pea (Pisum sativum) leaves (Doremus and Jagendorf, 1985) and was present in the cytosol 108

after overexpression in Arabidopsis protoplasts (Witz et al., 2012). Thus, UMP is generated 109

in the cytosol, while it appears that the responsible enzyme might have some affinity for the 110

chloroplast. Because ATCase (Figure 3B, #5) is feedback regulated by uridylates, in particular 111

UMP (Doremus and Jagendorf, 1985), the cytosolic uridine nucleotide pool must be tightly 112

connected to the plastidic pool. 113

CTP biosynthesis requires the phosphorylation of UMP to UTP (see below), which is the 114

substrate of CTP synthetase (CTPS, Figure 3B, #9). There are five CTPS isoenzymes in 115

Arabidopsis, and all reside in the cytosol. For one isoform (CTPS3) the activity and 116

stimulatory allosteric regulation by UTP and GTP has been recently shown (Daumann et al., 117

2018). Interestingly, some CTPS isoenzymes form filamentous aggregates, called 118

cytoophidia, inactivating the enzyme. In vitro, these are generated in particular in the 119

presence of CTP, indicating that the enzyme is feed-back regulated by this mechanism 120

(Daumann et al., 2018). Knockout mutants for each CTPS were characterized and except for 121

CTPS2, which showed a complete block of germination, no phenotypes were observed in the 122

single mutants (Daumann et al., 2018), indicating redundancy of the CTPS enzymes in most 123

situations. 124



5

The generation of nucleoside triphosphates 125

The final steps of UMP, GMP and CTP biosynthesis occur in the cytosol. With the exception 126

of CTP, which is directly synthesized from UTP, purine and pyrimidine nucleoside 127

triphosphate synthesis is achieved by phosphorylation of the respective monophosphates 128

(Figure 4). 129

The plastids and the mitochondria, which possess their own transcription and translation 130

machineries, must be supplied with ribonucleotides and deoxyribonucleotides from the 131

cytosol. Not much is known about (i) the phosphorylation state in which nucleotides are 132

taken up, (ii) which transporters are involved, (iii) if the concentrations of (desoxy) 133

nucleotides differ in the distinct cellular compartments and how this may be regulated –134

subcellular distributions have been estimated only for the adenylates (Stitt et al., 1982). 135

Describing the subcellular distribution of the enzymes involved in the last two steps of 136

mononucleotide phosphorylation can help in building hypotheses regarding the exact 137

nucleotide species imported into organelles. 138

The pyrimidine nucleotides, UMP and CMP, are phosphorylated by UMP kinases (UMK, 139

Figure 4, A and B, #12) Arabidopsis possess two evolutionarily distinct families of such 140

enzymes (i) UMKs related to adenylate kinases (AMKs) encoded by four genes and (ii) UMKs 141

related to eubacterial UMP kinases encoded by two genes. The AMK-like UMKs have not yet 142

been characterized, except for a biochemical analysis of UMK3 (At5g26667), which was 143

shown to utilize UMP and CMP as the best substrates (Zhou et al., 1998). These enzymes 144

have been predicted to reside in the cytosol and the mitochondria (Lange et al., 2008). From 145

the eubacterial UMP kinase family, one member called ‘plastid UMP kinase’ (PUMKIN, 146

At3g18680) was shown to be located in chloroplasts, and to have UMK activity in vitro. 147

Interestingly, the enzyme binds certain plastidic transcripts and is involved in plastid RNA 148

metabolism, which may not require its enzymatic function. Mutants are small and 149

compromised in plastid translation and photosynthetic performance (Schmid et al., 2019). 150

The orthologous enzyme in rice is localized in chloroplasts, participates in RNA metabolism, 151

and the corresponding loss-of-function mutants are pale green (Chen et al., 2018a; Zhu et 152

al., 2016). Additionally, they contain less UDP and more UMP (Dong et al., 2019) suggesting 153

that UMP phosphorylation in the chloroplast is functionally important. The phosphorylation 154

of TMP is not catalyzed by UMKs, but by a dedicated thymidine monophosphate kinase 155



6

(TMK, Figure 4B, #16). In Arabidopsis, a mitochondrial and a cytosolic version are generated 156

from a single gene by alternative splicing. Mutation of TMK leads to early seed abortion at 157

the zygote state (Ronceret et al., 2008). 158

The phosphorylation of GMP to GDP and dGMP to dGDP (Kumar et al., 2000) is catalyzed by 159

guanylate kinases (GMK, Figure 4C, #17) of which plants have two different types, a 160

cytosolic type (GKc), and a organelle type, dual targeted to plastids and mitochondria 161

(GKpm) (Figure 4C). The activity of GKpm in rice, pea, and Arabidopsis is regulated by 162

guanosine 3’, 5’-bisdiphosphate (ppGpp), a bacterial and plastid signaling molecule (Nomura 163

et al., 2014). Suppression of Arabidopsis GKpm (At3g06200) transcripts by RNA interference 164

(RNAi) results in a pale green or albino phenotype (Sugimoto et al., 2007) emphasizing the 165

importance of nucleotide monophosphate transport into organelles. Interestingly, the loss-166

of-function mutant of the rice GKpm gene is pale green, but does not exhibit DNA depletion 167

in the organelles, suggesting that deoxynucleotide and ribonucleotide metabolism are not 168

fully linked (Sugimoto et al., 2007). 169

Adenylate monophosphate is converted by adenylate kinase (AMK, Figure 4D, #18) into 170

adenosine diphosphate (ADP). Arabidopsis has seven AMK isoforms. AMK1, AMK2, and 171

AMK5 are located in the plastids, AMK3 and AMK4 reside in the cytosol, and AMK7 is 172

present in the mitochondria. For AMK1 a mitochondrial localization has also been observed 173

(Carrari et al., 2005; Lange et al., 2008). Because the plastids harbor the de novo synthesis 174

for AMP, they will not require net import of adenylates. Consistently, the loss-of-function 175

mutant for AMK2 has a bleached phenotype (Lange et al., 2008) suggesting that there is no 176

net ADP or ATP import to compensate for compromised adenlyate kinase activity in plastids. 177

A strong reduction of plastidic AMK activity in rice also results in an albino phenotype (Wei 178

et al., 2017). In potato (Solanum tuberosum), a reduction of plastidic AMK activity led to an 179

increase of the adenylate pool (AMP, ADP, ATP, and ADP-glucose) and increased starch 180

synthesis in tubers, but it is still not understood how plastidic AMK and adenylate de novo 181

synthesis are connected (Regierer et al., 2002). Interestingly, it was suggested that the 182

AMKs might contribute to protect RNA from random mis-incorporation of methyl-6 A marks. 183

AMKs are highly selective for AMP versus N6-methyl AMP released during the degradation 184

of RNA species carrying this abundant A modification. The selectivity of the AMKs possibly 185



7

prevents the formation of N6-methyl ATP, which is a substrate of RNA polymerase II (Chen 186

et al., 2018b). 187

Recently, a broad spectrum mononucleotide kinase only distantly related to the AMKs, but 188

with relatively high adenylate kinase activity was described (At5g60340). This enzyme was 189

localized in the nucleus and a knockout mutant was affected in stem elongation (Feng et al., 190

2012). 191

Plastids and mitochondria possess nucleoside monophosphate kinases for all nucleotides. 192

Thus, nucleoside monophosphates are probably imported into these organelles and are 193

phosphorylated to dinucleotides. Enzymes catalyzing the next step to trinucleotides, the 194

nucleoside diphosphate kinases (NDPKs), should therefore be found in plastids and 195

mitochondria as well as in the cytosol. This has been indeed observed (Luzarowski et al., 196

2017). The exact locations of the enzymes have been debated and a detailed phylogenetic 197

analysis suggests the presence of a fourth enzyme type in the endoplasmic reticulum (ER) 198

(Dorion and Rivoal, 2015). The NDPKs are multi-substrate enzymes accepting all nucleoside / 199

deoxynucleoside diphosphates (Zrenner et al., 2006) but there is a preference for generating 200

GTP (Kihara et al., 2011), which in the chloroplast may assist in repairing photosystem II 201

(Spetea and Lundin, 2012). Mutation of the gene for the plastidic NDPK in rice results in a 202

pale green phenotype and a lower photosynthetic rate (Zhou et al., 2017; Ye et al., 2016), 203

but since the chloroplast function is partially retained, there must be also nucleoside 204

triphosphate import into this organelle. Interestingly, NDPKs can also have moonlighting 205

activity as modulators of gene expression (Dorion and Rivoal, 2018). 206

Besides nucleotides, the nucleus and organelles need deoxynucleotides (dNTPs) for DNA 207

synthesis. Deoxynucleotide synthesis requires the reduction of the hydroxyl moiety on the 208

2’ carbon of the ribose by an enzyme complex called ribonucleotide reductase (RNR, Figure 209

4, #10). The RNR complex is comprised of two large regulatory (R1) and two small catalytic 210

(R2) subunits. Mutation of the major R2 subunit gene (tso2) results in lower dNTP 211

concentrations and abnormal plant development, while the additional mutation of a further 212

R2 subunit gene (Arabidopsis has three R2 subunit genes in total) is lethal (Wang and Liu, 213

2006). The substrates of RNR are the ribonucleotide diphosphates, suggesting that for CTP a 214

dedicated phosphatase might exist to support dCDP synthesis (Figure 4A). Alternatively, CDP 215

for dCTP synthesis might be generated from salvage of cytidine (see below). Interestingly, 216



8

RNR is subject to a complex allosteric regulation to adjust the correct dNTP pool sizes 217

(Sauge-Merle et al., 1999). In plants, RNR resides exclusively in the cytosol with the potential 218

to relocate to the nucleus upon exposure to UV radiation (Lincker et al., 2004). Especially 219

the plastidic DNA replication seems to rely strongly on sufficient RNR activity, because 220

partially compromising the function of the large RNR subunit by different mutations in the 221

corresponding gene resulted in reduced dNTP levels and impaired chloroplast division in 222

Arabidopsis (Garton et al., 2007). Consistently, chlorophyll biosynthesis in rice is reduced in 223

mutants of the small RNR subunit genes (Chen et al., 2015). All dNDPs can be synthesized 224

directly by RNR, except thymidine diphosphate, because it has no ribonucleotide 225

counterpart. Instead, RNR catalyzes the formation of dUDP from UDP (Figure 4B) and dUMP 226

is methylated at C5 to TMP catalyzed by thymidilate synthase. In Arabidopsis, three 227

enzymes were recently characterized as thymidylate synthases, which are also dihydrofolate 228

reductases (DHFR-TS, Figure 4B, #15) with only two isoforms displaying thymidylate 229

synthase activity (Gorelova et al., 2017). Interestingly, in roots all isoforms can reside either 230

in the cytosol, the nucleus, or the mitochondria depending on the developmental state of 231

the cell, but not in plastids. The two active isoforms seem to be redundant, since only a 232

double mutant of the respective genes is lethal, whereas single gene loss-of-function 233

mutants are phenotypically inconspicuous (Gorelova et al., 2017). The substrate for DHAFR-234

TS is dUMP (Gorelova et al., 2017), but the RNR provides dUDP. It is unknown which enzyme 235

links these two processes in vivo. An alternative dUMP source in mitochondria is the 236

deamination of dCMP as shown recently in rice (Niu et al., 2017; Xu et al., 2014). 237

238

SALVAGE AND DEGRADATION 239

Metabolic sources of nucleosides and nucleobases 240

Nucleosides and nucleobases can be released from nucleotides or nucleic acids in 241

metabolism (Figures 2, 5 and 6) or can be taken up from the environment (Girke et al., 242

2014), where they can occur in substantial amounts (Phillips et al., 1997). 243

The main metabolic source for most nucleosides is probably the turnover of RNA, in 244

particular in the vacuole. Vacuolar RNA degradation, for example of ribosomal RNA after 245

ribophagy (Floyd et al., 2015), generates nucleotides which likely are degraded to 246

nucleosides by vacuolar phosphatases. The details of this process have not been 247



9

investigated so far. The tonoplast membrane possesses a nucleoside exporter (equilabrative 248

nucleoside transporter 1, ENT1, At1g70330; Bernard et al., 2011) for the release of 249

nucleosides into the cytoplasm. For adenosine, the turnover of S-adenosyl methionine 250

(SAM), used for methylation reactions, is another important source (Figures 2 and 5). After 251

transfer of the methyl group from SAM, the resulting S-adenosyl homocysteine (SAH) is 252

hydrolyzed to homocysteine and adenosine (Sauter et al., 2013). 253

There are no strong sources for nucleobases in plant metabolism, except for adenine, which 254

is released during polyamine, nicotianamine, and ethylene biosynthesis (Sauter et al., 2013). 255

In all three pathways SAM is used and 5’-methylthioadenosine is generated, which is 256

hydrolyzed to 5-methylthioribose and adenine (Siu et al., 2008). Also the degradation of 257

cytokinins produces small amounts of adenine (Schmülling et al., 2003). Purine bases are 258

released from nucleic acids by spontaneous depurination (Barbado et al., 2018) resulting in 259

low amounts of adenine and guanine. The metabolic source of hypoxanthine is likely the 260

spontaneous deamination of adenine in DNA, resulting in a hypoxanthine base, which is 261

removed from the DNA by base excision repair (Karran and Lindahl, 1980). Similarly, the 262

non-enzymatic deamination of cytosine in DNA generates uracil, which is removed by base 263

excision repair (Figure 2). 264

Purine and Pyrimidine Salvage Metabolism 265

Nucleosides and nucleobases can be converted into nucleotides, which is called ‘salvage’ 266

(Figures 2, 5 and 6). In contrast to de novo biosynthesis of nucleotides, which generates 267

nucleotides from basic metabolites (see above), the salvage reactions recycle nucleobases 268

and nucleosides derived from metabolism or uptake (see previous section) to nucleotides. 269

Nucleobases react with activated phosphoribose (5-phosphoribosyl-1-pyrophosphate, PRPP) 270

to the respective nucleotides - a reaction catalyzed by phosphoribosyltransferases (PRTs). 271

Adenine phosphoribosyltransferase (APRT, Figure 5A, #19), hypoxanthine guanine 272

phosphoribosyltransferase (HGPRT, Figure 5A, #21) and uracil phosphoribosyltransferase 273

(UPRT, Figure 6, #39) are the three types of nucleobase-specific enzymes present in plants. 274

Nucleosides are phosphorylated to nucleoside monophosphates by kinases. Adenosine 275

kinase (ADK, Figure 5A, #20), inosine guanosine kinase (IGK, Figure 5A, #22), and uridine 276

cytidine kinase (UCK, Figure 6, #35) salvage ribonucleotides, whereas thymidine kinase (TK, 277



10

Figure 4B, #14) and deoxynucleoside kinase (dNK, Figure 4, #11) phosphorylate thymidine 278

and the other three deoxynucleosides, respectively. 279

Several salvage enzymes have a critical function for plant metabolism and their mutation 280

has severe consequences. Mutation of the gene for the main APRT (Figure 5, #19) activity, 281

APT1 (At1g27450), results in male sterility (Gaillard et al., 1998b), whereas a strong 282

downregulation increases the resistance to oxidative stress (Sukrong et al., 2012). Deletion 283

or strong downregulation of the gene for the main ADK (Figure 5, #20) enzyme, ADK1 284

(At3g09820), compromises transmethylation reactions, because the accumulating 285

adenosine inhibits S-adenosylhomocystein (SAH) hydrolase – an enzyme of the SAM cycle. It 286

has been shown that ADK1 and SAH hydrolase interact and partially reside in the nucleus 287

probably mediated by nuclear methyltransferases (Lee et al., 2012). Reduced 288

transmethylation causes a range of developmental abnormalities (Young et al., 2006; 289

Moffatt et al., 2002). Guanine and hypoxanthine salvage seems to be less critical, because 290

HGPRT (Figure 5, #21) mutants are phenotypically normal except for a slight delay in 291

germination (Schroeder et al., 2018; Liu et al., 2007). Mutation of HGPRT leads to guanine 292

but not hypoxanthine accumulation in vivo, probably reflecting that guanine can only be 293

salvaged, whereas hypoxanthine can also be degraded (Baccolini and Witte, 2019). Kinase 294

activity for inosine and guanosine has been measured in plant extracts (Deng and Ashihara, 295

2010; Katahira and Ashihara, 2006) (IGK, Figure 5A, #22) but the corresponding gene is still 296

unknown. Some evidence has been provided that the activity is associated with the 297

intermembrane space of mitochondria (Combes et al., 1989). 298

The only uracil salvage activity in Arabidopsis is located in plastids and encoded by UPP 299

(UPRT, Figure 6, #39). Mutation of UPP leads to growth arrest in the seedling stage and an 300

albino phenotype (Mainguet et al., 2009). Interestingly, it was recently shown that this 301

phenotype is unrelated to the lack of UPRT activity in the mutant, but is caused by the 302

absence of the UPP protein per se, demonstrating that uracil salvage does not play such an 303

essential role for Arabidopsis as previously thought (Ohler et al., 2019). Salvage of uridine is 304

more prominent than salvage of uracil in Arabidopsis. Uridine and cytidine salvage are 305

performed by dual-specific uridine and cytidine kinases (UCK) (Ohler et al., 2019). These 306

enzymes also possess a UPRT-like domain, but do not have UPRT activity (Chen and Thelen, 307

2011). Simultaneous mutation of UCK1 and UCK2 results in dwarf plants that fail to reach 308



11

maturity (Chen and Thelen, 2011). Previously, UCK1 and UCK2 were found to localize in 309

plastids (Chen and Thelen, 2011), but Ohler et al. (2019) demonstrated that these enzymes 310

reside in the cytosol, which was also confirmed in our laboratory (M. Chen and C.-P. Witte, 311

unpublished data). 312

Interestingly, deoxynucleoside-specific salvage enzymes also exist (Figure 4) and the 313

abrogation of thymidine salvage by thymidine kinase (TK, Figure 4B, #14) is lethal for plants 314

(Clausen et al., 2012). However, it is unclear, why thymidine salvage is of such importance. 315

Thymidine kinase occurs in the cytosol, the mitochondria, and the plastids (Xu et al., 2015) 316

and is of particular importance for chloroplast maintenance when germinating seedlings 317

turn autotrophic (Pedroza-García et al., 2019). The other deoxynucleosides are salvaged by 318

an enzyme with broad deoxynucleoside specificity (Clausen et al., 2012) (dNK, Figure 4, 319

#11), potentially associated with mitochondria (Clausen et al., 2014). 320

Purine Nucleotide Degradation 321

Instead of being salvaged, nucleobases and nucleosides can also be fully degraded by plants, 322

but for guanine, adenine, and adenosine a salvage reaction needs to precede degradation. 323

Adenine and adenosine first must be converted to AMP, which can then be deaminated by 324

AMP deaminase (AMPD, Figure 5A, #2) to IMP as a first step into degradation. This is 325

necessary because Arabidopsis and plants in general lack adenosine deaminase (Chen et al., 326

2018b; Dancer et al., 1997). Interestingly, for N6-methyl AMP, plants as well as many other 327

eukaryotes possess a special deaminase, called N6-methyl-AMP deaminase (MAPDA) (Chen 328

et al., 2018b). N6-methylated adenine is the most frequent modification in mRNA, but is 329

also present in other RNA species (Chen and Witte, 2019). MAPDA is phylogenetically 330

related to adenosine deaminases and hydrolyzes N6-methyl AMP to IMP removing the 331

aminomethyl group. This example shows that modified nucleotides must also have an 332

access route to general nucleotide degradation. 333

From IMP, the purine nucleotide degradation pathway cannot be entered directly in 334

Arabidopsis, but conversion to XMP and apparently even to GMP is required (Baccolini and 335

Witte, 2019). These recent results show that the route for AMP catabolism and the route for 336

GMP biosynthesis (partially) overlap. Therefore, branch points of both routes must be 337

controlled, but it is not yet clear how this is achieved. GMP dephoshorylation by a so far 338

unknown phosphatase (GMPP, Figure 5A, #23) initiates purine nucleotide catabolism. At the 339



12

stage of guanosine, salvage back to the nucleotide level via IGK (Figure 5A, #22) is still 340

possible, but the GMPP and IGK reactions might be spatially or temporarily separated to 341

avoid a futile cycle. The deamination of guanosine to xanthosine by guanosine deaminase 342

(GSDA, Figure 5A, #24; Dahncke and Witte, 2013) marks the point of no return, because 343

xanthosine cannot be salvaged and is dedicated for degradation (Yin et al., 2014). Although 344

xanthosine appears to be generated mainly by GSDA, there is strong evidence for an 345

alternative route directly from XMP to xanthosine catalyzed by an XMP phosphatase (XMPP, 346

Figure 5A, #25) (Baccolini and Witte, 2019). An XMP-specific phosphatase, which may 347

represent this XMPP, is currently under investigation in our laboratory. In summary, GMP 348

catabolism begins with dephosphorylation and deamination of guanosine, and most AMP is 349

apparently also degraded via GMP, while some might be dephosphorylated already at the 350

stage of XMP. 351

In clear contrast to purine metabolism in many other organisms, guanine is not an 352

intermediate of purine nucleotide catabolism in Arabidopsis and probably most plants. For 353

degradation, guanine must first be salvaged to GMP (Dahncke and Witte, 2013; Baccolini 354

and Witte, 2019). As well in contrast to purine metabolism in many other organisms, inosine 355

and hypoxanthine are not major intermediates of purine nucleotide catabolism in 356

Arabidopsis, because they are not derived from IMP dephosphorylation (Baccolini and 357

Witte, 2019), but possibly from t-RNA turnover and base excision repair of deaminated 358

adenine in DNA (Figure 5A). Because guanine and hypoxanthine do not play an important 359

role in purine nucleotide degradation, HGPRT (Figure 5A, #21) is decoupled from purine 360

catabolism in plants (Baccolini and Witte, 2019), which is in stark contrast to humans, where 361

mutation of HGPRT results in accumulation of purine nucleotide breakdown products and 362

severe phenotypic consequences (Lesch-Nyhan Syndrome) (Torres and Puig, 2007). 363

Purine catabolism can lead to the complete disintegration of the purine ring in plants 364

(Werner and Witte, 2011) (Figure 5B) to recycle nitrogen (Soltabayeva et al., 2018), but is 365

also used to generate the intermediates uric acid and especially allantoin, which counteract 366

stress by reducing reactive oxygen species (Brychkova et al., 2008; Irani and Todd, 2016; 367

Irani and Todd, 2018; Nourimand and Todd, 2019; Lescano et al., 2016; Watanabe et al., 368

2014; Casartelli et al., 2019; Ma et al., 2016). Sometimes also the accumulation of allantoate 369

has been observed (Alamillo et al., 2010). In tropical legumes like soybean (Glycine max) or 370



13

common bean (Phaseolus vulgaris), the ureides allantoin and allantoate are used as long 371

distance nitrogen transport compounds for the export of fixed nitrogen from the nodules 372

(Carter and Tegeder, 2016; Tegeder, 2014) mediated by the ureide permease (UPS) 373

transporters (Collier and Tegeder, 2012; Desimone et al., 2002; Schmidt et al., 2006). 374

Ureides also function in long distance transport in non-nodulated legumes (Diaz-Leal et al., 375

2012; Quiles et al., 2019) and are probably used in many plants for this purpose (Redillas et 376

al., 2019; Lescano et al., 2016). 377

It has recently been shown that xanthosine hydrolysis to xanthine and ribose is catalyzed by 378

a cytosolic nucleoside hydrolase heteromer consisting of nucleoside hydrolase 1 (NSH1, 379

Figure 5A, #26) and nucleoside hydrolase 2 (NSH2, Figure 5A, #27) in vivo (Baccolini and 380

Witte, 2019). NSH1 has only weak xanthosine and inosine but strong uridine hydrolase 381

activity (Jung et al., 2009; Jung et al., 2011; Baccolini and Witte, 2019; Riegler et al., 2011). 382

However, NSH1 is required to activate NSH2, which is the stronger xanthosine and inosine 383

hydrolase in the complex. Nucleoside catabolism is the major metabolic source of ribose, 384

which is recycled to ribose-5-phosphate by ribokinase in the plastids (Riggs et al., 2016; 385

Schroeder et al., 2018). Xanthine and hypoxanthine are catabolized by the same enzyme, 386

xanthine dehydrogenase (XDH, Figure 5A, #28; Urarte et al., 2015) finally to uric acid in the 387

cytosol. Arabidopsis has a second gene encoding XDH (At4g34900) with no apparent 388

xanthine dehydrogenase activity in vivo (Hauck et al., 2014). Interestingly, XDH has been 389

shown to play a dual role during powdery mildew pathogen attack on Arabidopsis (Ma et al., 390

2016). In the epidermis the enzyme is postulated to operate as an NADH oxidase generating 391

superoxide (Zarepour et al., 2010) to prevent fungal entry, whereas in the mesophyll it 392

works as a xanthine dehydrogenase producing urate, which is suggested to function as a 393

reactive oxygen species scavenger. It was proposed that by this mechanism the reactive 394

oxygen species are confined to the infection site. For further degradation, uric acid must be 395

imported into the peroxisomes, but molecular details about this import are still unknown. In 396

the peroxisome, urate is oxidized by urate oxidase (UOX, Figure 5B, #29) as well as 397

hydrolyzed and decarboxylated by allantoin synthase (ALNS, Figure 5B, #30) to (S)-allantoin 398

(Lamberto et al., 2010; Pessoa et al., 2010). Mutation of UOX leads to strong accumulation 399

of uric acid, which is deleterious for peroxisome maintenance in the embryo, leading to a 400

severe suppression of germination and seedling establishment (Hauck et al., 2014) – 401



14

surprisingly, accumulation of similar amounts of xanthine in Arabidopsis plants lacking XDH 402

does not lead to strong phenotypic alterations under standard growth conditions (Hauck et 403

al., 2014; Schroeder et al., 2018; Soltabayeva et al., 2018). Allantoin must be transported 404

from the peroxisomes to the ER for further degradation, but it is unclear how this is 405

achieved. One may speculate that allantoin accumulation under certain stress conditions 406

may be in part caused by altered peroxisome to ER transport efficiency. However, one 407

apparent reason for allantoin accumulation is an altered catalytic capacity for allantoin 408

generation and degradation under stress (Irani and Todd, 2016; Irani and Todd, 2018; 409

Lescano et al., 2016; Casartelli et al., 2019). 410

In the ER, allantoin is hydrolyzed by four enzymes (Figure 5B, #31 to #34) completely 411

releasing the ring nitrogen as ammonia (Werner et al., 2008; Werner et al., 2010; Serventi et 412

al., 2010; Todd and Polacco, 2006). These enzymes are also responsible for supplying the 413

shoot of tropical legumes with nitrogen exported from the nodules as allantoin and 414

allantoate (Werner et al., 2013; Díaz-Leal et al., 2014). 415

Pyrimidine Nucleotide Degradation 416

Pyrimidine nucleotide catabolism is initiated by UMP / CMP phosphatase(s) (UCPP, Figure 6, 417

#36) which have not yet been identified. Their activity might be temporarily and / or 418

spatially separated from uridine cytidine kinases (UCKs, Figure 6, #35; Ohler et al., 2019) to 419

avoid a futile cycle of pyrimidine nucleotide dephosphorylation and pyrimidine nucleoside 420

salvage. Cytidine is deaminated to uridine by a cytosolic cytidine deaminase (CDA, Figure 6, 421

#37). Interestingly, plants can neither degrade nor salvage the free base cytosine (Katahira 422

and Ashihara, 2002). Arabidopsis contains several copies of cytidine deaminase, but only 423

one copy is functional. The mutation of CDA results in smaller plants probably because 424

cytidine accumulation is toxic (Chen et al., 2016). Generally, the accumulation of nucleosides 425

can reduce plant performance as has been shown for GSDA (Figure 5A, #24) mutants 426

(Schroeder et al., 2018). Consistently, transgenic lines with increased vacuolar nucleoside 427

export (Bernard et al., 2011) are smaller than the wild type. 428

Uridine is hydrolyzed by the cytosolic nucleoside hydrolase 1 (NSH1, Figure 6, #26) to uracil 429

and ribose (Jung et al., 2009). NSH2 is not involved in uridine hydrolysis. NSH1 occurs in two 430

forms in vivo, either as a homomer (probably a homodimer: Kopecná et al., 2013) for uridine 431

hydrolysis, and as a heteromer interacting with NSH2 for xanthosine and inosine hydrolysis 432



15

(Baccolini and Witte, 2019). One should note that these nucleoside hydrolases usually do 433

not hydrolyze cytidine (Jung et al., 2009) or guanosine in vivo unless these compounds 434

accumulate in catabolic mutants (Chen et al., 2016; Dahncke and Witte, 2013; Baccolini and 435

Witte, 2019). Adenosine is a substrate of NSH1 and also of 5′-methylthioadenosine 436

nucleosidase 2 (MTAN2; Siu et al., 2008) in vitro, but both enzymes hydrolyze adenosine 437

with very low catalytic efficiency. The main adenosine hydrolytic activity of Arabidopsis 438

resides probably in the apoplast (see below). 439

The plastidic nucleobase transporter (PLUTO, Figure 6, #38) reallocates uracil probably in 440

symport with protons from the cytosol into the plastids for further metabolic conversion 441

