TSitologiya i Genetika 2025, vol. 59, no. 5, 70-84
Cytology and Genetics 2025, vol. 59, no. 5, 503–515, doi: https://www.doi.org/10.3103/S009545272505007X

Non-canonical structures in the genome of bovine foamy virus

Limanskaya O.Yu., Balak O.K., Limanskii A.P.

  1. National Scientific Center «Institute of Experimental and Clinical Veterinary Medicine», 83 G. Skovorody St., Kharkiv, 61023, Ukraine
  2. Kharkiv National Medical University, 4 Nauka Ave., Kharkiv, 61022, Ukraine
  3. Institute of Physiological Active Compounds, 58 Nauka Ave., Kharkiv, 61072, Ukraine

SUMMARY. Bioinformatics methods have been used to identify putative perfect G-quadruplexes (G4s) and three-way junctions (3WJs) in bovine foamy virus (BFV) genome. Artificial intelligence (AI) AlphaFold 3 was used to confirm putative G4s and 3WJs by building 3D models of these non-canonical structures. G4s are secondary structures formed by G-rich sequences. Multihelical 3WJs formed by three duplexes connected at the binding point and G4s are considered as alternative structures in DNA and RNA that differ from the classical double-stranded B-DNA. In the present paper, the localization map of putative conservative intramolecular G4s formed by two G-tetrads on the BFV genome was created. 7 putative conservative G-quadruplexes in the sense strand of BFV proviral DNA and 22 G4s in the antisense strand formed by two G-tetrads with G-score from 32 to 36 were found by the multiple alignment of 37 BFV isolates with complete genome. The density of G4s was 0.6 G4/kb for the sense strand of the BFV proviral DNA, while it was 1.8 G4/kb for the antisense strand. One conservative 3WJ motif with length of 73 bp with 100 % homology localized in the 5′-untranslated region and partially on the 5′-end of the gag gene was found for a set of 37 BFV isolates. The 3WJ structure in BFV RNA is stabilized by 20 complementary bp with a free energy ΔG of – 19.8 kcal/mol. The significance of this structure for BFV functioning has been proven. The use of AI AlphaFold 3 to build 3D models of theoretically determined perfect G4s and 3WJs in the BFV genome allowed us to reliably determine putative alternative structures in nucleic acids.

Keywords: alternative structure, bioinformatics, AlphaFold 3, bovine foamy virus, G-quadruplex (G4), three-way junction (3WJ), prediction of non-canonical structures

TSitologiya i Genetika
2025, vol. 59, no. 5, 70-84

Current Issue
Cytology and Genetics
2025, vol. 59, no. 5, 503–515,
doi: 10.3103/S009545272505007X

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References

Abramson, J., Adler, J., Dunger, J., et al., Accurate structure prediction of biomolecular interactions with AlphaFold 3, Nature, 2024, vol. 630, pp. 493–500. https://doi.org/10.1038/s41586-024-07487-w

Agarwal, T., Roy, S., Kumar, S., et al., In the sense of transcription regulation by G-quadruplexes: asymmetric effects in sense and antisense strands, Biochemistry, 2014, vol. 53, pp. 3711–3718. https://doi.org/10.1021/bi401451q

Bao, Q., Hotz-Wagenblatt, A., Betts, M.J., et al., Shared and cell type-specific adaptation strategies of Gag and Env yield high titer bovine foamy virus variants, Infect. Genet. Evol., 2020, vol. 82, p. 104287. https://doi.org/10.1016/j.meegid.2020.104287

Bedrat, A., Lacroix, L., and Mergny, J.L., Re-evaluation of G-quadruplex propensity with G4Hunter, Nucleic Acids Res., 2016, vol. 44, pp. 1746–1759. https://doi.org/10.1093/nar/gkw006

Belotserkovskii, B.P., Neil, A.J., Saleh, S.S., et al., Transcription blockage by homopurine DNA sequences: role of sequence composition and single-strand breaks, Nucleic Acids Res., vol. 201341, pp. 1817–1828. https://doi.org/10.1093/nar/gks1333

Brazda, V., Kolomaznık, J., Lysek, J., et al., G4Hunter web application: a web server for G-quadruplex prediction, Bioinformatics, 2019, vol. 35, pp. 3493–3495. https://doi.org/10.1093/bioinformatics/btz087

De Nicola, B., Lech, C.J., Heddi, B., et al., Structure and possible function of a G-quadruplex in the long terminal repeat of the proviral HIV-1 genome, Nucleic Acids Res., 2016, vol. 44, pp. 6442–6451. https://doi.org/10.1093/nar/gkw432

Evans, L., Kotar, A., Valentini, M., et al., Identification and characterisation of G-quadruplex DNA-forming sequences in the Pseudomonas aeruginosa genome, RSC Chem. Biol., 2023, vol. 4, pp. 94–100. https://doi.org/10.1039/d2cb00205a

