Human <scp>RIF</scp> 1 and protein phosphatase 1 stimulate <scp>DNA</scp> replication origin licensing but suppress origin activation

Name: Work Video:
Duration: 3 min 30 s
Shin‐ichiro Hiraga; Tony Ly; Javier Garzón; Zuzana Hořejšı́; Yoshinobu Ohkubo; Akinori Endo; Chikashi Obuse; Simon J. Boulton; Angus I. Lamond; Anne D. Donaldson
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4297], 'feedback.': [692], 'ToolsAdd': [693], 'favoritesDownload': [695], 'CitationsTrack': [696], 'CitationsPermissions': [697], 'ShareFacebookTwitterLinked': [698], 'InMendeleyWechatReddit': [699], 'Figures': [700], 'Info': [702], 'Abstract': [703], 'human': [705, 722, 788, 1095, 1517, 1701, 1741, 1769, 1811, 1874, 1925, 2174, 2201, 2297, 2398, 3109, 3164], 'controls': [708, 920, 2882, 3116], 'replication,': [710, 3882, 4016], 'molecular': [713], 'mechanism': [714], 'is': [715, 963, 981, 1086, 1110, 1223, 1345, 1450, 1478, 1503, 1559, 1569, 1717, 1790, 2294, 2724, 2767, 2791, 2809, 3111, 3670, 3827, 4018, 4175], 'largely': [716], 'unknown.': [717], 'Here,': [718, 1696], 'we': [719, 781, 1697, 1866, 2259, 2284, 2885, 2966, 3142, 3642, 3883], 'demonstrate': [720, 1767, 3082], 'that': [721, 737, 755, 853, 881, 940, 1300, 1422, 1473, 1699, 1714, 1768, 1789, 1806, 1836, 2142, 2173, 2246, 2275, 2281, 2663, 2718, 2803, 2869, 2880, 3013, 3049, 3066, 3083, 3108, 3115, 3644, 3677, 3929, 4216, 4267, 4282], 'negatively': [724, 866, 928], 'regulates': [725, 854, 1512], 'forming': [729], 'complex': [731, 878, 1049, 1102, 1178], 'with': [732, 761, 833, 879, 909, 991, 1005, 1068, 1248, 1366, 1385, 1425, 1516, 1652, 1668, 1706, 1819, 1880, 1935, 1987, 2013, 2035, 2042, 2105, 2129, 2150, 2166, 2178, 2192, 2231, 2277, 2388, 2397, 2550, 2635, 2680, 2807, 3071, 3092, 3158, 3202, 3242, 3293, 3534, 3624, 3659, 3720, 3729, 3766, 3800, 3856, 3906, 3970, 3981, 3994, 4012, 4044, 4085, 4131, 4168, 4245, 4272, 4340], '(PP1)': [736, 913, 1351], 'limits': [738, 882], 'phosphorylation-mediated': [739], 'MCM': [743, 752, 885, 925, 1066, 1101, 1177, 1393, 1465, 1476, 1728, 1761, 2303, 2319, 2375, 2449, 2526, 2564, 2592, 2696, 2728, 2837, 2862, 2875, 2883, 2893, 3044, 3068, 3090, 3117, 3126, 3140, 3149, 3322, 3532, 3577, 3682], 'replicative': [744], 'helicase.': [745], 'We': [746, 1268, 1734, 2120, 2213, 2305, 2895, 3171, 3523, 4002, 4025, 4149], 'identify': [747], 'specific': [748, 1809, 3122], 'residues': [749, 1879, 2003, 2654, 3180, 3218, 3232, 3264, 3287, 3527, 3540], 'on': [750, 1155, 1636, 1658, 1763, 2132, 2624, 2833, 2892, 2988, 3514, 3744, 3759, 3837, 4306, 4319], 'four': [751], 'helicase': [753, 1191, 1477, 3118, 3127, 3996], 'subunits': [754, 1162, 1179, 1355], 'show': [756, 1698, 2470, 3817, 3921], 'hyperphosphorylation': [757, 1440, 2446, 2562, 2706, 2738, 2813, 3046], 'upon': [758, 3313], 'depletion,': [760, 3201, 3565], 'regulatory': [763, 1161], 'N-terminal': [764, 1203], 'domain': [765], 'MCM4': [767, 2505, 2709, 2741, 2815, 2841, 2939, 2955, 2989, 3000, 3188, 3194, 3263, 3408], 'being': [768], 'particularly': [769, 1406], 'strongly': [770, 2867], 'affected.': [771], 'In': [772, 1196, 1335, 1543, 2159, 2323, 2730, 2776, 2798, 3299, 3896, 4226], 'addition': [773, 1955, 4045, 4151], 'role': [776, 786, 1317, 1774, 1780, 3131], 'in': [777, 790, 903, 954, 1012, 1033, 1080, 1094, 1235, 1240, 1255, 1279, 1307, 1320, 1326, 1332, 1342, 1500, 1505, 1508, 1522, 1541, 1617, 1692, 1732, 1743, 1775, 1802, 1862, 2004, 2027, 2218, 2269, 2314, 2353, 2494, 2534, 2597, 2669, 2690, 2726, 2751, 2770, 2781, 2785, 2847, 2850, 2906, 2936, 2954, 3134, 3332, 3492, 3496, 3521, 3655, 3663, 3820, 3840, 3853, 3880, 3888, 3926, 3989, 4021, 4058, 4094, 4160, 4182, 4221, 4241, 4249, 4331], 'limiting': [778, 1230, 1724, 4030], 'activation,': [780], 'discover': [782, 1736], 'an': [783, 1273, 1571, 1737, 3606, 3651, 4187, 4280], 'unexpected': [784, 1738], 'new': [785], 'RIF1-PP1': [789, 804, 891, 919, 932, 945, 1715, 1742, 1770, 2312], 'mediating': [791], 'efficient': [792], 'licensing.': [794, 831], 'Specifically,': [795], 'during': [796, 965, 1122, 1134, 1407, 1670], 'G1': [798, 904, 955, 1034, 1408], 'phase': [799, 1035, 1136, 1409, 3928, 4163], 'cell': [802, 966, 1007, 1082, 1111, 1124, 1156, 1461, 1926, 