{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T12:34:11Z","timestamp":1786624451607,"version":"build-2736575974"},"reference-count":102,"publisher":"Frontiers Media SA","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\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Oncol."],"abstract":"<jats:p>Myelodysplastic syndromes (MDS) are a heterogeneous group of disorders caused by abnormalities at the hematopoietic stem cell level and are associated with ineffective hematopoiesis, peripheral cytopenias, and a propensity for leukemic transformation. Recent research elucidates the key mechanisms and roles of ribosomal proteins (RPs), microRNAs (miRNAs), extracellular vesicles (EVs), and long non-coding RNAs (lncRNAs) in the pathophysiology of MDS. Mutations or altered expression of RP genes lead to ribosomal stress and activation of p53, resulting in impaired erythropoiesis and apoptosis of hematopoietic progenitors. Dysregulation of miRNAs modulates gene expression programs that are essential for hematopoietic differentiation and apoptosis. Moreover, EV-associated miRNAs mediate multiple processes, including intercellular communication and regulation of the bone marrow (BM) microenvironment, thereby contributing to DNA damage and clonal evolution. EV-associated miR-10a and miR-15a have been reported to induce DNA damage in HSCs. LncRNAs have emerged as promising tools for cancer diagnosis and prognostic biomarkers. They play critical roles in regulating malignant cell proliferation, apoptosis, and epigenetic modifications. This overview highlights the molecular complexity of MDS pathogenesis, offering insights to support stratified diagnosis and the development of targeted therapies.<\/jats:p>","DOI":"10.3389\/fonc.2026.1863527","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T12:00:46Z","timestamp":1786622446000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Ribosomal proteins, miRNAs, and extracellular vesicles in myelodysplastic syndromes: molecular regulation and biomarker potential"],"prefix":"10.3389","volume":"16","author":[{"given":"Gazmend","family":"Temaj","sequence":"first","affiliation":[{"name":"Faculty of Pharmacy, College UBT","place":["Prishtina, Kosovo"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarmistha","family":"Saha","sequence":"additional","affiliation":[{"name":"Department of Biotechnology, GLA University","place":["Mathura, Uttar Pradesh, India"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brigitta","family":"Buttari","sequence":"additional","affiliation":[{"name":"Department of Cardiovascular and Endocrine-Metabolic Diseases and Aging, Istituto Superiore di Sanit\u00e0","place":["Rome, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pelin","family":"Telkoparan-Akillilar","sequence":"additional","affiliation":[{"name":"Department of Medical Biology, Faculty of Medicine, Gazi University","place":["Ankara, T\u00fcrkiye"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Silvia","family":"Chichiarelli","sequence":"additional","affiliation":[{"name":"Department of Biochemical Sciences \u201cA. Rossi-Fanelli\u201d, Sapienza University of Rome","place":["Rome, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nexhibe","family":"Nuhii","sequence":"additional","affiliation":[{"name":"Department of Pharmacy, Faculty of Medical Sciences, State University of Tetovo","place":["Tetovo, North Macedonia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rifat","family":"Hadziselimovic","sequence":"additional","affiliation":[{"name":"Faculty of Science, University of Sarajevo","place":["Sarajevo, Bosnia and Herzegovina"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luciano","family":"Saso","sequence":"additional","affiliation":[{"name":"Department of Physiology and Pharmacology \u201cVittorio Erspamer\u201d, Sapienza University of Rome","place":["Rome, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,13]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cll.2011.08.005","article-title":"Myelodysplastic syndromes","volume":"31","author":"Odenike","year":"2011","journal-title":"Clin Lab Med"},{"key":"B2","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1038\/s41392-024-01735-1","article-title":"Extracellular vesicles as tools and targets in therapy for diseases","volume":"9","author":"Kumar","year":"2024","journal-title":"Signal Transd Targ Ther"},{"key":"B3","doi-asserted-by":"publisher","first-page":"595","DOI":"10.1042\/BST20190854","article-title":"MicroRNAs