336:
321:
403:. The steady state levels of 8-oxoguanine in DNA of several organs including the brain are significantly reduced in hMTH1-Tg over-expressing mice. Conversely, MTH1-null mice exhibit a significantly higher level of 8-oxo-dGTP accumulation than that of the wild type. Over-expression of hMTH1 prevents the age-dependent accumulation of DNA 8-oxoguanine that occurs in wild-type mice. The lower levels of oxidized guanines are associated with greater
2099:
3329:
332:, which is specific to 8-oxoguanine nucleotides. Human MTH1, however, has substrate specificity for a much broader range of oxidatively damaged nucleotides. The mechanism of hMTH1's broad specificity for these oxidized nucleotides is derived from their recognition in the enzyme's substrate binding pocket due to an exchange of protonation state between two nearby aspartate residues.
443:
to other human cells. Because a cancer cell divides much more rapidly than a normal human cell, it is far more in need of an enzyme like MTH1 that prevents fatal mutations during replication. This property of cancer cells could allow for monitoring of cancer treatment efficacy by measuring MTH1 expression. Development of suitable probes for this purpose is currently underway.
451:
older MTH1 inhibitors may be due to off-target cytotoxic effects. After revisiting the experiment, the original authors of this claim found that while the original MTH1 inhibitors in question lead to damaged nucleotides being incorporated into DNA, they demonstrate the others that do not induce toxicity fail to introduce the DNA lesion. Research into this topic is ongoing.
450:
or antiproliferation effects of MTH1 inhibition on cancer cells, even calling into question whether MTH1 truly does serve to remove oxidatively damaged nucleotides from a cell's nucleotide pool. One study of newly discovered MTH1 inhibitors suggests that these anticancer properties exhibited by the
442:
in cancer cells leads to incorporation of 8-oxo-dGTP and other oxidatively damaged nucleotides into the cell's DNA, damaging it and causing cell death. However, cancer cells have also been shown to benefit from use of MTH1. Cells from malignant breast tumors exhibit extreme MTH1 expression compared
1008:
Coskun E, Jaruga P, Jemth AS, Loseva O, Scanlan LD, Tona A, Lowenthal MS, Helleday T, Dizdaroglu M (September 2015). "Addiction to MTH1 protein results in intense expression in human breast cancer tissue as measured by liquid chromatography-isotope-dilution tandem mass spectrometry".
607:
Rudling, Axel; Gustafsson, Robert; Almlöf, Ingrid; Homan, Evert; Scobie, Martin; Warpman
Berglund, Ulrika; Helleday, Thomas; Stenmark, Pål; Carlsson, Jens (2017-10-12). "Fragment-Based Discovery and Optimization of Enzyme Inhibitors by Docking of Commercial Chemical Space".
1120:
Petrocchi A, Leo E, Reyna NJ, Hamilton MM, Shi X, Parker CA, Mseeh F, Bardenhagen JP, Leonard P, Cross JB, Huang S, Jiang Y, Cardozo M, Draetta G, Marszalek JR, Toniatti C, Jones P, Lewis RT (March 2016). "Identification of potent and selective MTH1 inhibitors".
1742:
Takahashi M, Maraboeuf F, Sakai Y, Yakushiji H, Mishima M, Shirakawa M, Iwai S, Hayakawa H, Sekiguchi M, Nakabeppu Y (May 2002). "Role of tryptophan residues in the recognition of mutagenic oxidized nucleotides by human antimutator MTH1 protein".
1499:
Furuichi M, Yoshida MC, Oda H, Tajiri T, Nakabeppu Y, Tsuzuki T, Sekiguchi M (December 1994). "Genomic structure and chromosome location of the human mutT homologue gene MTH1 encoding 8-oxo-dGTPase for prevention of A:T to C:G transversion".
200:, also known as oxidized purine nucleoside triphosphatase, or (2'-deoxy) ribonucleoside 5'-triphosphate pyrophosphohydrolase, or Nudix hydrolase 1 (NUDT1), or MutT homolog 1 (MTH1), or 7,8-dihydro-8-oxoguanine triphosphatase) is an
1951:
Bialkowski K, Kasprzak KS (September 2003). "Inhibition of 8-oxo-2'-deoxyguanosine 5'-triphosphate pyrophosphohydrolase (8-oxo-dGTPase) activity of the antimutagenic human MTH1 protein by nucleoside 5'-diphosphates".
473:
has been made available as a chemical probe to academic researchers. However AstraZeneca has found that neither AZ13792138 nor genetic knockdown of MTH1 displays any significant cytotoxicity to cancer cells.
1712:
Topp H, Armbrust S, Lengger C, Schöch G, Davies J, Stichler W, Manz F, Fusch C (June 2002). "Renal excretion of 8-oxo-7,8-dihydro-2(')-deoxyguanosine: degradation rates of RNA and metabolic rate in humans".
1197:
Kettle JG, Alwan H, Bista M, Breed J, Davies NL, Eckersley K, Fillery S, Foote KM, Goodwin L, Jones DR, Käck H, Lau A, Nissink JW, Read J, Scott JS, Taylor B, Walker G, Wissler L, Wylot M (March 2016).
1335:
1981:
Kamiya H, Yakushiji H, Dugué L, Tanimoto M, Pochet S, Nakabeppu Y, Harashima H (February 2004). "Probing the substrate recognition mechanism of the human MTH1 protein by nucleotide analogs".
351:, suggesting that it is involved in the sanitization of nucleotide pools both for nuclear and mitochondrial genomes. In plants, MTH1 has also been shown to enhance resistance to heat- and
1852:
Kennedy CH, Pass HI, Mitchell JB (June 2003). "Expression of human MutT homologue (hMTH1) protein in primary non-small-cell lung carcinomas and histologically normal surrounding tissue".
1772:
Liu Z, Wang LE, Strom SS, Spitz MR, Babaian RJ, DiGiovanni J, Wei Q (March 2003). "Overexpression of hMTH in peripheral lymphocytes and risk of prostate cancer: a case-control analysis".
808:
De Luca G, Ventura I, Sanghez V, Russo MT, Ajmone-Cat MA, Cacci E, Martire A, Popoli P, Falcone G, Michelini F, Crescenzi M, Degan P, Minghetti L, Bignami M, Calamandrei G (August 2013).
