Large‐scale image‐based profiling of single‐cell phenotypes in arrayed CRISPR‐Cas9 gene perturbation screens

Name: Work Video:
Duration: 3 min 30 s
Reinoud E. A. de Groot; Joel Lüthi; Helen Lindsay; René Holtackers; Lucas Pelkmans
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2392, 2409, 2560, 2638, 2952, 3054, 3279, 3378, 3410, 3504, 3923, 3973, 4161], 'screens': [17, 603, 691, 697], 'Reinoud': [18, 21, 150, 153, 284], 'de': [19, 22, 151, 154], 'Groot': [20, 23, 152, 155], 'orcid.org/0000-0002-1905-6501': [24, 156], 'Institute': [25, 47, 86, 107, 133, 157, 179, 218, 239, 265], 'Molecular': [27, 49, 88, 109, 135, 159, 181, 220, 241, 267, 298, 334], 'Life': [28, 50, 61, 89, 110, 136, 160, 182, 193, 221, 242, 268, 299, 310], 'Sciences,': [29, 51, 90, 111, 137, 161, 183, 222, 243, 269, 300], 'University': [30, 52, 69, 91, 112, 138, 162, 184, 201, 223, 244, 270, 301, 318], 'Zürich,': [32, 33, 54, 55, 71, 72, 93, 94, 114, 115, 140, 141, 164, 165, 186, 187, 203, 204, 225, 226, 246, 247, 272, 273, 303, 304, 320, 321], 'Switzerland': [34, 56, 73, 95, 116, 142, 166, 188, 205, 227, 248, 274, 305, 322], 'Search': [35, 74, 96, 117, 143, 167, 206, 228, 249, 275], 'for': [36, 75, 97, 118, 144, 168, 207, 229, 250, 276, 395, 451, 486, 557, 857, 1024, 1054, 1079, 1083, 1091, 1272, 1414, 1418, 1618, 1841, 1941, 2129, 2172, 2191, 2313, 2434, 2480, 2516, 2553, 2566, 2575, 2662, 2775, 3288, 3678, 3707, 3798, 3845], 'more': [37, 76, 98, 119, 145, 169, 208, 230, 251, 277, 1724, 3212, 3229, 3259, 3956], 'papers': [38, 77, 99, 120, 146, 170, 209, 231, 252, 278], 'by': [39, 78, 100, 121, 147, 171, 210, 232, 253, 279, 540, 570, 743, 848, 1116, 1126, 1229, 1241, 1258, 1338, 1467, 1487, 2073, 2435, 2856, 2934, 2967, 3121, 3142, 3248, 3657, 3937, 4079, 4188, 4220], 'this': [40, 79, 101, 122, 148, 172, 211, 233, 254, 280, 449, 681, 751, 1020, 1477, 1842, 1926, 2147, 2257, 2642, 2712, 2855, 2905, 2920, 3496, 3651], 'author': [41, 80, 102, 123, 149, 173, 212, 234, 255, 281, 362], 'Joel': [42, 44, 174, 176, 286], 'Lüthi': [43, 45, 175, 177], 'orcid.org/0000-0003-3023-170X': [46, 178], 'Systems': [57, 189, 335], 'Biology': [58, 190, 307, 336], 'PhD': [59, 191, 308], 'program,': [60, 192, 309], 'Science': [62, 194, 311], 'Zürich': [63, 67, 195, 199, 312, 316], 'Graduate': [64, 196, 313], 'School,': [65, 197, 314], 'ETH': [66, 198, 315], 'and': [68, 200, 292, 317, 343, 361, 382, 447, 456, 481, 522, 526, 576, 600, 607, 638, 667, 768, 851, 939, 1047, 1076, 1096, 1119, 1145, 1173, 1201, 1206, 1219, 1222, 1266, 1276, 1341, 1383, 1411, 1614, 1616, 1704, 1806, 1809, 1863, 1885, 1895, 1900, 1902, 1920, 1956, 1992, 2021, 2054, 2159, 2166, 2174, 2184, 2188, 2224, 2247, 2274, 2311, 2333, 2351, 2353, 2374, 2379, 2394, 2424, 2462, 2609, 2630, 2678, 2756, 2782, 2846, 2882, 2944, 2991, 2993, 3020, 3033, 3050, 3087, 3100, 3104, 3154, 3172, 3224, 3228, 3293, 3317, 3333, 3403, 3413, 3423, 3458, 3522, 3549, 3587, 3616, 3680, 3705, 3712, 3763, 3795, 3806, 3819, 3852, 3863, 3909, 3979, 4021, 4023, 4067, 4069, 4110, 4118, 4120, 4165, 4199, 4201, 4216, 4238], 'Helen': [81, 83, 213, 215, 288], 'Lindsay': [82, 84, 214, 216], 'orcid.org/0000-0002-3574-1541': [85, 217], 'René': [103, 105, 235, 237, 290], 'Holtackers': [104, 106, 236, 238], 'Lucas': [124, 128, 256, 260, 293], 'Pelkmans': [125, 129, 257, 261, 294], 'Corresponding': [126, 258], 'Author': [127, 259, 282], '[email': [130, 262, 332], 'protected]': [131, 263, 333], 'orcid.org/0000-0002-6754-9730': [132, 264], 'Information': [283], 'Groot1,': [285], 'Lüthi1,2,': [287], 'Lindsay1,': [289], 'Holtackers1': [291], '*,1': [295], '1Institute': [296], '2Systems': [306], '*Corresponding': [323], 'author.': [324], 'Tel:': [325], '+41': [326], '44': [327], '63': [328], '53': [329], '123;': [330], 'E-mail:': [331], '(2018)14:e8064https://doi.org/10.15252/msb.20178064': [337], 'PDFDownload': [338], 'PDF': [339], 'article': [341], 'text': [342, 1271, 1613], 'main': [344, 1612], 'figures.': [345], 'Peer': [346], 'ReviewDownload': [347], 'a': [348, 423, 433, 437, 470, 544, 585, 612, 631, 702, 844, 920, 996, 1084, 1093, 1097, 1104, 1132, 1245, 1285, 1289, 1309, 1399, 1403, 1433, 1437, 1502, 1506, 1539, 1543, 1576, 1580, 1670, 1746, 1751, 1761, 1767, 1817, 1887, 1963, 2011, 2342, 2355, 2382, 2414, 2419, 2467, 2484, 2491, 2496, 2544, 2558, 2570, 2595, 2668, 2679, 2696, 2719, 2730, 2746, 2750, 2821, 2837, 2861, 2900, 2914, 2923, 3004, 3059, 3266, 3302, 3353, 3555, 3592, 3727, 4058, 4088, 4095, 4168, 4204, 4229], 'summary': [349], 'the': [351, 396, 399, 457, 463, 529, 