and long non-coding RNAs as novel regulators of ribosome biogenesis","volume":"48","author":"McCool","year":"2020","journal-title":"Biochem Soc Trans"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1038\/nature06494","article-title":"Identification of RPS14 as a 5q- syndrome gene by RNA interference screen","volume":"451","author":"Ebert","year":"2008","journal-title":"Nature"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1007\/s00277-012-1486-2","article-title":"Over-expression of RPL23 in myelodysplastic syndromes is associated with apoptosis resistance of CD34+ cells and predicts poor prognosis and distinct response to CHG chemotherapy or decitabine","volume":"91","author":"Wu","year":"2012","journal-title":"Ann Hematol"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1182\/blood.2021014472","article-title":"Molecular characterization of mutant TP53 acute myeloid leukemia and high-risk myelodysplastic syndrome","volume":"139","author":"Grob","year":"2022","journal-title":"Blood"},{"key":"B7","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1038\/nm.2054","article-title":"Identification of miR-145 and miR-146a as mediators of the 5q\u2013 syndrome phenotype","volume":"16","author":"Starczynowski","year":"2010","journal-title":"Nat Med"},{"key":"B8","doi-asserted-by":"publisher","first-page":"31","DOI":"10.14715\/cmb\/2022.68.2.5","article-title":"Effect of miR-223-3p on cell pyroptosis in myelodysplastic syndrome and itsmechanism via regulating the expression of NLRP3","volume":"68","author":"Yin","year":"2022","journal-title":"Cell Mol Biol"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.3390\/cells12040658","article-title":"Extracellular vesicles and microRNA in myelodysplastic syndromes","volume":"12","author":"Meunier","year":"2023","journal-title":"Cells"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1038\/onc.2017.184","article-title":"LncRNA-mediated regulation of cell signaling in cancer","volume":"36","author":"Peng","year":"2017","journal-title":"Oncogene"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.3389\/fonc.2020.581007","article-title":"microRNAs biogenesis, functions and role in tumor angiogenesis","volume":"10","author":"Annese","year":"2020","journal-title":"Front Oncol"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1003018","article-title":"Virus-encoded microRNAs: an overview and a look to the future","volume":"8","author":"Kincaid","year":"2012","journal-title":"PloS Pathog"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(93)90529-Y","article-title":"The C. elegans heterochronic gene lin-4 encodes small RNAs with antisense complementarity to lin-14","volume":"75","author":"Lee","year":"1993","journal-title":"Cell"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(04)00034-0","article-title":"The 20 years it took to recognize the importance of tiny RNAs","volume":"116","author":"Ruvkun","year":"2004","journal-title":"Cell"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(93)90530-4","article-title":"Posttranscriptional regulation of the heterochronic gene lin-14 by lin-4 mediates temporal pattern formation in C. elegans","volume":"75","author":"Wightman","year":"1993","journal-title":"Cell"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3390\/toxins9060181","article-title":"Small and smaller\u2014sRNAs and microRNAs in the regulation of toxin gene expression in prokaryotic cells: a mini-review","volume":"9","author":"Bloch","year":"2017","journal-title":"Toxins"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-024-00805-0","article-title":"The biogenesis and regulation of animal microRNAs","volume":"26","author":"Kim","year":"2025","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1038\/nrm3838","article-title":"Regulation of microRNA biogenesis","volume":"15","author":"Ha","year":"2014","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B19","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1038\/nrg2843","article-title":"The widespread regulation of microRNA biogenesis, function and decay","volume":"11","author":"Krol","year":"2010","journal-title":"Nat Rev Genet"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1038\/nrg3462","article-title":"Argonaute proteins: functional insights and emerging roles","volume":"14","author":"Meister","year":"2013","journal-title":"Nat Rev Genet"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1038\/emboj.2013.44","article-title":"GW182 proteins cause PABP dissociation