2001 – 2006 Ph.D. candidate in the Department of Biomedical Sciences, State University of New York at Albany, Albany, New York. Advisor: Dr. Joachim Frank. Received Ph.D. degree in 2006.
1996 – 2000 Department of Biochemistry and Molecular Biology, College of Life Sciences, Peking University, China. Received B.S. degree in 2000.

2017/04 – present, Professor, School of Life Sciences, Peking University
2017/04 - present, PI, Peking-Tsinghua Center for Life Sciences
2008/12 – 2017/03, Assistant Professor, Associate Professor with tenure, Full Professor, School of Life Sciences, Tsinghua University
2007/01 – 2008/12, Research Associate, Howard Hughes Medical Institute, Columbia University Medical Center, New York, NY (2008/04-2008/12); Howard Hughes Medical Institute, Wadsworth Center, Albany, NY (2007/01-2008/03)
2006/08 – 2006/12, Research Affiliate, Health Research Inc., Wadsworth Center, Albany, New York

The general research interest of our laboratory is to understand mechanistic details of various cellular molecular machines, using cryo-EM technique as a major tool. In the past a few years, we have primarily focused on studying ribosome biogenesis in both prokaryotic and eukaryotic systems, trying to answer a few fundamental questions, such as “how the ribosome is made in the cell”, and “how ribosome biogenesis, or the translation capacity of a cell is regulated by various growth-control pathways”. The ongoing projects in the lab include:
1. Ribosome biogenesis and translation regulation.
2. Structure and mechanisms of DNA replication machinery.

1. Jiang, F., Shen, J., Cheng, J., Wang, X., Yang, J., Li, N., Gao, N., and Jin, Q. (2022). N-terminal signal peptides facilitate the engineering of PVC complex as a potent protein delivery system. Sci Adv 8, eabm2343.
2. Micic, J., Rodriguez-Galan, O., Babiano, R., Fitzgerald, F., Fernandez-Fernandez, J., Zhang, Y., Gao, N., Woolford, J.L., and de la Cruz, J. (2022). Ribosomal protein eL39 is important for maturation of the nascent polypeptide exit tunnel and proper protein folding during translation. Nucleic acids research.
3. Cheng, J., Li, N., Huo, Y., Dang, S., Tye, B.-K.#, Gao, N.#, and Zhai, Y.# (2022). Structural Insight into the MCM double hexamer activation by Dbf4-Cdc7 kinase. Nature communications 13, 1396.
4. Du, S., Wang, G., Zhang, Z., Ma, C., Gao, N.#, and Xiao, J.# (2022). Structural insights into how GlcNAc-1-phosphotransferase directs lysosomal protein transport. The Journal of biological chemistry 298, 101702.
5. Hu, X., Jiang, S., Xu, F., Zeng, C., Wang, X., Liu, W., Cheng, A., Ma, C., Gao, N., Zhao, Y., Dai, J., and Zhao, G. (2022). Engineering and functional analysis of yeast with a monotypic 40S ribosome subunit. PNAS 119, e2114445119.
6. Ma, C., Wang, C., Luo, D., Yan, L., Yang, W., Li, N., and Gao, N. (2022). Structural insights into the membrane microdomain organization by SPFH family proteins. Cell Res 32, 176-189.
7. Wang, X., Cheng, J., Shen, J., Liu, L., Li, N., Gao, N., Jiang, F., and Jin, Q. (2022). Characterization of Photorhabdus Virulence Cassette as a causative agent in the emerging pathogen Photorhabdus asymbiotica. Sci China Life Sci 65, 618-630.
8. Zhang, W.#, Li, Z., Sun, Y., Cui, P., Liang, J., Xing, Q., Wu, J., Xu, Y., Zhang, W., Zhang, Y.#, He, L.#, and Gao, N. (2022). Cryo-EM structure of Mycobacterium tuberculosis 50S ribosomal subunit bound with clarithromycin reveals dynamic and specific interactions with macrolides. Emerg Microbes Infect 11, 293-305.
