{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,26]],"date-time":"2026-10-26T13:34:03Z","timestamp":1793021643957,"version":"4.5.0"},"reference-count":95,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T00:00:00Z","timestamp":1790726400000},"content-version":"vor","delay-in-days":29,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000053","name":"National Eye Institute","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000053","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100001818","name":"Research to Prevent Blindness","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100001818","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Biophysical Journal"],"published-print":{"date-parts":[[2026,9]]},"DOI":"10.1016\/j.bpj.2026.09.043","type":"journal-article","created":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T23:07:35Z","timestamp":1790809655000},"update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["Mechanisms of protein transport between subcompartments of the cone photoreceptor outer segment membrane system"],"prefix":"10.1016","author":[{"given":"Alexey A.","family":"Tokarev","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9162-4924","authenticated-orcid":false,"given":"Peter D.","family":"Calvert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.bpj.2026.09.043_bib1","series-title":"Handbook of Biological Physics","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/S1383-8121(00)80008-1","article-title":"Phototransduction in vertebrate rods and cones: Molecular mechanisms of amplification, recovery and light adaptation","author":"Pugh","year":"2000"},{"key":"10.1016\/j.bpj.2026.09.043_bib2","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1016\/S0065-3233(03)63010-X","article-title":"Rhodopsin structure, dynamics, and activation: a perspective from crystallography, site-directed spin labeling, sulfhydryl reactivity, and disulfide cross-linking","volume":"63","author":"Hubbell","year":"2003","journal-title":"Adv. Protein Chem."},{"key":"10.1016\/j.bpj.2026.09.043_bib3","doi-asserted-by":"crossref","first-page":"878350","DOI":"10.3389\/fcell.2022.878350","article-title":"Patterning and Development of Photoreceptors in the Human Retina","volume":"10","author":"Hussey","year":"2022","journal-title":"Front. Cell Dev. Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib4","doi-asserted-by":"crossref","first-page":"4585","DOI":"10.1113\/JP282058","article-title":"Photoreceptor physiology and evolution: cellular and molecular basis of rod and cone phototransduction","volume":"600","author":"Lamb","year":"2022","journal-title":"J. Physiol."},{"key":"10.1016\/j.bpj.2026.09.043_bib5","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/0005-2736(89)90519-1","article-title":"Concentration effects on reactions in membranes: rhodopsin and transducin","volume":"979","author":"Saxton","year":"1989","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2026.09.043_bib6","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/0005-2728(93)90038-H","article-title":"Amplification and kinetics of the activation steps in phototransduction","volume":"1141","author":"Pugh","year":"1993","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2026.09.043_bib7","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1083\/jcb.75.2.490","article-title":"Turnover of rod photoreceptor outer segments: I. membrane addition and loss in relationship to temperature","volume":"75","author":"Hollyfield","year":"1977","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib8","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1083\/jcb.75.2.507","article-title":"Turnover of rod photoreceptor outer segments. II. Membrane addition and loss in relationship to light","volume":"75","author":"Besharse","year":"1977","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib9","first-page":"117","article-title":"Mammalian cones: disc shedding phagocytosis, and renewal","volume":"17","author":"Anderson","year":"1978","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib10","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1002\/cne.901900307","article-title":"Disc morphogenesis in vertebrate photoreceptors","volume":"190","author":"Steinberg","year":"1980","journal-title":"J. Comp. Neurol."