师资队伍
舒校坤
化学生物学
博士/教授/博导
shuxiaokun@fudan.edu.cn；shu2010x@gmail.com
上海市杨浦区淞沪路2005号复旦大学江湾校区化学楼
研究兴趣
分子生物技术与生命逻辑解析：以物理化学“第一性原理”驱动新一代荧光探针及化学遗传学工具设计为研究主线，旨在实现生命体内细胞信号网络的高时空分辨成像与动态操控，解码生命系统运行的底层逻辑，探索生物大分子凝聚体（相分离）在生理与病理过程中的作用机制，以此驱动针对“不可成药”靶点的原创药物研发与精准治疗。长期致力于物理、化学与生物学的深度交叉研究。
教育经历
University of Oregon，博士，生物物理，导师：Jim Remington 教授（2003-2007）
复旦大学，硕士，凝聚态物理，导师：蒋平 教授，车静光 教授（2000-2003）
四川大学，学士，理论物理（1996-2000）
工作经历
复旦大学，相辉学者，特聘教授，“百利天恒”冠名教授，化学生物技术智能研究所iCOBRA（Institute of Chemical and Open Biotechnology Research and Application）创始所长（2026.3-至今）
University of California – San Francisco，Herfindal Endowed Chair讲席教授 任命（2026）
University of California – San Francisco，药学院, 药物化学系，教授（2010-2026）
Howard Hughes Medical Institute，University of California – San Diego, Research Fellow，合作导师：钱永健（Roger Y. Tsien）教授（2008诺贝尔奖获得者）（2007-2010）
University of Oregon，Department of Physics, Institute of Molecular Biology, 助理研究员, 合作导师：Jim Remington 教授（2004-2007）
主持美国NIH项目（*为共同主持）
项目类别
项目年限
经费(万, 美元）
NCI R01: EML4-ALK凝聚体如何驱动肿瘤*
2023-2028
335
NIH 4DN U01: 蛋白质凝聚体操控与功能研究*
2020-2025
543
NCI R01: 癌蛋白相分离高分辨成像与操控
2021-2026
215
Maximizing Investigator Research Award
2019-2029
591
NIGMS R01: 激酶活性荧光探针设计与应用
2018-2022
136
NIGMS R01: 蛋白酶活性荧光探针设计与应用
2015-2019
125
NIH Director New Innovator Award
2012-2017
236
近三年代表性学术论文及成就（创新性、影响力、与系统性）
注：*为通讯作者，+为共同作者
2023-2026以通讯作者在Nat. Struct. Mol. Biol., Nat. Chem. Biol., Nat. Microbiol., 和PNAS，ACS Cent. Sci. 等领域内顶级期刊和高水平综合期刊上发表多篇具有重大影响力/里程碑学术论文。
简要总结：1）成像工具创新：基于热力学理论（如Flory-Huggins Theory）开发了系列原创性分子探针及化学遗传学工具，突破了对活细胞内信号传导与生物大分子凝聚体高时空分辨成像与精准调控的技术瓶颈。2）疾病机制解析：首次解析了细胞信号与生物大分子相分离驱动肿瘤发生的底层分子机制，为理解复杂生命现象与重大疾病机制提供了物理化学视角。3）新靶点发现：基于上述原创机制，定义并验证了多个具有转化潜力的新型药靶。4）临床转化：结合自研的原创工具，开展针对“不可成药”靶点在内的新型药靶的创制性药物研发，打通了从基础物理化学原理到临床转化应用的全链条。
1) Li X+, Tan SK+, Chung CI+, Hatstat AK, Zhao Q, Luo J, DeGrado WF*, Shu X*. Design of an ultrabright biosensor for dynamic imaging of kinase activity in cells. Proc Natl Acad Sci U S A. 2026. In Press
Innovation (创新): 基于热力学原理并结合蛋白质从头设计开发了激酶活性分子探针NOVARK，对活细胞内的激酶信号进行高时空分辨成像。