(Witz et al., 2012). PLUTO belongs to the Nucleobase:Cation Symporter 1 (NCS1) family and 442

transports guanine and adenine as well, albeit with lower efficiency than uracil. There are 443

indications that a thiamine precursor, hydroxymethylpyrimidine, is also a PLUTO substrate 444

(Beaudoin et al., 2018). Interestingly, it was recently reported that PLUTO orthologs from 445

two grasses do not transport uracil, but only adenine and guanine next to a few other 446

substrates (Rapp et al., 2016), indicating that uracil metabolism might be organized 447

differently in these species. However, one should note that definite evidence for a function 448

of PLUTO in uracil transport into plastids in vivo has not yet been presented in any plant. 449

Other transporters capable of uracil transport have been identified, but these are located in 450

the plasma membrane (Niopek-Witz et al., 2014; Schmidt et al., 2004). 451

In the plastid, there is a branch point: uracil can either be salvaged by UPRT (Figure 6, #39, 452

see above) or be degraded. When uracil is applied from outside, strong catabolic activity is 453

usually observed (Ashihara et al., 2001; Katahira and Ashihara, 2002). The first reaction, in 454

which the uracil ring is reduced to dihydrouracil by dihydropyrimidine dehydrogenase (DPYD 455

/ PYD1, Figure 5, #40) residing in plastids, was shown to be rate limiting (Tintemann et al., 456

1985). Compared to mammalian DPYD, the plant enzyme lacks C-terminal domains for 457

cofactor binding, which are involved in electron delivery to the active site. Therefore the 458

plant enzyme is probably incomplete and might require a so far unknown interaction 459

partner for activity. The loss of activity, when the enzyme is expressed in the cytosol instead 460

of the plastid, is in agreement with this hypothesis (Cornelius et al., 2011). Mutants of DPYD 461

/ PYD1 show delayed germination and a misregulation of ABA responsive genes, whereas 462

constitutive overexpression results in an increase in growth and seed number (Cornelius et 463



16

al., 2011). In the next enzymatic steps, the dihydrouracil ring is opened by 464

dihydropyrimidine hydrolase (DPYH / PYD2, Figure 6, #41) and then the carbamino group is 465

hydrolytically released by -ureidopropionase (-UP / PYD3, Figure 6, #42; Walsh et al., 466

2001) generating -alanine. Not only uracil but also 5-methyluracil (thymine) is degraded in 467

this pathway (Cornelius et al., 2011) resulting in -aminoisobutyrate instead of -alanine. 468

The first reaction (DPYD, Figure 6, #40) is located in the plastids, the second (DPYH, Figure 6, 469

#41) in the ER, and the third (-UP, Figure 6, #42) in the cytosol, but it is unclear why such a 470

distribution is favorable, and how the metabolites are shuttled to these different locations 471

(Zrenner et al., 2009). Recently, the combination of genome-wide association data with 472

correlation networks built from metabolite and transcriptome data identified an 473

aminotransferase correlated with -alanine. The corresponding mutants accumulated -474

alanine, indicating that this might be the missing -alanine aminotransferase (BAAT / PYD4, 475

Figure 6, #43) of pyrimidine catabolism in plants (Wu et al., 2016). 476

Pyrimidine catabolism is induced by nitrogen starvation and in senescence (Cornelius et al., 477

2011; Zrenner et al., 2009) suggesting that similar to purine nitrogen also pyrimidine 478

nitrogen is recycled by plants. When uracil is given as the sole nitrogen source, its 479

degradation can support the growth of Arabidopsis to a limited extent (Zrenner et al., 2009). 480

EXTRACELLUAR ATP 481

Extracellular ATP (eATP) is a signal molecule, which is either actively released upon a 482

stimulus by plant cells via exocytosis or transport, or which is derived from damaged cells 483

(Cao et al., 2014) (Figure 7). A plasma-membrane based nucleotide transporter belonging to 484

the mitochondrial carrier family (pmANT1, Figure 7, #44) is involved in ATP export with 485

physiological relevance at least in pollen (Rieder and Neuhaus, 2011). eATP plays a role in 486

stress responses and is perceived by the receptor-like kinase ‘does not respond to 487

nucleotides 1’ (DORN1, Figure 7, #50), which recognizes ATP, GTP, and ADP but not AMP 488

and adenosine (Choi et al., 2014). Interestingly, CTP and NAD are also sensed by plant cells 489

and a potential receptor for NAD has been identified recently (Wang et al., 2017). 490

The ATP signal might be quenched by an apoplastic apyrase (Riewe et al., 2008a), an enzyme 491

which hydrolyzes NTPs or NDPs to NMPs. Seven apyrases are encoded by the Arabidopsis 492

genome (APY1 to APY7) and APY1 and APY2 were believed to represent these extracellular 493



17

enzymes (Lim et al., 2014). However, by scanning the substrate spectra of the apyrases, only 494

APY3 (Figure 7, #45) showed strong activity with ATP (and other NTPs) but also APY5 and 495

APY6 were slightly active with NTPs (Chiu et al., 2015). These apyrases are therefore 496

possible candidates for the apolastic enzymes in Arabidopsis. However, other secreted 497

phosphatases might also be involved, for example members of the unspecific purple acid 498

phosphatases (Del Vecchio et al., 2014; Wang et al., 2011). 499

The AMP resulting from ATP dephosphorylation is hydrolyzed in the apoplast to adenosine 500

by a 5’ nucleotidase (Figure 7, #46). An AMP-specific extracellular 5’ nucleotidase associated 501

to the plasma membrane was purified from peanut (Arachis hypogaea) (Sharma et al., 1986; 502

Gupta and Sharma, 1996), but the corresponding gene has not been identified. Adenosine 503

can be either taken up via the adenosine proton symporter ‘equilibrative nucleoside 504

transporter 3’ (ENT3, Figure 7, #47) (Cornelius et al., 2012; Traub et al., 2007) or further 505

hydrolyzed by the apoplastic purine-specific nucleoside hydrolase 3 (NSH3, Figure 7, #48) 506

(Jung et al., 2011) to adenine and ribose. NSH3 hydrolyzes inosine more efficiently than 507

adenosine, whereas a cell wall bound nucleoside hydrolase of potato, probably the ortholog 508

of NSH3 in this plant, was highly specific for adenosine and did not hydrolyze inosine (Riewe 509

et al., 2008b). 510

Simultaneous genetic blockage of nucleoside uptake and hydrolysis leads to an 511

accumulation of adenosine and uridine in the apoplast, a reduction of photosystem II 512

efficiency, and a higher susceptibility to the necrotrophic fungus Botrytis cinerea possibly 513

caused by reduced expression of WRKY33 (Daumann et al., 2015), known to be essential for 514

Botrytis resistance (Liu et al., 2015). Treatment with eATP increases the resistance to 515

Botrytis (Tripathi et al., 2017) and the expression of WRKY33 and other defense related 516

genes is reduced in a dorn1 mutant and boosted in a DORN1 overexpression line upon 517

challenge with eATP (Jewell et al., 2019). Taken together it appears that adenosine 518

accumulation in the apoplast dampens the DORN1 mediated response, indicating that ENT3 519

and NSH3 are required to remove the breakdown products of eATP signaling. Also the 520

adenine resulting from adenosine hydrolysis by NSH3 is taken up by plant cells. It is not 521

entirely clear which transporters mediate this uptake. Possible candidates are azaguanine 522

resistant 1 and 2 (AZG1 and 2; Figure 7, #49), which have been shown to facilitate adenine 523

and guanine uptake into Arabidopsis seedlings (Mansfield et al., 2009), or members of the 524



18

nucleobase-ascorbate transporter (NAT) family (Niopek-Witz et al., 2014) as well as of the 525

purine permease (PUP) family (Girke et al., 2014). Interestingly, fungi seem to be able to 526

influence the purinergic signaling in the apoplast by interfering with apoplastic nucleotide 527

metabolism via the excretion of nucleotidases to improve colonization (Nizam et al., 2019). 528

CONNECTIONS TO CYTOKININ HOMEOSTASIS 529

The biosynthesis of the cytokinins involves the generation of N6-modified AMP, carrying an 530

isoprenoid group (Sakakibara, 2005). However, cytokinin ribotides or ribosides are inactive – 531

the free modified base is the active hormone binding to the receptors (Romanov et al., 532

2018; Yamada et al., 2001). The question arises whether the enzymes that are employed for 533

cytokinin homeostasis (activation from ribotides and inactivation to ribosides / ribotides) 534

are the same as for the metabolism of adenine nucleotides. 535

It was shown that a cytokinin ribotide-specific enzyme (cytokinin riboside 5′-536

monophosphate phosphoribohydrolase, called lonely guy (LOG)) can release the active 537

cytokinin from the ribotide (Kurakawa et al., 2007; Kuroha et al., 2009). A recent report 538

demonstrated that the mutation of the seven genes coding for functional LOGs in 539

Arabidopsis resulted in a phenotype that cannot be attenuated by exogenous cytokinin 540

ribotides, suggesting that the hydrolysis by LOGs is the main pathway of cytokinin activation 541

(Osugi et al., 2017). Therefore, the cytosolic nucleoside hydrolases do not seem to be 542

involved in cytokinin activation, although it could be shown that cytokinin ribosides are 543

substrates in vitro, catalyzed with comparatively low efficiency (Kopecná et al., 2013; Jung 544

et al., 2009). Consistently, cytokinin-related phenotypes were not observed in nucleoside 545

hydrolase mutants (Riegler et al., 2011). However, long-distance transport of cytokinins may 546

involve an activation of cytokinin ribosides / ribotides in the apoplast prior to uptake or 547

perception (Romanov et al., 2018). In Arabidopsis, a third nucleoside hydrolase (NSH3) that 548

is located in the apoplast (see section on extracellular ATP) has been shown to hydrolyze 549

adenosine, but cytokinin ribosides have not been assessed (Jung et al., 2011). Interestingly, 550

an apoplastic nucleoside phosphorylase was isolated from potato that converted cytokinin 551

ribosides to cytokinins and ribose-1-phosphate in the presence of phosphate, and can also 552

work in the synthesis direction of ribosides (Bromley et al., 2014). The enzyme preferred 553

cytokinins / cytokinin ribosides over adenine / adenosine as substrates and is supposedly 554

involved in cytokinin-mediated tuber endodormancy. Close homologs in Arabidopsis (e.g. 555



19

At4g28940) are predicted to be located in the apoplast (SUBA web server, Hooper et al., 556

2017), but have not yet been characterized. 557

The inactivation of cytokinins is inter alia achieved by transferring a phosphoribosyl moiety 558

on the active cytokinin resulting in an inactive cytokinin-ribotide. This reaction is at least 559

partially performed by an adenine phosphoribosyltransferase (APRT, Figure 5A, #19) 560

because (i) APT1 mutants convert cytokinins less efficiently to the respective ribotides 561

(Moffatt et al., 1991), (ii) APT1 loss-of-function plants contain more active cytokinins (Zhang 562

et al., 2013), and (iii) APT1 mutants have cytokinin-related phenotypes (Gaillard et al., 563

1998a). APT1 is also clearly involved in the salvage of adenine, which is less efficiently 564

converted to AMP in an APT1 mutant (Moffatt et al., 1991) and is slightly more abundant in 565

a mutant plant with reduced APT1 activity (Sukrong et al., 2012). In conclusion, APT1 566

participates in adenine and cytokinin metabolism and the question arises how one enzyme 567

can serve both distinct metabolic roles adequately. There is also some evidence that 568

adenosine kinase (ADK, Figure 5A, #20) contributes to cytokinin homeostasis, because plants 569

with reduced ADK activity show cytokinin-related phenotypes and contain more cytokinin 570

ribosides (Schoor et al., 2011). However, ADK is also involved in the maintenance of 571

transmethylation activity, therefore an indirect impact of reduced ADK activity on cytokinin 572

homeostasis cannot be fully excluded. 573

Conclusions 574

The synthesis, interconversion, and degradation of nucleotides is intrinsically linked with the 575

propagation and reading of genetic information, with energy metabolism including the 576

metabolic activation of many biomolecules, but also with methylation reactions, signal 577

transduction, the recycling of nitrogen, and the modification of oxidative stress. We are 578

getting closer to completing a full inventory of enzymes involved in plant nucleotide 579

metabolism, but we are far from understanding how these enzymes operate together to 580

achieve nucleotide homeostasis. There is only fragmentary information about regulation on 581

all levels, transport processes are incompletely defined, and the organization of nucleotide 582

metabolism on tissue and organ level is not well understood (see outstanding questions). 583

These issues need to be addressed to allow a better integration of nucleotide metabolism 584

into a molecular model of plant physiology. 585



20

586

Acknowledgements 587

The authors thank Henryk Straube for critical reading of the manuscript. 588

589

590

Figure Legends 591

Figure 1. Structural composition of nucleobases, nucleosides, and nucleotides. 592

For the nucleobases ‘R’ is simply a proton. For the nucleosides ‘R’ is a sugar moiety which 593

can be ribose or deoxyribose (carrying a proton instead of a hydroxyl group at the 2’ carbon 594

of the ribose). Nucleotides have up to three phosphate groups esterified to the hydroxyl 595

group of the 5’ carbon of the nucleoside sugar determining the prefix mono-, di-, and 596

triphosphate in the name of the molecule. The terminal phosphate always carries two 597

charges irrespective of the number of phosphates present. The pyrimidine nucleobases 598

(upper row) and the purine nucleobases (lower row) are shown with the groups attached to 599

the heterocycles highlighted in red (oxo groups), blue (amino groups), and grey shading 600

(methyl group). 601

602

Figure 2. Schematic overview of plant nucleotide metabolism. 603

Nucleotides are synthesized ‘de novo’ from precursor molecules listed in the upper left box 604

(PRPP, 5-phosphoribosyl-1-pyrophosphate). The phosphorylation of nucleoside 605

monophosphates via nucleoside diphosphates (NDPs) generates nucleoside triphosphates 606

(NTPs), which serve as building blocks for RNA synthesis and as precursors for the 607

biosynthesis of the metabolites shown in the center (S-adenosyl methionine (SAM), UDP-608

glucose (UDP-Glc), and NADH are given as examples). However, the nucleoside 609

triphosphates, in particular ATP and GTP, are not only precursors for other metabolites, but 610

are also essential stores of chemical energy in the phosphoanhydride bonds used in a 611

multitude of energetic coupling reactions, as well as important donors of phosphate in 612

kinase reactions (not shown). NDPs can be reduced to dNDPs (deoxynucleoside 613

diphosphates), which after phosphorylation to dNTPs serve as precursors for DNA 614

biosynthesis. RNA degradation in the cytosol releases nucleoside monophosphates, whereas 615

nucleosides are produced during vacuolar RNA degradation. Adenosine and adenine are 616

products of biochemical reactions involving S-adenosyl methionine (SAM). Non-enzymatic 617

decay (depurination) and enzymatic repair reactions result in nucleoside and nucleobase 618

release from DNA. Nucleobases and nucleosides can be recycled to nucleotides in so called 619

‘salvage’ reactions. Plants are also capable of full nucleotide degradation via certain 620

nucleosides and nucleobases releasing the nitrogen of the nucleobases as ammonia. 621

Figure 3. Purine and pyrimidine de novo biosynthesis. 622



21

(A) Purine de novo biosynthesis. (B) Pyrimidine de novo biosynthesis. Enzymes and 623

transporters: BT1 (1), brittle1; AMPD (2), AMP deaminase; IMPDH (3), IMP dehydrogenase; 624

GMPS (4), GMP synthetase; ATCase (5), asparate transcarbamoylase; DHOase (6), 625

dihydroorotatase; DHODH (7), dihydroorotate dehydrogenase; UMPS (8), UMP synthase; 626

CTPS (9), CTP synthetase. An anchor symbol denotes an association with the respective 627

membrane. 628

629

Figure 4. Synthesis of nucleoside and deoxynucleoside triphosphates. 630

Synthesis of (A) cytidylates, (B) uridylates and thymidylates, (C) guanylates, and (D) 631

adenylates. RNR (10), ribonucleotide reductase; dNK (11), deoxynucleoside kinase; UMK 632

(12), UMP kinase; NDPK (13), nucleoside diphosphate kinase; TK (14), thymidine kinase; 633

DHFR-TS (15), dihydrofolate reductase-thymidylate synthase; TMK (16), thymidylate kinase; 634

GMK (17), guanylate kinase; AMK (18), adenylate kinase. The subcellular locations where 635

enzymes with these activities are found are indicated. For TMK, a location in the plastids is 636

only assumed. The mononucleotides (AMP, GMP, UMP, and CMP) may also be derived from 637

salvage reactions (see Figures 5 and 6). 638

639

Figure 5. Salvage and degradation of purines. 640

(A) The reactions of purine nucleobase and nucleoside salvage as well as purine nucleotide 641

degradation, which overlaps partially with GMP synthesis. The salvage pathways are 642

highlighted by light grey shading, degradation reactions are encircled in dark gray. 643

Metabolites that can only undergo degradation and cannot be salvaged are shown with 644

brown shading. (B) Purine ring catabolism. The transport steps for urate and (S)-allantoin 645

are not shown explicitly. APRT (19), adenine phosphoribosyltransferase; ADK (20), 646

adenosine kinase; AMPD (2), AMP deaminase; IMPDH (3), IMP dehydrogenase; GMPS (4), 647

GMP synthetase; HGPRT (21), hypoxanthine guanine phosphoribosyltransferase; IGK (22), 648

inosine guanosine kinase; GMPP (23), GMP phosphatase; GSDA (24), guanosine deaminase; 649

XMPP (25), XMP phosphatase; NSH1 (26), nucleoside hydrolase 1; NSH2 (27), nucleoside 650

hydrolase 2; XDH (28), xanthine dehydrogenase; UOX (29), urate oxidase; ALNS (30), 651

allantoin synthase; ALN (31), allantoinase; AAH (32), allantoate amidohydrolase; UGAH (33), 652

ureidoglycine aminohydrolase; UAH (34), ureidoglycolate amidohydrolase. 653

654

Figure 6. Salvage and degradation of pyrimidines. 655

NC--alanine, N-carbamyl--alanine; malonate-SA, malonate semialdehyde. UCK (35), 656

uridine cytidine kinase; UCPP (36), UMP CMP phosphatase; CDA (37), cytidine deaminase; 657

NSH1 (26), nucleoside hydrolase 1; PLUTO (38), plastidic nucleobase transporter; UPRT (39), 658

uracil phosphoribosyltransferase; DPYD (40), dihydropyrimidine dehydrogenase; DPYH (41), 659

dihydropyrimidine hydrolase; β-UP (42), -ureidopropionase; BAAT (43) -alanine 660

aminotransferase. 661



22

662

Figure 7. Excretion, perception, and degradation of extracellular ATP 663

pmANT1 (44), plasma membrane adenine nucleotide transporter; APY3 (45), apyrase 3; 5‘ 664

NT (46), 5’ nucleotidase; ENT3 (47), equilibrative nucleoside transporter 3; NSH3 (48), 665

nucleoside hydrolase 3, AZG1/2 (49), azaguanine resistant 1 / azaguanine resistant 2, 666

DORN1 (50), does not respond to nucleotides 1 . 667

668

Literature cited 669

Alamillo JM, Diaz-Leal JL, Sanchez-Moran MV, Pineda M (2010) Molecular analysis of 670

ureide accumulation under drought stress in Phaseolus vulgaris L. Plant Cell Environ 33: 671

1828–1837. 672

Ashihara H, Loukanina N, Stasolla C, Thorpe TA (2001) Pyrimidine metabolism during 673

somatic embryo development in white spruce (Picea glauca). J Plant Physiol 158: 613–674

621. 675

Ashihara H, Stasolla C, Fujimura T, Crozier A (2018) Purine salvage in plants. 676

Phytochemistry 147: 89–124. 677

Atkins CA, Shelp BJ, Storer PJ (1985) Purification and properties of inosine monophosphate 678

oxidoreductase from nitrogen-fixing nodules of cowpea (Vigna-Unguiculata-L WALP). 679

Arch Biochem Biophys 236: 807–814. 680

Atkins CA, Smith, P. M. C., Storer PJ (1997) Reexamination of the intracellular localization of 681

de novo purine synthesis in cowpea nodules. Plant Physiol 113: 127–135. 682

Baccolini C, Witte C-P (2019) AMP and GMP catabolism in Arabidopsis converge on 683

xanthosine, which is degraded by a nucleoside hydrolase heterocomplex. Plant Cell 31: 684

734–751. 685

Bahaji A, Muñoz FJ, Ovecka M, Baroja-Fernández E, Montero M, Li J, Hidalgo M, Almagro 686

G, Sesma MT, Ezquer I, Pozueta-Romero J (2011a) Specific delivery of AtBT1 to 687

mitochondria complements the aberrant growth and sterility phenotype of homozygous 688

Atbt1 Arabidopsis mutants. Plant J 68: 1115–1121. 689

Bahaji A, Ovecka M, Bárány I, Risueño MC, Muñoz FJ, Baroja-Fernández E, Montero M, Li J, 690

Hidalgo M, Sesma MT, Ezquer I, Testillano PS, Pozueta-Romero J (2011b) Dual targeting 691

to mitochondria and plastids of AtBT1 and ZmBT1, two members of the mitochondrial 692

carrier family. Plant Cell Physiol 52: 597–609. 693

Barbado C, Cordoba-Canero D, Arizaa RR, Roldan-Arjona T (2018) Nonenzymatic release of 694

N7-methylguanine channels repair of abasic sites into an AP endonuclease-independent 695

pathway in Arabidopsis. Proc Natl Acad Sci USA 115: E916-E924. 696



23

Beaudoin GAW, Johnson TS, Hanson AD (2018) The PLUTO plastidial nucleobase 697

transporter also transports the thiamin precursor hydroxymethylpyrimidine. Biosci Rep 698

38. 699

Bernard C, Traub M, Kunz HH, Hach S, Trentmann O, Mohlmann T (2011) Equilibrative 700

nucleoside transporter 1 (ENT1) is critical for pollen germination and vegetative growth 701

in Arabidopsis. J Exp Bot 62: 4627–4637. 702

Bromley JR, Warnes BJ, Newell CA, Thomson JCP, James CM, Turnbull CGN, Hanke DE 703

(2014) A purine nucleoside phosphorylase in Solanum tuberosum L. (potato) with 704

specificity for cytokinins contributes to the duration of tuber endodormancy. Biochem J 705

458: 225–237. 706

Brychkova G, Alikulov Z, Fiuhr R, Sagi M (2008) A critical role for ureides in dark and 707

senescence-induced purine remobilization is unmasked in the Atxdh1 Arabidopsis 708

mutant. Plant J 54: 496–509. 709

Cao Y, Tanaka K, Nguyen CT, Stacey G (2014) Extracellular ATP is a central signaling 710

molecule in plant stress responses. Curr Opin Plant Biol 20: 82–87. 711

Carrari F, Coll-Garcia D, Schauer N, Lytovchenko A, Palacios-Rojas N, Balbo I, Rosso M, 712

Fernie AR (2005) Deficiency of a plastidial adenylate kinase in Arabidopsis results in 713

elevated photosynthetic amino acid biosynthesis and enhanced growth. Plant Physiol 714

137: 70–82. 715

Carter AM, Tegeder M (2016) Increasing nitrogen fixation and seed development in soybean 716

requires complex adjustments of nodule nitrogen metabolism and partitioning processes. 717

Current Biol 26: 2044–2051. 718

Casartelli A, Melino VJ, Baumann U, Riboni M, Suchecki R, Jayasinghe NS, Mendis H, 719

Watanabe M, Erban A, Zuther E, Hoefgen R, Roessner U, Okamoto M, Heuer S (2019) 720

Opposite fates of the purine metabolite allantoin under water and nitrogen limitations in 721

bread wheat. Plant Mol Biol 99: 477–497. 722

Chen F, Dong G, Ma X, Wang F, Zhang Y, Xiong E, Wu J, Wang H, Qian Q, Wu L, Yu Y (2018a) 723

UMP kinase activity is involved in proper chloroplast development in rice. Photosynth Res 724

137: 53–67. 725

Chen M, Herde M, Witte C-P (2016) Of the nine cytidine deaminase-like genes in 726

Arabidopsis, eight are pseudogenes and only one is required to maintain pyrimidine 727

homeostasis in vivo. Plant Physiol 171: 799–809. 728

Chen M, Urs MJ, Sánchez-González I, Olayioye MA, Herde M, Witte C-P (2018b) m6A RNA 729

degradation products are catabolized by an evolutionarily conserved N6-Methyl-AMP 730

deaminase in plant and mammalian cells. Plant Cell 30: 1511–1522. 731

Chen M, Witte C-P (2019) Functions and Dynamics of Methylation in Eukaryotic mRNA. In S 732

Jurga, J Barciszewski, eds, The DNA, RNA, and Histone Methylomes. Springer 733

International Publishing, Cham, pp. 333–351.Chen MJ, Thelen JJ (2011) Plastid uridine 734



24

salvage activity is required for photoassimilate allocation and partitioning in Arabidopsis. 735

Plant Cell 23: 2991–3006. 736

Chen X, Zhu L, Xin L, Du K, Ran X, Cui X, Xiang Q, Zhang H, Xu P, Wu X (2015) Rice stripe1-2 737

and stripe1-3 mutants encoding the small subunit of ribonucleotide reductase are 738

temperature sensitive and are required for chlorophyll biosynthesis. PloS One 10: 739

e0130172. 740

Chiu T-Y, Lao J, Manalansan B, Loqué D, Roux SJ, Heazlewood JL (2015) Biochemical 741

characterization of Arabidopsis apyrase family reveals their roles in regulating 742

endomembrane NDP/NMP homoeostasis. Biochem J 472: 43–54. 743

Choi J, Tanaka K, Cao Y, Qi Y, Qiu J, Liang Y, Lee SY, Stacey G (2014) Identification of a plant 744

receptor for extracellular ATP. Science 343: 290–294. 745

Clausen AR, Girandon L, Ali A, Knecht W, Rozpedowska E, Sandrini MP, Andreasson E, 746

Munch-Petersen B, Piskur J (2012) Two thymidine kinases and one multisubstrate 747

deoxyribonucleoside kinase salvage DNA precursors in Arabidopsis thaliana. FEBS J 279: 748

3889–3897. 749

Clausen AR, Mutahir Z, Munch-Petersen B, Piškur J (2014) Plants salvage 750

deoxyribonucleosides in mitochondria. Nucleosides, Nucleotides Nucleic Acids 33: 291–751

295. 752

Collier R, Tegeder M (2012) Soybean ureide transporters play a critical role in nodule 753

development, function and nitrogen export. Plant J 72: 355–367. 754

Combes A, Lafleuriel J, Lefloch F (1989) The Inosine-Guanosine Kinase-Activity of 755

Mitochondria in Tubers of Jerusalem Artichoke. Plant Physiol Biochem 27: 729–736. 756

Cornelius S, Traub M, Bernard C, Salzig C, Lang P, Mohlmann T (2012) Nucleoside transport 757

across the plasma membrane mediated by equilibrative nucleoside transporter 3 758

influences metabolism of Arabidopsis seedlings. Plant Biol 14: 696–705. 759

Cornelius S, Witz S, Rolletschek H, Mohlmann T (2011) Pyrimidine degradation influences 760

germination seedling growth and production of Arabidopsis seeds. J Exp Bot 62: 5623–761

5632. 762

Dahncke K, Witte CP (2013) Plant purine nucleoside catabolism employs a guanosine 763

deaminase required for the generation of xanthosine in Arabidopsis. Plant Cell 25: 4101–764

4109. 765

Dancer JE, Hughes RG, Lindell SD (1997) Adenosine-5’-phosphate deaminase. A novel 766

herbicide target. Plant Physiol 114: 119–129. 767

Daumann M, Fischer M, Niopek-Witz S, Girke C, Möhlmann T (2015) Apoplastic nucleoside 768

accumulation in Arabidopsis leads to reduced photosynthetic performance and increased 769

susceptibility against Botrytis cinerea. Front Plant Sci 6: 1158. 770



25

Daumann M, Hickl D, Zimmer D, DeTar RA, Kunz H-H, Möhlmann T (2018) Characterization 771

of filament-forming CTP synthases from Arabidopsis thaliana. Plant J: 96: 316–328. 772

Del Vecchio HA, Ying S, Park J, Knowles VL, Kanno S, Tanoi K, She Y-M, Plaxton WC (2014) 773

The cell wall-targeted purple acid phosphatase AtPAP25 is critical for acclimation of 774

Arabidopsis thaliana to nutritional phosphorus deprivation. Plant J 80: 569–581. 775

Deng WW, Ashihara H (2010) Profiles of purine metabolism in leaves and roots of Camellia 776

sinensis seedlings. Plant Cell Physiol 51: 2105–2118. 777

Desimone M, Catoni E, Ludewig U, Hilpert M, Schneider A, Kunze R, Tegeder M, Frommer 778

WB, Schumacher K (2002) A novel superfamily of transporters for allantoin and other oxo 779

derivatives of nitrogen heterocyclic compounds in Arabidopsis. Plant Cell 14: 847–856. 780

