(81) Fabian Höglsperger*, Bart E. Vos*, Arne D. Hofemeier, Maximilian D. Seyfried, Bastian Stövesand, Azadeh Alavizargar, Leon Topp, Andreas Heuer, Timo Betz** & Bart Jan Ravoo**, "Rapid and reversible optical switching of cell membrane area by an amphiphilic azobenzene", Nature Communications, June 23, 2023,
* shared first authorship, ** shared last authorship

(80) Lettermann, Leon, Alejandro Jurado, Timo Betz, Florentin Wörgötter, and Sebastian Herzog. “PhysNODE: Fusion of Data and Expert Knowledge for Modeling Dynamical Systems.” arXiv, May 16, 2023.

(79) Knotz, Gabriel, Till M. Muenker, Timo Betz, and Matthias Krüger. “Entropy Bound for Time Reversal Markers.” arXiv, February 15, 2023.

(78) Di Terlizzi, I., M. Gironella, D. Herráez-Aguilar, T. Betz, F. Monroy, M. Baiesi, and F. Ritort. “Variance Sum Rule for Entropy Production.” arXiv, February 16, 2023.

(77) Volpe, G.,...Muenker, T.M., Vos, B.E., Betz,... et al. "Roadmap for Optical Tweezers", Journal of Physics: Photonics 5, no. 2 (April 2023): 022501.


(76) Hofemeier, A.D., Muenker, T.M., Herkenrath, F., Ristau, M., Brandt, M., Shahriyari, M., Tiburcy, M., Zimmermann, W.H., Lenz, C., Mamchaoui, K., Bigot, A., Gilbert, P.M., Betz, T., Dystrophin is a mechanical tension modulator. bioRxiv 2022,

(75) Russo, A., Schürmann, H., Brandt, M., Scholz, K., Matos, A.L.L., Grill, D., Revenstorff, J., Rembrink, M., von Wulffen, M., Fischer-Riepe, L., Hanley, P.J., Häcker, H., Prünster, M., Sánchez-Madrid, F., Hermann, S., Klotz, L., Gerke, V., Betz, T., Vogl, T., Roth, J., Alarming and Calming: Opposing Roles of S100A8/S100A9 Dimers and Tetramers on Monocytes. 2022 Advanced Science, 2201505.

(74) Muenker, T. M.; Knotz, G.; Krüger, M.; Betz, T. Onsager Regression Characterizes Living Systems in Passive Measurements. bioRxiv 2022.

(73) Schürmann, H.; Abbasi, F.; Russo, A.; Hofemeier, A. D.; Brandt, M.; Roth, J.; Vogl, T.; Betz, T. Analysis of Monocyte Cell Tractions in 2.5D Reveals Mesoscale Mechanics of Podosomes during Substrate-Indenting Cell Protrusion. Journal of Cell Science 2022, 135 (10), jcs259042.

(72) Höglsperger, F.; Betz, T.; Ravoo, B. J. Reversible Photoresponsive Modulation of Osmotic Pressure via Macromolecular Host–Guest Interaction. ACS Macro Lett. 2022, 537–542.

(71) Kummer, D.; Steinbacher, T.; Thölmann, S.; Schwietzer, M. F.; Hartmann, C.; Horenkamp, S.; Demuth, S.; Peddibhotla, S. S. D.; Brinkmann, F.; Kemper, B.; Schnekenburger, J.; Brandt, M.; Betz, T.; Liashkovich, I.; Kouzel, I. U.; Shahin, V.; Corvaia, N.; Rottner, K.; Tarbashevich, K.; Raz, E.; Greune, L.; Schmidt, M. A.; Gerke, V.; Ebnet, K. A JAM-A–Tetraspanin–Αvβ5 Integrin Complex Regulates Contact Inhibition of Locomotion. Journal of Cell Biology 2022, 221 (4), e202105147.

(70) Long, H., Vos, B.E., Betz, T., Baker, B.M., Trappmann, B., Nonswelling and Hydrolytically Stable Hydrogels Uncover Cellular Mechanosensing in 3D. Advanced Science 2022, 9(12), 2105325.

(69) Brandt, M.; Gerke, V.; Betz, T. Human Endothelial Cells Display a Rapid Tensional Stress Increase in Response to Tumor Necrosis Factor-α. PLOS ONE 2022, 17 (6), e0270197.