1933, 2092, 2414, 3639, 3738, 3923, 3943, 4141, 4289, 4323], 'cycle,': [803, 1083], 'protects': [805, 893], 'origin-binding': [807], 'ORC1': [808, 894, 934, 942, 953, 1115, 1117, 1757], 'from': [810, 895, 1614, 1792, 2091, 2514, 2621, 3151, 3233], 'untimely': [811], 'phosphorylation': [812, 883, 922, 939, 1118, 1199, 1259, 1402, 1725, 2301, 2317, 2376, 2524, 2589, 2616, 2640, 2693, 2834, 2863, 2937, 2956, 2998, 3069, 3088, 3137, 3214, 3243, 3258, 3276, 3308, 3582], 'consequent': [814], 'degradation': [815, 1113, 1130], 'proteasome.': [818], 'Depletion': [819, 2497, 2536, 2609], 'or': [822, 989, 1420, 1689, 1939, 2038, 2110, 2241, 2341, 2392, 2401, 2428, 2431, 2465, 2483, 2509, 2540, 2742, 3156, 3537, 3596, 3602, 3619, 3724, 3730, 3770, 3787, 3894, 4344], 'inhibition': [823, 4274], 'PP1': [825, 880, 916, 1353, 1386, 1416, 1434, 1588, 1641, 1653, 1669, 1687, 1690, 1707, 1710, 1825, 1832, 1837, 1854, 1870, 1999, 2008, 2076, 2081, 2112, 2130, 2136, 2153, 2168, 2179, 2182, 2190, 2210, 2217, 2227, 2247, 2278, 2292, 2373, 2500, 2808, 2831, 2859, 2872, 2881, 2890, 2900, 2963, 2974, 3084, 3625, 3679, 3878], 'destabilizes': [826], 'ORC1,': [827], 'thereby': [828], 'reducing': [829], 'Consistent': [832], 'reduced': [834, 2924, 3686, 3965, 3995], 'licensing,': [836], 'RIF1-depleted': [837], 'cells': [838, 970, 1096, 1482, 1548, 1580, 2034, 2387, 2408, 2516, 2908, 3152, 3654, 3719, 3742, 3765, 3797, 3889, 3900, 3925, 3940, 3955, 3971, 4159, 4191, 4240, 4284, 4339], 'exhibit': [839, 1551], 'increased': [840, 2614, 2639, 3001, 3027, 3213, 3249, 3275, 3312, 3562, 4188], 'spacing': [841], 'between': [842], 'active': [843, 1189], 'origins.': [844, 1764], 'therefore': [847, 946, 982, 1396, 1457, 3061, 4026], 'acts': [848, 1379], 'as': [849, 1291, 1380, 1468, 1796, 1798, 1909, 2197, 2229, 2492, 2520, 2533, 2596, 2846, 3240, 3852, 3863, 4017, 4176, 4186, 4255], 'PP1-targeting': [851, 1382, 3113], 'subunit': [852, 1208, 1369, 1590, 3114], 'positively': [857], 'stimulating': [859, 1744], 'step,': [863], 'then': [865, 3781], 'counteracting': [868, 937], 'activation.': [871, 3688], 'Synopsis': [872], 'forms': [876], 'proteins': [886, 926, 1654, 1847, 1950, 1971, 2026, 2056, 2088, 2099, 2203, 2211, 2274, 2304, 2320, 2476, 2527, 2593, 2860, 3085, 3150, 3189], 'control': [888, 1278, 1344, 1463, 2300, 2403, 3159, 3631, 4224], 'S-phase': [889, 1165, 3700, 3714, 3954], 'progression.': [890], 'also': [892, 1735, 3026], 'phosphorylation-dependent': [896], 'degradation,': [897], 'ensure': [899, 2412], 'full': [900, 948], 'phase.': [905, 956], 'interacts': [908, 1818, 1986, 2128, 2177, 3623], 'via': [914, 1823], 'its': [915, 1128, 1709, 1824, 1998, 2080, 2181, 2622, 2830, 3500, 3877], 'interaction': [917, 1417, 1424, 1711, 1716, 1826, 1833, 1855, 1871, 2000, 2009, 2082, 2137, 2183, 2248, 2293, 2313, 2804, 2832, 3879], 'motifs.': [918, 2001, 2083, 2138, 2184], 'status': [923, 2590, 2694, 2835, 2839, 3138], 'regulate': [929, 3067], 'replication.': [931, 1487, 4225], 'stabilizes': [933], 'protein,': [935], 'CDK-mediated': [938, 1258], 'triggers': [941, 1127], 'proteasomal': [943, 1129], 'destruction.': [944], 'ensures': [947, 1302, 1474], 'protecting': [952], 'Introduction': [957], 'critical': [964, 1878], 'proliferation': [967, 2415], 'daughter': [969], 'inherit': [972], 'complete': [974], 'set': [975, 1930], 'genomic': [977], 'information.': [978], 'tightly': [983], 'controlled': [984], 'prevent': [986, 1431, 2812], 'either': [987, 1681, 1936, 2336, 2740, 3589, 3892, 4110], 'under-': [988], 'over-replication,': [990], 'establishment': [993, 1019], 'licensed': [995], 'origins': [998, 1030], 'their': [1000, 1160, 1607, 1615, 1905, 4293], 'subsequent': [1001], 'closely': [1003, 1583], 'coupled': [1004], 'other': [1006, 3530, 3942], 'cycle': [1008, 1125, 1462, 3640, 3739, 3944, 4142, 4291, 4324], 'events.': [1009], 'Best': [1010], 'understood': [1011], 'budding': [1013, 1197, 1236, 1308, 1327, 4022], 'yeast,': [1014, 1198, 1280, 1328, 1330, 1509], 'mechanisms': [1016, 1105, 1731], 'controlling': [1017, 1523, 1720, 2315, 2691, 2727], 'appear': [1022, 1537, 3220], 'essentially': [1023], 'conserved': [1024, 1504, 1791], 'throughout': [1025, 2416], 'eukaryotes': [1026], '12.': [1027], 