558, 580, 588, 628, 649, 664, 757, 813, 821, 926, 962, 986, 992, 1007, 1025, 1030, 1055, 1058, 1073, 1088, 1123, 1138, 1198, 1210, 1333, 1347, 1378, 1454, 1634, 1659, 1684, 1717, 1799, 1811, 1830, 1845, 1864, 1893, 1898, 1905, 1939, 1942, 1959, 1989, 1995, 1998, 2005, 2032, 2037, 2043, 2049, 2055, 2060, 2068, 2083, 2100, 2105, 2192, 2200, 2264, 2275, 2300, 2337, 2360, 2363, 2368, 2404, 2440, 2453, 2506, 2514, 2634, 2646, 2652, 2658, 2674, 2685, 2703, 2771, 2776, 2779, 2787, 2793, 2799, 2807, 2815, 2842, 2858, 2908, 2931, 2950, 2958, 2996, 3039, 3044, 3053, 3083, 3096, 3116, 3144, 3161, 3166, 3173, 3189, 3193, 3198, 3202, 3208, 3215, 3221, 3232, 3237, 3243, 3284, 3306, 3328, 3345, 3368, 3377, 3384, 3389, 3394, 3404, 3408, 3432, 3454, 3462, 3474, 3517, 3546, 3550, 3568, 3571, 3584, 3598, 3639, 3688, 3692, 3720, 3741, 3751, 3758, 3775, 3785, 3799, 3802, 3822, 3837, 3850, 3864, 3867, 3880, 3884, 3891, 3921, 3931, 3943, 3965, 3970, 3990, 4006, 4013, 4092, 4103, 4130, 4138, 4146, 4160, 4173, 4179], 'editorial': [352, 356], 'decision': [353, 357], 'including': [355, 3024], 'letters,': [358], 'reviewer': [359], 'comments': [360], 'responses': [363], 'to': [364, 367, 435, 477, 643, 699, 756, 771, 834, 840, 929, 998, 1018, 1028, 1072, 1460, 1475, 1652, 1682, 1692, 1749, 1813, 1924, 1987, 2023, 2145, 2259, 2306, 2328, 2330, 2439, 2452, 2456, 2458, 2605, 2651, 2656, 2692, 2798, 2887, 2919, 2930, 3007, 3128, 3147, 3179, 3252, 3261, 3314, 3348, 3383, 3494, 3545, 3583, 3649, 3723, 3730, 3750, 3912, 4049, 4172, 4208], 'feedback.': [365], 'ToolsAdd': [366], 'favoritesDownload': [368], 'CitationsTrack': [369], 'CitationsPermissions': [370], 'ShareFacebookTwitterLinked': [371], 'InMendeleyWechatReddit': [372], 'Figures': [373], '&': [374, 620, 953, 1836, 1860, 2803, 2876, 3873, 4082, 4153, 4182], 'Info': [375], 'Abstract': [376], 'High-content': [377], 'imaging': [378], 'using': [379, 502, 828, 1372, 1874, 2868, 3071, 3554, 3591], 'automated': [380, 829, 1267, 1633], 'microscopy': [381, 830], 'computer': [383, 1264, 2167], 'vision': [384, 1265, 2168], 'allows': [385, 893], 'multivariate': [386, 498, 532, 733, 967, 1888, 2762, 3117, 4017, 4211], 'phenotypes.': [390, 535, 737, 837, 2410], 'Here,': [391, 2726, 4091], 'we': [392, 1081, 1136, 1632, 1732, 1765, 2298, 2319, 2644, 2649, 2701, 2714, 2810, 2826, 2980, 3068, 3110, 3138, 3219, 3235, 3834, 3877, 3925, 3947], 'present': [393, 1015, 1825], 'methods': [394, 839, 1617], 'application': [397, 1056], 'CISPR-Cas9': [400], 'system': [401, 507, 651, 928, 1009, 1060, 1086], 'large-scale,': [403, 551, 1752], 'image-based,': [404], 'experiments.': [407, 2230], 'We': [408, 431, 468, 1014, 1049, 1107, 1195, 1648, 1668, 1706, 1743, 1794, 1839, 1876, 2255, 2616, 2767, 2852, 3002, 3037, 3163, 3840, 3897, 4227], 'show': [409, 1208, 2557, 4057], 'that': [410, 425, 473, 710, 773, 879, 882, 965, 1176, 1209, 1731, 1778, 1820, 1984, 2121, 2486, 2556, 2618, 2648, 2706, 2904, 2938, 3010, 3027, 3052, 3091, 3109, 3131, 3245, 3351, 3876, 3969, 3996, 4026, 4045, 4159, 4244], 'CRISPR-Cas9-mediated': [411, 1226, 1238], 'can': [414, 668, 707, 739, 774, 984, 1733, 1870, 2431, 3111], 'be': [415, 644, 740, 775, 985, 1872, 2244, 2432, 2946, 4135], 'achieved': [416], 'human': [418, 1112, 1485], 'tissue': [419, 1113], 'culture': [420, 1114], 'cells': [421, 485, 521, 523, 578, 606, 763, 842, 875, 900, 1115, 1144, 1153, 1160, 1178, 1188, 1205, 1216, 1282, 1323, 1371, 1396, 1430, 1486, 1499, 1526, 1536, 1563, 1573, 1600, 1685, 1797, 1812, 1880, 1899, 2157, 2170, 2216, 2302, 2335, 2376, 2399, 2464, 2541, 2555, 2568, 2579, 2592, 2600, 2694, 2705, 2728, 2738, 2743, 2758, 2769, 2785, 2812, 2830, 2843, 2859, 2885, 2896, 2928, 2940, 3047, 3156, 3402, 3406, 3439, 3443, 3468, 3472, 3605, 3609, 3621, 3682, 3932, 4000, 4056, 4086, 4141, 4215, 4240], 'timeframe': [424], 'is': [426, 508, 548, 769, 831, 975, 1101, 1212, 1363, 1852, 1954, 1979, 2000, 2008, 2478, 2623, 2818, 3258, 3301, 3380, 3552, 3755, 3975, 4027, 4072, 4076, 4186, 4206, 4217], 'compatible': [427], 'with': [428, 550, 747, 944, 1034, 1070, 1263, 1284, 1398, 1432, 1501, 1538, 1575, 1639, 1710, 1781, 1798, 1815, 1982, 2104, 2117, 2218, 2490, 2543, 2569, 2594, 2667, 2695, 2745, 2792, 2831, 2844, 2913, 3197, 3445, 3513, 3564, 3611, 3623, 3933, 4002, 4031], 'phenotyping.': [430], 'developed': [432, 556, 856, 1745], 'pipeline': [434, 1748], 'construct': [436], 'large-scale': [438, 688, 1931, 2193, 2267], 'library': [440, 450, 1754, 1768, 1935, 3287], '2,281': [442, 1771], 'sequence-verified': [443, 1756], 