Impact (影响): 为基于蛋白质从头设计原理开发激酶活性探针铺平了道路。这些探针可用于对活细胞内的激酶信号进行读取，从而解析生命系统的运行逻辑。
系统性：NOVARK激酶活性探针是对本课题组在2018年基于相分离热力学原理设计的全新激酶活性探针SPARK (发表于Mol. Cell) 的一次最新拓展。这些激酶活性探针是我们在实现生命体内细胞信号网络的高时空分辨成像工具开发方面系统性工作的一部分，其它还包括：1）蛋白酶活性探针，比如2015年开发的蛋白酶近红外荧光探针iCasper (发表于PNAS)；2018年开发的蛋白酶绿色荧光探针 FlipGFP (发表于JACS)；2）蛋白质相互作用分子探针（见下一页的系统性工作介绍）。
2) Yang J+, Chung CI+, Koach J, Liu H, Navalkar A, He H, Ma Z, Zhao Q, Yang X, He L, Mittag T, Shen Y, Weiss WA, Shu X*. MYC phase separation selectively modulates the transcriptome. Nat Struct Mol Biol. 2024 May 29. PMID: 38811792 PMCID: PMC11479839
Innovation (创新): 首次发现相分离本身对MYC靶向基因的调控具有高度选择性，对癌细胞增殖起关键作用。
Impact (影响): 定义并验证了MYC相分离是一个全新的潜在药物靶点，其优点是相分离抑制剂具有高度选择性。由于MYC对70%肿瘤发生或发展、以及侵袭、转移、耐药、重塑肿瘤微环境、免疫逃逸密切相关，此发现对肿瘤分子机制研究具有重大指导意义，并且提出了针对“不可成药”靶点药物研发的新范式。
系统性：此项工作，结合我们在致癌融合转录因子YAP-MAML2上的工作，共同揭示了相分离的全新生物学功能，加深了对生物大分子凝聚体及相分离在生理与病理过程中的作用机制理解，为研发“不可成药”靶点药物提供了全新理论依据。
3) Li X, Combs D, Salaita K, Shu X*. Polarized activity of the focal adhesion kinase within single focal adhesions during cell migration. Nat. Chem. Biol. 2023 Jun 22
Innovation (创新): 利用热力学的相分离机制开发了基因编码的黏着斑激酶（FAK）活性分子探针，能在活细胞和脊椎动物中可视化内源FAK 活性，并且具有高时空分辨率。首次揭示了 FAK在迁移细胞前缘高度极化的全新特点。
Impact (影响): 阐明了 FAK 如何充当力学传感器，在细胞迁移的黏着斑更新中扮演重要角色，对理解细胞迁移包括癌细胞在肿瘤中的迁移机制、及开发新型抑制剂具有重大指导意义。此探针对探索生命体运行的底层逻辑有重大价值。
4) Chung CI, Yang J, Yang X, Liu H, Ma Z, Szulzewsky F, Holland EC, Shen Y, Shu X*. Phase separation of YAP-MAML2 differentially regulates the transcriptome. Proc Natl Acad Sci U S A. 2024. 121(7): e2310430121.
Innovation (创新): 首次揭示致癌融合转录因子YAP-MAML2发生相分离并形成具有转录活性的生物大分子凝聚体。基于热力学Flory-Huggins Theory，通过增加溶质与溶剂的相互作用，开发了可逆的化学遗传学工具，发现相分离并非全面调控所有受控基因，而是高度选择性地影响一小部分目标基因。
Impact (影响): 加深了对生物分子凝聚体在基因转录中的作用理解，表明相分离在不同靶基因的选择性调控中扮演独特角色。
系统性：此次研究开发的化学遗传学工具 (SPARK-OFF)，可以精准溶解转录因子凝聚体。SPARK-OFF是我们在实现对生命体内对生物大分子凝聚体相分离操控工具开发方面系统性工作的一部分，另一个与之互补的工具是SPARK-ON，可以精准诱导转录因子相分离，形成凝聚体。相较于传统方法，我们工具的优势是可以在不引入突变、且不改变蛋白质丰度情况下精准操控相分离，广泛适用于生物大分子凝聚体的研究，对理解相分离的生物学功能具有重大意义。
5) Chung CI, Yang J, Shu X*. Chemogenetic Minitool for Dissecting the Roles of Protein Phase Separation. ACS Cent Sci. 2023 Jul 26; 9(7):1466-1479.