(68) Raghuraman, S., Schubert, A.-S., Bröker, S., Jurado, A., Müller, A., Brandt, M., Vos, B.E., Hofemeier, A.D., Abbasi, F., Stehling, M., Wittkowski, R., Ivaska, J., Betz, T., 2022. Pressure Drives Rapid Burst-Like Coordinated Cellular Motion from 3D Cancer Aggregates (Adv. Sci. 6/2022). Advanced Science 9, 2270037.


(67) Hurst S., Vos B., Brandt M., Betz T., 2021,
Intracellular softening and increased viscoelastic fluidity during division,
Nature Physics, 17, 1270-1276,

(66) BE Vos, T Betz, 2021, Designing a LEGO-based microscope for an educational setting, Proceedings OSA- ETOP, Th4B. 1,

(65) Andreu, I., Falcones, B., Hurst, S., Chahare, N., Quiroga, X., Roux, A.-L.L., Kechagia, Z., Beedle, A.E.M., Elósegui-Artola, A., Trepat, X., Farré, R., Betz, T., Almendros, I., Roca-Cusachs, P., 2021, The force loading rate drives cell mechanosensing through both reinforcement and fluidization.
Nature Communications, 12, 4229,

(64) V Zieger, T Betz, BE Vos, 2021,Combining light sheet microscopy and optical tweezers for simultaneous micromanipulation and 3D visualization
OSA- Optical Manipulation and Its Applications, AM2D. 3,

(63) Vos BE, Betz Blesa E, Betz T. 2021, Designing a high-resolution, LEGO-based microscope for an educational setting. The Biophysicist, 2021,

(62) Betz, T., (2021), Viscoelastic properties driving collective migration in zebrafish development,
in "Viscoelasticity and Collective Cell Migration". Academic Press, pp. 135–155.

(61) Olguin-Olguin, A.*, Aalto, A.*, Maugis, B., Boquet-Pujadas, A., Hofmann, D., Ermlich, L., Betz, T., Gov, N.S., Reichman-Fried, M., Raz, E. (2021)
Chemokine-biased robust self-organizing polarization of migrating cells in vivo.
Proceedings of the National Academy of Science USA, 118 (7) e2018480118,

(60) Hofemeier, A.D., Limon, T., Muenker, T.M., Wallmeyer, B., Jurado, A., Afshar, M.E., Ebrahimi, M., Tsukanov, R., Oleksiievets, N., Enderlein, J., Gilbert, P.M., Betz, T., 2021.
Global and local tension measurements in biomimetic skeletal muscle tissues reveals early mechanical homeostasis.
eLife 10, e60145.

(59) Alfred Nguyen, Matthias Brandt, Timo Betz,
Microchip based microrheology via Acoustic Force Spectroscopy shows that endothelial cell mechanics follows a fractional viscoelastic model
Lab on a Chip, 2021,


(58) Gross-Thebing, S*., Truszkowski, L*., Tenbrinck, D*+., Sánchez-Iranzo, H., Camelo, C., Westerich, K.J., Singh, A., Paul Maier, P., Prengel, J., Lange, P., Hüwel, J., Gaede, F., Sasse, R., Vos, E.B., Betz, T., Matis, M., Prevedel, R., Luschnig, S., Diz-Muñoz, A., Burger, M., Raz, E+. (2020)
Using migrating cells as probes to illuminate features in live embryonic tissues
Science Advances (6) 49, eabc5546.

(57) Allard, A.; Valentino, F.; Sykes, C.; Betz, T.; Campillo, C.
Fluctuations of a Membrane Nanotube Covered with an Actin Sleeve. Phys. Rev. E 2020, 102 (5), 052402.

(56) Grimaldi, C.; Schumacher, I.; Boquet-Pujadas, A.; Tarbashevich, K.; Vos, B. E.; Bandemer, J.; Schick, J.; Aalto, A.; Olivo-Marin, J.-C.; Betz, T.; Raz, E.
E-Cadherin Focuses Protrusion Formation at the Front of Migrating Cells by Impeding Actin Flow. Nature Communications 2020, 11 (1), 5397.

