Molecular crowding and RNA synergize to promote phase separation, microtubule interaction, and seeding of Tau condensates

Name: Work Video:
Duration: 3 min 30 s
Janine Hochmair; Christian Exner; Maximilian Franck; Alvaro Dominguez‐Baquero; Lisa Diez; H. Brognaro; Matthew L. Kraushar; Thorsten Mielke; Helena Radbruch; Senthilvelrajan Kaniyappan; Sven Falke; Eckhard Mandelkow�; Christian Betzel; Susanne Wegmann
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1561, 1604, 1617, 1668, 1674, 1679, 1690, 1733, 1740, 1775, 1791, 1813, 1838, 1910, 1951, 2012, 2037, 2041, 2055, 2081, 2112, 2131, 2154, 2159, 2176, 2211, 2236, 2244, 2249, 2268, 2284, 2328, 2353, 2407, 2441, 2465, 2474, 2493, 2508, 2564, 2577, 2630, 2642, 2654, 2692, 2695, 2716, 2726, 2745, 2771, 2810, 2912, 2924, 2932, 2946, 2952, 2958, 2989, 3016, 3139, 3163, 3170, 3226, 3233, 3247, 3286, 3292, 3349, 3376, 3379, 3400, 3404, 3413, 3461, 3500, 3506, 3527, 3554, 3559, 3583, 3592, 3596, 3603, 3606, 3619, 3691, 3700, 3790, 3889, 3893, 3908, 3937, 3945, 3974, 4154, 4162], 'full': [311, 779], 'name.': [313, 781], 'Resources': [315, 783], 'Sven': [323, 325, 791, 793, 981], 'Falke': [324, 326, 792, 794, 3276], 'orcid.org/0000-0003-3409-1791': [327, 795], 'Eckhard': [358, 360, 826, 828, 983], 'Mandelkow': [359, 361, 827, 829], 'orcid.org/0000-0003-4655-4829': [362, 830], 'Conceptualization,': [384, 425, 461, 852, 893, 929], 'Funding': [385, 428, 467, 853, 896, 935], 'acquisition,': [386, 429, 468, 854, 897, 936], 'Methodology,': [387, 430, 471, 855, 898, 939], 'Betzel': [401, 403, 869, 871], 'orcid.org/0000-0002-3879-5019': [404, 872], 'Supervision,': [427, 466, 895, 934], 'Susanne': [443, 447, 911, 915, 988], 'Wegmann': [444, 448, 912, 916, 989, 1932, 3928], 'Corresponding': [445, 913], 'Author': [446, 914, 958], '[email': [449, 917, 1068], 'protected]': [450, 918, 1069], 'orcid.org/0000-0002-5388-2479': [451, 919], 'Validation,': [469, 937], 'original': [474, 942], 'draft,': [475, 943], 'Project': [476, 944], 'administration,': [477, 945], 'Information': [959], 'Hochmair1,†,': [961], 'Exner2,†,': [963], 'Franck1,': [965], 'Dominguez-Baquero1,': [967], 'Diez1,': [969], 'Brognaro2,': [971], 'Kraushar3,': [974], 'Mielke3,': [976], 'Radbruch4,': [978], 'Kaniyappan5,6,': [980], 'Falke2,': [982], 'Mandelkow5,6,': [984], 'Betzel2': [986], '*,1': [990], '1German': [991], '2Institute': [999], '3Max': [1018], '4Institute': [1027], '5German': [1034], '6Department': [1042], '†': [1054], 'These': [1055, 1252, 2444], 'authors': [1056], 'contributed': [1057], 'equally': [1058], 'work': [1061], '*Corresponding': [1062], 'author.': [1063], 'Tel:': [1064], '+49': [1065], '30450539834;': [1066], 'E-mail:': [1067], 'EMBO': [1071], 'Journal': [1072], '(2022)41:e108882https://doi.org/10.15252/embj.2021108882': [1073], 'See': [1074], 'also:': [1075], 'VI': [1076], 'Wiersma': [1077], 'et': [1078, 1532, 1551, 1555, 1574, 1580, 1589, 1596, 1607, 1664, 1686, 1703, 1749, 1820, 1824, 1828, 1832, 1848, 1852, 1856, 1884, 1929, 1933, 1990, 1994, 1998, 2050, 2065, 2069, 2108, 2118, 2143, 2147, 2164, 2168, 2188, 2199, 2203, 3273, 3277, 3282, 3575, 3612, 3775, 3925, 3929, 4116], 'al': [1079], '(June': [1080], '2022)': [1081], 'PDFDownload': [1082], 'PDF': [1083], 'article': [1085], 'text': [1086], 'main': [1088], 'figures.': [1089], 'Peer': [1090], 'ReviewDownload': [1091], 'a': [1092, 1187, 1246, 1256, 1419, 1459, 1505, 1869, 1923, 2344, 2350, 2489, 2549, 2557, 2918, 3174, 3255, 3296, 3369, 3643, 3805, 3850, 3953, 4093], 'summary': [1093], 'editorial': [1096, 1100], 'decision': [1097, 1101], 'including': [1099, 1220, 1399, 1812, 2222, 2404], 'letters,': [1102], 'reviewer': [1103], 'comments': [1104], 'responses': [1107, 3734], 'feedback.': [1109], 'ToolsAdd': [1110], 'favoritesDownload': [1112], 'CitationsTrack': [1113], 'CitationsPermissions': [1114], 'Figures': [1115], 'Info': [1117], 'Abstract': [1118], 'Biomolecular': [1119], 'condensation': [1120, 1177, 1328, 1352, 1900, 2237, 2266, 2512, 2529, 2531, 2568, 2708, 3026, 3235, 3479, 3733, 3748, 3787, 