{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,12,9]],"date-time":"2025-12-09T18:11:49Z","timestamp":1765303909486,"version":"3.46.0"},"reference-count":52,"publisher":"AIP Publishing","issue":"4","funder":[{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["PTDC\/CTM-CTM\/3389\/2021"],"award-info":[{"award-number":["PTDC\/CTM-CTM\/3389\/2021"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDP\/04378\/2020"],"award-info":[{"award-number":["UIDP\/04378\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDB\/04378\/2020"],"award-info":[{"award-number":["UIDB\/04378\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["LA\/P\/0140\/2020"],"award-info":[{"award-number":["LA\/P\/0140\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["2022.11305.BD"],"award-info":[{"award-number":["2022.11305.BD"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["pubs.aip.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2025,12,1]]},"abstract":"<jats:p>Color modulation in cephalopods is driven by the coordinated action of specialized skin cells that contribute to both pigment-based coloration (chromatophores) and structural coloration (iridophores and leucophores). Reflectins, a family of proteins unique to cephalopods, are highly expressed in these biophotonic structures. In vivo, dynamic structural coloration is regulated by the reversible assembly of reflectin-based structures, which are triggered by phosphorylation events. In vitro studies have shown that reflectins can self-assemble into a variety of architectures from nano- to macro-scales, including colloidal assemblies, liquid\u2013liquid coacervates and hydrogels. Such assembly is responsive to chemical and physical stimuli, such as pH, ionic strength, and mechanical stress. In this review, we highlight the mechanisms of reflectin self-assembly in vitro, using both full-length sequences and truncated variants, and discuss emerging opportunities for engineering novel dynamic biomimetic materials.<\/jats:p>","DOI":"10.1063\/5.0294035","type":"journal-article","created":{"date-parts":[[2025,12,9]],"date-time":"2025-12-09T18:09:58Z","timestamp":1765303798000},"update-policy":"https:\/\/doi.org\/10.1063\/aip-crossmark-policy-page","source":"Crossref","is-referenced-by-count":0,"title":["Dynamic self-assembly of reflectins and their contribution to cephalopods coloration"],"prefix":"10.1063","volume":"6","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3517-9050","authenticated-orcid":false,"given":"C\u00e1tia Lopes","family":"Soares","sequence":"first","affiliation":[{"name":"Associate Laboratory i4HB \u2013 Institute for Health and Bioeconomy, School of Science and Technology, Universidade NOVA de Lisboa 1 , 2829-516 Caparica,","place":["Portugal"]},{"name":"UCIBIO \u2013 Applied Molecular Biosciences Unit, Department of Chemistry, School of Science and Technology, Universidade NOVA de Lisboa 2 , 2829-516 Caparica,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7416-7215","authenticated-orcid":false,"given":"Ana Margarida Gon\u00e7alves Carvalho","family":"Dias","sequence":"additional","affiliation":[{"name":"Associate Laboratory i4HB \u2013 Institute for Health and Bioeconomy, School of Science and Technology, Universidade NOVA de Lisboa 1 , 2829-516 Caparica,","place":["Portugal"]},{"name":"UCIBIO \u2013 Applied Molecular Biosciences Unit, Department of Chemistry, School of Science and Technology, Universidade NOVA de Lisboa 2 , 2829-516 Caparica,","place":["Portugal"]}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4586-3024","authenticated-orcid":false,"given":"Ana Cec\u00edlia Afonso","family":"Roque","sequence":"additional","affiliation":[{"name":"Associate Laboratory i4HB \u2013 Institute for Health and Bioeconomy, School of Science and Technology, Universidade NOVA de Lisboa 1 , 2829-516 Caparica,","place":["Portugal"]},{"name":"UCIBIO \u2013 Applied Molecular Biosciences Unit, Department of Chemistry, School of Science and Technology, Universidade NOVA de Lisboa 2 , 2829-516 Caparica,","place":["Portugal"]}]}],"member":"317","published-online":{"date-parts":[[2025,12,9]]},"reference":[{"issue":"35","key":"2025120913095170800_c1","doi-asserted-by":"publisher","first-page":"14862","DOI":"10.1039\/c3ra41096j","article-title":"Structural coloration in nature","volume":"3","year":"2013","journal-title":"RSC Adv."