Lim, E. J. et al. Systematic review and meta-analysis of the prevalence of chronic fatigue syndrome/myalgic encephalomyelitis (CFS/ME). J. Transl. Med. 18, 100. https://doi.org/10.1186/s12967-020-02269-0 (2020).
Vardaman, M. & Gilmour, S. Letter: Time to correct the record on the global burden of myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS). J. Transl. Med. 23, 331. https://doi.org/10.1186/s12967-025-06281-0 (2025).
Institute of Medicine. Beyond Myalgic Encephalomyelitis/Chronic Fatigue Syndrome: Redefining an Illness (The National Academies Press, 2015). https://doi.org/10.17226/19012.
Lim, E. J. & Son, C. G. Review of case definitions for myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS). J. Transl. Med. 18, 289. https://doi.org/10.1186/s12967-020-02455-0 (2020).
Sweetman, E. et al. Current research provides insight into the biological basis and diagnostic potential for myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS). Diagnostics (Basel) 9, 73. https://doi.org/10.3390/diagnostics9030073 (2019).
Nakatomi, Y. et al. Neuroinflammation in patients with chronic fatigue syndrome/myalgic encephalomyelitis: An 11C-(R)-PK11195 PET study. J. Nucl. Med. 55, 945–950. https://doi.org/10.2967/jnumed.113.131045 (2014).
Hvidberg, M. F., Brinith, L. S., Olesen, A. V., Peersen, K. D. & Ehlers, L. The health-related quality of life for patients with myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS). PLoS ONE 10, e0132421. https://doi.org/10.1371/journal.pone.0132421 (2015).
Walitt, B. et al. Deep phenotyping of post-infectious myalgic encephalomyelitis/chronic fatigue syndrome. Nat. Commun. 15, 907. https://doi.org/10.1038/s41467-024-45107-3 (2024).
Rasa, S. et al. Chronic viral infections in myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS). J. Transl. Med. 16, 268. https://doi.org/10.1186/s12967-018-1644-y (2018).
Hwang, J. H. et al. Evaluation of viral infection as an etiology of ME/CFS: A systematic review and meta-analysis. J. Transl. Med. 21, 763. https://doi.org/10.1186/s12967-023-04635-0 (2023).
Deumer, U.-S. et al. Myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS): An overview. J. Clin. Med. 10, 4786. https://doi.org/10.3390/jcm10204786 (2021).
Lande, A. et al. Human leukocyte antigen alleles associated with myalgic encephalomyelitis/chronic fatigue syndrome (ME/CFS). Sci. Rep. 10, 5267. https://doi.org/10.1038/s41598-020-62157-x (2020).
James, L. M. & Georgopoulos, A. P. At the root of 3 “long” diseases: Persistent antigens inflicting chronic damage on the brain and other organs in Gulf War Illness, Long-COVID-19, and Chronic Fatigue Syndrome. Neurosci. Insights https://doi.org/10.1177/26331055221114817 (2022).
Komaroff, A. L. & Lipkin, W. I. Insights from myalgic encephalomyelitis/chronic fatigue syndrome may help unravel the pathogenesis of postacute COVID-19 syndrome. Trends. Mol. Med. 27, 895–906. https://doi.org/10.1016/j.molmed.2021.06.002 (2021).
Komaroff, A. L. & Lipkin, W. I. ME/CFS and Long COVID share similar symptoms and biological abnormalities: Road map to the literature. Front. Med. (Lausanne) 10, 1187163. https://doi.org/10.3389/fmed.2023.1187163 (2023).
Chen, B. et al. Viral persistence, reactivation, and mechanisms of long COVID. Elife 12, e86015. https://doi.org/10.7554/eLife.86015 (2023).
Swank, Z. et al. Measurement of circulating viral antigens post-SARS-CoV-2 infection in a multicohort study. Clin. Microbiol. Infect. 30, 1599–1605. https://doi.org/10.1016/j.cmi.2024.09.001 (2024).
James, L. M. et al. Anthrax vaccination, Gulf War Illness, and Human Leukocyte Antigen (HLA). Vaccines (Basel). 12, 613. https://doi.org/10.3390/vaccines12060613 (2024).
James, L. M. & Georgopoulos, A. P. Persistent antigens hypothesis: The human leukocyte antigen (HLA) connection. J. Neurol. Neuromed. 3, 27–31. https://doi.org/10.29245/2572.942X/2018/6.1235 (2018).
Bai, N. A. & Richardson, C. S. Posttreatment Lyme disease syndrome and myalgic encephalomyelitis/chronic fatigue syndrome: A systematic review and comparison of pathogenesis. Chronic Dis. Transl. Med. 9, 183–190. https://doi.org/10.1002/cdt3.74 (2023).
Wong, K. H., Shapiro, E. D. & Soffer, G. K. A review of post-treatment lyme disease syndrome and chronic lyme disease for the practicing immunologist. Clin. Rev. Allergy Immunol. 62, 264–271. https://doi.org/10.1007/s12016-021-08906-w (2022).
Jutras, B. L. et al. Borrelia burgdorferi peptidoglycan is a persistent antigen in patients with Lyme arthritis. Proc. Natl. Acad. Sci. U S A. 116, 13498–13507. https://doi.org/10.1073/pnas.1904170116 (2019).
Istrail, S. et al. Comparative immunopeptidomics of humans and their pathogens. Proc. Natl. Acad. Sci. USA 101, 13268–13272. https://doi.org/10.1073/pnas.0404740101 (2004).
Grinde, B. Herpesviruses: Latency and reactivation–viral strategies and host response. J. Oral Microbiol. 5, 22766. https://doi.org/10.3402/jom.v5i0.22766 (2013).
