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The Structure and Function of DNA G-Quadruplexes
Open accessAbstract
Guanine-rich DNA sequences can fold into four-stranded, noncanonical secondary structures called G-quadruplexes (G4s). G4s were initially considered a structural curiosity, but recent evidence suggests their involvement in key genome functions such as transcription, replication, genome stability, and epigenetic regulation, together with numerous connections to cancer biology. Collectively, these advances have stimulated research probing G4 mechanisms and consequent opportunities for therapeutic intervention. Here, we provide a perspective on the structure and function of G4s with an emphasis on key molecules and methodological advances that enable the study of G4 structures in human cells. We also critically examine recent mechanistic insights into G4 biology and protein interaction partners and highlight opportunities for drug discovery.
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“Lastly, more serious DNA damages, such as double-strand breaks associated with non-B DNA structures like G4s, and the subsequent DNA repair can be mutagenic (7,27).”
From G-quadruplex-mediated genomic instability drives SNVs in cancer (2024) · cites this paper as [7]
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1 retracted
Zhang, S., Sun, H., Wang, L., Liu, Y., Chen, H., Li, Q., Guan, A., Liu, M., & Tang, Y. (2018) · Nucleic Acids Research · doi:10.1093/nar/gky665 · Cited by 158
RetractedHallmarks of Cancer: The Next Generation
Hanahan, D., & Weinberg, R. A. (2011) · Cell · doi:10.1016/j.cell.2011.02.013 · Cited by 66,590 · cited 2×
“achieve unlimited proliferation by protecting ends of chromosomes [23].”
BackgroundMolecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid
WATSON, J. D., & CRICK, F. H. C. (1953) · Nature · doi:10.1038/171737a0 · Cited by 13,584 · cited 1×
“The most widely recognised DNA structure is that of the classical DNA double helix [1], which defines a structural basis for the genetic code via defined base-pairing.”
BackgroundQuadruplex DNA: sequence, topology and structure
Burge, S., Parkinson, G. N., Hazel, P., Todd, A. K., & Neidle, S. (2006) · Nucleic Acids Research · doi:10.1093/nar/gkl655 · Cited by 2,404 · cited 1×
“ogies arising from different combinations of strand direction (Figure 1D–F), as well as length and loop composition [9,10].”
BackgroundSiddiqui-Jain, A., Grand, C. L., Bearss, D. J., & Hurley, L. H. (2002) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.182256799 · Cited by 2,274 · cited 9×
“A cocrystal structure of bos taurus DHX36 helicase bound to the MYC promoter G4 revealed a mode of binding in which the G4 makes contacts with a DHX36-specificmotif (DSM) and the C terminal OB-fold domain (Figure 4) [73].”
BackgroundInfluentialQuantitative visualization of DNA G-quadruplex structures in human cells
Biffi, G., Tannahill, D., McCafferty, J., & Balasubramanian, S. (2013) · Nature Chemistry · doi:10.1038/nchem.1548 · Cited by 2,173 · cited 3×
“structures in human cell lines upon ligand treatment together with depletion of the G4 resolving helicase FANCJ [41,42], to reveal increased numbers of G4 foci staining in nuclei after pyridostatin [41] or telomestatin ligand treatment and FANCJ depletion in chicken DT40 cells [42].”
BackgroundSelf-citationSen, D., & Gilbert, W. (1988) · Nature · doi:10.1038/334364a0 · Cited by 1,866 · cited 1×
“The finding that G4-rich telomeric repeats form G4 structures [6] suggested a mechanistic link with telomerase-mediated extension of telomeres, prompting an exploration of G4 stabilising ligands that may inhibit the growth of cancer cells by interfering with telomere maintenance (reviewed in [89]).”
BackgroundCrystal structure of parallel quadruplexes from human telomeric DNA
Parkinson, G., Lee, M. P. H., & Neidle, S. (2002) · Nature · doi:10.1038/nature755 · Cited by 1,863 · cited 1×
“Structural studies using X-ray crystallography and NMR spectroscopy have provided detailed insights into the structure of DNA G4s primarily based on the human telomeric repeat (Figure 1B,C) [11] or sequences derived from the promoter regions of certain human genes such as MYC or KIT [12,13].”
BackgroundPrevalence of quadruplexes in the human genome
Huppert, J. L., & Balasubramanian, S. (2005) · Nucleic Acids Research · doi:10.1093/nar/gki609 · Cited by 1,795 · cited 2×
“Based on biophysical studies on different G4 structures, algorithms using sequence motifs such as GR3NxGR3NxGR3NxGR3 were developed and deployed to predict putative G4 structures in genomic DNA [14,15].”
BackgroundMethodSelf-citationHelix formation by guanylic acid.
