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In this study, we show that TMV can still achieve long\u2010distance vascular transport in the absence of CP, but systemic infection becomes intermittent because the processes underlying \u201csystemic movement\u201d can fail at multiple, separable steps. By resolving movement into phloem loading, vascular translocation, and phloem exit, we show that CP deficiency does not completely block entry into or transport within the vasculature. Instead, CP loss imposes a strong, route\u2010specific defect in bicollateral phloem, disproportionately reducing use of the adaxial\/internal pathway associated with efficient upward spread, while downward movement through abaxial\/external phloem remains comparatively permissive. In systemic leaves, CP\u2010deficient TMV frequently produces vein\/phloem\u2010associated reporter patterns rather than broad lamina invasion, indicating that phloem exit\/unloading is an additional, independent bottleneck. Enhancing RNA silencing suppression markedly enhances systemic viral accumulation and symptoms. However, it does not proportionally increase lamina invasion. This finding further supports the idea that exit and establishment are the limiting steps. Finally, among several helper viruses tested, only cucumber mosaic virus consistently alleviates this exit barrier, enabling CP\u2010deficient TMV to escape the vasculature and establish lamina infection.<\/jats:p>","DOI":"10.1111\/tpj.71082","type":"journal-article","created":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T07:56:44Z","timestamp":1785830204000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Systemic infection of tobacco mosaic virus is limited by upward transport and phloem unloading without coat protein"],"prefix":"10.1111","volume":"127","author":[{"given":"Minhue","family":"Jung","sequence":"first","affiliation":[{"name":"Department of Agricultural Biotechnology Seoul National University  Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9066-6903","authenticated-orcid":false,"given":"Kook\u2010Hyung","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Agricultural Biotechnology Seoul National University  Seoul Republic of Korea"},{"name":"Research Institute of Agriculture and Life Sciences Seoul National University  Seoul Republic of Korea"},{"name":"Plant Genomics and Breeding Institute Seoul National University  Seoul 08826 Republic of Korea"},{"name":"Plant Health Center Seoul National University  Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,8,4]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.106.042473"},{"key":"e_1_2_9_3_1","unstructured":"Bozkurt T.(2012)Purification of effector complexes from plants: in planta co\u2010immunoprecipitation of effectors. 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