3800, 3847, 3865, 4033], 'neuronal': [1123, 1420, 1469, 1568, 1788, 2250], 'microtubule-associated': [1124, 1301], 'protein': [1125, 1302, 1416, 1424, 1798, 1926, 2483, 3550, 3813, 3875, 3890], '(MAPT)': [1127], 'can': [1128, 1548, 1654, 1707, 1906, 1972, 2003, 2079, 2103, 2534, 3178, 3579], 'be': [1129, 1806, 1907, 2005, 2128, 2535, 2556, 2571, 3346, 3580, 3802, 3880], 'induced': [1130, 1305, 2006, 2224, 2536, 2911, 3337, 3368, 3705], 'coacervation': [1132, 1163, 2124, 2306, 3430, 3707], 'with': [1133, 1164, 1186, 1228, 1377, 1408, 1467, 1944, 1980, 2054, 2135, 2240, 2307, 2325, 2364, 2382, 2384, 2387, 2621, 2629, 2843, 2848, 2917, 3117, 3202, 3589, 3708, 3822, 3952, 4055, 4112, 4132, 4197], 'polyanions': [1134, 1310, 2258, 2543, 3710, 3742], 'like': [1135, 1311, 1916, 2544, 3133], 'RNA,': [1136, 1165, 2254, 2385, 2804], 'or': [1137, 1313, 1511, 1920, 2228, 2386, 2541, 2659, 2807, 2906, 3334, 3812, 3815, 4141], 'molecular': [1139, 1160, 1237, 1265, 1315, 1346, 1518, 1611, 1691, 2296, 2427, 2538, 2594, 2631, 2866, 3167], 'crowding.': [1140, 1316, 2247], 'have': [1143, 1323, 1636, 1708, 1721, 1802, 1901, 1962, 3196], 'linked': [1145], 'both': [1147, 2183, 2287, 2574, 2591, 2669, 2808, 2907, 4053], 'functional': [1148], 'binding': [1150, 1423, 1814, 2155, 2177, 2315, 2355, 2747], 'pathological': [1152, 1215, 1967, 2084, 2498], 'aggregation': [1153, 1476, 1490, 1653, 1757, 1781, 2106], 'in': [1154, 1170, 1218, 1242, 1333, 1342, 1386, 1430, 1479, 1524, 1585, 1655, 1695, 1739, 1772, 1786, 1790, 1808, 1893, 1975, 1984, 2207, 2243, 2279, 2294, 2453, 2464, 2532, 2563, 2587, 2653, 2665, 2723, 2820, 2881, 2931, 2961, 2999, 3021, 3192, 3254, 3268, 3295, 3341, 3375, 3384, 3473, 3608, 3730, 3810, 3907, 4189], 'neurodegenerative': [1155], 'diseases.': [1156], 'We': [1157, 1206, 2291, 3703, 4151], 'find': [1158, 2292, 2417], 'that': [1159, 1209, 1226, 1255, 1320, 1406, 1425, 1626, 1712, 1871, 2175, 2282, 2418, 2685, 3123, 3716, 3726, 3741, 3752, 3873, 4075, 4153, 4186], 'two': [1166, 1895, 2738, 2776], 'conditions': [1167, 1181, 1322, 3728], 'likely': [1168, 2349], 'coexisting': [1169], 'cytosol,': [1172], 'enable': [1175, 2301, 3396, 3784, 3849], 'at': [1178, 1222, 1353, 1402, 1560, 1616, 2267, 2309, 2406, 2440, 2576, 2613, 2624, 2814, 2923, 2988, 3008, 3027, 3035, 3060, 3085, 3180, 3526, 3918, 3959, 4034, 4064, 4070, 4103, 4169], 'physiological': [1179, 1354, 2815], 'buffer': [1180, 1355, 3387], 'produce': [1184, 3836], 'strong': [1188, 2365], 'affinity': [1189, 2356], 'charged': [1191, 2017, 2057, 2360, 2728, 2778], 'surfaces.': [1192], 'During': [1193, 3285], 'condensate-mediated': [1194], 'polymerization,': [1196], 'their': [1197, 1381, 3650], 'synergy': [1198], 'enhances': [1199], 'bundling': [1200, 1332, 2342], 'spatial': [1202], 'arrangement': [1203], 'microtubules.': [1205], 'further': [1207, 2446], 'show': [1208, 1319], 'different': [1210, 1236, 1395, 1658, 1709, 1722, 1809, 1896, 2419, 2501, 3036, 3061, 3086, 4035], 'efficiently': [1213], 'induce': [1214, 1284, 1583, 2099, 3721, 4144], 'aggregates': [1217, 1341, 1446, 1523, 1633, 1770, 1988, 2086, 2434], 'cells,': [1219, 2122, 3779], 'accumulations': [1221, 1244, 1662, 1675, 1698, 2423, 2445], 'nuclear': [1224, 1404, 1562, 1586, 1618, 2401, 2408, 2433, 2442], 'envelope': [1225, 1405, 1563, 1619], 'correlate': [1227, 1407], 'nucleocytoplasmic': [1229, 1409, 1569, 2449], 'transport': [1230, 1410, 1570, 2450], 'deficits.': [1231, 1411], 'Fluorescent': [1232], 'lifetime': [1233], 'imaging': [1234], 'reveals': [1235], 'packing': [1238, 1692, 2428], 'densities': [1239], 'cellular': [1243, 1294, 1396, 1723, 2519], 'condensate-like': [1247, 1400], 'density': [1248], 'nuclear-envelope': [1250], 'Tau.': [1251, 2750], 'findings': [1253], 'suggest': [1254, 4152], 'complex': [1257, 2008], 'interplay': [1258], 'between': [1259, 1345, 1763, 1947, 3263, 4023, 4086], 'interaction': [1260], 'partners,': [1261], 'post-translational': [1262], 'modifications,': [1263], 'regulates': [1267], 'formation': [1269, 1286, 1366, 1841, 2082, 2113, 2463, 