{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T23:45:51Z","timestamp":1789429551219,"version":"build-2803163510"},"reference-count":109,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2021,6,17]],"date-time":"2021-06-17T00:00:00Z","timestamp":1623888000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001824","name":"Grantov\u00e1 Agentura \u010cesk\u00e9 Republiky","doi-asserted-by":"publisher","award":["21-03224S"],"award-info":[{"award-number":["21-03224S"]}],"id":[{"id":"10.13039\/501100001824","id-type":"DOI","asserted-by":"publisher"}]},{"name":"ERDF\/ESF, Centre for Research of Pathogenicity and Virulence of Parasites","award":["CZ.02.1.01\/0.0\/0.0\/16_019\/0000759"],"award-info":[{"award-number":["CZ.02.1.01\/0.0\/0.0\/16_019\/0000759"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Heme biosynthesis is essential for almost all living organisms. Despite its conserved function, the pathway\u2019s enzymes can be located in a remarkable diversity of cellular compartments in different organisms. This location does not always reflect their evolutionary origins, as might be expected from the history of their acquisition through endosymbiosis. Instead, the final subcellular localization of the enzyme reflects multiple factors, including evolutionary origin, demand for the product, availability of the substrate, and mechanism of pathway regulation. The biosynthesis of heme in the apicomonad Chromera velia follows a chimeric pathway combining heme elements from the ancient algal symbiont and the host. Computational analyses using different algorithms predict complex targeting patterns, placing enzymes in the mitochondrion, plastid, endoplasmic reticulum, or the cytoplasm. We employed heterologous reporter gene expression in the apicomplexan parasite Toxoplasma gondii and the diatom Phaeodactylum tricornutum to experimentally test these predictions. 5-aminolevulinate synthase was located in the mitochondria in both transfection systems. In T. gondii, the two 5-aminolevulinate dehydratases were located in the cytosol, uroporphyrinogen synthase in the mitochondrion, and the two ferrochelatases in the plastid. In P. tricornutum, all remaining enzymes, from ALA-dehydratase to ferrochelatase, were placed either in the endoplasmic reticulum or in the periplastidial space.<\/jats:p>","DOI":"10.3390\/ijms22126495","type":"journal-article","created":{"date-parts":[[2021,6,17]],"date-time":"2021-06-17T11:20:26Z","timestamp":1623928826000},"page":"6495","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":12,"title":["Using Diatom and Apicomplexan Models to Study the Heme Pathway of Chromera velia"],"prefix":"10.3390","volume":"22","author":[{"given":"Jitka","family":"Richtov\u00e1","sequence":"first","affiliation":[{"name":"Biology Centre CAS, Laboratory of Evolutionary Protistology, Institute of Parasitology, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"},{"name":"Faculty of Science, University of South Bohemia, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5909-2307","authenticated-orcid":false,"given":"Lilach","family":"Sheiner","sequence":"additional","affiliation":[{"name":"Welcome Centre for Integrative Parasitology, College of Medical, Veterinary and Life Sciences, Institute of Infection, Immunity and Inflammation, University of Glasgow, Glasgow G12 8QQ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5876-4391","authenticated-orcid":false,"given":"Ansgar","family":"Gruber","sequence":"additional","affiliation":[{"name":"Biology Centre CAS, Laboratory of Evolutionary Protistology, Institute of Parasitology, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1952-3894","authenticated-orcid":false,"given":"Shun-Min","family":"Yang","sequence":"additional","affiliation":[{"name":"Biology Centre CAS, Laboratory of Evolutionary Protistology, Institute of Parasitology, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"},{"name":"Faculty of Science, University of South Bohemia, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lud\u011bk","family":"Ko\u0159en\u00fd","sequence":"additional","affiliation":[{"name":"Department of Biochemistry, University of Cambridge, Cambridge CB2 1TN, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Boris","family":"Striepen","sequence":"additional","affiliation":[{"name":"Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, PA 19104, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4819-7319","authenticated-orcid":false,"given":"Miroslav","family":"Oborn\u00edk","sequence":"additional","affiliation":[{"name":"Biology Centre CAS, Laboratory of Evolutionary Protistology, Institute of Parasitology, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"},{"name":"Faculty of Science, University of South Bohemia, 370 05 \u010cesk\u00e9 Bud\u011bjovice, Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,6,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1146\/annurev.arplant.57.032905.105448","article-title":"Tetrapyrrole biosynthesis in higher plants","volume":"58","author":"Tanaka","year":"2007","journal-title":"Annu. 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