'are': [1031, 1169, 1209, 1361, 1483, 1638, 2011, 2067, 2678, 2762, 3506, 3930, 4123], 'established': [1032], 'through': [1036, 1103, 1640, 1708, 2079, 2180], 'stepwise': [1038], 'formation': [1039, 1183], 'pre-replication': [1042], 'complex,': [1043, 1187, 1729], 'composed': [1044], 'recognition': [1048, 1375], '(ORC)': [1050], 'minichromosome': [1052], 'maintenance': [1053], '(MCM)': [1054], 'hexamers': [1055], '3.': [1056, 1078, 3703], 'ORC': [1057], 'binds': [1058, 2075], 'first': [1059, 2896], 'potential': [1061, 2007], 'sites': [1063, 1200, 2644, 3207, 3259, 3309, 3583], 'recruits': [1065], 'complexes,': [1067], '"origin': [1070], 'licensing"': [1071], 'step': [1072, 1783], 'assisted': [1073], 'Cdc6': [1075], 'Cdt1': [1077], 'Later': [1079], 'prevented': [1087], 'cyclin-dependent': [1089], 'kinase': [1090, 1140, 1143], '(CDK)': [1091], 'activity,': [1092], 'which': [1093, 1168, 1558, 4135], 'inhibits': [1097], 'loading': [1099, 1266, 1762], 'several': [1104, 2270, 3567], '4.': [1106, 2495], 'One': [1107], 'these': [1109, 1428, 2324, 2760, 2800, 3080, 3550], 'cycle-dependent': [1112], 'protein:': [1116], 'cyclin': [1120], 'A–CDK2': [1121], 'S/G2': [1123], 'phases': [1126], '56.': [1131], 'Origin': [1132], 'S': [1135, 3618, 3656, 3927, 4162, 4242], 'requires': [1137], 'two': [1138, 2108, 2652, 3538, 3922], 'activities,': [1141], 'Dbf4-dependent': [1142], '(DDK)': [1144], 'CDK': [1146, 1153, 1239, 3591], '7.': [1147, 1195, 1267], 'activities': [1149], 'DDK': [1151, 1175, 1215, 1221, 1398, 1448, 2660, 3598, 3628, 4031, 4048, 4071, 4219, 4273], 'depend': [1154], 'cycle-controlled': [1157], 'accumulation': [1158, 3652, 4157, 4203, 4238], '(Dbf4': [1163], 'cyclins,': [1166], 'respectively),': [1167], 'upregulated': [1170], 'at': [1171, 1192, 1781, 1904, 1916, 1997, 2643, 3023, 3262, 3278, 3288, 3859, 3941], 'G1/S-phase': [1173], 'transition.': [1174], 'phosphorylates': [1176], 'promote': [1181], 'CMG': [1185], '(Cdc45-MCM-GINS)': [1186], 'forks': [1194], 'within': [1201, 1609, 3181, 3528, 4144], 'region': [1204], 'Mcm4': [1207], 'only': [1211, 3302, 3535, 3553], 'essential': [1212, 1759, 2295, 2810], 'targets': [1213, 1251, 2658], 'initiate': [1217], '8.': [1220], 'activity': [1222, 1435, 1449, 3629, 4032, 4072], 'proposed': [1224], 'be': [1226, 1808, 1840, 2236, 2245, 2657, 3062, 3222], 'one': [1227, 3303, 3536, 3554], 'factors': [1231], 'yeast': [1237, 1256, 1293, 1309, 1336, 1454, 1498, 1564, 1618, 1793, 4023], '910.': [1238], 'contrast': [1241], 'drives': [1242], 'recruitment': [1243], 'polymerase': [1246, 1265], 'ε,': [1247], 'relevant': [1250], 'most': [1252, 3197, 3579, 4286], 'clearly': [1253], 'identified': [1254, 1290, 2268, 3172, 3208, 3239, 3282, 3310, 3524], 'where': [1257, 3609], 'Sld2': [1261], 'Sld3': [1263], 'promotes': [1264], 'others': [1270, 3568], 'recently': [1271], 'discovered': [1272], 'additional': [1274], 'layer': [1275], 'mediated': [1281, 1346, 1639], '111213.': [1286, 1352, 1410, 4024], 'was': [1288, 1311, 2162, 2328, 2343, 2421, 2487, 2528, 2542, 2600, 2734, 2844, 3025, 3195, 3260, 3311, 3558, 3789, 4204], 'originally': [1289], 'telomeric': [1294], 'chromatin': [1295], 'component': [1296], '14.': [1297], 'discovery': [1299], 'late': [1303, 4161], 'telomeres': [1306], '1516': [1310], 'accompanied': [1312], 'studies': [1314], 'identifying': [1315], 'genomewide': [1321, 1525], 'regulation': [1322], 'fission': [1329], 'mammals': [1333], '171819202122.': [1334], 'cells,': [1337], 'function': [1339, 1496, 1788, 1801, 1843, 1858, 2687], 'catalytic': [1354, 1589], 'have': [1356, 1647], 'poor': [1357], 'substrate': [1358, 1374], 'specificity': [1359], 'intrinsically': [1362], 'promiscuous,': [1363], 'requiring': [1364], 'association': [1365], 'targeting': [1368, 1388, 1432, 1838], 'correct': [1371], 'localization': [1372, 2022, 2053], '2324.': [1376, 1599], 'Yeast': [1377], 'subunit,': [1383], 'interacting': [1384, 2230], 'it': [1389, 1627, 2243], 'dephosphorylate': [1391, 2874], 'complex.': [1394], 'Rif1-PP1': [1395, 1455], 'counteracts': [1397], 'maintain': [1400], 'low': [1401, 4194], 'levels': [1403, 3244, 3277], 'Mcm4,': [1405, 1442], 'Rif1': [1411, 1437, 2233], 'contains': [1412], 'series': [1414], 'motifs': [1418, 1429, 