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Despite their importance, the genetic changes that gave rise to the unique human form remain mostly unknown\n                    <jats:sup>2<\/jats:sup>\n                    . Here we systematically map the gene-regulatory changes that shaped human skeletal evolution. Using massively parallel reporter assays (MPRAs) in chondrocytes, we assayed 561,410 human-derived substitutions in promoters and enhancers, identifying 15,077 loci with human-specific regulatory activity. We then generated human\u2013ape hybrid cells and differentiated them into osteochondral progenitors. Integrating the hybrid cells with MPRA measurements produced genome-wide atlases of human-specific changes in\n                    <jats:italic>cis<\/jats:italic>\n                    -regulatory expression, and the sequence variants that drive them. These atlases reveal an extensive rewiring of the extracellular matrix (ECM), including a marked suppression of glycosaminoglycan (GAG) biosynthesis, leading to an approximately three-to-fourfold reduction in joint GAG content in humans compared with non-human apes. We find that this human-specific shift bears signatures of selection, and is likely to be a key contributor to the exceptional susceptibility of humans to degenerative skeletal diseases\n                    <jats:sup>3\u20135<\/jats:sup>\n                    . Together, our results reveal a coordinated evolutionary remodelling of the human skeletal ECM, and establish a comprehensive framework for dissecting the genetic basis of human skeletal biology.\n                  <\/jats:p>","DOI":"10.1038\/s41586-026-11053-x","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T15:05:38Z","timestamp":1790175938000},"page":"698-706","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["The gene-regulatory evolution of the human 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