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However, how VEGF-C balances lymphatic endothelial cells (LECs) proliferation versus sprouting to ensure functional vessel formation has remained unclear. Using high-fidelity conditional genetics and receptor-specific ligands, we uncover a requirement for the alternative receptor VEGFR2 in VEGF-C-VEGFR3\u2013driven lymphatic vessel sprouting. While activation of VEGFR2 alone fails to induce lymphangiogenesis, VEGFR2 loss abolishes LEC sprouting, but not proliferation, in response to VEGF-C. In contrast, deletion of the VEGFR3 downstream effector PI3K\u03b1 completely abrogates lymphangiogenesis. VEGFR2 is activated and found in proximity to VEGFR3 in LECs in vivo, with PI3K\u03b1 controlling their relative cell-surface availability and VEGF-C increasing VEGFR2 relative to VEGFR3, thereby priming LECs for sprouting. 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