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G-quadruplex ligands bind four-stranded structures that form in guanine-rich DNA and RNA sequences. Four guanines pair through Hoogsteen hydrogen bonds into a planar G-quartet, and stacked quartets are stabilized by potassium ions. Parallel four-stranded complexes of guanine-rich DNA were described in 1988. In 2013 a structure-specific antibody visualized G-quadruplexes in human cell nuclei, with numbers that changed during the cell cycle. Such structures occur at telomeres, in gene promoters and in ribosomal DNA. Pyridostatin, reported in 2008, stabilizes telomeric G-quadruplexes and triggers a DNA damage response there. 360A and MTR-106 are further stabilizing ligands, and BMVC-8C3O is a carbazole-based G-quadruplex ligand. Quarfloxin targets G-quadruplexes in ribosomal DNA and inhibits ribosomal RNA (rRNA) biogenesis.