Diaz-Leal JL, Galvez-Valdivieso G, Fernandez J, Pineda M, Alamillo JM (2012) 781

Developmental effects on ureide levels are mediated by tissue-specific regulation of 782

allantoinase in Phaseolus vulgaris L. J Exp Bot 63: 4095–4106. 783

Díaz-Leal JL, Torralbo F, Antonio Quiles F, Pineda M, Alamillo JM (2014) Molecular and 784

functional characterization of allantoate amidohydrolase from Phaseolus vulgaris. Physiol 785

Plant 152: 43–58. 786

Dong Q, Zhang Y-X, Zhou Q, Liu Q-E, Chen D-B, Wang H, Cheng S-H, Cao L-Y, Shen X-H 787

(2019) UMP Kinase Regulates Chloroplast Development and Cold Response in Rice. Int J 788

Mol Sci 20: 2107. 789

Doremus HD, Jagendorf AT (1985) Subcellular localization of the pathway of de novo 790

pyrimidine nucleotide biosynthesis in pea leaves. Plant Physiol 79: 856–861. 791

Dorion S, Rivoal J (2015) Clues to the functions of plant NDPK isoforms. Naunyn-792

Schmiedeberg’s Arch Pharmacol 388: 119–132. 793

Dorion S, Rivoal J (2018) Plant nucleoside diphosphate kinase 1: A housekeeping enzyme 794

with moonlighting activity. Plant Signal Behav 13: e1475804. 795

Feng X, Yang R, Zheng X, Zhang F (2012) Identification of a novel nuclear-localized adenylate 796

kinase 6 from Arabidopsis thaliana as an essential stem growth factor. Plant Physiol 797

Biochem 61: 180–186. 798

Floyd BE, Morriss SC, MacIntosh GC, Bassham DC (2015) Evidence for autophagy-799

dependent pathways of rRNA turnover in Arabidopsis. Autophagy 11: 2199–2212. 800

Gaillard C, Moffatt BA, Blacker M, Laloue M (1998a) Male sterility associated with APRT 801

deficiency in Arabidopsis thaliana results from a mutation in the gene APT1. Mol Gen 802

Genet 257: 348–353. 803

Gaillard C, Moffatt BA, Blacker M, Laloue M (1998b) Male sterility associated with APRT 804

deficiency in Arabidopsis thaliana results from a mutation in the gene APT1. Mol Gen 805

Genet 257: 348–353. 806



26

Garton S, Knight H, Warren GJ, Knight MR, Thorlby GJ (2007) Crinkled leaves 8 - a mutation 807

in the large subunit of ribonucleotide reductase - leads to defects in leaf development 808

and chloroplast division in Arabidopsis thaliana. Plant J 50: 118–127. 809

Girke C, Daumann M, Niopek-Witz S, Möhlmann T (2014) Nucleobase and nucleoside 810

transport and integration into plant metabolism. Front Plant Sci 5: 443. 811

Gorelova V, Lepeleire J de, van Daele J, Pluim D, Meï C, Cuypers A, Leroux O, Rébeillé F, 812

Schellens JHM, Blancquaert D, Stove CP, van der Straeten D (2017) Dihydrofolate 813

reductase/thymidylate synthase fine-tunes the folate status and controls redox 814

homeostasis in plants. Plant cell 29: 2831–2853. 815

Gupta A, Sharma CB (1996) Purification to homogeneity and characterization of plasma 816

membrane and Golgi apparatus-specific 5’-adenosine monophosphatases from peanut 817

cotyledons. Plant Sci 117: 65–74. 818

Han BW, Bingman CA, Mahnke DK, Bannen RM, Bednarek SY, Sabina RL, Phillips GN (2006) 819

Membrane association, mechanism of action, and structure of Arabidopsis embryonic 820

factor 1 (FAC1). J Biol Chem 81: 14939–14947. 821

Hauck OK, Scharnberg J, Escobar NM, Wanner G, Giavalisco P, Witte C-P 2014) Uric acid 822

accumulation in an Arabidopsis urate oxidase mutant impairs seedling establishment by 823

blocking peroxisome maintenance. Plant Cell 26: 3090–3100. 824

Hooper CM, Castleden IR, Tanz SK, Aryamanesh N, Millar AH (2017) SUBA4: the interactive 825

data analysis centre for Arabidopsis subcellular protein locations. Nucleic Acids Res 45: 826

D1064-D1074. 827

Hu D, Li Y, Jin W, Gong H, He Q, Li Y (2017) Identification and characterization of a plastidic 828

adenine nucleotide uniporter (OsBT1-3) required for chloroplast development in the 829

early leaf stage of rice. Sci Rep 7. 830

Irani S, Todd CD (2016) Ureide metabolism under abiotic stress in Arabidopsis thaliana. J 831

Plant Physiol 199: 87–95. 832

Irani S, Todd CD (2018) Exogenous allantoin increases Arabidopsis seedlings tolerance to 833

NaCl stress and regulates expression of oxidative stress response genes. J Plant Physiol 834

221: 43–50. 835

Ito J, Batth TS, Petzold CJ, Redding-Johanson AM, Mukhopadhyay A, Verboom R, Meyer 836

EH, Millar AH, Heazlewood JL (2011) Analysis of the Arabidopsis cytosolic proteome 837

highlights subcellular partitioning of central plant metabolism. J Proteome Res 10: 1571-838

1582. 839

Jewell JB, Sowders JM, He R, Willis MA, Gang DR, Tanaka K (2019) Extracellular ATP Shapes 840

a Defense-Related Transcriptome Both Independently and along with Other Defense 841

Signaling Pathways. Plant Physiol 179: 1144–1158. 842


https://www.ncbi.nlm.nih.gov/pubmed/?term=Ito%20J%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Batth%20TS%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Petzold%20CJ%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Redding-Johanson%20AM%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Mukhopadhyay%20A%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Verboom%20R%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Meyer%20EH%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Meyer%20EH%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Millar%20AH%5BAuthor%5D&cauthor=true&cauthor_uid=21166475

https://www.ncbi.nlm.nih.gov/pubmed/?term=Heazlewood%20JL%5BAuthor%5D&cauthor=true&cauthor_uid=21166475


27

Jung B, Florchinger M, Kunz HH, Traub M, Wartenberg R, Jeblick W, Neuhaus HE, 843

Mohlmann T (2009) Uridine-ribohydrolase is a key regulator in the uridine degradation 844

pathway of Arabidopsis. Plant Cell 21: 876–891. 845

Jung B, Hoffmann C, Mohlmann T (2011) Arabidopsis nucleoside hydrolases involved in 846

intracellular and extracellular degradation of purines. Plant J 65: 703–711. 847

Kafer C, Zhou L, Santoso D, Guirgis A, Weers B, Park S, Thornburg R (2004) Regulation of 848

pyrimidine metabolism in plants. Front Biosci-Landmrk 9: 1611–1625. 849

Karran P, Lindahl T (1980) Hypoxanthine in deoxyribonucleic acid: generation by heat-850

induced hydrolysis of adenine residues and release in free form by a deoxyribonucleic 851

acid glycosylase from calf thymus. Biochem 19: 6005–6011. 852

Katahira R, Ashihara H (2002) Profiles of pyrimidine biosynthesis, salvage and degradation 853

in disks of potato (Solanum tuberosum L.) tubers. Planta 215: 821–828. 854

Katahira R, Ashihara H (2006) Profiles of purine biosynthesis, salvage and degradation in 855

disks of potato (Solanum tuberosum L.) tubers. Planta 225: 115–126. 856

Kihara A, Saburi W, Wakuta S, Kim M-H, Hamada S, Ito H, Imai R, Matsui H (2011) 857

Physiological and biochemical characterization of three nucleoside diphosphate kinase 858

isozymes from rice (Oryza sativa L.). Biosci Biotechnol Biochem 75: 1740–1745. 859

Kirchberger S, Tjaden J, Neuhaus HE (2008) Characterization of the Arabidopsis Brittle1 860

transport protein and impact of reduced activity on plant metabolism. Plant J 56: 51–63. 861

Kopecná M, Blaschke H, Kopecny D, Vigouroux A, Koncitíková R, Novák O, Kotland O, 862

Strnad M, Moréra S, Schwartzenberg K von (2013) Structure and function of nucleoside 863

hydrolases from Physcomitrella patens and maize catalyzing the hydrolysis of purine, 864

pyrimidine, and cytokinin ribosides. Plant Physiol 163: 1568–1583. 865

Kumar V, Spangenberg O, Konrad M (2000) Cloning of the guanylate kinase homologues 866

AGK-1 and AGK-2 from Arabidopsis thaliana and characterization of AGK-1. Eur J Biochem 867

267: 606–615. 868

Kurakawa T, Ueda N, Maekawa M, Kobayashi K, Kojima M, Nagato Y, Sakakibara H, 869

Kyozuka J (2007) Direct control of shoot meristem activity by a cytokinin-activating 870

enzyme. Nature 445: 652 EP -. 871

Kuroha T, Tokunaga H, Kojima M, Ueda N, Ishida T, Nagawa S, Fukuda H, Sugimoto K, 872

Sakakibara H (2009) Functional analyses of LONELY GUY cytokinin-activating enzymes 873

reveal the importance of the direct activation pathway in Arabidopsis. Plant Cell 21: 874

3152–3169. 875

Lamberto I, Percudani R, Gatti R, Folli C, Petrucco S (2010) Conserved alternative splicing of 876

Arabidopsis transthyretin-like determines protein localization and S-allantoin synthesis in 877

peroxisomes. Plant Cell 22: 1564–1574. 878



28

Lange PR, Geserick C, Tischendorf G, Zrenner R (2008) Functions of chloroplastic adenylate 879

kinases in Arabidopsis. Plant Physiol 146: 492–504. 880

Lee S, Doxey AC, McConkey BJ, Moffatt BA (2012) Nuclear targeting of methyl-recycling 881

enzymes in Arabidopsis thaliana is mediated by specific protein interactions. Mol Plant 5: 882

231–248. 883

Leroch M, Kirchberger S, Haferkamp I, Wahl M, Neuhaus HE, Tjaden J (2005) Identification 884

and characterization of a novel plastidic adenine nucleotide uniporter from Solanum 885

tuberosum. J Biol Chem 280: 17992–18000. 886

Lescano CI, Martini C, González CA, Desimone M (2016) Allantoin accumulation mediated 887

by allantoinase downregulation and transport by ureide permease 5 confers salt stress 888

tolerance to Arabidopsis plants. Plant Mol Biol 91: 581-595 889

Lim MH, Wu J, Yao J, Gallardo IF, Dugger JW, Webb LJ, Huang J, Salmi ML, Song J, Clark G, 890

Roux SJ (2014) Apyrase suppression raises extracellular ATP levels and induces gene 891

expression and cell wall changes characteristic of stress responses. Plant Physiol 164: 892

2054–2067. 893

Lincker F, Philipps G, Chabouté M-E (2004) UV-C response of the ribonucleotide reductase 894

large subunit involves both E2F-mediated gene transcriptional regulation and protein 895

subcellular relocalization in tobacco cells. Nucleic Acids Res 32: 1430–1438. 896

Liu S, Kracher B, Ziegler J, Birkenbihl RP, Somssich IE (2015) Negative regulation of ABA 897

signaling by WRKY33 is critical for Arabidopsis immunity towards Botrytis cinerea. eLife 4: 898

e07295. 899

Liu XY, Qian WQ, Liu X, Qin HJ, Wang DW (2007) Molecular and functional analysis of 900

hypoxanthine-guanine phosphoribosyltransferase from Arabidopsis thaliana. New Phytol 901

175: 448–461. 902

Luzarowski M, Kosmacz M, Sokolowska E, Jasinska W, Willmitzer L, Veyel D, Skirycz A 903

(2017) Affinity purification with metabolomic and proteomic analysis unravels diverse 904

roles of nucleoside diphosphate kinases. J Exp Bot 68: 3487–3499. 905

Ma X, Wang W, Bittner F, Schmidt N, Berkey R, Zhang L, King H, Zhang Y, Feng J, Wen Y, 906

Tan L, Li Y, Zhang Q, Deng Z, Xiong X, Xiao S (2016) Dual and opposing roles of xanthine 907

dehydrogenase in defense-associated reactive oxygen species metabolism in Arabidopsis. 908

Plant Cell 28: 1108–1126. 909

Mainguet SE, Gakiere B, Majira A, Pelletier S, Bringel F, Guerard F, Caboche M, Berthome 910

R, Renou JP (2009) Uracil salvage is necessary for early Arabidopsis development. Plant J 911

60: 280–291. 912

Mansfield TA, Schultes NP, Mourad GS (2009) AtAzg1 and AtAzg2 comprise a novel family 913

of purine transporters in Arabidopsis. FEBS Lett 583: 481–486. 914

Moffatt B, Ashihara H (2002) Purine and pyrimdine nucleotide synthesis and metabolism, 915

The Arabidopsis book. American Society of Plant Biologists, Rockville, MD. 916



29

Moffatt B, Pethe C, Laloue M (1991) Metabolism of benzyladenine is impaired in a mutant 917

of Arabidopsis thaliana lacking adenine phosphoribosyltransferase activity. Plant Physiol 918

95: 900–908. 919

Moffatt BA, Stevens YY, Allen MS, Snider JD, Pereira LA, Todorova MI, Summers PS, 920

Weretilnyk EA, Martin-McCaffrey L, Wagner C (2002) Adenosine kinase deficiency is 921

associated with developmental abnormalities and reduced transmethylation. Plant 922

Physiol 128: 812–821. 923

Niopek-Witz S, Deppe J, Lemieux MJ, Möhlmann T (2014) Biochemical characterization and 924

structure-function relationship of two plant NCS2 proteins, the nucleobase transporters 925

NAT3 and NAT12 from Arabidopsis thaliana. Biochim Biophys Acta 1838: 3025–3035. 926

Niu M, Wang Y, Wang C, Lyu J, Wang Y, Dong H, Long W, Di Wang, Kong W, Wang L, Guo X, 927

Sun L, Hu T, Zhai H, Wang H, Wan J (2017) ALR encoding dCMP deaminase is critical for 928

DNA damage repair, cell cycle progression and plant development in rice. J Exp Bot 68: 929

5773–5786. 930

Nizam S, Qiang X, Wawra S, Nostadt R, Getzke F, Schwanke F, Dreyer I, Langen G, Zuccaro 931

A (2019) Serendipita indica E5′NT modulates extracellular nucleotide levels in the plant 932

apoplast and affects fungal colonization. EMBO Rep 20: e47430. 933

Nomura Y, Izumi A, Fukunaga Y, Kusumi K, Iba K, Watanabe S, Nakahira Y, Weber APM, 934

Nozawa A, Tozawa Y (2014) Diversity in guanosine 3’,5’-bisdiphosphate (ppGpp) 935

sensitivity among guanylate kinases of bacteria and plants. J Biol Chem 289: 15631–936

15641. 937

Nourimand M, Todd CD (2019) There is a direct link between allantoin concentration and 938

cadmium tolerance in Arabidopsis. Plant Physiol Biochem 135: 441–449. 939

Ohler L, Niopek-Witz S, Mainguet SE, Möhlmann T (2019) Pyrimidine salvage: physiological 940

functions and interaction with chloroplast biogenesis. Plant Physiol 180: 1816–1828 941

Osugi A, Kojima M, Takebayashi Y, Ueda N, Kiba T, Sakakibara H (2017) Systemic transport 942

of trans-zeatin and its precursor have differing roles in Arabidopsis shoots. Nat Plants 3: 943

17112. 944

Pedroza-García J-A, Nájera-Martínez M, Mazubert C, Aguilera-Alvarado P, Drouin-Wahbi J, 945

Sánchez-Nieto S, Gualberto JM, Raynaud C, Plasencia J (2019) Role of pyrimidine salvage 946

pathway in the maintenance of organellar and nuclear genome integrity. Plant J 97: 430–947

446. 948

Pessoa J, Sarkany Z, Ferreira-da-Silva F, Martins S, Almeida MR, Li JM, Damas AM (2010) 949

Functional characterization of Arabidopsis thaliana transthyretin-like protein. BMC Plant 950

Biol 10: 30. 951

Phillips DA, Joseph CM, Hirsch PR (1997) Occurrence of flavonoids and nucleosides in 952

agricultural soils. Appl Environ Microbiol 63: 4573–4577. 953



30

Quiles FA, Galvez-Valdivieso G, Guerrero-Casado J, Pineda M, Piedras P (2019) Relationship 954

between ureidic/amidic metabolism and antioxidant enzymatic activities in legume 955

seedlings. Plant Physiol Biochem 138: 1–8. 956

Rapp M, Schein J, Hunt KA, Nalam V, Mourad GS, Schultes NP (2016) The solute specificity 957

profiles of nucleobase cation symporter 1 (NCS1) from Zea mays and Setaria viridis 958

illustrate functional flexibility. Protoplasma 253: 611–623. 959

Redillas MCFR, Bang SW, Lee D‐K, Kim YS, Jung H, Chung PJ, Suh J‐W, Kim J‐K (2019) 960

Allantoin accumulation through overexpression of ureide permease1 improves rice 961

growth under limited nitrogen conditions. Plant Biotechnol J 17: 1289–1301. 962

Regierer B, Fernie AR, Springer F, Perez-Melis A, Leisse A, Koehl K, Willmitzer L, 963

Geigenberger P, Kossmann J (2002) Starch content and yield increase as a result of 964

altering adenylate pools in transgenic plants. Nat Biotechnol 20: 1256–1260. 965

Rieder B, Neuhaus HE (2011) Identification of an Arabidopsis plasma membrane-located 966

ATP transporter important for anther development. Plant Cell 23: 1932–1944. 967

Riegler H, Geserick C, Zrenner R (2011) Arabidopsis thaliana nucleosidase mutants provide 968

new insights into nucleoside degradation. New Phytol 191: 349–359. 969

Riewe D, Grosman L, Fernie AR, Wucke C, Geigenberger P (2008a) The potato-specific 970

apyrase is apoplastically localized and has influence on gene expression, growth, and 971

development. Plant Physiol 147: 1092–1109. 972

Riewe D, Grosman L, Fernie AR, Zauber H, Wucke C, Geigenberger P (2008b) A cell wall-973

bound adenosine nucleosidase is involved in the salvage of extracellular ATP in Solanum 974

tuberosum. Plant Cell Physiol 49: 1572–1579. 975

Riggs JW, Rockwell NC, Cavales PC, Callis J (2016) Identification of the plant ribokinase and 976

discovery of a role for Arabidopsis ribokinase in nucleoside metabolism. J Biol Chem 291: 977

22572–22582. 978

Romanov GA, Lomin SN, Schmülling T (2018) Cytokinin signaling: from the ER or from the 979

PM? That is the question! New Phytol 218: 41–53. 980

Ronceret A, Gadea-Vacas J, Guilleminot J, Lincker F, Delorme V, Lahmy S, Pelletier G, 981

Chabouté M-E, Devic M (2008) The first zygotic division in Arabidopsis requires de novo 982

transcription of thymidylate kinase. Plant J 53: 776–789. 983

Sabina RL, Paul AL, Ferl RJ, Laber B, Lindell SD (2007) Adenine nucleotide pool perturbation 984

is a metabolic trigger for AMP deaminase inhibitor-based herbicide toxicity. Plant Physiol 985

143: 1752–1760. 986

Sakakibara H (2005) Cytokinin biosynthesis and regulation. Vitam Horm 72: 271–287. 987

Sauge-Merle S, Falconet D, Fontecave M (1999) An active ribonucleotide reductase from 988

Arabidopsis thaliana - Cloning, expression and characterization of the large subunit. Eur J 989

Biochem 266: 62–69. 990



31

Sauter M, Moffatt B, Saechao MC, Hell R, Wirtz M (2013) Methionine salvage and S-991

adenosylmethionine: essential links between sulfur, ethylene and polyamine 992

biosynthesis. Biochem J 451: 145–154. 993

Schmid L-M, Ohler L, Möhlmann T, Brachmann A, Muiño JM, Leister D, Meurer J, Manavski 994

N (2019) PUMPKIN, the sole plastid UMP kinase, associates with group II introns and 995

alters their metabolism. Plant Physiol 179: 248–264. 996

Schmidt A, Baumann N, Schwarzkopf A, Frommer WB, Desimone M (2006) Comparative 997

studies on ureide permeases in Arabidopsis thaliana and analysis of two alternative splice 998

variants of AtUPS5. Planta 224: 1329–1340. 999

Schmidt A, Su YH, Kunze R, Warner S, Hewitt M, Slocum RD, Ludewig U, Frommer WB, 1000

Desimone M (2004) UPS1 and UPS2 from Arabidopsis mediate high affinity transport of 1001

uracil and 5-fluorouracil. J Biol Chem 279: 44817–44824. 1002

Schmülling T, Werner T, Riefler M, Krupková E, Bartrina y Manns I (2003) Structure and 1003

function of cytokinin oxidase/dehydrogenase genes of maize, rice, Arabidopsis and other 1004

species. J Plant Res 116: 241–252. 1005

Schoor S, Farrow S, Blaschke H, Lee S, Perry G, Schwartzenberg K von, Emery N, Moffatt B 1006

(2011) Adenosine kinase contributes to cytokinin interconversion in Arabidopsis. Plant 1007

Physiol 157: 659–672. 1008

Schroeder RY, Zhu A, Eubel H, Dahncke K, Witte C-P (2018) The ribokinases of Arabidopsis 1009

thaliana and Saccharomyces cerevisiae are required for ribose recycling from nucleotide 1010

catabolism, which in plants is not essential to survive prolonged dark stress. New Phytol 1011

217: 233–244. 1012

Serventi F, Ramazzina I, Lamberto I, Puggioni V, Gatti R, Percudani R (2010) Chemical basis 1013

of nitrogen recovery through the ureide pathway. Formation and hydrolysis of S-1014

ureidoglycine in plants and bacteria. ACS Chem Biol 5: 203–214. 1015

Sharma CB, Mittal R, Tanner W (1986) Purification and properties of a glycoprotein 1016

adenosine 5′-monophosphatase from the plasma membrane fraction of Arachis 1017

hypogaea cotyledons. Biochim Biophys Acta 884: 567–577. 1018

Shelp BJ, Atkins CA (1983) Role of Inosine monophosphate oxidoreductase in the formation 1019

of ureides in nitrogen-fixing nodules of cowpea (Vigna-unguiculata-L Walp). Plant Physiol 1020

72: 1029–1034. 1021

Sigel H, Operschall BP, Griesser R (2009) Xanthosine 5 ‘-monophosphate (XMP). Acid-base 1022

and metal ion-binding properties of a chameleon-like nucleotide. Chem Soc Rev 38: 1023

2465–2494. 1024

Siu KKW, Lee JE, Sufrin JR, Moffatt BA, McMillan M, Cornell KA, Isom C, Howell PL (2008) 1025

Molecular determinants of substrate specificity in plant 5’-methylthioadenosine 1026

nucleosidases. J Mol Biol 378: 112–128. 1027



32

Smith PM, Atkins CA (2002) Purine biosynthesis. Big in cell division, even bigger in nitrogen 1028

assimilation. Plant Physiol 128: 793–802. 1029

Soltabayeva A, Srivastava S, Kurmanbayeva A, Bekturova A, Fluhr R, Sagi M (2018) Early 1030

senescence in older leaves of low nitrate-grown Atxdh1 uncovers a role for purine 1031

catabolism in N supply. Plant Physiol 178: 1027–1044. 1032

Spetea C, Lundin B (2012) Evidence for nucleotide-dependent processes in the thylakoid 1033

lumen of plant chloroplasts—an update. FEBS Lett 586: 2946–2954. 1034

Stasolla C, Katahira R, Thorpe TA, Ashihara H (2003) Purine and pyrimidine nucleotide 1035

metabolism in higher plants. J Plant Physiol 160: 1271–1295. 1036

Stitt M, Lilley RM, Heldt HW (1982) Adenine-nucleotide levels in the cytosol, chloroplasts, 1037

and mitochondria of wheat leaf protoplasts. Plant Physiol 70: 971–977. 1038

Sugimoto H, Kusumi K, Noguchi K, Yano M, Yoshimura A, Iba K (2007) The rice nuclear 1039

gene, VIRESCENT 2, is essential for chloroplast development and encodes a novel type of 1040

guanylate kinase targeted to plastids and mitochondria. Plant J 52: 512–527. 1041

Sukrong S, Yun K-Y, Stadler P, Kumar C, Facciuolo T, Moffatt BA, Falcone DL (2012) 1042

Improved growth and stress tolerance in the Arabidopsis oxt1 mutant triggered by 1043

altered adenine metabolism. Mol Plant 5: 1310–1332. 1044

Tegeder M (2014) Transporters involved in source to sink partitioning of amino acids and 1045

ureides. Opportunities for crop improvement. J Exp Bot 65: 1865–1878. 1046

Tintemann H, Wasternack C, Benndorf R, Reinbothe H (1985) The rate-limiting step of 1047

uracil degradation in tomato cell-suspension cultures and Euglena-gracilis invivo studies. 1048

Comp Biochem Physiol, Part B: Biochem Mol Biol 82: 787–792. 1049

Todd CD, Polacco JC (2006) AtAAH encodes a protein with allantoate amidohydrolase 1050

activity from Arabidopsis thaliana. Planta 223: 1108–1113. 1051

Torres RJ, Puig JG (2007) Hypoxanthine-guanine phosophoribosyltransferase (HPRT) 1052

deficiency: Lesch-Nyhan syndrome. Orphanet J Rare Dis. 2: 48. 1053

Traub M, Florchinger M, Piecuch J, Kunz HH, Weise-Steinmetz A, Deitmer JW, Neuhaus HE, 1054

Mohlmann T (2007) The fluorouridine insensitive 1 (fur1) mutant is defective in 1055

equilibrative nucleoside transporter 3 (ENT3), and thus represents an important 1056

pyrimidine nucleoside uptake system in Arabidopsis thaliana. Plant J 49: 855–864. 1057

Tripathi D, Zhang T, Koo AJ, Stacey G, Tanaka K (2017) Extracellular ATP acts on jasmonate 1058

signaling to reinforce plant defense. Plant Physiol 176: 511–523. 1059

Ullrich A, Knecht W, Piskur J, Loffler M (2002) Plant dihydroorotate dehydrogenase differs 1060

significantly in substrate specificity and inhibition from the animal enzymes. FEBS Lett 1061

529: 346–350. 1062

Urarte E, Esteban R, Moran JF, Bittner F (2015) Established and proposed roles of xanthine 1063

oxidoreductase in oxidative and reductive pathways in plants. In KJ Gupta, AU 1064



33

Igamberdiev, eds, Reactive oxygen and nitrogen species signaling and communication in 1065

plants. Springer, Cham, Switzerland, pp. 15–42. 1066

Wagner KG, Backer AI (1992) Dynamics of nucleotides in plants studied on a cellular basis. 1067

In JK W, F M, eds, International Review of Cytology Vol. 134, pp. 1–84. 1068

Walsh TA, Green SB, Larrinua IM, Schmitzer PR (2001) Characterization of plant beta-1069

ureidopropionase and functional overexpression in Escherichia coli. Plant Physiol 125: 1070

1001–1011. 1071

Wang L, Li Z, Qian W, Guo W, Gao X, Huang L, Wang H, Zhu H, Wu JW,Wang D, Liu D (2011) 1072

The Arabidopsis purple acid phosphatase At-PAP10 is predominantly associated with the 1073

root surface and plays an important role in plant tolerance to phosphate limitation. Plant 1074

Physiol 157: 1283–1299. 1075

Wang C, Liu Z (2006) Arabidopsis ribonucleotide reductases are critical for cell cycle 1076

progression, DNA damage repair, and plant development. Plant Cell 18: 350–365. 1077

Wang C, Zhou M, Zhang X, Yao J, Zhang Y, Mou Z (2017) A lectin receptor kinase as a 1078

potential sensor for extracellular nicotinamide adenine dinucleotide in Arabidopsis 1079

thaliana. eLife 6: e25474 1080

Watanabe S, Matsumoto M, Hakomori Y, Takagi H, Shimada H, Sakamoto A (2014) The 1081

purine metabolite allantoin enhances abiotic stress tolerance through synergistic 1082

activation of abscisic acid metabolism. Plant Cell Environ 37: 1022–1036. 1083

Wei X, Song X, Wei L, Tang S, Sun J, Hu P, Cao X (2017) An epiallele of rice AK1 affects 1084

photosynthetic capacity. J Integr Plant Biol 59: 158–163. 1085

Werner AK, Medina-Escobar N, Zulawski M, Sparkes IA, Cao F-Q, Witte CP (2013) The 1086

ureide-degrading reactions of purine ring catabolism employ three amidohydrolases and 1087

one aminohydrolase in Arabidopsis, soybean, and rice. Plant Physiol 163: 672–681. 1088

Werner AK, Romeis T, Witte CP (2010) Ureide catabolism in Arabidopsis thaliana and 1089

Escherichia coli. Nat Chem Biol 6: 19–21. 1090

Werner AK, Sparkes IA, Romeis T, Witte CP (2008) Identification, biochemical 1091

characterization, and subcellular localization of allantoate amidohydrolases from 1092

Arabidopsis and soybean. Plant Physiol 146: 418–430. 1093

Werner AK, Witte CP (2011) The biochemistry of nitrogen mobilization: purine ring 1094

catabolism. Trends Plant Sci 16: 381–387. 1095

Witz S, Jung B, Furst S, Mohlmann T (2012) De novo pyrimidine nucleotide synthesis mainly 1096

occurs outside of plastids, but a previously undiscovered nucleobase importer provides 1097

substrates for the essential salvage pathway in Arabidopsis. Plant Cell 24: 1549–1559. 1098

