Present, and will not rely on the presence of E1, we infer that CG8878 usually acts at the ci regulatory region to impede the spread of heterochromatin into this region, most likely within a dose sensitive manner. Supporting Info Mutations within a Drosophila Putative Protein Kinase amine, G to facilitate amino acid alignment. Accession numbers provided in Acknowledgments We thank Scott Hanna and Tanya McCracken for their technical help. Author Contributions Conceived and made the experiments: AM JL. Performed the experiments: AM JL. Analyzed the data: AM JL. Contributed reagents/ materials/analysis tools: AM JL. Wrote the paper: AM JL. References 1. Eaton S, Kornberg TB Repression of ci-D in posterior compartments of Drosophila by engrailed. Genes Dev. 4: 10681077. two. Bushey D, Locke J Mutations in Su205 and Su37 Suppress PElement-Dependent Silencing in Drosophila melanogaster. Genetics, 168: 1395 1411 three. Sameny A, Locke J The P-element-induced silencing impact of KP transposons is dose dependent in Drosophila melanogaster. Genome 54: 75262 4. Sun FL, Cuaycong MH, Craig CA, Wallrath LL, Locke J, et al. The fourth chromosome of Drosophila melanogaster: interspersed euchromatic and heterochromatic domains. Proc Natl Acad Sci USA 97: 53405345. 5. Wallrath LL, Elgin SCR Position impact variegation in Drosophila is linked to an altered 1516647 chromatin structure. Genes & Development 9: 1263 1277 6. Chetverina D, Savitskaya E, Maksimenko O, Melnikova L, Zaytseva O, et al. Red flag on the white reporter: a versatile insulator abuts the white gene in Drosophila and is omnipresent in mini-white constructs. Nucleic Acids Research 36:929937 7. Dernburg AF, Broman KW, Fung JC, Marshall WF, Philips J, et al. Perturbation of Nuclear Architecture by Long-Distance Chromosome Interactions. Cell 85:745759 8. Csink AK, Henikoff S Genetic modification of heterochromatic association and nuclear organization in Drosophila. Nature 381: 529531. 9. Belyaeva ES, Koryakov DE, Pokholkova GV, Demakova OV, Zhimulev IF Cytological study of the brown dominant position impact. Chromosoma 106: 124132. 10. Lindsley DL, Zimm GG The Genome of Drosophila melanogaster. Academic Press, New York. 11. Ashburner M. Drosophila, A laboratory manual Cold Spring Harbor Press, Cold Spring Harbor, New York 12. Ephrussi B, Herold JL Studies of eye pigments of Drosophila. I. Methods of extraction and quantitative estimation of the pigment components. Genetics 29: 148175. 13. Hahn MW, Han MV, Han S-G Gene Family Evolution across 12 Drosophila Genomes. PLoS Genet 3: e197. 14. Qi H, Yao C, Cai W, Girton J, Johansen KM, et al Asator, a Tau-Tubulin Kinase Homolog in Drosophila Localizes to the Mitotic Spindle. Developmental Dynamics 238: 32483256. 15. Shin J, Chakraborty G, Bharatham N, Kang CB, Tochio N, et al. NMR Solution Structure of Human Vaccinia-related Kinase 1 Reveals the Cterminal Tail Essential for Its Structural Stability and Autocatalytic Activity. J Biol Chem. 286: 2213122138, 16. Brameier M, Andrea Krings A, MacCallum RM NucPred–Predicting nuclear localization of proteins. Bioinformatics 23: 11591160 17. Saito K, Sakaguchi Y, Suzuki T, Siomi H, Siomi MC Pimet, the Drosophila homolog of HEN1, mediates 29-O-methylation of Piwi- interacting RNAs at their 39 ends. Genes Dev. 21: 16031608 18. Horwich MD, Li C, Matranga C, Vagin V, Farley G, et al. The Drosophila RNA Methyltransferase, DmHen1, Modifies Germline piRNAs and SingleStranded siRNAs in RISC. Current Biology 17: 1265.Present, and doesn’t rely on the