from silenced miRNA targets in the absence of deadenylation","volume":"32","author":"Zekri","year":"2013","journal-title":"EMBO J"},{"key":"B22","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1038\/s41580-020-00315-9","article-title":"Gene regulation by long non-coding RNAs and its biological functions","volume":"22","author":"Statello","year":"2021","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B23","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1016\/j.gde.2020.03.002","article-title":"Gene regulation by non-coding RNAs in the 3D genome architecture","volume":"61","author":"Tachiwana","year":"2020","journal-title":"Curr Opin Genet Dev"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1101\/gad.1953310","article-title":"Canonical and alternate functions of the microRNA biogenesis machinery","volume":"24","author":"Chong","year":"2010","journal-title":"Genes Dev"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2018.00402","article-title":"Overview of microRNA biogenesis, mechanisms of actions, and circulation","volume":"9","author":"O\u2019Brien","year":"2018","journal-title":"Front Endocrinol"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-biophys-101922-072452","article-title":"Decoding and recoding of mRNA sequences by the ribosome","volume":"52","author":"Rodnina","year":"2023","journal-title":"Annu Rev Biophys"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a032664","article-title":"Translation in prokaryotes","volume":"10","author":"Rodnina","year":"2018","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1038\/nature07582","article-title":"Quality control by the ribosome following peptide bond formation","volume":"457","author":"Zaher","year":"2009","journal-title":"Nature"},{"key":"B29","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/j.trecan.2020.08.003","article-title":"Dysregulated ribosome biogenesis reveals therapeutic liabilities in cancer","volume":"7","author":"Bursa\u0107","year":"2021","journal-title":"Trends Cancer"},{"key":"B30","first-page":"1","article-title":"Beitraege zur Histologie und Aetiologie der Carconoms. Contributions to the histology and etiology of carcinomas","volume":"14","author":"Pianese","year":"1896","journal-title":"Beitr Pathol Anat Allgem Pathol"},{"key":"B31","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.bcp.2018.11.014","article-title":"Ribosome biogenesis: an emerging druggable pathway for cancer therapeutics","volume":"159","author":"Catez","year":"2019","journal-title":"Biochem Pharmacol"},{"key":"B32","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1038\/nrc.2017.104","article-title":"Ribosome biogenesis in cancer: new players and therapeutic avenues","volume":"18","author":"Pelletier","year":"2018","journal-title":"Nat Rev Cancer"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.3390\/biomedicines10092088","article-title":"Ribosome-directed therapies in cancer","volume":"10","author":"Temaj","year":"2022","journal-title":"Biomedicines"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1038\/5951","article-title":"The gene encoding ribosomal protein S19 is mutated in Diamond-Blackfan anaemia","volume":"21","author":"Draptchinskaia","year":"1999","journal-title":"Nat Genet"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1080\/17512433.2022.2098110","article-title":"Ribosomopathies and cancer: pharmacological implications","volume":"15","author":"Temaj","year":"2022","journal-title":"Expert Rev Clin Pharmacol"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1242\/dmm.020529","article-title":"Ribosomopathies: how a common root can cause a tree of pathologies","volume":"8","author":"Danilova","year":"2015","journal-title":"Dis Model Mech"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.3390\/genes12091392","article-title":"Treacher Collins syndrome: genetics, clinical features and management","volume":"12","author":"Marsza\u0142ek-Kruk","year":"2021","journal-title":"Genes"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.94.7.3110","article-title":"TCOF1 gene encodes a putative nucleolar phosphoprotein that exhibits mutations in Treacher Collins syndrome throughout its coding region","volume":"94","author":"Wise","year":"1997","journal-title":"Proc Natl Acad Sci"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1038\/s41390-022-02009-8","article-title":"Shwachman Diamond syndrome: narrow genotypic spectrum and variable clinical features","volume":"92","author":"Thompson","year":"2022","journal-title":"Pediatr