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The beet cyst nematode (\n                    <jats:italic>Heterodera schachtii<\/jats:italic>\n                    ) is a crucial pest in sugar beet (\n                    <jats:italic>Beta vulgaris<\/jats:italic>\n                    ssp.\n                    <jats:italic>vulgaris<\/jats:italic>\n                    ). While all species of the genus\n                    <jats:italic>Beta<\/jats:italic>\n                    are highly susceptible, the three species of the beet wild relative genus\n                    <jats:italic>Patellifolia<\/jats:italic>\n                    are entirely resistant. Recently, we cloned the\n                    <jats:italic>Hs4<\/jats:italic>\n                    gene from\n                    <jats:italic>P. procumbens,<\/jats:italic>\n                    which confers complete resistance. In this study, we aimed to determine whether putative\n                    <jats:italic>Hs4<\/jats:italic>\n                    orthologs exist in\n                    <jats:italic>Beta<\/jats:italic>\n                    and\n                    <jats:italic>Patellifolia<\/jats:italic>\n                    species. The\n                    <jats:italic>Hs4<\/jats:italic>\n                    gene consisted of 4999 bp, with six exons and five introns.\n                    <jats:italic>Patellifolia<\/jats:italic>\n                    species contain highly similar\n                    <jats:italic>Hs4<\/jats:italic>\n                    homologs. Single nucleotide polymorphisms and insertions\/deletions between accessions and species could be detected. We found an exonic integration of three bases, resulting in the addition of one amino acid. Interestingly, this variant was present in single accessions of all three\n                    <jats:italic>Patellifolia<\/jats:italic>\n                    species.\n                    <jats:italic>Beta vulgaris<\/jats:italic>\n                    ssp.\n                    <jats:italic>vulgaris<\/jats:italic>\n                    contains an\n                    <jats:italic>Hs4<\/jats:italic>\n                    homolog (\n                    <jats:italic>BvHs4<\/jats:italic>\n                    ) with 60 % protein identity to Hs4.\n                    <jats:italic>BvHs4<\/jats:italic>\n                    homologs were present in all\n                    <jats:italic>Beta<\/jats:italic>\n                    species analyzed. Further, we examined the expression patterns of\n                    <jats:italic>Hs4<\/jats:italic>\n                    and\n                    <jats:italic>BvHs4<\/jats:italic>\n                    homologs. While\n                    <jats:italic>Hs4<\/jats:italic>\n                    homologs from\n                    <jats:italic>Patellifolia<\/jats:italic>\n                    species are strongly expressed in roots,\n                    <jats:italic>BvHs4<\/jats:italic>\n                    homologs are expressed mainly in leaves. When the spatio-temporal expression of\n                    <jats:italic>Hs4<\/jats:italic>\n                    was examined, no response to nematode inoculation was observed. These results are highly relevant for searching for functional\n                    <jats:italic>Hs4<\/jats:italic>\n                    alleles and breeding nematode-resistant varieties.\n                  <\/jats:p>","DOI":"10.1038\/s41598-026-40666-5","type":"journal-article","created":{"date-parts":[[2026,2,28]],"date-time":"2026-02-28T04:23:51Z","timestamp":1772252631000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Subcellular localization and differential expression provide insights into the putative function of the nematode resistance gene Hs4"],"prefix":"10.1038","volume":"16","author":[{"given":"Annika","family":"Schildberg","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kevin","family":"Dorn","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Christian","family":"Jung","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2026,2,28]]},"reference":[{"key":"40666_CR1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0152456","volume":"11","author":"MM Romeiras","year":"2016","unstructured":"Romeiras, M. 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