244:
221:
118:
143:
495:
250:
149:
1189:
1096:
protein linked together to form a domain. The TMDs are usually not conserved across genomes due to its specificity and diversity in function as channels or ligand-binding controllers. However, NBDs are highly conserved across different genomes—an observation consistent with which it binds and hydrolyzes ATP. NBD binds
1279:-retinyl-phosphatidyl-ethanolamine (A2PE). After diurnal disk-shedding and phagocytosis of outer segment by RPE cells, A2PE is hydrolyzed inside the RPE phagolysosome to form A2E. Accumulation of A2E causes toxicity at the primary RPE level and secondary photoreceptor destruction in macular degenerations.
2108:
Allikmets R, Singh N, Sun H, Shroyer NF, Hutchinson A, Chidambaram A, Gerrard B, Baird L, Stauffer D, Peiffer A, Rattner A, Smallwood P, Li Y, Anderson KL, Lewis RA, Nathans J, Leppert M, Dean M, Lupski JR (March 1997). "A photoreceptor cell-specific ATP-binding transporter gene (ABCR) is mutated in
1941:
Beaty TH, Ruczinski I, Murray JC, Marazita ML, Munger RG, Hetmanski JB, Murray T, Redett RJ, Fallin MD, Liang KY, Wu T, Patel PJ, Jin SC, Zhang TX, Schwender H, Wu-Chou YH, Chen PK, Chong SS, Cheah F, Yeow V, Ye X, Wang H, Huang S, Jabs EW, Shi B, Wilcox AJ, Lie RT, Jee SH, Christensen K, Doheny KF,
1449:
Allikmets R, Singh N, Sun H, Shroyer NF, Hutchinson A, Chidambaram A, Gerrard B, Baird L, Stauffer D, Peiffer A, Rattner A, Smallwood P, Li Y, Anderson KL, Lewis RA, Nathans J, Leppert M, Dean M, Lupski JR (March 1997). "A photoreceptor cell-specific ATP-binding transporter gene (ABCR) is mutated in
1216:
knockout mouse has delayed dark adaptation but normal final rod threshold relative to controls. This suggests bulk transmembrane diffusion pathways that remove ATR/NR-PE from extracellular membranes. After bleaching the retina with strong light, ATR/NR-PE accumulates significantly in outer segments.
1095:
Previously known as the photoreceptor rim protein RmP or ABCR, the recently proposed ABCA4 structure consists of two transmembrane domains (TMDs), two large glycosylated extracytosolic domains (ECD), and two internal nucleotide binding domains (NBDs). One TMD spans across membranes with six units of
1313:
with multiple markers giving evidence of linkage and association at the genome-wide significance level. Although SNPs in this gene are associated with cleft lip/palate there is no functional or expression data to support it as the causal gene which may, instead, lie in a region adjacent to ABCA4. A
1116:
ligands and control channel access to its binding sites. Once an ATP is hydrolized at the NBDs of the channel, NBDs are brought together to tilt and modify TMDs to modulate ligand binding to the channel. A recently proposed model of retinoid transfer occurring as a result of alternating exposure of
1172:
to an NBD to bring two NBDs together and expose outer vestibule high affinity binding site located in TMD, (2) binding of NR-PE/ATR on extracellular side of the channel, (3) ATP hydrolysis promoting gate opening and movement of NR-PE/ATR across the membrane to the low-affinity binding site on the
1258:
disease causing progressive loss of photoreceptor cells. STGD is characterized by reduced visual acuity and color vision, loss of central (macular) vision, delayed dark adaptation, and accumulation of autofluorescent RPE lipofuscin. Removal of NR-PE/ATR appears to be significant in normal bleach
2457:
Gerber S, Rozet JM, van de Pol TJ, Hoyng CB, Munnich A, Blankenagel A, Kaplan J, Cremers FP (February 1998). "Complete exon-intron structure of the retina-specific ATP binding transporter gene (ABCR) allows the identification of novel mutations underlying
Stargardt disease".
2146:
Martínez-Mir A, Bayés M, Vilageliu L, Grinberg D, Ayuso C, del Río T, García-Sandoval B, Bussaglia E, Baiget M, Gonzàlez-Duarte R, Balcells S (February 1997). "A new locus for autosomal recessive retinitis pigmentosa (RP19) maps to 1p13-1p21".