},{"key":"10.1016\/j.bpj.2026.09.043_bib11","doi-asserted-by":"crossref","first-page":"2267","DOI":"10.1083\/jcb.105.5.2267","article-title":"Cone outer segment morphogenesis: Taper change and distal invaginations","volume":"105","author":"Eckmiller","year":"1987","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib12","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1016\/S1350-9462(96)00026-2","article-title":"Morphogenesis and renewal of cone outer segments","volume":"16","author":"Eckmiller","year":"1997","journal-title":"Prog. Retin. Eye Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib13","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1002\/cne.902870410","article-title":"Modulation of disk margin structure during renewal of cone outer segments in the vertebrate retina","volume":"287","author":"Corless","year":"1989","journal-title":"J. Comp. Neurol."},{"key":"10.1016\/j.bpj.2026.09.043_bib14","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1083\/jcb.201508093","article-title":"Discs of mammalian rod photoreceptors form through the membrane evagination mechanism","volume":"211","author":"Ding","year":"2015","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib15","doi-asserted-by":"crossref","first-page":"jcs229526","DOI":"10.1242\/jcs.229526","article-title":"The route of the visual receptor rhodopsin along the cilium","volume":"132","author":"Chadha","year":"2019","journal-title":"J. Cell Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib16","doi-asserted-by":"crossref","first-page":"1493","DOI":"10.1007\/s00424-021-02558-7","article-title":"Functional compartmentalization of photoreceptor neurons","volume":"473","author":"Malhotra","year":"2021","journal-title":"Pflugers Arch."},{"key":"10.1016\/j.bpj.2026.09.043_bib17","doi-asserted-by":"crossref","first-page":"636737","DOI":"10.3389\/fcell.2021.636737","article-title":"Compartmentalization of Photoreceptor Sensory Cilia","volume":"9","author":"Barnes","year":"2021","journal-title":"Front. Cell Dev. Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib18","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1083\/jcb.39.1.169","article-title":"The renewal of protein in retinal rods and cones","volume":"39","author":"Young","year":"1968","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib19","first-page":"222","article-title":"A difference between rods and cones in the renewal of outer segment protein","volume":"8","author":"Young","year":"1969","journal-title":"Investig. Ophthalmol."},{"key":"10.1016\/j.bpj.2026.09.043_bib20","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/0042-6989(72)90108-3","article-title":"The renewal of diffusely distributed protein in the outer segments of rods and cones","volume":"12","author":"Bok","year":"1972","journal-title":"Vision Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib21","doi-asserted-by":"crossref","first-page":"525","DOI":"10.1016\/j.bpj.2010.11.085","article-title":"A simplified mass-transfer model for visual pigments in amphibian retinal-cone outer segments","volume":"100","author":"Weber","year":"2011","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2026.09.043_bib22","doi-asserted-by":"crossref","first-page":"2697","DOI":"10.1016\/j.bpj.2012.04.052","article-title":"Cone outer segments: A biophysical model of membrane dynamics, shape retention, and lamella formation","volume":"102","author":"Corless","year":"2012","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2026.09.043_bib23","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/S0092-8674(00)00023-4","article-title":"Genetic evidence for selective transport of opsin and arrestin by kinesin-II in mammalian photoreceptors","volume":"102","author":"Marszalek","year":"2000","journal-title":"Cell"},{"key":"10.1016\/j.bpj.2026.09.043_bib24","doi-asserted-by":"crossref","first-page":"4866","DOI":"10.1096\/fj.15-275677","article-title":"Kinesin family 17 (osmotic avoidance abnormal-3) is dispensable for photoreceptor morphology and function","volume":"29","author":"Jiang","year":"2015","journal-title":"FASEB J."