851:"MTH1, an oxidized purine nucleoside triphosphatase, suppresses the accumulation of oxidative damage of nucleic acids in the hippocampal microglia during kainate-induced excitotoxicity"
898:
Tsuzuki T, Egashira A, Igarashi H, Iwakuma T, Nakatsuru Y, Tominaga Y, Kawate H, Nakao K, Nakamura K, Ide F, Kura S, Nakabeppu Y, Katsuki M, Ishikawa T, Sekiguchi M (September 2001).
438:. The enzyme's much-studied ability to sanitize a cell's nucleotide pool prevents it from developing mutations, including cancerous ones. Specifically, another study found that MTH1
427:
Studies have suggested that this enzyme plays a role in both preventing the formation of cancer cells and the proliferation of cancer cells. This makes it a topic of interest in
252:
2156:
446:
Disagreement exists concerning MTH1's functionality relative to prevention of DNA damage and cancer. Subsequent studies have had difficulty reproducing previously reported
211:. During DNA repair, the enzyme hydrolyses oxidized purines and prevents their addition onto the DNA chain. As such it has important role in aging and cancer development.
2011:
1093:
Samaranayake GJ, Troccoli CI, Huynh MQ, Win A, Ji D, Kool ET, Rai P (July 2017). "Towards a better understanding of MTH1 as a therapeutic target in RAS-driven cancer ".
957:
Gad H, Koolmeister T, Jemth AS, Eshtad S, Jacques SA, Ström CE, et al. (April 2014). "MTH1 inhibition eradicates cancer by preventing sanitation of the dNTP pool".
2036:
154:
317:
nucleoside triphosphates, such as 8-oxo-dGTP, 8-oxo-dATP, 2-oxo-dATP, 2-hydroxy-dATP, and 2-hydroxy rATP, to monophosphates, thereby preventing misincorporation.
173:
2098:
459:
MTH1 is a potential drug target to treat cancer, however there are conflicting results regarding the cytotoxicity of MTH1 inhibitors toward cancer cells.
2849:
2149:
2379:
339:
Examples of the protonation state exchange between MTH1 residues Asp-119 and Asp-120 that allows the enzyme to recognize oxidized nucleotides.
2078:
2348:
2142:
3048:
407:. The hMTH1-Tg animals have a significantly longer lifespan than their wild-type littermates. These findings provide a link between
766:"Ectopic expression of the human MutT-type Nudix hydrolase, hMTH1, confers enhanced tolerance to oxidative stress in arabidopsis"
593:
166:
328:
MutT enzymes in non-human organisms often have substrate specificity for certain types of oxidized nucleotides, such as that of
2939:
2019:
713:
Waz S, Nakamura T, Hirata K, Koga-Ogawa Y, Chirifu M, Arimori T, Tamada T, Ikemizu S, Nakabeppu Y, Yamagata Y (February 2017).
1531:"Intracellular localization of 8-oxo-dGTPase in human cells, with special reference to the role of the enzyme in mitochondria"
2934:
371:
269:. The enzyme is used in DNA repair, where it hydrolysis the oxidized purines and prevents their addition onto the DNA chain.
133:
109:
3204:
810:"Prolonged lifespan with enhanced exploratory behavior in mice overexpressing the oxidized nucleoside triphosphatase hMTH1"
715:"Structural and Kinetic Studies of the Human Nudix Hydrolase MTH1 Reveal the Mechanism for Its Broad Substrate Specificity"
2956:
1601:"Multi-forms of human MTH1 polypeptides produced by alternative translation initiation and single nucleotide polymorphism"
1336:"The Helleday Team presents potent MTH1 inhibitor 'Karonudib' and experimentally addresses the non-active MTH1 inhibitors"
370:, have been identified. Additional variants have been observed, but their full-length natures have not been determined. A
3004:
2321:
1566:"Regulation of expression of the human MTH1 gene encoding 8-oxo-dGTPase. Alternative splicing of transcription products"
431:, both as a potential method for healthy cells to prevent cancer and a weakness to target within existing cancer cells.
387:
A mouse model has been studied that over-expresses hMTH1-Tg (NUDT1). The hMTH1-Tg mice express high levels of the hMTH1
551:"Cloning and expression of cDNA for a human enzyme that hydrolyzes 8-oxo-dGTP, a mutagenic substrate for DNA synthesis"
2926:
1599:
Oda H, Taketomi A, Maruyama R, Itoh R, Nishioka K, Yakushiji H, Suzuki T, Sekiguchi M, Nakabeppu Y (November 1999).
3319:
416:
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3240:
3227:
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2526:
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2195:
2182:
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1046:"A Chimeric ATP-Linked Nucleotide Enables Luminescence Signaling of Damage Surveillance by MTH1, a Cancer Target"
463:
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127:
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32:
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320:
276:
266:
2640:
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1158:"Novel Class of Potent and Cellularly Active Inhibitors Devalidates MTH1 as Broad-Spectrum Cancer Target"
237:
178:
3122:
3041:
2770:
2678:
2124:
1679:"8-Chloro-dGTP, a hypochlorous acid-modified nucleotide, is hydrolyzed by hMTH1, the human MutT homolog"
434:
Eliminating the MTH1 gene in mice results in over three times more mice developing tumors compared to a
363:
102:
3194:
1529:
Kang D, Nishida J, Iyama A, Nakabeppu Y, Furuichi M, Fujiwara T, Sekiguchi M, Takeshige K (June 1995).
510:"Chromosomal location and some structural features of human clathrin light-chain genes (CLTA and CLTB)"
1883:"An oxidized purine nucleoside triphosphatase, MTH1, suppresses cell death caused by oxidative stress"
2064:
1242:
Warpman
Berglund U, Sanjiv K, Gad H, Kalderén C, Koolmeister T, Pham T, et al. (December 2016).
966:
911:
664:
296:
508:
Ponnambalam S, Jackson AP, LeBeau MM, Pravtcheva D, Ruddle FH, Alibert C, Parham P (December 1994).
3359:
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130:
54:
3091:
2990:
2556:
2134:
1797:
990:
435:
1644:"A molecular basis for the selective recognition of 2-hydroxy-dATP and 8-oxo-dGTP by human MTH1"
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1998:
1969:
1939:
1904:
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1730:
1700:
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Fujikawa K, Yakushiji H, Nakabeppu Y, Suzuki T, Masuda M, Ohshima H, Kasai H (February 2002).
1665:
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Sakai Y, Furuichi M, Takahashi M, Mishima M, Iwai S, Shirakawa M, Nakabeppu Y (March 2002).