1827, 1834, 1872, 2010], '("[R/K]x[V/I]x[FW]"': [1419], '"[S/G]IL[K/R]")': [1421], 'mediate': [1423], 'PP1.': [1426], 'Mutating': [1427], 'leads': [1438, 2560, 3649], 'allowing': [1443], 'even': [1446], 'when': [1447, 3968], 'abnormally': [1451], 'low.': [1452], 'module': [1456], 'contributes': [1458], 'strict': [1460], 'phosphorylation,': [1466, 2884, 3683], 'acting': [1467], 'repressor': [1470], 'initiation': [1472], 'kept': [1479], 'inactive': [1480], 'until': [1481], 'ready': [1484], 'begin': [1486], 'It': [1488], 'has': [1489, 1628, 1663, 1676, 2265], 'been': [1490, 1629, 1648, 1664, 1677, 2267], 'unclear': [1491, 1630], 'what': [1493], 'extent': [1494, 2773, 4281], 'regulating': [1501, 3135], 'metazoans.': [1506], 'As': [1507, 2749], 'mammalian': [1510, 1533, 1634, 1860], 'timing,': [1515], 'mouse': [1519, 1545], 'involved': [1521], 'program': [1527], '171922.': [1528], 'However,': [1529, 1674], 'some': [1530, 2768, 4056], 'effects': [1531, 1632, 2987, 4034], 'do': [1535], 'not': [1536, 1560, 1570, 1973, 1981, 2215, 2503, 2792, 3946], 'parallel': [1539], 'those': [1540], 'yeast.': [1542, 1733], 'particular,': [1544, 2777, 3226], 'embryonic': [1546], 'fibroblast': [1547], 'lacking': [1549], 'severe': [1552, 4105], 'sensitivity': [1553], 'replication-inhibiting': [1555], 'drugs': [1556], '25,': [1557], 'case': [1562, 4020], 'rif1': [1565], 'mutants': [1566], '1826': [1567], 'expected': [1572, 2198], 'consequence': [1573], 'removing': [1575], 'repressor.': [1578], 'Mammalian': [1579], 'possess': [1581], 'three': [1582, 1869, 2006, 2152, 2468, 2545, 2973, 3279, 3531, 3551], 'related': [1584], 'subtypes': [1585], '(α,': [1591, 2902], 'β,': [1592, 2903], 'γ)': [1594, 2905], 'encoded': [1595], 'separate': [1597], 'genes': [1598], 'does': [1602, 2502], 'contain': [1603], 'PP1-binding': [1604], 'motifs,': [1605, 1688, 1712], 'although': [1606, 2765, 3566], 'position': [1608], 'sequence': [1612, 3594], 'differs': [1613], 'arrangement': [1616], '27.': [1620, 1848], 'Partly': [1621], 'due': [1622, 3672, 3998, 4068], 'structural': [1625], 'divergence,': [1626], 'whether': [1631, 2126, 4006, 4028], 'interaction.': [1642, 2291], 'Drosophila': [1645, 1661], 'reported': [1649], 'interact': [1651, 1705, 2149, 2165, 2276], '282930313233,': [1655], 'based': [1657, 3743], 'co-overexpression': [1659], 'experiments,': [1660, 2325], 'suggested': [1665], 'act': [1667, 4220], 'fly': [1671], 'development': [1672], '33.': [1673], 'there': [1675, 2766], 'no': [1678, 2929], 'direct': [1679], 'investigation': [1680], 'importance': [1684, 1852, 2309], 'interaction,': [1691], 'metazoan': [1693], 'function.': [1695], 'can': [1704, 2147], 'important': [1718], 'paralleling': [1730], 'requirement': [1739], 'origins,': [1750], 'ensuring': [1752], 'G1-specific': [1754], 'stabilization': [1755], 'Our': [1765], 'results': [1766, 2677, 2761, 2801, 2865, 3081], 'plays': [1771], 'dual': [1773], 'control—having': [1777], 'repressive': [1779], '(a': [1787], 'mammals),': [1795], 'well': [1797], 'positive': [1800], 'supporting': [1803], 'may': [1807, 1839, 2234, 2244, 3060, 3613], 'cells.': [1812, 2030], 'Results': [1813], 'physically': [1817, 2148], 'evolutionarily': [1829], 'conservation': [1830], 'suggests': [1835, 4266], 'core': [1842], 'eukaryotic': [1845], 'To': [1849, 2877, 3128, 3869], 'investigate': [1850, 3871], 'control,': [1865], 'mutated': [1867, 1996], 'substituting': [1877], 'alanine': [1881], '(I292A,': [1882], 'F294A,': [1883], 'I2181A,': [1884], 'L2182A,': [1885], 'V2204A,': [1886], 'F2206A)': [1888], 'create': [1890, 2016], 'RIF1-pp1bs': [1892, 1897, 1940, 1949, 2161], 'allele': [1893, 1898], '(Fig': [1894, 1964, 2155, 2286, 2321, 2347, 2626, 2648, 2712, 2744, 2758, 2794, 2912, 2940, 2976, 3003, 3297, 3666, 3917, 3976, 4073, 4099, 4147, 4198, 4212, 4260], '1A).': [1895], 'This': [1896, 4171, 4263], 'wild-type': [1900, 1937], 'were': [1902, 1914, 1951, 1972, 2040, 2057, 2089, 2100, 2395, 2409, 2438, 2512, 2571, 2577, 2594, 2995, 3238, 3569, 3727, 3773, 3798, 3848, 3901, 4347], 'fused': [1903], 'N-termini': [1906], 'GFP': [1908, 2055, 2065], 'described': [1910, 2493, 4178], '34.': [1911, 2368], 'constructs': [1913, 2358, 2394, 3726, 3772, 4346], 'integrated': [1915, 