
Laboratoire Léon-Brillouin (LLB), UMR12 CEA, CNRS, Université Paris-Saclay, Gif-sur-Yvette, France
Tél. : +33 1 69 08 67 20
Email : sophie.combet@cnrs.fr
Function
CNRS Director of Research at the LLB
Scientific activity
I focus on the impact of ligand binding on protein structure through two primary projects: (i) the TSPO membrane protein, an essential neuroimaging marker with a high affinity for pharmacological ligands, and (ii) phycobiliproteins, such as C-phycocyanin, to investigate how interactions with bioactive ligands under pressure treatment influence their stability, with applications in the food industry.
I combine low-resolution techniques (SAXS/SANS), modeling approaches (e.g. ab initio, hybrid, interactive simulations), and higher-resolution structural methods (crystallography, cryo-EM) in close collaboration with I2BC (Saclay), IBPC (Paris), and MLZ (Germany).
Peer-reviewed publications
Marquette A, Nelson ARJ, Combet S*, Bechinger B. Charge-dependent insertion of the antimicrobial peptide magainin 2 into lipid monolayers probed by neutron reflectivity. BBA-Biomembranes, 1868 :184563, 2026. https://doi.org/10.1016/j.bbamem.2026.184563
Combet S*, dos Santos Morais R, Comte A, Cheron A, Barbet-Massin E, Müller-Landau H, Delalande O, Hubert JF, Charbonnier JB, Varela PF. Weak protein-bicelle binding quantification via surface-based DNA nanolevers. Eur Biophys J, 55 :379, 2026. https://doi.org/10.1007/s00249-026-01829-4
Jovanovic Z, Gligorijevic N, Annighöfer B, Dudzinski D, Levic S, Pavlovic V, Nikolic M, Brûlet A, Assifaoui A, Combet S*, Minic S. Structural and functional characteristics of beta-lactoglobulin/C-phycocyanin/starch composite gels induced by pressure. Food Hydrocol 171, 111791, 2026. https://doi.org/10.1016/j.foodhyd.2025.111791
Pozza A, Martel A, Moir M, Darwish TA, Wimalan K, Koutsioubas A, Combet S, Bonneté F. Unraveling ShuA detergent-induced colloidal behavior in solution : A comprehensive SEC-MALS, SAXS, and SANS study. Protein Science 34, e70258, 2025. https://doi.org/10.1002/pro.70258
Gligorijevic N, Svrzic N, Dojcinovic B, Stankovic D, Combet S, Minic S, Nikolic M. Optimizing alginate immobilization of food-derived C-phycocyanin : structural and functional characterization. J Serb Chem Soc, 13094, 1-14, 2025. https://doi.org/10.2298/JSC241025012G
Hotton C, Sakhawoth Y, Rollet AL, Sireix-Plénet J, Tea L, Combet S, Sharp M, Hoffmann I, Nallet F, Malikova N. Ion-specific effects in polyelectrolyte solutions : chain-chain interactions, chain rigidity and dynamics. Comptes Rendus de Chimie, 2025. https://doi.org/10.5802/crchim.326
Minic S, Velickovic L, Thureau A, Gligorijevic N, Brûlet A, Combet* S. Probing the structural stability of R-phycocyanin under pressure. Prot Sci 33:e5145, 2024. https://doi.org/10.1002/pro.5145
Bellanger T, Wien F, Combet S, Fernández Varela P, Weidmann S. The role of membrane physiology in sHSP Lo18-lipid interaction and lipochaperone activity. Sci Rep 14, 17048, 2024. https://doi.org/10.1038/s41598-024-67362-6
Saade C, Pozza A, Bonnete F, Finet S, Lutz-Bueno V, Tully MD, Varela PF, Lacapere JJ, Combet* S. Enhanced structure/function of mTSPO translocator in lipid :surfactant mixed micelles. Biochimie 224, 3, 2024. https://doi.org/10.1016/j.biochi.2024.04.008
Keshavarzi A, Asi Shirazi A, Korfanta R, Královic N, Klacsová M, Martínez JC, Teixeira J, Combet S, Uhríková D. Thermodynamic and structural study of budesonide – exogenous lung surfactant system. Int J Mol Sci 25, 2990, 2024. https://doi.org/10.3390/ijms25052990
Královic-Kanjaková N, Hubcík L, Asi Shirazi A, Klacsová M, Martínez JC, Combet S, Teixeira J, Uhríková D. Polymyxin B-enriched exogenous lung surfactant : thermodynamics and structure. Langmuir 40, 13, 6847, 2024. https://doi.org/10.1021/acs.langmuir.3c03746
Bellanger T, da Silva Barreira D, Wien F, Delarue P, Senet P, Rieu A, Neiers F, Fernández Varela P, Combet S, Weidmann S. Significant influence of four highly conserved amino-acids in lipochaperon-active sHsps on the structure and functions of the Lo18 protein. Sci Rep 13, 19036, 2023. https://doi.org/10.1038/s41598-023-46306-6
