The nucleus is a central coordinator of cell adaptation to environmental forces and preserving its architecture under mechanical challenges is crucial for cell functions. Nuclear mechanics are orchestrated by the Linker of Nucleoskeleton and Cytoskeleton (LINC) complexes. These assemblies of proteins exhibit low elastic moduli on their own, but collectively function as mechanotransducing hubs embedded in the nuclear envelope (NE). While considerable progress was made in determining the structure and composition of LINC complexes, the physical mechanisms by which they convey forces across the NE remain undefined. In recent work, we established that clustering of LINC components at the nanoscale is a central determinant of force transmission at the NE. To probe and define the physical mechanisms of force transmission by those LINC components, we have assembled a consortium of three US and French teams that have synergistic and complementary expertise in all aspects of the proposed works. Working together on this project we aim at: (i) identifying the molecular mechanisms governing the formation, maintenance and disassembly of LINC protein clusters as a function of forces applied to the NE, (ii) measuring forces exerted at these clusters with novel optical force sensors (OFS) and (iii) using these data to formulate novel physical models that define and predict how modulation of LINC complex clustering can induce local changes in membrane shape. The project will be fully integrated among the three teams, with experiments informing theory (and vice versa), collaborative sharing of reagents and coordinated work across specific aims. It will be implemented through a highly multidisciplinary approach that integrates super-resolution (SR) microscopy, single molecule (SM) tracking and FRET imaging, cellular nanomanipulation, engineering of split-fluorescent reporters and OFS, their calibration using DNA origami, and theoretical modeling. Overall, this work will offer a mechanistic understanding of nanoscale force transmission across the NE. It will provide novel perspectives into the physical principles that govern the functions of LINC mechanotransducing hubs and a broader comprehension of other force-transmitting complexes. Its outcomes will be of immediate interest to the field of mechanobiology and, more broadly, will contribute novel rationales for the design of new force-responsive nanomaterials.