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projects: chromatin enzymes
projects: chromaint factors

projects: DNA ladders
projects: polycistronic expression
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gallery: amino acids
gallery: secondary structure
gallery: selected proteins
gallery: protein/DNA complexes

background reading


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Gene regulation is fundamental to the proper functioning of a cell, and many cancers can be traced to abnormal gene regulation. Our goal is to understand how genes are regulated by combining genetic, biochemical and structural descriptions. Our expectation is that such information will contribute to the development of future therapeutic agents against cancer and other human diseases.

We combine biochemical and structural approaches, including X-ray crystallography and cryoelectron microscopy, in our studies of chromatin enzymes. The questions that we wish to address include:

1. how chromatin enzymes and transcription factors recognize nucleosomes and,
2. how gene activity is regulated on a chromatin template.

The two rate-limiting steps in structure determination of large complexes by X-ray crystallography are generating large quantities of high purity material and crystallizing the complexes. We have developed methods for reconstituting recombinant multicomponent complexes and we continue research to improve those methods. We are also investigating novel methods and tools for crystallizing multicomponent complexes.


October 2020: Song Tan named holder of the Verne M. Willaman Chair in Molecular Biology.

June 2020: Our paper describing the structure of the LSD1 histone demethylase in complex with the corepressor protein CoREST and the nucleosome is published in Molecular Cell (press release).

October 2019: Congratulations to Ryan Santilli and Joey Shibata on their 1st Place in their group in the Biochemistry-Molecular division of the 22nd UMBC Undergraduate Research Symposium!

April 2019: Congratulations to Ryan Santilli, Joey Shibata and Jack Williamson on their University Fellowships and Phi Kappa Phi Peter T. Luckie Award for Outstanding Research by a Junior at the 2019 Penn State Undergraduate Exhibition!

April 2018: Congratulations to Rosalie Sowers on her Goldwater Scholarship!

June 2017: Congratulations to alumnus Rob McGinty on his Pew Scholarship and Searle Scholarship!

May 2017: Our paper describing plasmids to prepare DNA ladders inexpensively is published in Scientific Reports. Read the news release and about the undergraduate researchers.

April 2015: Song Tan awarded the Penn State Faculty Scholar Medal

March 2015: Congratulations to Rob McGinty on his Dale F. Frey Award for Breakthrough Scientists!

November 2014: Ryan Henrici is awarded a Marshall Scholarship for graduate studies in the U.K.!

October 2014: Our paper describing the structure of the PRC1 ubiquitylation module/nucleosome complex is published in Nature (News & Views, press release).

October 2014: Song Tan awarded the Eberly College of Science C.I. Noll Award for Excellence in Teaching

April 2014: Congratulations to Brian Geary ('01) on his Penn State Alumni Achievement Award!

March 2014: Congratulations to Ryan Henrici on being named a Goldwater Scholar!

May 2012: Congratulations to undergrads Josh McAnulty and Zach Hostetler on their graduation, and a special congratulations to Zach on graduating as Student Marshal of the Eberly College of Science!

April 2012: Congratulations to Heather Agnew ('03) on her Penn State Alumni Achievement Award!

Jan 2012: Congratulations to Rob McGinty on his Damon Runyon Fellowship Award from the Damon Runyon Cancer Research Foundation!

Jan 2011: New video introduction to our research.

Nov 2009: We've moved to our new research space in the Center for Eukaryotic Gene Regulation on the 4th floor of North Frear Lab. You can check out pictures of the Opening Reception and the Center's research space.

The Penn State 38th Summer Symposium in Molecular Biology on the topic of Chromatin and Epigenetic Regulation of Transcription was held in 2019.