Gellert, M., Lipsett, M., & Davies, D. R. (1962) · Proceedings of the National Academy of Sciences of the United States of America · doi:10.1073/pnas.48.12.2013 · Cited by 1,430
Flagged first, then by citations
4 of the 121 references couldn’t be matched to a known paper.
Research works printed but not linked: 4
- Bang, 1910, Untersuchungen über die Guanylsäure, Biochem. Z., 26, 293
- Testorelli, 2003, Telomerase and cancer, J. Exp. Clin. Cancer Res., 22, 165
- Anders, 2014, Genome-wide localization of small molecules, Nat. Biotechnol., 32, 92, 10.1038/nbt.2776 · DOI
- Pagano, 2015, Identification of novel interactors of human telomeric G-quadruplex DNA, Chem. Commun., 51, 2964, 10.1039/C4CC07231F · DOI
Cited by
Sorted by citations
Items 201–300 of 706
Dynamics Studies of DNA with Non-canonical Structure Using NMR Spectroscopy
Oh, K., Kim, J., Park, C., & Lee, J. (2020) · International Journal of Molecular Sciences · doi:10.3390/ijms21082673 · Cited by 19
Identification of G-quadruplex nucleic acid structures by high-throughput sequencing: A review
Song, D., Luo, J., Duan, X., Jin, F., & Lu, Y. (2025) · International Journal of Biological Macromolecules · doi:10.1016/j.ijbiomac.2025.139896 · Cited by 17
Andreeva, D. V., Vedekhina, T., Gostev, A. S., Dezhenkova, L. G., Volodina, Y. L., Markova, A. A., Nguyen, M. T., Ivanova, O. M., Dolgusheva, V. A., Varizhuk, A. M., et al. (2024) · European Journal of Medicinal Chemistry · doi:10.1016/j.ejmech.2024.116222 · Cited by 17
Dynamics of G-Quadruplex Formation under Molecular Crowding
Aznauryan, M. G., & Birkedal, V. (2023) · The Journal of Physical Chemistry Letters · doi:10.1021/acs.jpclett.3c02453 · Cited by 13
Papageorgiou, A. C., Pospíšilová, M., Cibulka, J., Ashraf, R., Waudby, C. A., Kadeřávek, P., Maroz, V., Kubíček, K., Prokop, Z., Krejčí, L., et al. (2023) · Nature Communications · doi:10.1038/s41467-023-42503-z · Cited by 16
Soenarjo, A. L., Lan, Z., Sazanovich, I., Chan, Y.-S., Ringholm, M., Jha, A., & Klug, D. (2023) · Journal of the American Chemical Society · doi:10.1021/jacs.3c04044 · Cited by 14
Saha, P., Panda, D., & Dash, J. (2023) · Chemical Society Reviews · doi:10.1039/d3cs00166k · Cited by 38
Higher-order G-quadruplexes in promoters are untapped drug targets
Monsen, R. C. (2023) · Frontiers in Chemistry · doi:10.3389/fchem.2023.1211512 · Cited by 16
“Historically, the largest loop length has been taken to be seven nucleotides because of the destabilizing effect of large loops in vitro (Spiegel et al., 2020; Ravichandran et al., 2021).”
BackgroundInfluentialProteolysis-targeting chimeras in biotherapeutics: Current trends and future applications
Li, Q., Zhou, L., Qin, S., Huang, Z., Li, B., Li, B., Yang, M., Nice, E. C., Zhu, H., & Huang, C. (2023) · European Journal of Medicinal Chemistry · doi:10.1016/j.ejmech.2023.115447 · Cited by 17
The Intertwined Role of 8-oxodG and G4 in Transcription Regulation
Gorini, F., Ambrosio, S., Lania, L., Majello, B., & Amente, S. (2023) · International Journal of Molecular Sciences · doi:10.3390/ijms24032031 · Cited by 16
Teloxantron inhibits the processivity of telomerase with preferential DNA damage on telomeres
Maciejewska, N., Olszewski, M., Jurasz, J., Bagiński, M., Stasevych, M., Zvarych, V. I., Folini, M., & Zaffaroni, N. (2022) · Cell Death and Disease · doi:10.1038/s41419-022-05443-y · Cited by 14
“Most of the telomerase-targeting compounds are nonselective because G4 structures also occur in additional G-rich DNA sequences other than telomeres, which may cause high toxicity to normal cells [29].”
BackgroundHuang, Q., Wang, X., Cheng, A., Zhang, H., Yu, Q.-M., Shen, C.-F., Awadasseid, A., Zhao, X., Xiong, X., Wu, Y.-L., et al. (2022) · Biochemical Pharmacology · doi:10.1016/j.bcp.2022.115062 · Cited by 14
Shitikov, E. A., Bespiatykh, D., Malakhova, M. V., Bespyatykh, J., Bodoev, I. N., Vedekhina, T., Zaychikova, M. V., Veselovsky, V. A., Klimina, K. M., Ilina, E. N., et al. (2022) · Frontiers in Microbiology · doi:10.3389/fmicb.2022.817024 · Cited by 18
Kent, T., & Clynes, D. (2021) · Genes · doi:10.3390/genes12111734 · Cited by 16
“CFSs also characteristically have a paucity of replication origins and are often late replicating, limiting the opportunity for stalled replication forks to be rescued by converging forks (for extensive review on CFSs, see Reference [9]).”
