E. Arana, A. Vehlow, N. E. Harwood, E. Vigorito, R. Henderson et al., Activation of the Small GTPase Rac2 via the B Cell Receptor Regulates B Cell Adhesion and Immunological-Synapse Formation, Immunity, vol.28, issue.1, pp.88-99, 2008.

O. Bakke and B. Dobberstein, MHC class II-associated invariant chain contains a sorting signal for endosomal compartments, Cell, vol.63, issue.4, pp.707-716, 1990.

K. T. Bashour, A. Gondarenko, H. Chen, K. Shen, X. Liu et al., CD28 and CD3 have complementary roles in T-cell traction forces, Proceedings of the National Academy of Sciences of the United States of America, vol.111, pp.2241-2247, 2014.

R. Basu and M. Huse, Mechanical Communication at the Immunological Synapse, Trends in Cell Biology, xx, pp.1-14, 2016.

R. Basu, B. M. Whitlock, J. Husson, A. Le-floc'h, W. Jin et al., Cytotoxic T Cells Use Mechanical Force to Potentiate, Target Cell Killing. Cell, pp.1-11, 2016.
URL : https://hal.archives-ouvertes.fr/hal-01317871

F. D. Batista and N. E. Harwood, The who, how and where of antigen presentation to B cells, Nature Reviews. Immunology, vol.9, issue.1, pp.15-27, 2009.

N. Bertaux and D. Marguet, Dynamic multiple-target tracing to probe spatiotemporal cartography of cell membranes, vol.5, pp.687-694, 2008.
URL : https://hal.archives-ouvertes.fr/hal-00295014

N. Bufi, M. Saitakis, S. Dogniaux, O. Buschinger, A. Bohineust et al., Human Primary Immune Cells Exhibit Distinct Mechanical Properties that Are Modified by Inflammation, Biophysical Journal, vol.108, issue.9, pp.2181-2190, 2015.

J. P. Butler, I. M. Toli?-nørrelykke, B. Fabry, and J. J. Fredberg, Traction fields, moments, and strain energy that cells exert on their surroundings, American Journal of Physiology. Cell Physiology, vol.282, issue.3, pp.595-605, 2002.

J. C. Cambier, C. M. Pleiman, and M. R. Clark, Signal transduction by the B cell antigen receptor and its coreceptors, Annu Rev Immunol, vol.12, pp.457-486, 1994.

Y. R. Carrasco and F. D. Batista, B cell recognition of membrane-bound antigen: an exquisite way of sensing ligands, Current Opinion in Immunology, vol.18, issue.3, pp.286-91, 2006.

Y. R. Carrasco, S. J. Fleire, T. Cameron, M. L. Dustin, and F. D. Batista, LFA-1/ICAM-1 interaction lowers the threshold of B cell activation by facilitating B cell adhesion and synapse formation, Immunity, vol.20, issue.5, pp.589-99, 2004.

J. Champion, A. Walker, and S. Mitragotri, Role of particle size in phagocytosis of polymeric microspheres, Pharmaceutical Research, vol.25, issue.8, pp.1815-1821, 2008.

J. Cheng, A. Grassart, and D. G. Drubin, Myosin 1E coordinates actin assembly and cargo trafficking during clathrin-mediated endocytosis, Molecular Biology of the Cell, vol.23, issue.15, pp.2891-904, 2012.

K. Clark, M. Langeslag, C. G. Figdor, and F. N. Van-leeuwen, Myosin II and mechanotransduction: a balancing act, Trends in Cell Biology, vol.17, issue.4, pp.178-186, 2007.

M. A. Conti and R. S. Adelstein, Nonmuscle myosin II moves in new directions, Journal of Cell Science, vol.121, pp.11-18, 2008.

A. F. Cunningham, F. Gaspal, K. Serre, E. Mohr, I. R. Henderson et al., Salmonella induces a switched antibody response without germinal centers that impedes the extracellular spread of infection, Journal of Immunology, vol.178, issue.10, pp.6200-6207, 1950.

