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How plant importin\u2010\u03b1 maintains selectivity for endogenous cargos while accommodating diverse viral NLSs remains unclear. Here, we report the 1.88\u2009\u00c5 crystal structure of\n                    <jats:italic>Nicotiana benthamiana<\/jats:italic>\n                    importin\u2010\u03b11 lacking the autoinhibitory IBB domain (NbImp\u03b11\u0394IBB), and its complexes with three plant viral NLSs: BBSV CP, CMV\u2010fny 2b, and BBSV p7a, respectively. NbImp\u03b11 adopts the canonical armadillo repeat\u2010based superhelical fold with conserved major and minor NLS\u2010binding sites. All viral NLS peptides bind in a 2:1 stoichiometry and share a common orientation along the central groove (C to N direction), but exhibit distinct interaction strategies. BBSV CP and CMV\u2010fny 2b contain bipartite NLSs with reversed site usage, whereas BBSV p7a employs a monopartite NLS with repeated basic motifs to occupy both sites. Structural and functional analyses demonstrate that NbImp\u03b11 decodes the three\u2010dimensional spatial topology of basic residues rather than primary NLS sequences. The major binding site mediates high\u2010specificity initial anchoring, while the minor binding site provides plasticity for diverse NLS recognition. This dual\u2010site, topology\u2010based model explains selective yet flexible nuclear import and viral adaptation to plant constraints.\n                  <\/jats:p>","DOI":"10.1111\/tpj.71055","type":"journal-article","created":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T16:36:02Z","timestamp":1785256562000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Structures of\n                    <i>Nicotiana benthamiana<\/i>\n                    importin\u2010\u03b11 complexes reveal mechanisms for plant virus\n                    <scp>NLS<\/scp>\n                    recognition"],"prefix":"10.1111","volume":"127","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0185-9293","authenticated-orcid":false,"given":"Yunlong","family":"Liu","sequence":"first","affiliation":[{"name":"School of Life Sciences Tiangong University  Tianjin 300387 China"},{"name":"College of Biological Sciences China Agricultural 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