BackgroundNovel implications of a strictly monomorphic (GCC) repeat in the human PRKACB gene
Khamse, S., Jafarian, Z., Bozorgmehr, A., Tavakoli, M., Afshar, H., Keshavarz, M., Moayedi, R., & Ohadi, M. (2021) · Scientific Reports · doi:10.1038/s41598-021-99932-3 · Cited by 22
Shin, C.-R., Choi, E., Kim, G., Baek, S. Y., Park, B., Hwang, J., Jun, J., Kil, H.-J., Oh, H., Lee, K., et al. (2021) · Scientific Reports · doi:10.1038/s41598-021-91675-5 · Cited by 14
“HK Chillyang-myeon, Gangjin-gun, Jeonnam, South Korea 11 √ √ (10)a This study”
BackgroundHossain, K. A., Jurkowski, M., Czub, J., & Kogut, M. (2021) · Computational and Structural Biotechnology Journal · doi:10.1016/j.csbj.2021.04.039 · Cited by 13
Amjadi‐Moheb, F., Paniri, A., & Akhavan‐Niaki, H. (2021) · Cancer Research · doi:10.1158/0008-5472.can-20-3613 · Cited by 16
Gudanis, D., Kaniowski, D., Kulik, K., Baranowski, D., Gdaniec, Z., & Nawrot, B. (2020) · Cells · doi:10.3390/cells9112375 · Cited by 14
Street, S. T. G., Peñalver, P., O’Hagan, M., Hollingworth, G. J., Morales, J. C., & Galán, M. C. (2021) · Chemistry - A European Journal · doi:10.1002/chem.202100040 · Cited by 18
DNA G-quadruplexes: Structural and functional insights
Jonchhe, S., Lahiri, S., & Rothenberg, E. (2025) · DNA repair · doi:10.1016/j.dnarep.2025.103910 · Cited by 13
Sousa, V., Ladeira, B., Garanger, É., Lecommandoux, S., Meijer, E. W., Dankers, P. Y. W., Mano, J. F., & Borges, J. (2025) · ACS Nano · doi:10.1021/acsnano.5c10088 · Cited by 12
Sarma, S., Thakur, N., Varshney, N., Jha, H., & Sarma, T. K. (2024) · Communications Chemistry · doi:10.1038/s42004-024-01353-6 · Cited by 13
“Although bothDNAandGMParehighlywatersoluble,theG-quadruplexcanactasa hydrotropicsystembytrappingwaterinsolublemoleculesinthesmall cavitiesformedbyverticallystackedguaninetetrads 52 – 54 (Fig.”
Merlino, F., Marzano, S., Zizza, P., D’Aria, F., Grasso, N., Carachino, A., Iachettini, S., Biroccio, A., Fonzo, S. D., Grieco, P., et al. (2024) · Nucleic Acids Research · doi:10.1093/nar/gkae471 · Cited by 19
“The compelling evidence that G4 structures act, in concert with certain proteins ( 6–9 ), as active regulators of cancer-related genomic processes has made them an emerging research topic and an attractive target for gene regulation and anticancer drug de-sign ( 1 , 3 , 10–13 ).”
BackgroundG4Bank: A database of experimentally identified DNA G-quadruplex sequences
Zhong, H., Dong, M., & Gao, F. (2023) · Interdisciplinary Sciences Computational Life Sciences · doi:10.1007/s12539-023-00577-9 · Cited by 13
Yella, V. R., & Vanaja, A. (2023) · Biochimie · doi:10.1016/j.biochi.2023.06.002 · Cited by 13
Jenner, L. P., Peška, V., Fulnečková, J., & Sýkorová, E. (2022) · Genes · doi:10.3390/genes13091663 · Cited by 18
Liu, L., Zhu, L., Tong, H., Su, C., Wells, J. W., & Chalikian, T. V. (2022) · The Journal of Physical Chemistry B · doi:10.1021/acs.jpcb.2c04304 · Cited by 15
Singh, M., Gupta, R., Comez, L., Paciaroni, A., Rani, R., & Kumar, V. (2022) · Drug Discovery Today · doi:10.1016/j.drudis.2022.06.002 · Cited by 13
Ligands as Stabilizers of G-Quadruplexes in Non-Coding RNAs
Figueiredo, J., Santos, T., Miranda, A., Alexandre, D., Teixeira, B., Simões, P., Lopes-Nunes, J., & Cruz, C. (2021) · Molecules · doi:10.3390/molecules26206164 · Cited by 13
“rG4s are thermodynamically stable secondary structures in which the guanines, which are linked between themselves via Hoogsteen hydrogen bonds, engage the N1, N7, O6, and N2 atoms from each base (Figure 1) [6,7].”
