#：corresponding author.

Undergraduate students (hightlighted in purple)

Independent Career:

2026
Membrane-associated periodic skeleton regulates major forms of endocytosis in neurons through a signaling-driven positive feedback loop.

37. Membrane-associated periodic skeleton regulates major forms of endocytosis in neurons through a signaling-driven positive feedback loop.

J Fei , Y Zheng , C LaLonde , Y Tao , R Zhou#. Science Advances 12, eaeb0803 (2026)

Highlighted in Penn State News, EurekAlert, Medical Xpress

Journal cover
Actin depolymerization promotes axon regeneration by restoring axonal mitochondria transport in mouse models of optic neuropathy.

36. Actin depolymerization promotes axon regeneration by restoring axonal mitochondria transport in mouse models of optic neuropathy.

L Li , X Feng , F Fang , J Fei , J Lu , H Zhang , M Ždímalová , Y Yuan , Z Luo , H Huang , D Liu , F Huang , Y Zeng , F Bian , I.J You , L Liu , X Yan , M Li , Y Lin , Q Xie , A Krish , X Duan , J Goldberg , X Chen , M Braun , Z Lansky , Y Han , R Zhou , Y Hu#. Science Translational Medicine 18,eadw0908 (2026)

Journal cover
Coordination of cell organelles to promote metabolon formation.

35. Coordination of cell organelles to promote metabolon formation.

Z Sha , AM Pedley , TD Iles , S Zhang , JR Staub , R Zhou , SJ Benkovic# Proc. Natl. Acad. Sci. 123 (5) e2532504123 (2026)

Tunable Chemical and Optical Control of ER-Plasma Membrane Contact Site Geometry and Dynamics with High-Fidelity Visualization

34. Tunable Chemical and Optical Control of ER-Plasma Membrane Contact Site Geometry and Dynamics with High-Fidelity Visualization

S Zhang , J Fei , Y Zheng , R Zhou#. bioRxiv, under review at JACS Au (2026)

A Chemically Inducible Multimerization System for Tunable and Background-Free RTK Activation

33. A Chemically Inducible Multimerization System for Tunable and Background-Free RTK Activation

Y Zheng , J Fei , A Chakrabarti , R Zhou#. Analytical Chemistry 98 (2), 1328-1341 (2026)

"Zipper" Grammar of CTD Governs the Spatial Programing of the Transcription Cycle of RNA Polymerase II

32. "Zipper" Grammar of CTD Governs the Spatial Programing of the Transcription Cycle of RNA Polymerase II

Q Zhang , HA Hardtke , Y Zheng , A Gerber , MKV Ramani , E Escobar , J Brodbelt , R Zhou , YJ Zhang# bioRxiv, in revision at Nature Communications (2026)

2025
Membrane-associated periodic skeleton regulates major forms of endocytosis in neurons through a signaling-driven positive feedback loop.

31. Membrane-associated periodic skeleton regulates major forms of endocytosis in neurons through a signaling-driven positive feedback loop.

J Fei , Y Zheng , C LaLonde , Y Tao , R Zhou#. bioRxiv (2025)

CLaSSiNet: A Computational Framework for Quantitative Classification of Nanoscale Network Architectures from Super-Resolution Microscopy.

30. CLaSSiNet: A Computational Framework for Quantitative Classification of Nanoscale Network Architectures from Super-Resolution Microscopy.

Y Tao , R Zhou#. bioRxiv, under revision at JACS Au (2025)

Roles of histone chaperone Nap1 and histone acetylation in regulating phase-separation of chromatin arrays.

29. Roles of histone chaperone Nap1 and histone acetylation in regulating phase-separation of chromatin arrays.

J Gao , H Li , S Tan , R Zhou , TH Lee#. Nature Communications 16, 10672 (2025)

A chemically inducible multimerization system for tunable and background-free RTK activation.

28. A chemically inducible multimerization system for tunable and background-free RTK activation.

Y Zheng , J Fei , A Chakrabarti , R Zhou#. bioRxiv (2025)

FLEXTAG:A Small and Self-renewable Protein Labeling System for Anti-fading Multi-color Super-resolution Imaging.