9. Liu, Y., Ai, C., Gan, T., Wu, J., Jiang, Y., Liu, X., Lu, R., Gao, N., Li, Q., Ji, X., and Hu, J. (2021). Transcription shapes DNA replication initiation to preserve genome integrity. Genome Biol 22, 176.
10. Zheng, L., Zheng, Z., Li, X., Wang, G., Zhang, K., Wei, P., Zhao, J.#, and Gao, N#. (2021). Structural insight into the mechanism of energy transfer in cyanobacterial phycobilisomes. Nature communications 12, 5497.
11. Zhang, J., Zhang, D., Sun, L., Ji, G., Huang, X., Niu, T., Xu, J., Ma, C., Zhu, Y., Gao, N., Xu, W., and Sun, F. (2021). VHUT-cryo-FIB, a method to fabricate frozen hydrated lamellae from tissue specimens for in situ cryo-electron tomography. J Struct Biol 213, 107763.
12. Zheng, L., Liu, Z., Wang, Y., Yang, F., Wang, J., Huang, W., Qin, J., Tian, M., Cai, X., Liu, X., Mo, X., Gao, N.#, and Jia, D.# (2021). Cryo-EM structures of human GMPPA-GMPPB complex reveal how cells maintain GDP-mannose homeostasis. Nat Struct Mol Biol 28, 1-12.
13. Yang, H., Yu, Z., Chen, X., Li, J., Li, N., Cheng, J., Gao, N., Yuan, H.X., Ye, D., Guan, K.L., and Xu, Y. (2021). Structural insights into TSC complex assembly and GAP activity on Rheb. Nature communications 12, 339.
14. Zhao, T., Chen, Y.M., Li, Y., Wang, J., Chen, S., Gao, N.#, and Qian, W.# (2021). Disome-seq reveals widespread ribosome collisions that promote cotranslational protein folding. Genome Biol 22, 16.
15. Du, S., Cao, Y., Zhu, Q., Yu, P., Qi, F., Wang, G., Du, X., Bao, L., Deng, W., Zhu, H., Liu, J., Nie, J., Zheng, Y., Liang, H., Liu, R., Gong, S., Xu, H., Yisimayi, A., Lv, Q., Wang, B., He, R., Han, Y., Zhao, W., Bai, Y., Qu, Y., Gao, X., Ji, C., Wang, Q., Gao, N., Huang, W., Wang, Y., Xie, X.S., Su, X.D., Xiao, J., and Qin, C. (2020). Structurally Resolved SARS-CoV-2 Antibody Shows High Efficacy in Severely Infected Hamsters and Provides a Potent Cocktail Pairing Strategy. Cell 183, 1013-1023 e13.
16. Micic, J., Li, Y., Wu, S., Wilson, D., Tutuncuoglu, B., Gao, N.#, and Woolford, J.L., Jr.# (2020). Coupling of 5S RNP rotation with maturation of functional centers during large ribosomal subunit assembly. Nature communications 11, 3751.
17. Wilson, D.M., Li, Y., LaPeruta, A., Gamalinda, M., Gao, N.#, and Woolford, J.L., Jr.# (2020). Structural insights into assembly of the ribosomal nascent polypeptide exit tunnel. Nature communications 11, 5111.
18. Liang, X., Zuo, M.Q., Zhang, Y., Li, N., Ma, C., Dong, M.Q., and Gao, N.# (2020). Structural snapshots of human pre-60S ribosomal particles before and after nuclear export. Nature communications 11, 3542.
19. Cao, Y., Su, B., Guo, X., Sun, W., Deng, Y., Bao, L., Zhu, Q., Zhang, X., Zheng, Y., Geng, C., Chai, X., He, R., Li, X., Lv, Q., Zhu, H., Deng, W., Xu, Y., Wang, Y., Qiao, L., Tan, Y., Song, L., Wang, G., Du, X., Gao, N., Liu, J., Xiao, J., Su, X.D., Du, Z., Feng, Y., Qin, C., Qin, C., Jin, R., and Xie, X.S. (2020). Potent Neutralizing Antibodies against SARS-CoV-2 Identified by High-Throughput Single-Cell Sequencing of Convalescent Patients` B Cells. Cell 182, 73-84 e16.