BackgroundCarlsson, E. R., Fenger, M., Henriksen, T., Kjær, L. K., Worm, D., Hansen, D. L., Madsbad, S., & Poulsen, H. E. (2020) · PLoS ONE · doi:10.1371/journal.pone.0243918 · Cited by 16
“Guanine oxidation is known to destabilize the guanine-quadruplexes (so called G4’s) established in the single-strand stretches of promoters (thereby affecting gene transcription) and telomeres [9].”
BackgroundAhmed, A. A., & Neidle, S. (2020) · Molecules · doi:10.3390/molecules25225407 · Cited by 14
Deng, Y., Hu, J., Wang, J., Kuai, Z., Hao, F., He, Y., Liang, M., Song, A., Yin, C., Yi, H., et al. (2026) · ACS Applied Materials & Interfaces · doi:10.1021/acsami.5c20923 · Cited by 6
DNA 5mC methylation inhibits the formation of G-quadruplex structures in the genome
Niu, K., Xiang, L., Zhang, X., Li, X., Yao, T., Li, J., Zhang, C., Liu, J., Peng, Y., Xu, G., et al. (2025) · Genome biology · doi:10.1186/s13059-025-03678-4 · Cited by 13
Yang, J., Qin, G., Huang, B.-Y., Song, H.-L., Sun, J.-W., Postings, M. L., Scott, P., Zhao, C.-Q., Wang, C.-Y., Tan, W.-J., et al. (2024) · National Science Review · doi:10.1093/nsr/nwae388 · Cited by 12
“…modes are known or have been postulated [22 ], typical synthetic G4 ligands contain flat aromatic chromophores for π – π stacking with G-quartets, perhaps with side chains for interaction with loops and grooves of the G4s, and steric bulk that prevents intercalation between base pairs [23 ].”
Dey, A., Pandav, K., Nath, M., Barthwal, R., & Prasad, R. (2022) · Molecular Therapy — Nucleic Acids · doi:10.1016/j.omtn.2022.11.008 · Cited by 15
“The G4 structure in the telomeric region prevents the access of telomerase enzyme from causing apoptosis and consequent cell death.(2) Depending on the length of the G rich sequences, loop length/size, and glycosidic bond in the presence of metal ions, G-quadruplex DNA structures take on several topologies, such as an…”
BackgroundLópez‐Tena, M., Chen, S., & Winssinger, N. (2022) · Bioconjugate Chemistry · doi:10.1021/acs.bioconjchem.2c00292 · Cited by 14
Lee, J., & Ha, J. (2022) · Analytical Chemistry · doi:10.1021/acs.analchem.2c01000 · Cited by 12
Ruggiero, E., Frasson, I., Tosoni, E., Scalabrin, M., Perrone, R., Marušič, M., Plavec, J., & Richter, S. N. (2022) · ACS Infectious Diseases · doi:10.1021/acsinfecdis.1c00508 · Cited by 21
Methodological advances of bioanalysis and biochemical targeting of intracellular G‐quadruplexes
Teng, X., Dai, Y., & Li, J. (2022) · Exploration · doi:10.1002/exp.20210214 · Cited by 19
Yan, Y.-R., Zhao, D., Zhang, Q., Chang, Y.-Y., & Liu, M. (2022) · Journal of Analysis and Testing · doi:10.1007/s41664-021-00206-8 · Cited by 12
Brázda, V., Bohálová, N., & Bowater, R. (2021) · Gene · doi:10.1016/j.gene.2021.146058 · Cited by 12
Hache, F., & Changenet, P. (2021) · Chirality · doi:10.1002/chir.23359 · Cited by 12
Volumetric Properties of Four-Stranded DNA Structures
Chalikian, T., & Macgregor, R. (2021) · Biology · doi:10.3390/biology10080813 · Cited by 12
Soundarapandian, S., Alexander, A., Pillai, A. S., Enoch, I. V., & Yousuf, S. (2021) · Journal of Biomolecular Structure and Dynamics · doi:10.1080/07391102.2021.1911849 · Cited by 12
“Furthermore, G4 DNAs are detectable in cancer cell states compared with DNAs in their normal form, rendering the former as highly interesting targets in drug discovery (Jochen et al., 2020).”