'targeting': [445, 546, 1109, 1233, 1246, 1291, 1311, 1343, 1405, 1439, 1457, 1491, 1508, 1545, 1582, 1712, 1718, 1758, 1775, 2078, 2220, 2415, 2493, 2498, 2546, 2597, 3321, 3447, 3613, 3843, 3964, 4005], 'plasmids': [446, 1110, 1234, 1458, 1492, 1713, 1719, 1800, 2219, 3446, 3612, 3844], 'profiled': [448, 569, 3220, 3236], 'genes': [452, 635, 662, 709, 904, 1199, 1464, 1709, 1777, 2071, 2103, 2110, 2120, 2961, 3130], 'affecting': [453, 3290], 'cellular': [454, 805, 809, 2965, 2975, 2989, 3063, 3118, 3291], 'morphology': [455, 2990, 3223, 3292, 3521, 3641, 3673, 3808, 3893, 4109], 'subcellular': [458, 814, 1865], 'localization': [459, 1866], 'components': [461, 1869, 2223, 3326, 3452, 3848, 4177], 'nuclear': [464, 589, 822, 1831, 2185, 2800, 4108], 'pore': [465, 590, 823, 1832], 'complex': [466, 591, 824, 1833, 3217], '(NPC).': [467, 825], 'conceived': [469], 'machine-learning': [471], 'method': [472], 'harnesses': [474], 'genetic': [475, 602, 655, 671, 677, 1044, 1336, 1367, 1482], 'heterogeneity': [476, 2370], 'score': [478, 2357, 3011, 3300, 3356, 3753, 3787], 'perturbations': [480, 1368, 3023, 3026, 3090, 3289, 3337, 3350, 3514, 3565, 3725, 3830, 4047], 'identify': [482, 708, 1988, 3129], 'phenotypically': [483, 2397, 2507, 2624, 2675, 2708, 2909, 2942, 3045, 3441, 3470, 3603, 3944, 4224], 'perturbed': [484, 906, 1187, 1351, 1386, 1720, 2375, 2398, 2426, 2508, 2625, 2676, 2910, 3046, 3442, 3471, 3604, 3945, 4001, 4225], 'in-depth': [487], 'characterization': [488, 2519], 'effects.': [492, 679, 2639], 'This': [493, 738, 889, 2365, 2954, 3241, 4071, 4122], 'approach': [494, 1017, 1211, 1628, 2237, 2366, 2955], 'enables': [495, 2957], 'genome-scale': [496], 'CRISPR-Cas9.': [503], 'Synopsis': [504], 'The': [505, 646, 1183, 1354, 1973, 2002, 2474, 2510, 3298, 3341, 3374, 3627, 3665, 3716, 3768, 4041, 4156], 'applied': [509, 925], 'high-content': [511], 'screens.': [515], 'Computational': [516], 'classifiers': [517, 573, 2213, 2323, 2449, 2647, 2969, 2983, 3009, 3014, 3070, 3251, 3911], 'trained': [518, 2214, 2320, 2982, 3015, 3069, 3313, 3910], 'between': [519, 574, 2316, 2460, 2841, 3169, 3393, 3528, 3961, 3992], 'wild-type': [520, 1382, 2373, 2423, 2441, 2686, 2788, 2832, 2845, 2943], 'expressing': [524, 1327, 1524, 1561, 1598], 'Cas9': [525], 'gRNA': [527, 1095, 1609, 1637, 1650, 1737, 1974], 'enable': [528], 'single': [533, 545, 762, 1105, 1152, 1370, 1964, 2724, 3832, 4214], 'cell': [534, 566, 1074, 1268, 1883, 1913, 2160, 2442, 2454, 2482, 2622, 2654, 3222, 3520, 3640, 3672, 3807, 3892, 4105], 'mediated': [537], 'transient': [541, 1230, 1242, 1488], 'transfection': [542, 1118, 1231, 1243, 1307, 1489, 2412], 'plasmid': [547, 1773, 2416, 2748, 3400], 'combined': [549, 1102, 1262], 'profiling.': [553, 3642, 3839], 'Methods': [554], 'are': [555, 568, 905, 1179, 1256, 1409, 1689, 1779, 1968, 2018, 2126, 2252, 2450, 2550, 2580, 2601, 2671, 2707, 2734, 2759, 2897, 2941, 3330, 3338, 3530, 3589, 3747, 3955, 4009, 4100], 'construction': [559, 1943], 'screening': [563, 656, 704, 715, 731, 746, 871, 891, 1064, 1934, 1948, 2028, 2063, 2086, 3547, 3585], 'libraries.': [564], 'Single': [565, 1966, 2599], 'training': [571, 2968, 3249], 'computational': [572, 3250], 'transfected': [575, 1252, 1283, 1397, 1431, 1500, 1537, 1574, 1728, 1795, 2217, 2239, 2489, 2542, 2593, 2744, 3254, 3401, 3405, 3444, 3610, 3622, 3898], 'non-transfected': [577, 2248, 3256], 'from': [579, 764, 777, 1697, 2227, 2336, 2770, 3157, 3255, 3473, 3606, 3691, 3710, 3774, 3790, 3915, 3999], 'same': [581, 2338, 3966, 4007], 'well.': [582, 888, 2446, 3684], 'Profiling': [583], 'marker': [586, 1843, 2781], 'identifies': [592], 'several': [593, 1827], 'classes': [594], 'phenotypic': [596, 969, 2721, 3181, 3263, 3385], 'perturbations.': [597, 1045], 'Introduction': [598], 'Forward': [599], 'reverse': [601, 1117], 'mammalian': [605], 'model': [608, 859], 'organisms': [609], 'have': [610, 924, 2729, 2749, 3352, 3726], 'provided': [611], 'wealth': [613], 'information': [615, 803, 946], 'about': [616], 'function': [618], '(Boutros': [619, 907], 'Ahringer,': [621], '2008;': [622], 'Liberali': [623, 911, 2289], 'et': [624, 693, 718, 722, 726, 782, 786, 790, 794, 862, 866, 908, 912, 916, 949, 957, 1011, 1645, 1679, 1856, 1915, 2286, 2290, 2294, 2325, 3184, 4034, 4038, 4149], 'al,': [625, 694, 719, 723, 727, 783, 787, 791, 795, 863, 867, 909, 913, 917, 950, 958, 1012, 1646, 1680, 1857, 1916, 2287, 2291, 2295, 2326, 3185, 4035, 4039, 4150], '2015).': [626, 695, 1013, 4040], 'Nonetheless,': [627], 'role': [629], 