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However, their evolutionary trajectory about how blood cells emerged and diversified throughout animal history remains unclear. Here, we present a combination of bioinformatics and functional data that demonstrate that the metazoan blood cell program most likely originated through the repurposing of an ancestral premetazoan toolkit governed by\n                    <jats:italic toggle=\"yes\">Fos<\/jats:italic>\n                    . This primordial program established the macrophage-like initial blood cells at the metazoan root. Then, the first lineage bifurcation at the origin of Bilateria drove the emergence of a specialized mast\/killer lineage, characterized by acquisition of granular proteases for antiparasitic defense. Subsequent deuterostome\/vertebrate innovations branched T\/NK and erythrocyte\/thrombocyte lineages from mast cells while B cells derived from macrophages. Our data also show that a prototypic thymus formed at the gill edges of a chordate ancestor. In line with the evolutionary history, the modern hematopoietic pathway shows a vestige of the phylogeny; differentiation potentials of phylogenetically old cell lineages expressing\n                    <jats:italic toggle=\"yes\">Fos<\/jats:italic>\n                    such as macrophages and mast cells are widely retained, and ancient HSCs with limited lineage potentials have been inherited as origo-lineage progenitors. Our framework provides the history of blood cells showing an adaptive innovation built upon ancient unicellular foundations.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2528110123","type":"journal-article","created":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T18:07:33Z","timestamp":1779991653000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":3,"title":["Animals have expanded the evolutionary legacy of unicellular ancestors in blood cells"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3157-4168","authenticated-orcid":false,"given":"Yosuke","family":"Nagahata","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/00pqz0q24","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratory of Immunology, Institute for Life and Medical Sciences, Kyoto University","place":["Kyoto, 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