{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T12:36:31Z","timestamp":1786624591094,"version":"build-2736575974"},"reference-count":28,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T00:00:00Z","timestamp":1786579200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100003787","name":"Natural Science Foundation of Hebei Province","doi-asserted-by":"publisher","award":["C2024301017"],"award-info":[{"award-number":["C2024301017"]}],"id":[{"id":"10.13039\/501100003787","id-type":"DOI","asserted-by":"publisher"}]},{"name":"2025 Hebei Province Yanzhao Golden Platform Talent Gathering Programme"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["J. Fungi","JoF"],"abstract":"<jats:p>The SUN family protein MoSun4 in Magnaporthe oryzae has been previously implicated in mitophagy and has potential as a target for reducing rice blast, but its role as a secreted protein remains poorly understood. In this study, signal peptide prediction and yeast secretion assays confirmed MoSun4 signal peptide function, and co-localization revealed the localization of the extra-invasive hyphal membrane (EIHM)-associated apoplastic compartment or matrix, establishing MoSun4 as a secreted protein. In addition, deletion of the MoSUN4 gene reduced the hyphal growth, conidiation and virulence of M. oryzae. We further showed that MoSun4 is involved in regulating the expression of virulence-related genes, including multiple genes involved in cell wall degradation (eglC, eglD), secondary metabolism (gliK), and melanin biosynthesis (SDH1, BUF1, Cmr1, ALB1). Collectively, our study reveals that MoSun4 is a secreted protein that contributes to pathogenicity and is associated with altered expression of virulence-related genes, providing new insights into the functions of SUN family proteins.<\/jats:p>","DOI":"10.3390\/jof12080605","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T11:33:46Z","timestamp":1786620826000},"page":"605","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["MoSun4 Contributes to Pathogenicity and Is Associated with Altered Expression of Virulence-Related Genes in Magnaporthe oryzae"],"prefix":"10.3390","volume":"12","author":[{"given":"Huimin","family":"Li","sequence":"first","affiliation":[{"name":"Plant Protection Institute, Hebei Academy of Agriculture and Forestry Sciences, National Collection of Plant-Associated Microbes (Hebei), IPM Innovation Center of Hebei Province, International Science and Technology Joint Research Center on IPM of Hebei Province, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8182-6688","authenticated-orcid":false,"given":"Zhenhe","family":"Su","sequence":"additional","affiliation":[{"name":"Plant Protection Institute, Hebei Academy of Agriculture and Forestry Sciences, National Collection of Plant-Associated Microbes (Hebei), IPM Innovation Center of Hebei Province, International Science and Technology Joint Research Center on IPM of Hebei Province, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaomeng","family":"Liu","sequence":"additional","affiliation":[{"name":"Plant Protection Institute, Hebei Academy of Agriculture and Forestry Sciences, National Collection of Plant-Associated Microbes (Hebei), IPM Innovation Center of Hebei Province, International Science and Technology Joint Research Center on IPM of Hebei Province, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1435-1907","authenticated-orcid":false,"given":"Lemeng","family":"Dong","sequence":"additional","affiliation":[{"name":"Plant Hormone Biology Group, Swammerdam Institute for Life Sciences, University of Amsterdam, Science Park 904, 1098 XH Amsterdam, The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinggang","family":"Guo","sequence":"additional","affiliation":[{"name":"Plant Protection Institute, Hebei Academy of Agriculture and Forestry Sciences, National Collection of Plant-Associated Microbes (Hebei), IPM Innovation Center of Hebei Province, International Science and Technology Joint Research Center on IPM of Hebei Province, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,13]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1007\/s10529-012-0871-1","article-title":"Genetic engineering of crop plants for fungal resistance: Role of antifungal genes","volume":"34","author":"Ignacimuthu","year":"2012","journal-title":"Biotechnol. Lett."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/j.mib.2016.10.001","article-title":"Investigating the cell biology of plant infection by the rice blast fungus Magnaporthe oryzae","volume":"34","author":"Yan","year":"2016","journal-title":"Curr. Opin. Microbiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1111\/j.1364-3703.2011.00766.x","article-title":"Host-induced gene silencing: A tool for understanding fungal host interaction and for developing novel disease control strategies","volume":"13","author":"Nunes","year":"2012","journal-title":"Mol. Plant Pathol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Qu, Z., Chen, D., Hu, H., Liu, H., Zheng, L., Huang, J., Li, Y., Zhu, L., and Chen, X. (2026). Selectively targeting UDP-glucose 4-epimerase MoUGE1 for controlling rice blast disease. J. Adv. Res.","DOI":"10.1016\/j.jare.2026.01.072"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"3130","DOI":"10.1105\/tpc.110.077040","article-title":"HIGS: Host-induced gene silencing in the obligate biotrophic fungal pathogen Blumeria graminis","volume":"22","author":"Nowara","year":"2010","journal-title":"Plant Cell"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"796","DOI":"10.1111\/mpp.12342","article-title":"The Magnaporthe grisea species complex and plant pathogenesis","volume":"17","author":"Zhang","year":"2016","journal-title":"Mol. Plant Pathol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"582","DOI":"10.1016\/j.tim.2017.12.007","article-title":"Rise of a cereal killer: The biology of Magnaporthe oryzae biotrophic growth","volume":"26","author":"Fernandez","year":"2018","journal-title":"Trends Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"618","DOI":"10.1038\/s41477-024-01642-x","article-title":"Targeting Magnaporthe oryzae effector MoErs1 and host papain-like protease OsRD21 interaction to combat rice blast","volume":"10","author":"Liu","year":"2024","journal-title":"Nat. Plants"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1186\/s42483-024-00226-z","article-title":"Early molecular events in the interaction between Magnaporthe oryzae and rice","volume":"6","author":"Zhang","year":"2024","journal-title":"Phytopathol. Res."