D. Silva, N. S. Klein, and U. , Dynamics of B cells in germinal centres, Nature Reviews Immunology, vol.15, issue.3, pp.137-148, 2015.

L. Delamarre, M. Pack, H. Chang, I. Mellman, and E. S. Trombetta, Differential Lysosomal Proteolysis in Antigen-Presenting Cells Determines Antigen Fate, Science, vol.307, issue.5715, pp.1630-1634, 2005.

M. Dembo and Y. L. Wang, Stresses at the cell-to-substrate interface during locomotion of fibroblasts, Biophysical Journal, vol.76, issue.4, pp.77386-77394, 1999.

L. K. Denzin and P. Cresswell, HLA-DM induces clip dissociation from MHC class II ?? dimers and facilitates peptide loading, Cell, vol.82, issue.1, pp.90061-90067, 1995.

L. K. Denzin, D. B. Sant'angelo, C. Hammond, M. J. Surman, and P. Cresswell, Negative regulation by HLA-DO of MHC class II-restricted antigen processing, Science, vol.5335, pp.106-115, 1997.

D. Depoil, M. Weber, B. Treanor, S. J. Fleire, Y. R. Carrasco et al., Early events of B cell activation by antigen, Science Signaling, vol.2, issue.63, 2009.

C. Driessen, R. A. Bryant, A. M. Lennon-duménil, J. A. Villadangos, P. W. Bryant et al., Cathepsin S controls the trafficking and maturation of MHC class II molecules in dendritic cells, Journal of Cell Biology, vol.147, issue.4, pp.775-790, 1999.

S. J. Fleire, J. P. Goldman, Y. R. Carrasco, M. Weber, D. Bray et al., B cell ligand discrimination through a spreading and contraction response, Science, issue.5774, pp.738-779, 2006.

D. Georgess, I. Machuca-gayet, A. Blangy, and P. Jurdic, Podosome organization drives osteoclast-mediated bone resorption, vol.8, pp.192-204, 2014.

G. Giannone, B. Dubin-thaler, O. Rossier, Y. Cai, G. Jiang et al., , vol.128, pp.561-575, 2017.

S. F. Gonzalez, V. Lukacs-kornek, M. P. Kuligowski, L. Pitcher, S. E. Degn et al., Capture of influenza by medullary dendritic cells via SIGN-R1 is essential for humoral immunity in draining lymph nodes, Nature Immunology, vol.11, issue.5, pp.427-434, 2010.

C. C. Goodnow, J. Crosbie, S. Adelstein, T. B. Lavoie, S. J. Smith-gill et al., Altered immunoglobulin expression and functional silencing of self-reactive B lymphocytes in transgenic mice, Nature, vol.334, pp.676-682, 1988.

. Grakoui, S. K. Bromley, C. Sumen, M. M. Davis, S. Shaw et al., The immunological synapse: a molecular machine controlling T cell activation, Science, issue.5425, pp.221-227, 1999.

M. Gupta, B. R. Sarangi, J. Deschamps, Y. Nematbakhsh, A. Callan-jones et al., Adaptive rheology and ordering of cell cytoskeleton govern matrix rigidity sensing, Nature Communications, vol.6, p.7525, 2015.
URL : https://hal.archives-ouvertes.fr/hal-01226836

P. Gupta, R. Martin, H. Knö-lker, D. Nihalani, and D. K. Sinha, Myosin-1 inhibition by PClP affects membrane shape, cortical actin distribution and lipid droplet dynamics in early Zebrafish embryos, PLoS ONE, pp.1-21, 2017.

A. K. Harris, P. Wild, and D. Stopak, Silicone rubber substrata: a new wrinkle in the study of cell locomotion, Science, issue.4440, pp.177-186, 1980.

K. L. Hui, L. Balagopalan, L. E. Samelson, and A. Upadhyaya, Cytoskeletal forces during signaling activation in Jurkat T cells, Molecular Biology of the Cell, vol.26, pp.685-695, 2014.

J. Jacobelli, R. S. Friedman, M. A. Conti, A. Lennon-dumenil, M. Piel et al., Confinement-optimized three-dimensional T cell amoeboid motility is modulated via myosin IIA-regulated adhesions, Nature Immunology, vol.11, issue.10, pp.953-61, 2010.