Innovation (创新): 基于热力学Flory-Huggins Theory，通过增加溶质与溶质相互作用，首次开发了能够精准诱导特定蛋白质相分离的化学遗传学工具(SPARK-ON), 用于探索生物分子凝聚物中相分离如何调节转录因子的基因转录活性。
Impact (影响): SPARK-ON 可以在不引入突变、且不改变蛋白质丰度的情况下诱导相分离，为深入研究多种转录因子及其他生物分子凝聚物的相分离生物学功能提供了一个强有力的工具。克服了传统研究相分离方法的致命缺陷-通常依赖突变，但突变往往会改变转录因子自身的活性，导致难以区分是相分离还是突变直接影响了转录。
6) Yang J, Xiao Y, Lidsky PV, Wu CT, Bonser LR, Peng S, Garcia-Knight MA, Tassetto M, Chung CI, Li X, Nakayama T, Lee IT, Nayak JV, Ghias K, Hargett KL, Shoichet BK, Erle DJ, Jackson PK, Andino R*, Shu X*. Fluorogenic reporter enables identification of compounds that inhibit SARS-CoV-2. Nat Microbiol. 2023 Jan; 8(1):121-134.
Innovation (创新): 基于全新三级结构的荧光蛋白UnaG（不同于GFP)，开发了名为SURF的可逆的蛋白质-蛋白质相互作用荧光探针。证明SURF适用于高通量筛选，并成功发现了三种能阻断病毒在小鼠模型中复制的天然化合物。
Impact (影响): SURF系统首次能同时具有高时空分辨率、无须外源辅因子、由暗到亮荧光变化、能定量检测蛋白质之间的相互作用强度、定量测量抑制剂的活性。已成功应用于筛选“不可成药”靶点的抑制剂，包括MYC（未发表）。对在活细胞中研究蛋白质相互作用及开发抑制剂或结合剂具有重大应用价值。
系统性：SURF是我们在设计蛋白互作分子探针方面系统性工作的最新进展，前几项工作包括：1）2014年基于Singlet oxygen-mediated Triplet Energy Transfer原理的STET探针，突破了FRET在探测距离上的限制；2）2018年基于相分离原理设计的SPPIER (Separation of Phases-based Protein-protein Interaction Reporter) 探针，比其它基于GFP的探针在亮度上提升了10倍，而且其信号图案极易检测。
完整文章列表
注：*为通讯作者，+为共同作者
以第一或通讯作者在 Science, Molecular Cell, Nat. Struct. Mol. Biol., Nat. Methods, Nat. Chem. Biol., Nat. Microbiol. 等顶刊及其子刊发表多篇里程碑学术论文。
1. Shu X, Jiang P, Che J (2003) Surface alloying of immiscible metals induced by surface state shift. Surface Science 545, 199-210
2. Quillin ML, Anstrom DM, Shu X, O'Leary S, Kallio K, Chudakov DM, Remington SJ (2005) Kindling Fluorescent Protein from Anemonia sulcata: Dark-State Structure at 1.38 Å Resolution. Biochemistry 44, 5774-5787
3. Shu X, Shaner NC, Yarbrough CA, Tsien RY, Remington SJ. Novel chromophores and buried charges control color in mFruits. Biochemistry. 2006 Aug 15; 45(32):9639-47. PMID: 16893165
4. Shu X, Kallio K, Shi X, Abbyad P, Kanchanawong P, Childs W, Boxer SG, Remington SJ. (2007) Ultrafast Excited State Dynamics in The Green Fluorescent Protein Variant S65T/H148D. 1. Mutagenesis and Structural Studies. Biochemistry 46, 12005-12013
5. Shi X, Abbyad P, Shu X, Kallio K, Kanchanawong P, Childs W, Remington SJ, Boxer SG. (2007) Ultrafast Excited State Dynamics in The Green Fluorescent Protein Variant S65T/H148D. 2. Unusual Photophysical Properties. Biochemistry 46, 120014-12025
6. Leiderman P, Genosar L, Huppert D, Shu X, Remington SJ, Solntsev KM, Tolbert LM (2007) Ultrafast Excited State Dynamics in The Green Fluorescent Protein Variant S65T/H148D. 3. Short- and Long-time Dynamics of the Excited-state Proton Transfer. Biochemistry 46, 120026-12036
7. Shu X, Leiderman P, Gepshtein R, Smith NR, Kallio K, Huppert D, Remington SJ (2007) An alternative excited state proton transfer pathway in a green fluorescent protein variant S205V. Protein Science 16, 2703-2710