(55) Rheinlaender, J.; Dimitracopoulos, A.; Wallmeyer, B.; Kronenberg, N. M.; Chalut, K. J.; Gather, M. C.; Betz, T.; Charras, G.; Franze, K.
Cortical Cell Stiffness Is Independent of Substrate Mechanics. Nat Mater 2020, 19 (9), 1019–1025.

(54) Davidson, P. M.; Battistella, A.; Déjardin, T.; Betz, T.; Plastino, J.; Borghi, N.; Cadot, B.; Sykes, C.
Nesprin-2 Accumulates at the Front of the Nucleus during Confined Cell Migration. EMBO Rep. 2020, 21 (7), e49910.

(53) Colin, A.; Letort, G.; Razin, N.; Almonacid, M.; Ahmed, W.; Betz, T.; Terret, M.-E.; Gov, N. S.; Voituriez, R.; Gueroui, Z.; Verlhac, M.-H.
Active Diffusion in Oocytes Nonspecifically Centers Large Objects during Prophase I and Meiosis I. J. Cell Biol. 2020, 219 (3).

(52) Allard, A.; Bouzid, M.; Betz, T.; Simon, C.; Abou-Ghali, M.; Lemière, J.; Valentino, F.; Manzi, J.; Brochard-Wyart, F.; Guevorkian, K.; Plastino, J.; Lenz, M.; Campillo, C.; Sykes, C.
Actin Modulates Shape and Mechanics of Tubular Membranes.
Science Advances 2020, 6 (17), eaaz3050.


(51) Turlier, H.; Betz, T.
Unveiling the Active Nature of Living-Membrane Fluctuations and Mechanics.
Annual Review of Condensed Matter Physics 2019, 10 (1), 213–232.


(50) Wedlich-Söldner, R.; Betz, T.
Self-Organization: The Fundament of Cell Biology.
Phil. Trans. R. Soc. B 2018, 373 (1747), 20170103.

(49) Wallmeyer, B.; Trinschek, S.; Yigit, S.; Thiele, U.; Betz, T.
Collective Cell Migration in Embryogenesis Follows the Laws of Wetting.
Biophysical Journal 2018, 114 (1), 213–222.

(48) Betz, T. Physik Journal 
Gut Geschüttelt, Nicht Gerührt 
Pro-Physik.De. Physik Journal 2018, 17 (2), 29–34.

(47) Physics of Biological Membranes; Bassereau, P., Sens, P., Eds.; Springer International Publishing, 2018.

(46) Baschieri, F.; Dayot, S.; Elkhatib, N.; Ly, N.; Capmany, A.; Schauer, K.; Betz, T.; Vignjevic, D. M.; Poincloux, R.; Montagnac, G.
Frustrated Endocytosis Controls Contractility-Independent Mechanotransduction at Clathrin-Coated Structures.
Nature Communications 2018, 9 (1), 3825.

(45) Ahmed, W. W.; Fodor, É.; Almonacid, M.; Bussonnier, M.; Verlhac, M.-H.; Gov, N.; Visco, P.; van Wijland, F.; Betz, T.
Active Mechanics Reveal Molecular-Scale Force Kinetics in Living Oocytes.
Biophysical Journal 2018, 114 (7), 1667–1679.


(44) Rückerl, F.; Lenz, M.; Betz, T.; Manzi, J.; Martiel, J.-L.; Safouane, M.; Paterski-Boujemaa, R.; Blanchoin, L.; Sykes, C.
Adaptive Response of Actin Bundles under Mechanical Stress.
Biophys. J. 2017, 113 (5), 1072–1079.

(43) Peuhu, E.; Kaukonen, R.; Lerche, M.; Saari, M.; Guzmán, C.; Rantakari, P.; De Franceschi, N.; Wärri, A.; Georgiadou, M.; Jacquemet, G.; Mattila, E.; Virtakoivu, R.; Liu, Y.; Attieh, Y.; Silva, K. A.; Betz, T.; Sundberg, J. P.; Salmi, M.; Deugnier, M.-A.; Eliceiri, K. W.; Ivaska, J.
SHARPIN Regulates Collagen Architecture and Ductal Outgrowth in the Developing Mouse Mammary Gland.
EMBO J. 2017, 36 (2), 165–182.

