Hi, I’m Ben!
I am an Associate Research Scholar and former EMBO Postdoctoral Research Fellow in the Department of Chemistry at Princeton University where I work with Prof. Tom Muir on how cancer seizes control of gene regulation.
In 2021, I received my PhD in Chemical Biology, advised by Prof. Daniel Summerer, for developing proteins that recognize rare DNA modifications in the human genome.
ORCID · Google Scholar · GitHub · LinkedIn
Research interests
My work focuses on:
- How do small chemical changes in chromatin control cell-type-specific gene regulation?
- How do diseases exploit or disrupt epigenetic mechanisms?
I study these questions using chemical biology approaches, pooled genetic screens, and integrated analysis of multi-omics data, connecting molecular features to regulatory outcomes and potential therapeutic vulnerabilities.
Publications
Preprints
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HM-DyadCap – Capture and Mapping of 5-Hydroxymethylcytosine/5-Methylcytosine CpG Dyads in Mammalian DNA.bioRxiv 2025. doi:10.1101/2025.10.29.685270 DOI
Original Research
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Evolved Readers of 5‐Carboxylcytosine CpG Dyads Reveal a High Versatility of the Methyl‐CpG‐Binding Domain for Recognition of Noncanonical Epigenetic Marks.Angew. Chem. Int. Ed. 2024, 63(17), e202318837. doi:10.1002/anie.202318837 DOI PMID:38284298 PMID
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Epigenetic CpG duplex marks probed by an evolved DNA reader via a well-tempered conformational plasticity.Nucleic Acids Res. 2023, 51(12), 6495–6506. doi:10.1093/nar/gkad134 DOI PMID:36919612 PMID PMC10325892 PMC
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A translational repression reporter assay for the analysis of RNA-binding protein consensus sites.RNA Biol. 2023, 20(1), 85–94. doi:10.1080/15476286.2023.2192553 DOI PMID:36946649 PMID PMC10038052 PMC
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Imaging-Based In Situ Analysis of 5‑Methylcytosine at Low Repetitive Single Gene Loci with Transcription-Activator-Like Effector Probes.ACS Chem. Biol. 2023, 18(2), 230–236. doi:10.1021/acschembio.2c00857 DOI PMID:36693632 PMID PMC9942090 PMC
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A high-throughput effector screen identifies a novel small molecule scaffold for inhibition of ten-eleven translocation dioxygenase 2.RSC Med. Chem. 2022, 13(12), 1540–1548. doi:10.1039/d2md00186a DOI PMID:36545435 PMID PMC9749932 PMC
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Evolved DNA Duplex Readers for Strand-Asymmetrically Modified 5‑Hydroxymethylcytosine/5-Methylcytosine CpG Dyads.Impact: First affinity probes enabling selective enrichment of strand-asymmetric epigenetic DNA marks from genomic DNA
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Engineered TALE Repeats for Enhanced Imaging‐Based Analysis of Cellular 5‐Methylcytosine.
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Light‐Activation of DNA‐Methyltransferases.Angew. Chem. Int. Ed. 2021, 60(24), 13507–13512. doi:10.1002/anie.202103945 DOI PMID:33826797 PMID PMC8251764 PMC
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Light-Activatable TET-Dioxygenases Reveal Dynamics of 5‑Methylcytosine Oxidation and Transcriptome Reorganization.JACS 2020, 142(16), 7289–7294. doi:10.1021/jacs.0c01193 DOI PMID:32286069 PMIDImpact: First temporal control of TET dioxygenase activity in mammalian cells enabling time-resolved monitoring of methylation dynamics and transcriptome reorganization
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CRISPR-Cas12a–assisted PCR tagging of mammalian genes.J. Cell Biol. 2020, 219(6), e201910210. doi:10.1083/jcb.201910210 DOI PMID:32406907 PMID PMC7265327 PMCImpact: Extended our CASTLING approach for gene editing to mammalian cells with up to 60% in-frame tagging efficiency; now widely adopted for endogenous protein tagging studies
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Designer Receptors for Nucleotide‐Resolution Analysis of Genomic 5‐Methylcytosine by Cellular Imaging.Angew. Chem. Int. Ed. 2020, 59(23), 8927–8931. doi:10.1002/anie.202001935 DOI PMID:32167219 PMID PMC7318601 PMCImpact: First nucleotide-resolution imaging of 5mC in single cells; enabled direct correlation between epigenetic marks and transcription factor recruitment
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Complete Profiling of Methyl-CpG-Binding Domains for Combinations of Cytosine Modifications at CpG Dinucleotides Reveals Differential Read-out in Normal and Rett-Associated States.Impact: First comprehensive profiling of the human MBD protein family revealing differential recognition of oxidized CpG marks by Rett syndrome-associated MeCP2 mutations
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Pooled clone collections by multiplexed CRISPR-Cas12a-assisted gene tagging in yeast.Nat. Commun. 2019, 10(1), 2960. doi:10.1038/s41467-019-10816-7 DOI PMID:31273196 PMID PMC6609715 PMCImpact: CASTLING enables rapid construction of yeast clone libraries with >90% tagging efficiency; method adapted for mammalian gene tagging (Fueller et al. 2020, J Cell Biol) and cited in advanced Cas12a engineering studies
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Programmable Protein–DNA Cross-Linking for the Direct Capture and Quantification of 5‑Formylcytosine.JACS 2019, 141(24), 9453–9457. doi:10.1021/jacs.9b01432 DOI PMID:31180648 PMID
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Overcoming conservation in TALE–DNA interactions: a minimal repeat scaffold enables selective recognition of an oxidized 5-methylcytosine.