8. Shu X, Wang L, Colip L, Kallio K, Remington SJ (2009) Unique Interactions between the Chromophore and Glutamate 16 Lead to Far-red Emission in a Red Fluorescent Protein. Protein Science 18, 460-466
9. Shu X, Royant A, Lin MZ, Aguilera TA, Lev-Ram V, Steinbach PA, Tsien RY. Mammalian expression of infrared fluorescent proteins engineered from a bacterial phytochrome. Science. 2009 May 8; 324(5928):804-7. PMID: 19423828
10. TJ Deerinck, EA Bushong, V Lev-Ram, X Shu, RY Tsien, MH Ellisman. Enhancing Serial Block-Face Scanning Electron Microscopy to Enable High Resolution 3-D Nanohistology of Cells and Tissues. Microscopy and Microanalysis. 2010 Jul 1; 16(S2):1138-1139
11. Shu X, Lev-Ram V, Deerinck TJ, Qi Y, Ramko EB, Davidson MW, Jin Y, Ellisman MH, Tsien RY. A genetically encoded tag for correlated light and electron microscopy of intact cells, tissues, and organisms. PLoS Biol. 2011 Apr; 9(4):e1001041. PMID: 21483721
12. M Ellisman, X Shu, V Lev-Ram, T Deerinck, R Tsien, S Lamont, J Martinez, M Berlanga, E Bushong, M Martone, S Larson. Bridging Gaps in Imaging by Applying EM Tomography and Serial Block Face SEM, Including a New Genetically Encoded Tag for Correlated Light and 3D Electron Microscopy of Intact Cells, Tissues and Organisms: Integrating the Resulting Correlated Image Data Using the Whole Brain Catalog. Microscopy and Microanalysis. 2011 Jul 1; 17(S2):2-3
13. Ellisman MH, Deerinck TJ, Shu X, Sosinsky GE. Picking faces out of a crowd: genetic labels for identification of proteins in correlated light and electron microscopy imaging. Methods Cell Biol. 2012; 111:139-55. PMID: 228579276/8
14. T.J. Deerinck, E. Bushong, R. Tsien, M. Ellisman, X. Shu. Advances in Correlated Light and Electron Microscopy Imaging Probes. Microscopy and Microanalysis. 2012 Jul 1; 18(S2):834-835
15. Qi YB, Garren EJ, Shu X, Tsien RY, Jin Y. Photo-inducible cell ablation in Caenorhabditis elegans using the genetically encoded singlet oxygen generating protein miniSOG. Proc Natl Acad Sci U S A. 2012 May 8; 109(19):7499-504. PMID: 22532663
16. Ou HD, Kwiatkowski W, Deerinck TJ, Noske A, Blain KY, Land HS, Soria C, Powers CJ, May AP, Shu X, Tsien RY, Fitzpatrick JA, Long JA, Ellisman MH, Choe S, O'Shea CC. A Structural Basis for the Assembly and Functions of a Viral Polymer that Inactivates
Multiple Tumor Suppressors. Cell. 2012 Oct 12; 151(2):304-19. PMID: 23063122
17. Butko MT, Yang J, Geng Y, Kim HJ, Jeon NL, Shu X, Mackey MR, Ellisman MH, Tsien RY, Lin MZ. Fluorescent and photo-oxidizing TimeSTAMP tags track protein fates in light and electron microscopy. Nat Neurosci. 2012 Nov; 15(12):1742-51. PMID: 23103964
