{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T14:02:55Z","timestamp":1791468175052,"version":"4.3.4"},"reference-count":51,"publisher":"American Chemical Society (ACS)","issue":"40","license":[{"start":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T00:00:00Z","timestamp":1790380800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T00:00:00Z","timestamp":1790380800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T00:00:00Z","timestamp":1790380800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/100000181","name":"Air Force Office of Scientific Research","doi-asserted-by":"publisher","award":["FA9550-18-1-0395"],"award-info":[{"award-number":["FA9550-18-1-0395"]}],"id":[{"id":"10.13039\/100000181","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000181","name":"Air Force Office of Scientific Research","doi-asserted-by":"publisher","award":["FA9550-20-1-0324"],"award-info":[{"award-number":["FA9550-20-1-0324"]}],"id":[{"id":"10.13039\/100000181","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000181","name":"Air Force Office of Scientific Research","doi-asserted-by":"publisher","award":["FA9550-23-1-0583"],"award-info":[{"award-number":["FA9550-23-1-0583"]}],"id":[{"id":"10.13039\/100000181","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100021856","name":"Ministero dell'Universit? e della Ricerca","doi-asserted-by":"publisher","award":["CN_00000041"],"award-info":[{"award-number":["CN_00000041"]}],"id":[{"id":"10.13039\/501100021856","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100021856","name":"Ministero dell'Universit? e della Ricerca","doi-asserted-by":"publisher","award":["PE0000006"],"award-info":[{"award-number":["PE0000006"]}],"id":[{"id":"10.13039\/501100021856","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,8]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Aquaporin-4 (AQP4) is an integral membrane protein channel specific for water molecules. It is highly polarized in the endfeet of astrocytes in the central nervous system where it allows water\u2013ion homeostasis. AQP4 has two N-terminal isoforms: M1 (long) and M23 (short). Both isoforms retain their function but differ in their N-terminal lengths, which regulate AQP4 aggregation size. There are also long and short isoforms at the C-terminus. The long form has a 29-amino-acid extension (AQP4ex) that interacts with \u03b1-syntrophin, a component of the dystrophin-associated protein complex (DAPC). This interaction anchors AQP4 to the perivascularastrocyte endfeet through the DAPC, thereby maintaining its polarized localization at the blood\u2013brain barrier. This study uses peptides with the sequence of the extended region to demonstrate a strong interaction with \u03b1-syntrophin, bound to a glass surface via maleimide-PEG silane chemistry. It was observed that substituting alanine for threonine, four amino acids from the C-terminus, eliminates binding, and an alanine scan shows that the last six amino acids at the C-terminus also play a crucial role. Kinetic studies using time-lapse single-molecule fluorescence imaging were used to determine the Gibbs free energy of activation for both forward and reverse rates (\u0394G\u2021) using an Eyring analysis. Equilibrium constants (Keq) were measured between 15 and 37 \u00b0C, and standard-state thermodynamic free energies (\u0394G\u00b0), enthalpies (\u0394H\u00b0), and entropies (\u0394S\u00b0) were determined using a van\u2019t Hoff analysis. It was found that forward rates ranged from 4.5 \u00d7 104 to 1.0 \u00d7 105 M\u20131 s\u20131 depending on peptide length, with \u0394G\u2021forward = 45\u201346 kJ mol\u20131. Reverse rates ranged from 11.6 to 21.1 s\u20131 depending on peptide length, with \u0394G\u2021reverse = 65.2\u201367.0 kJ mol\u20131. Using a vant\u2019 Hoff analysis, the overall standard-state free energies were between \u221219 and \u221222 kJ mol\u20131. The work also describes how dynamic single-molecule imaging can be used to determine kinetic and thermodynamic properties.