2645, 2712, 2913, 3350, 3868, 3946, 3967, 4018, 4163], 'function': [1271, 3434], 'condensates.': [1274, 1957, 2028, 2091, 2371, 2484, 2856, 3627, 3702], 'Conditions': [1275], 'leading': [1276], 'prolonged': [1278], 'existence': [1279], 'may': [1283, 2261, 2555, 2569], 'seeding-competent': [1288, 2375], 'lead': [1291, 1600], 'distinct': [1293, 3842], 'accumulations.': [1296, 2521], 'SYNOPSIS': [1297], 'Condensation': [1298], 'is': [1304, 1418, 1527, 1620, 1676, 1872, 2011, 2031, 2073, 2298, 2377, 2699, 2852, 3543], 'presence': [1308, 2329, 2655], 'Here': [1317], 'we': [1318, 2213, 2275, 2416, 2598, 2683, 3237, 3309, 3402, 3739, 3943, 4028, 4181], 'these': [1321, 2486, 3970], 'synergistic': [1324], 'effects': [1325, 3145, 3227], 'Tau-dependent': [1330], 'vitro,': [1334, 1894], 'cells.': [1343, 2455], 'Synergy': [1344], 'enables': [1350, 2339, 4161], 'conditions.': [1356], 'Phosphorylation': [1357], 'content': [1360, 1519, 3814], 'change': [1361], 'Tau-condensate': [1362], 'surface': [1363, 2162, 2773], 'wetting,': [1364], 'organization': [1368, 2014], 'bundles.': [1371], 'develop': [1374], 'potential': [1376, 1969], 'time': [1378, 2990, 3271, 3528, 3597], 'regardless': [1379], 'physicochemical': [1382], 'origin,': [1383], 'even': [1385], 'absence': [1388, 2245, 2466, 3909], 'percolation': [1390, 2479], 'Aged': [1391], 'seed': [1394, 1756, 2462], 'accumulations,': [1398, 1542, 2403], 'Introduction': [1412], 'intrinsically': [1414], 'disorded': [1415], '(MT)': [1422], 'helps': [1426], 'stabilize': [1428, 3159], 'axons': [1429], 'central': [1432], 'nervous': [1433], 'system.': [1434], 'In': [1435, 1795, 2040, 2121, 2210, 2248, 3399, 3737, 3778, 3817, 3932, 4179], "Alzheimer's": [1436], 'disease': [1437, 1463], '(AD)': [1438], 'other': [1440, 2257, 3709, 3770], 'tauopathies,': [1441], 'accumulates': [1443], 'intraneuronally': [1444], 'as': [1447, 1508, 1683, 1835, 1837, 2139, 2150, 2152, 2191, 2193, 2651, 2662, 2664, 2758, 2960, 3020, 3052, 3077, 3102, 3357, 3535, 3766, 3804, 3828, 3882], 'neurofibrillary': [1448], 'tangles': [1449], '(NFTs)': [1450], 'into': [1451, 1955, 2021, 2061, 2395], 'highly': [1452], 'ordered': [1453], 'β-structured': [1454], 'aggregates,': [1456], 'which': [1457, 2010, 2471, 2942, 3244, 3344, 3496, 3578, 3661, 4119], 'are': [1458, 1465, 1482, 1728, 1753, 1774, 1784, 2596, 2943, 2957, 3136, 3497, 3513], 'hallmark': [1460], 'associated': [1466], 'progressive': [1468, 2478, 3785], 'death': [1470], '(Braak': [1471], 'Braak,': [1473], '1991).': [1474], 'spreading': [1478, 1762], 'brain': [1481, 1526, 1792], 'thought': [1483, 1730], 'involve': [1485], 'small': [1492, 3948, 4165], 'amounts': [1493], 'misfolded': [1495], '(Frost': [1497, 1595], 'Diamond,': [1499], '2010;': [1500], 'Dujardin': [1501], 'Hyman,': [1503], '2019),': [1504, 1582, 1942], 'mechanism': [1506], 'described': [1507, 1697, 3581], '"templated': [1509], 'misfolding"': [1510], '"seeded': [1512], 'aggregation".': [1513], 'Interestingly,': [1514, 2456], 'morphology': [1516, 1587, 1669], 'intraneuronal': [1521], 'variable': [1528], 'disease-specific': [1530], '(Ferrer': [1531], 'al,': [1533, 1552, 1556, 1575, 1581, 1590, 1597, 1608, 1665, 1687, 1704, 1750, 1821, 1825, 1829, 1833, 1849, 1853, 1857, 1885, 1930, 1934, 1991, 1995, 1999, 2051, 2066, 2070, 2109, 2119, 2144, 2148, 2165, 2169, 2189, 2200, 2204, 3274, 3278, 3283, 3576, 3613, 3776, 3926, 3930, 4117], '2014),': [1534, 1666], 'also,': [1536], 'subtle': [1538], 'forms': [1539, 1711, 2195], 'example': [1544, 3446, 3472], 'around': [1545], 'nucleus,': [1547], 'occur': [1549], '(Eftekharzadeh': [1550, 1573], '2018;': [1553, 1850, 1992], 'Cornelison': [1554], '2019).': [1557, 1609, 2071], 'accumulating': [1559], 'was': [1564, 3451, 3912, 4120], 'shown': [1565, 2956, 3051, 3076, 3101, 3534], 'inhibit': [1567], 'proteins': [1572], '2018)': [1576], '(Cornelison': [1579], 'changes': [1584, 3809], '(Sheffield': [1588], '2006)': [1591], 'lamin': [1593], 'folds': [1594], '2016),': [1598, 1688], 'MTs': [1602, 1818, 2181], 'invading': [1603], 