18. Asensio CS, Sirkis DW, Maas JW, Egami K, To TL, Brodsky FM, Shu X, Cheng Y, Edwards RH. Self-Assembly of VPS41 Promotes Sorting Required for Biogenesis of the Regulated Secretory Pathway. Dev Cell. 2013 Nov 25; 27(4):425-37. PMID: 24210660
19. Yu D, Gustafson WC, Han C, Lafaye C, Noirclerc-Savoye M, Ge WP, Thayer DA, Huang H, Kornberg TB, Royant A, Jan LY, Jan YN, Weiss WA, Shu X. An improved monomeric infrared fluorescent protein for neuronal and tumour brain imaging. Nat Commun. 2014; 5:3626. PMID: 24832154
20. To TL, Fadul MJ, Shu X. Singlet oxygen triplet energy transfer-based imaging technology for mapping protein-protein proximity in intact cells. Nat Commun. 2014; 5:4072. PMID: 24905026
21. To TL, Piggott BJ, Makhijani K, Yu D, Jan YN, Shu X. Rationally designed fluorogenic protease reporter visualizes spatiotemporal dynamics of apoptosis in vivo. Proc Natl Acad Sci U S A. 2015 Mar 17; 112(11):3338-43. PMID: 25733847
22. Yu D, Baird MA, Allen JR, Howe ES, Klassen MP, Reade A, Makhijani K, Song Y, Liu S, Murthy Z, Zhang SQ, Weiner OD, Kornberg TB, Jan YN, Davidson MW, Shu X. A naturally monomeric infrared fluorescent protein for protein labeling in vivo. Nat Methods. 2015 Aug; 12(8):763-5. PMID: 26098020
23. Yu D, Dong Z, Gustafson WC, Ruiz-González R, Signor L, Marzocca F, Borel F, Klassen MP, Makhijani K, Royant A, Jan YN, Weiss WA, Guo S, Shu X. Rational design of a monomeric and photostable far-red fluorescent protein for fluorescence imaging in vivo. Protein Sci. 2016 Feb;25(2):308-15. PMID: 26549191
24. To TL, Zhang Q, Shu X. Structure-guided design of a reversible fluorogenic reporter of protein-protein interactions. Protein Sci. 2016 Mar;25(3):748-53. PMID: 26690964
25. To TL, Medzihradszkya KF, Burlingame AL, DeGrado WF, Jo H, Shu X. Photoactivatable protein labeling by singlet oxygen mediated reactions. Bioorg Med Chem Lett. 2016 26, 3359-3363
26. To TL, Schepis A, Ruiz-González R, Zhang Q, Yu D, Dong Z, Coughlin SR, Shu X. Rational design of a GFP-based fluorogenic caspase reporter for imaging apoptosis in vivo. Cell Chem Biol. 2016 Jul 21; 23(7): 875-882
27. Feliks M, Lafaye C, Shu, X, Royant A, Field M. Structural Determinants of Improved Fluorescence in a family of Bacteriophytochrome-Based Infrared Fluorescent Proteins: Insights from Continuum Electrostatic Calculations and Molecular Dynamics Simulations. Biochemistry. 2016 Jul 29 (Epub ahead of print)
28. Rodriguez EA, Tran GN, Gross LA, Crisp JL, Shu X, Lin JY, Tsien RY. A far-red fluorescent protein evolved from a cyanobacterial phycobiliprotein. Nat Methods. 2016 Aug 1. doi: 10.1038/nmeth.3935. [Epub ahead of print]
29. Rodriguez EA*, Campbell RE*, Lin JY*, Lin MZ*, Miyawaki A*, Palmer AE*, Shu X*, Zhang J*, Tsien RY*. The growing and glowing toolbox of fluorescent and photoactive 7/8proteins. Trends in Biochemical Sciences. 2016 Nov 1. pii: S0968-0004(16)30173-6.