1620:
1612:
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Sakumi K, Furuichi M, Tsuzuki T, Kakuma T, Kawabata S, Maki H, Sekiguchi M (November 1993).
521:
439:
301:
2433:
1156:
Ellermann M, Eheim A, Rahm F, Viklund J, Guenther J, Andersson M, et al. (July 2017).
90:
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2856:
2670:
2370:
2190:
1881:
Yoshimura D, Sakumi K, Ohno M, Sakai Y, Furuichi M, Iwai S, Nakabeppu Y (September 2003).
428:
367:
1386:
66:
970:
915:
668:
651:
Tchou J, Kasai H, Shibutani S, Chung MH, Laval J, Grollman AP, Nishimura S (June 1991).
37:
3333:
3222:
3163:
2290:
1835:
1811:"A novel mechanism for preventing mutations caused by oxidation of guanine nucleotides"
1810:
1361:"MTH1, A Phase I, Study on Tumors Inhibition, First in Human, First in Class (MASTIFF)"
1311:
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288:
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567:
550:
3343:
3127:
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2834:
2444:
934:
900:"Spontaneous tumorigenesis in mice defective in the MTH1 gene encoding 8-oxo-dGTPase"
899:
687:
652:
412:
1801:
374:
that results in the production of an additional, longer isoform has been described.
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2714:
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994:
866:
447:
392:
348:
306:
1412:
Kettle JG, Alwan H, Bista M, Breed J, Davies NL, Eckersley K, et al. (2016).
1360:
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355:-induced oxidative stress, resulting in fewer dead cells and less accumulation of
1471:
1454:
1430:
1413:
1216:
1199:
1022:
849:
Kajitani K, Yamaguchi H, Dan Y, Furuichi M, Kang D, Nakabeppu Y (February 2006).
621:
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3235:
3071:
2915:
1414:"Potent and Selective Inhibitors of MTH1 Probe Its Role in Cancer Cell Survival"
1200:"Potent and Selective Inhibitors of MTH1 Probe Its Role in Cancer Cell Survival"
594:"Entrez Gene: NUDT1 nudix (nucleoside diphosphate linked moiety X)-type motif 1"
470:
255:
3328:
1174:
1157:
1134:
904:
Proceedings of the
National Academy of Sciences of the United States of America
657:
Proceedings of the
National Academy of Sciences of the United States of America
653:"8-oxoguanine (8-hydroxyguanine) DNA glycosylase and its substrate specificity"
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Yoshimura K, Ogawa T, Tsujimura M, Ishikawa K, Shigeoka S (September 2014).
731:
677:
404:
388:
344:
287:
during replication and transcription can cause mutations that may result in
220:
2037:"Call to bioscientists: choose and use your chemicaI probes very carefully"
2002:
1973:
1943:
1916:
Kamiya H, Dugué L, Yakushiji H, Pochet S, Nakabeppu Y, Harashima H (2003).
1908:
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1934:
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535:
1061:
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352:
2107:: Solution structure of the hMTH1, a nucleotide pool sanitization enzyme
978:
197:
78:
3011:
2980:
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2753:
2512:
2502:
2477:
2472:
2207:
1244:"Validation and development of MTH1 inhibitors for treatment of cancer"
826:
809:
469:
A potent and selective MTH1 inhibitor AZ13792138 has been developed by
97:
3287:
3057:
2888:
2530:
2507:
2497:
2467:
2462:
2457:
2452:
2409:
2399:
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2018:. American Association for the Advancement of Science. Archived from
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85:
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2995:
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400:
396:
284:
280:
1285:"MTH1 as a Chemotherapeutic Target: The Elephant in the Room"
462:
Karonudib, an MTH1 inhibitor, is currently being evaluated a
391:
that degrades 8-oxodGTP and 8-oxoGTP and therefore excludes
324:
Example hydrolysis of oxidized nucleotide catalyzed by MTH1
309:
in DNA, the protein encoded by this gene is an enzyme that
3026:
1564:
Oda H, Nakabeppu Y, Furuichi M, Sekiguchi M (July 1997).
1237:
1235:
241:
1367:. U.S. National Institutes of Health. 14 February 2017
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240:
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366:transcript variants, some of which encode distinct
172:
160:
148:
143:
120:
108:
96:
84:
72:
60:
48:
43:
31:
26:
21:
1455:"MutT Homolog 1 (MTH1): The Silencing of a Target"
246:
1918:"Substrate recognition by the human MTH1 protein"
1809:Ishibashi T, Hayakawa H, Sekiguchi M (May 2003).
1044:Ji D, Beharry AA, Ford JM, Kool ET (July 2016).
305:because of its ability to prevent occurrence of
343:The encoded protein is localized mainly in the
803:
801:
3042:
2150:
2072:
8:
1123:Bioorganic & Medicinal Chemistry Letters
466:clinical trial for safety and tolerability.
3049:
3035:
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2535:
2265:
2258:
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2135:
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140:
2127:at the U.S. National Library of Medicine
1933:
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1310:
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933:
923:
874:
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740:
730:
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676:
566:
525:
239:
2041:The Institute of Cancer Research, London
1283:Samaranayake GJ, Huynh M, Rai P (2017).
1050:Journal of the American Chemical Society
588:
586:
3324:
2094:
2012:"MTH1: From Hot Topic to Devalidation?"
1715:Archives of Biochemistry and Biophysics
500:
18:
7:
1954:Free Radical Biology & Medicine
1887:The Journal of Biological Chemistry
1854:Free Radical Biology & Medicine
1648:The Journal of Biological Chemistry
1570:The Journal of Biological Chemistry
1535:The Journal of Biological Chemistry
719:The Journal of Biological Chemistry
555:The Journal of Biological Chemistry
247:{\displaystyle \rightleftharpoons }
2043:. The Institute of Cancer Research
1922:Nucleic Acids Research. Supplement
14:
204:that in humans is encoded by the
3327:
2097:
2035:Workman, Paul (10 July 2017).
2016:Science Translational Medicine
1459:Journal of Medicinal Chemistry
1418:Journal of Medicinal Chemistry
1334:Thomas A (21 September 2016).