},{"key":"10.1016\/j.bpj.2026.09.043_bib25","doi-asserted-by":"crossref","first-page":"3049","DOI":"10.1242\/jcs.01167","article-title":"Quantification of the cytoplasmic spaces of living cells with EGFP reveals arrestin-EGFP to be in disequilibrium in dark adapted rod photoreceptors","volume":"117","author":"Peet","year":"2004","journal-title":"J. Cell Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib26","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1085\/jgp.200910322","article-title":"Diffusion of a soluble protein, photoactivatable GFP, through a sensory cilium","volume":"135","author":"Calvert","year":"2010","journal-title":"J. Gen. Physiol."},{"key":"10.1016\/j.bpj.2026.09.043_bib27","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1111\/j.1365-2818.2007.01715.x","article-title":"Fluorescence relaxation in 3D from diffraction-limited sources of PAGFP or sinks of EGFP created by multiphoton photoconversion","volume":"225","author":"Calvert","year":"2007","journal-title":"J. Microsc."},{"key":"10.1016\/j.bpj.2026.09.043_bib28","doi-asserted-by":"crossref","first-page":"1479","DOI":"10.1016\/j.febslet.2006.01.080","article-title":"Conserved cis-elements in the Xenopus red opsin promoter necessary for cone-specific expression","volume":"580","author":"Babu","year":"2006","journal-title":"FEBS Lett."},{"key":"10.1016\/j.bpj.2026.09.043_bib29","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/S0014-5793(98)00018-0","article-title":"Transgene expression in Xenopus rods","volume":"423","author":"Knox","year":"1998","journal-title":"FEBS Lett."},{"key":"10.1016\/j.bpj.2026.09.043_bib30","doi-asserted-by":"crossref","first-page":"3173","DOI":"10.1242\/dev.122.10.3173","article-title":"Transgenic Xenopus embryos from sperm nuclear transplantations reveal FGF signaling requirements during gastrulation","volume":"122","author":"Kroll","year":"1996","journal-title":"Development"},{"key":"10.1016\/j.bpj.2026.09.043_bib31","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0030101","article-title":"Rhodopsin Mutant P23H Destabilizes Rod Photoreceptor Disk Membranes","volume":"7","author":"Haeri","year":"2012","journal-title":"PLoS One"},{"key":"10.1016\/j.bpj.2026.09.043_bib32","doi-asserted-by":"crossref","first-page":"2831","DOI":"10.1083\/jcb.201711104","article-title":"Actin filaments partition primary cilia membranes into distinct fluid corrals","volume":"217","author":"Lee","year":"2018","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib33","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1073\/pnas.1115109109","article-title":"Steric volume exclusion sets soluble protein concentrations in photoreceptor sensory cilia","volume":"109","author":"Najafi","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2026.09.043_bib34","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1085\/jgp.201210818","article-title":"Impact of signaling microcompartment geometry on GPCR dynamics in live retinal photoreceptors","volume":"140","author":"Najafi","year":"2012","journal-title":"J. Gen. Physiol."},{"key":"10.1016\/j.bpj.2026.09.043_bib35","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1083\/jcb.140.4.821","article-title":"Rapid Diffusion of Green Fluorescent Protein in the Mitochondrial Matrix","volume":"140","author":"Partikian","year":"1998","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib36","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1016\/S0076-6879(99)02024-8","article-title":"Green fluorescent protein as a probe to study intracellular solute diffusion","volume":"302","author":"Verkman","year":"1999","journal-title":"Methods Enzymol."},{"key":"10.1016\/j.bpj.2026.09.043_bib37","first-page":"646","article-title":"Morphological components associated with frog cone outer segment disc margins","volume":"28","author":"Fetter","year":"1987","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib38","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1167\/iovs.11-8635","article-title":"Prominin-1 localizes to the open rims of outer segment lamellae in xenopus laevis rod and cone photoreceptors","volume":"53","author":"Han","year":"2012","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib39","article-title":"Distinct roles for prominin-1 and photoreceptor cadherin in outer segment disc morphogenesis in CRISPR-altered X. laevis","volume":"134","author":"Carr","year":"2021","journal-title":"J. Cell Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib40","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1016\/j.exer.2009.03.023","article-title":"Focus