*Co-corresponding authors
30. Makhijani K, To TL, Ruiz-González R, Lafaye C, Royant A, Shu X. Precision optogenetic tool for single-cell and multiple-cell ablation in a live animal model system. Cell Chem Biol. 2017 2017 Jan 19;24(1):110-119
31. To TL, Shu X. Detecting Activity at Different Length Scales: From Subdiffraction to Whole-Animal Activity. Biochemistry 2017 Oct 3;56(39):5163-5164
32. Zhang Q, Huang H, Zhang L, Wu R, Chung CI, Zhang SQ, Torra J, Schepis A, Coughlin SR, Kornberg TB, Shu X. (2018) Visualizing dynamics of cell signaling in vivo with a phase separation-based kinase reporter. Mol Cell 69, 334-346. PMCID: PMC5788022
33. Zhang SQ, Huang H, Yang J, Kratochvil HT, Lolicato M, Liu Y, Shu X*, Liu L*, DeGrado WF*. Designed peptides that assemble into cross-a amyloid-like structures. Nat Chem Biol. 2018 Sep; 14(9):870-875. PMID: 30061717. *Co-corresponding authors
34. Zhang Q, Zheng YW, Coughlin SR, Shu X. (2018) A rapid fluorogenic GPCR - betaarrestin interaction assay. Protein Sci. (Cover Story). 27, 874-79. PMCID: PMC5866937
35. Chung CI, Zhang Q, Shu X. Dynamic Imaging of Small Molecule Induced Protein-Protein Interactions in Living Cells with a Fluorophore Phase Transition Based Approach. Anal Chem. 2018 Dec 18; 90(24):14287-14293. PMCID: PMC6298840
36. Torra J, Lafaye C, Signor L, Aumonier S, Flors C, Shu X, Nonell S, Gotthard G, Royant A. Tailing miniSOG: structural bases of the complex photophysics of a flavin-binding singlet oxygen photosensitizing protein. Sci Rep. 2019 02 20; 9(1):2428. PMID: 30787421. PMCID: PMC6382843
37. Zhang Q, Schepis A, Huang H, Yang J, Ma W, Torra J, Zhang SQ, Yang L, Wu H, Nonell S, Dong Z, Kornberg TB, Coughlin SR, Shu X. Designing a Green Fluorogenic Protease Reporter by Flipping a Beta Strand of GFP for Imaging Apoptosis in Animals. J Am Chem Soc. 2019 03 20; 141(11):4526-4530. PMID: 30821975. PMCID: PMC6486793
38. Shu X. Imaging dynamic cell signaling invivo with new classes of fluorescent reporters. Curr Opin Chem Biol. 2020 Feb; 54:1-9. PMID: 31678813. PMCID: PMC7131868
39. Guo WH, Qi X, Yu X, Liu Y, Chung CI, Bai F, Lin X, Lu D, Wang L, Chen J, Su LH, Nomie KJ, Li F, Wang MC, Shu X, Onuchic JN, Woyach JA, Wang ML, Wang J. Enhancing intracellular accumulation and target engagement of PROTACs with reversible covalent chemistry. Nat Commun. 2020 08 26; 11(1):4268. PMID: 32848159. PMCID: PMC7450057
40. Li X, Lidsky P, Xiao Y, Wu CT, Garcia-Knight M, Yang J, Nakayama T, Nayak JV, Jackson PK, Andino R, Shu X. Ethacridine inhibits SARS-CoV-2 by inactivating viral particles. PLoS Pathog. 2021 Sep 03; 17(9):e1009898. PMID: 34478458. PMCID: PMC8445489
41. Lafaye C, Aumonier S, Torra J, Signor L, von Stetten D, Noirclerc-Savoye M, Shu X, Ruiz-González R, Gotthard G, Royant A, Nonell S. Riboflavin-binding proteins for singlet oxygen production. Photochem Photobiol Sci. 2022 Sep; 21(9):1545-1555. PMID: 35041199