'nucleoplasm': [1605], '(Paonessa': [1606], 'assembly': [1612, 1710], 'state': [1613], 'not': [1621, 2074, 2098], 'known,': [1622], 'but': [1623, 2102], 'it': [1624], 'appears': [1625], 'they': [1627, 1635, 1971], 'differ': [1628, 1701], 'from': [1629, 2473, 2770, 2854, 3138, 3469, 3558], 'amyloid-like': [1630], 'large': [1631, 3877], 'cytosolic': [1632, 2399, 2431], 'since': [1634, 1970], 'never': [1637], 'reported': [1639, 2644, 3768], 'investigations': [1641], 'pathology': [1644, 1761], 'using': [1645, 3015], 'amyloid': [1646], 'dyes.': [1647], 'Cell': [1648], 'models': [1649], 'seeded': [1651], 'part': [1656], 'recapitulate': [1657], 'types': [1659, 2397, 2420], '(Holmes': [1663], 'whereby': [1667], 'subcelluar': [1671], 'localization': [1672], 'governed': [1677], 'material': [1681], 'used': [1682, 1873, 2722, 3719, 3803], 'seeds': [1684], '(Kaufman': [1685], 'various': [1696, 1881], 'seems': [1699], '(Kaniyappan': [1702], '2020).': [1705, 1858, 2000, 2120, 2209], 'all': [1713, 2219, 2390, 3727], 'display': [1714, 2424], 'unique': [1715, 2345], 'biophysical': [1716], 'biochemical': [1718], 'characteristics': [1719], '(mis)functions.': [1724], 'Monomers': [1725], 'dimers': [1727], 'widely': [1729], 'resemble': [1732], 'soluble': [1734], '"native"': [1735], 'form': [1736, 1863, 2586, 3121, 3179, 4187], 'cytosol.': [1741, 2565], 'oligomers': [1743, 1846], 'seem': [1744, 2320], 'encode': [1746], 'neurotoxicity': [1747], '(Ward': [1748], '2012)': [1751], 'discussed': [1754], 'mediate': [1759], 'neurons': [1764], '(Gerson': [1765], 'Kayed,': [1767], '2013).': [1768], 'rich': [1771], 'β-structure': [1773], 'stable': [1776], 'end': [1777], 'product': [1778], 'processes': [1782, 2221], 'deposited': [1785], 'long-lasting': [1787], 'inclusions': [1789], '(Brion,': [1793], '1998).': [1794], 'addition,': [1796], 'liquid-like': [1797, 2027, 2062, 2647], 'suggested': [1804, 1964, 2174], 'involved': [1807], 'functions,': [1811, 2496], 'polymerization': [1816, 2334], '(Hernández-Vega': [1819, 2198], '2017;': [1822, 1931, 2067, 2201, 3275, 3927], 'Siahaan': [1823], '2019;': [1826, 1830, 2205, 3279], 'Tan': [1827, 2202], 'Zhang': [1831], '2020)': [1834, 2052], 'well': [1836, 2151, 2192, 2663], 'de': [1839], 'novo': [1840], 'competent': [1844], '(Wegmann': [1847, 1989], 'Boyko': [1851, 1993], '2020;': [1854, 1996], 'Kanaan': [1855, 1997], 'Liquid-like': [1859, 3549], 'via': [1864], 'liquid–liquid': [1865], 'separation': [1867], '(LLPS),': [1868], 'cells': [1875], 'assemble': [1877], 'membrane-less': [1878], 'organelles': [1879], 'functions': [1882], '(Hyman': [1883], '2014;': [1886], 'Mitrea': [1887], 'Kriwacki,': [1889], '2016).': [1890], 'From': [1891], 'studies': [1892], 'modes': [1897, 2425], 'identified.': [1903], 'LLPS': [1905, 2002, 2101, 2220, 2278, 2562, 2871, 2992, 3157, 3530, 3599, 3693, 3723, 4146, 4176], 'initiated': [1908], 'through': [1909, 2007, 2760, 2977, 3125, 3515, 3553, 3563, 3762], 'addition': [1911, 3363, 3481, 3938, 4138], 'macromolecular': [1913], 'agents': [1915, 2540], 'polyethylene': [1917, 2634], 'glycol': [1918, 2635], '(PEG)': [1919], 'dextrane': [1921], 'dilute': [1924, 3560], 'solution': [1927, 3269, 3298, 4190], '(Ambadipudi': [1928, 3924], '2018).': [1935, 2170, 3931], 'crowding-induced': [1937, 2089, 3831], 'colloid': [1938], 'osmotic': [1939], 'pressure': [1940], '(Mitchison,': [1941], 'together': [1943], 'attractive': [1945], 'forces': [1946, 2367], 'molecules,': [1949, 4027], 'triggers': [1950], 'de-mixing': [1952, 2811], 'liquid-dense': [1956], 'Crowding-induced': [1958], 'biomolecular': [1959, 2502, 3626, 3771], 'harbor': [1966], 'convert—at': [1973], 'least': [1974], 'vitro—into': [1976], 'oligomeric': [1977], 'species': [1979], 'properties': [1981], 'similar': [1982, 2087, 2378, 2396, 3370, 3731, 3980], 'vitro': [1985, 2533], 'generated': [1986], 'also': [2004, 3704], 'coacervation,': [2009], 'spontaneous': [2013, 3867, 3904, 3949, 4048], 'oppositely': [2016], '(parts': [2018], 'of)': [2019], 'biomolecules': [2020], 'higher': [2022], 'order': [2023], 