on molecules: RDS","volume":"89","author":"Conley","year":"2009","journal-title":"Exp. Eye Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib41","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.72817","article-title":"Determinants shaping the nanoscale architecture of the mouse rod outer segment","volume":"10","author":"P\u00f6ge","year":"2021","journal-title":"eLife"},{"key":"10.1016\/j.bpj.2026.09.043_bib42","doi-asserted-by":"crossref","first-page":"10577","DOI":"10.1074\/jbc.R100005200","article-title":"The Influence of Macromolecular Crowding and Macromolecular Confinement on Biochemical Reactions in Physiological Media","volume":"276","author":"Minton","year":"2001","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.bpj.2026.09.043_bib43","doi-asserted-by":"crossref","first-page":"1835","DOI":"10.1016\/j.bpj.2011.08.031","article-title":"Finite platelet size could be responsible for platelet margination effect","volume":"101","author":"Tokarev","year":"2011","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2026.09.043_bib44","series-title":"The Mathematics of Diffusion","author":"Crank","year":"1975"},{"key":"10.1016\/j.bpj.2026.09.043_bib45","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1042\/bst0230341","article-title":"The Control of Flux: 21 Years On","volume":"23","author":"Kacser","year":"1995","journal-title":"Biochem. Soc. Trans."},{"key":"10.1016\/j.bpj.2026.09.043_bib46","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1007\/BF01816949","article-title":"Control theory of metabolic channelling","volume":"143","author":"Kholodenko","year":"1995","journal-title":"Mol. Cell. Biochem."},{"key":"10.1016\/j.bpj.2026.09.043_bib47","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/0303-2647(75)90050-7","article-title":"Mathematicalanalysis of multienzyme systems. 2. Steady state and transient control","volume":"7","author":"Heinrich","year":"1975","journal-title":"Biosystems"},{"key":"10.1016\/j.bpj.2026.09.043_bib48","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1016\/S0022-5320(64)80084-8","article-title":"Receptor cell outer segment development and ultrastructure of the disk membranes in the retina of the tadpole (Rana pipiens)","volume":"11","author":"Nilsson","year":"1964","journal-title":"J. Ultrastruct. Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib49","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/0014-4835(86)90061-8","article-title":"A concentration of fucosylated glycoconjugates at the base of cone outer segments: Quantitative electron microscope autoradiography","volume":"42","author":"Anderson","year":"1986","journal-title":"Exp. Eye Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib50","first-page":"179","article-title":"Shifting distribution of autoradiographic label in cone outer segments and its implications for renewal (only abstract is available)","volume":"34","author":"Eckmiller","year":"1993","journal-title":"J. Hirnforsch."},{"key":"10.1016\/j.bpj.2026.09.043_bib51","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1126\/science.185.4149.457","article-title":"Lateral Diffusion of Visual Pigment in Rod Disk Membranes","volume":"185","author":"Liebman","year":"1974","journal-title":"Science"},{"key":"10.1016\/j.bpj.2026.09.043_bib52","series-title":"Birefringence, Dichroism and Rod Outer Segment Structure BT - Photoreceptor Optics","first-page":"199","author":"Liebman","year":"1975"},{"key":"10.1016\/j.bpj.2026.09.043_bib53","doi-asserted-by":"crossref","first-page":"2589","DOI":"10.1083\/jcb.105.6.2589","article-title":"Differences in the Protein Composition of Bovine Retinal Rod Outer Segment Disk and Plasma Membranes Isolated by a Ricin-Gold-Dextran Density Perturbation Method","volume":"105","author":"Molday","year":"1987","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib54","first-page":"50","article-title":"A rim-specific membrane protein of rod outer segment discs","volume":"28","author":"Molday","year":"1987","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib55","first-page":"5996","article-title":"Single molecule tracking of opsins in living Xenopus cone photoreceptors","volume":"67","author":"Lovy","year":"2026","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib56","series-title":"Mathematical Biology. I. An