T. Junt, E. A. Moseman, M. Iannacone, S. Massberg, P. Lang et al., Subcapsular sinus macrophages in lymph nodes clear lymphborne viruses and present them to antiviral B cells, Nature, vol.450, issue.7166, pp.110-114, 2007.

M. Kaksonen, C. P. Toret, and D. G. Drubin, Harnessing actin dynamics for clathrinmediated endocytosis, Nature Reviews. Molecular Cell Biology, vol.7, issue.6, pp.404-414, 2006.

R. Kasturi, C. Vasulka, and J. D. Johnson, Ca2+, caldesmon, and myosin light chain kinase exchange with calmodulin, Journal of Biological Chemistry, vol.268, issue.11, pp.7958-7964, 1993.

S. H. Kaufmann, Immunology's foundation: the 100-year anniversary of the Nobel Prize to Paul Ehrlich and Elie Metchnikoff, Nature Immunology, vol.9, issue.7, pp.705-712, 2008.

J. Larsson and S. Karlsson, The role of Smad signaling in hematopoiesis, Oncogene, vol.24, issue.9, pp.5676-5692, 2005.

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, issue.2, pp.357-368, 2014.

C. Liu, H. Miller, G. Orlowski, H. Hang, A. Upadhyaya et al., Actin reorganization is required for the formation of polarized B cell receptor signalosomes in response to both soluble and membrane-associated antigens, Journal of Immunology, vol.188, issue.7, pp.3237-3283, 1950.

C. Liu, H. Miller, S. Sharma, A. Beaven, A. Upadhyaya et al., Analyzing actin dynamics during the activation of the B cell receptor in live B cells, Biochemical and Biophysical Research Communications, vol.427, issue.1, pp.202-208, 2012.

V. Lotteau, L. Teyton, A. Peleraux, T. Nilsson, L. Karlsson et al., Lotteau AntiCLIP invariant chain.pdf, Nature, 1990.

I. C. Maclennan, From the dark zone to the light, Curr Biol, vol.4, issue.1, pp.17-25, 1994.

K. Mandal, I. Wang, E. Vitiello, L. A. Orellana, and M. Balland, Cell dipole behaviour revealed by ECM sub-cellular geometry, Nature Communications, vol.5, p.5749, 2014.
URL : https://hal.archives-ouvertes.fr/hal-02063891

M. Marsh and H. T. Mcmahon, The structural era of endocytosis, Science, vol.285, issue.5425, pp.215-220, 1999.

L. Martinez-pomares and S. Gordon, Antigen Presentation the Macrophage Way, Cell, vol.131, issue.4, pp.641-643, 2007.

D. W. Metzger, C. A. Metzger, L. A. Ling, J. S. Hurst, and V. H. Van-cleave, Preparative Isolation of Murine CD5 B Cells by Panning and Magnetic Beads, Annals of the New York Academy of Sciences, vol.651, issue.1, pp.75-77, 1992.

N. A. Mitchison, T-cell-B-cell cooperation, Nature Reviews Immunology, vol.4, issue.4, pp.308-312, 2004.

S. Mostowy and P. Cossart, Septins: the fourth component of the cytoskeleton, Nature Reviews. Molecular Cell Biology, vol.13, issue.3, pp.183-94, 2012.

T. Nagasawa, Microenvironmental niches in the bone marrow required for B-cell development, Nature Reviews. Immunology, vol.6, issue.2, pp.107-116, 2006.

E. Natkanski, W. Lee, B. Mistry, A. Casal, J. E. Molloy et al., B cells use mechanical energy to discriminate antigen affinities, Science, issue.6140, pp.1587-90, 2013.

D. Needham, The adenosinetriphosphatase activity of myosin preparations, Biochem J, vol.36, issue.1-2, pp.113-120, 1942.

G. J. Nossal, A. Abbot, J. Mitchell, and Z. Lummus, Antigens in immunity. XV. Ultrastructural features of antigen capture in primary and secondary lymphoid follicles, The Journal of Experimental Medicine, vol.127, issue.2, pp.277-90, 1968.