<\/jats:p>","DOI":"10.1021\/acs.jpcb.6c04328","type":"journal-article","created":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T22:01:26Z","timestamp":1790460086000},"page":"10427-10437","source":"Crossref","is-referenced-by-count":0,"title":["AQP4 Extended Region Binds to \u03b1-Syntrophin\nwith a Novel Binding Motif"],"prefix":"10.1021","volume":"130","author":[{"given":"Eric P.","family":"Jacobo","sequence":"first","affiliation":[{"name":"Washington State University , , P.O. Box 644630, , ,","place":["Pullman, WA, United States, 99164-4630"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael J.","family":"Martinez","sequence":"additional","affiliation":[{"name":"Washington State University , , P.O. Box 644630, , ,","place":["Pullman, WA, United States, 99164-4630"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5093-6662","authenticated-orcid":true,"given":"Barbara","family":"Barile","sequence":"additional","affiliation":[{"name":"University of Bari Aldo Moro , , ,","place":["Bari, Italy, 70124"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alessia","family":"Memeo","sequence":"additional","affiliation":[{"name":"University of Bari Aldo Moro , , ,","place":["Bari, Italy, 70124"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Grazia Paola","family":"Nicchia","sequence":"additional","affiliation":[{"name":"University of Bari Aldo Moro , , ,","place":["Bari, Italy, 70124"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2097-5051","authenticated-orcid":true,"given":"James A.","family":"Brozik","sequence":"additional","affiliation":[{"name":"Washington State University , , P.O. Box 644630, , ,","place":["Pullman, WA, United States, 99164-4630"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,9,26]]},"reference":[{"issue":"2","key":"2026100809094601400_cit1","doi-asserted-by":"publisher","first-page":"90","DOI":"10.3390\/cells8020090","article-title":"Aquaporin-4 Water Channel in the Brain and Its Implication\nfor Health and Disease","volume":"8","author":"Mader","year":"2019","journal-title":"Cells"},{"key":"2026100809094601400_cit2","doi-asserted-by":"publisher","first-page":"654865","DOI":"10.3389\/fgene.2021.654865","article-title":"Human Aquaporins: Functional Diversity and Potential\nRoles in Infectious and Non-infectious Diseases","volume":"12","author":"Azad","year":"2021","journal-title":"Front. Genet."},{"issue":"4","key":"2026100809094601400_cit3","doi-asserted-by":"publisher","first-page":"1036","DOI":"10.1016\/j.neuroscience.2009.08.019","article-title":"Aquaporin-4\nin brain and spinal cord oedema","volume":"168","author":"Saadoun","year":"2010","journal-title":"Neuroscience"},{"issue":"1","key":"2026100809094601400_cit4","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1093\/brain\/awab311","article-title":"Emerging\nroles for dynamic aquaporin-4 subcellular relocalization in CNS water\nhomeostasis","volume":"145","author":"Salman","year":"2022","journal-title":"Brain"},{"issue":"5","key":"2026100809094601400_cit5","doi-asserted-by":"publisher","first-page":"1603","DOI":"10.3390\/ijms21051603","article-title":"Regulation of AQP4 in the Central\nNervous System","volume":"21","author":"Vandebroek","year":"2020","journal-title":"Int. J. Mol. Sci."},{"issue":"1","key":"2026100809094601400_cit6","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1093\/brain\/awab311","article-title":"Emerging\nroles for dynamic aquaporin-4 subcellular relocalization in CNS water\nhomeostasis","volume":"145","author":"Salman","year":"2022","journal-title":"Brain"},{"issue":"1","key":"2026100809094601400_cit7","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1186\/1742-2094-5-22","article-title":"Neuromyelitis\noptica\npathogenesis and aquaporin 4","volume":"5","author":"Graber","year":"2008","journal-title":"J. Neuroinflammation"},{"issue":"1","key":"2026100809094601400_cit8","doi-asserted-by":"publisher","DOI":"10.1212\/NXI.0000000000000062","article-title":"Does MOG Ig-positive AQP4seronegative\nopticospinal\ninflammatory disease justify a diagnosis of NMO spectrum disorder?","volume":"2","author":"Zamvil","year":"2015","journal-title":"Neurol. Neuroimmunol. Neuroinflammation"},{"issue":"18","key":"2026100809094601400_cit9","doi-asserted-by":"publisher","first-page":"7437","DOI":"10.1073\/pnas.0902725106","article-title":"Crystal structure of human aquaporin\n4 at 1.8 A and its mechanism of conductance","volume":"106","author":"Ho","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"issue":"11","key":"2026100809094601400_cit10","doi-asserted-by":"publisher","first-page":"3010","DOI":"10.1016\/j.bpj.2009.09.017","article-title":"Reversible, temperaturedependent supramolecular assembly\nof aquaporin-4 orthogonal arrays in live cell membranes","volume":"97","author":"Crane","year":"2009","journal-title":"Biophys. J."},{"issue":"6","key":"2026100809094601400_cit11","doi-asserted-by":"publisher","first-page":"813","DOI":"10.1242\/jcs.042341","article-title":"Determinants of aquaporin-4 assembly\nin orthogonal\narrays revealed by live-cell single-molecule fluorescence imaging","volume":"122","author":"Crane","year":"2009","journal-title":"J. Cell Sci."},{"issue":"12","key":"2026100809094601400_cit12","doi-asserted-by":"publisher","first-page":"2936","DOI":"10.1016\/j.bpj.2011.05.012","article-title":"Model of Aquaporin-4 Supramolecular\nAssembly in Orthogonal Arrays Based on Heterotetrameric Association\nof M1-M23 Isoforms","volume":"100","author":"Jin","year":"2011","journal-title":"Biophys. 