Minic S, Annighöfer B, Milcic M, Maignen F, Brûlet A, Combet* S. The effects of biliverdin on pressure-induced unfolding of apomyoglobin : The specific role of Zn2+ ions. Int J Biol Macromol 245, 125549, 2023. https://doi.org/10.1016/j.ijbiomac.2023.125549
Velickovic L, Simovic A, Gligorijevic N, Thureau A, Obradovic M, Vasovic T, Sotiroudis G, Zoumpanioti M, Brûlet A, Cirkovic Velickovic T, Combet S, Nikolic M, Minic S. Exploring and strengthening the potential of R-phycocyanin from Nori flakes as a food colourant Food Chemistry 426, 136669, 2023. https://doi.org/10.1016/j.foodchem.2023.136669
Combet* S, Bonneté F, Finet S, Pozza A, Saade C, Martel A, Koutsioubas A, Lacapère JJ. Effect of amphiphilic environment on the solution structure of mouse TSPO translocator protein. Biochimie 205, 61-72, 2023. https://doi.org/10.1016/j.biochi.2022.11.014
Minic S, Annighöfer B, Hélary A, Sago L, Cornu D, Brûlet A, Combet* S. Structure of proteins under pressure: covalent binding effects of biliverdin on beta-lactoglobulin, Biophysical J, 121, 1, 2022. https://doi.org/10.1016/j.bpj.2022.06.003
Simovic A, Combet S, Cirkovic Velickovic T, Nikolic M, Minic S. Probing the stability of the food colourant R-phycoerythrin from dried Nori flakes. Food Chemistry 374 : 131780, 2022. https://doi.org/10.1016/j.foodchem.2021.131780
Kralovic-Kanjakova N, Hubcik L, Bucsi A, Klacsova M, Combet S, Teixeira J, Martinez JC, Uhrikova D. Calcium mediated DNA binding in non-lamellar structures formed by DOPG/glycerol monooleate. Chem Phys Lipids 239 : 105118, 2021. https://doi.org/10.1016/j.chemphyslip.2021.105118
Minic S, Annighöfer B, Hélary A, Hamdane D, Hui Bon Hoa G, Loupiac C, Brûlet A, Combet* S. Effect of ligands on HP-induced unfolding and oligomerization of β-lactoglobulin. Biophysical J 119 : 2262-2274, 2020. https://doi.org/10.1016/j.bpj.2020.10.019
Marichal L, Degrouard J, Gatin A, Raffray N, Aude JC, Boulard Y, Combet S, Cousin F, Labarre J, Renault JP, Pin S. The importance of protein size in protein-nanoparticle interactions: from protein corona to colloidal self-assembly. Langmuir 36:8218, 2020. https://dx.doi.org/10.1021/acs.langmuir.0c01334
Dzubak P, Gurska S, Bogdanova K, Uhrikova D, Kanjakova N, Combet S, Klunda T, Kolar M, Hajduch M, Polakova M. Antimicrobial and cytotoxic activity of (thio)alkyl hexopyranosides, nonionic glycolipid mimetics. Carbohydrate Res 488 : 107905, 2020. https://doi.org/10.1016/j.carres.2019.107905
Mias-Lucquin D, Dos Santos Morais R, Chéron A, Chenuel T, Pérez J, Lagarrigue M, Winder S, Alviset G, Le Rumeur E, Czjzek M, Combet S, Hubert JF, Delalande O. How the central domain of dystrophin bridges F-actin to sarcolemma lipids. J Struct Biol 209 : 107411, 2020. https://doi.org/10.1016/j.jsb.2019.107411
Annighöfer B, Hélary A, Brûlet A, Colas de la Noue A, Loupiac C, Combet* S. High pressure cell to investigate protein unfolding up to 600 MPa by small-angle neutron scattering. Rev Sci Instr 90 : 025106, 2019. https://doi.org/10.1063/1.5051765
Combet S, Cousin F, Rezaei H, Noinville S. Membrane interaction of off-pathway prion oligomers and lipid-induced on-pathway intermediates during prion conversion: a clue for neurotoxicity. Biochim Biophys Acta Biomembr 1861: 514, 2019. DOI: 10.1016/j.bbamem.2018.12.001
Dos Santos Morais R, Delalande O, Pérez J, Mias-Lucquin D, Lagarrigue M, Martel A, Molza AE, Chéron A, Raguénès-Nicol C, Chenuel T, Bondon A, Appavou MS, Le Rumeur E, Combet* S, Hubert JF. Human dystrophin structural changes upon binding to anionic membrane lipid. Biophys J 115: 1231, 2018. DOI: 10.1016/j.bpj.2018.07.039
Chevreuil M, Law-Hine D, Chen J, Bressanelli S, Combet S, Constantin D, Degrouard J, Möller J, Zeghal M, Tresset G. Nonequilibrium self-assembly dynamics of icosahedral viral capsids packaging genome or polyelectrolyte. Nature Comm 9: 3071, 2018. DOI: 10.1038/s41467-018-05426-8
Chen J, Chevreuil M, Combet S, Lansac Y, Tresset G. Investigating the thermal dissociation of viral capsid by lattice model. J Phys Condens Matter 29 : 474001, 2017. https://doi.org/10.1088/1361-648X/aa8d88