Wu S, Alseekh S, Cuadros-Inostroza A, Fusari CM, Mutwil M, Kooke R, Keurentjes JB, 1099

Fernie AR, Willmitzer L, Brotman Y (2016) Combined use of genome-wide association 1100



34

data and correlation networks unravels key regulators of primary metabolism in 1101

Arabidopsis thaliana. PLoS GENET 12. 1102

Xu J, Deng Y, Li Q, Zhu X, He Z (2014) STRIPE2 encodes a putative dCMP deaminase that 1103

plays an important role in chloroplast development in rice. J Genet Genomics 41: 539–1104

548. 1105

Xu J, Zhang HY, Xie CH, Xue HW, Dijkhuis P, Liu CM (2005) EMBRYONIC FACTOR 1 encodes 1106

an AMP deaminase and is essential for the zygote to embryo transition in Arabidopsis. 1107

Plant J 42: 743–756. 1108

Xu J, Zhang L, Yang D-L, Li Q, He Z (2015) Thymidine kinases share a conserved function for 1109

nucleotide salvage and play an essential role in Arabidopsis thaliana growth and 1110

development. New Phytol 208: 1089–1103. 1111

Yamada H, Suzuki T, Terada K, Takei K, Ishikawa K, Miwa K, Yamashino T, Mizuno T (2001) 1112

The Arabidopsis AHK4 histidine kinase is a cytokinin-binding receptor that transduces 1113

cytokinin signals across the membrane. Plant Cell Physiol 42: 1017–1023. 1114

Ye W, Hu S, Wu L, Ge C, Cui Y, Chen P, Wang X, Xu J, Ren D, Dong G, Qian Q, Guo L (2016) 1115

White stripe leaf 12 (WSL12), encoding a nucleoside diphosphate kinase 2 (OsNDPK2), 1116

regulates chloroplast development and abiotic stress response in rice (Oryza sativa L.). 1117

Mol Breeding 36: 57. 1118

Yin Y, Katahira R, Ashihara H (2014) Metabolism of purine nucleosides and bases in 1119

suspension-cultured Arabidopsis thaliana cells. Eur Chem Bull 3: 925–934. 1120

Young LS, Harrison BR, Narayana, M. U. M., Moffatt BA, Gilroy S, Masson PH (2006) 1121

Adenosine kinase modulates root gravitropism and cap morphogenesis in arabidopsis. 1122

Plant Physiol 142: 564–573. 1123

Zarepour M, Kaspari K, Stagge S, Rethmeier R, Mendel RR, Bittner F (2010) Xanthine 1124

dehydrogenase AtXDH1 from Arabidopsis thaliana is a potent producer of superoxide 1125

anions via its NADH oxidase activity. Plant Mol Biol 72: 301–310. 1126

Zhang T, Feng P, Li Y, Yu P, Yu G, Sang X, Ling Y, Zeng X, Li Y, Huang J, Zhang T, Zhao F, 1127

Wang N, Zhang C, Yang Z, Wu R, He G (2018) VIRESCENT-ALBINO LEAF 1 regulates leaf 1128

colour development and cell division in rice. J Exp Bot 69: 4791–4804. 1129

Zhang X, Chen Y, Lin X, Hong X, Zhu Y, Li W, He W, An F, Guo H (2013) Adenine 1130

phosphoribosyl transferase 1 is a key enzyme catalyzing cytokinin conversion from 1131

nucleobases to nucleotides in Arabidopsis. Mol Plant 6: 1661–1672. 1132

Zhou K, Xia J, Wang Y, Ma T, Li Z (2017) A Young Seedling Stripe2 phenotype in rice is 1133

caused by mutation of a chloroplast-localized nucleoside diphosphate kinase 2 required 1134

for chloroplast biogenesis. Genet Mol Biol 40: 630–642. 1135

Zhou L, Lacroute F, Thornburg R (1998) Cloning, expression in Escherichia coli, and 1136

characterization of Arabidopsis thaliana UMP/CMP kinase. Plant Physiol 117: 245–254. 1137



35

Zhu X, Guo S, Wang Z, Du Q, Xing Y, Zhang T, Shen W, Sang X, Ling Y, He G (2016) Map-1138

based cloning and functional analysis of YGL8, which controls leaf colour in rice (Oryza 1139

sativa). BMC Plant Biol 16: 134. 1140

Zrenner R, Ashihara H (2011) Nucleotide Metabolism. In H Ashihara, A Crozier, A 1141

Komamine, eds, Plant metabolism and biotechnology. Wiley, Cambridge, New York, 1142

pp. 135–162. 1143

Zrenner R, Riegler H, Marquard CR, Lange PR, Geserick C, Bartosz CE, Chen CT, Slocum RD 1144

(2009) A functional analysis of the pyrimidine catabolic pathway in Arabidopsis. New 1145

Phytol 183: 117–132. 1146

Zrenner R, Stitt M, Sonnewald U, Boldt R (2006) Pyrimidine and purine biosynthesis and 1147

degradation in plants. Annu Rev Plant Biol 57: 805–836. 1148

1149



ADVANCES

• The pathways of plant nucleotide metabolism have been better defined through the detailed analyses of mutants and the discovery of many new genes / proteins involved, for example the plastid uracil transporter, the nucleoside hydrolases, the CTP synthases, and guanosine deaminase.

• It has become clear that purine nucleotide catabolism may not only be involved in recycling nitrogen, but also in producing catabolic intermediates which dampen stress responses.

• Extracellular ATP has emerged as a new signaling molecule.

• Cells contain many modified nucleotides. A first enzyme for the degradation of a modified nucleotide, N6-methyl-AMP, has been discovered.



OUTSTANDING QUESTIONS

• How is nucleotide metabolism regulated (i) on the enzymatic level (ii) by transcriptional and post-transcriptional mechanisms (iii) by compartmentalization or organization in protein complexes (iv) by transport (v) by tissue-specific gene expression?

• Which transporters mediate purine and pyrimidine metabolite movement, specifically (i) catabolic intermediates between different cellular compartments (ii) nucleotides into the organelles (iii) metabolites over long distances?

• How are the nucleotide and deoxynucleotide species in the distinct cellular compartments balanced? Is this adjusted upon developmental and environmental stimuli, and how is this achieved?

• Which nucleotide phosphatases mediate dephosphorylation in vivo, for example dephosphorylation of mononucleotides to nucleosides?

• How are modified and damaged nucleotides degraded, and how do they re-enter nucleotide metabolism?



N

NN

N 6

6

5

79

1

NH2

HN

NN

N

O

H2N

HN

NN

N

O

HN

NR

R R R R

O

O

HN

N

O

O

H C3N

N

NH2

O

O

HNN

NN O

adenineadenosine / deoxyadenosineAMP / dAMP

uracil

R = HR = ribose /

dexoyriboseR = ribose monophosphate /

deoxyribose monophosphate

uridine / dexoyuridineUMP / dUMP

cytosinecytidine / deoxycytidineCMP / dCMP

hypoxanthineinosine / deoxyinosineIMP /dIMP

guanineguanosine / deoxyguanosineGMP /dGMP

xanthinexanthosine / deoxyxanthosineXMP /dXMP

thyminethymidine= deoxythymidineTMP= dTMP

2’3’

= ribose

nucleoside / deoxynucleoside

nucleoside monophosphate (nucleoside mononucleotide)

nucleoside diphosphate

nucleoside triphosphate

nucleotides

= dexoyribose

nucleobase

O

OH OHH

OPO

O

O

OP

O

O

OP

O

O

R R

Figure 1. Structural composi�on of nucleobases, nucleosides, and nucleo�des.

For the nucleobases ‘R’ is simply a proton. For the nucleosides ‘R’ is a sugar moiety which can be ribose or deoxyribose (carrying a proton instead of a hydroxyl group at the 2’ carbon of the ribose). Nucleo�des have up to three phosphate groups esterified to the hydroxyl group of the 5’ carbon of the nucleoside sugar determining the prefix mono- di- and triphosphate in the name of the molecule. The terminal phosphate always carries two charges irrespec�ve of the number of phosphates present. The pyrimidine nucleobases (upper row) and the purine nucleobases (lower row) are shown with the groups a�ached to the heterocycles highlighted in red (oxo groups), blue (amino groups), and grey shading (methyl group).



AMPPRPPglutamineaspartatebicarbonateglycineformyl-tetrahydrofolate

NDPsNTPs

dNDPsdNTPs

DNA

RNA

nucleobasesnucleosides

UDP-GlcNADHetc.

S-adenosylmethionine

GMPUMP CTP

de novobiosynthesis

degradation

SAM metabolismribose

carbon dioxideammoniaglyoxylatemalonate- semialdehyde

biosynthesisdegradation

degradation

biosynthesis

phosphorylation

degradation

salvage

reduction

(via dNDPs)

(to nucleosides) decay, repair

AAAAA

Figure 2. Schema�c overview of plant nucleo�de metabolism.

Nucleo�des are synthesized ‘de novo’ from precursor molecules listed in the upper le� box. The phosphoryla�on of nucleoside monophosphates via nucleoside diphosphates (NDPs) generates nucleoside triphosphates (NTPs), which serve as building blocks for RNA synthesis and as precursors for the biosynthesis of the metabolites shown in the center (S-adenosyl methionine, UDP-glucose, and NADH are given as examples). However, the nucleoside triphosphates, in par�cular ATP and GTP, are not only precursors for other metabolites, but are also essen�al stores of chemical energy in the phosphoanhydride bonds used in a mul�tude of energe�c coupling reac�ons, as well as important donors of phosphate in kinase reac�ons (not shown). NDPs can be reduced to dNDPs (deoxynucleoside diphosphates), which a�er phosphoryla�on to dNTPs serve as precursors for DNA biosynthesis. RNA degrada�on in the cytosol releases nucleoside monophosphates, whereas nucleosides are produced during vacuolar RNA degrada�on. Adenosine and adenine are products of biochemical reac�ons involving S-adenosyl methionine (SAM). Non-enzyma�c decay (depurina�on) and enzyma�c repair reac�ons result in nucleoside and nucleobase release from DNA. Nucleobases and nucleosides can be recycled to nucleo�des in so called ‘salvage’ reac�ons. Plants are also capable of full nucleo�de degrada�on via certain nucleosides and nucleobases releasing the nitrogen of the nucleobases as ammonia.



AMP AMP

PLASTIDSA B MITOCHONDRIA

outer membrane

IMP IMP

XMP

GMP

purine de novo biosynthesis

2 reactions

At4g32400At3g20330

At4g22930

At5g23300

At3g54470

At1g30820CTPS1 to 5

At3g12670At4g02120At4g20320At4g34890

2 reactions

carbamoylasparate

carbamoylasparate

dihydroorotate

dihydroorotate

?

?

( )

10 reactions

orotate

orotate

UMP

pyrimidine de novo biosynthesis

UTP CTP

AMPDAt2g38280

IMPDH

At1g79470At1g16350

GMPS

ATCase

DHODH

DHOase

At1g63660

BT1

?

?

CYTOSOL PLASTIDS CYTOSOL

UMPS

CTPS

1

2

4 8

9

7

6

5

3

Figure 3. Purine and pyrimidine de novo biosynthesis.

(A) Purine de novo biosynthesis. (B) Pyrimidine de novo biosynthesis. Enzymes and transporters: BT1 (1), bri�le1; AMPD (2), AMP deaminase; IMPDH (3), IMP dehydrogenase; GMPS (4), GMP synthetase; ATCase (5), asparate transcarbamoylase; DHOase (6), dihydroorotatase; DHODH (7), dihydroorotate dehydrogenase; UMPS (8), UMP synthase; CTPS (9), CTP synthetase. An anchor symbol denotes an associa�on with the respec�ve membrane.



GMP

GDP

GTP

GMK

de novovia IMP

de novo

from synthesis

NDPK

deoxy adenosine AMP

ADP ADPdADP

dAMP

dATPATP

dGDP

dGMP

dGTP

AMK

dNK

synthesis

salvage

RNR

substrate level and oxidativephosphorylation

GDPsynthesis

RNR

NDPK

At2g37250

At1g72040

At3g07800At5g23070

At5g59440

At2g16370At4g34570At2g21550

deoxy guanosine dNK

salvage

salvage

At1g72040

At5g47840At5g50370At5g63400At5g35170

At4g09320At5g63310

At3g27060At2g21790

At3g23580At5g40942

At4g11010At4g23900At4g23895

At2g39270At3g01820

At3g60180At4g25280At5g26667At3g60961At3g18680At3g10030

CYTOSOL CYTOSOL

CYTOSOLCYTOSOL

CYTOSOLPLASTIDSMITOCHONDRIA


de novovia UTP

de novo

AMK1 to 7

UMK1 to 6

NDPK1 to 5

RNR1RNR2 (tso2)RNR2ARNR2B

DC

BA

CMP

CDP

CTP CTP

UMK

NDPK

dCDP

dCMP

dCTP

CDPsynthesisRNR

deoxy cytidine dNK

salvage

At1g72040


RNR


CYTOSOLMITOCHONDRIA

UMP

UDP

dUDP dUMP TMP

TMP

TDP

TTPUTP

UMK

DHFR-TS

TMKTK

NDPK

?

?

thymidine

de novovia UTP

10

10

10

10

11 11

11

1212

1313

1313

14

15

16

1817

At2g41880

At3g06200At3g57550

GMK 1 to 3

Figure 4. Synthesis of nucleoside and deoxynucleoside triphosphates. 1

Synthesis of (A) cy�dylates, (B) uridylates and thymidylates, (C) guanylates, and (D) 2 adenylates. RNR (10), ribonucleo�de reductase; dNK (11), deoxynucleoside kinase; UMK (12), 3 UMP kinase; NDPK (13), nucleoside diphosphate kinase; TK (14), thymidine kinase; DHFR-TS 4 (15), dihydrofolate reductase-thymidylate synthase; TMK (16), thymidylate kinase; GMK (17), 5 guanylate kinase; AMK (18), adenylate kinase. The subcellular loca�ons where enzymes with 6 these ac�vi�es are found are indicated. For TMK, a loca�on in the plas�ds is only assumed. 7 The mononucleo�des (AMP, GMP, UMP, and CMP) may also be derived from salvage 8 reac�ons (see Figures 5 and 6). 9



IMPadenosine AMP guanosine

xanthosineinosine

hypoxantine xanthine

urate

ringcatabolism

XMP GMP

adenine guanine

At4g34890

At2g36310At1g05620

XDHXDH

NSH1NSH2

NSH1NSH2

GSDA

ADK IGK

IGK

IMPDH

HGPRT

HGPRT

GMPS

XMPP

AMPD

GMPP

At5g28050

At1g71750APRT

At1g71750

At2g38280 At1g63660At1g79470At1g16350

At3g09820

At1g27450At1g80050At4g22570At4g12440At5g11160

At5g03300

At?

At?

At?

At?

CYTOSOL

urate

HIU

OHCU

(S)-allantoin

At2g26230

allantoate

(S)-ureidoglycine

glyoxylate

(S)-ureidoglycolate

UOX

At5g43600UAH

At4g17050

At4g20070

At4g04955 ALN

AAHNH3

NH3

NH32

At5g58220 ALNS

At5g58220 ALNS

UGAH

PERO

XISOM

EEN

DO

PLASM

IC R

ETICU

LUM

2. SAM, ethylene biosynthesis1. SAM, polyamine + nicotianamine biosynthesis

4. cytokinin degradation5. uptake

4. uptake

2. SAM, methylation reactions 3. AMP phosphotransferase

1. (vacuolar) RNA turnover

3. spontaneous depurination

1. (vacuolar) RNA turnover2. uptake

1. spontaneous depurination2. uptake

t-RNA turnover

‚A‘ deamination in DNAand base excision repair

sources

NH

O

NH

NH2

O H O

NH2O-NH

HN

OO

NH

NH2

O H

NH

HN

ON

HN-O

O

NH

O

HOH O

NH2O-

A

B

19

20

21

21

22

22 23

2425

26

27

29

30

30

31

32

33

34

2828

2 3 4

Figure 5. Salvage and degrada�on of purines.

(A) The reac�ons of purine nucleobase and nucleoside salvage as well as purine nucleo�de degrada�on, which overlaps par�ally with GMP synthesis. The salvage pathways are highlighted by light grey shading, degrada�on reac�ons are encircled in dark gray. Metabolites that can only undergo degrada�on and cannot be salvaged are shown with brown shading. (B) Purine ring catabolism. The transport steps for urate and (S)-allantoin are not shown explicitly. APRT (19), adenine phosphoribosyltransferase; ADK (20), adenosine kinase; AMPD (2), AMP deaminase; IMPDH (3), IMP dehydrogenase; GMPS (4), GMP synthetase; HGPRT (21), hypoxanthine guanine phosphoribosyltransferase; IGK (22), inosine guanosine kinase; GMPP (23), GMP phosphatase; GSDA (24), guanosine deaminase; XMPP (25), XMP phosphatase; NSH1 (26), nucleoside hydrolase 1; NSH2 (27), nucleoside hydrolase 2; XDH (28), xanthine dehydrogenase; UOX (29), urate oxidase; ALNS (30), allantoin synthase; ALN (31), allantoinase; AAH (32), allantoate amidohydrolase; UGAH (33), ureidoglycine aminohydrolase; UAH (34), ureidoglycolate amidohydrolase.



PLASTIDS CYTOSOL

ER

UMPUMP

UCK1 to 5

At3g17810

At2g36310At3g53900 At2g19570

At5g03555

At5g12200

At5g64370

DPYH

DPYD

NSH1

UCPP

UPRT

β-UP

uracil uracil

uridine

CMP

cytidine

dihydrouracil

NC-β-alanine

malonate-SA

β-alanine

PLUTO

At?

CDA

UCK UCKUCPPAt?

At5g40870At3g27190At1g55810At4g26510At3g27440

H/

H

N

N O

OHCH

53

*H

/CH3

*

H2

H

HN

N

O

O

OH

H

/CH3*

H

O

H2N

OH

H

/CH3 H

O

O

O

H

H

1. (vacuolar) RNA turnover2. uptake

sources

2. uptake

1. ‚C‘ deamination in DNA and base excision repair

BAATAt?

3535 3636

37

38

26

41

42

43

39

40

Figure 6. Salvage and degrada�on of pyrimidines.

NC-β-alanine, N-carbamyl-β-alanine; malonate-SA, malonate semialdehyde. UCK (35), uridine cy�dine kinase; UCPP (36), UMP CMP phosphatase; CDA (37), cy�dine deaminase; NSH1 (26), nucleoside hydrolase 1; PLUTO (38), plas�dic nucleobase transporter; UPRT (39), uracil phosphoribosyltransferase; DPYD (40), dihydropyrimidine dehydrogenase; DPYH (41), dihydropyrimidine hydrolase; β-UP (42), β-ureidopropionase; BAAT (43) β-alanine aminotransferase.



ATPATP

AMP

APOPLAST

signals

CELL

At1g14240

At5g18860

At4g05120

At5g56450

exocytosis

rupture

At3g10960At5g50300

NSH3

?

?

?

DORN1

adenosine

adenine

5‘ NT

APY3

AZG1/2NSH3

ENT3

pmANT1

4544

46

48

47

49

50

Figure 7. Excre�on, percep�on, and degrada�on of extracellular ATP

pmANT1 (44), plasma membrane adenine nucleo�de transporter; APY3 (45), apyrase 3; 5‘ NT (46), 5’ nucleo�dase; ENT3 (47), equilibra�ve nucleoside transporter 3; NSH3 (48), nucleoside hydrolase 3, AZG1/2 (49), azaguanine resistant 1 / azaguanine resistant 2, DORN1 (50), does not respond to nucleo�des 1 .



Parsed CitationsAlamillo JM, Diaz-Leal JL, Sanchez-Moran MV, Pineda M (2010) Molecular analysis of ureide accumulation under drought stress inPhaseolus vulgaris L. Plant Cell Environ 33: 1828–1837.

Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Ashihara H, Loukanina N, Stasolla C, Thorpe TA (2001) Pyrimidine metabolism during somatic embryo development in white spruce(Picea glauca). J Plant Physiol 158: 613–621.


Ashihara H, Stasolla C, Fujimura T, Crozier A (2018) Purine salvage in plants. Phytochemistry 147: 89–124.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Atkins CA, Shelp BJ, Storer PJ (1985) Purification and properties of inosine monophosphate oxidoreductase from nitrogen-fixingnodules of cowpea (Vigna-Unguiculata-L WALP). Arch Biochem Biophys 236: 807–814.


Atkins CA, Smith, P. M. C., Storer PJ (1997) Reexamination of the intracellular localization of de novo purine synthesis in cowpeanodules. Plant Physiol 113: 127–135.


Baccolini C, Witte C-P (2019) AMP and GMP catabolism in Arabidopsis converge on xanthosine, which is degraded by a nucleosidehydrolase heterocomplex. Plant Cell 31: 734–751.


Bahaji A, Muñoz FJ, Ovecka M, Baroja-Fernández E, Montero M, Li J, Hidalgo M, Almagro G, Sesma MT, Ezquer I, Pozueta-Romero J(2011a) Specific delivery of AtBT1 to mitochondria complements the aberrant growth and sterility phenotype of homozygous Atbt1Arabidopsis mutants. Plant J 68: 1115–1121.


Bahaji A, Ovecka M, Bárány I, Risueño MC, Muñoz FJ, Baroja-Fernández E, Montero M, Li J, Hidalgo M, Sesma MT, Ezquer I, TestillanoPS, Pozueta-Romero J (2011b) Dual targeting to mitochondria and plastids of AtBT1 and ZmBT1, two members of the mitochondrialcarrier family. Plant Cell Physiol 52: 597–609.


Barbado C, Cordoba-Canero D, Arizaa RR, Roldan-Arjona T (2018) Nonenzymatic release of N7-methylguanine channels repair ofabasic sites into an AP endonuclease-independent pathway in Arabidopsis. Proc Natl Acad Sci USA 115: E916-E924.


Beaudoin GAW, Johnson TS, Hanson AD (2018) The PLUTO plastidial nucleobase transporter also transports the thiamin precursorhydroxymethylpyrimidine. Biosci Rep 38.


Bernard C, Traub M, Kunz HH, Hach S, Trentmann O, Mohlmann T (2011) Equilibrative nucleoside transporter 1 (ENT1) is critical forpollen germination and vegetative growth in Arabidopsis. J Exp Bot 62: 4627–4637.


Bromley JR, Warnes BJ, Newell CA, Thomson JCP, James CM, Turnbull CGN, Hanke DE (2014) A purine nucleoside phosphorylase inSolanum tuberosum L. (potato) with specificity for cytokinins contributes to the duration of tuber endodormancy. Biochem J 458: 225–237.


Brychkova G, Alikulov Z, Fiuhr R, Sagi M (2008) A critical role for ureides in dark and senescence-induced purine remobilization isunmasked in the Atxdh1 Arabidopsis mutant. Plant J 54: 496–509.


Cao Y, Tanaka K, Nguyen CT, Stacey G (2014) Extracellular ATP is a central signaling molecule in plant stress responses. Curr OpinPlant Biol 20: 82–87.



https://www.ncbi.nlm.nih.gov/pubmed?term=Alamillo JM%2C Diaz%2DLeal JL%2C Sanchez%2DMoran MV%2C Pineda M %282010%29 Molecular analysis of ureide accumulation under drought stress in Phaseolus vulgaris L%2E Plant Cell Environ 33%3A 1828%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Molecular analysis of ureide accumulation under drought stress in Phaseolus vulgaris L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Molecular analysis of ureide accumulation under drought stress in Phaseolus vulgaris L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Alamillo&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ashihara H%2C Loukanina N%2C Stasolla C%2C Thorpe TA %282001%29 Pyrimidine metabolism during somatic embryo development in white spruce %28Picea glauca%29%2E J Plant Physiol 158%3A 613%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Pyrimidine metabolism during somatic embryo development in white spruce (Picea glauca)&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=Pyrimidine metabolism during somatic embryo development in white spruce (Picea glauca)&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ashihara&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ashihara H%2C Stasolla C%2C Fujimura T%2C Crozier A %282018%29 Purine salvage in plants%2E Phytochemistry 147%3A 89%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purine salvage in plants&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=Purine salvage in plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ashihara&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Atkins CA%2C Shelp BJ%2C Storer PJ %281985%29 Purification and properties of inosine monophosphate oxidoreductase from nitrogen%2Dfixing nodules of cowpea %28Vigna%2DUnguiculata%2DL WALP%29%2E Arch Biochem Biophys 236%3A 807%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purification and properties of inosine monophosphate oxidoreductase from nitrogen-fixing nodules of cowpea (Vigna-Unguiculata-L WALP)&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=Purification and properties of inosine monophosphate oxidoreductase from nitrogen-fixing nodules of cowpea (Vigna-Unguiculata-L WALP)&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Atkins&as_ylo=1985&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Atkins CA%2C Smith%2C P%2E M%2E C%2E%2C Storer PJ %281997%29 Reexamination of the intracellular localization of de novo purine synthesis in cowpea nodules%2E Plant Physiol 113%3A 127%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Reexamination of the intracellular localization of de novo purine synthesis in cowpea nodules&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=Reexamination of the intracellular localization of de novo purine synthesis in cowpea nodules&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Atkins&as_ylo=1997&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Baccolini C%2C Witte C%2DP %282019%29 AMP and GMP catabolism in Arabidopsis converge on xanthosine%2C which is degraded by a nucleoside hydrolase heterocomplex%2E Plant Cell 31%3A 734%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=AMP and GMP catabolism in Arabidopsis converge on xanthosine, which is degraded by a nucleoside hydrolase heterocomplex&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=AMP and GMP catabolism in Arabidopsis converge on xanthosine, which is degraded by a nucleoside hydrolase heterocomplex&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Baccolini&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bahaji A%2C Mu�oz FJ%2C Ovecka M%2C Baroja%2DFern�ndez E%2C Montero M%2C Li J%2C Hidalgo M%2C Almagro G%2C Sesma MT%2C Ezquer I%2C Pozueta%2DRomero J %282011a%29 Specific delivery of AtBT1 to mitochondria complements the aberrant growth and sterility phenotype of homozygous Atbt1 Arabidopsis mutants%2E Plant J 68%3A 1115%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Specific delivery of AtBT1 to mitochondria complements the aberrant growth and sterility phenotype of homozygous Atbt1 Arabidopsis mutants&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=Specific delivery of AtBT1 to mitochondria complements the aberrant growth and sterility phenotype of homozygous Atbt1 Arabidopsis mutants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bahaji&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bahaji A%2C Ovecka M%2C B�r�ny I%2C Risue�o MC%2C Mu�oz FJ%2C Baroja%2DFern�ndez E%2C Montero M%2C Li J%2C Hidalgo M%2C Sesma MT%2C Ezquer I%2C Testillano PS%2C Pozueta%2DRomero J %282011b%29 Dual targeting to mitochondria and plastids of AtBT1 and ZmBT1%2C two members of the mitochondrial carrier family%2E Plant Cell Physiol 52%3A 597%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=&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=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bahaji&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Barbado C%2C Cordoba%2DCanero D%2C Arizaa RR%2C Roldan%2DArjona T %282018%29 Nonenzymatic release of N7%2Dmethylguanine channels repair of abasic sites into an AP endonuclease%2Dindependent pathway in Arabidopsis%2E Proc Natl Acad Sci USA 115%3A E916%2DE&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Nonenzymatic release of N7-methylguanine channels repair of abasic sites into an AP endonuclease-independent pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Nonenzymatic release of N7-methylguanine channels repair of abasic sites into an AP endonuclease-independent pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Barbado&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Beaudoin GAW%2C Johnson TS%2C Hanson AD %282018%29 The PLUTO plastidial nucleobase transporter also transports the thiamin precursor hydroxymethylpyrimidine%2E Biosci Rep&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The PLUTO plastidial nucleobase transporter also transports the thiamin precursor hydroxymethylpyrimidine.&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 PLUTO plastidial nucleobase transporter also transports the thiamin precursor hydroxymethylpyrimidine.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Beaudoin&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bernard C%2C Traub M%2C Kunz HH%2C Hach S%2C Trentmann O%2C Mohlmann T %282011%29 Equilibrative nucleoside transporter 1 %28ENT1%29 is critical for pollen germination and vegetative growth in Arabidopsis%2E J Exp Bot 62%3A 4627%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Equilibrative nucleoside transporter 1 (ENT1) is critical for pollen germination and vegetative growth 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=Equilibrative nucleoside transporter 1 (ENT1) is critical for pollen germination and vegetative growth in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bernard&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Bromley JR%2C Warnes BJ%2C Newell CA%2C Thomson JCP%2C James CM%2C Turnbull CGN%2C Hanke DE %282014%29 A purine nucleoside phosphorylase in Solanum tuberosum L%2E %28potato%29 with specificity for cytokinins contributes to the duration of tuber endodormancy%2E Biochem J 458%3A 225%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A purine nucleoside phosphorylase in Solanum tuberosum L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=A purine nucleoside phosphorylase in Solanum tuberosum L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Bromley&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Brychkova G%2C Alikulov Z%2C Fiuhr R%2C Sagi M %282008%29 A critical role for ureides in dark and senescence%2Dinduced purine remobilization is unmasked in the Atxdh1 Arabidopsis mutant%2E Plant J 54%3A 496%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A critical role for ureides in dark and senescence-induced purine remobilization is unmasked in the Atxdh1 Arabidopsis mutant&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=A critical role for ureides in dark and senescence-induced purine remobilization is unmasked in the Atxdh1 Arabidopsis mutant&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Brychkova&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Cao Y%2C Tanaka K%2C Nguyen CT%2C Stacey G %282014%29 Extracellular ATP is a central signaling molecule in plant stress responses%2E Curr Opin Plant Biol 20%3A 82%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Extracellular ATP is a central signaling molecule in plant stress responses&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=Extracellular ATP is a central signaling molecule in plant stress responses&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Cao&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1


Carrari F, Coll-Garcia D, Schauer N, Lytovchenko A, Palacios-Rojas N, Balbo I, Rosso M, Fernie AR (2005) Deficiency of a plastidialadenylate kinase in Arabidopsis results in elevated photosynthetic amino acid biosynthesis and enhanced growth. Plant Physiol 137:70–82.


