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Estimation of division rate in actual time. The time between gastrulation and diagnosis ( t D ) consists of a premalignant time span, up to the emergence of the MRCA ( t 1 ), and the expansion of the tumor thereafter ( t 2 ). Assuming exponential tumor growth and approximately 10 9 tumor cells at diagnosis, this yields an estimate for the division rate per tumor for the subset used to estimate mu eff . To obtain an estimate for the population level, these estimates are subsequently averaged across the cohort.

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