1. Chen L-L. The biogenesis and emerging roles of circular RNAs. Nature reviews Molecular cell biology. 2016;17(4):205-11. https://doi.org/ 10.1038/nrm.2015.32. [
DOI:10.1038/nrm.2015.32.]
2. De Fraipont F, Gazzeri S, Cho WC, Eymin B. Circular RNAs and RNA splice variants as biomarkers for prognosis and therapeutic response in the liquid biopsies of lung cancer patients. Frontiers in genetics. 2019;10:390. [
DOI:10.3389/fgene.2019.00390.]
3. Jeck WR, Sharpless NE. Detecting and characterizing circular RNAs. Nature biotechnology. 2014;32(5):453-61. [
DOI:10.1038/nbt.2890.]
4. Huang Y, Zhu Q. Mechanisms regulating abnormal circular RNA biogenesis in cancer. Cancers. 2021;13(16):4185. [
DOI:10.3390/cancers13164185.]
5. Liang Y, Liu N, Yang L, Tang J, Wang Y, Mei M. A brief review of circRNA biogenesis, detection, and function. Current genomics. 2021;22(7):485-95. [
DOI:10.2174/1389202922666210331130722.]
6. Hwang HJ, Kim YK. Molecular mechanisms of circular RNA translation. Experimental S molecular medicine. 2024;56(6):1272-80. [
DOI:10.1038/s12276-024- 01220-3.]
7. Welden JR, Stamm S. Pre-mRNA structures forming circular RNAs. Biochimica et Biophysica Acta (BBA)-Gene Regulatory Mechanisms. 2019;1862(11-12):194410. [
DOI:10.1016/j.bbagrm.2019.194410.]
8. Kristensen LS, Andersen MS, Stagsted LV, Ebbesen KK, Hansen TB, Kjems J. The biogenesis, biology and characterization of circular RNAs. Nature reviews genetics. 2019;20(11):675-91. [
DOI:10.1038/s41576-019-0158-7.]
9. Tagawa T, Gao S, Koparde VN, Gonzalez M, Spouge JL, Serquiña AP, et al. Discovery of Kaposi’s sarcoma herpesvirus-encoded circular RNAs and a human antiviral circular RNA. Proceedings of the National Academy of Sciences. 2018;115(50):12805-10. [
DOI:10.1073/pnas.1816183115.]
10. Xu F, Xiao Q, Du WW, Wang S, Yang BB. CircRNA: Functions, applications and prospects. Journal. 2024;14(Issue):1503. [
DOI:10.3390/biom14121503]
11. Zhang J, Wang H, Wu K, Zhan F, Zeng H. Dysregulated circRNA_100876 contributes to proliferation and metastasis of colorectal cancer by targeting microRNA-516b (miR- 516b). Cancer Biology S Therapy. 2020;21(8):733-40. [
DOI:10.1080/15384047.2020.1776075]
12. Liu X, Zhang Y, Zhou S, Dain L, Mei L, Zhu G. Circular RNA: An emerging frontier in RNA therapeutic targets, RNA therapeutics, and mRNA vaccines. Journal of Controlled Release. 2022;348:84-94. [
DOI:10.1016/j.jconrel.2022.05.043]
13. Petkovic S, Müller S. RNA circularization strategies in vivo and in vitro. Nucleic acids research. 2015;43(4):2454-65. [
DOI:10.1093/nar/gkv045]
14. Zhang X, Wu H, Hong X, Xiao Y, Zeng X. Circular RNA: From Non-coding Regulators to Functional Protein Encoders. Pharmaceutical Science Advances. 2025:100085. [
DOI:10.1016/j.pscia.2025.100085.]
15. Waldern JM, Taylor C, Giannetti CA, Irving PS, Allen SR, Zhu M, et al. Structural determinants of inverted Alu-mediated backsplicing revealed by-MaP and-JuMP. Nucleic Acids Research. 2025;53(9):gkaf433. [
DOI:10.1093/nar/gkaf433]
16. Zhang D, Ma Y, Naz M, Ahmed N, Zhang L, Zhou J-J, et al. Advances in CircRNAs in the Past Decade: Review of CircRNAs Biogenesis, Regulatory Mechanisms, and Functions in Plants. Genes. 2024;15(7):958. [
DOI:10.3390/genes15070958]
17. Huang A, Zheng H, Wu Z, Chen M, Huang Y. Circular RNA-protein interactions: functions, mechanisms, and identification. Theranostics. 2020;10(8):3503. [
DOI:10.7150/thno.42174]
18. Ebbesen KK, Hansen TB, Kjems J. Insights into circular RNA biology. RNA biology. 2017;14(8):1035-45. [
DOI:10.1080/15476286.2016.1271524]
19. Misir S, Wu N, Yang BB. Specific expression and functions of circular RNAs. Cell Death S Differentiation. 2022;29(3):481-91. [
DOI:10.1038/s41418-022- 00948-7.]
