{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T04:04:02Z","timestamp":1789790642710,"version":"4.0.1"},"reference-count":85,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T00:00:00Z","timestamp":1789689600000},"content-version":"vor","delay-in-days":17,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Physiologia Plantarum"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Long\u2010distance molecular transport contributes to coordination between locally stressed tissues and distal organs, but assigning causal systemic\u2010signaling functions to mobile RNAs and extracellular vesicle (EV)\u2010associated molecules remains experimentally challenging. This review critically evaluates the evidence required to distinguish molecular association, source assignment, physical movement, recipient activity, persistence, and causal function in sequential\u2010stress systems. We assess source\u2010resolved RNA tracking, graft\u2010based mobility assays, sequence\u2010assignment and contamination errors, molecular integrity, route validation, EV source\u2010fluid quality, particle identity and heterogeneity, cargo topology, vesicular versus non\u2010vesicular extracellular RNA, recipient uptake, and functional delivery. Recent studies on mobile\u2010mRNA assignment, stress\u2010responsive graft\u2010transmissible transcripts, plant EV proteomes and markers, EV isolation and characterization, and extracellular small\u2010RNA carrier states are integrated to identify which inferences are experimentally supported and which remain provisional. Sequential\u2010stress designs are considered separately because persistence must be distinguished from residual injury, prolonged acute responses, generalized sensitization, and stress\u2010order effects. The objective of this review is to establish reproducible experimental and analytical criteria for assigning systemic\u2010signaling functions to RNA and EV\u2010associated carriers. We therefore propose an evidence hierarchy progressing from association and source assignment to movement, recipient activity, persistence, necessity, sufficiency, and rescue. This framework defines the minimum evidence needed to advance from molecular detection to causal assignment in studies of systemic plant acclimation.<\/jats:p>","DOI":"10.1111\/ppl.71123","type":"journal-article","created":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T03:22:38Z","timestamp":1789788158000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Causal Validation of Mobile\n                    <scp>RNA<\/scp>\n                    and Extracellular Vesicle Signals in Plants: Experimental Frameworks for Sequential Stress Acclimation"],"prefix":"10.1111","volume":"178","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6864-2366","authenticated-orcid":false,"given":"Wajid","family":"Zaman","sequence":"first","affiliation":[{"name":"Department of Life Sciences Yeungnam University  Gyeongsan Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5562-6703","authenticated-orcid":false,"given":"Adnan","family":"Amin","sequence":"additional","affiliation":[{"name":"Department of Life Sciences Yeungnam University  Gyeongsan Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,18]]},"reference":[{"key":"e_1_2_14_2_1","doi-asserted-by":"crossref","DOI":"10.1093\/hr\/uhaf166","article-title":"Mango Pangenome Reveals Dramatic Impacts of Reference Bias on Population Genomic Analyses","volume":"12","author":"Ahmad B.","year":"2025","journal-title":"Horticulture Research"},{"issue":"1","key":"e_1_2_14_3_1","doi-asserted-by":"crossref","DOI":"10.1111\/ppl.14209","article-title":"Impact of Individual, Combined and Sequential Stress on Photosynthesis Machinery in Rice (\n                     Oryza sativa\n                   L)","volume":"176","author":"Anwar K.","year":"2024","journal-title":"Physiologia Plantarum"},{"key":"e_1_2_14_4_1","doi-asserted-by":"crossref","DOI":"10.1093\/plphys\/kiag141","article-title":"Comparative Proteomics Reveals Protein Signatures Shared Between and Unique to Bona Fide Plant EVs and Other Plant\u2010Derived Vesicles","volume":"200","author":"Bifolco M.","year":"2026","journal-title":"Plant Physiology"},{"key":"e_1_2_14_5_1","doi-asserted-by":"crossref","DOI":"10.3390\/plants15030439","article-title":"Characterization of Extracellular Vesicles From Cell Suspension Cultures of \n                     Coffea arabica\n                   L","volume":"15","author":"Bonaventura A.","year":"2026","journal-title":"Plants"},{"key":"e_1_2_14_6_1","doi-asserted-by":"crossref","first-page":"2389","DOI":"10.1093\/jxb\/erac512","article-title":"Extracellular RNA: Mechanisms of Secretion and Potential Functions","volume":"74","author":"Borniego M.","year":"2023","journal-title":"Journal of Experimental Botany"},{"key":"e_1_2_14_7_1","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2409090121","article-title":"Diverse Plant RNAs Coat Arabidopsis Leaves and Are Distinct From Apoplastic RNAs","volume":"122","author":"Borniego M.","year":"2024","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"e_1_2_14_8_1","doi-asserted-by":"crossref","DOI":"10.1002\/jev2.70101","article-title":"Identification of MARVELous Protein Markers for \n                     Phytophthora infestans\n                   Extracellular Vesicles","volume":"14","author":"Breen S.","year":"2025","journal-title":"Journal of Extracellular Vesicles"},{"key":"e_1_2_14_9_1","doi-asserted-by":"crossref","DOI":"10.1186\/s12870-026-09064-x","article-title":"Fine Scale Gene\u00a0Expression Dynamics at the Foliar Level Reveal Grapevine Strategies to Cope With Recurrent Drought Stress","volume":"26","author":"Canton M.","year":"2026","journal-title":"BMC Plant Biology"},{"key":"e_1_2_14_10_1","doi-asserted-by":"crossref","DOI":"10.1111\/tpj.70498","article-title":"Proteomic Snapshot of Pattern Triggered Immunity in the Arabidopsis Leaf Apoplast","volume":"123","author":"Chen H.\u2010C.","year":"2025","journal-title":"Plant Journal"},{"key":"e_1_2_14_11_1","doi-asserted-by":"crossref","DOI":"10.1002\/pld3.70087","article-title":"Streamline Protocol for Arabidopsis Apoplastic Fluid Isolation Enables a Detailed Proteomic View of the Plant Extracellular Space","volume":"9","author":"Chen K.\u2010E.","year":"2025","journal-title":"Plant Direct"},{"issue":"4","key":"e_1_2_14_12_1","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1111\/tpj.15730","article-title":"Systemic Silencing of an Endogenous Plant Gene by Two Classes of Mobile 21\u2010Nucleotide Artificial Small RNAs","volume":"110","author":"Cisneros A. 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