Nonequilibrium polymer models for chromatin

The cell nucleus is a dynamic environment where ATP-driven processes - like transcription, replication, and epigenetic modifications - continually drive the genome far from thermodynamic equilibrium. Recent interdisciplinary efforts combining cell biology and physics have introduced coarse-grained polymer models that reveal how these active processes shape chromosome organization in space and time. We review how these models have shed light on selected key features of nuclear function: the maintenance of epigenetic memory, the coupling between transcriptional activity and chromatin motion, and the emergence of replication factories. These approaches provide mechanistic insight and predictive power that are beyond experiments alone. We conclude by outlining future directions toward viewing the genome as an active polymer maintained far from equilibrium.