'significant': [632, 2278], 'proportion': [633, 1184], 'remains': [636], 'unknown': [637], 'additional': [639], 'functions': [641], 'remain': [642], 'elucidated.': [645], 'discovery': [647], 'CRISPR-Cas': [650], 'has': [652, 684], 'revolutionized': [653], 'functional': [654, 689, 1001, 1608, 1736, 2106, 2112, 2123, 2131, 2136], 'because,': [657], 'unlike': [658], 'RNAi,': [659], 'targets': [661], 'at': [663, 1190], 'DNA': [665, 994, 1805, 2173], 'level': [666], 'therefore': [669], 'generate': [670, 2024], 'null': [672], 'alleles,': [673], 'resulting': [674], 'complete': [676], 'For': [680, 3685, 4053], 'reason,': [682], 'been': [685], 'used': [686, 1078, 2299, 2645, 3347, 3637, 3722], 'genomic': [690], '(Shalem': [692, 1010], 'Most': [696, 3187], 'performed': [698], 'date': [700], 'employ': [701], 'pooled': [703, 730, 745, 845], 'strategy,': [705], 'which': [706, 902, 983, 1687, 1695, 2242, 2251, 2632, 2717, 2733, 2814, 3387, 3754, 3952, 4133, 4185], 'cause': [711], 'differential': [712], 'growth': [713], 'conditions': [716], '(Koike-Yusa': [717], '2013;': [720], 'Shalem': [721], '2014;': [724], 'Wang': [725], '2014).': [728, 1647], 'However,': [729, 3207], 'precludes': [732], 'partially': [741], 'overcome': [742], 'combining': [744], 'RNA-seq,': [749], 'but': [750, 933, 2825], 'does': [752], 'not': [753, 801, 960, 976, 2619, 4029], 'easily': [754], 'scale': [755, 938], 'thousands': [760, 765, 1042], 'perturbations,': [767], 'limited': [770, 936, 1004], 'features': [772, 1891, 2310, 2346, 2438, 2987, 3064, 3081, 3127, 3146, 3636, 3674, 3804, 3809, 3887, 3894, 3906, 3954, 4064, 4099, 4196], 'read': [776], 'RNA': [778], 'transcript': [779], 'profiles': [780, 3151, 3178, 3436, 3619, 3630, 3662, 3669, 3696, 3708, 3773, 3781, 3929, 3950, 3997, 4162, 4174], '(Adamson': [781], '2016;': [784, 788, 792, 951, 955, 1858], 'Dixit': [785], 'Jaitin': [789], 'Datlinger': [793], '2017).': [796, 868, 3186], 'Moreover,': [797], 'sequencing-based': [798], 'approaches': [799], 'do': [800], 'provide': [802], 'on': [804, 812, 1103, 2163, 2215, 2341, 2359, 2466, 2971, 2985, 3016, 3634, 3757, 3836, 3942], 'size': [806, 1894, 2098, 3375, 3538, 3576], 'or': [807, 811, 1003, 1288, 1325, 1402, 1436, 1505, 1542, 1579, 1661, 2076, 2134, 2495, 3324, 3440, 3449, 3469, 3480, 3671], 'morphology,': [808], 'microenvironment,': [810], 'organization': [815], 'intracellular': [817], 'structures': [818], 'such': [819, 836], 'as': [820, 1465, 2303, 3426, 3888, 3890, 3987, 3989, 4074], 'Image-based': [826, 3281, 3415], 'phenotyping': [827, 850, 896], 'ideally': [832], 'suited': [833], 'study': [835], 'Recently,': [838, 919], 'perturb': [841, 883, 1461], 'format,': [846, 932], 'followed': [847], 'situ': [853, 1674], 'genotyping': [854], 'were': [855, 935, 3631, 3676], 'prokaryotic': [858], 'systems': [860], '(Emanuel': [861], '2017;': [864], 'Lawson': [865], 'An': [869], 'alternative': [870], 'strategy': [872, 892, 1053], 'involves': [873], 'seeding': [874], 'multi-well': [877, 1971], 'plates': [878], 'contain': [880], 'reagents': [881], 'one': [884, 2888], 'specific': [885, 903, 2964, 2972], 'per': [887, 1154, 1302, 1448, 1881, 3048, 3683], 'detailed,': [894], 'populations': [898, 3019, 3607], '2015;': [910, 914, 1917, 2292], 'Caicedo': [915, 2293], '2016).': [918, 2296], 'number': [921, 2033, 2038, 2044, 2050, 2056, 2069, 2101, 2118], 'studies': [923], 'an': [930, 1016, 1051, 1062, 1945, 2025, 3072, 3854, 4126], 'these': [934, 1177, 2939, 3899, 3985, 3993], 'only': [940, 1871], 'obtained': [941, 1257, 3709, 3789, 3998], 'well-averaged': [942, 2386, 3270], 'readouts': [943, 4018], 'low': [945, 2751], 'content': [947], '(Hultquist': [948], 'Tan': [952], 'Martin,': [954], 'Strezoska': [956], '2017),': [959], 'realizing': [961], 'full': [963, 2369, 4014], 'potential': [964], 'could': [971, 4134], 'bring.': [972], 'Importantly,': [973, 2836], '100%': [977], 'effective': [978, 1213], 'all': [980, 1000, 1321, 2308, 3635, 3695], 'targeted': [981, 1124, 1137, 1197, 1707, 1882, 2072, 2095, 2653, 2715, 2773, 3482], 'cells,': [982, 1416, 1729, 2241, 2250, 2936, 3257, 3714, 3833, 3903, 3946], 'result': [987], 'in-frame': [989], 'repair': [990], 'CRISPR-Cas9-induced': [993, 1698], 'lesions,': [995], 'failure': [997], 'target': [999, 1653, 3028, 3092], 'alleles': [1002], 'efficacy': [1005, 1334, 1455, 1643], 'address': [1019, 2260, 2641], 'problem,': [1021], 'allowing': [1022], 'us': [1023], 'first': [1026, 1660, 2981], 'time': [1027], 'combine': [1029], 'power': [1031, 3823], 'high-content,': [1035], 'across': [1041, 