1204:Journal of Medicinal Chemistry
867:10.1523/jneurosci.4948-05.2006
610:Journal of Medicinal Chemistry
372:single-nucleotide polymorphism
1:
1966:10.1016/S0891-5849(03)00362-9
1866:10.1016/S0891-5849(03)00176-X
1757:10.1016/S0022-2836(02)00163-8
1727:10.1016/S0003-9861(02)00034-6
1696:10.1016/S0014-5793(02)02240-8
1107:10.1158/1538-7445.AM2017-5473
568:10.1016/S0021-9258(19)49494-5
2125:2-hydroxy-dATP+diphosphatase
1983:Journal of Molecular Biology
1745:Journal of Molecular Biology
1472:10.1021/acs.jmedchem.6b00283
1431:10.1021/acs.jmedchem.5b01760
1217:10.1021/acs.jmedchem.5b01760
1023:10.1016/j.dnarep.2015.05.008
622:10.1021/acs.jmedchem.7b01006
191:2-hydroxy-dATP diphosphatase
22:2-hydroxy-dATP diphosphatase
2927:Protein-synthesizing GTPase
2010:Lowe, Derek (7 July 2017).
855:The Journal of Neuroscience
770:Plant & Cell Physiology
3376:
1175:10.1021/acschembio.7b00370
1135:10.1016/j.bmcl.2016.02.026
417:DNA damage theory of aging
3205:Michaelis–Menten kinetics
2986:Guanylate-binding protein
2170:acid anhydride hydrolases
2092:
1995:10.1016/j.jmb.2003.12.060
1827:10.1038/sj.embor.embor838
139:
3097:Diffusion-limited enzyme
2541:Heterotrimeric G protein
2236:Phosphoadenylylsulfatase
2129:Medical Subject Headings
1774:Molecular Carcinogenesis
1583:10.1074/jbc.272.28.17843
1548:10.1074/jbc.270.24.14659
489:8-oxo-dGTP diphosphatase
277:nucleoside triphosphates
267:nucleoside triphosphates
2213:Thiamine-triphosphatase
732:10.1074/jbc.m116.749713
678:10.1073/pnas.88.11.4690
1900:10.1074/jbc.M306201200
1661:10.1074/jbc.M110566200
1617:10.1093/nar/27.22.4335
1605:Nucleic Acids Research
1514:10.1006/geno.1994.1657
1302:10.3390/cancers9050047
925:10.1073/pnas.191086798
527:10.1006/geno.1994.1650
340:
325:
248:
3190:Eadie–Hofstee diagram
3123:Allosteric regulation
2968:Polymerization motors
2679:Rho family of GTPases
1261:10.1093/annonc/mdw429
364:alternatively spliced
338:
323:
249:
3200:Lineweaver–Burk plot
1162:ACS Chemical Biology
1062:10.1021/jacs.6b02895
413:oxidative DNA damage
272:Misincorporation of
238:
2976:dynamin superfamily
1935:10.1093/nass/2.1.85
1393:. AstraZeneca. 2016
1340:Helleday Laboratory
979:10.1038/nature13181
971:2014Natur.508..215G
916:2001PNAS...9811456T
669:1991PNAS...88.4690T
347:, with some in the
3159:Enzyme superfamily
3092:Enzyme promiscuity
1365:ClinicalTrials.gov
1248:Annals of Oncology
827:10.1111/acel.12094
783:10.1093/pcp/pcu083
341:
326:
244:
230:2-hydroxy-dATP + H
3315:
3314:
3024:
3023:
3020:
3019:
2521:
2520:
2486:
2485:
2231:Adenylylsulfatase
2115:
2114:
1254:(12): 2275–2283.
616:(19): 8160–8169.
357:hydrogen peroxide
293:neurodegeneration
254:2-hydroxy-dAMP +
225:chemical reaction
188:
187:
184:
183:
103:metabolic pathway
3367:
3350:Nudix hydrolases
3332:
3331:
3323:
3195:Hanes–Woolf plot
3138:Enzyme activator
3133:Enzyme inhibitor
3107:Enzyme catalysis
3051:
3044:
3037:
3028:
2536:
2266:
2259:
2159:
2152:
2145:
2136:
2101:
2081:
2074:
2067:
2058:
2052:
2050:
2048:
2031:
2029:
2027:
2006:
1977:
1947:
1937:
1912:
1902:
1893:(39): 37965–73.
1877:
1848:
1838:
1805:
1786:10.1002/mc.10108
1768:
1738:
1708:
1698:
1673:
1663:
1638:
1628:
1595:
1585:
1576:(28): 17843–50.
1560:
1550:
1541:(24): 14659–65.
1525:
1485:
1484:
1474:
1453:Papeo G (2016).
1450:
1444:
1443:
1433:
1409:
1403:
1402:
1400:
1398:
1383:
1377:
1376:
1374:
1372:
1357:
1351:
1350:
1348:
1346:
1331:
1325:
1324:
1314:
1304:
1280:
1274:
1273:
1263:
1239:
1230:
1229:
1219:
1194:
1188:
1187:
1177:
1168:(8): 1986–1992.
1153:
1147:
1146:
1117:
1111:
1110:
1090:
1084:
1083:
1073:
1041:
1035:
1034:
1005:
999:
998:
965:(7495): 215–21.
954:
948:
947:
937:
927:
910:(20): 11456–61.
895:
889:
888:
878:
846:
840:
839:
829:
805:
796:
795:
785:
761:
755:
754:
744:
734:
725:(7): 2785–2794.
710:
701:
700:
690:
680:
648:
642:
641:
604:
598:
597:
590:
581:
580:
570:
561:(31): 23524–30.
546:
540:
539:
529:
505:
455:As a drug target
302:Escherichia coli
265:oxidized purine
253:
251:
250:
245:
141:
19:
16:Class of enzymes
3375:
3374:
3370:
3369:
3368:
3366:
3365:
3364:
3340:
3339:
3338:
3326:
3318:
3316:
3311:
3223:Oxidoreductases
3209:
3185:Enzyme kinetics
3173:
3169:List of enzymes
3142:
3111:
3082:Catalytic triad
3060:
3055:
3025:
3016:
2961:
2920:
2671:Ras superfamily
2660:
2644:
2624:
2570:
2560:
2552:
2517:
2482:
2438:
2419:
2365:
2322:Plasma membrane
2285:
2240:
2217:
2191:Pyrophosphatase
2177:
2163:
2121:
2116:
2111:
2108:
2102:
2088:
2085:
2055:
2046:
2044:
2034:
2025:
2023:
2022:on 10 July 2017
2009:
1980:
1950:
1915:
1880:
1860:(11): 1447–57.
1851:
1808:
1771:
1741:
1711:
1689:(1–3): 149–51.
1676:
1654:(10): 8579–87.
1641:
1611:(22): 4335–43.