Res"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.3390\/cells9102300","article-title":"How altered ribosome production can cause or contribute to human disease: the spectrum of ribosomopathies","volume":"9","author":"Venturi","year":"2020","journal-title":"Cells"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1016\/j.xjidi.2023.100240","article-title":"En route to targeted ribosome editing to replenish skin anchor protein LAMB3 in junctional epidermolysis bullosa","volume":"4","author":"Wimmer","year":"2024","journal-title":"JID Innov"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.3390\/cancers16132392","article-title":"The paradox of ribosomal insufficiency coupled with increased cancer: shifting the perspective from the cancer cell to the microenvironment","volume":"16","author":"D\u2019Andrea","year":"2024","journal-title":"Cancers"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2023.1194788","article-title":"Editorial: Inherited and acquired ribosomopathies: missing puzzle pieces","volume":"14","author":"Valli","year":"2023","journal-title":"Front Genet"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.3390\/cells10010110","article-title":"microRNAs mediated regulation of the ribosomal proteins and its consequences on the global translation of proteins","volume":"10","author":"Reza","year":"2021","journal-title":"Cells"},{"key":"B45","doi-asserted-by":"publisher","first-page":"1988","DOI":"10.1093\/nar\/gkad1235","article-title":"Discovery of novel microRNA mimic repressors of ribosome biogenesis","volume":"52","author":"Bryant","year":"2024","journal-title":"Nucleic Acids Res"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.3390\/diagnostics12030629","article-title":"The profile of microRNA expression in bone marrow in non-Hodgkin\u2019s lymphomas","volume":"12","author":"Veryaskina","year":"2022","journal-title":"Diagnostics"},{"key":"B47","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1038\/s41392-021-00728-8","article-title":"Ribosomal proteins and human diseases: molecular mechanisms and targeted therapy","volume":"6","author":"Kang","year":"2021","journal-title":"Signal Transd Targ Ther"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1631\/jzus.B2300904","article-title":"Mechanisms of ribosomopathy and phase separation-related ribosomopathy","volume":"26","author":"Pan","year":"2025","journal-title":"J Zhejiang Univ-Sci B"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.18632\/oncotarget.7127","article-title":"Tumor suppressor microRNAs are downregulated in myelodysplastic syndrome with spliceosome mutations","volume":"7","author":"Aslan","year":"2016","journal-title":"Oncotarget"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.4137\/CMBD.S16952","article-title":"Ribosomopathies: mechanisms of disease","volume":"7","author":"Nakhoul","year":"2014","journal-title":"Clin Med Insights Blood Disord"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.18632\/oncotarget.10293","article-title":"RPL27A is a target of miR-595 and may contribute to the myelodysplastic phenotype through ribosomal dysgenesis","volume":"7","author":"Alkhatabi","year":"2016","journal-title":"Oncotarget"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/j.pathol.2020.03.010","article-title":"The effect of miR-223 on cellular behaviour in non-5q myelodysplastic syndromes through targeting RPS14","volume":"52","author":"Nie","year":"2020","journal-title":"Pathol (Phila)"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1007\/s12185-017-2201-9","article-title":"Over-expression of miR-196b-5p is significantly associated with the progression of myelodysplastic syndrome","volume":"105","author":"Wen","year":"2017","journal-title":"Int J Hematol"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.3389\/frhem.2023.1184962","article-title":"MicroRNA profiling in aplastic anemia reveals similarities between secondary myelodysplastic syndromes arising from clonal progression and de novo MDS","volume":"2","author":"Chee","year":"2023","journal-title":"Front Hematol"},{"key":"B55","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.bbrc.2014.11.085","article-title":"MicroRNA-196b promotes cell proliferation and suppress cell differentiation in vitro","volume":"457","author":"Cao","year":"2015","journal-title":"Biochem Biophys Res Commun"},{"key":"B56","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1038\/s41419-022-04578-2","article-title":"LncRNA