1629, 4012, 4191], 'Results': [1046], 'Discussion': [1048], 'devised': [1050], 'experimental': [1052, 1155, 2236], 'format.': [1065], 'To': [1066, 1624, 2640, 2710, 2977, 3815, 3918], 'allow': [1067], 'maximum': [1068], 'flexibility': [1069], 'regard': [1071], 'line': [1075, 2512, 3343, 3718], 'assay': [1077], 'screening,': [1080], 'opted': [1082], 'one-component': [1085], 'where': [1087, 1997], 'coding': [1089, 3422], 'sequence': [1090, 1975, 2003], 'SpCas9,': [1092], 'chimeric': [1094], 'fluorescent': [1098, 1906], 'protein': [1099, 1808, 2998, 3226, 3295, 3524, 3861, 3869], '(tdTomato)': [1100], 'plasmid.': [1106, 1247, 1292, 1312, 1406, 1440, 1509, 1546, 1583, 2499, 2547, 2598], 'introduced': [1108], 'into': [1111, 1958, 1970, 2377, 2505, 2820, 2860, 3901], 'assayed': [1120, 1466], 'expression': [1121, 1148, 1166, 1249, 1279, 1722, 2607, 2777], 'quantitative': [1127, 1259], 'immunofluorescence': [1128, 1260, 1814], '(Fig': [1129, 1171, 1666, 1701, 1739, 1790, 2347, 2626, 2739, 2765, 2850, 2880, 3065, 3205, 3977, 4241], '1A).': [1130], 'As': [1131, 1760], 'proof': [1133, 1762], 'concept,': [1135], 'transferrin': [1139], 'receptor': [1140], '(TFRC)': [1141], 'HeLa': [1143, 1204, 1281, 1395, 1498, 1535, 1796, 2156, 2540, 3902], 'assessed': [1146], 'TFRC': [1147, 1165, 1277, 1290, 1300, 1310, 1317, 1392, 1404, 1426, 1438, 1444, 1593], 'approximately': [1150, 1878], '4,000': [1151, 1879], 'condition.': [1156], 'A': [1157, 1930, 1950, 2500], 'subpopulation': [1158], '(which': [1161], 'expresses': [1162], 'tdTomato)': [1163], 'loses': [1164], 'starting': [1167], '2': [1168, 2075], 'days': [1169, 1305, 1528, 1565, 1602], 'post-transfection': [1170], '1B': [1172], 'C),': [1174], 'indicating': [1175, 1730, 2903, 2937, 3968], 'functionally': [1180, 1384], 'genetically': [1181, 1186, 1350, 1385, 2245, 2253, 2425], 'perturbed.': [1182], 'increased': [1189], 'longer': [1191], 'times': [1192], 'after': [1193, 1306], 'transfection.': [1194], 'also': [1196, 2827, 4043], 'LAMP1': [1200, 1340, 1496, 1507, 1519, 1570, 1581], 'YAP1': [1202, 1342, 1533, 1544, 1556], 'additionally': [1207], 'U2OS': [1215, 1572], '(Figs': [1217, 3031, 3098, 4019, 4065, 4116, 4197], '1D': [1218], 'EV1A,': [1220], 'B': [1221], 'C).': [1223, 3980], 'Figure': [1224, 1479, 1928, 2149, 2388, 3273, 3498, 3653], '1.': [1225], 'Schematic': [1235, 1375, 1936, 2401], 'overview': [1236], 'tdTomato': [1248, 1274, 1326, 1523, 1560, 1597, 2606, 2780], '(magenta)': [1250, 1275], 'marks': [1251], 'cells.': [1253, 1330, 1353, 1387, 1420, 2427, 2577, 2725, 2911, 3319, 3765, 3917, 4226], 'Single-cell': [1254], 'measurements': [1255], '(green)': [1261, 1278], 'segmentation,': [1269], 'see': [1270, 1611], 'details.': [1273, 1619], 'control': [1286, 1400, 1434, 1503, 1540, 1577, 2492, 2531, 2747, 3018, 3336, 3399, 3438, 3467, 3625], 'plasmid,': [1287, 1401, 1435, 1504, 1541, 1578, 2494], 'Scale': [1293, 1421, 1510, 1547, 1584, 2177, 2204, 2583, 3484], 'bar,': [1294, 1422, 1511, 1548, 1585, 2178, 2205, 2584, 3485], '50': [1295, 1423, 1512, 1549, 1586, 2179, 2206, 2585], 'μm.': [1296, 1424, 1513, 1550, 1587, 2180, 2207, 2586, 3487], 'Quantification': [1297, 1331, 1425], 'normalized': [1299, 1316, 1517, 1554, 1591], 'staining': [1301, 1318, 1390, 1494, 1520, 1531, 1557, 1568, 1594, 2190, 2202, 2538, 2790, 2834, 2849, 3085, 3227, 3239, 3296, 3419, 3464, 3525, 3573, 3886, 4062, 4112, 4194, 4235], 'cell,': [1303, 2182], '1–4': [1304], 'Violin': [1313, 1441, 1514, 1551, 1588], 'plots': [1314, 1442, 1515, 1552, 1589], 'intensity': [1319, 1521, 1558, 1595, 1901, 2992, 4063], 'analysed': [1322, 2009, 3139], '(grey)': [1324], '(T(+),': [1328], 'magenta)': [1329], 'TFRC,': [1339], 'plasmids;': [1344], 'bars': [1345], 'indicate': [1346, 2109, 3108], 'percentage': [1348], 'T(+)': [1352, 1419, 1449, 2240, 2332, 2461, 2554, 2567, 2621, 2693, 2704, 2737, 2755, 2808, 2829, 2895, 2927, 3153, 3316, 3679, 3711, 3762, 3772, 3794, 3914], 'mean': [1355, 1518, 1555, 1592, 3040, 3149, 3434, 3599, 3617, 3628, 3666, 3779, 3927], '±': [1356], 'standard': [1357, 2527], 'deviation': [1358, 2528], 'three': [1360, 1711], 'independent': [1361, 2229, 3842, 3962], 'experiments': [1362], 'displayed.': [1364], 'Evaluation': [1365], 'bDNA': [1373, 1388], 'FISH.': [1374], 'representation': [1376, 1452, 1937, 2081, 2402, 3460, 3507, 3559], 'expected': [1379, 4073], 'phenotype': [1380, 2561, 3974, 4123], 'FISH': [1389], 'mRNA': [1393, 1427, 1445], 'Cell': [1407, 2548], 'outlines': [1408, 2549], 'indicated': [1410], 'colour-coded': [1412, 2768, 