(42) Hörner, F.; Meissner, R.; Polali, S.; Pfeiffer, J.; Betz, T.; Denz, C.; Raz, E.
Holographic Optical Tweezers-Based in Vivo Manipulations in Zebrafish Embryos.
J Biophotonics 2017, 10 (11), 1492–1501.

(41) Goudarzi, M.; Tarbashevich, K.; Mildner, K.; Begemann, I.; Garcia, J.; Paksa, A.; Reichman-Fried, M.; Mahabaleshwar, H.; Blaser, H.; Hartwig, J.; Zeuschner, D.; Galic, M.; Bagnat, M.; Betz, T.; Raz, E.
Bleb Expansion in Migrating Cells Depends on Supply of Membrane from Cell Surface Invaginations.
Dev. Cell 2017, 43 (5), 577-587.e5.

(40) Attieh, Y.; Clark, A. G.; Grass, C.; Richon, S.; Pocard, M.; Mariani, P.; Elkhatib, N.; Betz, T.; Gurchenkov, B.; Vignjevic, D. M.
Cancer-Associated Fibroblasts Lead Tumor Invasion through Integrin-Β3-Dependent Fibronectin Assembly.
J. Cell Biol. 2017, 216 (11), 3509–3520.


(39) Valentino, F.; Sens, P.; Lemière, J.; Allard, A.; Betz, T.; Campillo, C.; Sykes, C.
Fluctuations of a Membrane Nanotube Revealed by High-Resolution Force Measurements.
Soft Matter 2016, 12 (47), 9429–9435.

(38) Turlier, H.; Fedosov, D. A.; Audoly, B.; Auth, T.; Gov, N. S.; Sykes, C.; Joanny, J.-F.; Gompper, G.; Betz, T.
Equilibrium Physics Breakdown Reveals the Active Nature of Red Blood Cell Flickering.
Nat Phys 2016, 12 (5), 513–519.

(37) Kopanska, K. S.; Alcheikh, Y.; Staneva, R.; Vignjevic, D.; Betz, T.
Tensile Forces Originating from Cancer Spheroids Facilitate Tumor Invasion.
PLoS ONE 2016, 11 (6), e0156442.

(36) Kaukonen, R.; Mai, A.; Georgiadou, M.; Saari, M.; De Franceschi, N.; Betz, T.; Sihto, H.; Ventelä, S.; Elo, L.; Jokitalo, E.; Westermarck, J.; Kellokumpu-Lehtinen, P.-L.; Joensuu, H.; Grenman, R.; Ivaska, J.
Normal Stroma Suppresses Cancer Cell Proliferation via Mechanosensitive Regulation of JMJD1a-Mediated Transcription.
Nat Commun 2016, 7, 12237.

(35) Fodor, É.; Ahmed, W. W.; Almonacid, M.; Bussonnier, M.; Gov, N. S.; Verlhac, M.-H.; Betz, T.; Visco, P.; Wijland, F. van.
Nonequilibrium Dissipation in Living Oocytes.
EPL 2016, 116 (3), 30008.

(34) Caorsi, V.; Lemière, J.; Campillo, C.; Bussonnier, M.; Manzi, J.; Betz, T.; Plastino, J.; Carvalho, K.; Sykes, C.
Cell-Sized Liposome Doublets Reveal Active Tension Build-up Driven by Acto-Myosin Dynamics. Soft Matter 2016, 12 (29), 6223–6231.


(33) Kopanska, K. S.; Bussonnier, M.; Geraldo, S.; Simon, A.; Vignjevic, D.; Betz, T.
Quantification of Collagen Contraction in Three-Dimensional Cell Culture.
Methods Cell Biol. 2015, 125, 353–372.

(32) Gárate, F.; Betz, T.; Pertusa, M.; Bernal, R.
Time-Resolved Neurite Mechanics by Thermal Fluctuation Assessments.
Phys. Biol. 2015, 12 (6), 066020.

(31) Davis, J. R.; Luchici, A.; Mosis, F.; Thackery, J.; Salazar, J. A.; Mao, Y.; Dunn, G. A.; Betz, T.; Miodownik, M.; Stramer, B. M.
Inter-Cellular Forces Orchestrate Contact Inhibition of Locomotion.
Cell 2015, 161 (2), 361–373.