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Genome-wide C-SWAT library for high-throughput yeast genome tagging.Nat. Methods 2018, 15(8), 598–600. doi:10.1038/s41592-018-0045-8 DOI PMID:29988096 PMID
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Monomeric and Dimeric 68Ga-Labeled Bombesin Analogues for Positron Emission Tomography (PET) Imaging of Tumors Expressing Gastrin-Releasing Peptide Receptors (GRPrs).J. Med. Chem. 2018, 61(5), 2062–2074. doi:10.1021/acs.jmedchem.7b01856 DOI PMID:29432691 PMID
Reviews, Book Chapters, Monographs
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Deciphering strand-asymmetrically modified CpG dyads in the DNA double-helix.
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DNA Modifications, Methods and Protocols.Methods Mol. Biol. 2021, 2198, 381–399. doi:10.1007/978-1-0716-0876-0 DOI PMID:32822046 PMID
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Programmable tools for targeted analysis of epigenetic DNA modifications.Curr. Opin. Chem. Biol. 2021, 63, 1–10. doi:10.1016/j.cbpa.2021.01.002 DOI PMID:33588304 PMID
Protocols
Good protocols evolve over time, but printed copies often don’t. Here, I share version-controlled write-ups to help track changes. You can check whether your printed copy is current, or just browse for the latest method:
| Identifier | Title | Domain | Status | Last commit |
|---|---|---|---|---|
| SOP0001.1 | Isolation of nuclei from tissue culture cells | Cell biology | Public | 2025-11-03 |
| SOP0002.1 | Whole cell lysates from tissue culture cells | Cell biology | Internal | |
| SOP0003.1 | Native polyacrylamide gel electrophoresis | Biochemistry | Internal | |
| SOP0004.3 | Core histone expression and purification | Biochemistry | Internal | |
| SOP0005.2 | Histone octamer, tetramer, or dimer formation | Biochemistry | Internal | |
| SOP0006.2 | Reconstitution of mononucleosomes and nucleosome arrays | Biochemistry | Internal | |
| SOP0007.1 | Denaturating SDS polyacrylamide gel electrophoresis | Biochemistry | Internal | |
| SOP0008.1 | Western blotting | Biochemistry | Internal | |
| SOP0009.1 | Immunoblotting | Biochemistry | Internal | |
| SOP0010.1 | In-gel protein staining with Coomassie Brilliant Blue | Biochemistry | Internal | |
| SOP0011.1 | In-gel protein staining with silver nitrate | Biochemistry | Internal | |
| SOP0012.2 | Non-viral transfection of animal tissue culture cells | Tissue culture | Internal | |
| SOP0013.1 | Electroporation of animal tissue culture cells | Tissue culture | Internal | |
| SOP0014.1 | Counting cells | Tissue culture | Internal | |
| SOP0015.1 | Cultivating HeLa cells | Tissue culture | Internal | |
| SOP0016.1 | Cultivating MCF 10A cells | Tissue culture | Internal | 2025-10-25 |
| SOP0017.3 | Lentiviral generation of stable animal cell lines | Tissue culture | Internal | |
| SOP0018.1 | Generation of stable cell lines using DNA transposases | Tissue culture | Internal | |
| SOP0019.1 | Selection and screening of stable mammalian cell lines | Tissue culture | Internal | |
| SOP0020.1 | Plasmid preparation by alkaline lysis | Molecular biology | Internal | |
| SOP0021.1 | Purification of nucleic acids | Molecular biology | Internal | |
| SOP0022.1 | Purification of nucleic acids from agarose gels | Molecular biology | Internal | |
| SOP0023.1 | Separation of RNA from DNA with guanidinium thiocyanate | Molecular biology | Internal | |
| SOP0024.1 | Extraction of genomic DNA from tissue culture cells | Molecular biology | Internal | |
| SOP0025.1 | Nuclear extraction and fractionation of chromatin-associated proteins | Cell biology | Internal | |
| SOP0026.1 | Acidic extraction of histone proteins | Cell biology | Internal | |
| SOP0027.1 | Immunoprecipitation of nuclear and cytosolic proteins | Biochemistry | Internal | |
| SOP0028.2 | Cell cycle synchronization of mammalian cell lines | Tissue culture | Internal | |
| SOP0029.1 | Cell cycle analysis by fluorescence staining | Tissue culture | Internal | |
| SOP0030.1 | Making reagents and buffers | Core skills | Public | 2025-11-03 |
| SOP0031.1 | Designing and performing dilutions | Core skills | Internal | 2025-11-03 |
| SOP0032.1 | Making and running agarose gels | Molecular biology | Internal | |
| SOP0033.1 | Restriction digest, end modification, and ligation of DNA | Molecular biology | Internal | |
| SOP0034.1 | Transformation of plasmids into a bacterial host | Molecular biology | Internal | |
| SOP0035.1 | Writing a good protocol | Core skills | Internal | 2025-11-03 |
| SOP0036.1 | Banking, tracking, and sharing of materials | Core skills | Internal | 2025-11-03 |
| SOP0037.1 | Quantitative amplicon sequencing with unique molecular identifiers | Molecular biology | Draft | |
| SOP0038.1 | Design and cloning of CRISPR guide RNAs | Tissue culture | Internal | |
| SOP0039.1 | Qualitative polymerase chain reactions | Molecular biology | Internal | |
| SOP0101.1 | Version control using Git and GitHub | Data management | Public | 2025-11-03 |