2389], 'FRT': [1918], 'site': [1919, 3555], 'Flp-In': [1922, 2031, 2384, 3716, 3762, 4336], 'T-REx': [1923, 2032, 2385, 3717, 3763, 4337], '293': [1924, 2033, 2386, 3718, 3764, 4338], 'line,': [1927], 'creating': [1928], 'stable': [1932, 3721, 3767, 4341], 'lines': [1934], 'expressed': [1941, 1974, 2145, 2195, 2356, 2721], 'under': [1942], 'doxycycline-inducible': [1944], 'promoter.': [1945], 'Both': [1946], 'successfully': [1952, 2923], 'induced': [1953, 2344, 3790], 'doxycycline': [1957, 2045], '(DOX),': [1958], 'localized': [1960], 'nucleus': [1963], '1B).': [1965], 'Without': [1966], 'DOX': [1967, 2346, 2420, 3732, 3792], 'induction,': [1968], 'GFP-fused': [1970], '(confirmed': [1975], 'microscopy': [1977], 'Western': [1979, 2103, 2139, 2441, 2471, 2548, 2633, 3295], 'blotting:': [1980], 'shown).': [1982], 'Figure': [1983, 2369, 2824, 3702, 4301], '1.': [1984, 3319], 'isoforms': [1991, 2078, 2154, 2501, 2511, 2901, 2975, 3065, 3096], 'Construction': [1992], 'cDNA': [1995], 'Critical': [2002], 'all': [2005, 2151, 3227, 3575], 'substituted': [2012], 'alanine,': [2014], 'RIF-pp1bs': [2018], 'allele.': [2019], 'Expression': [2020], 'GFP-RIF1': [2024, 2127, 2146, 2338, 2357, 2430, 2444, 2464, 2702, 2722, 3648, 3712, 3893, 3958, 4234], 'fusion': [2025, 2723], 'stably': [2028], 'transfected': [2029, 2396], 'GFP,': [2036, 2084, 2337, 2390, 2427, 3722, 3768, 3785, 4342], 'GFP-RIF1,': [2037, 2085, 2196, 2391, 3723, 3769, 3786, 4343], 'GFP-RIF1-pp1bs': [2039, 2087, 2342, 2393, 2452, 2733, 3695, 3725, 3771, 3788, 4345], 'cultivated': [2041], 'μM': [2044, 3802, 3845], '(DOX)': [2046], '3': [2048], 'days,': [2049], 'expression': [2051, 2334, 2462, 2700, 3646, 3693, 3710, 3756, 3834, 3975, 4009, 4043, 4084, 4232, 4270, 4316], 'confirmed': [2058, 2543, 4124], 'fluorescence': [2060], 'microscopy.': [2061], 'Phase-contrast,': [2062], 'DAPI-stain,': [2063], 'images': [2066], 'shown.': [2068], 'Scale': [2069], 'bar': [2070], 'indicates': [2071], '25': [2072], 'μm.': [2073], 'recovered': [2090, 2228], 'extracts': [2093], 'using': [2094, 2477, 2517, 2909], 'GFP-Trap': [2095], 'beads,': [2096], 'co-purifying': [2098], 'analyzed': [2101, 2529, 2595, 2845, 3851, 3912], 'blotting': [2104, 2549], 'anti-GFP': [2106], '(upper': [2107], 'panels)': [2109, 2546], 'isoform-specific': [2111, 2518, 2551, 2910], 'antibodies': [2113, 2478, 2637], '(lower': [2114, 2544], 'panels).': [2115], 'Download': [2116, 2118, 2605, 2607, 2855, 2857, 3865, 3867], 'figure': [2117, 2606, 2856, 3866], 'PowerPoint': [2119, 2608, 2858, 3868], 'performed': [2121, 3143], 'co-immunoprecipitation': [2122], 'experiments': [2123, 2220], 'test': [2125], 'depending': [2131], 'presence': [2134, 3842, 4228], 'analysis': [2140, 2473, 2634, 3146, 3296, 3637, 3811, 4139], 'revealed': [2141, 2638], 'ectopically': [2144, 2194, 2355, 2720, 3890, 3956], '1C,': [2156], 'lane': [2157, 2715, 2747, 2789, 2796, 3006], '5).': [2158, 2797], 'contrast,': [2160, 2732], 'unable': [2163, 2735], 'any': [2167, 3077, 3699], 'isoform': [2169, 2931], '(lane': [2170, 3018, 3041], '6),': [2171], 'demonstrating': [2172, 2717], 'indeed': [2176], 'Only': [2185], 'small': [2187], 'fraction': [2188], 'co-immunoprecipitated': [2191], 'since': [2199], 'numerous': [2200, 3173], 'PP1-interacting': [2202], 'compete': [2204], 'limited': [2207], 'pool': [2208], '282935.': [2212], 'could': [2214, 2957], 'detect': [2216], 'pull-down': [2219], 'against': [2221, 2365], 'endogenous': [2222, 2232, 2326, 2366, 2459, 2611, 3776, 3962, 4210], 'RIF1.': [2223, 4001], 'amount': [2225], 'simply': [2235], 'below': [2237], 'our': [2238], 'detection': [2239], 'threshold,': [2240], 'alternatively,': [2242], 'causes': [2249, 3713], 'masking': [2250], 'epitope': [2254], 'recognized': [2255], 'antibody': [2258], 'used': [2260], 'pull-down.': [2262], 'Endogenous': [2263], 'however': [2266, 4235], 'large-scale': [2271], 'analyses': [2272], '28293132,': [2279], 'confirming': [2280], 'binding': [2283], 'observe': [2285], '1C)': [2287], 'reflects': [2288], 'physiological': [2290], 'next': [2306], 'tested': [2307, 2886, 2985, 4027], '2A).': [2322], 'depleted': [2329, 2513, 2897, 2996, 3153, 3774, 3960, 4348], 'siRNA': [2331, 2363, 2400, 2569, 2584, 2911, 2921, 