1164:(ATR) with phosphatidylethanolamine (PE), trapped inside the disk as charged species out to the cytoplasmic surface. Once transported, ATR is reduced to vitamin A and then transferred to retinal pigment epithelium to be recycled into 11-
2715:
Körschen HG, Beyermann M, Müller F, Heck M, Vantler M, Koch KW, Kellner R, Wolfrum U, Bode C, Hofmann KP, Kaupp UB (August 1999). "Interaction of glutamic-acid-rich proteins with the cGMP signalling pathway in rod photoreceptors".
2336:
Martínez-Mir A, Paloma E, Allikmets R, Ayuso C, del Rio T, Dean M, Vilageliu L, Gonzàlez-Duarte R, Balcells S (January 1998). "Retinitis pigmentosa caused by a homozygous mutation in the
Stargardt disease gene ABCR".
2419:
Nasonkin I, Illing M, Koehler MR, Schmid M, Molday RS, Weber BH (January 1998). "Mapping of the rod photoreceptor ABC transporter (ABCR) to 1p21-p22.1 and identification of novel mutations in
Stargardt's disease".
1493:
Nasonkin I, Illing M, Koehler MR, Schmid M, Molday RS, Weber BH (January 1998). "Mapping of the rod photoreceptor ABC transporter (ABCR) to 1p21-p22.1 and identification of novel mutations in
Stargardt's disease".
1263:
signaling that causes photoreceptors to degenerate. ABCA4 also mitigates long-term effects of accumulation of ATR that results in irreversible ATR binding to a second molecule of ATR and NR-PE to form dihydro-
1242:. From this experiment, it was concluded that ABCR has a significant role in clearing accumulation of ATR/NR-PE to prevent formation of A2E in extracellular photoreceptor surfaces during bleach recovery.
2637:
Maugeri A, van Driel MA, van de Pol DJ, Klevering BJ, van Haren FJ, Tijmes N, Bergen AA, Rohrschneider K, Blankenagel A, Pinckers AJ, Dahl N, Brunner HG, Deutman AF, Hoyng CB, Cremers FP (April 1999).
2639:"The 2588G→C mutation in the ABCR gene is a mild frequent founder mutation in the Western European population and allows the classification of ABCR mutations in patients with Stargardt disease"
2487:
Azarian SM, Megarity CF, Weng J, Horvath DH, Travis GH (June 1998). "The human photoreceptor rim protein gene (ABCR): genomic structure and primer set information for mutation analysis".
1993:"Expression and mutation analyses implicate ARHGAP29 as the etiologic gene for the cleft lip with or without cleft palate locus identified by genome-wide association on chromosome 1p22"
2384:
Cremers FP, van de Pol DJ, van Driel M, den
Hollander AI, van Haren FJ, Knoers NV, Tijmes N, Bergen AA, Rohrschneider K, Blankenagel A, Pinckers AJ, Deutman AF, Hoyng CB (March 1998).
1156:. Flippase is a transmembrane protein that "flips" its conformation to transport materials across a membrane. In the case of ABCA4, the flippase facilitates transfer of N-retinyl-
3058:
257:
156:
2604:"Retinal stimulates ATP hydrolysis by purified and reconstituted ABCR, the photoreceptor-specific ATP-binding cassette transporter responsible for Stargardt disease"
1112:: the two dimerized compartments of the channel are different from each other. When TMDs are situated in a membrane, they form a barrel-like structure permeable to
2972:
2303:
Allikmets R, Shroyer NF, Singh N, Seddon JM, Lewis RA, Bernstein PS, Peiffer A, Zabriskie NA, Li Y, Hutchinson A, Dean M, Lupski JR, Leppert M (September 1997).
1537:
Allikmets R, Shroyer NF, Singh N, Seddon JM, Lewis RA, Bernstein PS, Peiffer A, Zabriskie NA, Li Y, Hutchinson A, Dean M, Lupski JR, Leppert M (September 1997).
3235:
2877:
2845:
1117:
external and internal TMD ligand binding sites, all controlled by binding of ATP, is based on recent structural analyses of bacterial ABC transporters.
763:
2186:
Azarian SM, Travis GH (June 1997). "The photoreceptor rim protein is an ABC transporter encoded by the gene for recessive
Stargardt's disease (ABCR)".
3033:
79:
744:
1625:
van Meer G, Halter D, Sprong H, Somerharju P, Egmond MR (February 2006). "ABC lipid transporters: extruders, flippases, or flopless activators?".
1812:
1601:
1414:
2855:
1991:
Leslie EJ, Mansilla MA, Biggs LC, Schuette K, Bullard S, Cooper M, Dunnwald M, Lidral AC, Marazita ML, Beaty TH, Murray JC (November 2012).