S. L. Nutt, S. Vambrie, P. Steinlein, Z. Kozmik, . Rolink et al., Independent regulation of the two Pax5 alleles during B-cell development, Nature Genetics, vol.21, issue.4, pp.390-395, 1999.

D. Obino and A. Lennon-duménil, A critical role for cell polarity in antigen extraction, processing, and presentation by B lymphocytes, Advances in Immunology, vol.123, pp.51-67, 2014.

T. Okada, M. J. Miller, I. Parker, M. F. Krummel, M. Neighbors et al., Antigen-Engaged B Cells Undergo Chemotaxis toward the T Zone and Form Motile Conjugates with Helper T Cells, PLoS Biology, vol.3, issue.6, 2005.

Q. Pcr, Extrafollicular Activation of Lymph, Science, pp.1672-1676, 2006.

K. Rajewsky, Clonal selection and learning in the antibody system, 1996.

K. Reif, E. H. Ekland, L. Ohl, H. Nakano, M. Lipp et al., Balanced responsiveness to chemoattractants from adjacent zones determines Bcell position, Nature, issue.6876, pp.94-99, 2002.

J. Riedl, A. H. Crevenna, K. Kessenbrock, J. H. Yu, D. Neukirchen et al., Lifeact: a versatile marker to visualize F-actin, Nature Methods, vol.5, issue.7, pp.605-612, 2008.

R. J. Riese, P. R. Wolf, D. Brömme, L. R. Natkin, J. A. Villadangos et al., Essential role for cathepsin S in MHC class II -Associated invariant chain processing and peptide loading, Immunity, vol.4, issue.4, pp.80249-80255, 1996.

P. Roche and P. Cresswell, Proteolysis of the class II-associated invariant chain generates a peptide binding site in intracellular HLA-DR molecules, Journal of Immunology, vol.187, issue.3, pp.1076-1080, 1950.

K. L. Rock, S. I. Haber, D. Liano, B. Benacerraf, and A. K. Abbas, Antigen presentation by hapten???specific B lymphocytes III. Analysis of the immunoglobulin???dependent pathway of antigen presentation to interleukin 1???dependent T lymphocytes, European Journal of Immunology, vol.16, issue.11, pp.1407-1412, 1986.

B. Sabass, M. L. Gardel, C. M. Waterman, and U. S. Schwarz, High resolution traction force microscopy based on experimental and computational advances, Biophysical Journal, vol.94, issue.1, pp.207-227, 2008.

A. Saci and C. L. Carpenter, RhoA GTPase regulates B cell receptor signaling, Molecular Cell, vol.17, issue.2, pp.205-214, 2005.

K. Schauer, T. Duong, K. Bleakley, S. Bardin, M. Bornens et al., Probabilistic density maps to study global endomembrane organization, Nature Methods, vol.7, issue.7, pp.560-566, 2010.

J. W. Slot and H. J. Geuze, Cryosectioning and immunolabeling, Nature Protocols, vol.2, issue.10, pp.2480-2491, 2007.

W. Song, C. Liu, and A. Upadhyaya, The pivotal position of the actin cytoskeleton in the initiation and regulation of B cell receptor activation, Biochimica et Biophysica Acta, vol.1838, issue.2, pp.569-78, 2014.

K. M. Spillane and P. Tolar, B cell antigen extraction is regulated byphysical properties of antigen presenting cells, Journal of Cell Biology, vol.2, pp.1-19, 2016.

A. Stoddart, M. L. Dykstra, B. K. Brown, W. Song, S. K. Pierce et al., Lipid rafts unite signaling cascades with clathrin to regulate BCR internalization, Immunity, vol.17, issue.4, pp.416-424, 2002.

K. Suzuki, I. Grigorova, T. G. Phan, L. M. Kelly, and J. G. Cyster, Visualizing B cell capture of cognate antigen from follicular dendritic cells, The Journal of Experimental Medicine, vol.206, issue.7, pp.1485-93, 2009.