1598:
1563:
1528:
1498:
1494:
1492:Further reading
1489:
1488:
1452:
1451:
1447:
1411:
1410:
1406:
1396:
1394:
1391:Open Innovation
1385:
1384:
1380:
1370:
1368:
1359:
1358:
1354:
1344:
1342:
1333:
1332:
1328:
1282:
1281:
1277:
1241:
1240:
1233:
1196:
1195:
1191:
1155:
1154:
1150:
1119:
1118:
1114:
1092:
1091:
1087:
1043:
1042:
1038:
1007:
1006:
1002:
956:
955:
951:
897:
896:
892:
848:
847:
843:
807:
806:
799:
763:
762:
758:
712:
711:
704:
650:
649:
645:
606:
605:
601:
592:
591:
584:
548:
547:
543:
507:
506:
502:
497:
480:
457:
429:cancer research
425:
385:
380:
236:
235:
233:
217:
17:
12:
11:
5:
3373:
3371:
3363:
3362:
3357:
3352:
3342:
3341:
3337:
3336:
3313:
3312:
3310:
3309:
3296:
3283:
3270:
3257:
3244:
3231:
3217:
3215:
3211:
3210:
3208:
3207:
3202:
3197:
3192:
3187:
3181:
3179:
3175:
3174:
3172:
3171:
3166:
3161:
3156:
3150:
3148:
3147:Classification
3144:
3143:
3141:
3140:
3135:
3130:
3125:
3119:
3117:
3113:
3112:
3110:
3109:
3104:
3099:
3094:
3089:
3084:
3079:
3074:
3068:
3066:
3062:
3061:
3056:
3054:
3053:
3046:
3039:
3031:
3022:
3021:
3018:
3017:
3015:
3014:
3009:
3008:
3007:
3002:
2993:
2988:
2983:
2972:
2970:
2963:
2962:
2960:
2959:
2954:
2953:
2952:
2947:
2942:
2931:
2929:
2922:
2921:
2919:
2918:
2913:
2908:
2903:
2898:
2897:
2896:
2891:
2886:
2881:
2871:
2870:
2869:
2864:
2854:
2853:
2852:
2847:
2842:
2830:
2829:
2828:
2827:
2822:
2812:
2811:
2810:
2805:
2800:
2790:
2789:
2788:
2783:
2778:
2768:
2763:
2762:
2761:
2756:
2746:
2745:
2744:
2739:
2734:
2729:
2719:
2718:
2717:
2712:
2702:
2701:
2700:
2695:
2690:
2675:
2673:
2662:
2661:
2659:
2658:
2657:
2656:
2651:
2642:
2638:
2637:
2636:
2631:
2622:
2618:
2617:
2616:
2615:
2614:
2604:
2603:
2602:
2597:
2587:
2582:
2577:
2568:
2564:
2563:
2562:
2558:
2550:
2545:
2543:
2533:
2523:
2522:
2519:
2518:
2516:
2515:
2510:
2505:
2500:
2494:
2492:
2488:
2487:
2484:
2483:
2481:
2480:
2475:
2470:
2465:
2460:
2455:
2449:
2447:
2440:
2439:
2437:
2436:
2430:
2428:
2421:
2420:
2418:
2417:
2412:
2407:
2402:
2397:
2392:
2387:
2382:
2376:
2374:
2367:
2366:
2364:
2363:
2362:
2361:
2356:
2346:
2345:
2344:
2339:
2334:
2329:
2319:
2318:
2317:
2312:
2307:
2296:
2294:
2287:
2286:
2284:
2283:
2278:
2272:
2270:
2269:Cu++ (3.6.3.4)
2263:
2256:
2242:
2241:
2239:
2238:
2233:
2227:
2225:
2219:
2218:
2216:
2215:
2210:
2205:
2204:
2203:
2198:
2187:
2185:
2179:
2178:
2164:
2162:
2161:
2154:
2147:
2139:
2133:
2132:
2120:
2119:External links
2117:
2113:
2112:
2110:
2109:
2103:
2096:
2093:
2090:
2089:
2086:
2084:
2083:
2076:
2069:
2061:
2054:
2053:
2032:
2007:
1978:
1960:(6): 595–602.
1948:
1913:
1878:
1849:
1806:
1769:
1739:
1709:
1674:
1639:
1596:
1561:
1526:
1495:
1493:
1490:
1487:
1486:
1445:
1424:(6): 2346–61.
1404:
1378:
1352:
1326:
1275:
1231:
1210:(6): 2346–61.
1189:
1148:
1112:
1085:
1056:(29): 9005–8.
1036:
1000:
949:
890:
861:(6): 1688–98.
841:
820:(4): 695–705.
797:
776:(9): 1534–43.
756:
702:
663:(11): 4690–4.
