K. Matsui, J. J. Boniface, P. Steffner, P. A. Reay, and M. M. Davis, «Kinetics of T-cell receptor binding to peptide/I-Ek complexes: correlation of the dissociation rate with T-cell responsiveness, Proceedings of the National Academy of Sciences of the United States of America, vol.91, pp.12862-12866, 1994.

J. J. Lin, S. T. Low-nam, K. N. Alfieri, D. B. Mcaffee, N. C. Fay et al., Groves, «Mapping the stochastic sequence of individual ligand-receptor binding events to cellular activation: T cells act on the rare events, Science signaling, vol.12, p.2019

D. S. Lyons, S. A. Lieberman, J. Hampl, J. J. Boniface, Y. Chien et al.,

. Davis, A TCR binds to antagonist ligands with lower affinities and faster dissociation rates than to agonists.,» Immunity, vol.5, pp.53-61, 1996.

G. J. Kersh, E. N. Kersh, D. H. Fremont, and P. M. Allen, High-and low-potency ligands with similar affinities for the TCR: the importance of kinetics in TCR signaling.,» Immunity, vol.9, pp.817-826, 1998.

M. Aleksic, O. Dushek, H. Zhang, E. Shenderov, J. Chen et al.,

P. A. Coombs and . Merwe, «Dependence of T cell antigen recognition on T cell receptorpeptide MHC confinement time, » Immunity, vol.32, pp.163-174, 2010.

O. Dushek, M. Aleksic, R. J. Wheeler, H. Zhang, S. Cordoba et al.,

V. Chen, T. Cerundolo, D. Dong, P. A. Coombs, and . Merwe, «Antigen potency and maximal efficacy reveal a mechanism of efficient T cell activation, Science signaling, vol.4, p.2011

T. W. Mckeithan, Kinetic proofreading in T-cell receptor signal transduction, Proceedings of the National Academy of Sciences of the United States of America, vol.92, pp.5042-5046, 1995.

G. I. Bell, «Models for the specific adhesion of cells to cells, Science, vol.200, pp.618-627, 1978.

B. T. Marshall, M. Long, J. W. Piper, T. Yago, R. P. Mcever et al., «Direct observation of catch bonds involving cell-adhesion molecules, Nature, vol.423, pp.190-193, 2003.

J. B. Huppa, M. Axmann, M. A. Mörtelmaier, B. F. Lillemeier, E. W. Newell et al., «TCR-peptide-MHC interactions in situ show accelerated kinetics and increased affinity, Nature, vol.463, pp.963-967, 2010.

J. Huang, V. I. Zarnitsyna, B. Liu, L. J. Edwards, N. Jiang et al., «The kinetics of two-dimensional TCR and pMHC interactions determine T-cell responsiveness, Nature, vol.464, pp.932-936, 2010.

S. T. Kim, K. Takeuchi, Z. J. Sun, M. Touma, C. E. Castro et al., «The alphabeta T cell receptor is an anisotropic mechanosensor, The Journal of biological chemistry, vol.284, pp.31028-31037, 2009.

Y. Li, B. Chen, P. Wu, T. Cheng, L. Kao et al.,

R. Roffler and . Cutting, Edge: mechanical forces acting on T cells immobilized via the TCR complex can trigger TCR signaling, Journal of immunology, vol.184, pp.5959-5963, 1950.

K. H. Hu and M. J. Butte, «T cell activation requires force generation, The Journal of cell biology, vol.213, pp.535-542, 2016.

Y. Liu, L. Blanchfield, V. P. Ma, R. Andargachew, K. Galior et al., «DNA-based nanoparticle tension sensors reveal that T-cell receptors transmit defined pN forces to their antigens for enhanced fidelity, Proceedings of the National Academy of Sciences of the United States of America, vol.113, pp.5610-5615, 2016.