Medhasi, S. & Chantratita, N. Human leukocyte antigen (HLA) system: Genetics and association with bacterial and viral infections. J. Immunol. Res. 2022, 9710376. https://doi.org/10.1155/2022/9710376 (2022).
Trowsdale, J. & Knight, J. C. Major histocompatibility complex genomics and human disease. Annu. Rev. Genomics Hum. Genet. 14, 301–332. https://doi.org/10.1146/annurev-genom-091212-153455 (2013).
Hov, J. R. et al. Electrostatic modifications of the human leukocyte antigen-DR P9 peptide-binding pocket and risk to primary sclerosing cholangitis. Hepatology 53, 1967–1976. https://doi.org/10.1002/hep.24299 (2011).
Dendrou, C. A., Petersen, J., Rossjohn, J. & Fugger, L. HLA variation and disease. Nat. Rev. Immunol. 18, 325–339. https://doi.org/10.1038/nri.2017.143 (2018).
Twomey, R. et al. Chronic fatigue and postexertional malaise in people living with long COVID: An observational study. Phys. Ther. 102, pzac005. https://doi.org/10.1093/ptj/pzac005 (2022).
Jason, L. A. & Dorri, J. A. ME/CFS and post-exertional malaise among patients with long COVID. Neurol. Int. 15, 1–11. https://doi.org/10.3390/neurolint15010001 (2022).
Vernon, S. D. et al. Incidence and prevalence of Post-COVID-19 myalgic encephalomyelitis: A report from the observational RECOVER-Adult Study. J. Gen. Intern. Med. 40, 1085–1094. https://doi.org/10.1007/s11606-024-09290-9 (2025).
Augusto, D. G. & Hollenbach, J. A. HLA variation and antigen presentation in COVID-19 and SARS-CoV-2 infection. Curr. Opin. Immunol. 76, 102178. https://doi.org/10.1016/j.coi.2022.102178 (2022).
Augusto, D. G. et al. A common allele of HLA is associated with asymptomatic SARS-CoV-2 infection. Nature 620, 128–136. https://doi.org/10.1038/s41586-023-06331-x (2023).
Douillard, V. et al. Current HLA investigations on SARS-CoV-2 and perspectives. Front. Genet. 12, 774922. https://doi.org/10.3389/fgene.2021.774922 (2021).
Wolday, D. et al. HLA variation and SARS-CoV-2 specific antibody response. Viruses 15, 906. https://doi.org/10.3390/v15040906 (2023).
Lochhead, R. B., Strle, K., Arkivar, S. L., Weis, J. J. & Steere, A. C. Lyme arthritis: Linking infection, inflammation and autoimmunity. Nat. Rev. Rheumatol. 17, 449–461. https://doi.org/10.1038/s41584-021-00648-5 (2021).
Georgopoulos, A. P., James, L. M. & Peterson, P. K. Human Leukocyte Antigen (HLA) at the root of persistent antigens and long COVID. J. Immunol. Sci. 9, 1–3. https://doi.org/10.29245/2578-3009/2025/1.1257 (2025).
Pons-Fuster, E. et al. HLA-A* 03 may confer protection against long COVID through an enhanced immune response. Infect. Dis. Now 55, 105057. https://doi.org/10.1016/j.idnow.2025.105057 (2025).
Barquera, R. et al. Binding affinities of 438 HLA proteins to complete proteomes of seven pandemic viruses and distributions of strongest and weakest HLA peptide binders in populations worldwide. HLA. 96, 277–298. https://doi.org/10.1111/tan.13956 (2020).
Arrieta-Bolaños, E., Hernández-Zaragoza, D. I. & Barquera, R. An HLA map of the world: A comparison of HLA frequencies in 200 worldwide populations reveals diverse patterns for class I and class II. Front. Genet. 14, 866407. https://doi.org/10.3389/fgene.2023.866407 (2023).
Buhler, S. & Sanchez-Mazas, A. HLA DNA sequence variation among human populations: Molecular signatures of demographic and selective events. PLoS ONE 6, e14643. https://doi.org/10.1371/journal.pone.0014643 (2011).
Uniprot. Accessed on April 11, 2025. https://www.uniprot.org/uniprotkb
Reynisson, B., Alvarez, B., Paul, S., Peters, B. & Nielsen, M. NetMHCpan-41 and NetMHCIIpan-4.0: Improved predictions of MHC antigen presentation by concurrent motif deconvolution and integration of MS MHC eluted ligand data. Nucleic Acids Res. 48, W449–W454. https://doi.org/10.1093/nar/gkaa379 (2020).
IEDB Analysis Resource. Accessed on April 11, 2025. http://tools.iedb.org/mhci/
Charonis, S., James, L. M. & Georgopoulos, A. P. In silico assessment of binding affinities of three dementia-protective Human Leukocyte Antigen (HLA) alleles to nine human herpes virus antigens. Curr. Res. Transl. Med. 68, 211–216. https://doi.org/10.1016/j.retram.2020.06.002 (2020).
Charonis, S., Tsilibary, E. P. & Georgopoulos, A. SARS-CoV-2 virus and Human Leukocyte Antigen (HLA) Class II: Investigation in silico of binding affinities for COVID-19 protection and vaccine development. J. Immunol. Sci. 4, 12–23. https://doi.org/10.29245/2578-3009/2020/4.1198 (2020).
Charonis, S. A., Tsilibary, E. P. & Georgopoulos, A. P. In silico investigation of binding affinities between human leukocyte antigen class I molecules and SARS-CoV-2 virus spike and ORF1ab proteins. Explor. Immunol. 1, 16–26. https://doi.org/10.37349/ei.2021.00003 (2021).