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Hartwell was the first to discover that the products of specific genes are responsible for controlling the cycle of growth and division in cells. In experiments in yeast, he identified more than 100 genes specifically involved in cell-cycle control. By studying how yeast cells respond to damage caused by radiation, Hartwell developed the concept of checkpoints where cells pause, if need be, in the cell cycle to repair damage. It's now understood that these checkpoints are defective in most cancer cells. Hartwell also identified the specific yeast gene that controls passage through the first checkpoint in the cell cycle
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