'assemblies,': [2024], 'example,': [2026, 2127, 3827], 'This': [2029, 2751], 'driven': [2032], 'electrostatic': [2034, 2761, 3164, 4021, 4084, 4193], 'interactions': [2035, 3168, 4022, 4085, 4194], 'engaging': [2038], 'molecules.': [2039, 4088], 'case': [2042], 'Tau,': [2044, 2326, 2606, 3903], 'polyanionic': [2045, 2241, 2755], '(or': [2047], 'heparin;': [2048], 'Lin': [2049, 2068], 'co-condensate': [2053], 'positively': [2056, 2727], 'microtubule-assembly': [2059], 'domain': [2060], 'droplets': [2063, 3876], '(Zhang': [2064], 'It': [2072, 2171], 'known': [2075], 'whether': [2076, 2582], 'Tau:RNA': [2077, 2458, 3160, 3397, 3653], 'coacervates': [2078, 2459, 3161], 'trigger': [2080, 2104, 3156], 'Notably,': [2092, 3378, 4089], 'high': [2093, 2354], 'concentrations': [2094, 2816, 3919, 4037, 4126], 'do': [2097], '(Kampers': [2107], '1996)': [2110], 'without': [2111], 'liquid-condensed': [2115, 2194], 'intermediates': [2116], '(Lin': [2117], 'may,': [2125], 'important': [2129], 'co-condensation': [2132, 2324, 2838], 'RNA-containing': [2136], 'ribonucleoparticles': [2137], 'such': [2138, 2757, 3883], 'stress': [2140], 'granules': [2141], '(Vanderweyde': [2142], '2016;': [2145], 'Ash': [2146], '2021),': [2149, 3577], 'anionic': [2160], 'MT': [2161, 2333, 2341, 2361], '(Mukrasch': [2163], '2007;': [2166], 'Kadavath': [2167], 'involves': [2182], 'single': [2184, 3541], 'molecules': [2186, 3152], '(Janning': [2187], '2014)': [2190], 'reviewed': [2206], 'Mitchison,': [2208], 'following,': [2212, 3401], 'will': [2214], 'use': [2215, 3403], '"condensation"': [2216], 'describe': [2218, 3225], 'those': [2223], 'crowding,': [2226], 'polyanions,': [2227], 'both.': [2229], 'term': [2231], '"coacervation"': [2232], 'refers': [2233], 'solely': [2234], 'co-factors,': [2242], 'cytoplasm,': [2251], 'encounters': [2253], 'microtubules,': [2255], 'thus': [2260, 2570], 'undergo': [2262], 'crowding-': [2263], 'polyanion-induced': [2265], 'same': [2269, 2578, 2959, 3017], 'time.': [2270, 2579], 'To': [2271, 2580, 3224, 4013], 'mimic': [2272], 'scenario,': [2274], 'characterized': [2276], 'model': [2280, 2476, 3019], 'systems': [2281, 2667, 3698], 'consider': [2283], 'contribution': [2285, 3645, 3685], 'polyanions.': [2290], 'that,': [2293], 'fact,': [2295], 'necessary': [2299], 'phosphorylated': [2304], 'intracellular': [2310], 'ion': [2311, 3182, 4125], 'compositions.': [2312], 'partners': [2316], 'tubulin': [2319], 'compete': [2322], 'during': [2332, 2527, 2869, 3649, 3746], 'out': [2335], 'pronounced': [2340], 'geometric': [2346], 'bundle': [2347], 'arrangement,': [2348], 'result': [2351], '(wetting)': [2357], 'negatively': [2359, 2777], 'surfaces': [2362], 'combined': [2363], 'cohesive': [2366], 'inside': [2368, 2691, 3169], 'Tau:PEG:RNA': [2370, 2973, 3176, 3632], 'generation': [2373], 'PEG,': [2383, 2904, 3942], 'both,': [2388], 'aged': [2391], 'convert': [2394], 'envelope.': [2409, 2443], 'Combining': [2410], 'FRAP': [2411], 'FLIM': [2413], 'Tau-CFP,': [2415], 'reminiscent': [2429, 3984], 'versus': [2435], 'less': [2436, 3684, 3851], 'densely': [2437], 'packed': [2438], 'enhance': [2447], 'Tau-induced': [2448], 'deficits': [2451], 'present': [2452], 'Tau-expressing': [2454], 'binary': [2457], 'condensate': [2468, 2974, 3511, 3593, 3820, 3824], 'liquid-to-gel': [2469], 'transition,': [2470], 'deviates': [2472], 'current': [2475], 'formulated': [2480], 'RNA-binding': [2482], 'Collectively,': [2485], 'observations': [2487], 'provide': [2488], 'framework': [2490], 'explain': [2492], 'formation,': [2494], 'interactions,': [2495], 'activities': [2499], 'thereby': [2506, 3155], 'improve': [2507], 'understanding': [2509], 'how': [2511], 'contributes': [2513], 'biology': [2516], 'disease-related': [2518], 'Results': [2522], 'RNA.': [2545, 3487], 'cell': [2547], 'generally': [2550], 'crowded': [2551], 'environment,': [2552], 'potent': [2558], 'inducer': [2559, 2993, 3531], 'Cellular': [2566], 'influenced': [2572], 'factors': [2575], 'test': [2581, 3204, 4014], 'could': [2585, 3345, 3801, 3848], 'an': [2588, 2798, 