Zebrafish work was performed in accordance with AMC IACUC regulations. ENU mutagenesis was performed to create two independent clp1 mutations. Gene knockdown was performed with morpholino oligonucleotides (Gene-Tools). Lifespan analysis, in situ hybridization, TUNEL, RNA rescue, and immunostaining were performed with standard protocols. Additional detail can be found in the Extended Experimental Procedures.