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We were unable to generate a gro-1 clk-1 double mutant because of the proximity of these genes. Except for clk-3(qm38); clk-1(e2519) and clk-3(qm38); clk-2(qm37) double mutants, all Clk double mutants were maintained as lines with linkage group (LG) III mutations balanced over dpy-17(e164) because the double homozygous mutants did not produce viable lines. For analysis, double mutants were picked from the second homozygous generation.
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Because the age-1 gene has been best studied in strain TJ401 age-1(hx546) fer-15(b26) and the fer-75 mutation has no effect on life-span (9), we used this strain to generate an age-1 fer-15; gro-1 triple mutant strain.
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All mutations used in this study, except age-1(hx546), have scorable phenotypes distinct from their effects on life-span. These were used in the construction of multiple mutant strains. All genes used in the aging study are contained on LG III except age-1, fer-15, clk-3 (LG II), and daf-16 (LG I). We constructed strains containing unlinked mutations by generating heterozygotes with one of the mutations balanced over closely linked markers in trans, making the unbalanced mutation homozygous (as scored by its phenotype) and then making the second mutation homozygous by displacement of trans markers. Linked doubles between LG III mutations were constructed with the use of appropriate flanking markers to pick recombinants and then with the removal of those flanking markers by outcrossing. All double mutants constructed had developmental phenotypes fully consistent with their presumed genotypes. In addition, the presence of daf-16(m26) in the putative daf-16 double mutants was confirmed by a complementation test in which the ability to suppress daf-2(e1370) was scored. The age-1 fer-15; gro-1 strain develops slowly and is sterne at 25°C, confirming the presence of gro-1 and fer-15, respectively. Because this strain lives considerably longer than gro-1 and because fer-15 has no Known effect on life-span (9), we conclude that age-1 is also present in this strain. 18. We thank W. Lai for her expert help and T. Barnes and J. Ewbank for helpful discussion and for critical reading of the manuscript: Some nematode strains used in this work were provided by the Caenorhabditis Genetics Center, which is funded by the NIH National Center for Research Resources (NCRR). This work was supported by a Medical Research Council of Canada grant to S.H. and by fellowships to B.L. from the J. W. McConnell Foundation and from the Fonds pour la Formation de Chercheuss et l'Aide à la Recherche (FCAR), Québec.
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