3338], 'environment': [2589], 'where': [2590], 'present,': [2597], 'mixed': [2599], 'human': [2600, 2718], 'full-length': [2601], 'wild-type': [2602], '(5': [2604, 2817, 2845], 'µM': [2605, 2846, 3300, 3304, 3961, 3989, 4001, 4105], '2N4R': [2607], 'isoform,': [2608], '441': [2609], 'aa,': [2610], 'net': [2611], 'charge': [2612, 2625, 2785], 'pH': [2614, 2827], '7.4': [2615], '=': [2616, 2831, 2859, 2981, 3040, 3065, 3090, 3185, 3519, 3587, 3635, 3667, 3676, 3992, 3998, 4009], '+1.4': [2617], '(unphosphorylated);': [2618], 'Fig': [2619, 2674, 2962, 4011], '1A)': [2620, 4160], 'polyA': [2622, 2670, 2803, 3331, 3365], 'matching': [2626], 'concentrations,': [2627], 'agent': [2633, 2697], '(PEG8000;': [2636], '5%': [2637], '(w/vol)).': [2638], 'Light': [2639], 'microscopy': [2640, 2836, 3917], 'confirmed': [2641], 'previously': [2643, 3718, 3767, 3913], 'phases': [2649], '(observed': [2650], 'droplets)': [2652], '(Tau:RNA)': [2658], 'PEG': [2660, 2672, 2698, 2805, 3134, 3148, 3335, 3367, 4142], '(Tau:PEG),': [2661], 'tertiary': [2666], 'containing': [2668], '(Tau:PEG:RNA;': [2673], '1B).': [2675], 'Using': [2676], 'fluorescently': [2677], 'labeled': [2678], 'Tau-Dylight488,': [2679], 'polyA-Cy3,': [2680], 'PEG-Dylight647,': [2682], 'observed': [2684, 3310, 3829, 3914, 3944], 'become': [2689], 'enriched': [2690], 'condensates,': [2693, 3833, 3872], 'whereas': [2694], 'excluded': [2700, 2853, 3137, 3143], '(Fig': [2701, 3695, 3735, 3963, 4038, 4045, 4051, 4127, 4147, 4159, 4177], '1C).': [2702], 'Figure': [2703, 3424], '1.': [2704], 'Kinetics': [2705, 2968, 2995, 3509], 'mesoscopic': [2710, 2896, 2929, 4183], 'cluster': [2711, 2997, 4017], 'Domain': [2713], 'structure': [2714], 'longest': [2717], 'isoform': [2720], '(2N4R)': [2721], 'study:': [2725], 'C-terminal': [2729], 'half': [2730], 'contains': [2733], 'four': [2734], 'repeats': [2735], '(R1-R4)': [2736], 'proline-rich': [2739], 'regions': [2740], '(P1,': [2741], 'P2),': [2742], 'comprises': [2744], 'region': [2748, 2752], 'binds': [2753], 'macromolecules': [2756], 'interactions.': [2762], 'N-terminal': [2764, 2779], '~200': [2765], 'amino': [2766], 'acids': [2767], '(aa)': [2768], 'project': [2769], 'contain': [2775], 'inserts': [2780], '(N1,': [2781], 'N2).': [2782], 'homogenous': [2784], 'distribution': [2786, 2864, 3251], '(sliding': [2787], 'window': [2788], '10': [2790, 2832, 2860, 3005, 3043, 3392, 4108], 'aa)': [2791], 'along': [2792], 'its': [2793], 'aa': [2794], 'sequence': [2795], 'assigns': [2796], 'amphiphilic': [2799, 4155], 'character.': [2800], 'Addition': [2801, 2902, 3329], '(PEG8000),': [2806], 'induces': [2809], 'μM': [2818, 2879, 3010, 3013, 3922], '25': [2821, 2878], 'mM': [2822, 2825, 2885, 3003, 3006, 3389, 3393, 4066, 4072, 4109], 'HEPES,': [2823, 3004, 3390], 'DTT,': [2826], '7.4).': [2828], 'Scale': [2829, 2857], 'bars': [2830], 'μm.': [2833, 2861], 'Confocal': [2834], 'fluorescence': [2835], 'shows': [2837], 'polyA-Cy3': [2840], '(10:1,': [2841], 'polyA:polyA-Cy3)': [2842], '1%': [2849], 'Tau-DyLight488).': [2850], 'PEG8000-Dylight647': [2851], 'bar': [2858], 'Representative': [2862, 3475], 'size': [2863, 3250, 4041], 'assemblies': [2868], 'measured': [2872], 'time-resolved': [2874, 3239], 'DLS': [2875, 2940, 3494], '(trDLS).': [2876], 'At': [2877], 'low': [2882, 3000, 3385], 'salt': [2883, 3001, 3386, 4057], '(10': [2884], 'NaCl)': [2886, 3007, 3394], 'buffer,': [2887], 'trDLS': [2888, 2985, 3023, 3380, 3426, 3447, 3462, 3476, 3523, 3934, 4030], 'detects': [2889], 'monomers': [2891, 2927, 3084, 3111, 3312, 3557, 4134], '(radii': [2892, 2899], '~10': [2893], 'nm)': [2894, 3314, 3641], 'clusters': [2898, 2930, 3059, 3115, 3320, 3860, 3951, 3971, 4050, 4062, 4079, 4130, 4168, 4185], '~100–200': [2900], 'nm).': [2901], 'polyA,': [2905], '~15': [2909, 3288], 'min': [2910, 3289], 'radius': [2919, 3186, 3257, 3259, 3954, 4094], '>103': [2921], 'nm,': [2922, 3996, 4007], 'cost': [2925], 'solution.': [2933, 3377], 'point': [2935, 3489], 'bubble': [2936, 3490], 'sizes': [2937, 3491, 3610], 'correspond': [2938, 