Fay, M.M., Lyons, S.M., and Ivanov, P., RNA G-quadruplexes in biology: principles and molecular mechanisms, J. Mol. Biol., 2017, vol. 429, pp. 2127–2147. https://doi.org/10.1016/j.jmb.2017.05.017

Frasson, I., Nadai, M., and Richter, S.N., Conserved G-quadruplexes regulate the immediate early promoters of human alphaherpesviruses, Molecules, vol. 201924, p. 2375. https://doi.org/10.3390/molecules24132375

Guo, P., Erickson, S., and Anderson, D., A small viral RNA is required for in vitro packaging of bacteriophage phi 29 DNA, Science, 1987, vol. 236, pp. 690–694. https://doi.org/10.1126/science.3107124

Hall, T.A., BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT, Nucleic Acids Symp. Ser., 1999, vol. 41, pp. 95–98.

Hamann, M.V. and Lindemann, D., Foamy virus protein-nucleic acid interactions during particle morphogenesis, Viruses, 2016, vol. 8, p. 243. https://doi.org/10.3390/v8090243

Hechler, T., Materniak, M., Kehl, T., et al., Complete genome sequences of two novel European clade bovine foamy viruses from Germany and Poland, J. Virol, 2012, vol. 86, pp. 10905–10906. https://doi.org/10.1128/JVI.01875-12

Heng, X., Herrera, A.P., Song, Z., et al., Retroviral PBS-segment sequence and structure: Orchestrating early and late replication events, Retrovirology, 2024, vol. 21, p. 12. https://doi.org/10.1186/s12977-024-00646-x

Higgins, D., Thompson, J., Gibson, T., et al., CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice, Nucleic Acids Res., 1994, vol. 22, pp. 4673–4680. https://doi.org/10.1093/nar/22.22.4673

Jana, J. and Weisz, K., Thermodynamic stability of G-quadruplexes: impact of sequence and environment, ChemBioChem, 2021, vol. 22, pp. 2848–2856. https://doi.org/10.1002/cbic.202100127

Kardrmas, J.L., Ravin, A.J., and Leontis, N.B., Relative stabilities of DNA three-way, four-way and five-way junctions (multi-helix junction loops): unpaired nucleotides can be stabilizing or destabilizing, Nucleic Acids Res., 1995, vol. 23, pp. 2212–2222. https://doi.org/10.1093/nar/23.12.2212

Kikin, O., D’Antonio, L., and Bagga, P.S., QGRS Mapper: a web-based server for predicting G-quadruplexes in nucleotide sequences, Nucleic Acids Res, 2006, vol. 34, pp. W676–W82. https://doi.org/10.1093/nar/gkl253

Koirala, D., Shao, Y., Koldobskaya, Y., et al., A conserved RNA structural motif for organizing topology within picornaviral internal ribosome entry sites, Nat. Commun., 2019, vol. 10, p. 3629. https://doi.org/10.1038/s41467-019-11585-z

Kumar, S., Stecher, G., Suleski, M., Sanderford, M., Sharma, S., and Tamura, K., MEGA12: Molecular Evolutionary Genetics Analysis Version 12 for adaptive and green computing, Mol. Biol. Evol., 2024, vol. 41, pp. 1–9. https://doi.org/10.1093/molbev/msae263

Lindemann, D., Hütter, S., Wei, G., et al., The unique, the known, and the unknown of spumaretrovirus, Viruses, 2021, vol. 13, p. 105. https://doi.org/10.3390/v13010105

Lombardi, E.P. and Londono-Vallejo, A., A guide to computational methods for G-quadruplex prediction, Nucleic Acids Res., 2020, vol. 48, pp. 1–15. https://doi.org/10.1093/nar/gkz1097

Lu, H., Li, S., Chen, J., et al., Metal ions modulate the conformation and stability of a G-quadruplex with or without a small-molecular ligand, Metallomics, 2015, vol. 7, pp. 1508–1514. https://doi.org/10.1039/c5mt00188a

Mekata, H., Okagawa, T., Konnai, S., et al., Molecular epidemiology and whole-genome analysis of bovine foamy virus in Japan, Viruses 2021, vol. 13, p. 1017. https://doi.org/10.3390/v13061017

Menendez, C., Frees, S., and Bagga, P.S., QGRS-H Predictor: a web server for predicting homologous quadruplex forming G-rich sequence motifs in nucleotide sequences, Nucleic Acids Res., 2012, vol. 40, (Web Server issue), pp. W96–W103. https://doi.org/10.1093/nar/gks422

Monsen, R.C., Trent, J.O., and Chaires, J.B., G-quadruplex DNA: A longer story, Acc. Chem. Res., 2022, vol. 55, pp. 3242–3252. https://doi.org/10.1021/acs.accounts.2c00519

Nicoletto, G., Richter, S.N., and Frasson, I., Presence, location and conservation of putative G-quadruplex forming sequences in arboviruses infecting humans, Int. J. Mol. Sci., 2023, vol. 24, p. 9523. https://doi.org/10.3390/ijms24119523