42. Yang J, Xiao Y, Lidsky PV, Wu CT, Bonser LR, Peng S, Garcia-Knight MA, Tassetto M, Chung CI, Li X, Nakayama T, Lee IT, Nayak JV, Ghias K, Hargett KL, Shoichet BK, Erle DJ, Jackson PK, Andino R, Shu X. Fluorogenic reporter enables identification of compounds that inhibit SARS-CoV-2. Nat Microbiol. 2023 Jan; 8(1):121-134. PMID: 36604514. PMCID: PMC9831272
43. Li X, Combs JD, Salaita K, Shu X. Polarized focal adhesion kinase activity within a focal adhesion during cell migration. Nat Chem Biol. 2023 Dec; 19(12):1458-1468. PMID: 37349581. PMCID: PMC10732478
44. Li X, Chung CI, Yang J, Chaudhuri S, Munster PN, Shu X. ATM-SPARK: A GFP phase separation-based activity reporter of ATM. Sci Adv. 2023 Mar; 9(9):eade3760. PMID: 36857446. PMCID: PMC99771818/8
45. Dekker J, Alber F, Aufmkolk S, Beliveau BJ, Bruneau BG, Belmont AS, Bintu L, Boettiger A, Calandrelli R, Disteche CM, Gilbert DM, Gregor T, Hansen AS, Huang B, Huangfu D, Kalhor R, Leslie CS, Li W, Li Y, Ma J, Noble WS, Park PJ, Phillips-Cremins JE, Pollard KS, Rafelski SM, Ren B, Ruan Y, Shav-Tal Y, Shen Y, Shendure J, Shu X, Strambio-DeCastillia C, Vertii A, Zhang H, Zhong S. Spatial and temporal organization of the genome: Current state and future aims of the 4D nucleome project. Mol Cell. 2023 08 03;
83(15):2624-2640. PMID: 37419111. PMCID: PMC10528254
46. Chung CI, Yang J, Shu X. Chemogenetic Minitool for Dissecting the Roles of Protein Phase Separation. ACS Cent Sci. 2023 Jul 26; 9(7):1466-1479. PMID: 37521779. PMCID: PMC10375881
47. To TL, Li X, Shu X. Spying on SARS-CoV-2 with Fluorescent Tags and Protease Reporters. Viruses. 2023 09 27; 15(10). PMID: 37896782. PMCID: PMC10612051
48. Chung CI, Yang J, Yang X, Liu H, Ma Z, Szulzewsky F, Holland EC, Shen Y, Shu X. Phase separation of YAP-MAML2 differentially regulates the transcriptome. Proc Natl Acad Sci U S A. 2024 Feb 13; 121(7):e2310430121. PMID: 38315854. PMCID: PMC10873646
49. Yang J, Chung CI, Koach J, Liu H, Navalkar A, He H, Ma Z, Zhao Q, Yang X, He L, Mittag T, Shen Y, Weiss WA, Shu X. MYC phase separation selectively modulates the transcriptome. Nat Struct Mol Biol. 2024 May 29. PMID: 38811792 PMCID: PMC11479839
50. Wu G, Ma T, Hancock CE, Gonzalez S, Aryal B, Vaz S, Chan G, Palarca-Wong M, Allen N, Chung CI, Shu X, Liu Q. Opposing GPCR signaling programs protein intake setpoint in Drosophila. Cell. 2024 Aug 20. PMID: 39197448. PMCID: PMC11437785
51. Meyer K, Yserentant K, Cheloor-Kovilakam R, Ruff KM, Chung CI, Shu X, Huang B, Weiner OD. YAP charge patterning mediates signal integration through transcriptional co-condensates. Nat Commun. 2025 Aug 12; 16(1):7454. PMID: 40796733. PMCID: PMC12343768
52. Li X+, Tan SK+, Chung CI+, Hatstat AK, Zhao Q, Luo J, DeGrado WF*, Shu X*. Design of an ultrabright biosensor for dynamic imaging of kinase activity in cells. Proc Natl Acad Sci U S A. 2026. In Press
Book Chapter:
1. Tsz-Leung To, Xiaokun Shu. Chapter 4 Major methods and technologies for assessing cell death. 2022 Jan 1; 93-118