643:
599:
582:
541:
499:
498:
496:
493:
492:
491:
486:
479:
476:
456:
453:
424:
421:
384:
381:
379:
376:
289:carcinogenesis
259:
258:
243:
231:
223:the following
216:
213:
186:
185:
182:
181:
176:
170:
169:
164:
158:
157:
152:
146:
145:
137:
136:
125:
118:
117:
112:
106:
105:
100:
94:
93:
88:
82:
81:
76:
70:
69:
64:
58:
57:
52:
46:
45:
41:
40:
35:
29:
28:
24:
23:
15:
13:
10:
9:
6:
4:
3:
2:
3372:
3361:
3358:
3356:
3353:
3351:
3348:
3347:
3345:
3335:
3330:
3325:
3321:
3307:
3303:
3302:
3297:
3294:
3290:
3289:
3284:
3281:
3277:
3276:
3271:
3268:
3264:
3263:
3258:
3255:
3251:
3250:
3245:
3242:
3238:
3237:
3232:
3229:
3225:
3224:
3219:
3218:
3216:
3212:
3206:
3203:
3201:
3198:
3196:
3193:
3191:
3188:
3186:
3183:
3182:
3180:
3176:
3170:
3167:
3165:
3164:Enzyme family
3162:
3160:
3157:
3155:
3152:
3151:
3149:
3145:
3139:
3136:
3134:
3131:
3129:
3128:Cooperativity
3126:
3124:
3121:
3120:
3118:
3114:
3108:
3105:
3103:
3100:
3098:
3095:
3093:
3090:
3088:
3087:Oxyanion hole
3085:
3083:
3080:
3078:
3075:
3073:
3070:
3069:
3067:
3063:
3059:
3052:
3047:
3045:
3040:
3038:
3033:
3032:
3029:
3013:
3010:
3006:
3003:
3001:
2997:
2994:
2992:
2989:
2987:
2984:
2982:
2979:
2978:
2977:
2974:
2973:
2971:
2969:
2964:
2958:
2955:
2951:
2948:
2946:
2943:
2941:
2938:
2937:
2936:
2933:
2932:
2930:
2928:
2923:
2917:
2914:
2912:
2909:
2907:
2904:
2902:
2899:
2895:
2892:
2890:
2887:
2885:
2882:
2880:
2877:
2876:
2875:
2872:
2868:
2865:
2863:
2860:
2859:
2858:
2855:
2851:
2848:
2846:
2843:
2841:
2838:
2837:
2836:
2832:
2831:
2826:
2823:
2821:
2818:
2817:
2816:
2813:
2809:
2806:
2804:
2801:
2799:
2796:
2795:
2794:
2791:
2787:
2784:
2782:
2779:
2777:
2774:
2773:
2772:
2769:
2767:
2764:
2760:
2757:
2755:
2752:
2751:
2750:
2747:
2743:
2740:
2738:
2735:
2733:
2730:
2728:
2725:
2724:
2723:
2720:
2716:
2713:
2711:
2708:
2707:
2706:
2703:
2699:
2696:
2694:
2691:
2689:
2686:
2685:
2684:
2680:
2677:
2676:
2674:
2672:
2668:
2663:
2655:
2652:
2650:
2647:
2646:
2645:
2639:
2635:
2632:
2630:
2627:
2626:
2625:
2619:
2613:
2610:
2609:
2608:
2605:
2601:
2598:
2596:
2593:
2592:
2591:
2588:
2586:
2583:
2581:
2578:
2576:
2573:
2572:
2571:
2565:
2561:
2555:
2554:
2553:
2547:
2546:
2544:
2542:
2537:
2534:
2532:
2528:
2524:
2514:
2511:
2509:
2506:
2504:
2501:
2499:
2496:
2495:
2493:
2489:
2479:
2476:
2474:
2471:
2469:
2466:
2464:
2461:
2459:
2456:
2454:
2451:
2450:
2448:
2446:
2445:P-type ATPase
2441:
2435:
2432:
2431:
2429:
2426:
2422:
2416:
2413:
2411:
2408:
2406:
2403:
2401:
2398:
2396:
2393:
2391:
2388:
2386:
2383:
2381:
2378:
2377:
2375:
2372:
2368:
2360:
2357:
2355:
2352:
2351:
2350:
2347:
2343:
2340:
2338:
2335:
2333:
2330:
2328:
2325:
2324:
2323:
2320:
2316:
2313:
2311:
2308:
2306:
2303:
2302:
2301:
2298:
2297:
2295:
2292:
2288:
2282:
2279:
2277:
2274:
2273:
2271:
2267:
2264:
2260:
2257:
2255:
2251:
2247:
2243:
2237:
2234:
2232:
2229:
2228:
2226:
2224:
2220:
2214:
2211:
2209:
2206:
2202:
2199:
2197:
2194:
2193:
2192:
2189:
2188:
2186:
2184:
2180:
2175:
2171:
2167:
2160:
2155:
2153:
2148:
2146:
2141:
2140:
2137:
2130:
2126:
2123:
2122:
2118:
2106:
2100:
2095:
2091:
2082:
2077:
2075:
2070:
2068:
2063:
2062:
2059:
2042:
2038:
2033:
2021:
2017:
2013:
2008:
2004:
2000:
1996:
1992:
1989:(4): 843–50.
1988:
1984:
1979:
1975:
1971:
1967:
1963:
1959:
1955:
1949:
1945:
1941:
1936:
1931:
1927:
1923:
1919:
1914:
1910:
1906:
1901:
1896:
1892:
1888:
1884:
1879:
1875:
1871:
1867:
1863:
1859:
1855:
1850:
1846:
1842:
1837:
1832:
1828:
1824:
1821:(5): 479–83.
1820:
1816:
1812:
1807:
1803:
1799:
1795:
1791:
1787:
1783:
1779:
1775:
1770:
1766:
1762:
1758:
1754:
1751:(1): 129–39.
1750:
1746:
1740:
1736:
1732:
1728:
1724:
1720:
1716:
1710:
1706:
1702:
1697:
1692:
1688:
1684:
1680:
1675:
1671:
1667:
1662:
1657:
1653:
1649:
1645:
1640:
1636:
1632:
1627:
1622:
1618:
1614:
1610:
1606:
1602:
1597:
1593:
1589:
1584:
1579:
1575:
1571:
1567:
1562:
1558:
1554:
1549:
1544:
1540:
1536:
1532:
1527:
1523:
1519:
1515:
1511:
1508:(3): 485–90.
1507:
1503:
1497:
1496:
1491:
1482:
1478:
1473:
1468:
1465:(6): 2343–5.
1464:
1460:
1456:
1449:
1446:
1441:
1437:
1432:
1427:
1423:
1419:
1415:
1408:
1405:
1392:
1388:
1382:
1379:
1366:
1362:
1356:
1353:
1341:
1337:
1330:
1327:
1322:
1318:
1313:
1308:
1303:
1298:
1294:
1290:
1286:
1279:
1276:
1271:
1267:
1262:
1257:
1253:
1249:
1245:
1238:
1236:
1232:
1227:
1223:
1218:
1213:
1209:
1205:
1201:
1193:
1190:
1185:
1181:
1176:
1171:
1167:
1163:
1159:
1152:
1149:
1144:
1140:
1136:
1132:
1129:(6): 1503–7.
