{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T19:53:56Z","timestamp":1792785236499,"version":"4.5.0"},"reference-count":65,"publisher":"National Academy of Sciences","issue":"16","license":[{"start":{"date-parts":[[2026,4,16]],"date-time":"2026-04-16T00:00:00Z","timestamp":1776297600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"MOST | National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32230066 32400399 U24A20432"],"award-info":[{"award-number":["32230066 32400399 U24A20432"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"MOST | National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2023YFC2606800 2025YFC2609100"],"award-info":[{"award-number":["2023YFC2606800 2025YFC2609100"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Initiative Scientific Research Program, Institute of Zoology, CAS","award":["2023IOZ010 2023IOZ020"],"award-info":[{"award-number":["2023IOZ010 2023IOZ020"]}]},{"name":"Open Bidding Program of the National Forestry and Grassland Administration","award":["202401"],"award-info":[{"award-number":["202401"]}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,4,21]]},"abstract":"<jats:p>\n                    Pathogens frequently employ vector-manipulation strategies to enhance their transmission efficiency. Exosomes are increasingly recognized as mediators of interspecific communication between pathogens and their vectors. However, the mechanisms by which plant pathogenic nematode exosomes mediate cross-kingdom manipulation of vector development remain largely unexplored. Here, we demonstrate that the plant pathogenic nematode (\n                    <jats:italic toggle=\"yes\">Bursaphelenchus xylophilus<\/jats:italic>\n                    ), transmitted by the vector beetle (\n                    <jats:italic toggle=\"yes\">Monochamus alternatus<\/jats:italic>\n                    ), secretes exosomes containing microRNAs (miRNAs) that remodel the tracheal development of its vector. Upon nematode entry into the trachea, the beetle\u2019s tracheal diameter was markedly enlarged. Notably, exosome-like vesicles released from dispersal nematodes were internalized by the tracheal epithelial cells. Moreover, exosome-derived Bx-miR-71-5p directly activates\n                    <jats:italic toggle=\"yes\">Notch<\/jats:italic>\n                    expression, a key regulator of cell proliferation and differentiation.\n                    <jats:italic toggle=\"yes\">Notch<\/jats:italic>\n                    suppresses the expression of matrix metalloproteinases 3 (\n                    <jats:italic toggle=\"yes\">Mmp3<\/jats:italic>\n                    ), a critical enzyme for extracellular matrix (ECM) degradation, thereby promoting continuous ECM accumulation. Nanomaterial-mediated delivery of Bx-miR-71-5p to the vector beetle trachea upregulates the\n                    <jats:italic toggle=\"yes\">Notch<\/jats:italic>\n                    gene, leading to significant increases in ECM thickness and tracheal diameter, which consequently enhances nematode load. Collectively, this study identifies nematode exosomes as the delivery vehicle for Bx-miR-71-5p and defines a Notch\u2013Mmp3\u2013ECM axis through which pathogen signals remodel tracheal architecture to enhance vector competence and nematode load. These findings highlight that exosomes-mediated miRNA delivery may present a conserved \u201ctoolkit\u201d that can tune vector traits, and ultimately facilitates pathogen transmission efficiency during invasion.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2600386123","type":"journal-article","created":{"date-parts":[[2026,4,16]],"date-time":"2026-04-16T15:59:39Z","timestamp":1776355179000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":2,"title":["Plant pathogenic nematode exosomes remodel vector tracheae to enhance pathogen transmission"],"prefix":"10.1073","volume":"123","author":[{"given":"Yue","family":"Chang","sequence":"first","affiliation":[{"name":"State Key Laboratory of Animal Biodiversity Conservation and Integrated Pest Management, Institute of Zoology, Chinese Academy of Sciences","place":["Beijing, China"]},{"name":"University of Chinese Academy of Sciences","place":["Beijing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiao","family":"Zhou","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Animal Biodiversity Conservation and Integrated Pest Management, Institute of Zoology, Chinese Academy of Sciences","place":["Beijing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fangyuan","family":"Ye","sequence":"additional","affiliation":[{"name":"College of Life Sciences\/Hebei