Y. Feng, K. N. Brazin, E. Kobayashi, R. J. Mallis, E. L. Reinherz et al., Mechanosensing drives acuity of , javax.xml.bind.JAXBElement@682108ad, T-cell recognition, vol.114, p.2017

B. Liu, W. Chen, B. D. Evavold, and C. Zhu, «Accumulation of dynamic catch bonds between TCR and agonist peptide-MHC triggers T cell signaling, » Cell, vol.157, pp.357-368, 2014.

J. Hong, S. P. Persaud, S. Horvath, P. M. Allen, B. D. Evavold et al., Situ TCR-Peptide-Bound MHC Class II Kinetics Determine Functions of CD4+

T. Cells, Journal of immunology, vol.195, pp.3557-3564, 1950.

E. M. Kolawole, R. Andargachew, B. Liu, J. R. Jacobs, and B. D. Evavold, «2D Kinetic Analysis of TCR and CD8 Coreceptor for LCMV GP33 Epitopes, Frontiers in immunology, vol.9, p.2348, 2018.

L. V. Sibener, R. A. Fernandes, E. M. Kolawole, C. B. Carbone, F. Liu et al., Isolation of a Structural Mechanism for Uncoupling T Cell Receptor Signaling from Peptide-MHC Binding, vol.174, pp.672-687, 2018.

D. K. Das, Y. Feng, R. J. Mallis, X. Li, D. B. Keskin et al.,

G. Wang, E. L. Wagner, M. J. Reinherz, and . Lang, «Force-dependent transition in the T-cell receptor ?-subunit allosterically regulates peptide discrimination and pMHC bond lifetime, Proceedings of the National Academy of Sciences of the United States of America, vol.112, pp.1517-1522, 2015.

B. Liu, W. Chen, K. Natarajan, Z. Li, D. H. Margulies et al., «The cellular environment regulates in situ kinetics of T-cell receptor interaction with peptide major histocompatibility complex, European journal of immunology, vol.45, pp.2099-2110, 2015.

P. Robert, M. Aleksic, O. Dushek, V. Cerundolo, P. Bongrand et al., «Kinetics and mechanics of two-dimensional interactions between T cell receptors and different activating ligands, Biophysical journal, vol.102, p.2012

P. Robert, A. Nicolas, S. Aranda-espinoza, P. Bongrand, and L. Limozin, «Minimal encounter time and separation determine ligand-receptor binding in cell adhesion, Biophysical journal, vol.100, pp.2642-2651, 2011.

L. Limozin, P. Bongrand, and P. Robert, «A Rough Energy Landscape to Describe Surface-Linked Antibody and Antigen Bond Formation, » Scientific reports, vol.6, p.2016

P. Robert, L. Limozin, A. Pierres, and P. Bongrand, «Biomolecule association rates do not provide a complete description of bond formation, Biophysical journal, vol.96, pp.4642-4650, 2009.

N. J. Burroughs, Z. Lazic, and P. A. Merwe, «Ligand detection and discrimination by spatial relocalization: A kinase-phosphatase segregation model of TCR activation, Biophysical journal, vol.91, pp.1619-1629, 2006.

T. A. Springer, «Adhesion receptors of the immune system, Nature, vol.346, pp.425-434, 1990.

H. Yang, X. Gou, Y. Wang, T. M. Fahmy, A. Y. Leung et al., A dynamic model of chemoattractant-induced cell migration, Biophysical journal, vol.108, pp.1645-1651, 2015.

K. Ayu, M. G. Grinfeldt, S. V. Levin, and A. D. Smilgavichus, «Local mechanical oscillations of the cell surface within the range 0.2-30 Hz, European biophysics journal : EBJ, vol.19, pp.93-99, 1990.