3492], 'amplitudes,': [2941, 3495], 'proportional': [2944, 3498], 'intensity': [2947, 3501], 'scattered': [2949, 3503], 'light': [2950, 3241, 3360, 3504, 3885, 3916], 'respective': [2953, 3507], 'particles.': [2954, 3508], '2D': [2963], '0': [2965], '<t<45': [2966], 'min.': [2967], 'Tau:PEG,': [2970], 'Tau:RNA,': [2971], 'growth': [2975, 2998, 3512, 3620, 3629, 3754, 3761, 3781, 3821], 'assessed': [2976, 3514], 'power': [2978, 3516, 3584], 'law,': [2979, 3517], 'R': [2980, 3518, 3586, 3590], 'a*tb,': [2982, 3520, 3588], 'fitting': [2983, 3018, 3521], 'data': [2986, 3045, 3050, 3070, 3075, 3095, 3100, 3427, 3438, 3450, 3471, 3524], 'starting': [2987, 3525], 'addition.': [2994, 3532], '(25': [3002, 3299, 3388], '30': [3009, 4104], '50': [3012, 3303, 4000], '(E).': [3022], 'Tau:PEG': [3025, 3033, 3306, 3655, 3697, 4043], 'increasing': [3028, 4056, 4124], 'NaCl': [3029, 3037, 3062, 3087, 4036, 4067], 'concentrations.': [3030, 3038, 3063, 3088, 3183], 'Radius': [3031, 3056, 3081], 'N': [3039, 3064, 3089], '3': [3041, 3068, 3093], 'experiments,': [3042, 3067, 3092], 'last': [3044, 3069, 3094], 'points': [3046, 3071, 3096], 'per': [3047, 3072, 3097], 'experiments': [3048, 3073, 3098], 'shown,': [3049, 3074, 3099], 'mean': [3053, 3078, 3103, 3536], '±': [3054, 3079, 3104, 3537, 3639, 3671, 3680, 3994, 4005, 4097, 4101], 'SEM.': [3055, 3080, 3105], '2–3': [3066, 3091], 'Model': [3106], 'condensation:': [3109], '(grey': [3112], 'circles)': [3113, 3436], 'coacervate': [3116], '(pink)': [3119], 'grow': [3124, 3552], 'droplet': [3126, 3564], 'fusion': [3127, 3763], '(Oswald': [3128], 'ripening).': [3129], 'agents,': [3132], '(blue),': [3135], 'dense': [3140, 3171], 'phase.': [3141, 3172], 'Inset:': [3142], 'volume': [3144], 'exerted': [3146], 'force': [3149], 'individual': [3150], 'interact,': [3154], 'against': [3162], 'shielding': [3165], 'As': [3173], 'result,': [3175], 'cytosol-like': [3181], 'R0': [3184], 'hydration.': [3188], 'information:': [3190], 'H,': [3193], 'I,': [3194], 'J': [3195], 'compared': [3198, 3689], 'one-way': [3200], 'ANOVA': [3201], 'Tukey': [3203], 'multiple': [3206], 'comparison.': [3207], '*P': [3208], '<': [3209, 3212, 3215, 3218, 4171], '0.05,': [3210], '**P': [3211], '0.01,': [3213], '***P': [3214], '0.001,': [3216], '****P': [3217], '0.0001.': [3219], 'Download': [3220, 3222, 3545, 3547], 'figure': [3221, 3546], 'PowerPoint': [3223, 3548], 'process,': [3236], 'performed': [3238, 4029], 'dynamic': [3240, 3806], 'scattering': [3242], '(trDLS),': [3243], 'allows': [3245], 'following': [3246], 'evolution': [3248], 'particles': [3253], 'particle': [3256, 3293, 3342, 3373, 3609], '(R0,': [3258], 'hydration)': [3261], 'range': [3262], '100': [3264, 4065], '105': [3266], 'nm': [3267, 3957, 4099], 'over': [3270], '(Schubert': [3272], 'Van': [3280], 'Lindt': [3281, 3774], '2021).': [3284, 3777], 'measuring': [3291], 'radii': [3294, 3374, 3467], 'Tau:polyA,': [3302], 'Tau:PEG:polyA),': [3308], '(RMono~10': [3313], 'spontaneously': [3316, 4076, 4188], 'formed': [3317, 4063, 4077], '"mesoscopic"': [3318], '(RClust~100–200': [3321], 'nm;': [3322, 3673], 'Figs': [3323], '1D': [3324, 4148], 'EV1A': [3326], 'B).': [3328], '(50': [3332], 'µg/ml)': [3333], '(5%)': [3336], 'immediate': [3339], 'increase': [3340, 3371, 3607], 'radius,': [3343, 3594], 'attributed': [3347], '"microscopic"': [3352], '(RCond~1,000': [3355], 'nm),': [3356, 3682], 'judged': [3358], 'microscopy.': [3361], 'Simultaneous': [3362], 'measurements': [3381], 'were': [3382, 3717], 'recorded': [3383, 3452], 'pH7.4,': [3391], 'coacervation.': [3398], 'descriptions': [3405], 'monomers,': [3406], 'clusters,': [3407], 'refer': [3411], 'degree': [3414], 'assembling.': [3417], 'Click': [3418], 'here': [3419], 'expand': [3421], 'figure.': [3423], 'EV1.': [3425], 'Averaged': [3431], 'experimental': [3432], 'autocorrelation': [3433, 3470], '(black': [3435], 'fit': [3439], 'applying': [3440], 'CONTIN': [3441], 'analysis': [3442], '(red': [3443], 'line)': [3444], 'data.': [3448], 'Similar': [3449], 'fitted': [3454], 