presence of E1, we infer that CG8878 ordinarily acts in the ci regulatory region to impede the spread of heterochromatin into this region, probably in a dose sensitive manner. Supporting Details Mutations within a Drosophila Putative Protein Kinase amine, G to facilitate amino acid alignment. Accession numbers provided in Acknowledgments We thank Scott Hanna and Tanya McCracken for their technical assistance. Author Contributions Conceived and created the experiments: AM JL. Performed the experiments: AM JL. Analyzed the data: AM JL. Contributed reagents/ materials/analysis tools: AM JL. Wrote the paper: AM JL. References 1. Eaton S, Kornberg TB Repression of ci-D in posterior compartments of Drosophila by engrailed. Genes Dev. 4: 10681077. two. Bushey D, Locke J Mutations in Su205 and Su37 Suppress PElement-Dependent Silencing in Drosophila melanogaster. Genetics, 168: 1395 1411 three. Sameny A, Locke J The P-element-induced silencing impact of KP transposons is dose dependent in Drosophila melanogaster. Genome 54: 75262 four. Sun FL, Cuaycong MH, Craig CA, Wallrath LL, Locke J, et al. The fourth chromosome of Drosophila melanogaster: interspersed euchromatic and heterochromatic domains. Proc Natl Acad Sci USA 97: 53405345. 5. Wallrath LL, Elgin SCR Position effect variegation in Drosophila is associated with an altered 1516647 chromatin structure. Genes & Development 9: 1263 1277 6. Chetverina D, Savitskaya E, Maksimenko O, Melnikova L, Zaytseva O, et al. Red flag on the white reporter: a versatile insulator abuts the white gene in Drosophila and is omnipresent in mini-white constructs. Nucleic Acids Research 36:929937 7. Dernburg AF, Broman KW, Fung JC, Marshall WF, Philips J, et al. Perturbation of Nuclear Architecture by Long-Distance Chromosome Interactions. Cell 85:745759 8. Csink AK, Henikoff S Genetic modification of heterochromatic association and nuclear organization in Drosophila. Nature 381: 529531. 9. Belyaeva ES, Koryakov DE, Pokholkova GV, Demakova OV, Zhimulev IF Cytological study of the brown dominant position impact. Chromosoma 106: 124132. 10. Lindsley DL, Zimm GG The Genome of Drosophila melanogaster. Academic Press, New York. 11. Ashburner M. Drosophila, A laboratory manual Cold Spring Harbor Press, Cold Spring Harbor, New York 12. Ephrussi B, Herold JL Studies of eye pigments of Drosophila. I. Methods of extraction and quantitative estimation of the pigment components. Genetics 29: 148175. 13. Hahn MW, Han MV, Han S-G Gene Family Evolution across 12 Drosophila Genomes. PLoS Genet 3: e197. 14. Qi H, Yao C, Cai W, Girton J, Johansen KM, et al Asator, a Tau-Tubulin Kinase Homolog in Drosophila Localizes to the Mitotic Spindle. Developmental Dynamics 238: 32483256. 15. Shin J, Chakraborty G, Bharatham N, Kang CB, Tochio N, et al. NMR Solution Structure of Human Vaccinia-related Kinase 1 Reveals the Cterminal Tail Essential for Its Structural Stability and Autocatalytic Activity. J Biol Chem. 286: 2213122138, 16. Brameier M, Andrea Krings A, MacCallum RM NucPred–Predicting nuclear localization of proteins. Bioinformatics 23: 11591160 17. Saito K, Sakaguchi Y, Suzuki T, Siomi H, Siomi MC Pimet, the Drosophila homolog of HEN1, mediates 29-O-methylation of Piwi- interacting RNAs at their 39 ends. Genes Dev. 21: 16031608 18. Horwich MD, Li C, Matranga C, Vagin V, Farley G, et al. The Drosophila RNA Methyltransferase, DmHen1, Modifies Germline piRNAs and SingleStranded siRNAs in RISC. Current Biology 17: 1265.

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