  National Institutes of Health logo Pew Scholars Program logo
Our research is supported by the National Institutes of Health.
We gratefully acknowledge past support from the Pew Scholars Program in the Biomedical Sciences.
Members of our Lab: Contact Info:
PI Song Tan 468A North Frear Laboratory
Postdoctoral Fellows Priit Eek University Park, PA 16802
  Othman Rechice (814) 865-3355 (office)
  Jiang Zhu (814) 865-3354 (lab)
Grad Students Jose Espinola-Lopez
    Erik Leith  
Research Associates Geoffrey Fox
  Szu-yu Kuan
Undergraduates Dan Guld

Selected Publications:

  • Kim, S., J. Zhu, N. Yennawar, P. Eek and S. Tan (2020) Crystal structure of the LSD1/CoREST histone demethylase bound to its nucleosome substrate, Mol. Cell, 78:903-914 (abstract)

  • Sun, J., M. Paduch, S. Kim, R.M. Kramer, A.F. Barrios, V. Lu, J. Luke, S. Usatyuk, A.A. Kossakoff and S. Tan (2018) Structural basis for activation of SAGA histone acetyltransferase Gcn5 by partner subunit Ada2, PNAS,115:10010-10015. (abstract)

  • Henrici, R.C., T.J. Pecen, J.L. Johnston and S. Tan (2017) The pPSU plasmids for generating DNA molecular weight markers, Scientific Reports, 7:2438. (abstract)

  • McGinty, R.K., R.D. Makde and S. Tan (2016). Preparation, crystallization, and structure determination of chromatin enzyme/nucleosome complexes, Method Enzymol, 573:43-65. (abstract)

  • Girish, T.S., R.K. McGinty and S. Tan (2016). Multivalent interactions by the Set8 histone methyltransferase with its nucleosome substrate, J. Mol Biol., 428:1531-1543. (abstract)

  • Liokatis, S., R. Klingberg, S. Tan and D. Schwarzer (2016). Differentially isotope-labeled nucleosomes to study asymmetric histone modification crosstalk by time-resolved NMR spectroscopy, Angew. Chem., doi: 10.1002/anie.201601938. (abstract)

  • Jennings, M.J., A.F. Barrios and S. Tan (2016). Elimination of truncated recombinant protein expressed in Escherichia coli by removing cryptic translation initiation site, Prot. Expr. Purif., 121:17-21. (abstract)

  • McGinty, R.K. and S. Tan (2016) Recognition of the nucleosome by chromatin factors and enzymes, Curr. Opin. Struct. Biol., 21:128-136.

  • Kuo, Y.-M., R.A. Ryan, S. Tan, J. Cote and A. Andrews (2015). Site specificity analysis of Piccolo NuA4-mediated acetylation for different histone complexes, Biochem. J., 472:239-248. (abstract)

  • Kim, S., N. Chatterjee, M.J. Jennings, B. Bartholomew, and S. Tan (2015). Extranucleosomal DNA enhances the activity of the LSD1/CoREST histone demetthylase complex. Nucl. Acid Research, 43:4868-4880. (abstract)  (pdf link, 6.1 MB)

  • McGinty, R.K, and S. Tan (2015). Nucleosome structure and function. Chem Reviews, 115:2255-2273.

  • McGinty, R.K, Henrici, R.C. and S. Tan (2014). Crystal structure of the PRC1 ubiquitylation module bound to the nucleosome. Nature, 514:591-596. (abstract)

  • McGinty, R.K. and S. Tan (2014). Histones, Nucleosomes, and Chromatin Structure. In Fundamentals of Chromatin, J.L. Workman and S.M. Abmayr, ed (New York: Springer).

  • Makde, R.D. and S. Tan (2013). Strategies for crystallizing a chromatin protein in complex with the nucleosome core particle. Anal. Biochem., 442:138-145 (abstract).

  • Tan, S. and K. Nagai (2013). Protein-nucleic interactions, 'I have a cunning plan...'. Curr. Opin. Struct. Biol., 23:1-3.

  • Huang, J and S. Tan (2013). Piccolo NuA4-Catalyzed Acetylation of Nucleosomal Histones: Critical Roles of an Esa1 Tudor/Chromo Barrel Loop and an Epl1 Enhancer of Polycomb A (EPcA) Basic Region. Mol. Cell. Bio., 33:159-169. (abstract)

  • S. Tan (2012). Deciphering how the chromatin factor RCC1 recognizes the nucleosome: the importance of individuals in the scientific discovery processs. Biochem. Soc. Trans., 40:351-356. (abstract)

  • Chittuluru, J.R., Y. Chaban, J. Monnet-Saksouk, M. J. Carrozza, V. Sapountzi, W. Selleck, J. Huang, M. Cramet, S. Allard, G. Cai1, J. L. Workman, M.J. Fried, S. Tan, J. Cote and F. J. Asturias (2011). Structure and nucleosome interaction of the NuA4 and Piccolo-NuA4 histone acetyltransferase complexes. Nature Str. Mol. Biol., 39:8378-8391. (abstract)