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Commun."},{"issue":"5","key":"2026100809094601400_cit17","doi-asserted-by":"publisher","first-page":"790","DOI":"10.1002\/glia.23126","article-title":"Translational read through generates\nnew astrocyte AQP4 isoforms that modulate supramolecular clustering,\nglial endfeet localization, and water transport","volume":"65","author":"De Bellis","year":"2017","journal-title":"Glia"},{"issue":"4","key":"2026100809094601400_cit18","doi-asserted-by":"publisher","first-page":"2106","DOI":"10.1073\/pnas.0437946100","article-title":"An alpha-syntrophin-dependent\npool of AQP4 in astroglial end-feet confers bidirectional water flow\nbetween blood and brain","volume":"100","author":"Amiry-Moghaddam","year":"2003","journal-title":"Proc. Natl. Acad. 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U.S.A."},{"issue":"5","key":"2026100809094601400_cit21","doi-asserted-by":"publisher","first-page":"675","DOI":"10.1016\/0092-8674(95)90344-5","article-title":"Three muscular\ndystrophies: loss of cytoskeleton-extracellular matrix linkage","volume":"80","author":"Campbell","year":"1995","journal-title":"Cell"},{"issue":"2","key":"2026100809094601400_cit22","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1083\/jcb.139.2.375","article-title":"Animal models for\nmuscular dystrophy show different patterns of sarcolemmal disruption","volume":"139","author":"Straub","year":"1997","journal-title":"J. Cell Biol."},{"issue":"8","key":"2026100809094601400_cit23","doi-asserted-by":"publisher","first-page":"879","DOI":"10.1002\/glia.20347","article-title":"Assembly\nof a perivascular astrocyte protein scaffold at the mammalian blood-brain\nbarrier is dependent on alpha-syntrophin","volume":"53","author":"Bragg","year":"2006","journal-title":"Glia"},{"key":"2026100809094601400_cit24","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1016\/S0896-6273(00)80894-6","article-title":"Maturation and maintenance\nof the neuromuscular synapse:\nGenetic evidence for roles of the dystrophin--glycoprotein complex","volume":"25","author":"Grady","year":"2000","journal-title":"Neuron"},{"key":"2026100809094601400_cit25","doi-asserted-by":"publisher","first-page":"1691","DOI":"10.1021\/acs.jpcb.1c09358","article-title":"Thermodynamic Driving\nForces of Redox-Dependent CPR\nInsertion into Biomimetic Endoplasmic Reticulum Membranes","volume":"126","author":"Martinez","year":"2022","journal-title":"J. Phys. Chem. B"},{"issue":"12","key":"2026100809094601400_cit26","doi-asserted-by":"publisher","first-page":"3038","DOI":"10.1021\/acs.jpcb.5b11804","article-title":"Substrate\nDependent Native Luminescence from Cytochromes P450 3A4, 2C9, and\nP450cam","volume":"120","author":"Barnaba","year":"2016","journal-title":"J. Phys. Chem. B"},{"issue":"1","key":"2026100809094601400_cit27","doi-asserted-by":"publisher","first-page":"320","DOI":"10.1021\/la104133m","article-title":"Single Particle\nTracking Reveals\nCorralling of a Transmembrane Protein in a Double-Cushioned Lipid\nBilayer Assembly","volume":"27","author":"Poudel","year":"2011","journal-title":"Langmuir"},{"issue":"3","key":"2026100809094601400_cit28","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1021\/acs.jpcb.3c04529","article-title":"Thermodynamics and\nS-Palmitoylation Dependence of Interactions between Human Aquaporin-4\nM1 Tetramers in Model Membranes","volume":"128","author":"Carder","year":"2024","journal-title":"J. Phys. Chem.\nB"},{"issue":"1","key":"2026100809094601400_cit29","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1006\/jcis.1996.0217","article-title":"Methods of Digital\nVideo Microscopy for Colloidal Studies","volume":"179","author":"Crocker","year":"1996","journal-title":"J. Colloid\nInterface Sci."},{"key":"2026100809094601400_cit30","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1016\/j.ymeth.2016.09.016","article-title":"TrackMate:\nAn open and extensible platform for single-particle tracking","volume":"115","author":"Tinevez","year":"2017","journal-title":"Methods"},{"issue":"7","key":"2026100809094601400_cit31","doi-asserted-by":"publisher","first-page":"829","DOI":"10.1038\/s41592-022-01507-1","article-title":"TrackMate 7: integrating\nstate-of-the-art segmentation algorithms into tracking pipelines","volume":"19","author":"Ershov","year":"2022","journal-title":"Nat. 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Biol."