BackgroundMiclot, T., Corbier, C., Terenzi, A., Hognon, C., Grandemange, S., Barone, G., & Monari, A. (2020) · bioRxiv · doi:10.1101/2020.11.26.399741 · Cited by 12
Mgs1 function at G-quadruplex structures during DNA replication
Paeschke, K., & Burkovics, P. (2020) · Current Genetics · doi:10.1007/s00294-020-01128-1 · Cited by 11
Piccolo, M., Russo, C., Arciuolo, V., Ferraro, M. G., Abbate, V., Porzio, A. D., Cinquegrana, E., Leva, F. S. D., Pagano, B., Randazzo, A., et al. (2025) · Journal of Medicinal Chemistry · doi:10.1021/acs.jmedchem.4c01665 · Cited by 12
A Compendium of G-Flipon Biological Functions That Have Experimental Validation
Herbert, A. G. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms251910299 · Cited by 13
“This finding is true for the binding of the SP1 transcription factor to its B-DNA cognate sequence and a c-MYC parallel GQ [107] and many other proteins that bind both to GQs and to a B-DNA motif [21].”
BackgroundRecent Developments in Mitochondrial G‐Quadruplex Recognising Fluorescent Probes: A Review
Kumar, P., Pandith, A., Tseng, C., & Burnouf, T. (2023) · Journal of Photochemistry and Photobiology C Photochemistry Reviews · doi:10.1016/j.jphotochemrev.2023.100619 · Cited by 14
Research progress of small-molecule drugs in targeting telomerase in human cancer and aging
Shen, Z., Wang, Y., Wang, G., Gu, W. L., Zhao, S., Hu, X., Liu, W., Cai, Y., Ma, Z., Gautam, R. K., et al. (2023) · Chemico-Biological Interactions · doi:10.1016/j.cbi.2023.110631 · Cited by 12
The effect of side chain variations on quinazoline-pyrimidine G-quadruplex DNA ligands.
Bhuma, N., Chand, K., Andréasson, M., Mason, J., Das, R., Patel, A. K., Öhlund, D., & Chorell, E. (2023) · European journal of medicinal chemistry · doi:10.1016/j.ejmech.2023.115103 · Cited by 11
“It has been shown that G4 DNA can play vital roles in central biological processes such as control of replication, [5,6] transcription, [7,8] genome stability, [9,10] and epigenetic regulation [11 – 13].”
Clovis, N. S., Alam, P., Chand, A. K., Sardana, D., Khan, M. F., & Sen, S. (2022) · Journal of Photochemistry and Photobiology A Chemistry · doi:10.1016/j.jphotochem.2022.114432 · Cited by 15
Frasson, I., Soldà, P., Nadai, M., Tassinari, M., Scalabrin, M., Gokhale, V. M., Hurley, L. H., & Richter, S. N. (2022) · Antiviral Research · doi:10.1016/j.antiviral.2022.105432 · Cited by 13
“Guanine quadruplexes (G4s) are non-canonical nucleic acids structures, which are being been widely investigated for their unique structural and functional features (Spiegel et al., 2020).”
BackgroundG-quadruplexes Mark Sites of Methylation Instability Associated with Ageing and Cancer
Rauchhaus, J., Robinson, J., Monti, L., & Antonio, M. D. (2022) · Genes · doi:10.3390/genes13091665 · Cited by 23
“The most well studied of these structures is the G-quadruplex (G4)—a folded state of DNA that arises when guanine(G)-rich sequences assemble into G-quartets—stabilised by Hoogsteen hydrogen bonds and interactions with metal cations such as K+ (Figure 1A) [4].”
BackgroundPlatella, C., Napolitano, E., Riccardi, C., Musumeci, D., & Montesarchio, D. (2022) · ChemistryOpen · doi:10.1002/open.202200090 · Cited by 11
Role and Regulation of Pif1 Family Helicases at the Replication Fork
Malone, E. G., Thompson, M. D., & Byrd, A. K. (2022) · International Journal of Molecular Sciences · doi:10.3390/ijms23073736 · Cited by 14
“G-quadruplexes (G4) are secondary structures in DNA that form from intra- or intermolecular guanosine hydrogen bonding, forming highly stable planar tetrads stabilized by Hoogsteen hydrogen bonds [49].”
BackgroundOB-Folds and Genome Maintenance: Targeting Protein–DNA Interactions for Cancer Therapy
Par, S., Vaides, S., VanderVere-Carozza, P. S., Pawelczak, K. S., Stewart, J. A., & Turchi, J. J. (2021) · Cancers · doi:10.3390/cancers13133346 · Cited by 11
“G4s consist of four repeats of at least three guanosines to form a four stranded structure through Hoogsten base pairing [43].”
BackgroundPhenoxazine-based scaffold for designing G4-interacting agents
Цветков, В. Б., Varizhuk, A. M., Lizunova, S. A., Nikolenko, T. A., Ivanov, I. A., Severov, V. V., Belyaev, E. S., Shitikov, E. A., Pozmogova, G. E., & Aralov, A. V. (2020) · Organic & Biomolecular Chemistry · doi:10.1039/d0ob00983k · Cited by 13
Modulation of histone modifications and G-quadruplex structures by G-quadruplex-binding proteins.