20. Wang, W., Li, W., Ge, X., Yan, K., Mandava, C.S., Sanyal, S.#, and Gao, N.# (2020). Loss of a single methylation in 23S rRNA delays 50S assembly at multiple late stages and impairs translation initiation and elongation. PNAS 117, 15609-15619.
21. Wang, Y., Wang, G., Li, Y., Zhu, Q., Shen, H., Gao, N., and Xiao, J. (2020). Structural insights into secretory immunoglobulin A and its interaction with a pneumococcal adhesin. Cell Res 30, 602-609.
22. Zhang, B., Jin, Q., Xu, L., Li, N., Meng, Y., Chang, S., Zheng, X., Wang, J., Chen, Y., Neculai, D., Gao, N., Zhang, X., Yang, F., Guo, J., and Ye, S. (2020). Cooperative transport mechanism of human monocarboxylate transporter 2. Nature communications 11, 2429.
23. Jin, Q., Zhang, B., Zheng, X., Li, N., Xu, L., Xie, Y., Song, F., Bhat, E.A., Chen, Y., Gao, N., Guo, J., Zhang, X., and Ye, S. (2020). Cryo-EM structures of human pannexin 1 channel. Cell Res 30, 449-451.
24. Li, Y., Wang, G., Li, N., Wang, Y., Zhu, Q., Chu, H., Wu, W., Tan, Y., Yu, F., Su, X.D., Gao, N., and Xiao, J. (2020). Structural insights into immunoglobulin M. Science 367, 1014-1017.
25. Ye, Y., Wu, H., Chen, K., Clapier, C.R., Verma, N., Zhang, W., Deng, H., Cairns, B.R.#, Gao, N.#, and Chen, Z.# (2019). Structure of the RSC complex bound to the nucleosome. Science 366, 838-843.
26. Zheng, L., Li, Y., Li, X., Zhong, Q., Li, N., Zhang, K., Zhang, Y., Chu, H., Ma, C., Li, G.#, Zhao, J.#, and Gao, N#. (2019). Structural and functional insights into the tetrameric photosystem I from heterocyst-forming cyanobacteria. Nat Plants 5, 1087-1097.
27. Dong, D., Zheng, L., Lin, J., Zhang, B., Zhu, Y., Li, N., Xie, S., Wang, Y., Gao, N.#, and Huang, Z.# (2019). Structural basis of assembly of the human T cell receptor-CD3 complex. Nature 573, 546-552.
28. Ma, C., Wu, X., Sun, D., Park, E., Catipovic, M.A., Rapoport, T.A.#, Gao, N.#, and Li, L.# (2019). Structure of the substrate-engaged SecA-SecY protein translocation machine. Nature communications 10, 2872.
29. Zhu, X., Zhang, W., Guo, J., Zhang, X., Li, L., Wang, T., Yan, J., Zhang, F., Hou, B., Gao, N., Gao, G.F., and Zhou, X. (2019). Noc4L-Mediated Ribosome Biogenesis Controls Activation of Regulatory and Conventional T Cells. Cell Rep 27, 1205-1220 e1204.
30. Dong, L., Guan, X., Li, N., Zhang, F., Zhu, Y., Ren, K., Yu, L., Zhou, F., Han, Z., Gao, N., and Huang, Z. (2019). An anti-CRISPR protein disables type V Cas12a by acetylation. Nat Struct Mol Biol 26, 308-314.
31. Wang, J., Wang, J., Hu, M., Wu, S., Qi, J., Wang, G., Han, Z., Qi, Y., Gao, N., Wang, H.W., Zhou, J.M., and Chai, J. (2019). Ligand-triggered allosteric ADP release primes a plant NLR complex. Science 364, eaav5868.
32. Jiang, F., Li, N., Wang, X., Cheng, J., Huang, Y., Yang, Y., Yang, J., Cai, B., Wang, Y.P., Jin, Q.#, and Gao, N.# (2019). Cryo-EM Structure and Assembly of an Extracellular Contractile Injection System. Cell 177, 370-383 e315.