3230:
1740:"Insights into the function of Rim protein in photoreceptors and etiology of Stargardt's disease from the phenotype in abcr knockout mice"
1396:
2901:
243:
3208:
2818:
2386:"Autosomal recessive retinitis pigmentosa and cone-rod dystrophy caused by splice site mutations in the Stargardt's disease gene ABCR"
970:
1787:
Molday RS, Beharry S, Ahn J, Zhong M (2006). "Binding of N-Retinylidene-Pe to BACA4 and a Model for its
Transport Across Membranes".
963:
220:
1335:
1299:
1239:
1048:
1435:
1383:
1362:
2870:
1379:
50:, AW050280, Abc10, Abcr, D430003I15Rik, RmP, ARMD2, CORD3, FFM, RP19, STGD, STGD1, ATP binding cassette subfamily A member 4
2830:
2561:"Genotype/Phenotype analysis of a photoreceptor-specific ATP-binding cassette transporter gene, ABCR, in Stargardt disease"
1254:
gene are known to cause the autosomal-recessive disease
Stargardt macular dystrophy (STGD), which is a hereditary juvenile
142:
117:
1271:-retinyl-phosphatidyl-ethanolamine (A2PE-H2). A2PE-H2 traps ATR and accumulates in outer segments to further oxidize into
1358:
59:
2270:"A photoreceptor cell-specific ATP-binding transporter gene (ABCR) is mutated in recessive Stargardt macular dystrophy"
2231:
Sun H, Nathans J (September 1997). "Stargardt's ABCR is localized to the disc membrane of retinal rod outer segments".
256:
155:
2761:
Zhang K, Garibaldi DC, Kniazeva M, Albini T, Chiang MF, Kerrigan M, Sunness JS, Han M, Allikmets R (December 1999).
2894:
249:
148:
2863:
2524:
Rozet JM, Gerber S, Souied E, Perrault I, Châtelin S, Ghazi I, Leowski C, Dufier JL, Munnich A, Kaplan J (1998).
2803:
2682:"Variation of clinical expression in patients with Stargardt dystrophy and sequence variations in the ABCR gene"
2812:
1157:
808:
67:
2559:
Lewis RA, Shroyer NF, Singh N, Allikmets R, Hutchinson A, Li Y, Lupski JR, Leppert M, Dean M (February 1999).
1315:
789:
1185:) to release the bound ligand. The channel is then ready to transfer another molecule of NR-PE/ATR again.
1169:
1097:
3063:
1287:
1174:
2725:
2195:
1634:
1295:
1255:
1060:
131:
2836:
2824:
2808:
46:
2749:
2512:
2445:
2372:
2256:
2219:
2134:
1769:
1668:
1519:
1475:
1291:
1084:
91:
1585:
1577:
946:
925:
899:
878:
2763:"A novel mutation in the ABCR gene in four patients with autosomal recessive Stargardt disease"
1944:"Evidence for gene-environment interaction in a genome wide study of nonsyndromic cleft palate"
2784:
2741:
2703:
2668:
2625:
2590:
2547:
2504:
2475:
2437:
2407:
2364:
2324:
2291:
2248:
2211:
2174:
2126:
2096:
2061:
2022:
1973:
1923:
1874:
1818:
1808:
1761:
1717:
1660:
1607:
1597:
1558:
1511:
1467:
1056:
39:
2886:
2774:
2733:
2693:
2658:
2650:
2615:
2580:
2572:
2537:
2496:
2467:
2429:
2397:
2354:
2346:
2316:
2281:
2240:
2203:
2164:
2156:
2118:
2086:
2053:
2012:
2004:
1963:
1955:
1913:
1905:
1864:
1854:
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1751:
1707:
1699:
1650:
1642:
1589:
1550:
1503:
1459:
336:
267:
211:
166:
1067:
gene transcribes a large retina-specific protein with two transmembrane domains (TMD), two
494:
87:
2897:
2890:
1796:
1789:
1326:
gene to be the likely causal gene playing a role in nonsyndromic cleft lip and/or palate.