Carter AM, Tegeder M (2016) Increasing nitrogen fixation and seed development in soybean requires complex adjustments of nodulenitrogen metabolism and partitioning processes. Current Biol 26: 2044–2051.


Casartelli A, Melino VJ, Baumann U, Riboni M, Suchecki R, Jayasinghe NS, Mendis H, Watanabe M, Erban A, Zuther E, Hoefgen R,Roessner U, Okamoto M, Heuer S (2019) Opposite fates of the purine metabolite allantoin under water and nitrogen limitations inbread wheat. Plant Mol Biol 99: 477–497.


Chen F, Dong G, Ma X, Wang F, Zhang Y, Xiong E, Wu J, Wang H, Qian Q, Wu L, Yu Y (2018a) UMP kinase activity is involved in properchloroplast development in rice. Photosynth Res 137: 53–67.


Chen M, Herde M, Witte C-P (2016) Of the nine cytidine deaminase-like genes in Arabidopsis, eight are pseudogenes and only one isrequired to maintain pyrimidine homeostasis in vivo. Plant Physiol 171: 799–809.


Chen M, Urs MJ, Sánchez-González I, Olayioye MA, Herde M, Witte C-P (2018b) m6A RNA degradation products are catabolized by anevolutionarily conserved N6-Methyl-AMP deaminase in plant and mammalian cells. Plant Cell 30: 1511–1522.


Chen M, Witte C-P (2019) Functions and Dynamics of Methylation in Eukaryotic mRNA. In S Jurga, J Barciszewski, eds, The DNA, RNA,and Histone Methylomes. Springer International Publishing, Cham, pp. 333–351.Chen MJ, Thelen JJ (2011) Plastid uridine salvageactivity is required for photoassimilate allocation and partitioning in Arabidopsis. Plant Cell 23: 2991–3006.


Chen X, Zhu L, Xin L, Du K, Ran X, Cui X, Xiang Q, Zhang H, Xu P, Wu X (2015) Rice stripe1-2 and stripe1-3 mutants encoding the smallsubunit of ribonucleotide reductase are temperature sensitive and are required for chlorophyll biosynthesis. PloS One 10: e0130172.


Chiu T-Y, Lao J, Manalansan B, Loqué D, Roux SJ, Heazlewood JL (2015) Biochemical characterization of Arabidopsis apyrase familyreveals their roles in regulating endomembrane NDP/NMP homoeostasis. Biochem J 472: 43–54.


Choi J, Tanaka K, Cao Y, Qi Y, Qiu J, Liang Y, Lee SY, Stacey G (2014) Identification of a plant receptor for extracellular ATP. Science343: 290–294.


Clausen AR, Girandon L, Ali A, Knecht W, Rozpedowska E, Sandrini MP, Andreasson E, Munch-Petersen B, Piskur J (2012) Twothymidine kinases and one multisubstrate deoxyribonucleoside kinase salvage DNA precursors in Arabidopsis thaliana. FEBS J 279:3889–3897.


Clausen AR, Mutahir Z, Munch-Petersen B, Piškur J (2014) Plants salvage deoxyribonucleosides in mitochondria. Nucleosides,Nucleotides Nucleic Acids 33: 291–295.


Collier R, Tegeder M (2012) Soybean ureide transporters play a critical role in nodule development, function and nitrogen export.Plant J 72: 355–367.


Combes A, Lafleuriel J, Lefloch F (1989) The Inosine-Guanosine Kinase-Activity of Mitochondria in Tubers of Jerusalem Artichoke.Plant Physiol Biochem 27: 729–736.



https://www.ncbi.nlm.nih.gov/pubmed?term=Carrari F%2C Coll%2DGarcia D%2C Schauer N%2C Lytovchenko A%2C Palacios%2DRojas N%2C Balbo I%2C Rosso M%2C Fernie AR %282005%29 Deficiency of a plastidial adenylate kinase in Arabidopsis results in elevated photosynthetic amino acid biosynthesis and enhanced growth%2E Plant Physiol 137%3A 70%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Deficiency of a plastidial adenylate kinase in Arabidopsis results in elevated photosynthetic amino acid biosynthesis and enhanced growth&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Deficiency of a plastidial adenylate kinase in Arabidopsis results in elevated photosynthetic amino acid biosynthesis and enhanced growth&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Carrari&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Carter AM%2C Tegeder M %282016%29 Increasing nitrogen fixation and seed development in soybean requires complex adjustments of nodule nitrogen metabolism and partitioning processes%2E Current Biol 26%3A 2044%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Increasing nitrogen fixation and seed development in soybean requires complex adjustments of nodule nitrogen metabolism and partitioning processes&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=Increasing nitrogen fixation and seed development in soybean requires complex adjustments of nodule nitrogen metabolism and partitioning processes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Carter&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Casartelli A%2C Melino VJ%2C Baumann U%2C Riboni M%2C Suchecki R%2C Jayasinghe NS%2C Mendis H%2C Watanabe M%2C Erban A%2C Zuther E%2C Hoefgen R%2C Roessner U%2C Okamoto M%2C Heuer S %282019%29 Opposite fates of the purine metabolite allantoin under water and nitrogen limitations in bread wheat%2E Plant Mol Biol 99%3A 477%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Opposite fates of the purine metabolite allantoin under water and nitrogen limitations in bread wheat&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Opposite fates of the purine metabolite allantoin under water and nitrogen limitations in bread wheat&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Casartelli&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chen F%2C Dong G%2C Ma X%2C Wang F%2C Zhang Y%2C Xiong E%2C Wu J%2C Wang H%2C Qian Q%2C Wu L%2C Yu Y %282018a%29 UMP kinase activity is involved in proper chloroplast development in rice%2E Photosynth Res 137%3A 53%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=UMP kinase activity is involved in proper chloroplast development in rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=UMP kinase activity is involved in proper chloroplast development in rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chen&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chen M%2C Herde M%2C Witte C%2DP %282016%29 Of the nine cytidine deaminase%2Dlike genes in Arabidopsis%2C eight are pseudogenes and only one is required to maintain pyrimidine homeostasis in vivo%2E Plant Physiol 171%3A 799%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Of the nine cytidine deaminase-like genes in Arabidopsis, eight are pseudogenes and only one is required to maintain pyrimidine homeostasis in vivo&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=Of the nine cytidine deaminase-like genes in Arabidopsis, eight are pseudogenes and only one is required to maintain pyrimidine homeostasis in vivo&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chen&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chen M%2C Urs MJ%2C S�nchez%2DGonz�lez I%2C Olayioye MA%2C Herde M%2C Witte C%2DP %282018b%29 m6A RNA degradation products are catabolized by an evolutionarily conserved N6%2DMethyl%2DAMP deaminase in plant and mammalian cells%2E Plant Cell 30%3A 1511%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=m6A RNA degradation products are catabolized by an evolutionarily conserved N6-Methyl-AMP deaminase in plant and mammalian 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=m6A RNA degradation products are catabolized by an evolutionarily conserved N6-Methyl-AMP deaminase in plant and mammalian cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chen&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chen M%2C Witte C%2DP %282019%29 Functions and Dynamics of Methylation in Eukaryotic mRNA%2E In S Jurga%2C J Barciszewski%2C eds%2C The DNA%2C RNA%2C and Histone Methylomes%2E Springer International Publishing%2C Cham%2C pp%2E 333%26%23x2013%3B351%2EChen MJ%2C Thelen JJ %282011%29 Plastid uridine salvage activity is required for photoassimilate allocation and partitioning in Arabidopsis%2E Plant Cell 23%3A 2991%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Functions and Dynamics of Methylation in Eukaryotic mRNA&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=Functions and Dynamics of Methylation in Eukaryotic mRNA&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chen&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chen X%2C Zhu L%2C Xin L%2C Du K%2C Ran X%2C Cui X%2C Xiang Q%2C Zhang H%2C Xu P%2C Wu X %282015%29 Rice stripe1%2D2 and stripe1%2D3 mutants encoding the small subunit of ribonucleotide reductase are temperature sensitive and are required for chlorophyll biosynthesis%2E PloS One 10%3A e&dopt=abstract


https://scholar.google.com/scholar?as_q=Rice stripe1-2 and stripe1-3 mutants encoding the small subunit of ribonucleotide reductase are temperature sensitive and are required for chlorophyll biosynthesis&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=Rice stripe1-2 and stripe1-3 mutants encoding the small subunit of ribonucleotide reductase are temperature sensitive and are required for chlorophyll biosynthesis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chen&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Chiu T%2DY%2C Lao J%2C Manalansan B%2C Loqu� D%2C Roux SJ%2C Heazlewood JL %282015%29 Biochemical characterization of Arabidopsis apyrase family reveals their roles in regulating endomembrane NDP%2FNMP homoeostasis%2E Biochem J 472%3A 43%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Biochemical characterization of Arabidopsis apyrase family reveals their roles in regulating endomembrane NDP/NMP homoeostasis&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=Biochemical characterization of Arabidopsis apyrase family reveals their roles in regulating endomembrane NDP/NMP homoeostasis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Chiu&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Choi J%2C Tanaka K%2C Cao Y%2C Qi Y%2C Qiu J%2C Liang Y%2C Lee SY%2C Stacey G %282014%29 Identification of a plant receptor for extracellular ATP%2E Science 343%3A 290%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Identification of a plant receptor for extracellular ATP&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=Identification of a plant receptor for extracellular ATP&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Choi&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Clausen AR%2C Girandon L%2C Ali A%2C Knecht W%2C Rozpedowska E%2C Sandrini MP%2C Andreasson E%2C Munch%2DPetersen B%2C Piskur J %282012%29 Two thymidine kinases and one multisubstrate deoxyribonucleoside kinase salvage DNA precursors in Arabidopsis thaliana%2E FEBS J 279%3A 3889%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Two thymidine kinases and one multisubstrate deoxyribonucleoside kinase salvage DNA precursors in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Two thymidine kinases and one multisubstrate deoxyribonucleoside kinase salvage DNA precursors in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Clausen&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Clausen AR%2C Mutahir Z%2C Munch%2DPetersen B%2C Piškur J %282014%29 Plants salvage deoxyribonucleosides in mitochondria%2E Nucleosides%2C Nucleotides Nucleic Acids 33%3A 291%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Plants salvage deoxyribonucleosides in mitochondria&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=Plants salvage deoxyribonucleosides in mitochondria&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Clausen&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Collier R%2C Tegeder M %282012%29 Soybean ureide transporters play a critical role in nodule development%2C function and nitrogen export%2E Plant J 72%3A 355%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Soybean ureide transporters play a critical role in nodule development, function and nitrogen export&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=Soybean ureide transporters play a critical role in nodule development, function and nitrogen export&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Collier&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Combes A%2C Lafleuriel J%2C Lefloch F %281989%29 The Inosine%2DGuanosine Kinase%2DActivity of Mitochondria in Tubers of Jerusalem Artichoke%2E Plant Physiol Biochem 27%3A 729%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The Inosine-Guanosine Kinase-Activity of Mitochondria in Tubers of Jerusalem Artichoke&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 Inosine-Guanosine Kinase-Activity of Mitochondria in Tubers of Jerusalem Artichoke&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Combes&as_ylo=1989&as_allsubj=all&hl=en&c2coff=1


Cornelius S, Traub M, Bernard C, Salzig C, Lang P, Mohlmann T (2012) Nucleoside transport across the plasma membrane mediated byequilibrative nucleoside transporter 3 influences metabolism of Arabidopsis seedlings. Plant Biol 14: 696–705.


Cornelius S, Witz S, Rolletschek H, Mohlmann T (2011) Pyrimidine degradation influences germination seedling growth and productionof Arabidopsis seeds. J Exp Bot 62: 5623–5632.


Dahncke K, Witte CP (2013) Plant purine nucleoside catabolism employs a guanosine deaminase required for the generation ofxanthosine in Arabidopsis. Plant Cell 25: 4101–4109.


Dancer JE, Hughes RG, Lindell SD (1997) Adenosine-5'-phosphate deaminase. A novel herbicide target. Plant Physiol 114: 119–129.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Daumann M, Fischer M, Niopek-Witz S, Girke C, Möhlmann T (2015) Apoplastic nucleoside accumulation in Arabidopsis leads toreduced photosynthetic performance and increased susceptibility against Botrytis cinerea. Front Plant Sci 6: 1158.


Daumann M, Hickl D, Zimmer D, DeTar RA, Kunz H-H, Möhlmann T (2018) Characterization of filament-forming CTP synthases fromArabidopsis thaliana. Plant J: 96: 316–328.


Del Vecchio HA, Ying S, Park J, Knowles VL, Kanno S, Tanoi K, She Y-M, Plaxton WC (2014) The cell wall-targeted purple acidphosphatase AtPAP25 is critical for acclimation of Arabidopsis thaliana to nutritional phosphorus deprivation. Plant J 80: 569–581.


Deng WW, Ashihara H (2010) Profiles of purine metabolism in leaves and roots of Camellia sinensis seedlings. Plant Cell Physiol 51:2105–2118.


Desimone M, Catoni E, Ludewig U, Hilpert M, Schneider A, Kunze R, Tegeder M, Frommer WB, Schumacher K (2002) A novelsuperfamily of transporters for allantoin and other oxo derivatives of nitrogen heterocyclic compounds in Arabidopsis. Plant Cell 14:847–856.


Diaz-Leal JL, Galvez-Valdivieso G, Fernandez J, Pineda M, Alamillo JM (2012) Developmental effects on ureide levels are mediated bytissue-specific regulation of allantoinase in Phaseolus vulgaris L. J Exp Bot 63: 4095–4106.


Díaz-Leal JL, Torralbo F, Antonio Quiles F, Pineda M, Alamillo JM (2014) Molecular and functional characterization of allantoateamidohydrolase from Phaseolus vulgaris. Physiol Plant 152: 43–58.


Dong Q, Zhang Y-X, Zhou Q, Liu Q-E, Chen D-B, Wang H, Cheng S-H, Cao L-Y, Shen X-H (2019) UMP Kinase Regulates ChloroplastDevelopment and Cold Response in Rice. Int J Mol Sci 20: 2107.


Doremus HD, Jagendorf AT (1985) Subcellular localization of the pathway of de novo pyrimidine nucleotide biosynthesis in pea leaves.Plant Physiol 79: 856–861.


Dorion S, Rivoal J (2015) Clues to the functions of plant NDPK isoforms. Naunyn-Schmiedeberg's Arch Pharmacol 388: 119–132.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Dorion S, Rivoal J (2018) Plant nucleoside diphosphate kinase 1: A housekeeping enzyme with moonlighting activity. Plant SignalBehav 13: e1475804.


Feng X, Yang R, Zheng X, Zhang F (2012) Identification of a novel nuclear-localized adenylate kinase 6 from Arabidopsis thaliana as an www.plantphysiol.orgon April 24, 2020 - Published by Downloaded from Copyright © 2019 American Society of Plant Biologists. All rights reserved.

https://www.ncbi.nlm.nih.gov/pubmed?term=Cornelius S%2C Traub M%2C Bernard C%2C Salzig C%2C Lang P%2C Mohlmann T %282012%29 Nucleoside transport across the plasma membrane mediated by equilibrative nucleoside transporter 3 influences metabolism of Arabidopsis seedlings%2E Plant Biol 14%3A 696%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Nucleoside transport across the plasma membrane mediated by equilibrative nucleoside transporter 3 influences metabolism of Arabidopsis seedlings&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=Nucleoside transport across the plasma membrane mediated by equilibrative nucleoside transporter 3 influences metabolism of Arabidopsis seedlings&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Cornelius&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Cornelius S%2C Witz S%2C Rolletschek H%2C Mohlmann T %282011%29 Pyrimidine degradation influences germination seedling growth and production of Arabidopsis seeds%2E J Exp Bot 62%3A 5623%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Pyrimidine degradation influences germination seedling growth and production of Arabidopsis seeds&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=Pyrimidine degradation influences germination seedling growth and production of Arabidopsis seeds&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Cornelius&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Dahncke K%2C Witte CP %282013%29 Plant purine nucleoside catabolism employs a guanosine deaminase required for the generation of xanthosine in Arabidopsis%2E Plant Cell 25%3A 4101%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Plant purine nucleoside catabolism employs a guanosine deaminase required for the generation of xanthosine 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=Plant purine nucleoside catabolism employs a guanosine deaminase required for the generation of xanthosine in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Dahncke&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Dancer JE%2C Hughes RG%2C Lindell SD %281997%29 Adenosine%2D5%27%2Dphosphate deaminase%2E A novel herbicide target%2E Plant Physiol 114%3A 119%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Adenosine-5'-phosphate deaminase&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=Adenosine-5'-phosphate deaminase&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Dancer&as_ylo=1997&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Daumann M%2C Fischer M%2C Niopek%2DWitz S%2C Girke C%2C M�hlmann T %282015%29 Apoplastic nucleoside accumulation in Arabidopsis leads to reduced photosynthetic performance and increased susceptibility against Botrytis cinerea%2E Front Plant Sci&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Apoplastic nucleoside accumulation in Arabidopsis leads to reduced photosynthetic performance and increased susceptibility against Botrytis cinerea&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=Apoplastic nucleoside accumulation in Arabidopsis leads to reduced photosynthetic performance and increased susceptibility against Botrytis cinerea&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Daumann&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Daumann M%2C Hickl D%2C Zimmer D%2C DeTar RA%2C Kunz H%2DH%2C M�hlmann T %282018%29 Characterization of filament%2Dforming CTP synthases from Arabidopsis thaliana%2E Plant J%3A 96%3A 316%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Characterization of filament-forming CTP synthases from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Characterization of filament-forming CTP synthases from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Daumann&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Del Vecchio HA%2C Ying S%2C Park J%2C Knowles VL%2C Kanno S%2C Tanoi K%2C She Y%2DM%2C Plaxton WC %282014%29 The cell wall%2Dtargeted purple acid phosphatase AtPAP25 is critical for acclimation of Arabidopsis thaliana to nutritional phosphorus deprivation%2E Plant J 80%3A 569%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The cell wall-targeted purple acid phosphatase AtPAP25 is critical for acclimation of Arabidopsis thaliana to nutritional phosphorus deprivation&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 cell wall-targeted purple acid phosphatase AtPAP25 is critical for acclimation of Arabidopsis thaliana to nutritional phosphorus deprivation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Del&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Deng WW%2C Ashihara H %282010%29 Profiles of purine metabolism in leaves and roots of Camellia sinensis seedlings%2E Plant Cell Physiol 51%3A 2105%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Profiles of purine metabolism in leaves and roots of Camellia sinensis seedlings&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=Profiles of purine metabolism in leaves and roots of Camellia sinensis seedlings&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Deng&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Desimone M%2C Catoni E%2C Ludewig U%2C Hilpert M%2C Schneider A%2C Kunze R%2C Tegeder M%2C Frommer WB%2C Schumacher K %282002%29 A novel superfamily of transporters for allantoin and other oxo derivatives of nitrogen heterocyclic compounds in Arabidopsis%2E Plant Cell 14%3A 847%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A novel superfamily of transporters for allantoin and other oxo derivatives of nitrogen heterocyclic compounds 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 novel superfamily of transporters for allantoin and other oxo derivatives of nitrogen heterocyclic compounds in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Desimone&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Diaz%2DLeal JL%2C Galvez%2DValdivieso G%2C Fernandez J%2C Pineda M%2C Alamillo JM %282012%29 Developmental effects on ureide levels are mediated by tissue%2Dspecific regulation of allantoinase in Phaseolus vulgaris L%2E J Exp Bot 63%3A 4095%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Diaz-Leal&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=Developmental effects on ureide levels are mediated by tissue-specific regulation of allantoinase in Phaseolus vulgaris L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Developmental effects on ureide levels are mediated by tissue-specific regulation of allantoinase in Phaseolus vulgaris L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Diaz-Leal&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=D�az%2DLeal JL%2C Torralbo F%2C Antonio Quiles F%2C Pineda M%2C Alamillo JM %282014%29 Molecular and functional characterization of allantoate amidohydrolase from Phaseolus vulgaris%2E Physiol Plant 152%3A 43%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Díaz-Leal&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=Molecular and functional characterization of allantoate amidohydrolase from Phaseolus vulgaris&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=Molecular and functional characterization of allantoate amidohydrolase from Phaseolus vulgaris&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Díaz-Leal&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Dong Q%2C Zhang Y%2DX%2C Zhou Q%2C Liu Q%2DE%2C Chen D%2DB%2C Wang H%2C Cheng S%2DH%2C Cao L%2DY%2C Shen X%2DH %282019%29 UMP Kinase Regulates Chloroplast Development and Cold Response in Rice%2E Int J Mol Sci&dopt=abstract

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

https://scholar.google.com/scholar?as_q=UMP Kinase Regulates Chloroplast Development and Cold Response in Rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=UMP Kinase Regulates Chloroplast Development and Cold Response in Rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Dong&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Doremus HD%2C Jagendorf AT %281985%29 Subcellular localization of the pathway of de novo pyrimidine nucleotide biosynthesis in pea leaves%2E Plant Physiol 79%3A 856%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Subcellular localization of the pathway of de novo pyrimidine nucleotide biosynthesis in pea leaves&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=Subcellular localization of the pathway of de novo pyrimidine nucleotide biosynthesis in pea leaves&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Doremus&as_ylo=1985&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Dorion S%2C Rivoal J %282015%29 Clues to the functions of plant NDPK isoforms%2E Naunyn%2DSchmiedeberg%27s Arch Pharmacol 388%3A 119%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Clues to the functions of plant NDPK isoforms&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=Clues to the functions of plant NDPK isoforms&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Dorion&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Dorion S%2C Rivoal J %282018%29 Plant nucleoside diphosphate kinase 1%3A A housekeeping enzyme with moonlighting activity%2E Plant Signal Behav 13%3A e&dopt=abstract

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


https://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Dorion&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1


essential stem growth factor. Plant Physiol Biochem 61: 180–186.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Floyd BE, Morriss SC, MacIntosh GC, Bassham DC (2015) Evidence for autophagy-dependent pathways of rRNA turnover inArabidopsis. Autophagy 11: 2199–2212.


Gaillard C, Moffatt BA, Blacker M, Laloue M (1998a) Male sterility associated with APRT deficiency in Arabidopsis thaliana results froma mutation in the gene APT1. Mol Gen Genet 257: 348–353.


Gaillard C, Moffatt BA, Blacker M, Laloue M (1998b) Male sterility associated with APRT deficiency in Arabidopsis thaliana results froma mutation in the gene APT1. Mol Gen Genet 257: 348–353.


Garton S, Knight H, Warren GJ, Knight MR, Thorlby GJ (2007) Crinkled leaves 8 - a mutation in the large subunit of ribonucleotidereductase - leads to defects in leaf development and chloroplast division in Arabidopsis thaliana. Plant J 50: 118–127.


Girke C, Daumann M, Niopek-Witz S, Möhlmann T (2014) Nucleobase and nucleoside transport and integration into plant metabolism.Front Plant Sci 5: 443.


Gorelova V, Lepeleire J de, van Daele J, Pluim D, Meï C, Cuypers A, Leroux O, Rébeillé F, Schellens JHM, Blancquaert D, Stove CP,van der Straeten D (2017) Dihydrofolate reductase/thymidylate synthase fine-tunes the folate status and controls redox homeostasis inplants. Plant cell 29: 2831–2853.


Gupta A, Sharma CB (1996) Purification to homogeneity and characterization of plasma membrane and Golgi apparatus-specific 5'-adenosine monophosphatases from peanut cotyledons. Plant Sci 117: 65–74.


Han BW, Bingman CA, Mahnke DK, Bannen RM, Bednarek SY, Sabina RL, Phillips GN (2006) Membrane association, mechanism ofaction, and structure of Arabidopsis embryonic factor 1 (FAC1). J Biol Chem 81: 14939–14947.


Hauck OK, Scharnberg J, Escobar NM, Wanner G, Giavalisco P, Witte C-P 2014) Uric acid accumulation in an Arabidopsis urate oxidasemutant impairs seedling establishment by blocking peroxisome maintenance. Plant Cell 26: 3090–3100.


Hooper CM, Castleden IR, Tanz SK, Aryamanesh N, Millar AH (2017) SUBA4: the interactive data analysis centre for Arabidopsissubcellular protein locations. Nucleic Acids Res 45: D1064-D1074.


Hu D, Li Y, Jin W, Gong H, He Q, Li Y (2017) Identification and characterization of a plastidic adenine nucleotide uniporter (OsBT1-3)required for chloroplast development in the early leaf stage of rice. Sci Rep 7.


Irani S, Todd CD (2016) Ureide metabolism under abiotic stress in Arabidopsis thaliana. J Plant Physiol 199: 87–95.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Irani S, Todd CD (2018) Exogenous allantoin increases Arabidopsis seedlings tolerance to NaCl stress and regulates expression ofoxidative stress response genes. J Plant Physiol 221: 43–50.


Ito J, Batth TS, Petzold CJ, Redding-Johanson AM, Mukhopadhyay A, Verboom R, Meyer EH, Millar AH, Heazlewood JL (2011) Analysisof the Arabidopsis cytosolic proteome highlights subcellular partitioning of central plant metabolism. J Proteome Res 10: 1571-1582.