27. FLEXTAG:A Small and Self-renewable Protein Labeling System for Anti-fading Multi-color Super-resolution Imaging.

H Zhang , Y Yao , X Wang , Y Zheng , S Zhang , Y Tao , H Yang , Y Wang , M Liu , M Feric , R Zhou#. bioRxiv, Accepted at Nature Communications (2025)

2023
Watching biomolecules stride in real time.

26. Watching biomolecules stride in real time.

J Fei , R Zhou#. Science 379, 986-987 (2023)

2022
Proteomic and functional analyses of the periodic membrane skeleton in neurons.

25. Proteomic and functional analyses of the periodic membrane skeleton in neurons.

R Zhou# , B Han , R Nowak , Y Lu , E Heller , C Xia , AH Chishti , VM Fowler , X Zhuang#. Nature Communications 13, 3196 (2022)

Prior to Penn State:

Selected publications:

Membrane-associated periodic skeleton is a signaling platform for RTK transactivation in neurons.

24. Membrane-associated periodic skeleton is a signaling platform for RTK transactivation in neurons.

R Zhou , B Han , C Xia , X Zhuang. Science 365, 929-934 (2019)

Highlighted in Nature Reviews Neuroscience 20, 647 (2019): A platform for coordinated signalling

Junction resolving enzymes use multivalency to keep the Holliday junction dynamic.

23. Junction resolving enzymes use multivalency to keep the Holliday junction dynamic.

R Zhou# (co-corresponding author) , O Yang , AC Déclais et al. Nature Chemical Biology 15, 269-275 (2019)

Highlighted in Nature Chemical Biology 15, 209-210 (2019): Freedom of movement

Visualizing and discovering cellular structures with super-resolution microscopy.

22. Visualizing and discovering cellular structures with super-resolution microscopy.

Y Sigal , R Zhou , X Zhuang. Science 361, 880-887 (2018)

Structural organization of the actin-spectrin–based membrane skeleton in dendrites and soma of neurons

21. Structural organization of the actin-spectrin–based membrane skeleton in dendrites and soma of neurons

B Han , R Zhou# (co-first author / co-corresponding author) , C Xia , X Zhuang#. PNAS 114,6678-6685 (2017)

Real-time imaging of translation on single mRNA transcripts in live cells.

20. Real-time imaging of translation on single mRNA transcripts in live cells.

C Wang , B Han , R Zhou , X Zhuang. Cell 165, 990-1001 (2016)

Prevalent presence of periodic actin-spectrin-based membrane skeleton in a broad range of neuronal cell types and animal species.

19. Prevalent presence of periodic actin-spectrin-based membrane skeleton in a broad range of neuronal cell types and animal species.

J He , R Zhou (co-first author) , Z Wu et al. PNAS 113, 6029-6034 (2016)

Asymmetric unwrapping of nucleosomes under tension directed by DNA local flexibility.

18. Asymmetric unwrapping of nucleosomes under tension directed by DNA local flexibility.

TTM Ngo , Q Zhang , R Zhou , JG Yodh , T Ha. Cell 160, 1135-1144 (2015)

Highlighted in Cell 160, 1139-1140 (2015): A Molecular Take on Aesop’s The Oak and the Reeds.

Periodic DNA patrolling underlies diverse functions of Pif1 on R-loops and G-rich DNA.

17. Periodic DNA patrolling underlies diverse functions of Pif1 on R-loops and G-rich DNA.

R Zhou , J Zhang , ML Bochman , VA Zakian , T Ha. eLife 3, e02190 (2014)

Highlighted in eLife 3, e02854 (2014): DNA Helicases: Molecular watchdogs on genome patrol

Detecting intramolecular conformational dynamics of single molecules in short distance range with sub-nanometer sensitivity.

16. Detecting intramolecular conformational dynamics of single molecules in short distance range with sub-nanometer sensitivity.