311:
1318:, rare coding sequence variants, craniofacial specific expression, and interactions with
1168:-retinal. This alternating access-release model for ABCA4 has four steps: (1) binding of
2729:
2199:
1638:
2967:
2663:
2638:
2585:
2560:
2017:
1992:
1968:
1943:
1918:
1893:
1869:
1842:
1712:
1687:
1072:
2779:
2762:
2207:
2057:
1756:
1739:
1593:
678:
673:
668:
663:
658:
653:
648:
643:
638:
622:
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612:
607:
602:
597:
592:
576:
571:
566:
561:
556:
551:
546:
541:
536:
3224:
2849:
1688:"Focus on molecules: ABCA4 (ABCR)--an import-directed photoreceptor retinoid flipase"
1149:
1068:
523:
2516:
2449:
2138:
1791:
Binding of N-retinylidene-PE to ABCA4 and a model for its transport across membranes
1773:
1672:
1578:"ABCR: Rod photoreceptor-specific ABC transporter responsible for Stargardt disease"
1523:
1479:
2753:
2305:"Mutation of the Stargardt disease gene (ABCR) in age-related macular degeneration"
2223:
1539:"Mutation of the Stargardt disease gene (ABCR) in age-related macular degeneration"
1310:
329:
108:
71:
2376:
2320:
2260:
1646:
1554:
1305:
The GENEVA Cleft
Consortium study first identified ABCA4 as being associated with
2044:
MacDonald IM (June 2005). "Genetic aspects of age-related macular degeneration".
95:
2680:
Fishman GA, Stone EM, Grover S, Derlacki DJ, Haines HL, Hockey RR (April 1999).
1419:
National Center for
Biotechnology Information, U.S. National Library of Medicine
1401:
National Center for Biotechnology Information, U.S. National Library of Medicine
1109:
2698:
2681:
1703:
412:
1076:
228:
125:
75:
1804:
1079:-binding domains (NBD). The ABCA4 protein is almost exclusively expressed in
2620:
2603:
2542:
2526:"Spectrum of ABCR gene mutations in autosomal recessive macular dystrophies"
2525:
2075:"Normalization and subtraction: two approaches to facilitate gene discovery"
1306:
1178:
1141:
1133:
1101:
1055:. The gene was first cloned and characterized in 1997 as a gene that causes
1052:
708:
472:
350:
295:
282:
194:
181:
83:
2788:
2745:
2707:
2672:
2629:
2471:
2160:
2065:
2026:
1977:
1927:
1878:
1859:
1822:
1765:
1721:
1664:
1611:
2594:
2551:
2508:
2500:
2479:
2441:
2433:
2411:
2402:
2385:
2368:
2328:
2295:
2252:
2215:
2178:
2130:
2100:
1894:"Cleft lip and palate: understanding genetic and environmental influences"
1562:
1515:
1507:
1471:
1010:
1005:
3186:
1795:. Advances in Experimental Medicine and Biology. Vol. 572. pp.
1738:
Weng J, Mata NL, Azarian SM, Tzekov RT, Birch DG, Travis GH (July 1999).
1323:
1153:
1129:
1113:
994:
853:
834:
2286:
2269:
1436:"Entrez Gene: ABCA4 ATP-binding cassette, sub-family A (ABC1), member 4"
1173:
intracellular portion of TMD, and (4) release of adenosine diphosphate (
1104:
to carry out change in conformation of the ABC transporter. Transcribed
2359:
2169:
2122:
2091:
2074:
2008:
1959:
1463:
1161:
1034:
820:
775:
2350:
2244:
1843:"Retinopathy in mice induced by disrupted all-trans-retinal clearance"
1655:
3087:
3082:
3077:
3012:
2951:
2946:
1148:
to other ABCs, cellular localization of ABCA4, and analyses of ABCA4
1080:
978:
730:
2304:
1909:
1538:
2654:
2576:
3194:
3190:
3181:
3176:
3171:
3155:
3150:
3134:
3118:
3113:
3108:
3103:
3071:
3067:
3053:
3048:
3043:
3038:
3028:
3007:
3002:
2997:
2992:
2987:
2941:
2936:
2926:
2921:
2916:
2737:
1260:
1187:
1997:
Birth Defects Research. Part A, Clinical and Molecular Teratology
693:
689:
2981:
2976:
1319:
1041:
63:
2859:
1152:
suggest that ABCA4 may function as an inward-directed retinoid
2804:
GeneReviews/NIH/NCBI/UW entry on Retinitis Pigmentosa Overview
1188:
503:
1841:
Maeda A, Maeda T, Golczak M, Palczewski K (September 2008).
1051:
gene sub-family A (ABC1) found exclusively in multicellular
1892:
Dixon MJ, Marazita ML, Beaty TH, Murray JC (March 2011).