Jewell JB, Sowders JM, He R, Willis MA, Gang DR, Tanaka K (2019) Extracellular ATP Shapes a Defense-Related Transcriptome Both www.plantphysiol.orgon April 24, 2020 - Published by Downloaded from Copyright © 2019 American Society of Plant Biologists. All rights reserved.

https://www.ncbi.nlm.nih.gov/pubmed?term=Feng X%2C Yang R%2C Zheng X%2C Zhang F %282012%29 Identification of a novel nuclear%2Dlocalized adenylate kinase 6 from Arabidopsis thaliana as an essential stem growth factor%2E Plant Physiol Biochem 61%3A 180%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Identification of a novel nuclear-localized adenylate kinase 6 from Arabidopsis thaliana as an essential stem growth factor&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=Identification of a novel nuclear-localized adenylate kinase 6 from Arabidopsis thaliana as an essential stem growth factor&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Feng&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Floyd BE%2C Morriss SC%2C MacIntosh GC%2C Bassham DC %282015%29 Evidence for autophagy%2Ddependent pathways of rRNA turnover in Arabidopsis%2E Autophagy 11%3A 2199%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Evidence for autophagy-dependent pathways of rRNA turnover 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=Evidence for autophagy-dependent pathways of rRNA turnover in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Floyd&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Gaillard C%2C Moffatt BA%2C Blacker M%2C Laloue M %281998a%29 Male sterility associated with APRT deficiency in Arabidopsis thaliana results from a mutation in the gene APT1%2E Mol Gen Genet 257%3A 348%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Male sterility associated with APRT deficiency in Arabidopsis thaliana results from a mutation in the gene APT1&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=Male sterility associated with APRT deficiency in Arabidopsis thaliana results from a mutation in the gene APT1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gaillard&as_ylo=1998&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Gaillard C%2C Moffatt BA%2C Blacker M%2C Laloue M %281998b%29 Male sterility associated with APRT deficiency in Arabidopsis thaliana results from a mutation in the gene APT1%2E Mol Gen Genet 257%3A 348%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Male sterility associated with APRT deficiency in Arabidopsis thaliana results from a mutation in the gene APT1&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=Male sterility associated with APRT deficiency in Arabidopsis thaliana results from a mutation in the gene APT1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gaillard&as_ylo=1998&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Garton S%2C Knight H%2C Warren GJ%2C Knight MR%2C Thorlby GJ %282007%29 Crinkled leaves 8 %2D a mutation in the large subunit of ribonucleotide reductase %2D leads to defects in leaf development and chloroplast division in Arabidopsis thaliana%2E Plant J 50%3A 118%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Crinkled leaves 8 - a mutation in the large subunit of ribonucleotide reductase - leads to defects in leaf development and chloroplast division in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Crinkled leaves 8 - a mutation in the large subunit of ribonucleotide reductase - leads to defects in leaf development and chloroplast division in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Garton&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Girke C%2C Daumann M%2C Niopek%2DWitz S%2C M�hlmann T %282014%29 Nucleobase and nucleoside transport and integration into plant metabolism%2E Front Plant Sci&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Nucleobase and nucleoside transport and integration into plant metabolism&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=Nucleobase and nucleoside transport and integration into plant metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Girke&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Gorelova V%2C Lepeleire J de%2C van Daele J%2C Pluim D%2C Me� C%2C Cuypers A%2C Leroux O%2C R�beill� F%2C Schellens JHM%2C Blancquaert D%2C Stove CP%2C van der Straeten D %282017%29 Dihydrofolate reductase%2Fthymidylate synthase fine%2Dtunes the folate status and controls redox homeostasis in plants%2E Plant cell 29%3A 2831%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Dihydrofolate reductase/thymidylate synthase fine-tunes the folate status and controls redox homeostasis in plants&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=Dihydrofolate reductase/thymidylate synthase fine-tunes the folate status and controls redox homeostasis in plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gorelova&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Gupta A%2C Sharma CB %281996%29 Purification to homogeneity and characterization of plasma membrane and Golgi apparatus%2Dspecific 5%27%2Dadenosine monophosphatases from peanut cotyledons%2E Plant Sci 117%3A 65%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purification to homogeneity and characterization of plasma membrane and Golgi apparatus-specific 5'-adenosine monophosphatases from peanut cotyledons&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=Purification to homogeneity and characterization of plasma membrane and Golgi apparatus-specific 5'-adenosine monophosphatases from peanut cotyledons&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Gupta&as_ylo=1996&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Han BW%2C Bingman CA%2C Mahnke DK%2C Bannen RM%2C Bednarek SY%2C Sabina RL%2C Phillips GN %282006%29 Membrane association%2C mechanism of action%2C and structure of Arabidopsis embryonic factor 1 %28FAC1%29%2E J Biol Chem 81%3A 14939%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Membrane association, mechanism of action, and structure of Arabidopsis embryonic factor 1 (FAC1)&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=Membrane association, mechanism of action, and structure of Arabidopsis embryonic factor 1 (FAC1)&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Han&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hauck OK%2C Scharnberg J%2C Escobar NM%2C Wanner G%2C Giavalisco P%2C Witte C%2DP 2014%29 Uric acid accumulation in an Arabidopsis urate oxidase mutant impairs seedling establishment by blocking peroxisome maintenance%2E Plant Cell 26%3A 3090%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Witte C-P 2014) Uric acid accumulation in an Arabidopsis urate oxidase mutant impairs seedling establishment by blocking peroxisome maintenance&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=Witte C-P 2014) Uric acid accumulation in an Arabidopsis urate oxidase mutant impairs seedling establishment by blocking peroxisome maintenance&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hauck&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hooper CM%2C Castleden IR%2C Tanz SK%2C Aryamanesh N%2C Millar AH %282017%29 SUBA4%3A the interactive data analysis centre for Arabidopsis subcellular protein locations%2E Nucleic Acids Res 45%3A D1064%2DD&dopt=abstract

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


https://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hooper&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Hu D%2C Li Y%2C Jin W%2C Gong H%2C He Q%2C Li Y %282017%29 Identification and characterization of a plastidic adenine nucleotide uniporter %28OsBT1%2D3%29 required for chloroplast development in the early leaf stage of rice%2E Sci Rep&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Identification and characterization of a plastidic adenine nucleotide uniporter (OsBT1-3) required for chloroplast development in the early leaf stage of rice.&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Identification and characterization of a plastidic adenine nucleotide uniporter (OsBT1-3) required for chloroplast development in the early leaf stage of rice.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Hu&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Irani S%2C Todd CD %282016%29 Ureide metabolism under abiotic stress in Arabidopsis thaliana%2E J Plant Physiol 199%3A 87%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Ureide metabolism under abiotic stress in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Ureide metabolism under abiotic stress in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Irani&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Irani S%2C Todd CD %282018%29 Exogenous allantoin increases Arabidopsis seedlings tolerance to NaCl stress and regulates expression of oxidative stress response genes%2E J Plant Physiol 221%3A 43%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Exogenous allantoin increases Arabidopsis seedlings tolerance to NaCl stress and regulates expression of oxidative stress response genes&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Exogenous allantoin increases Arabidopsis seedlings tolerance to NaCl stress and regulates expression of oxidative stress response genes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Irani&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ito J%2C Batth TS%2C Petzold CJ%2C Redding%2DJohanson AM%2C Mukhopadhyay A%2C Verboom R%2C Meyer EH%2C Millar AH%2C Heazlewood JL %282011%29 Analysis of the Arabidopsis cytosolic proteome highlights subcellular partitioning of central plant metabolism%2E J Proteome Res&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Analysis of the Arabidopsis cytosolic proteome highlights subcellular partitioning of central plant metabolism&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=Analysis of the Arabidopsis cytosolic proteome highlights subcellular partitioning of central plant metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ito&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1


Independently and along with Other Defense Signaling Pathways. Plant Physiol 179: 1144–1158.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Jung B, Florchinger M, Kunz HH, Traub M, Wartenberg R, Jeblick W, Neuhaus HE, Mohlmann T (2009) Uridine-ribohydrolase is a keyregulator in the uridine degradation pathway of Arabidopsis. Plant Cell 21: 876–891.


Jung B, Hoffmann C, Mohlmann T (2011) Arabidopsis nucleoside hydrolases involved in intracellular and extracellular degradation ofpurines. Plant J 65: 703–711.


Kafer C, Zhou L, Santoso D, Guirgis A, Weers B, Park S, Thornburg R (2004) Regulation of pyrimidine metabolism in plants. FrontBiosci-Landmrk 9: 1611–1625.


Karran P, Lindahl T (1980) Hypoxanthine in deoxyribonucleic acid: generation by heat-induced hydrolysis of adenine residues andrelease in free form by a deoxyribonucleic acid glycosylase from calf thymus. Biochem 19: 6005–6011.


Katahira R, Ashihara H (2002) Profiles of pyrimidine biosynthesis, salvage and degradation in disks of potato (Solanum tuberosum L.)tubers. Planta 215: 821–828.


Katahira R, Ashihara H (2006) Profiles of purine biosynthesis, salvage and degradation in disks of potato (Solanum tuberosum L.)tubers. Planta 225: 115–126.


Kihara A, Saburi W, Wakuta S, Kim M-H, Hamada S, Ito H, Imai R, Matsui H (2011) Physiological and biochemical characterization ofthree nucleoside diphosphate kinase isozymes from rice (Oryza sativa L.). Biosci Biotechnol Biochem 75: 1740–1745.


Kirchberger S, Tjaden J, Neuhaus HE (2008) Characterization of the Arabidopsis Brittle1 transport protein and impact of reducedactivity on plant metabolism. Plant J 56: 51–63.


Kopecná M, Blaschke H, Kopecny D, Vigouroux A, Koncitíková R, Novák O, Kotland O, Strnad M, Moréra S, Schwartzenberg K von(2013) Structure and function of nucleoside hydrolases from Physcomitrella patens and maize catalyzing the hydrolysis of purine,pyrimidine, and cytokinin ribosides. Plant Physiol 163: 1568–1583.


Kumar V, Spangenberg O, Konrad M (2000) Cloning of the guanylate kinase homologues AGK-1 and AGK-2 from Arabidopsis thalianaand characterization of AGK-1. Eur J Biochem 267: 606–615.


Kurakawa T, Ueda N, Maekawa M, Kobayashi K, Kojima M, Nagato Y, Sakakibara H, Kyozuka J (2007) Direct control of shoot meristemactivity by a cytokinin-activating enzyme. Nature 445: 652 EP -.


Kuroha T, Tokunaga H, Kojima M, Ueda N, Ishida T, Nagawa S, Fukuda H, Sugimoto K, Sakakibara H (2009) Functional analyses ofLONELY GUY cytokinin-activating enzymes reveal the importance of the direct activation pathway in Arabidopsis. Plant Cell 21: 3152–3169.


Lamberto I, Percudani R, Gatti R, Folli C, Petrucco S (2010) Conserved alternative splicing of Arabidopsis transthyretin-like determinesprotein localization and S-allantoin synthesis in peroxisomes. Plant Cell 22: 1564–1574.


Lange PR, Geserick C, Tischendorf G, Zrenner R (2008) Functions of chloroplastic adenylate kinases in Arabidopsis. Plant Physiol 146:492–504.

Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title www.plantphysiol.orgon April 24, 2020 - Published by Downloaded from

Copyright © 2019 American Society of Plant Biologists. All rights reserved.

https://www.ncbi.nlm.nih.gov/pubmed?term=Jewell JB%2C Sowders JM%2C He R%2C Willis MA%2C Gang DR%2C Tanaka K %282019%29 Extracellular ATP Shapes a Defense%2DRelated Transcriptome Both Independently and along with Other Defense Signaling Pathways%2E Plant Physiol 179%3A 1144%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Extracellular ATP Shapes a Defense-Related Transcriptome Both Independently and along with Other Defense Signaling Pathways&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=Extracellular ATP Shapes a Defense-Related Transcriptome Both Independently and along with Other Defense Signaling Pathways&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jewell&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jung B%2C Florchinger M%2C Kunz HH%2C Traub M%2C Wartenberg R%2C Jeblick W%2C Neuhaus HE%2C Mohlmann T %282009%29 Uridine%2Dribohydrolase is a key regulator in the uridine degradation pathway of Arabidopsis%2E Plant Cell 21%3A 876%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Uridine-ribohydrolase is a key regulator in the uridine degradation pathway of Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Uridine-ribohydrolase is a key regulator in the uridine degradation pathway of Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jung&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Jung B%2C Hoffmann C%2C Mohlmann T %282011%29 Arabidopsis nucleoside hydrolases involved in intracellular and extracellular degradation of purines%2E Plant J 65%3A 703%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Arabidopsis nucleoside hydrolases involved in intracellular and extracellular degradation of purines&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Arabidopsis nucleoside hydrolases involved in intracellular and extracellular degradation of purines&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Jung&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kafer C%2C Zhou L%2C Santoso D%2C Guirgis A%2C Weers B%2C Park S%2C Thornburg R %282004%29 Regulation of pyrimidine metabolism in plants%2E Front Biosci%2DLandmrk 9%3A 1611%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Regulation of pyrimidine metabolism in plants&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Regulation of pyrimidine metabolism in plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kafer&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Karran P%2C Lindahl T %281980%29 Hypoxanthine in deoxyribonucleic acid%3A generation by heat%2Dinduced hydrolysis of adenine residues and release in free form by a deoxyribonucleic acid glycosylase from calf thymus%2E Biochem 19%3A 6005%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Hypoxanthine in deoxyribonucleic acid: generation by heat-induced hydrolysis of adenine residues and release in free form by a deoxyribonucleic acid glycosylase from calf thymus&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=Hypoxanthine in deoxyribonucleic acid: generation by heat-induced hydrolysis of adenine residues and release in free form by a deoxyribonucleic acid glycosylase from calf thymus&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Karran&as_ylo=1980&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Katahira R%2C Ashihara H %282002%29 Profiles of pyrimidine biosynthesis%2C salvage and degradation in disks of potato %28Solanum tuberosum L%2E%29 tubers%2E Planta 215%3A 821%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Profiles of pyrimidine biosynthesis, salvage and degradation in disks of potato (Solanum tuberosum L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Profiles of pyrimidine biosynthesis, salvage and degradation in disks of potato (Solanum tuberosum L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Katahira&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Katahira R%2C Ashihara H %282006%29 Profiles of purine biosynthesis%2C salvage and degradation in disks of potato %28Solanum tuberosum L%2E%29 tubers%2E Planta 225%3A 115%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Profiles of purine biosynthesis, salvage and degradation in disks of potato (Solanum tuberosum L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Profiles of purine biosynthesis, salvage and degradation in disks of potato (Solanum tuberosum L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Katahira&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kihara A%2C Saburi W%2C Wakuta S%2C Kim M%2DH%2C Hamada S%2C Ito H%2C Imai R%2C Matsui H %282011%29 Physiological and biochemical characterization of three nucleoside diphosphate kinase isozymes from rice %28Oryza sativa L%2E%29%2E Biosci Biotechnol Biochem 75%3A 1740%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Physiological and biochemical characterization of three nucleoside diphosphate kinase isozymes from rice (Oryza sativa L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Physiological and biochemical characterization of three nucleoside diphosphate kinase isozymes from rice (Oryza sativa L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kihara&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kirchberger S%2C Tjaden J%2C Neuhaus HE %282008%29 Characterization of the Arabidopsis Brittle1 transport protein and impact of reduced activity on plant metabolism%2E Plant J 56%3A 51%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Characterization of the Arabidopsis Brittle1 transport protein and impact of reduced activity on plant metabolism&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=Characterization of the Arabidopsis Brittle1 transport protein and impact of reduced activity on plant metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kirchberger&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kopecn� M%2C Blaschke H%2C Kopecny D%2C Vigouroux A%2C Koncit�kov� R%2C Nov�k O%2C Kotland O%2C Strnad M%2C Mor�ra S%2C Schwartzenberg K von %282013%29 Structure and function of nucleoside hydrolases from Physcomitrella patens and maize catalyzing the hydrolysis of purine%2C pyrimidine%2C and cytokinin ribosides%2E Plant Physiol 163%3A 1568%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Structure and function of nucleoside hydrolases from Physcomitrella patens and maize catalyzing the hydrolysis of purine, pyrimidine, and cytokinin ribosides&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=Structure and function of nucleoside hydrolases from Physcomitrella patens and maize catalyzing the hydrolysis of purine, pyrimidine, and cytokinin ribosides&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kopecná&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kumar V%2C Spangenberg O%2C Konrad M %282000%29 Cloning of the guanylate kinase homologues AGK%2D1 and AGK%2D2 from Arabidopsis thaliana and characterization of AGK%2D1%2E Eur J Biochem 267%3A 606%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Cloning of the guanylate kinase homologues AGK-1 and AGK-2 from Arabidopsis thaliana and characterization of AGK-1&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Cloning of the guanylate kinase homologues AGK-1 and AGK-2 from Arabidopsis thaliana and characterization of AGK-1&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kumar&as_ylo=2000&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kurakawa T%2C Ueda N%2C Maekawa M%2C Kobayashi K%2C Kojima M%2C Nagato Y%2C Sakakibara H%2C Kyozuka J %282007%29 Direct control of shoot meristem activity by a cytokinin%2Dactivating enzyme%2E Nature 445%3A 652 EP&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Direct control of shoot meristem activity by a cytokinin-activating enzyme&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=Direct control of shoot meristem activity by a cytokinin-activating enzyme&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kurakawa&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Kuroha T%2C Tokunaga H%2C Kojima M%2C Ueda N%2C Ishida T%2C Nagawa S%2C Fukuda H%2C Sugimoto K%2C Sakakibara H %282009%29 Functional analyses of LONELY GUY cytokinin%2Dactivating enzymes reveal the importance of the direct activation pathway in Arabidopsis%2E Plant Cell 21%3A 3152%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Functional analyses of LONELY GUY cytokinin-activating enzymes reveal the importance of the direct activation pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Functional analyses of LONELY GUY cytokinin-activating enzymes reveal the importance of the direct activation pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Kuroha&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Lamberto I%2C Percudani R%2C Gatti R%2C Folli C%2C Petrucco S %282010%29 Conserved alternative splicing of Arabidopsis transthyretin%2Dlike determines protein localization and S%2Dallantoin synthesis in peroxisomes%2E Plant Cell 22%3A 1564%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Conserved alternative splicing of Arabidopsis transthyretin-like determines protein localization and S-allantoin synthesis in peroxisomes&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=Conserved alternative splicing of Arabidopsis transthyretin-like determines protein localization and S-allantoin synthesis in peroxisomes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lamberto&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Lange PR%2C Geserick C%2C Tischendorf G%2C Zrenner R %282008%29 Functions of chloroplastic adenylate kinases in Arabidopsis%2E Plant Physiol 146%3A 492%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Functions of chloroplastic adenylate kinases 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=Functions of chloroplastic adenylate kinases in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lange&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1


Google Scholar: Author Only Title Only Author and Title

Lee S, Doxey AC, McConkey BJ, Moffatt BA (2012) Nuclear targeting of methyl-recycling enzymes in Arabidopsis thaliana is mediated byspecific protein interactions. Mol Plant 5: 231–248.


Leroch M, Kirchberger S, Haferkamp I, Wahl M, Neuhaus HE, Tjaden J (2005) Identification and characterization of a novel plastidicadenine nucleotide uniporter from Solanum tuberosum. J Biol Chem 280: 17992–18000.


Lescano CI, Martini C, González CA, Desimone M (2016) Allantoin accumulation mediated by allantoinase downregulation and transportby ureide permease 5 confers salt stress tolerance to Arabidopsis plants. Plant Mol Biol 91: 581-595


Lim MH, Wu J, Yao J, Gallardo IF, Dugger JW, Webb LJ, Huang J, Salmi ML, Song J, Clark G, Roux SJ (2014) Apyrase suppressionraises extracellular ATP levels and induces gene expression and cell wall changes characteristic of stress responses. Plant Physiol164: 2054–2067.


Lincker F, Philipps G, Chabouté M-E (2004) UV-C response of the ribonucleotide reductase large subunit involves both E2F-mediatedgene transcriptional regulation and protein subcellular relocalization in tobacco cells. Nucleic Acids Res 32: 1430–1438.


Liu S, Kracher B, Ziegler J, Birkenbihl RP, Somssich IE (2015) Negative regulation of ABA signaling by WRKY33 is critical forArabidopsis immunity towards Botrytis cinerea. eLife 4: e07295.


Liu XY, Qian WQ, Liu X, Qin HJ, Wang DW (2007) Molecular and functional analysis of hypoxanthine-guanine phosphoribosyltransferasefrom Arabidopsis thaliana. New Phytol 175: 448–461.


Luzarowski M, Kosmacz M, Sokolowska E, Jasinska W, Willmitzer L, Veyel D, Skirycz A (2017) Affinity purification with metabolomic andproteomic analysis unravels diverse roles of nucleoside diphosphate kinases. J Exp Bot 68: 3487–3499.


Ma X, Wang W, Bittner F, Schmidt N, Berkey R, Zhang L, King H, Zhang Y, Feng J, Wen Y, Tan L, Li Y, Zhang Q, Deng Z, Xiong X, Xiao S(2016) Dual and opposing roles of xanthine dehydrogenase in defense-associated reactive oxygen species metabolism in Arabidopsis.Plant Cell 28: 1108–1126.


Mainguet SE, Gakiere B, Majira A, Pelletier S, Bringel F, Guerard F, Caboche M, Berthome R, Renou JP (2009) Uracil salvage isnecessary for early Arabidopsis development. Plant J 60: 280–291.


Mansfield TA, Schultes NP, Mourad GS (2009) AtAzg1 and AtAzg2 comprise a novel family of purine transporters in Arabidopsis. FEBSLett 583: 481–486.


Moffatt B, Ashihara H (2002) Purine and pyrimdine nucleotide synthesis and metabolism, The Arabidopsis book. American Society ofPlant Biologists, Rockville, MD.


Moffatt B, Pethe C, Laloue M (1991) Metabolism of benzyladenine is impaired in a mutant of Arabidopsis thaliana lacking adeninephosphoribosyltransferase activity. Plant Physiol 95: 900–908.


Moffatt BA, Stevens YY, Allen MS, Snider JD, Pereira LA, Todorova MI, Summers PS, Weretilnyk EA, Martin-McCaffrey L, Wagner C(2002) Adenosine kinase deficiency is associated with developmental abnormalities and reduced transmethylation. Plant Physiol 128:812–821.



https://www.ncbi.nlm.nih.gov/pubmed?term=Lee S%2C Doxey AC%2C McConkey BJ%2C Moffatt BA %282012%29 Nuclear targeting of methyl%2Drecycling enzymes in Arabidopsis thaliana is mediated by specific protein interactions%2E Mol Plant 5%3A 231%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Nuclear targeting of methyl-recycling enzymes in Arabidopsis thaliana is mediated by specific protein interactions&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=Nuclear targeting of methyl-recycling enzymes in Arabidopsis thaliana is mediated by specific protein interactions&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lee&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Leroch M%2C Kirchberger S%2C Haferkamp I%2C Wahl M%2C Neuhaus HE%2C Tjaden J %282005%29 Identification and characterization of a novel plastidic adenine nucleotide uniporter from Solanum tuberosum%2E J Biol Chem 280%3A 17992%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Identification and characterization of a novel plastidic adenine nucleotide uniporter from Solanum tuberosum&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=Identification and characterization of a novel plastidic adenine nucleotide uniporter from Solanum tuberosum&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Leroch&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Lescano CI%2C Martini C%2C Gonz�lez CA%2C Desimone M %282016%29 Allantoin accumulation mediated by allantoinase downregulation and transport by ureide permease 5 confers salt stress tolerance to Arabidopsis plants%2E Plant Mol Biol&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Allantoin accumulation mediated by allantoinase downregulation and transport by ureide permease 5 confers salt stress tolerance to Arabidopsis plants&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=Allantoin accumulation mediated by allantoinase downregulation and transport by ureide permease 5 confers salt stress tolerance to Arabidopsis plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lescano&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Lim MH%2C Wu J%2C Yao J%2C Gallardo IF%2C Dugger JW%2C Webb LJ%2C Huang J%2C Salmi ML%2C Song J%2C Clark G%2C Roux SJ %282014%29 Apyrase suppression raises extracellular ATP levels and induces gene expression and cell wall changes characteristic of stress responses%2E Plant Physiol 164%3A 2054%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Apyrase suppression raises extracellular ATP levels and induces gene expression and cell wall changes characteristic of stress responses&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=Apyrase suppression raises extracellular ATP levels and induces gene expression and cell wall changes characteristic of stress responses&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lim&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Lincker F%2C Philipps G%2C Chabout� M%2DE %282004%29 UV%2DC response of the ribonucleotide reductase large subunit involves both E2F%2Dmediated gene transcriptional regulation and protein subcellular relocalization in tobacco cells%2E Nucleic Acids Res 32%3A 1430%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=UV-C response of the ribonucleotide reductase large subunit involves both E2F-mediated gene transcriptional regulation and protein subcellular relocalization in tobacco 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=UV-C response of the ribonucleotide reductase large subunit involves both E2F-mediated gene transcriptional regulation and protein subcellular relocalization in tobacco cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Lincker&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Liu S%2C Kracher B%2C Ziegler J%2C Birkenbihl RP%2C Somssich IE %282015%29 Negative regulation of ABA signaling by WRKY33 is critical for Arabidopsis immunity towards Botrytis cinerea%2E eLife 4%3A e&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Negative regulation of ABA signaling by WRKY33 is critical for Arabidopsis immunity towards Botrytis cinerea&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=Negative regulation of ABA signaling by WRKY33 is critical for Arabidopsis immunity towards Botrytis cinerea&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Liu&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Liu XY%2C Qian WQ%2C Liu X%2C Qin HJ%2C Wang DW %282007%29 Molecular and functional analysis of hypoxanthine%2Dguanine phosphoribosyltransferase from Arabidopsis thaliana%2E New Phytol 175%3A 448%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Molecular and functional analysis of hypoxanthine-guanine phosphoribosyltransferase from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Molecular and functional analysis of hypoxanthine-guanine phosphoribosyltransferase from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Liu&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Luzarowski M%2C Kosmacz M%2C Sokolowska E%2C Jasinska W%2C Willmitzer L%2C Veyel D%2C Skirycz A %282017%29 Affinity purification with metabolomic and proteomic analysis unravels diverse roles of nucleoside diphosphate kinases%2E J Exp Bot 68%3A 3487%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Affinity purification with metabolomic and proteomic analysis unravels diverse roles of nucleoside diphosphate kinases&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=Affinity purification with metabolomic and proteomic analysis unravels diverse roles of nucleoside diphosphate kinases&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Luzarowski&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ma X%2C Wang W%2C Bittner F%2C Schmidt N%2C Berkey R%2C Zhang L%2C King H%2C Zhang Y%2C Feng J%2C Wen Y%2C Tan L%2C Li Y%2C Zhang Q%2C Deng Z%2C Xiong X%2C Xiao S %282016%29 Dual and opposing roles of xanthine dehydrogenase in defense%2Dassociated reactive oxygen species metabolism in Arabidopsis%2E Plant Cell 28%3A 1108%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Dual and opposing roles of xanthine dehydrogenase in defense-associated reactive oxygen species metabolism 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=Dual and opposing roles of xanthine dehydrogenase in defense-associated reactive oxygen species metabolism in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ma&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Mainguet SE%2C Gakiere B%2C Majira A%2C Pelletier S%2C Bringel F%2C Guerard F%2C Caboche M%2C Berthome R%2C Renou JP %282009%29 Uracil salvage is necessary for early Arabidopsis development%2E Plant J 60%3A 280%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Uracil salvage is necessary for early Arabidopsis 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=Uracil salvage is necessary for early Arabidopsis development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Mainguet&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Mansfield TA%2C Schultes NP%2C Mourad GS %282009%29 AtAzg1 and AtAzg2 comprise a novel family of purine transporters in Arabidopsis%2E FEBS Lett 583%3A 481%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=AtAzg1 and AtAzg2 comprise a novel family of purine transporters 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=AtAzg1 and AtAzg2 comprise a novel family of purine transporters in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Mansfield&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Moffatt B%2C Ashihara H %282002%29 Purine and pyrimdine nucleotide synthesis and metabolism%2C The Arabidopsis book%2E American Society of Plant Biologists%2C Rockville%2C MD&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purine and pyrimdine nucleotide synthesis and metabolism, The Arabidopsis book.&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=Purine and pyrimdine nucleotide synthesis and metabolism, The Arabidopsis book.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Moffatt&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Moffatt B%2C Pethe C%2C Laloue M %281991%29 Metabolism of benzyladenine is impaired in a mutant of Arabidopsis thaliana lacking adenine phosphoribosyltransferase activity%2E Plant Physiol 95%3A 900%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Metabolism of benzyladenine is impaired in a mutant of Arabidopsis thaliana lacking adenine phosphoribosyltransferase activity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Metabolism of benzyladenine is impaired in a mutant of Arabidopsis thaliana lacking adenine phosphoribosyltransferase activity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Moffatt&as_ylo=1991&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Moffatt BA%2C Stevens YY%2C Allen MS%2C Snider JD%2C Pereira LA%2C Todorova MI%2C Summers PS%2C Weretilnyk EA%2C Martin%2DMcCaffrey L%2C Wagner C %282002%29 Adenosine kinase deficiency is associated with developmental abnormalities and reduced transmethylation%2E Plant Physiol 128%3A 812%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Adenosine kinase deficiency is associated with developmental abnormalities and reduced transmethylation&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=Adenosine kinase deficiency is associated with developmental abnormalities and reduced transmethylation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Moffatt&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1


Niopek-Witz S, Deppe J, Lemieux MJ, Möhlmann T (2014) Biochemical characterization and structure-function relationship of two plantNCS2 proteins, the nucleobase transporters NAT3 and NAT12 from Arabidopsis thaliana. Biochim Biophys Acta 1838: 3025–3035.


Niu M, Wang Y, Wang C, Lyu J, Wang Y, Dong H, Long W, Di Wang, Kong W, Wang L, Guo X, Sun L, Hu T, Zhai H, Wang H, Wan J (2017)ALR encoding dCMP deaminase is critical for DNA damage repair, cell cycle progression and plant development in rice. J Exp Bot 68:5773–5786.


Nizam S, Qiang X, Wawra S, Nostadt R, Getzke F, Schwanke F, Dreyer I, Langen G, Zuccaro A (2019) Serendipita indica E5′NTmodulates extracellular nucleotide levels in the plant apoplast and affects fungal colonization. EMBO Rep 20: e47430.


Nomura Y, Izumi A, Fukunaga Y, Kusumi K, Iba K, Watanabe S, Nakahira Y, Weber APM, Nozawa A, Tozawa Y (2014) Diversity inguanosine 3',5'-bisdiphosphate (ppGpp) sensitivity among guanylate kinases of bacteria and plants. J Biol Chem 289: 15631–15641.


Nourimand M, Todd CD (2019) There is a direct link between allantoin concentration and cadmium tolerance in Arabidopsis. PlantPhysiol Biochem 135: 441–449.


Ohler L, Niopek-Witz S, Mainguet SE, Möhlmann T (2019) Pyrimidine salvage: physiological functions and interaction with chloroplastbiogenesis. Plant Physiol 180: 1816–1828


Osugi A, Kojima M, Takebayashi Y, Ueda N, Kiba T, Sakakibara H (2017) Systemic transport of trans-zeatin and its precursor havediffering roles in Arabidopsis shoots. Nat Plants 3: 17112.


Pedroza-García J-A, Nájera-Martínez M, Mazubert C, Aguilera-Alvarado P, Drouin-Wahbi J, Sánchez-Nieto S, Gualberto JM, Raynaud C,Plasencia J (2019) Role of pyrimidine salvage pathway in the maintenance of organellar and nuclear genome integrity. Plant J 97: 430–446.


Pessoa J, Sarkany Z, Ferreira-da-Silva F, Martins S, Almeida MR, Li JM, Damas AM (2010) Functional characterization of Arabidopsisthaliana transthyretin-like protein. BMC Plant Biol 10: 30.


Phillips DA, Joseph CM, Hirsch PR (1997) Occurrence of flavonoids and nucleosides in agricultural soils. Appl Environ Microbiol 63:4573–4577.