20. Czubak K, Sedehizadeh S, Kozlowski P, Wojciechowska M. An overview of circular RNAs and their implications in myotonic dystrophy. International journal of molecular sciences. 2019;20(18):4385. [
DOI:10.3390/ijms20184385]
21. Conn SJ, Pillman KA, Toubia J, Conn VM, Salmanidis M, Phillips CA, et al. The RNA binding protein quaking regulates formation of circRNAs. Cell. 2015;160(6):1125-34. [
DOI:10.1016/j.cell.2015.02.014]
22. Luo Y, Blechingberg J, Fernandes AM, Li S, Fryland T, Børglum AD, et al. EWS and FUS bind a subset of transcribed genes encoding proteins enriched in RNA regulatory functions. BMC genomics. 2015;16(1):929. https://doi.org/ 10.1186/s12864-015- 2125-9. [
DOI:10.1186/s12864-015- 2125-9.]
23. Ho JS, Di Tullio F, Schwarz M, Low D, Incarnato D, Gay F, et al. HNRNPM controls circRNA biogenesis and splicing fidelity to sustain cancer cell fitness. Elife. 2021;10:e59654. [
DOI:10.7554/eLife.59654.]
24. Feng J, Zhou J, Lin Y, Huang W. hnRNP A1 in RNA metabolism regulation and as a potential therapeutic target. Frontiers in Pharmacology. 2022;13:986409.
25. Wu J, Qi X, Liu L, Hu X, Liu J, Yang J, et al. Emerging epigenetic regulation of circular RNAs in human cancer. Molecular therapy Nucleic acids. 2019;16:589-96. [
DOI:10.1016/j.omtn.2019.04.011]
26. Tarrero LC, Ferrero G, Miano V, De Intinis C, Ricci L, Arigoni M, et al. Luminal breast cancer-specific circular RNAs uncovered by a novel tool for data analysis. Oncotarget. 2018;9(18):14580. [
DOI:10.18632/oncotarget.24522.]
27. Huang S, Han Y, Liu Y, Bu L, Wu C, Rao Z, et al. Super-enhancer-mediated circRNAs exhibit high splicing circularization diversity and transcriptional activity. Nucleic Acids Research. 2025;53(11):gkaf505. [
DOI:10.1093/nar/gkaf505]
28. Zhang M, Xu K, Fu L, Wang Q, Chang Z, Zou H, et al. Revealing epigenetic factors of circRNA expression by machine learning in various cellular contexts. Iscience. 2020;23(12). https://doi.org/ 10.1016/j.isci.2020.101842 [
DOI:10.1016/j.isci.2020.101842]
29. Zhang C, Cui H, Huang C, Kong F, Yang Q, Miao P, et al. Interactions of circRNAs with methylation: An important aspect of circRNA biogenesis and function. Molecular Medicine Reports. 2022;25(5):1-15. https://doi.org/ 10.3892/mmr.2022.12685 [
DOI:10.3892/mmr.2022.12685]
30. Zeng W, Zhu J-F, Guo J, Huang G-J, Ai L-S, Zeng Y, et al. m6A-modified circFNDC3B inhibits colorectal cancer stemness and metastasis via RNF41-dependent ASB6 degradation.Cell death S disease. 2022;13(11):1008. [
DOI:10.1038/s41419-022-05451- y.]
31. Amelimojarad M, Amelimojarad M. Regulatory mechanism of circular RNAs in brain and neurodegenerative diseases. Frontiers in Molecular Neuroscience. 2025;18:1507575. [
DOI:10.3389/fnmol.2025.1507575]
32. Golovina E, Eaton C, Cox V, Andel J, Savvulidi Vargova K. Mechanism of Action of circRNA/miRNA Network in DLBCL. Non-coding RNA. 2025;11(2):22. [
DOI:10.3390/ncrna11020022]
33. Karimi R, Javandoost E, Asadmasjedi N, Atashi A, Soleimani A, Behzadifard M. Circular RNAs: history, metabolism, mechanisms of function, and regulatory roles at a glance. Annals of Medicine and Surgery. 2025;87(1):141-50. [
DOI:10.1097/MS9.0000000000002761]
34. Yuan H, Liao X, Hu D, Guan D, Tian M. Back to the Origin: Mechanisms of circRNA- Directed Regulation of Host Genes in Human Disease. Non-coding RNA. 2024;10(5):49. [
DOI:10.3390/ncrna10050049]
35. Gao Y, Li C, Ji T, Yu K, Gao X. The biological function and mechanism of action of circRNA as a potential target in colorectal cancer. Critical Reviews in Oncology/Hematology. 2025:104828. 10.1016/j.critrevonc.2025.104828.