3160, 3334], '34,': [2332], '(wild': [2339], 'type)': [2340], '2A': [2348, 2627, 2649, 2713, 2745, 2787], '(i)).': [2349, 4170], 'Synonymous': [2350], 'base': [2351], 'substitutions': [2352], 'make': [2359], 'them': [2360], 'resistant': [2361], 'targeted': [2364], '2.': [2370], 'limit': [2374, 2861, 3086], '(i)': [2377, 2496, 4167], 'Outline': [2378], 'experiment.': [2380, 2854], 'On': [2381, 2405, 2433], 'Day': [2382], '1,': [2383], 'non-targeting': [2402], 'siRNA.': [2404], 'day': [2406, 2434], '2,': [2407], 'diluted': [2410], 'continuous': [2413], 'experiment,': [2418], 'added': [2422, 3858], 'induce': [2424], 'transcription': [2425], 'siRNA-resistant': [2429], 'GFP-RIF1-pp1bs.': [2432, 2466, 3895], '4,': [2435], 'chromatin-enriched': [2436], 'samples': [2437, 4330], 'prepared': [2439, 3849], 'blotting.': [2442], '(ii)': [2443, 2553], 'prevents': [2445], 'chromatin-associated': [2448, 2475, 2618, 3148], 'proteins,': [2450, 3091, 3141, 3552], 'while': [2451], 'cannot.': [2453], 'Upper': [2454], 'panel': [2455], 'confirms': [2456], 'removal': [2457], 'Lower': [2467], 'panels': [2469], 'blot': [2472], 'recognizing': [2479], 'MCM4,': [2480, 2619], 'phosphorylated': [2481, 2484, 2664, 2782, 3176, 3283, 3490, 3526, 3539, 3617], 'MCM2-S40,': [2482], 'MCM2-S53.': [2485], 'Loading': [2486, 2599], 'normalized': [2488, 2601], 'total': [2490, 2583, 2603], 'single': [2499, 2930], 'affect': [2504], 'phosphorylation.': [2506, 2729, 2894, 2990, 3119], 'PP1α,': [2507, 2538, 3037], 'PP1β,': [2508, 2539, 3038], 'PP1γ': [2510, 2541, 2559, 2994, 3031, 3040, 3054, 3059, 3095], 'HEK293': [2515, 2907, 3165], 'siRNAs': [2519, 2576], 'indicated,': [2521], 'then,': [2523], 'after': [2530, 3782], '2': [2531, 3734], 'days': [2532, 3795], '(A).': [2535, 2598], 'antibodies.': [2552], 'Simultaneous': [2554], 'depletion': [2555, 2891, 2927, 3017, 3035, 3271, 3315, 3587, 3753, 3831, 4208, 4312], 'PP1α': [2557, 2992, 3029, 3052, 3057, 3093], 'proteins.': [2565, 2697, 2876], 'Double': [2566], 'triple': [2568], 'transfections': [2570], 'performed.': [2572], 'Equal': [2573], 'amounts': [2574], 'mixed': [2578], 'give': [2580, 4136], 'constant': [2582], 'concentration': [2585], '(50': [2586], 'nM),': [2587], 'protein.': [2604], 'significantly': [2613, 3002], 'evident': [2620], 'retardation': [2623], 'SDS–PAGE': [2625, 2670], '(ii),': [2628, 2650, 2714, 2746, 2788, 2978, 3005], 'lanes': [2629, 2915, 2945, 2979], '2–4': [2630], '6).': [2632], 'phospho-specific': [2636], 'MCM2': [2642, 2665, 2711, 2743, 2843, 3022, 3338], 'Ser40': [2645, 2784], 'Ser53': [2647, 3024], 'lower': [2651], 'panels),': [2653], 'known': [2655], '3637.': [2661], 'Note': [2662, 3823], 'runs': [2666], 'slightly': [2667], 'faster': [2668], 'than': [2671, 3518, 3560, 4106], 'unphosphorylated': [2673], 'form': [2674], '37.': [2675], 'consistent': [2679, 3292, 3658, 3993], 'previous': [2681], 'observations': [2682, 4122], '19': [2683], 'confirm': [2685, 2878], 'Importantly,': [2698], 'ectopic': [2699, 3645, 3692, 3755, 3833, 3973, 4007, 4036, 4041, 4083, 4231, 4268, 4315], 'fully': [2703], 'suppressed': [2704], 'both': [2708], '5),': [2716], 'functional': [2725, 2959], 'sharp': [2731], '7).': [2748, 3019], 'shown': [2750, 3495, 3520], 'biological': [2753, 2852], 'repeat': [2754], 'experiment': [2757], 'EV1),': [2759], 'generally': [2763, 3245], 'reproducible': [2764, 2793], 'variability': [2769], 'apparent': [2772, 2779], 'effects.': [2775], 'reduction': [2780, 3988, 4057, 4093, 4248], 'Mcm2': [2783], 'Fig': [2786, 2848, 2918, 2948, 2982, 4165, 4332], '4': [2790, 3168], 'EV1,': [2795], 'general,': [2799], 'imply': [2802, 3107], 'MCM2.': [2817], 'Click': [2818, 4295], 'here': [2819, 4296], 'expand': [2821, 4298], 'figure.': [2823, 4300], 'EV1.': [2825], 'Effect': [2826, 3751, 3829, 4303, 4310], 'proteinsPhosphorylation': [2838], '2B,': [2849], 'duplicate': [2853], 'above': [2866], 'suggest': [2868], 'directs': [2871, 3678], 'directly': [2879], 'effect': [2888, 3669, 3873, 4108, 4172], 'individual': [2899], '2B': [2913, 2941, 2977, 3004], '(i),': [2914, 2942], '1–5;': [2916], 'Appendix': [2917, 2947, 2981], 'S1).': [2919], 'Although': [2920], 'treatment': [2922, 4053, 4087, 4111], 'each': [2925, 3182, 3512, 3543, 