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Impairment of mismatch repair, homologous recombination deficiencies associated with BRCA1/2 loss, polymerase proofreading defects (e.g., POLE mutations), and carcinogenic insults such as tobacco smoke, ultraviolet radiation, and prior chemotherapy collectively accelerate mutation accumulation and diversify tumor cell populations ( TP53 , APC , RB1 ) and oncogenes (e.g., KRAS , BRAF , PIK3CA , MYC amplification) that establish and sustain the malignant state ( EGFR T790M or ABL T315I, which may exist at low frequencies prior to treatment and expand under selective pressure ( 2.3.2 Chromosomal instability and copy number heterogeneity Beyond point mutations, chromosomal instability (CIN) constitutes a major engine of genetic heterogeneity through recurrent whole-chromosome gains and losses, segmental amplifications, deletions, and structural rearrangements

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