36. Zhang N, Wang X, Li Y, Lu Y, Sheng C, Sun Y, et al. Mechanisms and therapeutic implications of gene expression regulation by circRNA-protein interactions in cancer. Communications Biology. 2025;8(1):77. [
DOI:10.1038/s42003-024-07383-z]
37. Wang Y, Cui Y, Li X, Jin S-H, Wang H, Gaipl US, et al. CircRNAs: functions and emerging roles in cancer and immunotherapy. BMC medicine. 2025;23(1):477. [
DOI:10.1186/s12916-025-04306-5]
38. Hatzimanolis O, Sykes AM, Cristino AS. Circular RNAs in neurological conditions–computational identification, functional validation, and potential clinical applications. Molecular Psychiatry. 2025:1-24.
39. Zhang J, Zhou R, Zhang H, Peng Y, Meng J, Xi W, et al. Decoding circRNA translation: challenges and advances in computational method development. Frontiers in Genetics. 2025;16:1654305. [
DOI:10.3389/fgene.2025.1654305]
40. Margvelani G, Maquera KAA, Welden JR, Rodgers DW, Stamm S. Translation of circular RNAs. Nucleic acids research. 2025;53(1):gkae1167. [
DOI:10.1093/nar/gkae1167.]
41. Austin CP. Translating translation. Nature Reviews Drug Discovery. 2018;17(7):455- 6.
42. Zhang S, Wang W, Wu X, Zhou X. Regulatory roles of circular RNAs in coronary artery disease. Molecular therapy Nucleic acids. 2020;21:172-9. https://doi.org/ 10.1016/j.omtn.2020.05.024. [
DOI:10.1016/j.omtn.2020.05.024.]
43. Liu X, Wang X, Li J, Hu S, Deng Y, Yin H, et al. Identification of mecciRNAs and their roles in the mitochondrial entry of proteins. Science China Life sciences. 2020;63(10):1429-49. [
DOI:10.1007/s11427-020-1631-9]
44. Liu X, Yang Y, Shan G. Identification and detection of mecciRNAs. Methods. 2021;196:147-52. [
DOI:10.1016/j.ymeth.2021.02.006.]
45. Ren B, Guan M-X, Zhou T, Cai X, Shan G. Emerging functions of mitochondria- encoded noncoding RNAs. Trends in Genetics. 2023;39(2):125-39. [
DOI:10.1016/j.tig.2022.08.004.]
46. Garrido P, Casas-Benito A, Larrayoz IM, Narro-Íñiguez J, Rubio-Mediavilla S, Zozaya E, et al. Expression of Mitochondrial Long Non-Coding RNAs, MDL1 and MDL1AS, Are Good Prognostic and/or Diagnostic Biomarkers for Several Cancers, Including Colorectal Cancer. Cancers. 2024;16(5):960. [
DOI:10.3390/cancers16050960.]
47. Feng X-Y, Zhu S-X, Pu K-J, Huang H-J, Chen Y-Q, Wang W-T. New insight into circRNAs: characterization, strategies, and biomedical applications. Experimental hematology S oncology. 2023;12(1):91. [
DOI:10.1186/s40164-023-00451- w.]
48. Hua X, Yu L, Zhu H, Zhu Y, Fan G, Zhou G. Research progress of circRNAs in bone- related diseases. Frontiers in Oncology. 2025;15:1481322. [
DOI:10.3389/fonc.2025.1481322]
49. Zhang Z, Wang Z. Cellular functions and biomedical applications of circular RNAs: Cellular functions and biomedical applications of circular RNAs. Acta Biochimica et Biophysica Sinica. 2024;57(1):157. [
DOI:10.3724/abbs.2024241]
50. Yang L, Yi Y, Mei Z, Huang D, Tang S, Hu L, et al. Circular RNAs in cancer stem cells: Insights into their roles and mechanisms. International Journal of Molecular Medicine. 2025;55(3):50. [
DOI:10.3892/ijmm.2025.5491]
51. Gao L, Fan J, He J, Fan W, Che X, Wang X, et al. Circular RNA as Diagnostic and Prognostic Biomarkers in Hematological Malignancies: Systematic Review. Technology in Cancer Research S Treatment. 2024;23:15330338241285149. https://doi.org/ 10.1177/15330338241285149. [
DOI:10.1177/15330338241285149.]
52. Xiao Y, Qiu M, Tan C, Huang W, Hu S, Jiang X, et al. Systematic analysis of circRNA biomarkers for diagnosis, prognosis and therapy in colorectal cancer. Frontiers in Genetics. 2022;13:938672. [
DOI:10.3389/fgene.2022.938672]
53. Zhang Y, Wang Y, Su X, Wang P, Lin W. The value of circulating circular RNA in cancer diagnosis, monitoring, prognosis, and guiding treatment. Frontiers in oncology. 2021;11:736546. [
DOI:10.3389/fonc.2021.736546.]