Figure 1. 5-Hydroxymethylcytosine (5hmC) favors co-transcriptional R-loop formation.
(A) Native or modified deoxyribonucleotides (dCTPs) were incorporated upon PCR amplification into DNA fragments with sequences from the transcription termination region of ACTB (ACTB P1 and ACTB P2) or APOE. (B) Incorporation of dCTP variants confirmed by immunoblotting using specific antibodies against 5-methylcytosine (5mC), 5hmC, and double-stranded DNA (dsDNA). (C) R-loops formed upon in vitro transcription reactions were detected by immunoblotting using the S9.6 antibody. RNase H-treated in vitro transcription reaction products (RH+) serve as negative controls. All data are representative of seven independent experiments with similar results. (D) S9.6 immunoblots were quantified and the R-loop levels normalized against the levels detected in the reaction products of DNA templates containing native C. Data represent the mean and standard deviation (SD) from seven independent experiments. *p<0.05, **p<0.01, and ***p<0.001, two-tailed Student’s t-test. (E) In vitro transcription reaction products of ACTB P2 templates were visualized using atomic force microscopy (AFM). R-loop structures obtained from 5hmC-containing ACTB P2 transcription in the absence (RH-) or presence (RH+) of RNase H are shown. R-loops present in the transcription reaction products of C, 5mC, or 5hmC-containing ACTB P2 templates were counted in a minimum of 80 filaments observed in three individual AFM experiments.