BC200\/miR-150-5p\/MYB positive feedback loop promotes the Malignant proliferation of myelodysplastic syndrome","volume":"13","author":"Liu","year":"2022","journal-title":"Cell Death Dis"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.3390\/ncrna10060054","article-title":"Extracellular vesicle lncRNAs as key biomolecules for cell-to-cell communication and circulating cancer biomarkers","volume":"10","author":"Papoutsoglou","year":"2024","journal-title":"Non-Coding RNA"},{"key":"B58","doi-asserted-by":"publisher","first-page":"894","DOI":"10.3324\/haematol.2018.208926","article-title":"Long noncoding RNAs of single hematopoietic stem and progenitor cells in healthy and dysplastic human bone marrow","volume":"104","author":"Wu","year":"2019","journal-title":"Haematologica"},{"key":"B59","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1016\/j.ncrna.2020.12.002","article-title":"Role of miRNAs and lncRNAs in hematopoietic stem cell differentiation","volume":"6","author":"Ghafouri-Fard","year":"2021","journal-title":"Non-coding RNA Res"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2025.116688","article-title":"Ribosomal protein control of hematopoietic stem cell transformation through regulation of metabolism","volume":"44","author":"Harris","year":"2025","journal-title":"Cell Rep"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1182\/blood.2021011992","article-title":"Long noncoding RNAs: emerging regulators of normal and Malignant hematopoiesis","volume":"138","author":"Qiu","year":"2021","journal-title":"Blood"},{"key":"B62","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1016\/j.celrep.2019.03.080","article-title":"lncRNA Spehd regulates hematopoietic stem and progenitor cells and is required for multilineage differentiation","volume":"27","author":"Del\u00e1s","year":"2019","journal-title":"Cell Rep"},{"key":"B63","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1038\/s41580-023-00694-9","article-title":"Transcription regulation by long non-coding RNAs: mechanisms and disease relevance","volume":"25","author":"Ferrer","year":"2024","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1016\/j.sbi.2024.102866","article-title":"Long non-coding RNAs in the nucleolus: Biogenesis, regulation, and function","volume":"87","author":"Han","year":"2024","journal-title":"Curr Opin Struct Biol"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1093\/narcan\/zcae017","article-title":"The impact of ribosome biogenesis in cancer: from proliferation to metastasis","volume":"6","author":"Hwang","year":"2024","journal-title":"NAR Cancer"},{"key":"B66","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.4103\/1673-5374.358615","article-title":"Long-noncoding RNAs as epigenetic regulators in neurodegenerative diseases","volume":"18","author":"Ruffo","year":"2023","journal-title":"Neural Regener Res"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.3390\/cancers15194810","article-title":"The role of non-coding RNAs in myelodysplastic neoplasms","volume":"15","author":"Georgoulis","year":"2023","journal-title":"Cancers"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1182\/bloodadvances.2017008284","article-title":"A 4-lncRNA scoring system for prognostication of adult myelodysplastic syndromes","volume":"1","author":"Yao","year":"2017","journal-title":"Blood Adv"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2024.1414269","article-title":"Ribosome heterogeneity in development and disease","volume":"12","author":"Gao","year":"2024","journal-title":"Front Cell Dev Biol"},{"key":"B70","doi-asserted-by":"publisher","first-page":"20230379","DOI":"10.1098\/rstb.2023.0379","article-title":"Impacts of ribosomal RNA sequence variation on gene expression and phenotype","volume":"380","author":"Welfer","year":"2025","journal-title":"Philos Trans R Soc B Biol Sci"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.1126\/science.aau6977","article-title":"The biology , function , and biomedical applications of exosomes","volume":"367","author":"Kalluri","year":"2020","journal-title":"Science"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1038\/nrg.2016.4","article-title":"Enhancers as non-coding RNA transcription units: recent insights and future perspectives","volume":"17","author":"Li","year":"2016","journal-title":"Nat Rev Genet"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-022-00460-3","article-title":"Challenges and directions in studying cell\u2013cell communication by