  • Chatterjee, N., D. Sinha, M. Lemma-Dechassa, S. Tan, M. A. Shogren-Knaak, B. Bartholomew (2011). Histone H3 tail acetylation modulates ATP-dependent remodeling through multiple mechanisms. Nucl. Acid Research, 39:8378-8391. (abstract)

  • Charles, G.M., C. Chen, S. C. Shih, S. R. Collins, P. Beltrao, X. Zhang, T. Sharma, S. Tan, A. L. Burlingame, N. J. Krogan, H. D. Madhani, G. J. Narlikar (2011). Site-specific acetylation mark on an essential chromatin-remodeling complex promotes resistance to replication stress. PNAS, 108:10620-10625. (abstract)

  • Tan, S. and C.A. Davey (2011) Nucleosome stuctural studies. Curr. Opin. Struct. Biol., 21:128-136. (abstract)

  • Tan, S. and K. Nagai (2011) Diversity makes for a rich tapestry. Curr. Opin. Struct. Biol., 21:81-82.

  • Makde, R.D., J.R. England, H. Yennawar and S. Tan (2010). Structure of RCC1 chromatin factor bound to the nucleosome core particle. Nature, 467:562-566. (abstract)

  • England, J.R., J. Huang, M.J. Jennings, R. D. Makde, and S. Tan (2010). RCC1 uses a conformationally diverse loop region to interact with the nucleosome: a model for the RCC1-nucleosome complex. J. Mol Biol., 398:518-529. (abstract)

  • Choy, J.S., S. Wei, J.Y. Lee, S. Tan, S. Chu and T.H. Lee (2010) DNA methylation increases nucleosome compaction and rigidity, J. Am. Chem. Soc., 132:1782-1783. (abstract)

  • Psathas, J.N., S. Zheng, S. Tan and J. Reese (2009) Set2-dependent K36 methylation is regulated by novel intratail interactions within H3, Mol. Cell. Biol., 29:6413-6426. (abstract)

  • S. Tan and J. Reese (editors) (2009) BBA - Gene Regulatory Mechanisms Special Issue on Epigenetics, 1789:1-68.

  • Selleck, W. and S. Tan (2008) Recombinant protein complex expression in E. coli, Curr Protocols Protein Sci, Chapter 5, unit 5.21. (abstract)

  • Altaf, M., R.T. Utley, N. Lacoste, S. Tan, S.D. Briggs, J. Cote (2007) Interplay of chromatin modifiers on a short basic patch of histone H4 tail defines the boundary of telomeric heterochromatin, Mol. Cell, 28:1002-1014. (abstract)

  • Barrios, A., W. Selleck, B. Hnatkovich, R. Kramer, D. Sermwittayawong and S. Tan (2007) Expression and purification of recombinant yeast Ada2/Ada3/Gcn5 and Piccolo NuA4 histone acetyltransferase complexes, Methods, 41:271-277. (abstract)

  • Berndsen, C.E., W. Selleck, S.J. McBryant, J.C. Hansen, S. Tan and J.M. Denu (2007) Nucleosome recognition by the Piccolo NuA4 histone acetyltransferase complex, Biochemistry, 46:2091-2099. (abstract)

  • Berndsen, C.E., B.N. Albaugh, S. Tan and J.M. Denu (2007) Catalytic mechanism of a MYST family histone acetyltransferase, Biochemistry, 46:623-629, Accelerated Publication. (abstract)

  • Sermwittayawong, D. and S. Tan (2006) SAGA binds TBP via its Spt8 subunit in competition with DNA, implications for TBP recruitment, EMBO Journal, 25:3791-3800. (abstract)
  • Doyon Y., C. Cayrou, M. Ullah, A.J. Landry, V. Cote, W. Selleck, W.S. Lane, S. Tan, W.J. Yang, X.J. and J. Cote (2006) ING Tumor Suppressor Proteins Are Critical Regulators of Chromatin Acetylation Required for Genome Expression and Perpetuation, Mol. Cell, 21:51-64. (abstract)

