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. Author manuscript; available in PMC: 2026 Jan 26.
Published in final edited form as: Trends Genet. 2025 Aug 13;42(2):163–176. doi: 10.1016/j.tig.2025.07.006

Table 1.

Summary of existing methodologies for identification of Z-DNA

Category Subtype Methodology Refs
Biophysical methods Spectroscopy CD [1,35]
IR spectroscopy [6]
Raman spectroscopy (including surface-enhanced Raman) [7,8]
NMR Solution-state NMR [7,9,10]
X-ray diffraction Single-crystal X-ray diffraction [2,11]
Fluorescence techniques FRET (Z-to-B transition monitoring) [12]
smFRET [13]
Hydrogen exchange kinetics HDX [14]
Thermodynamic analysis ITC [15]
DSC [15]
Biochemical methods Electrophoresis Native/denaturing gel electrophoresis [1620]
Torque spectroscopy Single-molecule torque measurements [21,22]
Ultracentrifugation Sedimentation analysis [23]
Membrane-binding assays Nitrocellulose filtration [24]
Ligand-based detection Ruthenium complex groove binding [24]
Nuclease sensitivity DNase I/restriction enzyme cleavage assays [25,26]
Chemical probes Bromine/OsO4/DEPC modification [27,28]
Antibody/protein based High-throughput profiling ZαADAR1 ChAP-seq [34]
Zaa-FLAG ChIP-seq [30]
Z22 antibody-based ZIP-seq/ChIP-seq/CUT&TAG [35,36]
Computational prediction Algorithmic tools Z-Hunter [32]
DeepZ (machine learning) [29,32]
Z-DNABERT/ZSeeker [33]
nBMST [29]
Protein-independent methods Genome-wide mapping ssDNA mapping + PZFS overlap [37]
ViCAR Hi-C adaptation [38,39]