1128:
1124:
1116:
1113:
1108:
1104:
1100:
1096:
1089:
1086:
1081:
1077:
1072:
1067:
1063:
1059:
1055:
1051:
1047:
1040:
1037:
1032:
1028:
1024:
1020:
1016:
1012:
1004:
1001:
996:
992:
988:
984:
980:
976:
972:
968:
964:
960:
953:
950:
945:
941:
936:
931:
926:
921:
917:
913:
909:
905:
901:
894:
891:
886:
882:
877:
872:
868:
864:
860:
856:
852:
845:
842:
837:
833:
828:
823:
819:
815:
811:
804:
802:
798:
793:
789:
784:
779:
775:
771:
767:
760:
757:
752:
748:
743:
738:
733:
728:
724:
720:
716:
709:
707:
703:
698:
694:
689:
684:
679:
674:
670:
666:
662:
658:
654:
647:
644:
639:
635:
631:
627:
623:
619:
615:
611:
603:
600:
595:
589:
587:
583:
578:
574:
569:
564:
560:
556:
552:
545:
542:
537:
533:
528:
523:
519:
515:
511:
504:
501:
494:
490:
487:
485:
482:
481:
477:
475:
472:
467:
465:
460:
454:
452:
449:
444:
441:
437:
436:control group
432:
430:
422:
420:
418:
414:
410:
406:
402:
398:
394:
390:
382:
377:
375:
373:
369:
365:
360:
358:
354:
350:
346:
337:
333:
331:
322:
318:
316:
312:
308:
304:
303:
298:
294:
290:
286:
282:
278:
275:
270:
268:
264:
257:
229:
228:
227:
226:
222:
214:
212:
210:
207:
203:
199:
196:
192:
180:
177:
175:
171:
168:
165:
163:
159:
156:
153:
151:
147:
142:
138:
135:
132:
129:
126:
123:
119:
116:
113:
111:
107:
104:
101:
99:
95:
92:
89:
87:
83:
80:
79:NiceZyme view
77:
75:
71:
68:
65:
63:
59:
56:
53:
51:
47:
42:
39:
36:
34:
30:
25:
20:
3301:Translocases
3298:
3285:
3272:
3259:
3246:
3236:Transferases
3233:
3220:
3077:Binding site
2814:
2748:
2704:
2682:
2667:Small GTPase
2281:Wilson/ATP7B
2276:Menkes/ATP7A
2104:
2045:. Retrieved
2040:
2024:. Retrieved
2020:the original
2015:
1986:
1982:
1957:
1953:
1925:
1921:
1890:
1886:
1857:
1853:
1818:
1815:EMBO Reports
1814:
1780:(3): 123–9.
1777:
1773:
1748:
1744:
1718:
1714:
1686:
1683:FEBS Letters
1682:
1651:
1647:
1608:
1604:
1573:
1569:
1538:
1534:
1505:
1501:
1462:
1458:
1448:
1421:
1417:
1407:
1395:. Retrieved
1390:
1387:"AZ13792138"
1381:
1369:. Retrieved
1364:
1355:
1343:. Retrieved
1339:
1329:
1292:
1288:
1278:
1251:
1247:
1207:
1203:
1192:
1165:
1161:
1151:
1126:
1122:
1115:
1101:(13): 5473.
1098:
1094:
1088:
1053:
1049:
1039:
1014:
1010:
1003:
962:
958:
952:
907:
903:
893:
858:
854:
844:
817:
813:
773:
769:
759:
722:
718:
660:
656:
646:
613:
609:
602:
558:
554:
544:
520:(3): 440–4.
517:
513:
503:
468:
461:
458:
445:
433:
426:
393:8-oxoguanine
386:
361:
349:mitochondria
342:
329:
327:
307:8-oxoguanine
300:
271:
260:
219:This enzyme
218:
205:
190:
189:
67:BRENDA entry
3072:Active site
2991:Mitofusin-1
2966:3.6.5.5-6:
2935:Prokaryotic
2087:PDB gallery
1928:(2): 85–6.
1721:(1): 31–7.
471:AstraZeneca
261:The enzyme
256:diphosphate
55:IntEnz view
27:Identifiers
3360:DNA repair
3344:Categories
3275:Isomerases
3249:Hydrolases
3116:Regulation
2957:Eukaryotic
2590:Transducin
2427:(3.6.3.10)
2166:Hydrolases
1095:Cancer Res
1017:: 101–10.
1011:DNA Repair
814:Aging Cell
495:References
440:inhibition
311:hydrolyzes
263:hydrolyses
124:structures
91:KEGG entry
3154:EC number
2925:3.6.5.3:
2665:3.6.5.2:
2607:Gustducin
2539:3.6.5.1:
2373:(3.6.3.9)
2293:(3.6.3.8)
2196:Inorganic
1295:(5): 47.
630:1520-4804
448:cytotoxic
405:longevity
389:hydrolase
345:cytoplasm
313:oxidized
242:⇌
221:catalyses
44:Databases
3355:EC 3.6.1
3178:Kinetics
3102:Cofactor
3065:Activity
2201:Thiamine
2003:15095864
1974:12957652
1944:12903117
1909:12857738
1874:12757855
1845:12717453
1802:26364277
1794:12619034
1765:12051941
1735:12051680
1705:11852070
1670:11756418
1635:10536140
1502:Genomics
1481:26924380
1440:26878898
1321:28481306
1270:27827301
1226:26878898
1184:28679043
1143:26898335
1080:27413803
1031:26202347
987:24695224
944:11572992
885:16467516
836:23648059
792:24928220
751:28035004
638:28929756
514:Genomics
478:See also
378:Research
368:isoforms
362:Several
353:paraquat
297:isolated
295:. First
274:oxidized
215:Function
198:3.6.1.56
179:proteins
167:articles
155:articles
128:RCSB PDB
38:3.6.1.56
3334:Biology
3288:Ligases
3058:Enzymes
3012:Tubulin
2981:Dynamin
2833:other:
2513:Katanin
2503:Kinesin
2478:ATP13A3
2473:ATP13A2
2208:Apyrase
2047:19 July
2026:19 July
1836:1319193
1592:9211940
1557:7782328
1522:7713500
1397:19 July
1371:19 July
1345:19 July
1312:5447957
1289:Cancers
1071:5500214
995:4468358
967:Bibcode
912:Bibcode
876:6793619
742:5314174
697:2052552
665:Bibcode
577:8226881
536:7713494
464:phase I
330:E. coli
283:and/or
115:profile
98:MetaCyc
3320:Portal
3262:Lyases
2889:ARL13B
2749:RhoBTB
2643:α12/13
2531:GTPase
2508:Myosin
2498:Dynein
2468:ATP12A
2463:ATP11B
2458:ATP10A
2453:ATP8B1
2443:Other
2415:ATP1B4
2410:ATP1B3
2405:ATP1B2
2400:ATP1B1
2395:ATP1A4
2390:ATP1A3
2385:ATP1A2
2380:ATP1A1
2371:Na+/K+
2359:ATP2C2
2354:ATP2C1
2342:ATP2B4
2337:ATP2B3
2332:ATP2B2
2327:ATP2B1
2315:ATP2A3
2310:ATP2A2
2305:ATP2A1
2254:ATPase
2131:(MeSH)
2001:
1972:
1942:
1907:
1872:
1843:
1833:
1800:
1792:
1763:
1733:
1703:
1668:
1633:
1626:148714
1623:
1590:
1555:
1520:
1479:
1438:
1319:
1309:
1268:
1224:
1182:
1141:
1078:
1068:
1029:
993:
985:
959:Nature
942:
932:
883:
873:
834:
790:
749:
739:
695:
685:
636:
628:
575:
534:
484:NUDT15
423:Cancer
409:ageing
315:purine
202:enzyme
162:PubMed
144:Search
134:PDBsum
74:ExPASy
62:BRENDA
50:IntEnz
33:EC no.
