#：corresponding author.

Undergraduate students (hightlighted in purple)

Independent Career:

2025
Coordination of cell organelles to promote metabolon formation.

34. Coordination of cell organelles to promote metabolon formation.

Z Sha , AM Pedley , TD Iles , S Zhang , JR Staub , R Zhou , SJ Benkovic# Under review at PNAS (2025)

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

33. “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# under review at Nature Communications. (2025)

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

32. 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.

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

Y Tao , R Zhou#. Submitted. (2025)

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

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

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

Actin Depolymerization Promotes Axonal Mitochondria Transport for Axon Regeneration and Neuroprotection.

29. Actin Depolymerization Promotes Axonal Mitochondria Transport for Axon Regeneration and Neuroprotection.

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#. Under Revision at Science Translational Medicine. (2025)

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

28. 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 (2025)

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

27. 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 (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:

2019
Membrane-associated periodic skeleton is a signaling platform for RTK transactivation in neurons.

34. 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.

33. 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

2018
Visualizing and discovering cellular structures with super-resolution microscopy.

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

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

2017
Structural organization of the actin-spectrin–based membrane skeleton in dendrites and soma of neurons

31. 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)

2016
Real-time imaging of translation on single mRNA transcripts in live cells.

30. 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.

29. 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)

2015
Asymmetric unwrapping of nucleosomes under tension directed by DNA local flexibility.

28. 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.

2014
Periodic DNA patrolling underlies diverse functions of Pif1 on R-loops and G-rich DNA.

27. 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

2011
SSB functions as a sliding platform that migrates on DNA via reptation.

26. 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)

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

25. 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)

2010
Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics.

24. 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

2007
Fluorescence-force spectroscopy maps two-dimensional reaction landscape of the Holliday junction.

23. 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. publisher 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)