1582:
Vertebrate Phototransduction and the Visual Cycle, Part A
1144:, approximately at 1:120. Comparisons between mammalian
319:
1217:
This accumulation leads to formation of toxic cationic
552:
ATPase-coupled intramembrane lipid transporter activity
1100:
molecules (ATP) to utilize the high-energy inorganic
484:
3164:
3143:
3127:
3096:
3021:
2960:
2909:
1023:
ATP-binding cassette, sub-family A (ABC1), member 4
939:
918:
892:
871:
2073:Bonaldo MF, Lennon G, Soares MB (September 1996).
1788:
1375:
1373:
1371:
1354:
1352:
1350:
1282:Additional diseases that may link to mutations in
1584:. Methods in Enzymology. Vol. 315. pp.
537:ATPase-coupled transmembrane transporter activity
266:
165:
1380:GRCm38: Ensembl release 89: ENSMUSG00000028125
2871:
1059:, an autosomal recessive disease that causes
8:
1942:Pugh EW, Ling H, Scott AF (September 2011).
1836:
1834:
1832:
1128:is localized to outer segment disk edges of
1733:
1731:
1359:GRCh38: Ensembl release 89: ENSG00000198691
2878:
2864:
2856:
2846:United States National Library of Medicine
2602:Sun H, Molday RS, Nathans J (March 1999).
1083:localizing in outer segment disk edges of
704:
562:phosphatidylethanolamine flippase activity
519:
307:
206:
103:
2811:at the U.S. National Library of Medicine
2778:
2697:
2662:
2619:
2584:
2541:
2401:
2358:
2285:
2168:
2090:
2016:
1967:
1917:
1868:
1858:
1755:
1711:
1654:
2844:This article incorporates text from the
2109:recessive Stargardt macular dystrophy".
1450:recessive Stargardt macular dystrophy".
1430:
1428:
1160:(NR-PE), a covalent adduct of all-trans
598:intracellular membrane-bounded organelle
1346:
16:Mammalian protein found in Homo sapiens
20:
613:integral component of plasma membrane
271:
232:
227:
170:
129:
124:
7:
1259:recovery and to mitigate persistent
2608:The Journal of Biological Chemistry
1847:The Journal of Biological Chemistry
2643:American Journal of Human Genetics
2565:American Journal of Human Genetics
2530:European Journal of Human Genetics
1238:), which causes human dry and wet
1037:which in humans is encoded by the
936:
915:
889:
868:
844:
825:
799:
780:
754:
735:
489:
427:choroid plexus of fourth ventricle
407:
345:
324:
14:
3236:ATP-binding cassette transporters
2767:American Journal of Ophthalmology
2046:Canadian Journal of Ophthalmology
644:phospholipid transfer to membrane
547:phospholipid transporter activity
1336:ATP-binding cassette transporter
1322:support a role for the adjacent
1300:age-related macular degeneration
1240:age-related macular degeneration
1049:ATP-binding cassette transporter
654:phototransduction, visible light
493:
255:
248:
242:
219:
154:
147:
141:
116:
2268:Allikmets R (September 1997).
659:photoreceptor cell maintenance
593:integral component of membrane
504:More reference expression data
473:More reference expression data
1:
2780:10.1016/S0002-9394(99)00236-6
2321:10.1126/science.277.5333.1805
2208:10.1016/S0014-5793(97)00517-6
2058:10.1016/S0008-4182(05)80071-7
1757:10.1016/S0092-8674(00)80602-9
1686:Sullivan JM (November 2009).
1647:10.1016/j.febslet.2005.12.019
1594:10.1016/S0076-6879(00)15888-4
1555:10.1126/science.277.5333.1805
240:
139:
1140:is expressed much less than
431:Epithelium of choroid plexus
377:Epithelium of choroid plexus
3231:Genes on human chromosome 1
2895:membrane transport proteins
2823:human gene location in the
618:photoreceptor disc membrane
608:photoreceptor outer segment
3252:
2835:human gene details in the
2699:10.1001/archopht.117.4.504
1704:10.1016/j.exer.2009.03.005
669:phospholipid translocation
649:retinoid metabolic process
577:lipid transporter activity
435:retinal pigment epithelium
361:retinal pigment epithelium
3209:ABC transporter disorders
3204:
2686:Archives of Ophthalmology
1692:Experimental Eye Research
1576:Sun H, Nathans J (2000).
