Illustration from Hansel, Crowder et al., proposing the mechanism for cytoskeletal tension coupling between cellular microenvironment and the nuclear membrane for flat (A) and silicon nanoneedle (B) substrates. Cells on flat surfaces can readily form focal adhesions on strong cytoskeletal tension, activating Yes-associated protein (YAP). Nanoneedles disrupts focal adhesion formation, inhibiting cytoskeletal tension, reducing YAP activity. Simultaneously, direct perturbation of the nucleus results in the formation of lamin protein caps, and a physical decoupling of the two main types of lamin protein in the nucleus. Adapted under the terms of CC BY license.[34] Copyright 2019, The Authors.