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Though operationally defined as  cis- regulatory elements, enhancers can also communicate with promoters on a separate homolog in  trans , a mechanism that has been suggested to account for the ability of certain alleles of the same gene to complement one another in a process otherwise known as transvection. This homolog-pairing dependent process is facilitated in Drosophila by chromatin-associated pairing proteins, many of which remain unknown and their mechanism of action uncharacterized. Here we have tested the role of the  gypsy  chromatin insulator in facilitating pairing and communication between enhancers and promoters in  trans  using a transgenic  eGFP  reporter system engineered to allow for targeted deletions in the  vestigial  Boundary Enhancer ( vg BE) and the  hsp70  minimal promoter, along with one or two flanking  gypsy  elements. We found a modest 2.5-3x increase in  eGFP  reporter levels from homozygotes carrying an intact copy of the reporter on each homolog compared to unpaired hemizygotes, although this behavior was independent of  gypsy . However, detectable levels of GFP protein along the DV wing boundary in trans-heterozygotes lacking a single enhancer and promoter was only observed in the presence of two flanking  gypsy  elements. Our results demonstrate that  gypsy  can stimulate enhancer-promoter communication in  trans  throughout the genome in a context-dependent manner, likely through modulation of local chromatin dynamics once pairing has been established by other elements and highlights chromatin structure as the master regulator of this phenomenon.

The Drosophila gypsy Insulator Supports Transvection in the Presence of the vestigial Enhancer