Oyoshi, T., & Masuzawa, T. (2020) · Biochemical and Biophysical Research Communications - BBRC · doi:10.1016/j.bbrc.2020.02.178 · Cited by 11
Quadruplexes with a grain of salt: influence of cation type and concentration on DNA G4 stability
Cucchiarini, A., Kledus, F., Luo, Y., Brázda, V., & Mergny, J. (2025) · European Biophysics Journal · doi:10.1007/s00249-025-01772-w · Cited by 10
“G-quadruplexes or G4s are guanine-rich nucleic acids composed of four strands (DNA or RNA) and stabilized by non-covalent interactions: intra-quartet hydrogen bonds, π–π stacking between aromatic bases, and cation coordination (Spiegel et al. 2020).”
DNA G-quadruplex structures act as functional elements in α- and β-globin enhancers
Doyle, C., Herka, K., Flynn, S. M., Melidis, L., Dhir, S., Schoenfelder, S., Tannahill, D., & Balasubramanian, S. (2025) · Genome biology · doi:10.1186/s13059-025-03627-1 · Cited by 10
“G4s form in guanine-rich regions through guanine-guanine Hoogsteen base pairing to form G-tetrads, which assemble into stacks stabilized by cations to form a folded G4 structure [7] (Fig.”
Self-citationHerbert, A. G., Cherednichenko, O., Lybrand, T. P., Egli, M., & Poptsova, M. S. (2025) · International Journal of Molecular Sciences · doi:10.3390/ijms26062422 · Cited by 12
The iMab antibody selectively binds to intramolecular and intermolecular i-motif structures
Ruggiero, E., Marušič, M., Zanin, I., Martinez, C. D. P., Christ, D., Plavec, J., & Richter, S. N. (2024) · bioRxiv · doi:10.1093/nar/gkae1305 · Cited by 10
“G4s occur within guanine (G)-rich regions when two or more G-tetrads, which are associations of four Gs, self-stack (2).”
Geng, Y., Liu, C., Xu, N., Suen, M. C., Miao, H., Xie, Y., Zhang, B., Chen, X., Song, Y., Wang, Z., et al. (2024) · Nucleic Acids Research · doi:10.1093/nar/gkae473 · Cited by 12
“G-quadruplexes are secondary structures formed by the stacking of two or more G-tetrads in guanine-rich regions of DNA or RNA ( 14 ).”
BackgroundSynthetic Peptides: Promising Modalities for the Targeting of Disease‐Related Nucleic Acids
Ellenbroek, B. D., Kahler, J. P., Evers, S. R., & Pomplun, S. J. (2024) · Angewandte Chemie International Edition · doi:10.1002/anie.202401704 · Cited by 15
Bisoi, A., Sarkar, S., & Singh, P. C. (2024) · International Journal of Biological Macromolecules · doi:10.1016/j.ijbiomac.2024.131050 · Cited by 10
The intricate relationship of G-Quadruplexes and bacterial pathogenicity islands
Lyu, B., & Song, Q.-S. (2024) · bioRxiv · doi:10.7554/elife.91985 · Cited by 10
Andréasson, M., Donzel, M., Abrahamsson, A., Berner, A., Doimo, M., Quiroga, A., Eriksson, A., Chao, Y.-K., Overman, J., Pemberton, N., et al. (2024) · Journal of Medicinal Chemistry · doi:10.1021/acs.jmedchem.3c02127 · Cited by 10
Construction of DNA aggregates in cell milieu for bio‐interference
Guo, Y.-F., Liu, J., Huang, Y., Guo, X.-C., & Yang, D.-Y. (2023) · Aggregate · doi:10.1002/agt2.389 · Cited by 10
Kondo, J., & Nakamura, S. (2023) · Journal of Chemical Education · doi:10.1021/acs.jchemed.2c00964 · Cited by 10
Mass Spectrometry for Assessing Protein–Nucleic Acid Interactions
He, X., Chen, X., & Wang, Y. (2023) · Analytical Chemistry · doi:10.1021/acs.analchem.2c04353 · Cited by 12
Zheng, Y., Zhao, J., Zhang, L., & Wang, W.-Z. (2023) · Sensors and actuators. B, Chemical · doi:10.1016/j.snb.2023.133418 · Cited by 10
Fang, S., Liu, S., Yang, D., Yang, L., Hu, C., & Wan, J. (2022) · Frontiers in Genetics · doi:10.3389/fgene.2022.957023 · Cited by 12
“G-quadruplexes, which are also called G4 structures, are non-canonical secondary organizations formed by folding of guanine-rich DNA or RNA sequences (Spiegel et al., 2020).”