},{"issue":"99","key":"2025120913095170800_c2","doi-asserted-by":"publisher","first-page":"13448","DOI":"10.1039\/D1CC04386B","article-title":"Bio-inspired structural colors and their applications","volume":"57","year":"2021","journal-title":"Chem. Commun."},{"key":"2025120913095170800_c3","doi-asserted-by":"publisher","first-page":"111019","DOI":"10.1016\/j.dyepig.2022.111019","article-title":"Review of recent advancements in the biomimicry of structural colors","volume":"210","year":"2023","journal-title":"Dyes Pigm."},{"key":"2025120913095170800_c4","doi-asserted-by":"publisher","first-page":"66","DOI":"10.3390\/biomimetics7020066","article-title":"At the intersection of natural structural coloration and bioengineering","volume":"7","year":"2022","journal-title":"Biomimetics"},{"key":"2025120913095170800_c5","doi-asserted-by":"publisher","first-page":"S149","DOI":"10.1098\/rsif.2008.0366.focus","article-title":"Mechanisms and behavioural functions of structural coloration in cephalopods","volume":"6","year":"2009","journal-title":"J. R. Soc. Interface"},{"issue":"24","key":"2025120913095170800_c6","doi-asserted-by":"publisher","first-page":"15238","DOI":"10.1074\/jbc.M115.638254","article-title":"Structures, organization, and function of reflectin proteins in dynamically tunable reflective cells","volume":"290","year":"2015","journal-title":"J. Biol. Chem."},{"issue":"5","key":"2025120913095170800_c7","doi-asserted-by":"publisher","first-page":"793","DOI":"10.1016\/j.biomaterials.2009.10.038","article-title":"The role of protein assembly in dynamically tunable bio-optical tissues","volume":"31","year":"2010","journal-title":"Biomaterials"},{"issue":"44","key":"2025120913095170800_c8","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1098\/rsif.2009.0299","article-title":"Changes in reflectin protein phosphorylation are associated with dynamic iridescence in squid","volume":"7","year":"2010","journal-title":"J. R. Soc. Interface"},{"issue":"8","key":"2025120913095170800_c9","doi-asserted-by":"publisher","first-page":"4058","DOI":"10.1074\/jbc.M115.686014","article-title":"Cyclable condensation and hierarchical assembly of metastable reflectin proteins, the drivers of tunable biophotonics","volume":"291","year":"2016","journal-title":"J. Biol. Chem."},{"issue":"45","key":"2025120913095170800_c10","doi-asserted-by":"publisher","first-page":"16804","DOI":"10.1074\/jbc.RA119.010339","article-title":"Calibration between trigger and color: Neutralization of a genetically encoded coulombic switch and dynamic arrest precisely tune reflectin assembly","volume":"294","year":"2019","journal-title":"J. Biol. Chem."},{"issue":"173","key":"2025120913095170800_c11","doi-asserted-by":"publisher","first-page":"20200774","DOI":"10.1098\/rsif.2020.0774","article-title":"Electrochemistry as a surrogate for protein phosphorylation: Voltage-controlled assembly of reflectin A1","volume":"17","year":"2020","journal-title":"J. R. Soc. Interface"},{"issue":"204","key":"2025120913095170800_c12","doi-asserted-by":"publisher","first-page":"20230183","DOI":"10.1098\/rsif.2023.0183","article-title":"Voltage-calibrated, finely tunable protein assembly","volume":"20","year":"2023","journal-title":"J. R. Soc. Interface"},{"issue":"16","key":"2025120913095170800_c13","doi-asserted-by":"publisher","first-page":"8954","DOI":"10.3390\/ijms25168954","article-title":"Protein charge neutralization is the proximate driver dynamically tuning reflectin assembly","volume":"25","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"2025120913095170800_c14","doi-asserted-by":"publisher","first-page":"123736","DOI":"10.1016\/j.seppur.2023.123736","article-title":"A scalable method to purify reflectins from inclusion bodies","volume":"315","year":"2023","journal-title":"Sep. Purif. Technol."