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Base pairs are two nucleotide bases interacting via hydrogen bonding.

Bulge is a nonpaired nucleotide base in a double‐stranded nucleic acid.

Hybridisation is a process where two complementary strands interact via base pairing.

Nucleic acid conformation is the three‐dimensional structural arrangement of a double‐helical nucleic acid.

Nucleic acid modification is the incorporation, removal or substitution of a chemical group on the natural occurring nucleic acids.

Melting curves are the optical changes of a nucleic acid or protein‐like molecules as a function of temperature.

Stability of nucleic acids is the resistance of nucleic acid duplex to undergo unfolding, proportional to the magnitude of Δ G .

Thermal stability is the ability of a nucleic acid duplex to resist temperature denaturation.

Unfolding thermodynamic profile is the free energy (Δ G ∘ ), enthalpy (Δ H ) and entropy (Δ S ) parameters associated with the unfolding of a nucleic acid or protein molecule.

Cooperativity refers to the number of base pairs that are broken simultaneously in the unfolding of a nucleic acid or protein.

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Bottom: A*–C and G*–T against G–C. The asterisk marks the minor tautomer.","type":"text","version":1}],"direction":null,"format":"","indent":0,"type":"paragraph","version":1,"textFormat":0,"textStyle":""},{"children":[{"detail":0,"format":0,"mode":"normal","style":"","text":"All base pairs in this figure have the same shape, so the A–C and G–T mismatches are excellent mimics of Watson-Crick base pairs. Around 1 in 10⁴ molecules of the canonical bases exist as minor tautomers at any time. The probability of a nucleobase being a minor tautomer during replication and also during proofreading is 10⁻⁴ × 10⁻⁴ = 10⁻⁸. This is the error rate during copying of the human genome. Evolution has 'chosen' DNA bases that have one tautomer that predominates. 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This modified DNA may be amplified by DNA polymerases with high efficiency and greater than 99.9% fidelity. 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They are required to recognise short complementary DNA or RNA sequences embedded in longer nucleic acid stretches. High selectivity for the correct target and tight binding are important, and this cannot always be achieved by unmodified oligonucleotides.","type":"text","version":1}],"direction":null,"format":"","indent":0,"type":"paragraph","version":1,"textFormat":0,"textStyle":""},{"children":[{"detail":0,"format":0,"mode":"normal","style":"","text":"Duplex stability can be increased by increasing the number of inter-base hydrogen bonds, by increasing base stacking, and by introducing positive charges into the nucleotides. 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Examples are locked nucleic acid (LNA), which is conformationally locked in the A-form; hexitol nucleic acid (HNA); peptide nucleic acid (PNA), which is uncharged; and morpholino, which is uncharged or cationic.","type":"text","version":1}],"direction":null,"format":"","indent":0,"type":"paragraph","version":1,"textFormat":0,"textStyle":""}],"direction":null,"format":"","indent":0,"type":"root","version":1}},"link":{"type":"reference","appearance":"default"},"id":"6aa69f3d55f3858475d4d3e0"}],"plexBand":"auto","id":"6aa69f2255f3858475d4d3df"},{"blockType":"content","layout":"stacked","sectionHeading":"The major groove","columnHeadingStyle":"heading","columns":[{"size":"full","richText":{"root":{"children":[{"children":[{"detail":0,"format":0,"mode":"normal","style":"","text":"A–T, T–A, C–G and G–C are different in the major groove of DNA. The major groove provides an accessible surface for protein binding and recognition. 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In a parallel triplex the triplex-forming oligonucleotide (TFO) runs in the same direction as the purine strand of the duplex.","type":"text","version":1}],"direction":null,"format":"","indent":0,"type":"paragraph","version":1,"textFormat":0,"textStyle":""},{"children":[{"detail":0,"format":0,"mode":"normal","style":"","text":"Advantages: it recognises duplex sequence without denaturation, and is a means of recognising double-stranded genomic DNA. Disadvantages: pH dependence (the C⁺·G–C triad), low affinity and limited sequence recognition.","type":"text","version":1}],"direction":null,"format":"","indent":0,"type":"paragraph","version":1,"textFormat":0,"textStyle":""}],"direction":null,"format":"","indent":0,"type":"root","version":1}},"link":{"type":"reference","appearance":"default"},"id":"6aa69fbb55f3858475d4d3e4"}],"plexBand":"auto","id":"6aa69fad55f3858475d4d3e3"},{"blockType":"content","layout":"stacked","sectionHeading":"G-quadruplex