BackgroundVirgilio, A., Pecoraro, A., Benigno, D., Russo, A., Russo, G., Esposito, V., & Galeone, A. (2022) · International Journal of Molecular Sciences · doi:10.3390/ijms23115952 · Cited by 10
“Even though the significance of G4 structures in living cells was the subject of controversy in the past, the collection of more recent experimental data corroborates the presence and relevance of these conformations in living cells [13,14].”
BackgroundCan aggressive cancers be identified by the “aggressiveness” of their chromatin?
Gurova, K. V. (2022) · BioEssays · doi:10.1002/bies.202100212 · Cited by 9
“moters that can form G-quadruplex structures when not covered by nucleosomes.[75] Other barriers are the transcription termination sites (TTSs) of actively transcribed genes and RNAPII at the TTS.”
BackgroundNucleases and Co-Factors in DNA Replication Stress Responses
Nickoloff, J. A., Sharma, N., Taylor, L. E., Allen, S. J., & Hromas, R. (2022) · DNA · doi:10.3390/dna2010006 · Cited by 9
Yadav, K., Sardana, D., Shweta, H., Clovis, N. S., & Sen, S. (2022) · Journal of Physical Chemistry B · doi:10.1021/acs.jpcb.1c09349 · Cited by 10
Valle-Orero, J., Rieu, M., Tran, P. L. T., Joubert, A., Raj, S., Allemand, J., Croquette, V., & Boulé, J. (2022) · bioRxiv · doi:10.1093/nar/gkac667 · Cited by 10
Probing GFP Chromophore Analogs as Anti-HIV Agents Targeting LTR-III G-Quadruplex
Ryazantsev, D., Myshkin, M., Alferova, V., Tsvetkov, V., Shustova, E., Kamzeeva, P., Kovalets, P. V., Zaitseva, E., Baleeva, N., Zatsepin, T., et al. (2021) · Biomolecules · doi:10.3390/biom11101409 · Cited by 10
Климанова, Е. А., Sidorenko, S. V., Abramicheva, P. A., Tverskoi, A. M., Orlov, S. N., & Lopina, O. D. (2020) · International Journal of Molecular Sciences · doi:10.3390/ijms21217992 · Cited by 9
“In addition, experimental data show that G-quadruplexes are more frequent in cancer cells in comparison with normal cells [49].”
BackgroundPrice, D. A., Kartje, Z. J., Hughes, J. A., Hill, T. D., Loth, T. M., Watts, J., Gagnon, K. T., & Moran, S. D. (2020) · ChemBioChem · doi:10.1002/cbic.202000136 · Cited by 10
Bartosik, A. R., Hou, P., Tang, S., Vaughn, J. P., Smaldino, P. J., Ratan, A., Mayo, M. W., & Wang, Y. (2025) · Nucleic Acids Research · doi:10.1093/nar/gkaf621 · Cited by 13
G-quadruplex and i-motif DNA structures form in the promoter of the key innate immune adaptor MYD88
Brown, S. L., Swafford, K., McCrury, M., Nasrin, F., Gragg, C. Q., Chavan, A., Choudhury, S. R., Dickerhoff, J., Yang, D., & Kendrick, S. L. (2025) · Cell Reports Physical Science · doi:10.1016/j.xcrp.2025.102560 · Cited by 8
“Similar to other G4/iM-forming sequences, 17,27 Py31WT sits in one of two CpG islands in the MYD88 promoter (Figure 4A).”
G-quadruplex structures in 16S rRNA regions correlate with thermal adaptation in prokaryotes
Lyu, B., Niu, K.-K., Anderson, D., Feng, Q.-L., & Song, Q.-S. (2025) · Nucleic Acids Research · doi:10.1093/nar/gkaf042 · Cited by 9
Benassi, A., Peñalver, P., Pérez-Soto, M., Pirota, V., Freccero, M., Morales, J., & Doria, F. (2024) · Journal of Medicinal Chemistry · doi:10.1021/acs.jmedchem.4c00135 · Cited by 9
Ye, H.-F., Zhang, H., Xiang, J., Shen, G., Yang, F., Wang, F., Wang, J., & Tang, Y.-L. (2023) · International Journal of Biological Macromolecules · doi:10.1016/j.ijbiomac.2023.127825 · Cited by 9
Yuwen, X.-Y., Feng, Y., Huang, W., Li, X., & Lai, G.-S. (2022) · Social Science Research Network · doi:10.2139/ssrn.4159823 · Cited by 9
Stabilization of G-Quadruplex-Duplex Hybrid Structures Induced by Minor Groove-Binding Drugs
Scott, L., & Chalikian, T. (2022) · Life · doi:10.3390/life12040597 · Cited by 9
Giorgio, E. D., Ferino, A., Choudhary, H., Löffler, P. M. G., D’Este, F., Rapozzi, V., Tikhomirov, A. S., Shchekotikhin, A. E., Vogel, S., & Xodo, L. E. (2022) · Journal of Photochemistry and Photobiology B Biology · doi:10.1016/j.jphotobiol.2022.112449 · Cited by 13
Porzio, A. D., Galli, U., Amato, J., Zizza, P., Iachettini, S., Iaccarino, N., Marzano, S., Santoro, F., Brancaccio, D., Carotenuto, A., et al. (2021) · International Journal of Molecular Sciences · doi:10.3390/ijms222111959 · Cited by 14
“Indeed, they may offer telomere protectio and bl ck replication fork progression [4], while in gen promo ers, they may r present a physical obsta le in the way of the RNA polymer se machinery, thus affecting the expression of the downstream gene [13].”
