Nuclear speckle fusion via long-range directional motion regulates speckle morphology after transcriptional inhibition

Although the formation of RNA-protein bodies has been studied intensively, their mobility and how their number and size are regulated are still poorly understood. Here, we show significantly increased mobility of nuclear speckles after transcriptional inhibition, including long-range directed motion of one speckle towards another speckle, terminated by speckle fusion, over distances up to 4 μm and with velocities between 0.2 μm/min and 1.5 μm/min. Frequently, three or even four speckles follow very similar paths, with new speckles appearing along the path followed by a preceding speckle. Speckle movements and fusion events contribute to fewer, but larger, speckles after transcriptional inhibition. These speckle movements are not actin dependent, but occur within chromatin-depleted channels enrichedwith small granules containingthe specklemarker proteinSON. Similar longrange speckle movements and fusion events were observed after heat shockor heavymetal stress,and during lateG2 and earlyprophase.Our observations suggest a mechanism for long-range, directional nuclear specklemovements, contributingto overall regulationof nuclear speckle number and size as well as overall nuclear organization.