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1002\/1097-0061(200007)16:10<905::AID-YEA584>3.0.CO;2-1","article-title":"The \u2018SUN\u2019 family: Yeast SUN4\/SCW3 is involved in cell septation","volume":"16","author":"Mouassite","year":"2000","journal-title":"Yeast"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/j.femsyr.2004.05.001","article-title":"Uth1p: A yeast mitochondrial protein at the crossroads of stress, degradation and cell death","volume":"5","author":"Camougrand","year":"2004","journal-title":"FEMS Yeast Res."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1111\/1574-6968.12049","article-title":"Beyond the wall: Candida albicans secret(e)s to survive","volume":"338","author":"Sorgo","year":"2013","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Kuznetsov, E., Ku Erov\u00e1, H., V\u00e1chov\u00e1, L.E., and Palkov\u00e1, Z. (2013). SUN family proteins Sun4p, Uth1p and Sim1p are secreted from Saccharomyces cerevisiae and produced dependently on oxygen level. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0073882"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"P\u00e9rez-Hern\u00e1ndez, A., Gonz\u00e1lez, M., Gonz\u00e1lez, C., van Kan, J.A.L., and Brito, N. (2017). BcSUN1, a B. cinerea SUN-family protein, is involved in virulence. Front. Microbiol., 8.","DOI":"10.3389\/fmicb.2017.00035"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Yu, M., Yu, J., Cao, H., Song, T., Pan, X., Qi, Z., Du, Y., Zhang, R., Huang, S., and Liu, W. (2021). SUN-family protein UvSUN1 regulates the development and virulence of Ustilaginoidea virens. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.739453"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"13387","DOI":"10.1074\/jbc.M112.440172","article-title":"SUN proteins belong to a novel family of \u03b2-(1,3)-glucan-modifying enzymes involved in fungal morphogenesis","volume":"288","author":"Gastebois","year":"2013","journal-title":"J. Biol. Chem."},{"key":"ref_17","first-page":"e13439","article-title":"MoAti1 mediates mitophagy by facilitating recruitment of MoAtg8 to promote invasive growth in Magnaporthe oryzae Mol","volume":"25","author":"Shi","year":"2024","journal-title":"Plant Pathol."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Li, H., Mo, P., Zhang, J., Xie, Z., Liu, X., Chen, H., Yang, L., Liu, M., Zhang, H., and Wang, P. (2023). Methionine biosynthesis enzyme MoMet2 is required for rice blast fungus pathogenicity by promoting virulence gene expression via reducing 5mC modification. PLoS Genet., 19.","DOI":"10.1371\/journal.pgen.1010927"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1111\/j.1574-6968.2002.tb11046.x","article-title":"Dual cell wall\/mitochondria localization of the \u2019SUN\u2019 family proteins","volume":"207","author":"Velours","year":"2002","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"2686","DOI":"10.1016\/j.jia.2023.06.028","article-title":"Disruption of non-classically secreted protein (MoMtp) compromised conidiation, stress homeostasis, and pathogenesis of Magnaporthe oryzae","volume":"23","author":"Batool","year":"2024","journal-title":"J. Integr. Agric."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"4881","DOI":"10.1111\/1462-2920.15335","article-title":"Cleavage of PrePL by Lon promotes growth and pathogenesis in Magnaporthe oryzae","volume":"23","author":"Dang","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2056","DOI":"10.1128\/EC.00285-07","article-title":"Candida albicans Sun41p, a putative glycosidase, is involved in morphogenesis, cell wall biogenesis, and biofilm formation","volume":"6","author":"Hiller","year":"2007","journal-title":"Eukaryot. Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1111\/j.1567-1364.2009.00601.x","article-title":"The Saccharomyces SUN gene, UTH1, is involved in cell wall biogenesis","volume":"10","author":"Ritch","year":"2010","journal-title":"FEMS Yeast Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1525","DOI":"10.1128\/EC.3.6.1525-1532.2004","article-title":"Cellular localization and role of kinase activity of PMK1 in Magnaporthe grisea","volume":"3","author":"Bruno","year":"2004","journal-title":"Eukaryot. Cell"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1038\/nmeth.2089","article-title":"NIH Image to ImageJ: 25 years of image analysis","volume":"9","author":"Schneider","year":"2012","journal-title":"Nat. Methods"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1038\/s41587-019-0036-z","article-title":"SignalP 5.0 improves signal peptide predictions using deep neural networks","volume":"37","author":"Tsirigos","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"e2839","DOI":"10.21769\/BioProtoc.2839","article-title":"Functional evaluation of the signal peptides of secreted proteins","volume":"8","author":"Yin","year":"2018","journal-title":"Bio-Protocol"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1101","DOI":"10.1038\/nprot.2008.73","article-title":"Analyzing real-time PCR data by the comparative C T method","volume":"3","author":"Schmittgen","year":"2008","journal-title":"Nat. Protoc."}],"container-title":["Journal of Fungi"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2309-608X\/12\/8\/605\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T12:10:26Z","timestamp":1786623026000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2309-608X\/12\/8\/605"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,13]]},"references-count":28,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["jof12080605"],"URL":"https:\/\/doi.org\/10.3390\/jof12080605","relation":{},"ISSN":["2309-608X"],"issn-type":[{"value":"2309-608X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,13]]}}}