},{"key":"2026100809094601400_cit40","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1186\/1478-811X-8-8","article-title":"PDZ. domains and\ntheir binding partners: structure,\nspecificity, and modification","volume":"8","author":"Lee","year":"2010","journal-title":"Cell Commun.\nSignal."},{"key":"2026100809094601400_cit41","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.crstbi.2021.01.001","article-title":"Integrated analysis of Shank1 PDZ interactions with\nC-terminal and internal binding motifs","volume":"3","author":"Ali","year":"2021","journal-title":"Curr.\nRes. Struct. Biol."},{"issue":"2","key":"2026100809094601400_cit42","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1063\/1.1749604","article-title":"The Activated\nComplex in Chemical Reactions","volume":"3","author":"Henry","year":"1935","journal-title":"J. Chem. Phys."},{"key":"2026100809094601400_cit43","doi-asserted-by":"publisher","first-page":"790","DOI":"10.1002\/glia.23126","article-title":"Translational readthrough generates\nnew astrocyte AQP4 isoforms that modulate supramolecular clustering,\nglial endfeet localization, and water transport","volume":"65","author":"De Bellis","year":"2017","journal-title":"Glia"},{"key":"2026100809094601400_cit44","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1016\/bs.ircmb.2018.05.013","article-title":"Emerging Themes\nin PDZ Domain Signaling: Structure,\nFunction, and Inhibition","volume":"343","author":"Liu","year":"2019","journal-title":"Int. Rev. Cell Mol.\nBiol."},{"issue":"4","key":"2026100809094601400_cit45","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1038\/nrn3468","article-title":"Aquaporin water\nchannels in the nervous system","volume":"14","author":"Papadopoulos","year":"2013","journal-title":"Nat. Rev. Neurosci."},{"key":"2026100809094601400_cit46","doi-asserted-by":"publisher","first-page":"103080","DOI":"10.1016\/j.sbi.2025.103080","article-title":"Entropy, enthalpy,\nand evolution: Adaptive trade-offs\nin protein binding thermodynamics","volume":"94","author":"Rosemary","year":"2025","journal-title":"Curr. Opin.\nStruct. Biol."},{"key":"2026100809094601400_cit47","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1146\/annurev-biophys-083012-130318","article-title":"Entropy-enthalpy\ncompensation: role and ramifications\nin biomolecular ligand recognition and design","volume":"42","author":"Chodera","year":"2013","journal-title":"Annu. Rev. Biophys."},{"key":"2026100809094601400_cit48","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1186\/s40478-019-0707-5","article-title":"AQP4ex is crucial for\nthe anchoring of AQP4 at the astrocyte end-feet and for neuromyelitis\noptica antibody binding","volume":"7","author":"Palazzo","year":"2019","journal-title":"Acta Neuropathol. Commun."},{"issue":"1","key":"2026100809094601400_cit49","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1186\/s40478-024-01870-4","article-title":"Crucial role of Aquaporin-4\nextended\nisoform in brain water Homeostasis and Amyloid-\u03b2 clearance:\nimplications for Edema and neurodegenerative diseases","volume":"12","author":"Abbrescia","year":"2024","journal-title":"Acta Neuropathol. Commun."},{"issue":"4","key":"2026100809094601400_cit50","doi-asserted-by":"publisher","first-page":"1543","DOI":"10.1152\/physrev.00011.2013","article-title":"Physiological roles of aquaporin-4\nin brain","volume":"93","author":"Nagelhus","year":"2013","journal-title":"Physiol. Rev."},{"issue":"3","key":"2026100809094601400_cit51","doi-asserted-by":"publisher","first-page":"1117","DOI":"10.3390\/ijms23031117","article-title":"Disassembly\nand Mislocalization of AQP4 in Incipient Scar Formation after Experimental\nStroke","volume":"23","author":"Banitalebi","year":"2022","journal-title":"Int. J. Mol. Sci."}],"container-title":["The Journal of Physical Chemistry B"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acs.jpcb.6c04328","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/jpcbfk\/article-pdf\/130\/40\/10427\/68668164\/acs.jpcb.6c04328.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/jpcbfk\/article-pdf\/130\/40\/10427\/68668164\/acs.jpcb.6c04328.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T13:30:07Z","timestamp":1791466207000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/jpcbfk\/article\/130\/40\/10427\/5439443\/AQP4-Extended-Region-Binds-to-Syntrophin-with-a"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,26]]},"references-count":51,"journal-issue":{"issue":"40","published-online":{"date-parts":[[2026,9,26]]},"published-print":{"date-parts":[[2026,10,8]]}},"URL":"https:\/\/doi.org\/10.1021\/acs.jpcb.6c04328","relation":{},"ISSN":["1520-6106","1520-5207"],"issn-type":[{"value":"1520-6106","type":"print"},{"value":"1520-5207","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,26]]}}}