},{"issue":"7","key":"2025120913095170800_c15","doi-asserted-by":"publisher","first-page":"596","DOI":"10.1038\/nchem.1960","article-title":"Bulk protonic conductivity in a cephalopod structural protein","volume":"6","year":"2014","journal-title":"Nat. Chem."},{"issue":"1","key":"2025120913095170800_c16","doi-asserted-by":"publisher","first-page":"278","DOI":"10.1021\/acsami.5b08717","article-title":"Reflectin as a material for neural stem cell growth","volume":"8","year":"2016","journal-title":"ACS Appl. Mater. Interfaces"},{"issue":"39","key":"2025120913095170800_c17","doi-asserted-by":"publisher","first-page":"5621","DOI":"10.1002\/adma.201301472","article-title":"Reconfigurable infrared camouflage coatings from a cephalopod protein","volume":"25","year":"2013","journal-title":"Adv. Mater."},{"issue":"1","key":"2025120913095170800_c18","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1007\/s00441-007-0384-8","article-title":"Malleable skin coloration in cephalopods: Selective reflectance, transmission and absorbance of light by chromatophores and iridophores","volume":"329","year":"2007","journal-title":"Cell Tissue Res."},{"key":"2025120913095170800_c19","volume-title":"A Review of Visual Perception Mechanisms That Regulate Rapid Adaptive Camouflage in Cuttlefish","year":"2015"},{"issue":"1","key":"2025120913095170800_c20","doi-asserted-by":"publisher","first-page":"1004","DOI":"10.1038\/s41467-019-08891-x","article-title":"Dynamic pigmentary and structural coloration within cephalopod chromatophore organs","volume":"10","year":"2019","journal-title":"Nat Commun."},{"issue":"95","key":"2025120913095170800_c21","doi-asserted-by":"publisher","first-page":"20140106","DOI":"10.1098\/rsif.2014.0106","article-title":"Experimental determination of refractive index of condensed reflectin in squid iridocytes","volume":"11","year":"2014","journal-title":"J. R. Soc. Interface"},{"issue":"32","key":"2025120913095170800_c22","doi-asserted-by":"publisher","first-page":"3980","DOI":"10.1002\/adfm.201203705","article-title":"Bright white scattering from protein spheres in color changing, flexible cuttlefish skin","volume":"23","year":"2013","journal-title":"Adv. Funct. Mater."},{"issue":"19","key":"2025120913095170800_c23","doi-asserted-by":"publisher","first-page":"3733","DOI":"10.1242\/jeb.090415","article-title":"Dynamic biophotonics: Female squid exhibit sexually dimorphic tunable leucophores and iridocytes","volume":"216","year":"2013","journal-title":"J. Exp. Biol."},{"issue":"51","key":"2025120913095170800_c24","doi-asserted-by":"publisher","first-page":"11541","DOI":"10.1021\/acs.jpcb.0c07598","article-title":"The dynamism of intrinsically disordered proteins: Binding-induced folding, amyloid formation, and phase separation","volume":"124","year":"2020","journal-title":"J. Phys. Chem. B"},{"issue":"5655","key":"2025120913095170800_c25","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1126\/science.1091288","article-title":"Reflectins: The unusual proteins of squid reflective tissues","volume":"303","year":"2004","journal-title":"Science"},{"key":"2025120913095170800_c26","doi-asserted-by":"publisher","first-page":"69","DOI":"10.3389\/fchem.2019.00069","article-title":"Squid-inspired tandem repeat proteins: Functional fibers and films","volume":"7","year":"2019","journal-title":"Front. Chem."},{"issue":"10","key":"2025120913095170800_c27","doi-asserted-by":"publisher","first-page":"104801","DOI":"10.1063\/1.4985758","article-title":"Molecular mechanism of reflectin's tunable biophotonic control: Opportunities and limitations for new optoelectronics","volume":"5","year":"2017","journal-title":"APL Mater."