33. Yan, L., Wu, H., Li, X., Gao, N.#, and Chen, Z. # (2019). Structures of the ISWI-nucleosome complex reveal a conserved mechanism of chromatin remodeling. Nat Struct Mol Biol 26, 258-266.
34. Li, N., Lam, W.H., Zhai, Y.#, Cheng, J., Cheng, E., Zhao, Y., Gao, N.#, and Tye, B.K.# (2018). Structure of the origin recognition complex bound to DNA replication origin. Nature 559, 217-222.
35. Liang, K., Li, N., Wang, X., Dai, J., Liu, P., Wang, C., Chen, X.W., Gao, N., and Xiao, J. (2018). Cryo-EM structure of human mitochondrial trifunctional protein. Proceedings of the National Academy of Sciences of the United States of America 115, 7039-7044.
36. Yuan, Y., Du, C., Sun, C., Zhu, J., Wu, S., Zhang, Y., Ji, T., Lei, J., Yang, Y., Gao, N.#, and Nie, G.# (2018). Chaperonin-GroEL as a Smart Hydrophobic Drug Delivery and Tumor Targeting Molecular Machine for Tumor Therapy. Nano Lett 18, 921-928.
37. Yu, J., Zhang, B., Zhang, Y., Xu, C.Q., Zhuo, W., Ge, J., Li, J., Gao, N.#, Li, Y.#, and Yang, M.# (2018). A binding-block ion selective mechanism revealed by a Na/K selective channel. Protein & Cell 9, 629-639.
38. Li, Z., Ge, X., Zhang, Y., Zheng, L., Sanyal, S.#, and Gao, N.# (2018). Cryo-EM structure of Mycobacterium smegmatis ribosome reveals two unidentified ribosomal proteins close to the functional centers. Protein & Cell 9, 384-388.
39. Zhang, S., Li, N., Zeng, W., Gao, N.# and Yang, M.# (2017). Cryo-EM structures of the mammalian endo-lysosomal TRPML1 channel elucidate the combined regulation mechanism. Protein & Cell 8, 834-847.
40. Zhang, Y., Xiao, Z., Zou, Q., Fang, J., Wang, Q., Yang, X.#, and Gao, N.# (2017). Ribosome Profiling Reveals Genome-wide Cellular Translational Regulation upon Heat Stress in Escherichia coli. Genomics, Proteomics & Bioinformatics 15, 324-330.
41. Zhai, Y.#, Li, N., Jiang, H., Huang, X., Gao, N.#, and Tye, B.K.# (2017). Unique Roles of the Non-identical MCM Subunits in DNA Replication Licensing. Molecular Cell 67, 168-179.
42. Li, Z., Guo, Q., Zheng, L., Ji, Y., Xie, Y.-T., Lai, D.-H., Lun, Z.-R., Suo, X., and Gao, N.# (2017). Cryo-EM structures of the 80S ribosomes from human parasites Trichomonas vaginalis and Toxoplasma gondii. Cell Research 27, 1275-1288.
43. Peng, R., Xu, Y., Zhu, T., Li, N., Qi, J., Chai, Y., Wu, M., Zhang, X., Shi, Y., Wang, P.#, Wang, J.#, Gao, N.#, and Gao, G.F.# (2017). Alternate binding modes of anti-CRISPR viral suppressors AcrF1/2 to Csy surveillance complex revealed by cryo-EM structures. Cell Research 27, 853-864.
44. Biedka, S., Wu, S., LaPeruta, A.J., Gao, N., and Woolford, J.L., Jr. (2017). Insights into remodeling events during eukaryotic large ribosomal subunit assembly provided by high-resolution cryo-EM structures. RNA Biol 14, 1306-1313
45. Ma, C., Kurita, D., Li, N., Chen, Y., Himeno, H.#, and Gao, N.# (2017). Mechanistic insights into the alternative translation termination by ArfA and RF2. Nature 541, 550-553.
46. Ma, C., Wu, S., Li, N., Chen, Y., Yan, K., Li, Z., Zheng, L., Lei, J., Woolford, J.L., Jr.#, and Gao, N.# (2017). Structural snapshot of cytoplasmic pre-60S ribosomal particles bound by Nmd3, Lsg1, Tif6 and Reh1. Nat Struct Mol Biol 24, 214-220.