3331, 3339, 3703], 'white': [1413], 'T(−)': [1415, 2249, 2301, 2334, 2463, 2576, 2757, 2784, 2884, 2935, 3155, 3318, 3681, 3713, 3764, 3796, 3916], 'magenta': [1417, 3704], 'spots': [1428], 'spot': [1446], 'counts': [1447], 'cell.': [1450, 2665], 'Heatmap': [1451], 'designed': [1459], '26': [1462, 1708], 'selected': [1463, 1649, 2090, 3450, 3509, 3846], 'smFISH.': [1468], 'Download': [1469, 1471, 1620, 1622, 2139, 2141, 2231, 2233, 2612, 2614, 3488, 3490, 3643, 3645, 3811, 3813], 'figure': [1470, 1621, 2140, 2232, 2613, 3489, 3644, 3812], 'PowerPoint': [1472, 1623, 2142, 2234, 2615, 3491, 3646, 3814], 'Click': [1473, 1922, 2143, 3492, 3647], 'here': [1474, 1923, 2144, 3493, 3648], 'expand': [1476, 1925, 2146, 3495, 3650], 'figure.': [1478, 1927, 2148, 3497, 3652], 'EV1.': [1480], 'Functional': [1481], 'Immunofluorescence': [1493, 1530, 1567, 2534, 3456], '(T(+))': [1525, 1562, 1599], '4': [1527, 1564, 1601], 'post-transfection.': [1529, 1566, 1603], 'Rational': [1604], 'selection': [1605, 1635], 'highly': [1607, 4010], 'sequences,': [1610], 'material': [1615], 'systematically': [1625], 'test': [1626], 'our': [1627, 2690, 3140, 3817, 3939], 'multiple': [1630, 4192], 'genes,': [1631, 2096], 'sequences': [1638, 1651, 1738], 'high': [1640, 2697, 2731, 2838, 2915, 3934], 'predicted': [1641, 2475, 2659], 'on-target': [1642], '(Doench': [1644], 'separate,': [1654], 'expressed': [1655], 'exons,': [1656], 'while': [1657, 2894, 2922], 'avoiding': [1658], 'last': [1662], 'exons': [1663], 'transcripts': [1665, 1688], 'EV1D).': [1667], 'employed': [1669], 'single-molecule': [1671], 'fluorescence': [1672], 'hybridization': [1675], '(smFISH)': [1676], 'technique': [1677], '(Battich': [1678], '2013)': [1681], 'detect': [1683, 3262, 4209], 'depleted': [1690], 'due': [1691], 'nonsense-mediated': [1693], 'decay,': [1694], 'results': [1696, 2417, 3107, 3818], 'frameshift': [1699], 'mutations': [1700], '1E,': [1702], 'F': [1703], 'G).': [1705], 'each.': [1714], '72%': [1715], 'than': [1725, 3013, 3265, 3357], '30%': [1726], 'reliably': [1734], 'select': [1735, 3008, 3349, 3724], '1H,': [1740], 'Table': [1741, 3035], 'EV1).': [1742, 1793], 'subsequently': [1744, 3164], 'cost-effective': [1747], 'produce': [1750], 'plasmids.': [1759, 2079, 3626], 'principle,': [1764], 'constructed': [1766, 3841], 'consisting': [1769], 'transfection-grade': [1772], 'preparations': [1774], '1,457': [1776], 'annotated': [1780], 'ontology': [1783], '(GO)': [1784], 'terms': [1785, 3511, 3534, 3543, 3562, 3581], 'various': [1787], 'post-translational': [1788], 'modifications': [1789], 'EV2,': [1791], 'Dataset': [1792], '384-well': [1802], 'plates,': [1803], 'stained': [1804, 1840, 2171], 'total': [1807, 2175, 2997, 3174, 3225, 3294, 3523, 3776], 'subjected': [1810], 'mAb414,': [1816], 'monoclonal': [1818], 'antibody': [1819], 'binds': [1821], 'phenylalanine–glycine': [1822], '(FG)': [1823], 'repeats': [1824], 'subunits': [1828, 3030, 3057, 3094, 3213, 3230, 3323, 3615], '(NPC;': [1834], 'Davis': [1835], 'Blobel,': [1837, 2804, 4083], '1986).': [1838, 2805, 4084], 'because': [1844, 2754], 'regulation': [1846, 3865], 'NPC': [1848, 1868, 2222, 3097, 3233, 3329, 3838, 3851, 4180], 'assembly': [1849], 'interphase': [1851], 'incompletely': [1853], 'understood': [1854], '(Otsuka': [1855], 'Weberruss': [1859], 'Antonin,': [1861, 4183], '2016)': [1862], 'investigated': [1873], 'microscopy.': [1875], 'imaged': [1877], 'population': [1884, 2421, 2443, 2455, 2655, 2774], 'extracted': [1886, 3904], 'set': [1889, 2343, 2468, 3061, 3075, 4015, 4097], 'describing': [1892], 'shape': [1896], 'texture': [1903, 2994, 4195], 'markers': [1907], 'specified': [1909], 'subregions': [1910], 'every': [1912, 1977, 2445, 2481, 2620, 2663], '(Stoeger': [1914], 'Fig': [1918], 'EV3A': [1919], 'B).': [1921], 'EV2.': [1929], 'workflow': [1940], 'library.': [1949, 2029, 2064, 2087], 'pool': [1951], 'oligos': [1953], 'synthesized': [1955], 'cloned': [1957], 'vector': [1960], 'backbone': [1961], 'reaction.': [1965, 2014], 'colonies': [1967, 2017, 2053], 'picked': [1969, 2035], 'plates.': [1972], 'colony': [1978, 1999], 'PCR': [1980, 2006], 'amplified': [1981], 'primers': [1983], 'introduce': [1985], 'barcodes': [1986], 'row,': [1990], 'column': [1991], 'plate': [1993], 'well': [1996, 2339, 2485, 3049, 3159, 3889, 3988], 'located.': [2001], 'products': [2007], 'deep': [2012], 'sequencing': [2013], 'Unique,': [2015], 'mutation-free': [2016, 2040, 2047], 'selected,': [2019], 're-arrayed': [2020, 2052], 'miniprepped': [2022], 'Representation': [2030], 