Ojha, M., Vogt, J., Das, N.K., et al., Structure of saguaro cactus virus 3’ translational enhancer mimics 5’ cap for eIF4E binding, Proc. Natl. Acad. Sci. U. S. A., 2024, vol. 121, p. e2313677121. https://doi.org/10.1073/pnas.2313677121

Ouellet, J., Melcher, S., Iqbal, A., et al., Structure of the three-way helical junction of the hepatitis C virus IRES element, RNA, 2010, vol. 16, pp. 1597–1609. https://doi.org/10.1261/rna.2158410

Piazza, A., Cui, X., Adrian, M., et al., Non-Canonical G-quadruplexes cause the hCEB1 minisatellite instability in Saccharomyces cerevisiae, eLife, 2017, vol. 6, p. e26884. https://doi.org/10.7554/eLife.26884

Piekna-Przybylska, D., Sullivan, M.A., Sharma, G., et al., U3 region in the HIV-1 genome adopts a G-quadruplex structure in its RNA and DNA sequence, Biochemistry, 2014, vol. 53, pp. 2581–2593. https://doi.org/10.1021/bi4016692

Pinto-Santini, D.M., Stenbak, C.R., and Linial, M.L., Foamy virus zoonotic infections, Retrovirology, 2017, vol. 14, p. 55. https://doi.org/10.1186/s12977-017-0379-9

Rethwilm, A. and Bodem, J., Evolution of foamy viruses: the most ancient of all retroviruses, Viruses, 2013, vol. 5, pp. 2349–2374. https://doi.org/10.3390/v5102349

Song, K., Li, B., Li, H., et al., The characterization of G-quadruplexes in Tobacco genome and their function under abiotic stress, Int. J. Mol. Sci., 2024, vol. 25, p. 4331. https://doi.org/10.3390/ijms25084331

Song, Z., Gremminger, T., Singh, G., et al., The three-way junction structure of the HIV-1 PBS-segment binds host enzyme important for viral infectivity, Nucleic Acids Res., 2021, vol. 49, pp. 5925–5942. https://doi.org/10.1093/nar/gkab342

Stefos, G.C., Theodorou, G., and Politis, I., Genomic landscape, polymorphism and possible LINE-associated delivery of G-quadruplex motifs in the bovine genes, Genomics, 2022, vol. 114, p. 110272. https://doi.org/10.1016/j.ygeno.2022.110272

Varshney, D., Spiegel, J., Zyner, K., et al., The regulation and functions of DNA and RNA G-quadruplexes, Nat. Rev. Mol. Cell Biol., 2020, vol. 21, pp. 459–474. https://doi.org/10.1038/s41580-020-0236-x

Wang, S.R., Zhang, Q.Y., Wang, J.Q., et al., Chemical targeting of a G-quadruplex RNA in the Ebola virus L gene, Cell Chem. Biol., 2016, vol. 23, pp. 1113–1122. https://doi.org/10.1016/j.chembiol.2016.07.01

Wang, J., Huang, H., Zhao, K., et al., G-quadruplex in hepatitis B virus pregenomic RNA promotes its translation, J. Biol. Chem., 2023, vol. 299, p. 105151. https://doi.org/10.1016/j.jbc.2023.105151

Whisnant, A.W., Kehl, T., Bao, Q., et al., Identification of novel, highly expressed retroviral microRNAs in cells infected by bovine foamy virus, J. Virol., 2014, vol. 88, pp. 4679–4686. https://doi.org/10.1128/JVI.03587-13

Wolfe, A.L., Singh, K., Zhong, Y., et al., RNA G-quadruplexes cause eIF4A-dependent oncogene translation in cancer, Nature, 2014, vol. 513, pp. 65–70. https://doi.org/10.1038/nature13485

Wu, B., Girard, F., van Buuren, B., et al., Global structure of a DNA three-way junction by solution NMR: towards prediction of 3H fold, Nucleic Acids Res., 2004, vol. 32, pp. 3228–3239. https://doi.org/10.1093/nar/gkh645

Wu, Y.F., Tan, J., Su, Y., et al., Transcription factor AP1 modulates the internal promoter activity of bovine foamy virus, Virus Res., 2010, vol. 147, pp. 139–144. https://doi.org/10.1016/j.virusres.2009.10.005

Xu, X.J. and Chen, S.J., A method to predict the structure and stability of RNA/RNA complexes, Methods Mol. Biol., 2016, vol. 1490, pp. 63–72. https://doi.org/10.1007/978-1-4939-6433-85

Zhang, H., Endrizzi, J.A., Shu, Y., et al., Crystal structure of 3WJ core revealing divalent ion-promoted thermostability and assembly of the Phi29 hexameric motor pRNA, RNA, vol. 19, pp. 1226–1237. https://doi.org/10.1261/rna.037077.112

Zuker, M., Mfold web server for nucleic acid folding and hybridization prediction, Nucleic Acids Res., 2003, vol. 31, pp. 3406–3415. https://doi.org/10.1093/nar/gkg595