47. Zhai, Y., Cheng, E., Wu, H., Li, N., Yung, P.Y., Gao, N.#, and Tye, B.K.# (2017). Open-ringed structure of the Cdt1-Mcm2-7 complex as a precursor of the MCM double hexamer. Nat Struct Mol Biol 24, 300-308.
48. Li, N., Wu, J.X., Ding, D., Cheng, J., Gao, N.#, and Chen, L.# (2017). Structure of a Pancreatic ATP-Sensitive Potassium Channel. Cell 168, 101-110 e110.
49. Wu, S., Tan, D., Woolford, J.L., Jr., Dong, M.Q., and Gao, N.# (2017). Atomic modeling of the ITS2 ribosome assembly subcomplex from cryo-EM together with mass spectrometry-identified protein-protein crosslinks. Protein Science 26, 103-112.
50. Gu, J., Wu, M., Guo, R., Yan, K., Lei, J., Gao, N.#, and Yang, M.# (2016) The architecture of mammalian respirasome, Nature, 537, 639-643.
51. Kong, M.Y., Yan, K.G., Ma, C.Y., and Gao, N. # (2016). Distinct Binding and Enzymatic Activities of Two Ribosome-dependent NTPases YchF and YihA. Progress in Biochemistry and Biophysics 43, 570-578.
52. Tutuncuoglu, B., Jakovljevic, J., Wu, S., Gao, N., and Woolford, J.L., Jr. (2016). The N-terminal extension of yeast ribosomal protein L8 is involved in two major remodeling events during late nuclear stages of 60S ribosomal subunit assembly. RNA 22, 1386-1399.
53. Wu, S., Tutuncuoglu, B., Yan, K., Brown, H., Zhang, Y., Tan, D., Gamalinda, M., Yuan, Y., Li, Z., Jakovljevic, J., Ma, C., Lei, J., Dong, M.-Q., Woolford, J.L.#, and Gao, N.# (2016). Diverse roles of assembly factors revealed by structures of late nuclear pre-60S ribosomes. Nature 534, 133-137.
54. Du, W., Su, Q.P., Chen, Y., Zhu, Y., Jiang, D., Rong, Y., Zhang, S., Zhang, Y., Ren, H., Zhang, C., Wang, X., Gao, N., Wang, Y., Sun, L., Sun, Y., and Yu, L. (2016). Kinesin 1 Drives Autolysosome Tubulation. Developmental cell 37, 326-336.
55. Dong, D., Ren, K., Qiu, X., Zheng, J., Guo, M., Guan, X., Liu, H., Li, N., Zhang, B., Yang, D., Ma, C., Wang, S., Wu, D., Ma, Y., Fan, S., Wang, J., Gao, N., and Huang, Z. (2016). The crystal structure of Cpf1 in complex with CRISPR RNA. Nature 532, 522-526.
56. Ding, Y.H., Fan, S.B., Li, S., Feng, B.Y., Gao, N., Ye, K., He, S.M., and Dong, M.Q. (2016). Increasing the Depth of Mass-Spectrometry-Based Structural Analysis of Protein Complexes through the Use of Multiple Cross-Linkers. Analytical chemistry 88, 4461-4469.
57. Tan, D., Li, Q., Zhang, M.-J., Liu, C., Ma, C., Zhang, P., Ding, Y.-H., Fan, S.-B., Tao, L., Yang, B., Li, X., Ma, S., Liu, J., Feng, B., Liu, X., Wang, H.-W., He, S.-M., Gao, N., Ye, K., Dong, M.-Q., and Lei, X. (2016). Trifunctional cross-linker for mapping protein-protein interaction networks and comparing protein conformational states. eLife 5, e12509.
58. Ma, C., Yan, K., Tan, D., Li, N., Zhang, Y., Yuan, Y., Li, Z., Dong, M.Q., Lei, J., and Gao, N.# (2016). Structural dynamics of the yeast Shwachman-Diamond syndrome protein (Sdo1) on the ribosome and its implication in the 60S subunit maturation. Protein & Cell 7, 187-200.
