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.
Current research
I study how cancer-associated mutations in histone proteins disrupt nucleosome function. In healthy tissue, histone proteins form protein–DNA complexes, the nucleosomes, that govern access to the underlying genetic information. Although there are roughly 16 million nucleosomes for every copy of the human genome, there are only a few dozen per gene which can be enzymatically modified to signal different gene activity states. Defective nucleosomes can cause aberrant gene expression or silencing, promoting hallmark cancer phenotypes such as unrestrained proliferation, migration, and invasion.
A key challenge is distinguishing innocuous from detrimental histone mutations. To address this, I create and screen libraries of cancer-associated histones in cells, use chemical synthesis to reconstitute defined nucleosome substrates, and integrate screening data with structural and clinical information to identify functionally meaningful patterns.
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 look up recipes, 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 | 2026-03-03 |
| SOP0002.1 | Whole cell lysates from tissue culture cells | Cell biology | Public | 2026-03-03 |
| SOP0003.1 | Native polyacrylamide gel electrophoresis | Biochemistry | Public | 2026-02-28 |
| SOP0004.3 | Core histone expression and purification | Biochemistry | Internal | 2026-03-03 |
| SOP0005.2 | Histone octamer, tetramer, or dimer formation | Biochemistry | Public | 2026-03-03 |
| SOP0006.2 | Reconstitution of mononucleosomes and nucleosome arrays | Biochemistry | Internal | 2026-03-03 |
| SOP0007.1 | Denaturing SDS polyacrylamide gel electrophoresis | Biochemistry | Public | 2026-03-03 |
| SOP0008.1 | Western blotting | Biochemistry | Public | 2026-03-03 |
| SOP0009.1 | Immunoblotting | Biochemistry | Public | 2026-03-03 |
| SOP0010.1 | In-gel protein staining with Coomassie Brilliant Blue | Biochemistry | Public | 2026-03-03 |
| SOP0011.1 | In-gel protein staining with silver nitrate | Biochemistry | Public | 2026-03-03 |
| SOP0012.2 | Non-viral transfection of animal tissue culture cells | Tissue culture | Public | 2026-03-03 |
| SOP0013.1 | Electroporation of animal tissue culture cells | Tissue culture | Internal | 2026-02-28 |
| SOP0014.1 | Counting cells | Tissue culture | Public | 2026-02-27 |
| SOP0015.1 | Cultivating HeLa cells | Tissue culture | Public | 2026-02-28 |
| SOP0016.1 | Cultivating MCF 10A cells | Tissue culture | Public | 2026-03-03 |
| SOP0017.3 | Lentiviral generation of stable animal cell lines | Tissue culture | Public | 2026-03-03 |
| SOP0018.1 | Generation of stable cell lines using DNA transposases | Tissue culture | Public | 2026-02-28 |
| SOP0019.1 | Selection and screening of stable mammalian cell lines | Tissue culture | Public | 2026-02-27 |
| SOP0020.1 | Plasmid preparation by alkaline lysis | Molecular biology | Public | 2026-02-27 |
| SOP0021.1 | Purification of nucleic acids | Molecular biology | Public | 2026-02-28 |
| SOP0022.1 | Purification of nucleic acids from agarose gels | Molecular biology | Public | 2026-02-28 |
| SOP0023.1 | Separation of RNA from DNA with guanidinium thiocyanate | Molecular biology | Public | 2026-03-03 |
| SOP0024.1 | Extraction of genomic DNA from tissue culture cells | Molecular biology | Public | 2026-03-03 |
| SOP0025.1 | Nuclear extraction and fractionation of chromatin-associated proteins | Cell biology | Public | 2026-03-03 |
| SOP0026.1 | Acidic extraction of histone proteins | Cell biology | Public | 2026-02-28 |
| SOP0027.1 | Immunoprecipitation of nuclear and cytosolic proteins | Biochemistry | Internal | 2026-03-03 |
| SOP0028.2 | Cell cycle synchronization of mammalian cell lines | Tissue culture | Internal | 2026-03-03 |
| SOP0029.1 | Cell cycle analysis by fluorescence staining | Tissue culture | Internal | 2026-03-03 |
| SOP0030.1 | Making reagents and buffers | Core skills | Public | 2026-02-28 |
| SOP0031.1 | Designing and performing dilutions | Core skills | Public | 2026-02-28 |
| SOP0032.1 | Making and running agarose gels | Molecular biology | Public | 2026-03-03 |
| SOP0033.1 | Restriction digest, end modification, and ligation of DNA | Molecular biology | Public | 2026-02-27 |
| SOP0034.1 | Transformation of plasmids into a bacterial host | Molecular biology | Public | 2026-03-03 |
| SOP0035.1 | Writing a good protocol | Core skills | Internal | 2026-03-03 |
| SOP0036.1 | Banking, tracking, and sharing of materials | Core skills | Internal | 2026-02-25 |
| SOP0038.1 | Design and cloning of CRISPR guide RNAs | Tissue culture | Public | 2026-02-28 |
| SOP0039.1 | Qualitative polymerase chain reactions | Molecular biology | Public | 2026-03-03 |
| SOP0101.1 | Version control using Git and GitHub | Data management | Public | 2026-02-28 |