Quiles FA, Galvez-Valdivieso G, Guerrero-Casado J, Pineda M, Piedras P (2019) Relationship between ureidic/amidic metabolism andantioxidant enzymatic activities in legume seedlings. Plant Physiol Biochem 138: 1–8.


Rapp M, Schein J, Hunt KA, Nalam V, Mourad GS, Schultes NP (2016) The solute specificity profiles of nucleobase cation symporter 1(NCS1) from Zea mays and Setaria viridis illustrate functional flexibility. Protoplasma 253: 611–623.


Redillas MCFR, Bang SW, Lee D‐K, Kim YS, Jung H, Chung PJ, Suh J‐W, Kim J‐K (2019) Allantoin accumulation throughoverexpression of ureide permease1 improves rice growth under limited nitrogen conditions. Plant Biotechnol J 17: 1289–1301.


Regierer B, Fernie AR, Springer F, Perez-Melis A, Leisse A, Koehl K, Willmitzer L, Geigenberger P, Kossmann J (2002) Starch contentand yield increase as a result of altering adenylate pools in transgenic plants. Nat Biotechnol 20: 1256–1260.


Rieder B, Neuhaus HE (2011) Identification of an Arabidopsis plasma membrane-located ATP transporter important for antherdevelopment. Plant Cell 23: 1932–1944.


https://www.ncbi.nlm.nih.gov/pubmed?term=Niopek%2DWitz S%2C Deppe J%2C Lemieux MJ%2C M�hlmann T %282014%29 Biochemical characterization and structure%2Dfunction relationship of two plant NCS2 proteins%2C the nucleobase transporters NAT3 and NAT12 from Arabidopsis thaliana%2E Biochim Biophys Acta 1838%3A 3025%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Niopek-Witz&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=Biochemical characterization and structure-function relationship of two plant NCS2 proteins, the nucleobase transporters NAT3 and NAT12 from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Biochemical characterization and structure-function relationship of two plant NCS2 proteins, the nucleobase transporters NAT3 and NAT12 from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Niopek-Witz&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Niu M%2C Wang Y%2C Wang C%2C Lyu J%2C Wang Y%2C Dong H%2C Long W%2C Di Wang%2C Kong W%2C Wang L%2C Guo X%2C Sun L%2C Hu T%2C Zhai H%2C Wang H%2C Wan J %282017%29 ALR encoding dCMP deaminase is critical for DNA damage repair%2C cell cycle progression and plant development in rice%2E J Exp Bot 68%3A 5773%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=ALR encoding dCMP deaminase is critical for DNA damage repair, cell cycle progression and plant development in rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=ALR encoding dCMP deaminase is critical for DNA damage repair, cell cycle progression and plant development in rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Niu&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Nizam S%2C Qiang X%2C Wawra S%2C Nostadt R%2C Getzke F%2C Schwanke F%2C Dreyer I%2C Langen G%2C Zuccaro A %282019%29 Serendipita indica E5�?�NT modulates extracellular nucleotide levels in the plant apoplast and affects fungal colonization%2E EMBO Rep 20%3A e&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Serendipita indica E5â�?²NT modulates extracellular nucleotide levels in the plant apoplast and affects fungal colonization&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=Serendipita indica E5â�?²NT modulates extracellular nucleotide levels in the plant apoplast and affects fungal colonization&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Nizam&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Nomura Y%2C Izumi A%2C Fukunaga Y%2C Kusumi K%2C Iba K%2C Watanabe S%2C Nakahira Y%2C Weber APM%2C Nozawa A%2C Tozawa Y %282014%29 Diversity in guanosine 3%27%2C5%27%2Dbisdiphosphate %28ppGpp%29 sensitivity among guanylate kinases of bacteria and plants%2E J Biol Chem 289%3A 15631%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Diversity in guanosine 3',5'-bisdiphosphate (ppGpp) sensitivity among guanylate kinases of bacteria and plants&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=Diversity in guanosine 3',5'-bisdiphosphate (ppGpp) sensitivity among guanylate kinases of bacteria and plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Nomura&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Nourimand M%2C Todd CD %282019%29 There is a direct link between allantoin concentration and cadmium tolerance in Arabidopsis%2E Plant Physiol Biochem 135%3A 441%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=There is a direct link between allantoin concentration and cadmium tolerance 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=There is a direct link between allantoin concentration and cadmium tolerance in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Nourimand&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ohler L%2C Niopek%2DWitz S%2C Mainguet SE%2C M�hlmann T %282019%29 Pyrimidine salvage%3A physiological functions and interaction with chloroplast biogenesis%2E Plant Physiol 180%3A 1816%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Pyrimidine salvage: physiological functions and interaction with chloroplast biogenesis.&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Pyrimidine salvage: physiological functions and interaction with chloroplast biogenesis.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ohler&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Osugi A%2C Kojima M%2C Takebayashi Y%2C Ueda N%2C Kiba T%2C Sakakibara H %282017%29 Systemic transport of trans%2Dzeatin and its precursor have differing roles in Arabidopsis shoots%2E Nat Plants&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Systemic transport of trans-zeatin and its precursor have differing roles in Arabidopsis shoots&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=Systemic transport of trans-zeatin and its precursor have differing roles in Arabidopsis shoots&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Osugi&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Pedroza%2DGarc�a J%2DA%2C N�jera%2DMart�nez M%2C Mazubert C%2C Aguilera%2DAlvarado P%2C Drouin%2DWahbi J%2C S�nchez%2DNieto S%2C Gualberto JM%2C Raynaud C%2C Plasencia J %282019%29 Role of pyrimidine salvage pathway in the maintenance of organellar and nuclear genome integrity%2E Plant J 97%3A 430%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Pedroza-García&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=Role of pyrimidine salvage pathway in the maintenance of organellar and nuclear genome integrity&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=Role of pyrimidine salvage pathway in the maintenance of organellar and nuclear genome integrity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Pedroza-García&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Pessoa J%2C Sarkany Z%2C Ferreira%2Dda%2DSilva F%2C Martins S%2C Almeida MR%2C Li JM%2C Damas AM %282010%29 Functional characterization of Arabidopsis thaliana transthyretin%2Dlike protein%2E BMC Plant Biol&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Functional characterization of Arabidopsis thaliana transthyretin-like protein&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=Functional characterization of Arabidopsis thaliana transthyretin-like protein&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Pessoa&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Phillips DA%2C Joseph CM%2C Hirsch PR %281997%29 Occurrence of flavonoids and nucleosides in agricultural soils%2E Appl Environ Microbiol 63%3A 4573%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Occurrence of flavonoids and nucleosides in agricultural soils&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=Occurrence of flavonoids and nucleosides in agricultural soils&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Phillips&as_ylo=1997&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Quiles FA%2C Galvez%2DValdivieso G%2C Guerrero%2DCasado J%2C Pineda M%2C Piedras P %282019%29 Relationship between ureidic%2Famidic metabolism and antioxidant enzymatic activities in legume seedlings%2E Plant Physiol Biochem 138%3A 1%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Relationship between ureidic/amidic metabolism and antioxidant enzymatic activities in legume seedlings&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=Relationship between ureidic/amidic metabolism and antioxidant enzymatic activities in legume seedlings&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Quiles&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Rapp M%2C Schein J%2C Hunt KA%2C Nalam V%2C Mourad GS%2C Schultes NP %282016%29 The solute specificity profiles of nucleobase cation symporter 1 %28NCS1%29 from Zea mays and Setaria viridis illustrate functional flexibility%2E Protoplasma 253%3A 611%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The solute specificity profiles of nucleobase cation symporter 1 (NCS1) from Zea mays and Setaria viridis illustrate functional flexibility&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 solute specificity profiles of nucleobase cation symporter 1 (NCS1) from Zea mays and Setaria viridis illustrate functional flexibility&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Rapp&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Redillas MCFR%2C Bang SW%2C Lee D�?�K%2C Kim YS%2C Jung H%2C Chung PJ%2C Suh J�?�W%2C Kim J�?�K %282019%29 Allantoin accumulation through overexpression of ureide permease1 improves rice growth under limited nitrogen conditions%2E Plant Biotechnol J 17%3A 1289%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Allantoin accumulation through overexpression of ureide permease1 improves rice growth under limited nitrogen conditions&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=Allantoin accumulation through overexpression of ureide permease1 improves rice growth under limited nitrogen conditions&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Redillas&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Regierer B%2C Fernie AR%2C Springer F%2C Perez%2DMelis A%2C Leisse A%2C Koehl K%2C Willmitzer L%2C Geigenberger P%2C Kossmann J %282002%29 Starch content and yield increase as a result of altering adenylate pools in transgenic plants%2E Nat Biotechnol 20%3A 1256%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Starch content and yield increase as a result of altering adenylate pools in transgenic plants&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=Starch content and yield increase as a result of altering adenylate pools in transgenic plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Regierer&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1



Riegler H, Geserick C, Zrenner R (2011) Arabidopsis thaliana nucleosidase mutants provide new insights into nucleoside degradation.New Phytol 191: 349–359.


Riewe D, Grosman L, Fernie AR, Wucke C, Geigenberger P (2008a) The potato-specific apyrase is apoplastically localized and hasinfluence on gene expression, growth, and development. Plant Physiol 147: 1092–1109.


Riewe D, Grosman L, Fernie AR, Zauber H, Wucke C, Geigenberger P (2008b) A cell wall-bound adenosine nucleosidase is involved inthe salvage of extracellular ATP in Solanum tuberosum. Plant Cell Physiol 49: 1572–1579.


Riggs JW, Rockwell NC, Cavales PC, Callis J (2016) Identification of the plant ribokinase and discovery of a role for Arabidopsisribokinase in nucleoside metabolism. J Biol Chem 291: 22572–22582.


Romanov GA, Lomin SN, Schmülling T (2018) Cytokinin signaling: from the ER or from the PM? That is the question! New Phytol 218:41–53.


Ronceret A, Gadea-Vacas J, Guilleminot J, Lincker F, Delorme V, Lahmy S, Pelletier G, Chabouté M-E, Devic M (2008) The first zygoticdivision in Arabidopsis requires de novo transcription of thymidylate kinase. Plant J 53: 776–789.


Sabina RL, Paul AL, Ferl RJ, Laber B, Lindell SD (2007) Adenine nucleotide pool perturbation is a metabolic trigger for AMP deaminaseinhibitor-based herbicide toxicity. Plant Physiol 143: 1752–1760.


Sakakibara H (2005) Cytokinin biosynthesis and regulation. Vitam Horm 72: 271–287.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Sauge-Merle S, Falconet D, Fontecave M (1999) An active ribonucleotide reductase from Arabidopsis thaliana - Cloning, expressionand characterization of the large subunit. Eur J Biochem 266: 62–69.


Sauter M, Moffatt B, Saechao MC, Hell R, Wirtz M (2013) Methionine salvage and S-adenosylmethionine: essential links betweensulfur, ethylene and polyamine biosynthesis. Biochem J 451: 145–154.


Schmid L-M, Ohler L, Möhlmann T, Brachmann A, Muiño JM, Leister D, Meurer J, Manavski N (2019) PUMPKIN, the sole plastid UMPkinase, associates with group II introns and alters their metabolism. Plant Physiol 179: 248–264.


Schmidt A, Baumann N, Schwarzkopf A, Frommer WB, Desimone M (2006) Comparative studies on ureide permeases in Arabidopsisthaliana and analysis of two alternative splice variants of AtUPS5. Planta 224: 1329–1340.


Schmidt A, Su YH, Kunze R, Warner S, Hewitt M, Slocum RD, Ludewig U, Frommer WB, Desimone M (2004) UPS1 and UPS2 fromArabidopsis mediate high affinity transport of uracil and 5-fluorouracil. J Biol Chem 279: 44817–44824.


Schmülling T, Werner T, Riefler M, Krupková E, Bartrina y Manns I (2003) Structure and function of cytokinin oxidase/dehydrogenasegenes of maize, rice, Arabidopsis and other species. J Plant Res 116: 241–252.


Schoor S, Farrow S, Blaschke H, Lee S, Perry G, Schwartzenberg K von, Emery N, Moffatt B (2011) Adenosine kinase contributes tocytokinin interconversion in Arabidopsis. Plant Physiol 157: 659–672.

Pubmed: Author and Title www.plantphysiol.orgon April 24, 2020 - Published by Downloaded from Copyright © 2019 American Society of Plant Biologists. All rights reserved.

https://www.ncbi.nlm.nih.gov/pubmed?term=Rieder B%2C Neuhaus HE %282011%29 Identification of an Arabidopsis plasma membrane%2Dlocated ATP transporter important for anther development%2E Plant Cell 23%3A 1932%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Identification of an Arabidopsis plasma membrane-located ATP transporter important for anther 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=Identification of an Arabidopsis plasma membrane-located ATP transporter important for anther development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Rieder&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Riegler H%2C Geserick C%2C Zrenner R %282011%29 Arabidopsis thaliana nucleosidase mutants provide new insights into nucleoside degradation%2E New Phytol 191%3A 349%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Arabidopsis thaliana nucleosidase mutants provide new insights into nucleoside degradation&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Arabidopsis thaliana nucleosidase mutants provide new insights into nucleoside degradation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Riegler&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Riewe D%2C Grosman L%2C Fernie AR%2C Wucke C%2C Geigenberger P %282008a%29 The potato%2Dspecific apyrase is apoplastically localized and has influence on gene expression%2C growth%2C and development%2E Plant Physiol 147%3A 1092%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The potato-specific apyrase is apoplastically localized and has influence on gene expression, growth, and development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=The potato-specific apyrase is apoplastically localized and has influence on gene expression, growth, and development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Riewe&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Riewe D%2C Grosman L%2C Fernie AR%2C Zauber H%2C Wucke C%2C Geigenberger P %282008b%29 A cell wall%2Dbound adenosine nucleosidase is involved in the salvage of extracellular ATP in Solanum tuberosum%2E Plant Cell Physiol 49%3A 1572%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A cell wall-bound adenosine nucleosidase is involved in the salvage of extracellular ATP in Solanum tuberosum&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 cell wall-bound adenosine nucleosidase is involved in the salvage of extracellular ATP in Solanum tuberosum&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Riewe&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Riggs JW%2C Rockwell NC%2C Cavales PC%2C Callis J %282016%29 Identification of the plant ribokinase and discovery of a role for Arabidopsis ribokinase in nucleoside metabolism%2E J Biol Chem 291%3A 22572%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Identification of the plant ribokinase and discovery of a role for Arabidopsis ribokinase in nucleoside metabolism&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=Identification of the plant ribokinase and discovery of a role for Arabidopsis ribokinase in nucleoside metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Riggs&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Romanov GA%2C Lomin SN%2C Schm�lling T %282018%29 Cytokinin signaling%3A from the ER or from the PM%3F That is the question%21 New Phytol 218%3A 41%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Cytokinin signaling: from the ER or from the PM&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Cytokinin signaling: from the ER or from the PM&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Romanov&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ronceret A%2C Gadea%2DVacas J%2C Guilleminot J%2C Lincker F%2C Delorme V%2C Lahmy S%2C Pelletier G%2C Chabout� M%2DE%2C Devic M %282008%29 The first zygotic division in Arabidopsis requires de novo transcription of thymidylate kinase%2E Plant J 53%3A 776%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The first zygotic division in Arabidopsis requires de novo transcription of thymidylate kinase&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 first zygotic division in Arabidopsis requires de novo transcription of thymidylate kinase&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ronceret&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sabina RL%2C Paul AL%2C Ferl RJ%2C Laber B%2C Lindell SD %282007%29 Adenine nucleotide pool perturbation is a metabolic trigger for AMP deaminase inhibitor%2Dbased herbicide toxicity%2E Plant Physiol 143%3A 1752%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Adenine nucleotide pool perturbation is a metabolic trigger for AMP deaminase inhibitor-based herbicide toxicity&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=Adenine nucleotide pool perturbation is a metabolic trigger for AMP deaminase inhibitor-based herbicide toxicity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sabina&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sakakibara H %282005%29 Cytokinin biosynthesis and regulation%2E Vitam Horm 72%3A 271%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Cytokinin biosynthesis and regulation&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Cytokinin biosynthesis and regulation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sakakibara&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sauge%2DMerle S%2C Falconet D%2C Fontecave M %281999%29 An active ribonucleotide reductase from Arabidopsis thaliana %2D Cloning%2C expression and characterization of the large subunit%2E Eur J Biochem 266%3A 62%26%23x&dopt=abstract

https://scholar.google.com/scholar?as_q=&num=10&as_occt=any&as_sauthors=Sauge-Merle&hl=en&c2coff=1

https://scholar.google.com/scholar?as_q=An active ribonucleotide reductase from Arabidopsis thaliana - Cloning, expression and characterization of the large subunit&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=An active ribonucleotide reductase from Arabidopsis thaliana - Cloning, expression and characterization of the large subunit&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sauge-Merle&as_ylo=1999&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sauter M%2C Moffatt B%2C Saechao MC%2C Hell R%2C Wirtz M %282013%29 Methionine salvage and S%2Dadenosylmethionine%3A essential links between sulfur%2C ethylene and polyamine biosynthesis%2E Biochem J 451%3A 145%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Methionine salvage and S-adenosylmethionine: essential links between sulfur, ethylene and polyamine biosynthesis&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=Methionine salvage and S-adenosylmethionine: essential links between sulfur, ethylene and polyamine biosynthesis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sauter&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schmid L%2DM%2C Ohler L%2C M�hlmann T%2C Brachmann A%2C Mui�o JM%2C Leister D%2C Meurer J%2C Manavski N %282019%29 PUMPKIN%2C the sole plastid UMP kinase%2C associates with group II introns and alters their metabolism%2E Plant Physiol 179%3A 248%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=PUMPKIN, the sole plastid UMP kinase, associates with group II introns and alters their metabolism&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=PUMPKIN, the sole plastid UMP kinase, associates with group II introns and alters their metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schmid&as_ylo=2019&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schmidt A%2C Baumann N%2C Schwarzkopf A%2C Frommer WB%2C Desimone M %282006%29 Comparative studies on ureide permeases in Arabidopsis thaliana and analysis of two alternative splice variants of AtUPS5%2E Planta 224%3A 1329%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Comparative studies on ureide permeases in Arabidopsis thaliana and analysis of two alternative splice variants of AtUPS5&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=Comparative studies on ureide permeases in Arabidopsis thaliana and analysis of two alternative splice variants of AtUPS5&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schmidt&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schmidt A%2C Su YH%2C Kunze R%2C Warner S%2C Hewitt M%2C Slocum RD%2C Ludewig U%2C Frommer WB%2C Desimone M %282004%29 UPS1 and UPS2 from Arabidopsis mediate high affinity transport of uracil and 5%2Dfluorouracil%2E J Biol Chem 279%3A 44817%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=UPS1 and UPS2 from Arabidopsis mediate high affinity transport of uracil and 5-fluorouracil&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=UPS1 and UPS2 from Arabidopsis mediate high affinity transport of uracil and 5-fluorouracil&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schmidt&as_ylo=2004&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schm�lling T%2C Werner T%2C Riefler M%2C Krupkov� E%2C Bartrina y Manns I %282003%29 Structure and function of cytokinin oxidase%2Fdehydrogenase genes of maize%2C rice%2C Arabidopsis and other species%2E J Plant Res 116%3A 241%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Structure and function of cytokinin oxidase/dehydrogenase genes of maize, rice, Arabidopsis and other species&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=Structure and function of cytokinin oxidase/dehydrogenase genes of maize, rice, Arabidopsis and other species&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schmülling&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schoor S%2C Farrow S%2C Blaschke H%2C Lee S%2C Perry G%2C Schwartzenberg K von%2C Emery N%2C Moffatt B %282011%29 Adenosine kinase contributes to cytokinin interconversion in Arabidopsis%2E Plant Physiol 157%3A 659%26%23x&dopt=abstract



Schroeder RY, Zhu A, Eubel H, Dahncke K, Witte C-P (2018) The ribokinases of Arabidopsis thaliana and Saccharomyces cerevisiae arerequired for ribose recycling from nucleotide catabolism, which in plants is not essential to survive prolonged dark stress. New Phytol217: 233–244.


Serventi F, Ramazzina I, Lamberto I, Puggioni V, Gatti R, Percudani R (2010) Chemical basis of nitrogen recovery through the ureidepathway. Formation and hydrolysis of S-ureidoglycine in plants and bacteria. ACS Chem Biol 5: 203–214.


Sharma CB, Mittal R, Tanner W (1986) Purification and properties of a glycoprotein adenosine 5′-monophosphatase from the plasmamembrane fraction of Arachis hypogaea cotyledons. Biochim Biophys Acta 884: 567–577.


Shelp BJ, Atkins CA (1983) Role of Inosine monophosphate oxidoreductase in the formation of ureides in nitrogen-fixing nodules ofcowpea (Vigna-unguiculata-L Walp). Plant Physiol 72: 1029–1034.


Sigel H, Operschall BP, Griesser R (2009) Xanthosine 5 '-monophosphate (XMP). Acid-base and metal ion-binding properties of achameleon-like nucleotide. Chem Soc Rev 38: 2465–2494.


Siu KKW, Lee JE, Sufrin JR, Moffatt BA, McMillan M, Cornell KA, Isom C, Howell PL (2008) Molecular determinants of substratespecificity in plant 5'-methylthioadenosine nucleosidases. J Mol Biol 378: 112–128.


Smith PM, Atkins CA (2002) Purine biosynthesis. Big in cell division, even bigger in nitrogen assimilation. Plant Physiol 128: 793–802.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Soltabayeva A, Srivastava S, Kurmanbayeva A, Bekturova A, Fluhr R, Sagi M (2018) Early senescence in older leaves of low nitrate-grown Atxdh1 uncovers a role for purine catabolism in N supply. Plant Physiol 178: 1027–1044.


Spetea C, Lundin B (2012) Evidence for nucleotide-dependent processes in the thylakoid lumen of plant chloroplasts-an update. FEBSLett 586: 2946–2954.


Stasolla C, Katahira R, Thorpe TA, Ashihara H (2003) Purine and pyrimidine nucleotide metabolism in higher plants. J Plant Physiol 160:1271–1295.


Stitt M, Lilley RM, Heldt HW (1982) Adenine-nucleotide levels in the cytosol, chloroplasts, and mitochondria of wheat leaf protoplasts.Plant Physiol 70: 971–977.


Sugimoto H, Kusumi K, Noguchi K, Yano M, Yoshimura A, Iba K (2007) The rice nuclear gene, VIRESCENT 2, is essential for chloroplastdevelopment and encodes a novel type of guanylate kinase targeted to plastids and mitochondria. Plant J 52: 512–527.


Sukrong S, Yun K-Y, Stadler P, Kumar C, Facciuolo T, Moffatt BA, Falcone DL (2012) Improved growth and stress tolerance in theArabidopsis oxt1 mutant triggered by altered adenine metabolism. Mol Plant 5: 1310–1332.


Tegeder M (2014) Transporters involved in source to sink partitioning of amino acids and ureides. Opportunities for cropimprovement. J Exp Bot 65: 1865–1878.


Tintemann H, Wasternack C, Benndorf R, Reinbothe H (1985) The rate-limiting step of uracil degradation in tomato cell-suspensioncultures and Euglena-gracilis invivo studies. Comp Biochem Physiol, Part B: Biochem Mol Biol 82: 787–792.

Pubmed: Author and Title www.plantphysiol.orgon April 24, 2020 - Published by Downloaded from Copyright © 2019 American Society of Plant Biologists. All rights reserved.

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

https://scholar.google.com/scholar?as_q=Adenosine kinase contributes to cytokinin interconversion 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=Adenosine kinase contributes to cytokinin interconversion in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schoor&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Schroeder RY%2C Zhu A%2C Eubel H%2C Dahncke K%2C Witte C%2DP %282018%29 The ribokinases of Arabidopsis thaliana and Saccharomyces cerevisiae are required for ribose recycling from nucleotide catabolism%2C which in plants is not essential to survive prolonged dark stress%2E New Phytol 217%3A 233%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The ribokinases of Arabidopsis thaliana and Saccharomyces cerevisiae are required for ribose recycling from nucleotide catabolism, which in plants is not essential to survive prolonged dark stress&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 ribokinases of Arabidopsis thaliana and Saccharomyces cerevisiae are required for ribose recycling from nucleotide catabolism, which in plants is not essential to survive prolonged dark stress&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Schroeder&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Serventi F%2C Ramazzina I%2C Lamberto I%2C Puggioni V%2C Gatti R%2C Percudani R %282010%29 Chemical basis of nitrogen recovery through the ureide pathway%2E Formation and hydrolysis of S%2Dureidoglycine in plants and bacteria%2E ACS Chem Biol 5%3A 203%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Chemical basis of nitrogen recovery through the ureide pathway&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=Chemical basis of nitrogen recovery through the ureide pathway&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Serventi&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sharma CB%2C Mittal R%2C Tanner W %281986%29 Purification and properties of a glycoprotein adenosine 5�?�%2Dmonophosphatase from the plasma membrane fraction of Arachis hypogaea cotyledons%2E Biochim Biophys Acta 884%3A 567%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purification and properties of a glycoprotein adenosine 5â�?²-monophosphatase from the plasma membrane fraction of Arachis hypogaea cotyledons&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=Purification and properties of a glycoprotein adenosine 5â�?²-monophosphatase from the plasma membrane fraction of Arachis hypogaea cotyledons&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sharma&as_ylo=1986&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Shelp BJ%2C Atkins CA %281983%29 Role of Inosine monophosphate oxidoreductase in the formation of ureides in nitrogen%2Dfixing nodules of cowpea %28Vigna%2Dunguiculata%2DL Walp%29%2E Plant Physiol 72%3A 1029%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Role of Inosine monophosphate oxidoreductase in the formation of ureides in nitrogen-fixing nodules of cowpea (Vigna-unguiculata-L Walp)&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=Role of Inosine monophosphate oxidoreductase in the formation of ureides in nitrogen-fixing nodules of cowpea (Vigna-unguiculata-L Walp)&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Shelp&as_ylo=1983&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sigel H%2C Operschall BP%2C Griesser R %282009%29 Xanthosine 5 %27%2Dmonophosphate %28XMP%29%2E Acid%2Dbase and metal ion%2Dbinding properties of a chameleon%2Dlike nucleotide%2E Chem Soc Rev 38%3A 2465%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Xanthosine 5 '-monophosphate (XMP)&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=Xanthosine 5 '-monophosphate (XMP)&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sigel&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Siu KKW%2C Lee JE%2C Sufrin JR%2C Moffatt BA%2C McMillan M%2C Cornell KA%2C Isom C%2C Howell PL %282008%29 Molecular determinants of substrate specificity in plant 5%27%2Dmethylthioadenosine nucleosidases%2E J Mol Biol 378%3A 112%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Molecular determinants of substrate specificity in plant 5'-methylthioadenosine nucleosidases&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=Molecular determinants of substrate specificity in plant 5'-methylthioadenosine nucleosidases&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Siu&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Smith PM%2C Atkins CA %282002%29 Purine biosynthesis%2E Big in cell division%2C even bigger in nitrogen assimilation%2E Plant Physiol 128%3A 793%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purine biosynthesis&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=Purine biosynthesis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Smith&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Soltabayeva A%2C Srivastava S%2C Kurmanbayeva A%2C Bekturova A%2C Fluhr R%2C Sagi M %282018%29 Early senescence in older leaves of low nitrate%2Dgrown Atxdh1 uncovers a role for purine catabolism in N supply%2E Plant Physiol 178%3A 1027%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Early senescence in older leaves of low nitrate-grown Atxdh1 uncovers a role for purine catabolism in N supply&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=Early senescence in older leaves of low nitrate-grown Atxdh1 uncovers a role for purine catabolism in N supply&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Soltabayeva&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Spetea C%2C Lundin B %282012%29 Evidence for nucleotide%2Ddependent processes in the thylakoid lumen of plant chloroplasts%2Dan update%2E FEBS Lett 586%3A 2946%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Evidence for nucleotide-dependent processes in the thylakoid lumen of plant chloroplasts-an update&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=Evidence for nucleotide-dependent processes in the thylakoid lumen of plant chloroplasts-an update&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Spetea&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Stasolla C%2C Katahira R%2C Thorpe TA%2C Ashihara H %282003%29 Purine and pyrimidine nucleotide metabolism in higher plants%2E J Plant Physiol 160%3A 1271%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Purine and pyrimidine nucleotide metabolism in higher plants&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=Purine and pyrimidine nucleotide metabolism in higher plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Stasolla&as_ylo=2003&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Stitt M%2C Lilley RM%2C Heldt HW %281982%29 Adenine%2Dnucleotide levels in the cytosol%2C chloroplasts%2C and mitochondria of wheat leaf protoplasts%2E Plant Physiol 70%3A 971%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Adenine-nucleotide levels in the cytosol, chloroplasts, and mitochondria of wheat leaf protoplasts&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=Adenine-nucleotide levels in the cytosol, chloroplasts, and mitochondria of wheat leaf protoplasts&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Stitt&as_ylo=1982&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sugimoto H%2C Kusumi K%2C Noguchi K%2C Yano M%2C Yoshimura A%2C Iba K %282007%29 The rice nuclear gene%2C VIRESCENT 2%2C is essential for chloroplast development and encodes a novel type of guanylate kinase targeted to plastids and mitochondria%2E Plant J 52%3A 512%26%23x&dopt=abstract

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


https://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sugimoto&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Sukrong S%2C Yun K%2DY%2C Stadler P%2C Kumar C%2C Facciuolo T%2C Moffatt BA%2C Falcone DL %282012%29 Improved growth and stress tolerance in the Arabidopsis oxt1 mutant triggered by altered adenine metabolism%2E Mol Plant 5%3A 1310%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Improved growth and stress tolerance in the Arabidopsis oxt1 mutant triggered by altered adenine metabolism&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Improved growth and stress tolerance in the Arabidopsis oxt1 mutant triggered by altered adenine metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Sukrong&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Tegeder M %282014%29 Transporters involved in source to sink partitioning of amino acids and ureides%2E Opportunities for crop improvement%2E J Exp Bot 65%3A 1865%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Transporters involved in source to sink partitioning of amino acids and ureides&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=Transporters involved in source to sink partitioning of amino acids and ureides&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tegeder&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Tintemann H%2C Wasternack C%2C Benndorf R%2C Reinbothe H %281985%29 The rate%2Dlimiting step of uracil degradation in tomato cell%2Dsuspension cultures and Euglena%2Dgracilis invivo studies%2E Comp Biochem Physiol%2C Part B%3A Biochem Mol Biol 82%3A 787%26%23x&dopt=abstract



Todd CD, Polacco JC (2006) AtAAH encodes a protein with allantoate amidohydrolase activity from Arabidopsis thaliana. Planta 223:1108–1113.


