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Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,10,27]]},"abstract":"<jats:p>\n                    Most nonmammalian tetrapods exhibit more than three phalanges in their posterior digits (III\u2013V), whereas mammals are restricted to no more than three. So far, the molecular mechanism underlying this evolutionary reduction and the functions of\n                    <jats:italic toggle=\"yes\">Lhx2<\/jats:italic>\n                    and\n                    <jats:italic toggle=\"yes\">Lhx9<\/jats:italic>\n                    in digit development remain unclear. Here we show divergent evolution of\n                    <jats:italic toggle=\"yes\">Lhx2<\/jats:italic>\n                    and\n                    <jats:italic toggle=\"yes\">Lhx9<\/jats:italic>\n                    expression patterns in embryonic limbs across tetrapods. We further demonstrate that although ectopic overexpression of each gene in chick embryos delayed limb skeletal development, only\n                    <jats:italic toggle=\"yes\">Lhx9<\/jats:italic>\n                    reduced the phalanx number in the posterior digits. Transcriptomic analysis showed that\n                    <jats:italic toggle=\"yes\">Lhx2<\/jats:italic>\n                    regulates pathways for chondrocyte and osteoblast differentiation, and\n                    <jats:italic toggle=\"yes\">Lhx9<\/jats:italic>\n                    preferentially influences skeletal segmentation via the bone morphogenetic protein (BMP) pathway. The unique function and differential expression of\n                    <jats:italic toggle=\"yes\">Lhx9<\/jats:italic>\n                    in the limb buds between mammals and nonmammalian tetrapods suggest it may have acquired a role in suppressing phalanx number, driving the evolutionary reduction of phalanx number in mammals.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2605261123","type":"journal-article","created":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T18:01:52Z","timestamp":1792519312000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":0,"title":["<i>Lhx9<\/i>\n                    specifically modulates phalanx number, whereas\n                    <i>Lhx2<\/i>\n                    constrains digit length in 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