Y. Jung, I. Riven, S. W. Feigelson, E. Kartvelishvily, K. Tohya et al., «Three-dimensional localization of T-cell receptors in relation to microvilli using a combination of superresolution microscopies, Proceedings of the National Academy of Sciences of the United States of America, vol.113, p.2016

J. Husson, K. Chemin, A. Bohineust, C. Hivroz, and N. Henry, «Force generation upon T cell receptor engagement, PloS one, vol.6, p.2011

K. T. Bashour, A. Gondarenko, H. Chen, K. Shen, X. Liu et al.,

. Kam, Proceedings of the National Academy of Sciences of the United States of America, vol.111, pp.2241-2246, 2014.

F. Pincet and J. Husson, «The solution to the streptavidin-biotin paradox: the influence of history on the strength of single molecular bonds, Biophysical journal, vol.89, pp.4374-4381, 2005.

V. Lo-schiavo, P. Robert, L. Limozin, and P. Bongrand, «Quantitative modeling assesses the contribution of bond strengthening, rebinding and force sharing to the avidity of biomolecule interactions, PloS one, vol.7, p.44070, 2012.

Y. V. Pereverzev, O. V. Prezhdo, M. Forero, E. V. Sokurenko, and W. E. Thomas, «The two-pathway model for the catch-slip transition in biological adhesion, Biophysical journal, vol.89, pp.1446-1454, 2005.

W. E. Thomas, E. Trintchina, M. Forero, V. Vogel, and E. V. Sokurenko, «Bacterial adhesion to target cells enhanced by shear force, » Cell, vol.109, pp.913-923, 2002.

C. González, P. Chames, B. Kerfelec, D. Baty, P. Robert et al., «Nanobody-CD16 Catch Bond Reveals NK Cell Mechanosensitivity.,» Biophysical journal, vol.116, p.2019

C. D. Paschall, W. H. Guilford, and M. B. Lawrence, «Enhancement of L-selectin, but not P-selectin, bond formation frequency by convective flow, Biophysical journal, vol.94, pp.1034-1045, 2008.

K. C. Johnson and W. E. Thomas, «How Do We Know when Single-Molecule Force Spectroscopy Really Tests Single Bonds?, » Biophysical journal, vol.114, pp.2032-2039, 2018.

O. Dushek and P. A. Merwe, «An induced rebinding model of antigen discrimination, Trends in immunology, vol.35, pp.153-158, 2014.

R. M. Pielak, G. P. O'donoghue, J. J. Lin, K. N. Alfieri, N. C. Fay et al., Groves, «Early T cell receptor signals globally modulate ligand:receptor affinities during antigen discrimination, Proceedings of the National Academy of Sciences of the United States of America, vol.114, p.2017

J. Hong, C. Ge, P. Jothikumar, Z. Yuan, B. Liu et al., Singer et C. Zhu, «A TCR mechanotransduction signaling loop induces negative selection in the thymus, Nature immunology, vol.11, 2018.

K. N. Brazin, R. J. Mallis, D. K. Das, Y. Feng, W. Hwang et al., Structural Features of the ??TCR Mechanotransduction Apparatus That Promote pMHC Discrimination, vol.6, p.441, 2015.

P. Rosa, N. Gopalakrishnan, H. Ibrahim, M. Haug, and Ø. Halaas, The intercell dynamics of T cells and dendritic cells in a lymph node, vol.16, p.2016

A. Brodovitch, L. Limozin, P. Bongrand, and A. Pierres, «Use of TIRF to Monitor T-Lymphocyte Membrane Dynamics with Submicrometer and Subsecond Resolution, Cellular and molecular bioengineering, vol.8, issue.11, pp.178-186, 2015.

M. J. Taylor, K. Husain, Z. J. Gartner, S. Mayor, and R. D. Vale, DNA-Based T Cell Receptor Reveals a Role for Receptor Clustering in Ligand Discrimination, » Cell, vol.169, issue.11, p.2017

A. Pierres, A. M. Benoliel, C. Zhu, and P. Bongrand, «Diffusion of microspheres in shear flow near a wall: use to measure binding rates between attached molecules, Biophys J, vol.81, issue.11, pp.25-42, 2001.