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He made further advancements in defining the molecular pathway of programmed cell death, and has identified several key components, including: EGL-1, a protein which activates apoptosis by inhibiting CED-9; transcription factors ces-1 and ces-2, and ced-8, which controls the timing of cell death. He continued working on heterochronic mutants and other aspects of cell lineage, and established lines of research in signal transduction, morphogenesis, and neural development. Horvitz has collaborated with Victor Ambros

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