Torres RJ, Puig JG (2007) Hypoxanthine-guanine phosophoribosyltransferase (HPRT) deficiency: Lesch-Nyhan syndrome. Orphanet JRare Dis. 2: 48.


Traub M, Florchinger M, Piecuch J, Kunz HH, Weise-Steinmetz A, Deitmer JW, Neuhaus HE, Mohlmann T (2007) The fluorouridineinsensitive 1 (fur1) mutant is defective in equilibrative nucleoside transporter 3 (ENT3), and thus represents an important pyrimidinenucleoside uptake system in Arabidopsis thaliana. Plant J 49: 855–864.


Tripathi D, Zhang T, Koo AJ, Stacey G, Tanaka K (2017) Extracellular ATP acts on jasmonate signaling to reinforce plant defense. PlantPhysiol 176: 511–523.


Ullrich A, Knecht W, Piskur J, Loffler M (2002) Plant dihydroorotate dehydrogenase differs significantly in substrate specificity andinhibition from the animal enzymes. FEBS Lett 529: 346–350.


Urarte E, Esteban R, Moran JF, Bittner F (2015) Established and proposed roles of xanthine oxidoreductase in oxidative and reductivepathways in plants. In KJ Gupta, AU Igamberdiev, eds, Reactive oxygen and nitrogen species signaling and communication in plants.Springer, Cham, Switzerland, pp. 15–42.


Wagner KG, Backer AI (1992) Dynamics of nucleotides in plants studied on a cellular basis. In JK W, F M, eds, International Review ofCytology Vol. 134, pp. 1–84.


Walsh TA, Green SB, Larrinua IM, Schmitzer PR (2001) Characterization of plant beta-ureidopropionase and functional overexpressionin Escherichia coli. Plant Physiol 125: 1001–1011.


Wang L, Li Z, Qian W, Guo W, Gao X, Huang L, Wang H, Zhu H, Wu JW,Wang D, Liu D (2011) The Arabidopsis purple acid phosphataseAt-PAP10 is predominantly associated with the root surface and plays an important role in plant tolerance to phosphate limitation. PlantPhysiol 157: 1283–1299.


Wang C, Liu Z (2006) Arabidopsis ribonucleotide reductases are critical for cell cycle progression, DNA damage repair, and plantdevelopment. Plant Cell 18: 350–365.


Wang C, Zhou M, Zhang X, Yao J, Zhang Y, Mou Z (2017) A lectin receptor kinase as a potential sensor for extracellular nicotinamideadenine dinucleotide in Arabidopsis thaliana. eLife 6: e25474


Watanabe S, Matsumoto M, Hakomori Y, Takagi H, Shimada H, Sakamoto A (2014) The purine metabolite allantoin enhances abioticstress tolerance through synergistic activation of abscisic acid metabolism. Plant Cell Environ 37: 1022–1036.


Wei X, Song X, Wei L, Tang S, Sun J, Hu P, Cao X (2017) An epiallele of rice AK1 affects photosynthetic capacity. J Integr Plant Biol 59:158–163.


Werner AK, Medina-Escobar N, Zulawski M, Sparkes IA, Cao F-Q, Witte CP (2013) The ureide-degrading reactions of purine ringcatabolism employ three amidohydrolases and one aminohydrolase in Arabidopsis, soybean, and rice. Plant Physiol 163: 672–681.



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

https://scholar.google.com/scholar?as_q=The rate-limiting step of uracil degradation in tomato cell-suspension cultures and Euglena-gracilis invivo studies&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 rate-limiting step of uracil degradation in tomato cell-suspension cultures and Euglena-gracilis invivo studies&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tintemann&as_ylo=1985&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Todd CD%2C Polacco JC %282006%29 AtAAH encodes a protein with allantoate amidohydrolase activity from Arabidopsis thaliana%2E Planta 223%3A 1108%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=AtAAH encodes a protein with allantoate amidohydrolase activity from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=AtAAH encodes a protein with allantoate amidohydrolase activity from Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Todd&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Torres RJ%2C Puig JG %282007%29 Hypoxanthine%2Dguanine phosophoribosyltransferase %28HPRT%29 deficiency%3A Lesch%2DNyhan syndrome%2E Orphanet J Rare Dis&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Hypoxanthine-guanine phosophoribosyltransferase (HPRT) deficiency: Lesch-Nyhan syndrome&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=Hypoxanthine-guanine phosophoribosyltransferase (HPRT) deficiency: Lesch-Nyhan syndrome&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Torres&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Traub M%2C Florchinger M%2C Piecuch J%2C Kunz HH%2C Weise%2DSteinmetz A%2C Deitmer JW%2C Neuhaus HE%2C Mohlmann T %282007%29 The fluorouridine insensitive 1 %28fur1%29 mutant is defective in equilibrative nucleoside transporter 3 %28ENT3%29%2C and thus represents an important pyrimidine nucleoside uptake system in Arabidopsis thaliana%2E Plant J 49%3A 855%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The fluorouridine insensitive 1 (fur1) mutant is defective in equilibrative nucleoside transporter 3 (ENT3), and thus represents an important pyrimidine nucleoside uptake system in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=The fluorouridine insensitive 1 (fur1) mutant is defective in equilibrative nucleoside transporter 3 (ENT3), and thus represents an important pyrimidine nucleoside uptake system in Arabidopsis thaliana&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Traub&as_ylo=2007&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Tripathi D%2C Zhang T%2C Koo AJ%2C Stacey G%2C Tanaka K %282017%29 Extracellular ATP acts on jasmonate signaling to reinforce plant defense%2E Plant Physiol 176%3A 511%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Extracellular ATP acts on jasmonate signaling to reinforce plant defense&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=Extracellular ATP acts on jasmonate signaling to reinforce plant defense&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Tripathi&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ullrich A%2C Knecht W%2C Piskur J%2C Loffler M %282002%29 Plant dihydroorotate dehydrogenase differs significantly in substrate specificity and inhibition from the animal enzymes%2E FEBS Lett 529%3A 346%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Plant dihydroorotate dehydrogenase differs significantly in substrate specificity and inhibition from the animal enzymes&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=Plant dihydroorotate dehydrogenase differs significantly in substrate specificity and inhibition from the animal enzymes&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ullrich&as_ylo=2002&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Urarte E%2C Esteban R%2C Moran JF%2C Bittner F %282015%29 Established and proposed roles of xanthine oxidoreductase in oxidative and reductive pathways in plants%2E In KJ Gupta%2C AU Igamberdiev%2C eds%2C Reactive oxygen and nitrogen species signaling and communication in plants%2E Springer%2C Cham%2C Switzerland%2C pp%2E 15%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Established and proposed roles of xanthine oxidoreductase in oxidative and reductive pathways in plants.&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=Established and proposed roles of xanthine oxidoreductase in oxidative and reductive pathways in plants.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Urarte&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wagner KG%2C Backer AI %281992%29 Dynamics of nucleotides in plants studied on a cellular basis%2E In JK W%2C F M%2C eds%2C International Review of Cytology Vol%2E 134%2C pp%2E 1%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Dynamics of nucleotides in plants studied on a cellular basis&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=Dynamics of nucleotides in plants studied on a cellular basis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wagner&as_ylo=1992&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Walsh TA%2C Green SB%2C Larrinua IM%2C Schmitzer PR %282001%29 Characterization of plant beta%2Dureidopropionase and functional overexpression in Escherichia coli%2E Plant Physiol 125%3A 1001%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Characterization of plant beta-ureidopropionase and functional overexpression in Escherichia coli&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=Characterization of plant beta-ureidopropionase and functional overexpression in Escherichia coli&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Walsh&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wang L%2C Li Z%2C Qian W%2C Guo W%2C Gao X%2C Huang L%2C Wang H%2C Zhu H%2C Wu JW%2CWang D%2C Liu D %282011%29 The Arabidopsis purple acid phosphatase At%2DPAP10 is predominantly associated with the root surface and plays an important role in plant tolerance to phosphate limitation%2E Plant Physiol 157%3A 1283%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The Arabidopsis purple acid phosphatase At-PAP10 is predominantly associated with the root surface and plays an important role in plant tolerance to phosphate limitation&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=The Arabidopsis purple acid phosphatase At-PAP10 is predominantly associated with the root surface and plays an important role in plant tolerance to phosphate limitation&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wang&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wang C%2C Liu Z %282006%29 Arabidopsis ribonucleotide reductases are critical for cell cycle progression%2C DNA damage repair%2C and plant development%2E Plant Cell 18%3A 350%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Arabidopsis ribonucleotide reductases are critical for cell cycle progression, DNA damage repair, and 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=Arabidopsis ribonucleotide reductases are critical for cell cycle progression, DNA damage repair, and plant development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wang&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wang C%2C Zhou M%2C Zhang X%2C Yao J%2C Zhang Y%2C Mou Z %282017%29 A lectin receptor kinase as a potential sensor for extracellular nicotinamide adenine dinucleotide in Arabidopsis thaliana%2E eLife 6%3A e&dopt=abstract


https://scholar.google.com/scholar?as_q=A lectin receptor kinase as a potential sensor for extracellular nicotinamide adenine dinucleotide in Arabidopsis thaliana.&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=A lectin receptor kinase as a potential sensor for extracellular nicotinamide adenine dinucleotide in Arabidopsis thaliana.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wang&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Watanabe S%2C Matsumoto M%2C Hakomori Y%2C Takagi H%2C Shimada H%2C Sakamoto A %282014%29 The purine metabolite allantoin enhances abiotic stress tolerance through synergistic activation of abscisic acid metabolism%2E Plant Cell Environ 37%3A 1022%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The purine metabolite allantoin enhances abiotic stress tolerance through synergistic activation of abscisic acid metabolism&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 purine metabolite allantoin enhances abiotic stress tolerance through synergistic activation of abscisic acid metabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Watanabe&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wei X%2C Song X%2C Wei L%2C Tang S%2C Sun J%2C Hu P%2C Cao X %282017%29 An epiallele of rice AK1 affects photosynthetic capacity%2E J Integr Plant Biol 59%3A 158%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=An epiallele of rice AK1 affects photosynthetic capacity&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=An epiallele of rice AK1 affects photosynthetic capacity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wei&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Werner AK%2C Medina%2DEscobar N%2C Zulawski M%2C Sparkes IA%2C Cao F%2DQ%2C Witte CP %282013%29 The ureide%2Ddegrading reactions of purine ring catabolism employ three amidohydrolases and one aminohydrolase in Arabidopsis%2C soybean%2C and rice%2E Plant Physiol 163%3A 672%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The ureide-degrading reactions of purine ring catabolism employ three amidohydrolases and one aminohydrolase in Arabidopsis, soybean, and rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=The ureide-degrading reactions of purine ring catabolism employ three amidohydrolases and one aminohydrolase in Arabidopsis, soybean, and rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Werner&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1


Werner AK, Romeis T, Witte CP (2010) Ureide catabolism in Arabidopsis thaliana and Escherichia coli. Nat Chem Biol 6: 19–21.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Werner AK, Sparkes IA, Romeis T, Witte CP (2008) Identification, biochemical characterization, and subcellular localization of allantoateamidohydrolases from Arabidopsis and soybean. Plant Physiol 146: 418–430.


Werner AK, Witte CP (2011) The biochemistry of nitrogen mobilization: purine ring catabolism. Trends Plant Sci 16: 381–387.Pubmed: Author and TitleGoogle Scholar: Author Only Title Only Author and Title

Witz S, Jung B, Furst S, Mohlmann T (2012) De novo pyrimidine nucleotide synthesis mainly occurs outside of plastids, but a previouslyundiscovered nucleobase importer provides substrates for the essential salvage pathway in Arabidopsis. Plant Cell 24: 1549–1559.


Wu S, Alseekh S, Cuadros-Inostroza A, Fusari CM, Mutwil M, Kooke R, Keurentjes JB, Fernie AR, Willmitzer L, Brotman Y (2016)Combined use of genome-wide association data and correlation networks unravels key regulators of primary metabolism inArabidopsis thaliana. PLoS GENET 12.


Xu J, Deng Y, Li Q, Zhu X, He Z (2014) STRIPE2 encodes a putative dCMP deaminase that plays an important role in chloroplastdevelopment in rice. J Genet Genomics 41: 539–548.


Xu J, Zhang HY, Xie CH, Xue HW, Dijkhuis P, Liu CM (2005) EMBRYONIC FACTOR 1 encodes an AMP deaminase and is essential forthe zygote to embryo transition in Arabidopsis. Plant J 42: 743–756.


Xu J, Zhang L, Yang D-L, Li Q, He Z (2015) Thymidine kinases share a conserved function for nucleotide salvage and play an essentialrole in Arabidopsis thaliana growth and development. New Phytol 208: 1089–1103.


Yamada H, Suzuki T, Terada K, Takei K, Ishikawa K, Miwa K, Yamashino T, Mizuno T (2001) The Arabidopsis AHK4 histidine kinase is acytokinin-binding receptor that transduces cytokinin signals across the membrane. Plant Cell Physiol 42: 1017–1023.


Ye W, Hu S, Wu L, Ge C, Cui Y, Chen P, Wang X, Xu J, Ren D, Dong G, Qian Q, Guo L (2016) White stripe leaf 12 (WSL12), encoding anucleoside diphosphate kinase 2 (OsNDPK2), regulates chloroplast development and abiotic stress response in rice (Oryza sativa L.).Mol Breeding 36: 57.


Yin Y, Katahira R, Ashihara H (2014) Metabolism of purine nucleosides and bases in suspension-cultured Arabidopsis thaliana cells.Eur Chem Bull 3: 925–934.


Young LS, Harrison BR, Narayana, M. U. M., Moffatt BA, Gilroy S, Masson PH (2006) Adenosine kinase modulates root gravitropism andcap morphogenesis in arabidopsis. Plant Physiol 142: 564–573.


Zarepour M, Kaspari K, Stagge S, Rethmeier R, Mendel RR, Bittner F (2010) Xanthine dehydrogenase AtXDH1 from Arabidopsisthaliana is a potent producer of superoxide anions via its NADH oxidase activity. Plant Mol Biol 72: 301–310.


Zhang T, Feng P, Li Y, Yu P, Yu G, Sang X, Ling Y, Zeng X, Li Y, Huang J, Zhang T, Zhao F, Wang N, Zhang C, Yang Z, Wu R, He G (2018)VIRESCENT-ALBINO LEAF 1 regulates leaf colour development and cell division in rice. J Exp Bot 69: 4791–4804.


Zhang X, Chen Y, Lin X, Hong X, Zhu Y, Li W, He W, An F, Guo H (2013) Adenine phosphoribosyl transferase 1 is a key enzymecatalyzing cytokinin conversion from nucleobases to nucleotides in Arabidopsis. Mol Plant 6: 1661–1672.



https://www.ncbi.nlm.nih.gov/pubmed?term=Werner AK%2C Romeis T%2C Witte CP %282010%29 Ureide catabolism in Arabidopsis thaliana and Escherichia coli%2E Nat Chem Biol 6%3A 19%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Ureide catabolism in Arabidopsis thaliana and Escherichia coli&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=Ureide catabolism in Arabidopsis thaliana and Escherichia coli&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Werner&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Werner AK%2C Sparkes IA%2C Romeis T%2C Witte CP %282008%29 Identification%2C biochemical characterization%2C and subcellular localization of allantoate amidohydrolases from Arabidopsis and soybean%2E Plant Physiol 146%3A 418%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Identification, biochemical characterization, and subcellular localization of allantoate amidohydrolases from Arabidopsis and soybean&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=Identification, biochemical characterization, and subcellular localization of allantoate amidohydrolases from Arabidopsis and soybean&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Werner&as_ylo=2008&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Werner AK%2C Witte CP %282011%29 The biochemistry of nitrogen mobilization%3A purine ring catabolism%2E Trends Plant Sci 16%3A 381%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=The biochemistry of nitrogen mobilization: purine ring catabolism&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 biochemistry of nitrogen mobilization: purine ring catabolism&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Werner&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Witz S%2C Jung B%2C Furst S%2C Mohlmann T %282012%29 De novo pyrimidine nucleotide synthesis mainly occurs outside of plastids%2C but a previously undiscovered nucleobase importer provides substrates for the essential salvage pathway in Arabidopsis%2E Plant Cell 24%3A 1549%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=De novo pyrimidine nucleotide synthesis mainly occurs outside of plastids, but a previously undiscovered nucleobase importer provides substrates for the essential salvage pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=De novo pyrimidine nucleotide synthesis mainly occurs outside of plastids, but a previously undiscovered nucleobase importer provides substrates for the essential salvage pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Witz&as_ylo=2012&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Wu S%2C Alseekh S%2C Cuadros%2DInostroza A%2C Fusari CM%2C Mutwil M%2C Kooke R%2C Keurentjes JB%2C Fernie AR%2C Willmitzer L%2C Brotman Y %282016%29 Combined use of genome%2Dwide association data and correlation networks unravels key regulators of primary metabolism in Arabidopsis thaliana%2E PLoS GENET&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Combined use of genome-wide association data and correlation networks unravels key regulators of primary metabolism in Arabidopsis thaliana.&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Combined use of genome-wide association data and correlation networks unravels key regulators of primary metabolism in Arabidopsis thaliana.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Wu&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Xu J%2C Deng Y%2C Li Q%2C Zhu X%2C He Z %282014%29 STRIPE2 encodes a putative dCMP deaminase that plays an important role in chloroplast development in rice%2E J Genet Genomics 41%3A 539%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=STRIPE2 encodes a putative dCMP deaminase that plays an important role in chloroplast development in rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=STRIPE2 encodes a putative dCMP deaminase that plays an important role in chloroplast development in rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Xu&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Xu J%2C Zhang HY%2C Xie CH%2C Xue HW%2C Dijkhuis P%2C Liu CM %282005%29 EMBRYONIC FACTOR 1 encodes an AMP deaminase and is essential for the zygote to embryo transition in Arabidopsis%2E Plant J 42%3A 743%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=EMBRYONIC FACTOR 1 encodes an AMP deaminase and is essential for the zygote to embryo transition in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=EMBRYONIC FACTOR 1 encodes an AMP deaminase and is essential for the zygote to embryo transition in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Xu&as_ylo=2005&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Xu J%2C Zhang L%2C Yang D%2DL%2C Li Q%2C He Z %282015%29 Thymidine kinases share a conserved function for nucleotide salvage and play an essential role in Arabidopsis thaliana growth and development%2E New Phytol 208%3A 1089%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Thymidine kinases share a conserved function for nucleotide salvage and play an essential role in Arabidopsis thaliana growth and development&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Thymidine kinases share a conserved function for nucleotide salvage and play an essential role in Arabidopsis thaliana growth and development&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Xu&as_ylo=2015&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Yamada H%2C Suzuki T%2C Terada K%2C Takei K%2C Ishikawa K%2C Miwa K%2C Yamashino T%2C Mizuno T %282001%29 The Arabidopsis AHK4 histidine kinase is a cytokinin%2Dbinding receptor that transduces cytokinin signals across the membrane%2E Plant Cell Physiol 42%3A 1017%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=The Arabidopsis AHK4 histidine kinase is a cytokinin-binding receptor that transduces cytokinin signals across the membrane&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=The Arabidopsis AHK4 histidine kinase is a cytokinin-binding receptor that transduces cytokinin signals across the membrane&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Yamada&as_ylo=2001&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Ye W%2C Hu S%2C Wu L%2C Ge C%2C Cui Y%2C Chen P%2C Wang X%2C Xu J%2C Ren D%2C Dong G%2C Qian Q%2C Guo L %282016%29 White stripe leaf 12 %28WSL12%29%2C encoding a nucleoside diphosphate kinase 2 %28OsNDPK2%29%2C regulates chloroplast development and abiotic stress response in rice %28Oryza sativa L%2E%29%2E Mol Breeding&dopt=abstract

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

https://scholar.google.com/scholar?as_q=White stripe leaf 12 (WSL12), encoding a nucleoside diphosphate kinase 2 (OsNDPK2), regulates chloroplast development and abiotic stress response in rice (Oryza sativa L&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=White stripe leaf 12 (WSL12), encoding a nucleoside diphosphate kinase 2 (OsNDPK2), regulates chloroplast development and abiotic stress response in rice (Oryza sativa L&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Ye&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Yin Y%2C Katahira R%2C Ashihara H %282014%29 Metabolism of purine nucleosides and bases in suspension%2Dcultured Arabidopsis thaliana cells%2E Eur Chem Bull 3%3A 925%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Metabolism of purine nucleosides and bases in suspension-cultured Arabidopsis thaliana 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=Metabolism of purine nucleosides and bases in suspension-cultured Arabidopsis thaliana cells&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Yin&as_ylo=2014&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Young LS%2C Harrison BR%2C Narayana%2C M%2E U%2E M%2E%2C Moffatt BA%2C Gilroy S%2C Masson PH %282006%29 Adenosine kinase modulates root gravitropism and cap morphogenesis in arabidopsis%2E Plant Physiol 142%3A 564%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Adenosine kinase modulates root gravitropism and cap morphogenesis 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=Adenosine kinase modulates root gravitropism and cap morphogenesis in arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Young&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zarepour M%2C Kaspari K%2C Stagge S%2C Rethmeier R%2C Mendel RR%2C Bittner F %282010%29 Xanthine dehydrogenase AtXDH1 from Arabidopsis thaliana is a potent producer of superoxide anions via its NADH oxidase activity%2E Plant Mol Biol 72%3A 301%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Xanthine dehydrogenase AtXDH1 from Arabidopsis thaliana is a potent producer of superoxide anions via its NADH oxidase activity&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Xanthine dehydrogenase AtXDH1 from Arabidopsis thaliana is a potent producer of superoxide anions via its NADH oxidase activity&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zarepour&as_ylo=2010&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zhang T%2C Feng P%2C Li Y%2C Yu P%2C Yu G%2C Sang X%2C Ling Y%2C Zeng X%2C Li Y%2C Huang J%2C Zhang T%2C Zhao F%2C Wang N%2C Zhang C%2C Yang Z%2C Wu R%2C He G %282018%29 VIRESCENT%2DALBINO LEAF 1 regulates leaf colour development and cell division in rice%2E J Exp Bot 69%3A 4791%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=VIRESCENT-ALBINO LEAF 1 regulates leaf colour development and cell division in rice&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=VIRESCENT-ALBINO LEAF 1 regulates leaf colour development and cell division in rice&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhang&as_ylo=2018&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zhang X%2C Chen Y%2C Lin X%2C Hong X%2C Zhu Y%2C Li W%2C He W%2C An F%2C Guo H %282013%29 Adenine phosphoribosyl transferase 1 is a key enzyme catalyzing cytokinin conversion from nucleobases to nucleotides in Arabidopsis%2E Mol Plant 6%3A 1661%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Adenine phosphoribosyl transferase 1 is a key enzyme catalyzing cytokinin conversion from nucleobases to nucleotides 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=Adenine phosphoribosyl transferase 1 is a key enzyme catalyzing cytokinin conversion from nucleobases to nucleotides in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhang&as_ylo=2013&as_allsubj=all&hl=en&c2coff=1


Zhou K, Xia J, Wang Y, Ma T, Li Z (2017) A Young Seedling Stripe2 phenotype in rice is caused by mutation of a chloroplast-localizednucleoside diphosphate kinase 2 required for chloroplast biogenesis. Genet Mol Biol 40: 630–642.


Zhou L, Lacroute F, Thornburg R (1998) Cloning, expression in Escherichia coli, and characterization of Arabidopsis thaliana UMP/CMPkinase. Plant Physiol 117: 245–254.


Zhu X, Guo S, Wang Z, Du Q, Xing Y, Zhang T, Shen W, Sang X, Ling Y, He G (2016) Map-based cloning and functional analysis of YGL8,which controls leaf colour in rice (Oryza sativa). BMC Plant Biol 16: 134.


Zrenner R, Ashihara H (2011) Nucleotide Metabolism. In H Ashihara, A Crozier, A Komamine, eds, Plant metabolism and biotechnology.Wiley, Cambridge, New York, pp. 135–162.


Zrenner R, Riegler H, Marquard CR, Lange PR, Geserick C, Bartosz CE, Chen CT, Slocum RD (2009) A functional analysis of thepyrimidine catabolic pathway in Arabidopsis. New Phytol 183: 117–132.


Zrenner R, Stitt M, Sonnewald U, Boldt R (2006) Pyrimidine and purine biosynthesis and degradation in plants. Annu Rev Plant Biol 57:805–836.



https://www.ncbi.nlm.nih.gov/pubmed?term=Zhou K%2C Xia J%2C Wang Y%2C Ma T%2C Li Z %282017%29 A Young Seedling Stripe2 phenotype in rice is caused by mutation of a chloroplast%2Dlocalized nucleoside diphosphate kinase 2 required for chloroplast biogenesis%2E Genet Mol Biol 40%3A 630%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=A Young Seedling Stripe2 phenotype in rice is caused by mutation of a chloroplast-localized nucleoside diphosphate kinase 2 required for chloroplast biogenesis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=A Young Seedling Stripe2 phenotype in rice is caused by mutation of a chloroplast-localized nucleoside diphosphate kinase 2 required for chloroplast biogenesis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhou&as_ylo=2017&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zhou L%2C Lacroute F%2C Thornburg R %281998%29 Cloning%2C expression in Escherichia coli%2C and characterization of Arabidopsis thaliana UMP%2FCMP kinase%2E Plant Physiol 117%3A 245%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Cloning, expression in Escherichia coli, and characterization of Arabidopsis thaliana UMP/CMP kinase&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=Cloning, expression in Escherichia coli, and characterization of Arabidopsis thaliana UMP/CMP kinase&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhou&as_ylo=1998&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zhu X%2C Guo S%2C Wang Z%2C Du Q%2C Xing Y%2C Zhang T%2C Shen W%2C Sang X%2C Ling Y%2C He G %282016%29 Map%2Dbased cloning and functional analysis of YGL8%2C which controls leaf colour in rice %28Oryza sativa%29%2E BMC Plant Biol&dopt=abstract

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


https://scholar.google.com/scholar?as_q=&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zhu&as_ylo=2016&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zrenner R%2C Ashihara H %282011%29 Nucleotide Metabolism%2E In H Ashihara%2C A Crozier%2C A Komamine%2C eds%2C Plant metabolism and biotechnology%2E Wiley%2C Cambridge%2C New York%2C pp%2E 135%26%23x&dopt=abstract

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

https://scholar.google.com/scholar?as_q=Nucleotide Metabolism.&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=Nucleotide Metabolism.&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zrenner&as_ylo=2011&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zrenner R%2C Riegler H%2C Marquard CR%2C Lange PR%2C Geserick C%2C Bartosz CE%2C Chen CT%2C Slocum RD %282009%29 A functional analysis of the pyrimidine catabolic pathway in Arabidopsis%2E New Phytol 183%3A 117%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=A functional analysis of the pyrimidine catabolic pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_publication=&as_yhi=&as_allsubj=all&hl=en&lr=&c2coff=1

https://scholar.google.com/scholar?as_q=A functional analysis of the pyrimidine catabolic pathway in Arabidopsis&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zrenner&as_ylo=2009&as_allsubj=all&hl=en&c2coff=1

https://www.ncbi.nlm.nih.gov/pubmed?term=Zrenner R%2C Stitt M%2C Sonnewald U%2C Boldt R %282006%29 Pyrimidine and purine biosynthesis and degradation in plants%2E Annu Rev Plant Biol 57%3A 805%26%23x&dopt=abstract


https://scholar.google.com/scholar?as_q=Pyrimidine and purine biosynthesis and degradation in plants&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=Pyrimidine and purine biosynthesis and degradation in plants&num=10&btnG=Search+Scholar&as_occt=any&as_sauthors=Zrenner&as_ylo=2006&as_allsubj=all&hl=en&c2coff=1


nucleotide metabolism in plants - plant physiology1 1 author for contact: claus-peter witte 1,...

Documents