},{"key":"2025120913095170800_c28","doi-asserted-by":"publisher","first-page":"101644","DOI":"10.1016\/j.mtbio.2025.101644","article-title":"Cephalopod proteins for bioinspired and sustainable biomaterials design","volume":"31","year":"2025","journal-title":"Mater. Today Bio"},{"issue":"10","key":"2025120913095170800_c29","doi-asserted-by":"publisher","first-page":"2673","DOI":"10.1021\/acs.langmuir.9b03632","article-title":"Reflectin proteins bind and reorganize synthetic phospholipid vesicles","volume":"36","year":"2020","journal-title":"Langmuir"},{"issue":"52","key":"2025120913095170800_c30","doi-asserted-by":"publisher","first-page":"32891","DOI":"10.1073\/pnas.2009044117","article-title":"Structure, self-assembly, and properties of a truncated reflectin variant","volume":"117","year":"2020","journal-title":"Proc. Nat. Acad. Sci."},{"issue":"1745","key":"2025120913095170800_c31","doi-asserted-by":"publisher","first-page":"4243","DOI":"10.1098\/rspb.2012.1374","article-title":"Neural control of tuneable skin iridescence in squid","volume":"279","year":"2012","journal-title":"Proc. R. Soc. B"},{"issue":"11","key":"2025120913095170800_c32","doi-asserted-by":"publisher","first-page":"1759","DOI":"10.1242\/jeb.00955","article-title":"The role of muscarinic receptors and intracellular Ca2+ in the spectral reflectivity changes of squid iridophores","volume":"207","year":"2004","journal-title":"J. Exp. Biol."},{"issue":"7","key":"2025120913095170800_c33","doi-asserted-by":"publisher","first-page":"2552","DOI":"10.1073\/pnas.1217260110","article-title":"Membrane invaginations facilitate reversible water flux driving tunable iridescence in a dynamic biophotonic system","volume":"110","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"2025120913095170800_c34","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1039\/D4MA00788C","article-title":"Kinetics of charge-dependent reversible condensation of reflectin nanostructures","volume":"6","year":"2025","journal-title":"Mater. Adv."},{"issue":"3","key":"2025120913095170800_c35","doi-asserted-by":"publisher","first-page":"108277","DOI":"10.1016\/j.jbc.2025.108277","article-title":"Charge screening and hydrophobicity drive progressive assembly and liquid\u2013liquid phase separation of reflectin protein","volume":"301","year":"2025","journal-title":"J. Biol. Chem."},{"issue":"7","key":"2025120913095170800_c36","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1038\/nmat1930","article-title":"The self-organizing properties of squid reflectin protein","volume":"6","year":"2007","journal-title":"Nat. Mater."},{"issue":"33","key":"2025120913095170800_c37","doi-asserted-by":"publisher","first-page":"13279","DOI":"10.1039\/D4SC03299C","article-title":"Reversible and size-controlled assembly of reflectin proteins using a charged azobenzene photoswitch","volume":"15","year":"2024","journal-title":"Chem. Sci."},{"issue":"5","key":"2025120913095170800_c38","doi-asserted-by":"publisher","first-page":"1214","DOI":"10.1110\/ps.051840806","article-title":"pK values of the ionizable groups of proteins","volume":"15","year":"2006","journal-title":"Protein Sci."},{"issue":"2","key":"2025120913095170800_c39","doi-asserted-by":"publisher","first-page":"978","DOI":"10.1021\/acsbiomaterials.2c00088","article-title":"Squid skin cell-inspired refractive index mapping of cells, vesicles, and nanostructures","volume":"9","year":"2023","journal-title":"ACS Biomater. Sci. Eng."},{"issue":"10","key":"2025120913095170800_c40","doi-asserted-by":"publisher","first-page":"101113","DOI":"10.1063\/5.0214285","article-title":"Bulk proton conduction in films from a truncated reflectin variant","volume":"12","year":"2024","journal-title":"APL Mater."},{"key":"2025120913095170800_c41","doi-asserted-by":"publisher","first-page":"100049","DOI":"10.1016\/j.supmat.2023.100049","article-title":"Liquid-liquid phase separation: Fundamental physical principles, biological implications, and applications in supramolecular materials engineering","volume":"2","year":"2023","journal-title":"Supramol. Mater."