3821], 'isoform,': [2926], 'caused': [2932, 3014, 3043, 3050, 3324, 3696, 4055, 4088, 4155, 4236], 'major': [2934, 3064, 3100], 'change': [2935], 'top': [2943], 'panel,': [2944], '3–5;': [2946], 'S2A).': [2949], 'absence': [2951], 'changes': [2953], 'reflect': [2958], 'redundancy': [2960], 'among': [2961], 'isoforms.': [2964], 'Therefore,': [2965], 'carried': [2967], 'out': [2968], 'combined': [2969, 4271], 'depletions': [2970], '6–11;': [2980], 'S2B)': [2983], 'When': [2991], 'simultaneously,': [2997], '9),': [3007], 'level': [3010, 3510, 4195], 'close': [3011], 'Phosphorylation': [3020, 3192], 'double': [3032, 3055], 'depletion.': [3033, 3056, 3252], 'Triple': [3034], '11)': [3042], 'similar': [3047, 3254], 'levels,': [3070], 'PP1β': [3072], 'making': [3073], 'smaller': [3075], 'if': [3076, 3698], 'contribution.': [3078], 'Overall,': [3079], 'probably': [3097], 'playing': [3098], 'role.': [3101], 'Taken': [3102], 'together,': [3103], 'data': [3105], 'presented': [3106], 'down-regulates': [3121], 'phosphosites': [3123], 'explore': [3129], 'comparative': [3144], 'phosphoproteomic': [3145], 'treated': [3157], '(using': [3161], 'asynchronously': [3162, 3898], 'growing': [3163, 3899], 'cells;': [3166], 'see': [3167], 'details).': [3170], '(≥': [3174], '7)': [3175], 'serine': [3177, 3229, 3286], 'threonine': [3179, 3231], 'MCM2,': [3185, 3269], 'MCM3,': [3186, 3300], '(Table': [3190, 3224, 3266, 3316, 3572], '1).': [3191, 3267, 3317, 3573], 'affected': [3198, 3219, 3257, 3571, 3584], 'nine': [3203], '14': [3206], 'showing': [3209, 4326], 'substantially': [3210, 3274], '(>': [3211], 'twofold)': [3212], 'levels.': [3215], 'Interestingly,': [3216], 'clustered': [3223], '1)—in': [3225], 'five': [3228], 'positions': [3234], '23': [3235, 3409], '34': [3237, 3425], 'phosphorylated,': [3241], 'four-': [3246], 'sixfold': [3248], 'A': [3253], 'clustering': [3255], 'found': [3261, 3643, 4150], '131–145': [3265], 'For': [3268, 3574], 'led': [3272], 'seven': [3281], 'residues,': [3284], '40': [3289, 3347], '53': [3291, 3355, 3429], '2).': [3298], 'however,': [3301], '(Thr728)': [3304], '10': [3307, 3844], 'Table': [3318], 'Changes': [3320, 3331, 3505], 'phosphorylations': [3323], 'depleting': [3326], 'Protein': [3328], 'Position': [3329], 'Sequencea': [3330], 'Fold': [3335], 'changesb': [3336], 'Log2': [3337], '26': [3339, 3413], 'PLTSSPG': [3340], '1.30': [3341], '0.38': [3342], '27': [3343], 'LTSSPGR': [3344, 3352], '0.76': [3345], '−0.40': [3346], 'ALTSSPG': [3348], '2.04': [3349], '1.03': [3350], '41': [3351], '1.01': [3353], '0.02': [3354], 'EDESEGL': [3356], '2.55': [3357], '1.35': [3358], '108': [3359], 'LTASQRE': [3360], '4.43': [3361], '2.15': [3362], '139': [3363], 'LYDSDEE': [3364], '0.81': [3365], '−0.30': [3366], 'MCM3': [3367], '163': [3368], 'SDLTTLV': [3369], '1.29': [3370], '0.37': [3371], '164': [3372], 'DLTTLVA': [3373], '1.40': [3374], '0.48': [3375], '668': [3376], 'KKRSEDE': [3377], '0.73': [3378], '−0.46': [3379], '672': [3380], 'EDESETE': [3381], '0.89': [3382], '−0.17': [3383], '674': [3384], 'ESETEDE': [3385], '0.88': [3386], '−0.18': [3387], '708': [3388], 'YDPYDFS': [3389], '1.24': [3390], '0.31': [3391], '711': [3392], 'YDFSDTE': [3393], '1.07': [3394], '0.10': [3395], '713': [3396], 'FSDTEEE': [3397], '0.92': [3398, 3443], '−0.12': [3399], '722': [3400], 'QVHTPKT': [3401], '1.38': [3402], '0.47': [3403], '728': [3404], 'TADSQET': [3405], '2.05': [3406], '1.04': [3407], 'PAQTPRS': [3410], '5.63': [3411], '2.49': [3412], 'TPRSEDA': [3414], '3.97': [3415], '1.99': [3416], '31': [3417], 'DARSSPS': [3418], '5.60': [3419, 3423], '2.48': [3420, 3424], '32': [3421], 'ARSSPSQ': [3422], 'SSPSQRR': [3426], '5.55': [3427], '2.47': [3428], 'PMPTSPG': [3430], '2.07': [3431], '1.05': [3432], '54': [3433], 'MPTSPGV': [3434], '1.62': [3435], '0.70': [3436], '61': [3437], 'DLQSPAA': [3438], '1.76': [3439], '0.82': [3440], '87': [3441], 'FDVSSPL': [3442], '−0.13': [3444], '88': [3445], 'DVSSPLT': [3446], '0.74': [3447], '−0.43': [3448], '120': [3449], 'DLGSAQK': [3450], '1.42': [3451], '0.50': [3452], '131': [3453], 'DLQSDGA': [3454], '2.75': [3455], '1.46': [3456], '142': [3457], 'IVASEQS': [3458], '6.10': [3459], '2.61': [3460], '145': [3461], 