'colonies,': [2036], 'identified': [2041, 3021, 3088, 3191, 3200, 3211, 3515, 3566, 3878], 'gRNAs,': [2042, 2048], 'unique': [2046], 'constructs': [2058, 3900], 'Pie': [2065], 'chart': [2066], 'representing': [2067], '1,': [2074], '3': [2077, 2525, 3736], 'Network': [2080, 3506, 3558], 'Nodes': [2088, 2125, 3699, 3746], 'represent': [2089, 3700], 'Gene': [2091], 'Ontology': [2092], 'annotations': [2093, 2132, 2137], 'node': [2097], 'represents': [2099, 3539, 3577], 'annotation,': [2107], 'edges': [2108], 'sharing': [2111], 'annotations.': [2113, 2124], 'Edge': [2114], 'thickness': [2115], 'scales': [2116], 'share': [2122, 3535], 'colour': [2127, 2602], 'coded': [2128, 2603], 'phosphorylation-related': [2130], '(red)': [2133], 'ubiquitylation-related': [2135], '(blue).': [2138], 'EV3.': [2150], 'Nucleus': [2158], 'segmentation': [2161, 2187], 'based': [2162, 2340, 2358, 2465, 2970, 2984, 3633, 3756], 'image': [2164, 2535], 'processing': [2165], 'protein.': [2176], 'Nucleus,': [2181], 'cytoplasm': [2183], 'periphery': [2186, 2801], 'mAb414': [2189, 2201, 2537, 2789, 2816, 2833, 2848, 3084, 3238, 3418, 3463, 3572, 3667, 3803, 3885, 4061, 4080, 4115, 4193, 4234], 'experiment': [2198], 'pattern.': [2203, 2835, 3420, 3574], 'Scatterplot': [2208], 'misclassification': [2210, 3307], 'errors': [2211], 'HSPA5,': [2221, 3476], 'non-targeting': [2225, 2530, 3017, 3358, 3372, 3398, 3624, 3731, 3744], 'controls': [2226, 2305, 3359, 3732], 'two': [2228, 2261], 'Our': [2235], 'generates': [2238], 'may': [2243, 4124], 'perturbed,': [2246], 'wild-type.': [2254, 2709], 'leveraged': [2256], 'aspect': [2258], 'challenges': [2262], 'analysis': [2265, 2635, 2691, 3195, 3204, 3210, 3940], 'experiments;': [2270], 'technical': [2271, 2314], 'well-to-well': [2272, 2429], 'variation': [2273], 'identification': [2276, 2395], 'effects': [2280], 'high-dimensional': [2282, 2865], 'datasets': [2284], '(Loo': [2285], '2007;': [2288], 'First,': [2297], 'in-well': [2304], 'standardize': [2307], 'correct': [2312], 'variability': [2315, 2430], 'wells.': [2317], 'Second,': [2318], 'logistic': [2321, 3310], 'regression': [2322, 2448, 3311], '(Friedman': [2324], '2010)': [2327], 'attempt': [2329, 2457], 'categorize': [2331], '2A,': [2348], 'Tables': [2349, 3102], 'EV2': [2350], 'EV3)': [2352], 'calculated': [2354, 2479, 3038, 3165, 3392, 3553, 3590, 3632, 3677, 3926], 'classification': [2356, 2504, 2683, 3299, 3355, 3369, 3752, 3786], 'accuracy': [2361], 'classifier.': [2364], 'takes': [2367], 'among': [2371], 'both': [2372, 3883], 'account': [2378], 'thus': [2380], 'addresses': [2381], 'major': [2383], 'limitation': [2384], 'approaches.': [2387], '2.': [2389], 'A.': [2400], 'Transient': [2411], 'mixed': [2420], 'Technical': [2428], 'accounted': [2433], 'standardizing': [2436], 'Logistic': [2447], 'fitted': [2451, 2650], 'distinguish': [2459, 3253], 'features.': [2471, 2976, 3240, 3297, 3526], 'B,': [2472, 3766], 'C.': [2473, 3664], 'value': [2476, 2660, 2681], '(PV)': [2477, 2661], 'was': [2487, 3697, 3782], 'transiently': [2488], 'NUP160': [2497, 2545, 2596, 2772, 3479, 4164], 'positive': [2501, 2669, 2698], 'PV': [2502, 2571, 2610, 2670, 2699, 2753, 2839, 2916, 3395], 'indicates': [2503, 2513, 2682, 3344, 3719], 'class.': [2509, 2687], 'dotted': [2511, 3342, 3717], 'threshold': [2515, 3006, 3346, 3424, 3721], 'further': [2517, 2853, 3820, 3919], '[PV': [2520], '>': [2521, 2563], '0.62': [2522], '(mean': [2523], '+': [2524, 3362, 3735], '×': [2526, 3364, 3737], 'cells)].': [2532], 'D.': [2533, 3767], 'coloured': [2551], 'orange': [2552], '(PV': [2562], '0.62),': [2564], 'red': [2565], '<': [2572], '0.62,': [2573], 'blue': [2574], 'Missegmented': [2578], 'outlined': [2581], 'grey.': [2582], 'E,': [2587], 'F.': [2588], 'tSNE': [2589], 'projection': [2590, 2863], 'according': [2604, 3382, 3749], '(E)': [2608], '(F).': [2611], 'observed': [2617], '1C,': [2627], 'D,': [2628, 4068, 4119, 4200], 'G': [2629], 'H),': [2631], 'complicates': [2633], 'issue,': [2643], 'calculate': [2657], 'individual': [2664], 'Cells': [2666, 2912], 'classified': [2672], 'class': [2677], 'negative': [2680], 'By': [2688], 'limiting': [2689], 'value,': [2700], 'discard': [2702], 'illustrate': [2711, 2978], 'point,': [2713], 'NUP160,': [2716], 'causes': [2718], 'strong': [2720], 'effect': [2722], 'many': [2727], 'PV,': [2732], 'almost': [2735], 'exclusively': [2736, 2917], '2C).': [2740], 'In': [2741, 3136], 'contrast,': [2742], 'absolute': [2752], 'indistinguishable': [2760], 'feature': [2763, 2866, 2892, 3041, 