Introduction","author":"Murray","year":"2002"},{"key":"10.1016\/j.bpj.2026.09.043_bib57","doi-asserted-by":"crossref","first-page":"2185","DOI":"10.1016\/0042-6989(73)90221-6","article-title":"Optical function of human peripheral cones","volume":"13","author":"Miller","year":"1973","journal-title":"Vision Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib58","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1016\/S0014-4835(61)80018-3","article-title":"The fine structure of the extrafoveal receptors of the rhesus monkey","volume":"1","author":"Cohen","year":"1961","journal-title":"Exp. Eye Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib59","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1126\/science.147.3653.57","article-title":"Foveal Receptors of the Monkey Retina: Fine Structure","volume":"147","author":"Dowling","year":"1965","journal-title":"Science"},{"key":"10.1016\/j.bpj.2026.09.043_bib60","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1016\/0042-6989(70)90001-5","article-title":"Further studies on the question of the patency of saccules in outer segments of vertebrate photoreceptors","volume":"10","author":"Cohen","year":"1970","journal-title":"Vision Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib61","doi-asserted-by":"crossref","unstructured":"Tokarev A., Ratto N., Volpert V. Mathematical modeling of thrombin generation and wave propagation: from simple to complex models and backwards. BIOMAT 2018 International Symposium on Mathematical and Computational Biology. Springer; 2019. p. 1\u201322. https:\/\/link.springer.com\/chapter\/10.1007\/978-3-030-23433-1_1.","DOI":"10.1007\/978-3-030-23433-1_1"},{"key":"10.1016\/j.bpj.2026.09.043_bib62","first-page":"1717","article-title":"Lateral diffusion of rhodopsin in photoreceptor membrane: a reappraisal","volume":"15","author":"Govardovskii","year":"2009","journal-title":"Mol. Vis."},{"key":"10.1016\/j.bpj.2026.09.043_bib63","first-page":"788","article-title":"Quantification of interlamellar transport of intrinsic membrane proteins in living Xenopus cone photoreceptor outer segments","volume":"66","author":"Lovy","year":"2025","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"10.1016\/j.bpj.2026.09.043_bib64","doi-asserted-by":"crossref","first-page":"3111","DOI":"10.1073\/pnas.72.8.3111","article-title":"Brownian motion in biological membranes","volume":"72","author":"Saffman","year":"1975","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2026.09.043_bib65","doi-asserted-by":"crossref","first-page":"4317","DOI":"10.1073\/pnas.79.14.4317","article-title":"Lateral and rotational diffusion of bacteriorhodopsin in lipid bilayers: Experimental test of the Saffman-Delbruck equations","volume":"79","author":"Peters","year":"1982","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2026.09.043_bib66","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1016\/S0006-3495(82)84467-6","article-title":"Lateral diffusion of gramicidin C in phospholipid multibilayers. Effects of cholesterol and high gramicidin concentration","volume":"40","author":"Tank","year":"1982","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2026.09.043_bib67","doi-asserted-by":"crossref","first-page":"12650","DOI":"10.1021\/ja902853g","article-title":"Lateral Diffusion of Membrane Proteins","volume":"131","author":"Ramadurai","year":"2009","journal-title":"J. Am. Chem. Soc."},{"key":"10.1016\/j.bpj.2026.09.043_bib68","doi-asserted-by":"crossref","first-page":"4308","DOI":"10.1021\/acs.jpclett.7b01758","article-title":"Diffusion of Integral Membrane Proteins in Protein-Rich Membranes","volume":"8","author":"Javanainen","year":"2017","journal-title":"J. Phys. Chem. Lett."},{"key":"10.1016\/j.bpj.2026.09.043_bib69","doi-asserted-by":"crossref","first-page":"2630","DOI":"10.1021\/acs.jctc.3c00060","article-title":"Protein Crowding and Cholesterol Increase Cell Membrane Viscosity in a Temperature Dependent Manner","volume":"19","author":"F\u00e1bi\u00e1n","year":"2023","journal-title":"J. Chem. Theory Comput."},{"key":"10.1016\/j.bpj.2026.09.043_bib70","doi-asserted-by":"crossref","first-page":"989","DOI":"10.1016\/S0006-3495(87)83291-5","article-title":"Lateral diffusion in an archipelago. The effect of mobile obstacles","volume":"52","author":"Saxton","year":"1987","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2026.09.043_bib71","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/S1383-8121(00)80007-X","article-title":"Chapter 4 Ion channels of vertebrate photoreceptors","volume":"3","author":"Molday","year":"2000","journal-title":"Handb. Biol. Phys."