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Chua for the KOLF2.1J line; Dr. Nicholas Maragakis for the CS0002, CS25, and CS0201 lines; and Dr. Bipasha Mukherjee-Clavin for the CS83iCTR, 02623, and 01582 lines. Their generous contributions of hiPSC resources were essential to the completion of this study. The authors are grateful to Dr. Peter Guida and the support team of the Biology Department at Brookhaven National Laboratory for their invaluable assistance. The authors also thank Dr. Mike Sivertz and the NSRL operations staff for their essential support during the particle radiation experiments. 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An untargeted AAVS1 allele produces a approximately 500 bp fragment. PCR products were purified using QIAquick Gel extraction kit (28 706, Qiagen) following the manufacturer\u2019s protocol. Generation of Cortical Brain Organoids: Cortical brain organoids were generated from hiPSCs as previously described. Briefly, hiPSCs were dissociated into single cells using TrypLE, and 5.0 \u00d7 10^5 cells were seeded per well in ultra-low attachment U-bottom 96-well plates (Corning) to form uniform aggregates in mTeSR Plus medium (STEMCELL Technologies) supplemented with 50 uM Y-27632 (BioGems). On day 2, two-thirds of the medium was replaced every other day with fresh mTeSR Plus medium lacking Y-27632. When aggregates reached a diameter of 350 - 450 um, each aggregate was transferred to 24-well plates (Corning) containing cortical induction medium [DMEM/F12, N2 (1:200), B27 (1:100), 0.2 mg/mL glucose, 0.15 ug/mL cAMP, 0.5% NEAA, 1% L-alanyl-Lglutamine, 10 ug/mL heparin, 180 nM LDN-193189, 500 nM A83-01, 10 ug/mL IWR-1] and maintained for 4\u20135 days. Medium was changed every 3 days. Upon neuroectodermal differentiation, as indicated by brightened aggregate surfaces and radial neuroepithelial morphology, aggregates were embedded in growth factor-reduced Geltrex (Thermo Fisher Scientific) droplets. Geltrex-embedded aggregates were transferred to low-attachment 6-cm petri dishes containing organoid differentiation medium [DMEM-F12 with N2 supplement (1:200), B27 supplement (1:100), glucose (0.2 mg/mL), cAMP (0.15 ug/mL), 0.5% NEAA, 1% L-alanylL-glutamine, insulin (2.5 ug/mL)]. 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