formation","columnHeadingStyle":"heading","columns":[{"size":"full","richText":{"root":{"children":[{"type":"upload","version":3,"format":"","id":"6aa6a00ca6cd8f4f4e664be2","fields":null,"relationTo":"media","value":{"createdAt":"2026-09-13T12:27:24.328Z","updatedAt":"2026-09-13T12:29:26.419Z","internalOnly":false,"prefix":"media","url":"/api/media/file/fig19-g-quartet.png?prefix=media","filename":"fig19-g-quartet.png","mimeType":"image/png","filesize":91415,"width":1560,"height":781,"focalX":50,"focalY":50,"sizes":{"thumbnail":{"url":"/api/media/file/fig19-g-quartet-300x150.png?prefix=media","width":300,"height":150,"mimeType":"image/png","filesize":19965,"filename":"fig19-g-quartet-300x150.png"},"square":{"url":"/api/media/file/fig19-g-quartet-500x500.png?prefix=media","width":500,"height":500,"mimeType":"image/png","filesize":32118,"filename":"fig19-g-quartet-500x500.png"},"small":{"url":"/api/media/file/fig19-g-quartet-600x300.png?prefix=media","width":600,"height":300,"mimeType":"image/png","filesize":46794,"filename":"fig19-g-quartet-600x300.png"},"medium":{"url":"/api/media/file/fig19-g-quartet-900x451.png?prefix=media","width":900,"height":451,"mimeType":"image/png","filesize":75161,"filename":"fig19-g-quartet-900x451.png"},"large":{"url":"/api/media/file/fig19-g-quartet-1400x701.png?prefix=media","width":1400,"height":701,"mimeType":"image/png","filesize":131450,"filename":"fig19-g-quartet-1400x701.png"},"xlarge":{"url":null,"width":null,"height":null,"mimeType":null,"filesize":null,"filename":null},"og":{"url":"/api/media/file/fig19-g-quartet-1200x630.png?prefix=media","width":1200,"height":630,"mimeType":"image/png","filesize":112549,"filename":"fig19-g-quartet-1200x630.png"}},"thumbnailURL":"/api/media/file/fig19-g-quartet-300x150.png","alt":"A G-quartet held by hydrogen bonding and base stacking, and a G-quadruplex folded from a single DNA strand","id":"6aa696ac3a773b5151d6108b"}},{"children":[{"detail":0,"format":0,"mode":"normal","style":"","text":"A G-quartet, held by hydrogen bonding and base stacking, and a G-quadruplex folded from a single strand of DNA:","type":"text","version":1}],"direction":null,"format":"","indent":0,"type":"paragraph","version":1,"textFormat":0,"textStyle":""},{"type":"sequence","version":1,"sense":"...GGGTTAGGGTTAGGGTTA...","antisense":null,"duplexOffset":null,"uid":"","sourceUid":null},{"children":[{"detail":0,"format":0,"mode":"normal","style":"","text":"Single strands of DNA can fold into G-quadruplexes, which are stabilised by G-quartets. In the human genome these structures form and stabilise the ends of chromosomes. They also block gene expression in a reversible manner. The metal ion is essential: K⁺ is the best fit and the most stabilising, but Na⁺ is also possible. 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Understanding the basic components and structural features of DNA and RNA provides a foundation for studying more advanced nucleic-acid technologies. Small changes in nucleic-acid structure can have significant effects on their biological properties and applications. 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Here, we report the state of art of the triplex-based antigene strategy 50 years after the discovery of such a structure, and we show the importance of the actual applications and the main challenges that we still have ahead of us.","DOI":"10.1093/nar/gkn493","type":"journal-article","created":{"date-parts":[[2008,8,1]],"date-time":"2008-08-01T20:34:31Z","timestamp":1217622871000},"page":"5123-5138","source":"Crossref","is-referenced-by-count":326,"title":["The triple helix: 50 years later, the outcome"],"prefix":"10.1093","volume":"36","author":[{"given":"Maria","family":"Duca","sequence":"first","affiliation":[{"name":"LCMBA CNRS UMR6001, University of Nice-Sophia Antipolis, Parc Valrose, 06108 NICE Cedex 2"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pierre","family":"Vekhoff","sequence":"additional","affiliation":[{"name":"UMR 5153 CNRS, Muséum National d’Histoire Naturelle USM0503"},{"name":"INSERM UR565, 43 rue Cuvier, 75231 Paris Cedex 05"},{"name":"University Pierre et Marie Curie, Paris 75005, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kahina","family":"Oussedik","sequence":"additional","affiliation":[{"name":"UMR 5153 CNRS, Muséum National d’Histoire Naturelle USM0503"},{"name":"INSERM UR565, 43 rue Cuvier, 75231 Paris Cedex 05"},{"name":"University Pierre et Marie Curie, Paris 75005, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ludovic","family":"Halby","sequence":"additional","affiliation":[{"name":"UMR 5153 CNRS, Muséum National d’Histoire Naturelle USM0503"},{"name":"INSERM UR565, 43 rue Cuvier, 75231 Paris Cedex 05"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paola B.","family":"Arimondo","sequence":"additional","affiliation":[{"name":"UMR 5153 CNRS, Muséum National d’Histoire Naturelle USM0503"},{"name":"INSERM UR565, 43 rue Cuvier, 75231 Paris Cedex 05"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2008,8,1]]},"reference":[{"key":"2021080317501543700_B1","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1038/172156a0","article-title":"Evidence for 2-chain helix in crystalline structure of sodium deoxyribonucleate","volume":"172","author":"Franklin","year":"1953","journal-title":"Nature"},{"key":"2021080317501543700_B2","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1038/171737a0","article-title":"Molecular structure of nucleic acids; a structure for deoxyribose nucleic acid","volume":"171","author":"Watson","year":"1953","journal-title":"Nature"},{"key":"2021080317501543700_B3","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1038/171738a0","article-title":"Molecular structure of deoxypentose nucleic acids","volume":"171","author":"Wilkins","year":"1953","journal-title":"Nature"},{"key":"2021080317501543700_B4","doi-asserted-by":"crossref","first-page":"2023","DOI":"10.1021/ja01565a074","article-title":"Formation of a three-stranded polynucleotide molecule","volume":"79","author":"Felsenfeld","year":"1957","journal-title":"J. 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