},{"key":"10.1016\/j.bpj.2026.09.043_bib72","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.bbamem.2008.08.022","article-title":"Ordering effects of cholesterol and its analogues","volume":"1788","author":"R\u00f3g","year":"2009","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"10.1016\/j.bpj.2026.09.043_bib73","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.plipres.2005.02.001","article-title":"The role of cholesterol in rod outer segment membranes","volume":"44","author":"Albert","year":"2005","journal-title":"Prog. Lipid Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib74","doi-asserted-by":"crossref","first-page":"20727","DOI":"10.1016\/S0021-9258(17)45275-6","article-title":"Cholesterol modulation of photoreceptor function in bovine retinal rod outer segments","volume":"265","author":"Boesze-Battaglia","year":"1990","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.bpj.2026.09.043_bib75","doi-asserted-by":"crossref","first-page":"8151","DOI":"10.1016\/S0021-9258(18)83162-3","article-title":"Cholesterol heterogeneity in bovine red outer segment disk membranes","volume":"264","author":"Boesze-Battaglia","year":"1989","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.bpj.2026.09.043_bib76","doi-asserted-by":"crossref","first-page":"e80059","DOI":"10.1371\/journal.pone.0080059","article-title":"Regulation of rhodopsin-eGFP distribution in transgenic Xenopus rod outer segments by light","volume":"8","author":"Haeri","year":"2013","journal-title":"PLoS One"},{"key":"10.1016\/j.bpj.2026.09.043_bib77","first-page":"E2937","article-title":"In vivo optophysiology reveals that G-protein activation triggers osmotic swelling and increased light scattering of rod photoreceptors","volume":"114","author":"Zhang","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.bpj.2026.09.043_bib78","doi-asserted-by":"crossref","first-page":"336","DOI":"10.1016\/0005-2736(75)90275-8","article-title":"X-ray diffraction studies of retinal rods. II. Light effect on the osmotic properties","volume":"382","author":"Chabre","year":"1975","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2026.09.043_bib79","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/S0022-5320(65)80016-8","article-title":"The ultrastructure of the receptor outer segments in the retina of the leopard frog (Rana pipiens)","volume":"12","author":"Nilsson","year":"1965","journal-title":"J. Ultrastruct. Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib80","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1038\/235044a0","article-title":"X-ray Diffraction from Outer Segments of Visual Cells in Intact Eyes of the Frog","volume":"235","author":"Webb","year":"1972","journal-title":"Nature"},{"key":"10.1016\/j.bpj.2026.09.043_bib81","doi-asserted-by":"crossref","first-page":"917","DOI":"10.1083\/jcb.200612010","article-title":"Three-dimensional architecture of murine rod outer segments determined by cryoelectron tomography","volume":"177","author":"Nickell","year":"2007","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib82","doi-asserted-by":"crossref","first-page":"906","DOI":"10.1038\/223906a0","article-title":"Structure of frog photoreceptor membranes","volume":"223","author":"Blaurock","year":"1969","journal-title":"Nature"},{"key":"10.1016\/j.bpj.2026.09.043_bib83","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1038\/newbio244118a0","article-title":"Light induced changes of ionic flux in the retinal rod","volume":"244","author":"Chabre","year":"1973","journal-title":"Nat. New Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib84","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/0005-2736(75)90274-6","article-title":"X-ray diffraction studies of retinal rods. I. Structure of the disc membrane, effect of illumination","volume":"382","author":"Chabre","year":"1975","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.bpj.2026.09.043_bib85","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/262266a0","article-title":"Neutron diffraction studies of retinal rod outer segment membranes","volume":"262","author":"Saibil","year":"1976","journal-title":"Nature"},{"key":"10.1016\/j.bpj.2026.09.043_bib86","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/0022-2836(80)90319-8","article-title":"Neutron diffraction analysis of the structure of rod photoreceptor membranes in intact retinas","volume":"137","author":"Yeager","year":"1980","journal-title":"J. Mol. Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib87","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1098\/rspb.1978.0036","article-title":"The photoreceptors and pigment epithelim of the adult Xenopus retina: morphology and outer segment renewal","volume":"201","author":"Kinney","year":"1978","journal-title":"Proc.R.Soc.Lond.B."},{"key":"10.1016\/j.bpj.2026.09.043_bib88","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/cne.902570102","article-title":"Structural features of the terminal loop region of frog retinal rod outer segment disk membranes: I. Organization of Lipid Components","volume":"257","author":"Corless","year":"1987","journal-title":"J. Comp. Neurol."},{"key":"10.1016\/j.bpj.2026.09.043_bib89","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1016\/j.exer.2015.05.014","article-title":"Preparation of Xenopus laevis retinal cryosections for electron microscopy","volume":"136","author":"Tam","year":"2015","journal-title":"Exp. Eye Res."},{"key":"10.1016\/j.bpj.2026.09.043_bib90","doi-asserted-by":"crossref","first-page":"1029","DOI":"10.1016\/j.cell.2012.10.038","article-title":"Three-dimensional architecture of the rod sensory cilium and its disruption in retinal neurodegeneration","volume":"151","author":"Gilliam","year":"2012","journal-title":"Cell"},{"key":"10.1016\/j.bpj.2026.09.043_bib91","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1016\/j.str.2015.01.015","article-title":"Higher-order architecture of rhodopsin in intact photoreceptors and its implication for phototransduction kinetics","volume":"23","author":"Gunkel","year":"2015","journal-title":"Structure"},{"key":"10.1016\/j.bpj.2026.09.043_bib92","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/S0006-3495(62)86847-7","article-title":"In Situ Microspectrophotometric Studies on the Pigments of Single Retinal Rods","volume":"2","author":"Liebman","year":"1962","journal-title":"Biophys. J."},{"key":"10.1016\/j.bpj.2026.09.043_bib93","doi-asserted-by":"crossref","first-page":"7761","DOI":"10.1021\/bi0155091","article-title":"Advances in determination of a high-resolution three-dimensional structure of rhodopsin, a model of G-protein-coupled receptors (GPCRs)","volume":"40","author":"Teller","year":"2001","journal-title":"Biochemistry"},{"key":"10.1016\/j.bpj.2026.09.043_bib94","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1007\/s11538-024-01274-4","article-title":"Nonlinear Regression Modelling: A Primer with Applications and Caveats","volume":"86","author":"O\u2019Brien","year":"2024","journal-title":"Bull. Math. Biol."},{"key":"10.1016\/j.bpj.2026.09.043_bib95","doi-asserted-by":"crossref","first-page":"2941","DOI":"10.1016\/j.bpj.2025.07.036","article-title":"Inferring diffusion, reaction, and exchange parameters from imperfect FRAP","volume":"124","author":"Lorenzetti","year":"2025","journal-title":"Biophys. J."}],"container-title":["Biophysical Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0006349526006892?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0006349526006892?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,26]],"date-time":"2026-10-26T12:44:40Z","timestamp":1793018680000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0006349526006892"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":95,"alternative-id":["S0006349526006892"],"URL":"https:\/\/doi.org\/10.1016\/j.bpj.2026.09.043","relation":{},"ISSN":["0006-3495"],"issn-type":[{"value":"0006-3495","type":"print"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Mechanisms of protein transport between subcompartments of the cone photoreceptor outer segment membrane system","name":"articletitle","label":"Article Title"},{"value":"Biophysical Journal","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.bpj.2026.09.043","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Author(s). Published by Elsevier Inc. on behalf of Biophysical Society.","name":"copyright","label":"Copyright"}]}}