3119, 3150, 3177, 3271, 3435, 3600, 3618, 3629, 3661, 3668, 3701, 3780, 3928, 3949, 4051], 'space': [2764, 2867, 3120], '2B).': [2766], 'PV.': [2783, 3935], 'display': [2786, 4087], 'pattern,': [2791, 3086], 'majority': [2794], 'signal': [2796, 2817], 'localized': [2797], '(Davis': [2802, 4081], 'Within': [2806], 'population,': [2809], 'observe': [2811, 4228], 'mislocalized': [2819, 2847], 'few': [2822, 4230], 'bright': [2823, 4231], 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[3005], 'better': [3012], '49': [3022], '14': [3025], 'proteasome': [3029, 3056, 3216, 3322, 3614], '3A': [3032], 'EV4A,': [3034], 'EV4).': [3036], 'values': [3042, 3601, 4093], 'discovered': [3051], 'changes': [3058, 3264], 'broad': [3060, 4096], 'EV4C).': [3066], 'Next,': [3067], 'entirely': [3073], 'features,': [3078], 'namely': [3079], '118': [3080], 'nine': [3089], 'structural': [3093, 3325, 3451, 3847], '3B': [3099], 'EV4B,': [3101], 'EV3': [3103], 'EV5).': [3105, 3206], 'These': [3106], 'profile': [3112, 3171], 'dimensions': [3114], 'selecting': [3122], 'affect': [3132], 'distinct': [3133, 4169], 'biological': [3134], 'processes.': [3135], 'addition,': [3137], 'screen': [3141], 'well-averaging': [3143], 'obtain': [3148, 3948], 'each': [3158, 3170, 3686], 'experiment.': [3162], 'Mahalanobis': [3167, 3689, 3769], 'distance': [3168, 3690, 3729, 3742, 3770], 'distribution': [3175, 3693, 3777], 'quantify': [3180], 'dissimilarity': [3182], '(Caicedo': [3183], 'hits': [3190, 3199], 'between-well': [3194, 3267, 3658], 'overlap': [3196], 'within-well': [3203, 3209, 3246, 3759, 3791], 'supports': [3242], 'notion': [3244], 'profiling,': [3247], 'sensitive': [3260], 'comparison': [3268, 3659, 3760, 3792], 'profiles.': [3272, 4052], '3.': [3274], 'linear': [3303], 'transformation': [3304], 'error': [3308], 'models': [3312], 'classify': [3315], 'Perturbations': [3320], 'purple': [3332], 'green.': [3334], 'Non-targeting': [3335], 'brown.': [3340], 'higher': [3354], '(third': [3360, 3733], 'quartile': [3361, 3734], '1.5': [3363], 'interquartile': [3365, 3738], 'range': [3366, 3739], 'scores': [3370], 'controls).': [3373, 3745], 'nodes': [3379, 3529], 'scaled': [3381, 3748], 'score,': [3386], 'reflects': [3388], 'KS': [3390], 'statistic': [3391], 'distributions': [3396], 'respective': [3409], '(see': [3411], 'Materials': [3412], 'Methods).': [3414], 'Colour': [3421], 'calculation': [3425], '(A).': [3428], 'Hierarchical': [3429, 3595], 'clustering': [3430, 3596, 3983], 'standardized': [3433], 'HSPA5': [3448, 4139], 'NPC.': [3455], 'images': [3457, 4222], 'schematic': [3459], 'pattern': [3465, 4113], 'NUP62,': [3475], 'NUP133,': [3477], 'NUP107,': [3478], 'NUP98': [3481, 4166], 'populations.': [3483], '10': [3486], 'EV4.': [3499], 'GO': [3510, 3533, 3542, 3561, 3580], 'associated': [3512, 3563], 'Edges': [3527], 'formed': [3531], 'if': [3532], 'genes.': [3536], 'Node': [3537, 3575], 'enrichment': [3540, 3578], 'relative': [3544, 3582], 'library,': [3548, 3586], 'P-value': [3551], 'hypergeometric': [3556, 3593], 'test.': [3557, 3594], 'P-values': [3588], 'EV5.': [3654], 'Phenotypic': [3655], 'A,': [3663], '(A)': [3670], '(C)': [3675], 'profile,': [3687], 'calculated.': [3698], 'profiles,': [3702, 3986, 3994], 'grey': [3706], 'respectively.': [3715], 'large': [3728], 'plotted': [3783], 'against': [3784], '(as': [3788], 'cells)': [3797], '(B)': [3805], '(D).': [3810], 'validate': [3816], 'explore': [3821], 'focused': [3835], 'HSPA5/Bip,': [3853], 'ER': [3855, 3860], 'chaperone': [3856], 'luminal': [3859], 'folding': [3862], 'unfolded': [3868], 'response': [3870], '(UPR;': [3871], 'Pfaffenbach': [3872, 4152], 'Lee,': [3874, 4154], '2011)': [3875], 'EV6).': [3896], 'EV7)': [3908], 'separate': [3913], 'characterize': [3920], 'phenotypes,': [3924], 'Notably,': [3936], 'focussing': [3938], 'specifically': [3941], 'phenotype-relevant': [3953], 'pronounced': [3957], 'without': [3958, 4210], 'reducing': [3959], 'correlations': [3960], 'gRNAs': [3963, 4004], 'gene,': [3967], 'true': [3971], 'revealed': [3976, 3995, 4158], 'EV6A': [3978, 4022], 'Strikingly,': [3981], 'hierarchical': [3982], 'correlation': [3991], 'similar': [4011], '3C': [4020, 4066, 4117, 4198], 'D),': [4024], 'something': [4025], 'generally': [4028], 'realized': [4030], 'RNAi': [4032], '(Collinet': [4033], '2010;': [4036], 'Singh': [4037], 'clustergram': [4042, 4157], 'demonstrates': [4044], 'lead': [4048], 'instance,': [4054], 'NUP62-targeted': [4055], 'reduction': [4059], 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