P. Tolar, Cytoskeletal control of B cell responses to antigens, Nature Reviews Immunology, 2017.

P. Tolar, J. Hanna, P. D. Krueger, and S. K. Pierce, The constant region of the membrane immunoglobulin mediates B cell-receptor clustering and signaling in response to membrane antigens, Immunity, vol.30, issue.1, pp.44-55, 2009.

P. Tolar, H. W. Sohn, and S. K. Pierce, The initiation of antigen-induced B cell antigen receptor signaling viewed in living cells by fluorescence resonance energy transfer, Nature Immunology, vol.6, issue.11, pp.1168-1176, 2005.

P. Tolar, H. W. Sohn, and S. K. Pierce, Viewing the antigen-induced initiation of Bcell activation in living cells, Immunological Reviews, vol.221, pp.64-76, 2008.

P. Tolar and K. M. Spillane, Force generation in B-cell synapses: mechanisms coupling B-cell receptor binding to antigen internalization and affinity discrimination, Advances in immunology, vol.123, 2014.

B. Treanor, D. Depoil, A. Bruckbauer, and F. D. Batista, Dynamic cortical actin remodeling by ERM proteins controls BCR microcluster organization and integrity, The Journal of Experimental Medicine, vol.208, issue.5, pp.1055-68, 2011.

B. Treanor, D. Depoil, A. Gonzalez-granja, P. Barral, M. Weber et al., The membrane skeleton controls diffusion dynamics and signaling through the B cell receptor, Immunity, vol.32, issue.2, pp.187-99, 2010.

F. Tsai, G. Kuo, S. Chang, and P. Tsai, Ca²⁺ signaling in cytoskeletal reorganization, cell migration, and cancer metastasis, BioMed Research International, 2015.

F. C. Tsai and T. Meyer, Ca 2+ pulses control local cycles of lamellipodia retraction and adhesion along the front of migrating cells, Current Biology, vol.22, issue.9, pp.837-842, 2012.

F. Vascotto, D. Lankar, G. Faure-andre, P. Vargas, J. Diaz et al., The actin-based motor protein myosin II regulates MHC class II trafficking and BCR-driven antigen presentation, J Cell Biol, vol.176, issue.7, pp.1007-1019, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00316389

M. Vicente-manzanares, X. Ma, R. S. Adelstein, and A. R. Horwitz, Non-muscle myosin II takes centre stage in cell adhesion and migration, Nature Reviews. Molecular Cell Biology, vol.10, issue.11, pp.778-90, 2009.

Z. Wan, X. Chen, H. Chen, Q. Ji, Y. Chen et al., The activation of IgM-or isotype-switched IgG-and IgE-BCR exhibits distinct mechanical force sensitivity and threshold, vol.4, pp.1-24, 2015.

Z. Wan, S. Zhang, Y. Fan, K. Liu, F. Du et al., B cell activation is regulated by the stiffness properties of the substrate presenting the antigens, Journal of Immunology, vol.190, issue.9, pp.4661-75, 1950.

X. Wang and T. Ha, Defining single molecular forces required to activate integrin and notch signaling, Science, issue.6135, pp.991-995, 2013.

M. Weber, B. Treanor, D. Depoil, H. Shinohara, N. E. Harwood et al., Phospholipase C-gamma2 and Vav cooperate within signaling microclusters to propagate B cell spreading in response to membrane-bound antigen, The Journal of Experimental Medicine, vol.205, issue.4, pp.853-868, 2008.

B. Wickstead and K. Gull, The evolution of the cytoskeleton, Journal of Cell Biology, vol.194, issue.4, pp.513-525, 2011.

M. Yuseff, A. Reversat, D. Lankar, J. Diaz, I. Fanget et al., Polarized secretion of lysosomes at the B cell synapse couples antigen extraction to processing and presentation, Immunity, vol.35, issue.3, pp.361-74, 2011.

S. Zhang, X. Zhou, R. A. Lang, and F. Guo, RhoA of the Rho family small GTPases is essential for B lymphocyte development, PLoS ONE, vol.7, issue.3, 2012.

,