'each': [3456], '20': [3457], 's': [3458], 'interval': [3459], 'measurements.': [3463], 'Distribution': [3464], 'hydrodynamic': [3466], 'derived': [3468], '(A).': [3474], 'upon': [3480, 4137], 'heparin,': [3483], 'tRNA,': [3484, 3712], 'polyU': [3486, 3714], 'SD.': [3538], 'A': [3539], 'representative': [3540], 'experiment': [3542], 'show.': [3544], 'recruitment': [3555], 'coalescence': [3565], '(Ostwald': [3566], 'ripening;': [3567], 'Lifshitz': [3568], 'Slyozov,': [3570], '1961;': [3571], 'Siggia,': [3572], '1979;': [3573], 'Lee': [3574], 'law': [3585], 'being': [3591], 't': [3595], 'initiation,': [3600], 'b': [3602, 3615, 3666, 3675, 3997, 4008], 'coarseningexponent': [3604], 'describing': [3605], '(Berry': [3611], '2018);': [3614], '~': [3616], '1/3': [3617], 'describes': [3618], 'Ostwald': [3622, 3647, 3687, 3758, 3986], 'ripening': [3623, 3648, 3688, 3759, 3987], 'most': [3625], 'kinetics': [3630, 3755, 3968, 3979], '(b': [3634], '0.44;': [3636], 'R25min:': [3637, 3669, 3678], '1167': [3638], '103': [3640], 'indicated': [3642], 'major': [3644], 'formation.': [3651], 'Binary': [3652], 'showed': [3657], 'fast': [3658], 'initial': [3659], 'growth,': [3660], 'then': [3662], 'slowed': [3663], 'down': [3664], '(polyA:': [3665], '0.15;': [3668], '1460': [3670], '78': [3672, 3681], 'PEG:': [3674], '0.18;': [3677], '886': [3679], 'indicating': [3683, 4074], 'triple': [3692], 'system': [3694], '1E).': [3696], 'produced': [3699], 'smallest': [3701], '(heparin,': [3711], 'RNA)': [3715], 'found': [3725, 3740], 'resulted': [3729], 'EV1C).': [3736], 'summary,': [3738, 4180], 'producing': [3749], 'exhibit': [3753], 'indicative': [3756], '(=': [3760], 'condensates)': [3765], '(Van': [3773], 'behavior': [3782], 'would': [3783, 3834], 'limited': [3788, 3823, 3840], 'availability': [3791], 'and/or': [3794], 'RNA;': [3795], 'type': [3797, 3845], 'response': [3807], 'concentration.': [3816, 4058], 'contrast,': [3818], 'slow': [3819], 'fusion,': [3825], 'rapidly': [3835], 'size;': [3843], 'responsive': [3852], 'on/off': [3853], 'switch': [3854], 'condensation.': [3857], 'Mesoscopic': [3858], 'below': [3861, 4080], 'csat': [3862, 3897, 4081], 'microscopic': [3864, 3870, 3905, 4145, 4175], 'liquid': [3871], 'is,': [3874], 'enough': [3878], 'identified': [3881, 4182], 'microscopy,': [3886], 'occurs': [3887], 'when': [3888], 'concentration': [3891, 3976], 'exceeds': [3892], 'critical': [3894], 'saturation': [3895], 'concentration,': [3896], '(Shin': [3898], 'Brangwynne,': [3900], '2017).': [3901], 'For': [3902], 'condensation,': [3906], 'cofactors,': [3911], '~50–100': [3921], 'our': [3933], 'measurements,': [3935], 'before': [3936], '~150–250': [3956], 'already': [3958], '25–50': [3960], '1D).': [3964], 'Size': [3965], 'depended': [3972, 4019, 4082], 'followed': [3978], 'coacervates,': [3983], '(30': [3988], 'Tau:': [3990, 4002], 'R12min': [3991], '173.0': [3993], '10.7': [3995], '0.30;': [3999], 'R12min:': [4003], '230.8': [4004], '22.7': [4006], '0.45;': [4010], '1F).': [4012], 'if': [4015], 'within': [4025], '1G).': [4039], '1H)': [4046], '1I)': [4052], 'decreased': [4054], 'Only': [4059], 'few': [4060], 'none': [4069], '150': [4071], 'NaCl,': [4073, 4110], 'monomeric': [4090, 4198], 'had': [4092], '9.6': [4096], '1.0': [4098], '(mean': [4100], 'SD)': [4102], 'consistent': [4111], 'previous': [4113], 'reports': [4114], '(Mylonas': [4115], '2008),': [4118], 'rather': [4121], 'insensitive': [4122], '1J).': [4128], 'coexisted': [4131], 'disappeared': [4136], 'G).': [4150], 'character': [4156], '(=mesoscopic)': [4166], 'c': [4170], 'csat,': [4172], 'therefore': [4173], 'preceding': [4174], '1K).': [4178], 'based': [4191], 'coexist': [4196], 'T': [4199]}, 'cited_by_api_url': 'https://api.openalex.org/works?filter=cites:W4221061896', 'counts_by_year': [{'year': 2024, 'cited_by_count': 15}, {'year': 2023, 'cited_by_count': 24}, {'year': 2022, 'cited_by_count': 9}], 'updated_date': '2024-12-27T01:51:17.071101', 'created_date': '2022-04-03'}
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