BackgroundKumara, G. S. R., Pandtih, A., & Seo, Y. J. (2021) · Bioorganic & Medicinal Chemistry Letters · doi:10.1016/j.bmcl.2021.128462 · Cited by 9
The fellowship of the RING: BRCA1, its partner BARD1 and their liaison in DNA repair and cancer
Russi, M., Marson, D., Fermeglia, A., Aulić, S., Fermeglia, M., Laurini, E., & Pricl, S. (2021) · Pharmacology & Therapeutics · doi:10.1016/j.pharmthera.2021.108009 · Cited by 14
Folding Dynamics of DNA G-Quadruplexes Probed by Millisecond Temperature Jump Circular Dichroism.
Laouer, K., Schmid, M., Wien, F., Changenet, P., & Hache, F. (2021) · Journal of Physical Chemistry B · doi:10.1021/acs.jpcb.1c01993 · Cited by 9
New perspectives in cancer drug development: computational advances with an eye to design
Castelli, M., Serapian, S. A., Marchetti, F., Triveri, A., Pirota, V., Torielli, L., Collina, S., Doria, F., Freccero, M., & Colombo, G. (2021) · RSC Medicinal Chemistry · doi:10.1039/d1md00192b · Cited by 13
Monti, P., Brázda, V., Bohálová, N., Porubiaková, O., Menichini, P., Speciale, A., Bocciardi, R., Inga, A., & Fronza, G. (2021) · Genes · doi:10.3390/genes12020277 · Cited by 9
“G4 motifs are evolutionarily conserved from bacteria to human, confirming the importance of their formation in vivo [54].”
BackgroundNavarro, A., Benabou, S., Eritja, R., & Gargallo, R. (2020) · International Journal of Biological Macromolecules · doi:10.1016/j.ijbiomac.2020.05.062 · Cited by 9
Saha, P., Kumar, Y. P., Das, T., Müller, D., Bessi, I., Schwalbe, H., & Dash, J. (2019) · Bioconjugate chemistry · doi:10.1021/acs.bioconjchem.9b00655 · Cited by 9
How DNA secondary structures drive replication fork instability
Sethi, A., Fernández-Casañas, M., Delpino, B., & Coster, G. (2025) · DNA repair · doi:10.1016/j.dnarep.2025.103913 · Cited by 7
Development of Antitumor Drugs Targeting c-MYC Pu27 and KRAS G-Quadruplexes.
Mohamed, H. I., Song, Z., He, Y., & Gao, C. (2025) · Journal of Medicinal Chemistry · doi:10.1021/acs.jmedchem.3c00955 · Cited by 8
Metadata
- Authors
- Spiegel, J., Adhikari, S., & Balasubramanian, S.
- Affiliations
- Cancer Research UK (GB)
- University of Cambridge (GB)
- Corresponding author
- Balasubramanian, Shankar
- Venue
- Trends in Chemistry
- Publisher
- Elsevier BV
- Published
- 2020-02
- Volume
- 2
- Issue
- 2
- Pages
- 123-136
- ISSN
- 2589-5974 (ISSN-L)
- PMC
- PMC7472594
- OpenAlex
- W2966334511
- Cited by
- 920
- References
- 121
- Citations by year
- 2026: 111 · 2025: 157 · 2024: 158 · 2023: 144 · 2022: 151 · 2021: 122 · 2020: 74
- FWCI
- 7.73
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Trends in Chemistry (publishedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by-nc-nd)
- PubMed (publishedVersion)
- Apollo (University of Cambridge) (submittedVersion, cc-by-nc-nd)
- Apollo
- Funders
- Wellcome Trust
- Cancer Research UK
- Horizon 2020 Framework Programme
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › DNA and Nucleic Acid Chemistry
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Advanced biosensing and bioanalysis techniques
- Health Sciences › Medicine › Oncology › Metal complexes synthesis and properties
- Keywords
- G-quadruplex, guanine-rich DNA, genome stability, transcription, DNA replication, epigenetic regulation, cancer, G-quadruplex-binding proteins
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