59. Zhang, D., Yan, K., Liu, G., Song, G., Luo, J., Shi, Y., Cheng, E., Wu, S., Jiang, T., Lou, J., Gao, N. #, and Qin, Y.# (2016). EF4 disengages the peptidyl-tRNA CCA end and facilitates back-translocation on the 70S ribosome. Nat Struct Mol Biol 23, 125-131.
60. Zhang, J., Pan, X., Yan, K, Sun, S., Gao, N.# and Sui, S-F#. (2015). Mechanisms of ribosome stalling by SecM at multiple elongation steps. eLife, DOI: 10.7554/eLife.09684
61. Zhang, Y., Mandava, C.S., Cao, W., Li, X., Zhang, D., Li, N., Zhang, Y., Zhang, X., Qin, Y., Mi, K., Lei, J.#, Sanyal, S.#, and Gao, N.# (2015). HflX is a ribosome-splitting factor rescuing stalled ribosomes under stress conditions. Nat Struct Mol Biol 22, 906-913.
62. Zhang, D., Yan, K., Zhang, Y., Liu, G., Cao, X., Song, G., Xie, Q.#, Gao, N.#, and Qin, Y.# (2015). New insights into the enzymatic role of EF-G in ribosome recycling. Nucleic Acids Res 43, 10525-10533.
63. Ge, J., Li, W., Zhao, Q., Li, N., Chen, M., Zhi, P., Li, R., Gao, N.#, Xiao, B.#, and Yang, M.# (2015). Architecture of the mammalian mechanosensitive Piezo1 channel. Nature 527, 64-69.
64. Mi, N., Chen, Y., Wang, S., Chen, M., Zhao, M., Yang, G., Ma, M., Su, Q., Luo, S., Shi, J., Xu, J., Guo, Q., Gao, N., Sun, Y., Chen, Z., and Yu, L. (2015). CapZ regulates autophagosomal membrane shaping by promoting actin assembly inside the isolation membrane. Nat Cell Biol 17, 1112-1123.
65. Li, N., Zhai, Y.#, Zhang, Y., Li, W., Yang, M., Lei, J., Tye, B.K.#, and Gao, N#. (2015). Structure of the eukaryotic MCM complex at 3.8 A. Nature 524, 186-191.
66. Sun, C., Yuan, Y., Xu, Z., Ji, T., Tian, Y., Wu, S., Lei, J., Li, J.#, Gao, N.#, and Nie, G#. (2015). Fine-tuned H-ferritin nanocage with multiple gold clusters as near-infrared kidney specific targeting nanoprobe. Bioconjugate chemistry 26, 193-196.
67. Zhang, Y., Ma, C., Yuan, Y., Zhu, J., Li, N., Chen, C., Wu, S., Yu, L., Lei, J.#, and Gao, N.# (2014). Structural basis for interaction of a cotranslational chaperone with the eukaryotic ribosome. Nat Struct Mol Biol 21, 1042-1046.
68. Zhang, X., Yan, K., Zhang, Y., Li, N., Ma, C., Li, Z., Zhang, Y., Feng, B., Liu, J., Sun, Y., Xu, Y., Lei, J.#, and Gao, N.# (2014). Structural insights into the function of a unique tandem GTPase EngA in bacterial ribosome assembly. Nucleic Acids Res 42, 13430-13439.
69. Feng, B., Mandava, C.S., Guo, Q., Wang, J., Cao, W., Li, N., Zhang, Y., Zhang, Y., Wang, Z., Wu, J., Sanyal, S.#, Lei, J.#, and Gao, N.# (2014). Structural and functional insights into the mode of action of a universally conserved Obg GTPase. PLoS Biol 12, e1001866.
70. Yang, Z., Guo, Q., Goto, S., Chen, Y., Li, N., Yan, K., Zhang, Y., Muto, A., Deng, H., Himeno, H., Lei, J.#, and Gao, N.# (2014). Structural insights into the assembly of the 30S ribosomal subunit in vivo: functional role of S5 and location of the 17S rRNA precursor sequence. Protein Cell 5, 394-407.
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Laboratory Introduction