Dos Santos Morais R, Delalande O, Pérez J, Mouret L, Bondon A, Martel A, Appavou MS, Le Rumeur E, Hubert JF, Combet* S. Contrast-matched isotropic bicelles : a versatile tool to specifically probe the solution structure of peripheral membrane proteins using SANS. Langmuir 33 : 6572-6580, 2017. https://doi.org/10.1021/acs.langmuir.7b01369
Golub M, Combet S, Wieland DCF, Soloviov D, Kuklin A, Lokstein H, Schmitt FJ, Olliges R, Hecht M, Eckert HJ, Pieper J. Solution structure and excitation energy transfer in phycobiliproteins of Acaryochloris marina investigated by small angle scattering. Biochim Biophys Acta 1858 : 318-324, 2017. http://dx.doi.org/10.1016/j.bbabio.2017.01.010
Spinozzi F, Ceccone G, Moretti P, Campanella G, Ferrero C, Combet S, Ojea-Jimenez I, Ghigna P. Structural and thermodynamic properties of nanoparticle-protein complexes : a combined SAXS and SANS study. Langmuir 33 : 2248-2256, 2017. https://doi.org/10.1021/acs.langmuir.6b04072
Le Coeur C, Combet* S, Carrot G, Busch P, Teixeira J, Longeville S. Conformation of the polyethylene glycol chains in diPEGylated hemoglobin specifically probed by SANS : correlation with PEG length and in vivo efficiency. Langmuir 31 : 8402-8410, 2015. https://doi.org/10.1021/acs.langmuir.5b01121
Drappier C, Oliveira H, Sandre O, Ibarboure E, Combet S, Garanger E, Lecommandoux S. Self-assembled core-shell micelles from peptide-b-polymer molecular chimeras towards structure-activity relationships. Faraday Disc 166, 83-100, 2013. https://doi.org/10.1039/c3fd00098b
Koutsioubas A, Lairez D, Combet S, Fadda GC, Longeville S, Zalczer G. Crowding effects on helix-coil transition : beyond entropic stabilization. J Chem Phys 136, 215101-7, 2012. https://dx.doi.org/10.1063/1.4723871
Combet* S, Zanotti JM. Further evidence that hydration water is the main «driving force» of protein dynamics: a neutron scattering study on perdeuterated C-phycocyanin. Phys Chem Chem Phys 14 : 4927-4934, 2012. https://doi.org/10.1039/c2cp23725c
Zanotti JM, Combet S, Klimko S, Longeville S, Coneggo F. Present and future of the quasi-elastic neutron spectroscopy at LLB. More than simply samples : devices. Neutron News 22 : 24-27, 2011. https://dx.doi.org/10.1080/10448632.2011.617284
Cousin F, Gummel J, Combet S, Boué F. The model Lysozyme-PSSNa system for electrostatic complexation : similarities and differences with complex coacervation. Advances in Colloid and Interface Science 167 : 71-84, 2011. https://doi.org/10.1016/j.cis.2011.05.007
Combet* S, Zanotti JM, Bellissent-Funel MC. Temperature- and hydration-dependent internal dynamics of stripped human erythrocyte vesicles studied by incoherent neutron scattering. Biochim Biophys Acta-General Subjects 1810 : 202-210, 2011. https://doi.org/10.1016/j.bbagen.2010.10.009
Combet* S, Pieper J, Coneggo F, Ambroise JP, Bellissent-Funel MC, Zanotti JM. Coupling of laser excitation and inelastic neutron scattering : attempt to probe light-induced dynamics of C-phycocyanin. Eur Biophys J 37 : 693-700, 2008. https://doi.org/10.1007/s00249-008-0320-1
Combet* S, Gouraud S, Gobin R, Berthonaud V, Geelen G, Corman B, Verbavatz JM. Aquaporin-2 downregulation in the medulla of aging rats is posttranscriptional and can be corrected by dehydration. Am J Physiol Renal Physiol 294 : F1408-F1414, 2008. https://doi.org/10.1152/ajprenal.00437.2007
Köper I, Combet* S, Petry W, Bellissent-Funel MC. Dynamics of C-phycocyanin in various deuterated trehalose/water environments measured by quasielastic and elastic neutron scattering. Eur Biophys J 37 : 739-748, 2008. https://doi.org/10.1007/s00249-007-0248-x
Combet S, Geffroy N, Berthonaud V, Dick B, Teillet L, Verbavatz JM, Corman B, Trinh-Trang-Tan MM. Correction of age-related polyuria by dDAVP : molecular analysis of aquaporins and urea transporters. Am J Physiol Renal Physiol 284 : F199-F208, 2003. https://doi.org/10.1152/ajprenal.00167.2002
Ferrier ML, Combet S, van Landschoot M, Stoenoiu M, Cnops Y, Lameire N, Devuyst O. Inhibition of nitric oxide synthase reverses changes in peritoneal permeability in a rat model of acute peritonitis. Kidney Int 60 : 2343-2350, 2001.