3214:Types
2945:EF-Tu
2884:SAR1B
2867:RAB27
2862:RAB23
2815:RhoDF
2705:RhoUV
2688:CDC42
2683:Cdc42
2669:>
2654:GNA13
2649:GNA12
2634:GNA11
2623:αq/11
2612:GNAT3
2600:GNAT2
2595:GNAT1
2585:GNAI3
2580:GNAI2
2575:GNAI1
2527:3.6.5
2491:3.6.4
2434:ATP4A
2425:H+/K+
2300:SERCA
2262:3.6.3
2246:3.6.3
2223:3.6.2
2183:3.6.1
1798:S2CID
991:S2CID
935:58751
688:51731
415:(see
395:from
383:Aging
299:from
279:into
206:NUDT1
110:PRIAM
3306:list
3299:EC7
3293:list
3286:EC6
3280:list
3273:EC5
3267:list
3260:EC4
3254:list
3247:EC3
3241:list
3234:EC2
3228:list
3221:EC1
3005:OPA1
2998:and
2950:EF-G
2940:IF-2
2906:Rheb
2894:ARL6
2879:ARF6
2850:NRAS
2845:KRAS
2840:HRAS
2825:RhoD
2820:RhoF
2766:RhoH
2742:RhoG
2727:Rac1
2715:RhoV
2710:RhoU
2693:TC10
2629:GNAQ
2349:SPCA
2176:3.6)
2105:1iry
2049:2017
2028:2017
1999:PMID
1970:PMID
1940:PMID
1905:PMID
1870:PMID
1841:PMID
1790:PMID
1761:PMID
1731:PMID
1701:PMID
1666:PMID
1631:PMID
1588:PMID
1553:PMID
1518:PMID
1477:PMID
1436:PMID
1399:2017
1373:2017
1347:2017
1317:PMID
1266:PMID
1222:PMID
1180:PMID
1139:PMID
1076:PMID
1027:PMID
983:PMID
940:PMID
881:PMID
832:PMID
788:PMID
747:PMID
693:PMID
634:PMID
626:ISSN
573:PMID
532:PMID
411:and
399:and
209:gene
174:NCBI
131:PDBe
86:KEGG
3000:MX2
2996:MX1
2916:RGK
2911:Rap
2901:Ran
2874:Arf
2857:Rab
2835:Ras
2793:Rnd
2771:Rho
2722:Rac
2698:TCL
2559:olf
2291:Ca+
1991:doi
1987:336
1962:doi
1930:doi
1895:doi
1891:278
1862:doi
1831:PMC
1823:doi
1782:doi
1753:doi
1749:319
1723:doi
1719:402
1691:doi
1687:512
1656:doi
1652:277
1621:PMC
1613:doi
1578:doi
1574:272
1543:doi
1539:270
1510:doi
1467:doi
1426:doi
1307:PMC
1297:doi
1256:doi
1212:doi
1170:doi
1131:doi
1103:doi
1066:PMC
1058:doi
1054:138
1019:doi
975:doi
963:508
930:PMC
920:doi
871:PMC
863:doi
822:doi
778:doi
737:PMC
727:doi
723:292
683:PMC
673:doi
618:doi
563:doi
559:268
522:doi
419:).
401:RNA
397:DNA
291:or
285:RNA
281:DNA
150:PMC
122:PDB
3346::
2681::
2569:αi
2551:αs
2529::
2252::
2174:EC
2168::
2039:.
2014:.
1997:.
1985:.
1968:.
1958:35
1956:.
1938:.
1924:.
1920:.
1903:.
1889:.
1885:.
1868:.
1858:34
1856:.
1839:.
1829:.
1817:.
1813:.
1796:.
1788:.
1778:36
1776:.
1759:.
1747:.
1729:.
1717:.
1699:.
1685:.
1681:.
1664:.
1650:.
1646:.
1629:.
1619:.
1609:27
1607:.
1603:.
1586:.
1572:.
1568:.
1551:.
1537:.
1533:.
1516:.
1506:24
1504:.
1475:.
1463:59
1461:.
1457:.
1434:.
1422:59
1420:.
1416:.
1389:.
1363:.
1338:.
1315:.
1305:.
1291:.
1287:.
1264:.
1252:27
1250:.
1246:.
1234:^
1220:.
1208:59
1206:.
1202:.
1178:.
1166:12
1164:.
1160:.
1137:.
1127:26
1125:.
1099:77
1097:.
1074:.
1064:.
1052:.
1048:.
1025:.
1015:33
1013:.
989:.
981:.
973:.
961:.
938:.
928:.
918:.
908:98
906:.
902:.
879:.
869:.
859:26
857:.
853:.
830:.
818:12
816:.
812:.
800:^
786:.
774:55
772:.
768:.
745:.
735:.
721:.
717:.
705:^
691:.
681:.
671:.
661:88
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655:.
632:.
624:.
614:60
612:.
585:^
571:.
557:.
553:.
530:.
518:24
516:.
512:.
359:.
234:O
195:EC
3322::
3308:)
3304:(
3295:)
3291:(
3282:)
3278:(
3269:)
3265:(
3256:)
3252:(
3243:)
3239:(
3230:)
3226:(
3050:e
3043:t
3036:v
2808:3
2803:2
2798:1
2786:C
2781:B
2776:A
2759:2
2754:1
2737:3
2732:2
2641:G
2621:G
2567:G
2557:G
2549:G
2250:4
2248:-
2172:(
2158:e
2151:t
2144:v
2080:e
2073:t
2066:v
2051:.
2030:.
2005:.
1993::
1976:.
1964::
1946:.
1932::
1926:2
1911:.
1897::
1876:.
1864::
1847:.
1825::
1819:4
1804:.
1784::
1767:.
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1737:.
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1707:.
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1672:.
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1272:.
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