},{"key":"2025120913095170800_c42","doi-asserted-by":"publisher","first-page":"e24331","DOI":"10.1002\/pep2.24331","article-title":"A compositional view comparing modern biological condensates and primitive phase-separated compartments","volume":"115","year":"2023","journal-title":"Peptide Sci."},{"key":"2025120913095170800_c43","doi-asserted-by":"publisher","first-page":"3065","DOI":"10.1016\/j.bpj.2024.07.004","article-title":"A colloidal model for the equilibrium assembly and liquid-liquid phase separation of the reflectin A1 protein","volume":"123","year":"2024","journal-title":"Biophys J."},{"issue":"1","key":"2025120913095170800_c44","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1007\/BF00571425","article-title":"Physiological color change in squid iridophores","volume":"259","year":"1990","journal-title":"Cell Tissue Res."},{"key":"2025120913095170800_c45","unstructured":"R.\n            Gordon\n          , see https:\/\/escholarship.org\/uc\/item\/75d61930 for \u201cFour is not a crowd: Unexpected multiphase separation of reflectin proteins\u201d (University of California, Santa Barbara, 2023)."},{"issue":"2","key":"2025120913095170800_c46","doi-asserted-by":"publisher","first-page":"430","DOI":"10.3390\/molecules26020430","article-title":"A global review on short peptides: Frontiers and perspectives","volume":"26","year":"2021","journal-title":"Molecules"},{"issue":"4","key":"2025120913095170800_c47","doi-asserted-by":"publisher","first-page":"254","DOI":"10.1002\/polb.23204","article-title":"Recombinant reflectin-based optical materials","volume":"51","year":"2013","journal-title":"J. Polym. Sci., Part B: Polym. Phys."},{"issue":"18","key":"2025120913095170800_c48","doi-asserted-by":"publisher","first-page":"2833","DOI":"10.1016\/j.cub.2017.07.061","article-title":"Origin of the reflectin gene and hierarchical assembly of its protein","volume":"27","year":"2017","journal-title":"Curr. Biol."},{"key":"2025120913095170800_c49","doi-asserted-by":"publisher","first-page":"1054347","DOI":"10.3389\/fchem.2022.1054347","article-title":"Solvent modulation in peptide sub-microfibers obtained by solution blow spinning","volume":"10","year":"2022","journal-title":"Front. Chem."},{"key":"2025120913095170800_c50","doi-asserted-by":"publisher","first-page":"1267563","DOI":"10.3389\/fchem.2023.1267563","article-title":"Hierarchical self-assembly of a reflectin-derived peptide","volume":"11","year":"2023","journal-title":"Front. Chem."},{"issue":"38","key":"2025120913095170800_c51","doi-asserted-by":"publisher","first-page":"8405","DOI":"10.1002\/adma.201601666","article-title":"Self-assembly of the cephalopod protein reflectin","volume":"28","year":"2016","journal-title":"Adv. Mater."},{"issue":"18","key":"2025120913095170800_c52","doi-asserted-by":"publisher","first-page":"20938","DOI":"10.1021\/acsami.0c18929","article-title":"Long-range proton transport in films from a reflectin-derived polypeptide","volume":"13","year":"2021","journal-title":"ACS Appl. Mater. Interfaces"}],"container-title":["Biophysics Reviews"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.aip.org\/aip\/bpr\/article-pdf\/doi\/10.1063\/5.0294035\/20833235\/041302_1_5.0294035.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.aip.org\/aip\/bpr\/article-pdf\/doi\/10.1063\/5.0294035\/20833235\/041302_1_5.0294035.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,9]],"date-time":"2025-12-09T18:10:00Z","timestamp":1765303800000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.aip.org\/bpr\/article\/6\/4\/041302\/3374503\/Dynamic-self-assembly-of-reflectins-and-their"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,1]]},"references-count":52,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2025,12,1]]}},"URL":"https:\/\/doi.org\/10.1063\/5.0294035","relation":{},"ISSN":["2688-4089"],"issn-type":[{"value":"2688-4089","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2025,12]]},"published":{"date-parts":[[2025,12,1]]},"article-number":"041302"}}