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Genes in multicellular organisms are expressed as part of a developmental program that is largely dependent on self-perpetuating higher-order chromatin states. The mechanism of establishing and maintaining these epigenetic events is well studied in  Drosophila . The first known example of an epigenetic effect was that of (PEV) in  Drosophila , which has been shown to be due to gene silencing via heterochromatin formation. We are investigating a process similar to Position Effect Variegation (PEV) using a mini- w  transgene, called  Pci , inserted in the upstream regulatory region of  ci . The mini- white   +  transgene in  Pci  is expressed throughout the adult eye; however, when other  P  or  KP  elements are present, a variegated eye phenotype results indicating random  w   +  silencing during development. This  P  element dependent silencing (PDS) can be modified by the haplo-suppressors/triplo-enhancers,  Su(var)205  and  Su(var)3–7 , indicating that these heterochromatic modifiers also act dose dependently in PDS. Here we use a spontaneous derivative mutation of  Pci  called  Pci E1   ( E1 ) that variegates like PDS in the absence of  P  elements, presumably due to an adjacent  gypsy  element insertion, to screen for second-site modifier mutations that enhance variable silencing of  white   +  in  E1 . We isolated 7 mutations in  CG8878 , an essential gene, that enhance the  E1  variegated phenotype.  CG8878 , a previously uncharacterized gene, potentially encodes a serine/threonine kinase whose closest  Drosophila  paralogue,  ballchen  ( nhk-1 ), phosphorylates histones. These mutant alleles enhance both PDS at  E1  and Position Effect Variegation (PEV) at  w m4  , indicating a previously unknown common silencing mechanism between the two.

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Mutations in CG8878, a Novel Putative Protein Kinase, Enhance P Element Dependent Silencing (PDS) and Position Effect Variegation (PEV) in Drosophila melanogaster