extracellular vesicles","volume":"23","author":"Van Niel","year":"2022","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B74","doi-asserted-by":"publisher","first-page":"20230382","DOI":"10.1098\/rstb.2023.0382","article-title":"Built differently or defective: can RNA exosomopathies cause ribosome heterogeneity","volume":"380","author":"Bressman","year":"2025","journal-title":"Philos Trans R Soc B Biol Sci"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.3390\/ijms241411458","article-title":"Two \u201cEdges\u201d in our knowledge on the functions of ribosomal proteins: The revealed contributions of their regions to translation mechanisms and the issues of their extracellular transport by exosomes","volume":"24","author":"Ochkasova","year":"2023","journal-title":"Int J Mol Sci"},{"key":"B76","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1093\/cvr\/cvac031","article-title":"Methods for the identification and characterization of extracellular vesicles in cardiovascular studies: from exosomes to microvesicles","volume":"119","author":"Davidson","year":"2023","journal-title":"Cardiovasc Res"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.2147\/IJN.S264498","article-title":"Exosome: A review of its classification, isolation techniques, storage, diagnostic and targeted therapy applications","volume":"15","author":"Zhang","year":"2020","journal-title":"Int J Nanomed"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.1038\/s41576-020-0270-8","article-title":"The evolving metabolic landscape of chromatin biology and epigenetics","volume":"21","author":"Dai","year":"2020","journal-title":"Nat Rev Genet"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1618088114","article-title":"Phosphoproteins in extracellular vesicles as candidate markers for breast cancer","volume":"114","author":"Chen","year":"2017","journal-title":"Proc Natl Acad Sci"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1038\/s41375-020-0738-8","article-title":"Extracellular vesicles from myelodysplastic mesenchymal stromal cells induce DNA damage and mutagenesis of hematopoietic stem cells through miRNA transfer","volume":"34","author":"Meunier","year":"2020","journal-title":"Leukemia"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.1007\/s11626-025-01098-z","article-title":"Therapeutic potential of miR-10a overexpressing mesenchymal stem cell\u2013derived extracellular vesicles in modulating inflammation in collagen-induced arthritis","volume":"61","author":"Bai","year":"2025","journal-title":"Vitro Cell Dev Biol - Anim"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2021.629893","article-title":"MicroRNA-15a carried by mesenchymal stem cell-derived extracellular vesicles inhibits the immune evasion of colorectal cancer cells by regulating the KDM4B\/HOXC4\/PD-L1 axis","volume":"9","author":"Liu","year":"2021","journal-title":"Front Cell Dev Biol"},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.3389\/fonc.2024.1410656","article-title":"MicroRNA dysregulation in myelodysplastic syndromes: implications for diagnosis, prognosis, and therapeutic response","volume":"14","author":"Micheva","year":"2024","journal-title":"Front Oncol"},{"key":"B84","doi-asserted-by":"publisher","first-page":"4","DOI":"10.32607\/actanaturae.11209","article-title":"MicroRNAs in the myelodysplastic syndrome","volume":"13","author":"Veryaskina","year":"2021","journal-title":"Acta Naturae"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.3892\/ol.2020.11282","article-title":"Downregulation of extracellular vesicle microRNA-101 derived from bone marrow mesenchymal stromal cells in myelodysplastic syndrome with disease progression","volume":"19","author":"Saitoh","year":"2020","journal-title":"Oncol Lett"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.3324\/haematol.2017.182824","article-title":"Circulating exosomal microRNAs in acquired aplastic anemia and myelodysplastic syndromes","volume":"103","author":"Giudice","year":"2018","journal-title":"Haematologica"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.3324\/haematol.2017.183335","article-title":"Extracellular vesicles in the hematopoietic microenvironment","volume":"103","author":"Butler","year":"2018","journal-title":"Haematologica"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.3390\/biomedicines13102353","article-title":"MicroRNAs in BM-MSC-derived extracellular vesicles promote angiogenesis: An in vitro model