Strukturdifferenzierungen in Y-chromosom von Drosophila hydei: the unique morphology of the Y chromosomal lampbrush loops Threads results from 'coaxial shells' formed by different satellite-specific subregions within megabase-sized transcripts.

Abstract

The results of pulsed-field gel electrophoresis (PFGE) analysis and two-colour transcript fluorescence in situ hybridization (FISH) for the three Threads-specific DNA satellites YLII, YLI and rally are in support of long-range clustering of these sequence families within the subterminal region on the long arm of the Y chromosome of Drosophila hydei. On the basis of the linear arrangement of at least four extended clusters of satellite-specific sequences, the loop morphology of wild-type and several mutant Threads can be explained by assumption of a single Threads-specific transcription unit comprising about 5.1 Mb of repetitive DNA located between the Pseudonucleolus and the Nucleolus organizer. Transcription is unidirectional from the Pseudonucleolus towards the terminally located Nucleolus organizer. Transcripts most likely start in front of or within the 3.2 Mb region of YLII-related sequences, pass through subsequent blocks of 1.2 and 0.3 Mb of YLI- and rally-related sequences, respectively, and cease within the region of a smaller block of YLI-related repeats. The megabase-sized transcripts remain physically linked to the DNA axis and their extended satellite-specific regions form coaxial clouds or shells around the central DNA axis. In this way each cluster of earlier-transcribed sequences generates a cloud or shell on top of the later-transcribed ones. According to this model of 'satellite-specific coaxial shells' the tube-like morphology and other peculiarities of the Y chromosomal lampbrush loops Threads can be explained as a result of satellite-specific RNA superstructures and/or formation of extended ribonucleoprotein (RNP) complexes between clusters of satellite-specific transcripts and specific proteins. On the basis of this model the specific morphology of several Threads mutants can be interpreted as the result of large interstitial or terminal deletions that alter the total length of the Threads-specific transcription unit without exerting other major effects on principal features of the transcription process along the Threads.