Combet S, Teillet L, Geelen G, Pitrat B, Gobin R, Nielsen S, Trinh-Trang-Tan MM, Corman B, Verbavatz JM. Food restriction prevents age-related polyuria by vasopressin-dependent recruitment of aquaporin-2. Am J Physiol Renal Physiol 281 : F1123-F1131, 2001. https://doi.org/10.1152/ajprenal.0139.2001
Combet S, Ferrier ML, van Landschoot M, Moulin P, Stoenoiu M, Moulin P, Miyata T, Lameire N, Devuyst O. Chronic uremia induces permeability changes, increased nitric oxide synthase expression, and structural modifications in the peritoneum. J Am Soc Nephrol 12 : 2146-2157, 2001. https://doi.org/10.1681/ASN.V12102146
Devuyst O, Combet S, Cnops Y, Stoenoiu. Regulation of NO synthase isoforms in the peritoneum : implications for ultrafiltration failure in peritoneal dialysis. Nephrol Dial Transplant 16 : 675-678, 2001. https://doi.org/10.1093/ndt/16.3.675
Combet S, Miyata T, Moulin P, Pouthier D, Goffin E, Devuyst O. Vascular proliferation and enhanced expression of endothelial nitric oxide synthase in human peritoneum exposed to long-term peritoneal dialysis. J Am Soc Nephrol 11 : 717-728, 2000. https://doi.org/10.1681/ASN.V114717
Combet S, Balligand JL, Lameire N, Goffin E, Devuyst O. A specific method for measurement of nitric oxide synthase enzymatic activity in peritoneal biopsies. Kidney Int 57 : 332-338, 2000. https://doi.org/10.1046/j.1523-1755.2000.00839.x
Combet S, van Landschoot M, Moulin P, Piech A, Verbavatz JM, Goffin E, Balligand JL, Lameire N, Devuyst O. Regulation of aquaporin-1 and nitric oxide synthase isoforms in a rat model of acute peritonitis. J Am Soc Nephrol, 10 : 2185-2196, 1999. https://doi.org/10.1681/ASN.V10102185
Goffin E, Combet S, Jamar F, Cosyns JP, Devuyst O. Expression of aquaporin-1 in a long-term peritoneal dialysis patient with impaired transcellular water transport. Am J Kidney Dis, 33 : 383-398, 1999. https://doi.org/10.1016/s0272-6386(99)70317-3
Book chapters
Combet* S. An introduction to neutrons for biology. In : Neutrons & Biology, Combet S and Schirò G ed., EPJ Web of Conferences vol. 236-JDN 24, 2020. https://doi.org/10.1051/epjconf/202023601001
Devuyst O, Combet S, Balligand JL, Goffin E, Verbavatz JM. Expression and regulation of aquaporin-1 and endothelial nitric oxide synthase in relationship with water permeability across the peritoneum. In : Molecular Biology and Physiology of Water and Solute Transport, Hohmann and Nielsen ed., Kluwer Academic/Plenum Publishers, pp. 69-75, New York, 2000.
Combet S, Geffroy N, Teillet L, Berthonaud V, Gobin R, Beckmann A, Corman B, Verbavatz JM. Renal aquaporin expression in aging rat : effect of food restriction. In : Molecular Biology and Physiology of Water and Solute Transport, Hohmann and Nielsen ed., Kluwer Academic/Plenum Publishers, New York, pp. 63-68, 2000.
PhD theses
Combet S. Rôle des aquaporines et de l’oxyde nitrique dans la physiopathologie du rein et du péritoine. PhD thesis, Université Paris 6, 27 septembre 2001.
Combet S. Molecular mechanisms of peritoneal permeability : interaction between nitric oxide and the water channel aquaporin-1. PhD thesis, Université catholique de Louvain (UCL), Belgique, 20 avril 2001.