study","volume":"13","author":"Kusakabe","year":"2025","journal-title":"Biomedicines"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.1016\/j.febslet.2014.04.014","article-title":"p53 and ribosome biogenesis stress: The essentials","volume":"588","author":"Golomb","year":"2014","journal-title":"FEBS Lett"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1016\/j.ymgme.2015.07.004","article-title":"Biology of the bone marrow microenvironment and myelodysplastic syndromes","volume":"116","author":"Rankin","year":"2015","journal-title":"Mol Genet Metab"},{"key":"B91","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1038\/s41421-022-00505-z","article-title":"Extracellular vesicles transfer chromatin-like structures that induce non-mutational dysfunction of p53 in bone marrow stem cells","volume":"9","author":"Ghanam","year":"2023","journal-title":"Cell Discov"},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2025.1540313","article-title":"PIWI proteins and piRNAs: key regulators of stem cell biology","volume":"13","author":"Claro-Linares","year":"2025","journal-title":"Front Cell Dev Biol"},{"key":"B93","doi-asserted-by":"publisher","first-page":"1566","DOI":"10.1016\/j.cell.2019.11.022","article-title":"A translation-activating function of MIWI\/piRNA during mouse spermiogenesis","volume":"179","author":"Dai","year":"2019","journal-title":"Cell"},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0605506103","article-title":"MIWI associates with translational machinery and PIWI-interacting RNAs (piRNAs) in regulating spermatogenesis","volume":"103","author":"Grivna","year":"2006","journal-title":"Proc Natl Acad Sci"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.1101\/gad.245514.114","article-title":"Transgenerationally inherited piRNAs trigger piRNA biogenesis by changing the chromatin of piRNA clusters and inducing precursor processing","volume":"28","author":"Le Thomas","year":"2014","journal-title":"Genes Dev"},{"key":"B96","doi-asserted-by":"publisher","DOI":"10.3390\/ncrna10020019","article-title":"Functions of RNAi pathways in ribosomal RNA regulation","volume":"10","author":"Shatskikh","year":"2024","journal-title":"Non-Coding RNA"},{"key":"B97","doi-asserted-by":"publisher","first-page":"RP104295","DOI":"10.7554\/eLife.104295","article-title":"Meioc-Piwil1 complexes regulate rRNA transcription for differentiation of spermatogonial stem cells","volume":"14","author":"Kawasaki","year":"2025","journal-title":"eLife"},{"key":"B98","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.69322","article-title":"Haploinsufficiency of the essential gene Rps12 causes defects in erythropoiesis and hematopoietic stem cell maintenance","volume":"12","author":"Folgado-Marco","year":"2023","journal-title":"eLife"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.5045\/br.2014.49.4.216","article-title":"Pathogenesis of myelodysplastic syndromes: an overview of molecular and non-molecular aspects of the disease","volume":"49","author":"Visconte","year":"2014","journal-title":"Blood Res"},{"key":"B100","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2010-11-318584","article-title":"Reduced ribosomal protein gene dosage and p53 activation in low-risk myelodysplastic syndrome","volume":"118","author":"McGowan","year":"2011","journal-title":"Blood"},{"key":"B101","doi-asserted-by":"publisher","first-page":"2323","DOI":"10.1038\/s41598-017-02403-x","article-title":"Ribosomal protein L23 negatively regulates cellular apoptosis via the RPL23\/Miz-1\/c-Myc circuit in higher-risk myelodysplastic syndrome","volume":"7","author":"Qi","year":"2017","journal-title":"Sci Rep"},{"key":"B102","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1080\/10245332.2016.1193962","article-title":"Deregulated microRNA expression and its pathogenetic implications for myelodysplastic syndromes","volume":"21","author":"Kuang","year":"2016","journal-title":"Hematology"}],"container-title":["Frontiers in Oncology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fonc.2026.1863527\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T12:00:47Z","timestamp":1786622447000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fonc.2026.1863527\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,13]]},"references-count":102,"alternative-id":["10.3389\/fonc.2026.1863527"],"URL":"https:\/\/doi.org\/10.3389\/fonc.2026.1863527","relation":{},"ISSN":["2234-943X"],"issn-type":[{"value":"2234-943X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,13]]},"article-number":"1863527"}}