(30) Almonacid, M.; Ahmed, W. W.; Bussonnier, M.; Mailly, P.; Betz, T.; Voituriez, R.; Gov, N. S.; Verlhac, M.-H.
Active Diffusion Positions the Nucleus in Mouse Oocytes.
Nat. Cell Biol. 2015, 17 (4), 470–479.

(29) Ahmed, W. W.; Fodor, É.; Betz, T.
Active Cell Mechanics: Measurement and Theory.
Biochim. Biophys. Acta 2015, 1853 (11 Pt B), 3083–3094.

(28) Ahmed, W. W.; Betz, T.
Dynamic Cross-Links Tune the Solid–Fluid Behavior of Living Cells.
PNAS 2015, 112 (21), 6527–6528.


(27) Silva, M. S. e; Stuhrmann, B.; Betz, T.; Koenderink, G. H.
Time-Resolved Microrheology of Actively Remodeling Actomyosin Networks.
New J. Phys. 2014, 16 (7), 075010. https://doi.org/10.1088/1367-2630/16/7/075010.

(26) Peukes, J.; Betz, T.
Direct Measurement of the Cortical Tension during the Growth of Membrane Blebs.
Biophys J 2014, 107 (8), 1810–1820.

(25) Elkhatib, N.; Neu, M. B.; Zensen, C.; Schmoller, K. M.; Louvard, D.; Bausch, A. R.; Betz, T.; Vignjevic, D. M.
Fascin Plays a Role in Stress Fiber Organization and Focal Adhesion Disassembly.
Current Biology 2014, 24 (13), 1492–1499.

(24) Dasgupta, S.; Auth, T.; Gov, N. S.; Satchwell, T. J.; Hanssen, E.; Zuccala, E. S.; Riglar, D. T.; Toye, A. M.; Betz, T.; Baum, J.; Gompper, G.
Membrane-Wrapping Contributions to Malaria Parasite Invasion of the Human Erythrocyte.
Biophys. J. 2014, 107 (1), 43–54.

(23) Bussonnier, M.; Carvalho, K.; Lemière, J.; Joanny, J.-F.; Sykes, C.; Betz, T.
Mechanical Detection of a Long-Range Actin Network Emanating from a Biomimetic Cortex.
Biophys J 2014, 107 (4), 854–862.


(22) Lemière, J.; Guevorkian, K.; Campillo, C.; Sykes, C.; Betz, T.
α-Hemolysin Membrane Pore Density Measured on Liposomes.
Soft Matter 2013, 9 (11), 3181–3187.

(21) Lafaurie-Janvore, J.; Maiuri, P.; Wang, I.; Pinot, M.; Manneville, J.-B.; Betz, T.; Balland, M.; Piel, M.
ESCRT-III Assembly and Cytokinetic Abscission Are Induced by Tension Release in the Intercellular Bridge.
Science 2013, 339 (6127), 1625–1629.

(20) Carvalho, K.; Lemière, J.; Faqir, F.; Manzi, J.; Blanchoin, L.; Plastino, J.; Betz, T.; Sykes, C.
Actin Polymerization or Myosin Contraction: Two Ways to Build up Cortical Tension for Symmetry Breaking.
Phil. Trans. R. Soc. B 2013, 368 (1629), 20130005.


(19) Betz, T.; Sykes, C.
Time Resolved Membrane Fluctuation Spectroscopy.
Soft Matter 2012, 8(19), 5317–5326.


(18) Fink, J.; Carpi, N.; Betz, T.; Betard, A.; Chebah, M.; Azioune, A.; Bornens, M.; Sykes, C.; Fetler, L.; Cuvelier, D.; Piel, M.
External Forces Control Mitotic Spindle Positioning.
Nat Cell Biol 2011, 13 (7), 771–778.

(17) Betz, T.; Koch, D.; Lu, Y.-B.; Franze, K.; Käs, J. A.
Growth Cones as Soft and Weak Force Generators.
PNAS 2011, 108 (33), 13420–13425.


(16) Smith, D.; Gentry, B.; Stuhrmann, B.; Huber, F.; Strehle, D.; Brunner, C.; Koch, D.; Steinbeck, M.; Betz, T.; KäS, J. A.
The Cytoskeleton: An Active Polymer-Based Scaffold.
Biophys. Rev. Lett. 2009, 04 (01 & 02), 179.