'SEQSLGQ': [3462], '7.49': [3463], '2.91': [3464], 'MCM5': [3465], '498': [3466], 'WDETKGE': [3467], '1.69': [3468, 3472], '0.75': [3469, 3473], '512': [3470], 'TILSRFD': [3471], 'MCM6': [3474], '689': [3475], 'HADSPAP': [3476], '1.22': [3477], '0.29': [3478], '762': [3479], 'EIDSEEE': [3480], '5.30': [3481], '2.41': [3482], 'MCM7': [3483], '500': [3484], 'PRRSLEQ': [3485], '1.93': [3486], '0.95': [3487], 'Relevant': [3489], 'residue': [3491, 3612], 'bold': [3493], 'text,': [3494], 'context': [3498], 'six': [3501], 'surrounding': [3502], 'residues.': [3503], 'b': [3504], 'standardized': [3507], 'overall': [3509, 4132], 'chromatin.': [3515], 'Increases': [3516], 'greater': [3517], 'twofold': [3519, 3561], 'bold.': [3522], 'fewer': [3525], 'subunits,': [3533, 3578], 'observed': [3541], 'MCM5,': [3545], 'MCM6,': [3546], 'MCM7.': [3548], 'Among': [3549], '(MCM6': [3556], 'Ser762)': [3557], 'marginally': [3570], 'match': [3588], 'target': [3592, 3599], 'consensus': [3593, 3600], '[S/TP]': [3595], '[S': [3601], 'T': [3603], 'followed': [3604, 4313], 'acidic': [3607, 3611], 'residue,': [3608], 'correspond': [3614], 'previously': [3616, 4177], 'T]': [3620], '3839.': [3621], 'opposes': [3627], 'rate': [3633, 3887, 3992, 4067, 4098, 4251, 4277], 'By': [3634], 'flow': [3635, 3749, 3809, 3914], 'cytometry': [3636, 3810], 'distribution,': [3641, 4325], 'phase,': [3657, 4243], 'mild': [3661], 'defect': [3662], 'synthesis': [3665, 3708, 3839, 3886, 3991, 4066, 4254, 4309, 4321], '3A).': [3667], 'likely': [3671], 'excess': [3674, 4000], 'restrict': [3681], 'leading': [3684], 'Supporting': [3689], 'explanation,': [3691], 'little': [3697], 'accumulation.': [3701, 3715], 'affects': [3705], 'cellular': [3706, 4064, 4307], 'Ectopic': [3709], 'grown': [3728], 'without': [3731, 3972], 'days.': [3735], 'Plot': [3736], 'shows': [3737], 'distribution': [3740, 4143], 'content': [3746], 'measured': [3747, 4256], 'cytometry.': [3750, 3915], 'synthesis.': [3761, 4120], 'siRNA,': [3779], 'day,': [3784], 'addition.': [3793], 'Two': [3794], 'later,': [3796], 'pulse-labeled': [3799, 3902], '20': [3801], 'EdU': [3803, 3813, 3818, 3910, 3933, 3948, 3966, 4059, 4096, 4129, 4197, 4258], '15': [3805, 3904], 'min': [3806, 3905], 'prior': [3807], 'incorporation.': [3814], 'Overlaid': [3815], 'histograms': [3816], 'incorporation': [3819, 3911, 3967, 4097, 4130, 4259], 'condition.': [3822], 'x-axis': [3825], 'scale': [3826], 'bi-exponential.': [3828], 'XL413.': [3846], 'Cells': [3847], '(B)': [3854], 'XL413': [3857, 4050, 4052, 4086, 4153, 4174], 'same': [3861], 'time': [3862], 'DOX.': [3864], 'further': [3870], 'examined': [3884], 'expressing': [3891, 3957], 'brief,': [3897], '5-ethynyl-2′-deoxyuridine': [3907], '(EdU),': [3908], 'Histograms': [3916], '3B': [3918, 4333], 'C)': [3920], 'populations:': [3924], 'incorporating': [3931, 3947, 4192], 'significant': [3932], '(black': [3934, 3949], 'plot,': [3935, 3950], '"EdU': [3936, 3951], 'positive"': [3937], 'peak),': [3938], 'stages': [3945], 'negative"': [3952], 'peak).': [3953], '(and': [3959], 'RIF1)': [3963], 'showed': [3964], 'compared': [3969], '3B,': [3977], 'compare': [3978, 4075], 'red': [3979, 4101], 'plot': [3980], 'black': [3982, 4076], 'blue': [3984], 'plots),': [3985], 'implying': [3986], 'wished': [4003], 'examine': [4005], 'interferes': [4010], 'specifically': [4011], 'DDK-dependent': [4013], 'steps': [4014], 'simultaneously': [4029], 'exacerbates': [4033], 'expression,': [4038], 'combining': [4040], 'inhibitor': [4049], '40.': [4051], 'alone': [4054, 4154], 'incorporation,': [4060], 'reflecting': [4061], 'repression': [4062], 'impaired': [4070], '3C,': [4074, 4100, 4199], 'gray': [4078, 4200], 'histogram': [4079, 4185], 'profiles).': [4080], 'Combining': [4081], 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'cited_by_count': 8}, {'year': 2023, 'cited_by_count': 19}, {'year': 2022, 'cited_by_count': 16}, {'year': 2021, 'cited_by_count': 21}, {'year': 2020, 'cited_by_count': 19}, {'year': 2019, 'cited_by_count': 22}, {'year': 2018, 'cited_by_count': 24}, {'year': 2017, 'cited_by_count': 7}], 'updated_date': '2025-01-12T03:51:11.895873', 'created_date': '2017-01-26'}
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