R Zhou , S Kunzelmann , M Webb , T Ha. Nano Letters 11, 5482-5488 (2011)

SSB functions as a sliding platform that migrates on DNA via reptation.

15. SSB functions as a sliding platform that migrates on DNA via reptation.

R Zhou , AG Kozlov , R Roy , J Zhang , S Korolev , TM Lohman , T Ha. Cell 146, 222-232 (2011)

Selected as: the Cell PaperFlick (video) of the issue (Watch on Youtube)

Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics.

14. Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics.

C Grashoff , BD Hoffman , MD Brenner , R Zhou , M Parsons , MT Yang , M A. McLean , SG Sligar , CS Chen , T Ha , MA Schwartz. Nature 466, 263–266 (2010)

Highlighted in Nature 466, 192–193 (2010): Sensing tension

Fluorescence-force spectroscopy maps two-dimensional reaction landscape of the Holliday junction.

13. Fluorescence-force spectroscopy maps two-dimensional reaction landscape of the Holliday junction.

S Hohng , R Zhou , MK Nahas , J Yu , K Schulten , DMJ Lilley , T Ha. Science 318, 279-283 (2007)

Highlighted in Nature Methods 4, 986-987 (2007): Fluorescence-force spectroscopy: watch your tether

Other publications:

12. βII-spectrin promotes mouse brain connectivity through stabilizing axonal plasma membranes and enabling axonal organelle transport.

DN Lorenzo , A Badea , R Zhou , et al. PNAS 116, 15686-15695 (2019)

11. Spider silk peptide is a compact, linear nanospring ideal for intracellular tension sensing.

MD Brenner , R Zhou , et al. Nano Letters 16, 2096-2102 (2016)

10. Developmental mechanism of the periodic membrane skeleton in axons.

G Zhong , J He , R Zhou , D. Lorenzo , HP Babcock , V Bennett , X Zhuang. eLife 3, e04581 (2015)

9. Structural mechanisms of PriA-mediated DNA replication restart.

B Bhattacharyya , NP George , TM Thurmes , R Zhou , N Jani , SR Wessel , SJ Sandler , T Ha , JL Keck. PNAS 111, 1373-1378 (2014)

8. An improved surface passivation method for single-molecule studies.

B Hua , KY Han , R Zhou , H Kim , X Shi , SC Abeysirigunawardena , A Jain , D. Singh , V. Aggarwal , SA Woodson , T Ha. Nature Methods 11, 1233-1236 (2014)

7. The Telomere Capping Complex CST Has an Unusual Stoichiometry, Makes Multipartite Interaction with G-Tails, and Unfolds Higher-Order G-Tail Structures.

NF Lue , R Zhou , L Chico , N Mao , O Steinberg-Neifach , T Ha. PLoS Genetics 9, e1003145 (2013)

6. The SOSS1 single-stranded DNA binding complex promotes DNA end resection in concert with Exo1.

SH Yang , R Zhou , J Campbell , J Chen , T Ha , TT Paull. EMBO Journal 32, 126-132 (2013)

5. PriC-mediated DNA replication restart requires PriC complex formation with the single-stranded DNA-binding protein.

SR Wessel , AH Marceau , SC Massoni , R Zhou , T Ha , SJ Sandler , JL Keck. Journal of Biological Chemistry 288, 17569-17578 (2013)

4. Single molecule analysis of Thermus thermophilus SSB protein dynamics on single-stranded DNA.

J Zhang , R Zhou , J Inoue , T Mikawa , T Ha. Nucleic Acids Research 42, 3821-3832 (2013)

3. Single molecule analysis of SSB dynamics on DNA.

R Zhou , T Ha. Methods in Molecular Biology 922, 85-100 (2012)

2. Forcing a connection: Impacts of single-molecule force spectroscopy on in vivo tension sensing.

MD Brenner , R Zhou , T Ha. Biopolymers 95, 332-344 (2011)

1. Fluorescence-force spectroscopy at the single molecule level.

R Zhou , M Schlierf , T Ha. Methods in Enzymology 475C, 405-426 (2010)