Basic Science Center for Biotic Interactions, Institute of Life Science and Green Development, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongxia","family":"Zhang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Animal Biodiversity Conservation and Integrated Pest Management, Institute of Zoology, Chinese Academy of Sciences","place":["Beijing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianghua","family":"Sun","sequence":"additional","affiliation":[{"name":"College of Life Sciences\/Hebei Basic Science Center for Biotic Interactions, Institute of Life Science and Green Development, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3706-1267","authenticated-orcid":false,"given":"Lilin","family":"Zhao","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Animal Biodiversity Conservation and Integrated Pest Management, Institute of Zoology, Chinese Academy of Sciences","place":["Beijing, China"]},{"name":"University of Chinese Academy of Sciences","place":["Beijing, China"]},{"name":"Shanxi Fen River Plain Farmland Shelterbelt Ecosystem National Observation Research Station","place":["Jinzhong, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2026,4,16]]},"reference":[{"key":"e_1_3_5_1_2","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aaz1659"},{"key":"e_1_3_5_2_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41396-020-0720-5"},{"key":"e_1_3_5_3_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.jare.2021.03.010"},{"key":"e_1_3_5_4_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0063602"},{"key":"e_1_3_5_5_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1199410"},{"key":"e_1_3_5_6_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2222040120"},{"key":"e_1_3_5_7_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41396-021-01010-z"},{"key":"e_1_3_5_8_2","doi-asserted-by":"publisher","DOI":"10.1038\/nature.2014.15011"},{"key":"e_1_3_5_9_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ento.48.091801.112722"},{"key":"e_1_3_5_10_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-14953-2"},{"key":"e_1_3_5_11_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms6488"},{"key":"e_1_3_5_12_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms8321"},{"key":"e_1_3_5_13_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2207848119"},{"key":"e_1_3_5_14_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.aar4142"},{"key":"e_1_3_5_15_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-40887-6"},{"key":"e_1_3_5_16_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2025.115635"},{"key":"e_1_3_5_17_2","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201211138"},{"key":"e_1_3_5_18_2","doi-asserted-by":"publisher","DOI":"10.15252\/embr.201439363"},{"key":"e_1_3_5_19_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-04791-8"},{"key":"e_1_3_5_20_2","doi-asserted-by":"publisher","DOI":"10.15252\/embj.2020105057"},{"key":"e_1_3_5_21_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2016.01.043"},{"key":"e_1_3_5_22_2","doi-asserted-by":"publisher","DOI":"10.1002\/1873-3468.70055"},{"key":"e_1_3_5_23_2","doi-asserted-by":"publisher","DOI":"10.1111\/cmi.12672"},{"key":"e_1_3_5_24_2","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1102053"},{"key":"e_1_3_5_25_2","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1000541"},{"key":"e_1_3_5_26_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00001-21"},{"key":"e_1_3_5_27_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2022.111527"},{"key":"e_1_3_5_28_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms12341"},{"key":"e_1_3_5_29_2","doi-asserted-by":"publisher","DOI":"10.1111\/1749-4877.12512"},{"key":"e_1_3_5_30_2","doi-asserted-by":"publisher","DOI":"10.1002\/mbo3.415"},{"key":"e_1_3_5_31_2","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.812414"},{"key":"e_1_3_5_32_2","doi-asserted-by":"publisher","DOI":"10.3390\/ijms17060778"},{"key":"e_1_3_5_33_2","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2022.890949"},{"key":"e_1_3_5_34_2","doi-asserted-by":"publisher","DOI":"10.1603\/0022-0493(2004)097[0941:ANMFLB]2.0.CO;2"},{"key":"e_1_3_5_35_2","unstructured":"N. Ido \u201cSome observations on biology and morphology of the pine sawyer\u201d in Transactions of the 23rd Annual Meeting of the Kansai Branch of the Japanese Forestry Society Japanese Forestry Society Kansai Branch Ed. (Japanese Forestry Society Kansai Branch 1972) vol. 23 pp. 180\u2013182."