(15) Franze, K.; Gerdelmann, J.; Weick, M.; Betz, T.; Pawlizak, S.; Lakadamyali, M.; Bayer, J.; Rillich, K.; Gögler, M.; Lu, Y.-B.
Neurite Branch Retraction Is Caused by a Threshold-Dependent Mechanical Impact.
Biophysical Journal 2009, 97 (7), 1883–1890.

(14) Betz, T.; Lenz, M.; Joanny, J.-F.; Sykes, C.
ATP-Dependent Mechanics of Red Blood Cells.
Proceedings of the National Academy of Sciences 2009, 106 (36), 15320–15325.

(13) Betz, T.; Koch, D.; Lim, D.; Käs, J. A.
Stochastic Actin Polymerization and Steady Retrograde Flow Determine Growth Cone Advancement.
Biophysical Journal 2009, 96 (12), 5130–5138.


(12) Gögler, M.; Betz, T.; Käs, J. A.
Simultaneous Manipulation and Detection of Living Cell Membrane Dynamics.
Optics Letters 2007, 32 (13), 1893.

(11) Ehrlicher, A.; Betz, T.; Stuhrmann, B.; Gögler, M.; Koch, D.; Franze, K.; Lu, Y.; Käs, J. Optical Neuronal Guidance. Methods Cell Biol 2007, 83, 495–520.

(10) Betz, T.; Koch, D.; Stuhrmann, B.; Ehrlicher, A.; Käs, J.
Statistical Analysis of Neuronal Growth: Edge Dynamics and the Effect of a Focused Laser on Growth Cone Motility.
New J. Phys. 2007, 9 (11), 426–426.


(9) Stuhrmann, B.; Jahnke, H.-G.; Schmidt, M.; Jähn, K.; Betz, T.; Müller, K.; Rothermel, A.; Käs, J.; Robitzki, A. A.
Versatile Optical Manipulation System for Inspection, Laser Processing, and Isolation of Individual Living Cells.
Review of Scientific Instruments 2006, 77 (6), 063116.

(8) Rudolph, T.; Zimmer, K.; Betz, T.
Microstructuring of UV-Transparent Functionalised Films on Glass by Excimer Laser Irradiation. Materials Science and Engineering: C 2006, 26 (5), 1131–1135.

(7) Kamanyi, A.; Ngwa, W.; Betz, T.; Wannemacher, R.; Grill, W.
Combined Phase-Sensitive Acoustic Microscopy and Confocal Laser Scanning Microscopy. Ultrasonics 2006, 44 (Supplement 1), e1295–e1300.

(6) Betz, T.; Lim, D.; Käs, J. A.
Neuronal Growth: A Bistable Stochastic Process.
Phys. Rev. Lett. 2006, 96 (9), 098103.

2005 and before

(5) Stuhrmann, B.; Gögler, M.; Betz, T.; Ehrlicher, A.; Koch, D.; Käs, J.
Automated Tracking and Laser Micromanipulation of Motile Cells.
Review of Scientific Instruments 2005, 76 (3), 035105.

(4) Betz, T.; Teipel, J.; Koch, D.; Härtig, W.; Guck, J.; Käs, J.; Giessen, H.
Excitation beyond the Monochromatic Laser Limit: Simultaneous 3-D Confocal and Multiphoton Microscopy with a Tapered Fiber as White-Light Laser Source.
J Biomed Opt 2005, 10 (5), 054009.

(3) Koch, D.; Betz, T.; Ehrlicher, A.; Gogler, M.; Stuhrmann, B.; Kas, J.
Optical Control of Neuronal Growth. In Optical Trapping and Optical Micromanipulation International Society for Optics and Photonics, 2004; Vol. 5514, pp 428–436.

(2) Betz, T.; Kas, J.; Ehrlicher, A.; Raizen, M.; Milner, V.
Optical Cell Guidance Method and Apparatus.
Patent No: US20010333283P 20011114; WO2002US36456 20021113 2003.

(1) Ehrlicher, A.; Betz, T.; Stuhrmann, B.; Koch, D.; Milner, V.; Raizen, M. G.; Käs, J.
Guiding Neuronal Growth with Light.
Proceedings of the National Academy of Sciences of the United States of America 2002, 99 (25), 16024–16028.

Special Thanks to all the Funding!