  • Selleck, W., I. Fortin, D. Sermwittayawong, J. Cote and S. Tan (2005) The yeast Piccolo NuA4 HAT complex requires the Enhancer of Polycomb A and chromo domains to acetylate nucleosomes, Mol. Cell. Biol., 25:5535-5542. (abstract)

  • Lichty, J.L., J.L. Malecki, H.D. Agnew, D.J. Michelson-Horowitz and S. Tan (2005) Comparison of Affinity Tags for Protein Purification, Prot. Expr. Purif., 41:98-105. (abstract)
  • Tan, S., R.C. Kern and W. Selleck (2005) The pST44 polycistronic expression system for producing protein complexes in Escherichia coli, Prot. Expr. Purif., 40:385-395. (abstract)

  • Doyon, Y., W. Selleck, W.S. Lane, S. Tan and J. Côté (2004) Structural and functional conservation of the NuA4 histone acetyltransferase complex from yeast to human, Mol. Cell. Biol., 24:1884-1896. (abstract)

  • Boudreault, A.A. , D. Cronier, W. Selleck, N. Lacoste, R.T. Utley, S. Allard, J. Savard, W.S. Lane, S. Tan and J. Côté (2003) Yeast Enhancer of polycomb defines global Esa1-dependent acetylation of chromatin, Genes & Dev, 17:1415-1428. (abstract)

  • Bleichenbacher, M.M., S. Tan and T.J. Richmond (2003) Novel Interactions Between the Components of Human and Yeast TFIIA/TBP/DNA Complexes, J. Mol Biol., 332:783-793. (abstract)

  • Boyer, L.A., M.R. Langer, K.A. Crowley, S. Tan, J.M. Denu, and C.L. Peterson (2002) Essential role for the SANT domain in the functioning of multiple chromatin remodeling enzymes. Mol. Cell, 10:935-942. (abstract)

  • Balasubramanian, R., M.G. Pray-Grant, W. Selleck, P.A. Grant and S. Tan (2002) Role of the Ada2 and Ada3 transcriptional coactivators in histone acetylation, J. Biol. Chem., 277: 7989-7995. (abstract)

  • Selleck, W., R. Howley, J. Fang, V. Podolny, M.G. Fried, S. Buratowski and S. Tan (2001) A histone fold TAF octamer within the yeast TFIID transcriptional coactivator, Nature Struct. Biol., 8:695-700. (abstract)
  • Brown, C.E., L. Howe, K. Sousa, S.C. Alley, M.J. Carrozza, S. Tan and J.L. Workman (2001) Recruitment of HAT complexes by direct activator interactions with the ATM-related Tra1 subunit, Science, 292: 2333-2337. (abstract)
  • Tan, S. (2001) A Modular Polycistronic Expression System for Overexpressing Protein Complexes in Escherichia Coli, Prot. Expr. Purif., 21: 224-234. (abstract)
  • Gaiser, F., S. Tan and T.J. Richmond (2000) Novel Dimerization Fold of RAP30/RAP74 in Human TFIIF at 1.7 Å Resolution , J. Mol Biol., 302: 1119-1127. (abstract)
  • Tan, S., Y. Hunziker, L. Pellegrini and T.J. Richmond (2000) Crystallization of the yeast MATalpha2/MCM1/DNA ternary complex: general methods and principles for protein/DNA cocrystallization, J. Mol Biol., 297: 947-959. (reprint)
  • Tan, S. and T.J. Richmond (1998) Crystal structure of the yeast MATalpha2/MCM1/DNA ternary complex, Nature, 391: 660-666. (abstract)
  • Tan, S., Y. Hunziker, D.F. Sargent and T.J. Richmond (1996) Crystal structure of a yeast TFIIA/TBP/DNA complex, Nature, 381: 127-134. (abstract)
  • Pellegrini, L, S. Tan and T.J. Richmond (1995) Structure of serum response factor core bound to DNA, Nature, 376: 490-498. (abstract)

video introduction | projects: | chromatin enzymes | chromatin factors | DNA ladders | polycistronic expression | picture & movie gallery: | amino acids | secondary structure | selected proteins | protein/DNA complexes |
background reading | reprints | positions |
lab pictures |

This page is maintained by Song Tan: (814-865-3355)
Department of Biochemistry & Molecular Biology, 108 Althouse Lab, University Park, PA 16802

This page was last updated on October 12, 2020.

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