},{"key":"e_1_3_5_36_2","first-page":"96","article-title":"Prediction of emergence of the pine sawyer adult through continuous observations of larval growth by the log-cutting method","volume":"25","author":"Kishi Y.","year":"1976","unstructured":"Y. Kishi, Prediction of emergence of the pine sawyer adult through continuous observations of larval growth by the log-cutting method. For. Pests 25, 96\u201398 (1976).","journal-title":"For. Pests"},{"key":"e_1_3_5_37_2","unstructured":"Y. Takizawa \u201cEcology of the pine sawyer in Tohoku District (VIII). Periods of pupal and adult stages in the pupal cell under natural temperature conditions in Morioka City\u201d in Transactions of the 31st Annual Meeting of the Tohoku Branch of the Japanese Forestry Society Japanese Forestry Society Tohoku Branch Ed. (Japanese Forestry Society Tohoku Branch 1979) vol. 31 pp. 156\u2013157 (1979)."},{"key":"e_1_3_5_38_2","unstructured":"N. Enda \u201cDuration of Monochamus alternatus HOPE from pupation to emergence\u201d in Transactions of the 32nd Annual Meeting of the Kanto Branch of the Japanese Forestry Society Japanese Forestry Society Kanto Branch Ed. (Japanese Forestry Society Kanto Branch Niigata 1980) vol. 32 pp. 91\u201392."},{"key":"e_1_3_5_39_2","doi-asserted-by":"publisher","DOI":"10.1163\/156854101753250782"},{"key":"e_1_3_5_40_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.en.29.010184.000555"},{"key":"e_1_3_5_41_2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.84621"},{"key":"e_1_3_5_42_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijpharm.2023.123090"},{"key":"e_1_3_5_43_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.jchromb.2023.123982"},{"key":"e_1_3_5_44_2","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M502017200"},{"key":"e_1_3_5_45_2","doi-asserted-by":"publisher","DOI":"10.1080\/20013078.2020.1766192"},{"key":"e_1_3_5_46_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2110097119"},{"key":"e_1_3_5_47_2","doi-asserted-by":"publisher","DOI":"10.1093\/femsre\/fuab044"},{"key":"e_1_3_5_48_2","doi-asserted-by":"publisher","DOI":"10.1186\/s13287-020-01656-0"},{"key":"e_1_3_5_49_2","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2020.00076"},{"key":"e_1_3_5_50_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0046661"},{"key":"e_1_3_5_51_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-05274-6"},{"key":"e_1_3_5_52_2","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.abe0357"},{"key":"e_1_3_5_53_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-30995-0"},{"key":"e_1_3_5_54_2","doi-asserted-by":"publisher","DOI":"10.1186\/s12951-021-01225-4"},{"key":"e_1_3_5_55_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-024-00767-3"},{"key":"e_1_3_5_56_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.97.19.10295"},{"key":"e_1_3_5_57_2","doi-asserted-by":"publisher","DOI":"10.1163\/156854100509402"},{"key":"e_1_3_5_58_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1521098113"},{"key":"e_1_3_5_59_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1149460"},{"key":"e_1_3_5_60_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2010.01.007"},{"key":"e_1_3_5_61_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0707594105"},{"key":"e_1_3_5_62_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1319683111"},{"key":"e_1_3_5_63_2","unstructured":"X. Tang . RNA-Seq of pine sawyer beetle. GSA (Genome Sequence Archive). https:\/\/ngdc.cncb.ac.cn\/gsa\/browse\/CRA006464. Accessed 30 March 2026."},{"key":"e_1_3_5_64_2","unstructured":"Y. Chang . Pine Wood Nematode Small RNAome. GSA (Genome Sequence Archive). https:\/\/ngdc.cncb.ac.cn\/gsa\/browse\/CRA040045. Deposited 17 March 2026."},{"key":"e_1_3_5_65_2","unstructured":"Y. Chang Monochamus alternatus Transcriptome. GSA (Genome Sequence Archive). https:\/\/ngdc.cncb.ac.cn\/gsa\/browse\/CRA040052. Deposited 16 May 2025."}],"updated-by":[{"DOI":"10.1073\/pnas.2622410123","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T00:00:00Z","timestamp":1784073600000}}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2600386123","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T16:16:04Z","timestamp":1784132164000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/10.1073\/pnas.2600386123"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,16]]},"references-count":65,"journal-issue":{"issue":"16","published-print":{"date-parts":[[2026,4,21]]}},"alternative-id":["10.1073\/pnas.2600386123"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2600386123","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,16]]},"assertion":[{"value":"2026-01-12","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-03-05","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-04-16","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2600386123"}}