The cytoskeleton, an interconnected network of filamentous polymers and regulatory proteins, plays a crucial role in maintaining cell shape, facilitating intracellular transport, and enabling cell movement. Recent studies have highlighted how both internal and external forces can influence the cytoskeleton, affecting local mechanical properties and cellular behavior. The cytoskeleton's ability to generate, transmit, and respond to mechanical signals over various timescales is a key area of research. Long-lived cytoskeletal structures may act as epigenetic determinants of cell shape, function, and fate, potentially integrating past interactions with the mechanical microenvironment to influence future cellular behavior. The review discusses the three main types of cytoskeletal polymers—actin filaments, microtubules, and intermediate filaments—and their roles in cell mechanics. It also explores how the cytoskeleton establishes long-range order, transmits mechanical forces, and senses the mechanical microenvironment. The authors emphasize the importance of understanding the collective behavior of cytoskeletal structures rather than individual filaments, and they highlight the potential for cytoskeletal structures to record and influence cellular behavior over time. Finally, they discuss the implications of these findings for understanding normal and diseased cell behavior, as well as the need for new models and experimental techniques to further explore the link between physical force and cell behavior.The cytoskeleton, an interconnected network of filamentous polymers and regulatory proteins, plays a crucial role in maintaining cell shape, facilitating intracellular transport, and enabling cell movement. Recent studies have highlighted how both internal and external forces can influence the cytoskeleton, affecting local mechanical properties and cellular behavior. The cytoskeleton's ability to generate, transmit, and respond to mechanical signals over various timescales is a key area of research. Long-lived cytoskeletal structures may act as epigenetic determinants of cell shape, function, and fate, potentially integrating past interactions with the mechanical microenvironment to influence future cellular behavior. The review discusses the three main types of cytoskeletal polymers—actin filaments, microtubules, and intermediate filaments—and their roles in cell mechanics. It also explores how the cytoskeleton establishes long-range order, transmits mechanical forces, and senses the mechanical microenvironment. The authors emphasize the importance of understanding the collective behavior of cytoskeletal structures rather than individual filaments, and they highlight the potential for cytoskeletal structures to record and influence cellular behavior over time. Finally, they discuss the implications of these findings for understanding normal and diseased cell behavior, as well as the need for new models and experimental techniques to further explore the link between physical force and cell behavior.