Active processes shape and move the genome and nucleoplasm

Active processes are omnipresent in the cell nucleus. From genome-based activities such as transcription and replication, to metabolic activity in the nucleoplasm and its liquid condensates, the nucleus is far from thermodynamic equilibrium. The nucleus’ active nature leads to many emergent properties of its principal components — the genome and the nucleoplasm — ranging from genome’s compartmentalization, coherent chromatin motions, emergent rheology, to non-equilibrium liquid–liquid phase separations of the nucleoplasm. Its rich phenomenology makes the nucleus well-suited to bring new non-equilibrium physics to light, which in turn reveals its complex underlying physiology. Understanding the role of active processes in the spatial organization and dynamics of the genome and the nucleoplasm is critical for new insights into the function of the human genome.