1415:"Mouse PubMed Reference:"
1397:"Human PubMed Reference:"
1047:ABCA4 is a member of the
1009:
1004:
1000:
993:
977:
971:Chr 3: 121.84 – 121.97 Mb
958:
943:
922:
911:
896:
875:
864:
851:
847:
832:
828:
819:
806:
802:
787:
783:
774:
761:
757:
742:
738:
729:
714:
707:
703:
687:
522:
518:
501:
492:
483:
470:
419:
410:
357:
348:
318:
310:
306:
289:
276:
239:
218:
209:
205:
188:
175:
138:
115:
106:
102:
57:
54:
44:
37:
32:
28:
23:
2813:Medical Subject Headings
2390:Human Molecular Genetics
1805:10.1007/0-387-32442-9_64
1158:phosphatidylethanolamine
2621:10.1074/jbc.274.12.8269
2543:10.1038/sj.ejhg.5200221
1898:Nature Reviews Genetics
1316:genome wide association
964:Chr 1: 93.99 – 94.12 Mb
679:transmembrane transport
2472:10.1006/geno.1997.5164
2161:10.1006/geno.1996.4528
1860:10.1074/jbc.M804505200
1209:
1098:adenosine triphosphate
423:neural layer of retina
2501:10.1007/s004390050765
2434:10.1007/s004390050649
1508:10.1007/s004390050649
1288:fundus flavimaculatus
1246:Clinical significance
1230:-retinyl-ethanolamine
1200:-retinyl-ethanolamine
1191:
623:endoplasmic reticulum
2809:ABCA4+protein,+human
1948:Genetic Epidemiology
1296:retinitis pigmentosa
1256:macular degeneration
1061:macular degeneration
639:response to stimulus
572:transporter activity
273:3 G1|3 52.94 cM
234:Chromosome 3 (mouse)
132:Chromosome 1 (human)
2837:UCSC Genome Browser
2825:UCSC Genome Browser
2730:1999Natur.400..761K
2403:10.1093/hmg/7.3.355
2287:10.1038/ng0997-122a
2200:1997FEBSL.409..247A
1639:2006FEBSL.580.1171V
389:right lobe of liver
2848:, which is in the
2123:10.1038/ng0397-236
2092:10.1101/gr.6.9.791
2009:10.1002/bdra.23076
1960:10.1002/gepi.20595
1464:10.1038/ng0397-236
1292:cone-rod dystrophy
1221:-pyridinium salt,
1210:
1085:rod photoreceptors
809:ENSMUSG00000028125
632:Biological process
586:Cellular component
542:nucleotide binding
530:Molecular function
439:epithelium of lens
381:tail of epididymis
373:right uterine tube
3216:
3215:
2887:Membrane proteins
2351:10.1038/ng0198-11
2245:10.1038/ng0997-15
1814:978-0-387-28464-4
1603:978-0-12-182216-3
1057:Stargardt disease
1020:
1019:
1016:
1015:
989:
988:
954:
953:
933:
932:
907:
906:
886:
885:
860:
859:
841:
840:
815:
814:
796:
795:
770:
769:
751:
750:
699:
698:
664:visual perception
514:
513:
510:
509:
479:
478:
466:
465:
404:
403:
385:corpus epididymis
302:
301:
201:
200:
96:ABCA4 - orthologs
3243:
2891:carrier proteins
2880:
2873:
2866:
2857:
2792:
2782:
2757:
2711:
2701:
2676:
2666:
2633:
2623:
2598:
2588:
2555:
2545:
2520:
2483:
2453:
2415:
2405:
2380:
2362:
2332:
2315:(5333): 1805–7.
2299:
2289:
2264:
2227:
2182:
2172:
2142:
2104:
2094:
2069:
2031:
2030:
2020:
1988:
1982:
1981:
1971:
1938:
1932:
1931:
1921:
1889:
1883:
1882:
1872:
1862:
1853:(39): 26684–93.
1838:
1827:
1826:
1794:
1784:
1778:
1777:
1759:
1735:
1726:
1725:
1715:
1683:
1677:
1676:
1658:
1622:
1616:
1615:
1573:
1567:
1566:
1549:(5333): 1805–7.
1534:
1528:
1527:
1490:
1484:
1483:
1446:
1440:
1439:
1432:
1423:
1422:
1411:
1405:
1404:
1393:
1387:
1377:
1366:
1356:
1177:) and inorganic
1025:, also known as
1002:
1001:
973:
966:
949:
937:
928:
916:
912:RefSeq (protein)
902:
890:
881:
869:
845:
826:
800:
781:
755:
736:
705:
520:
506:
497:
490:
475:
415:
413:Top expressed in
408:
353:
351:Top expressed in
346:
325:
308:
298:
285:
274:
259:
252:
246:
235:
223:
207:
197:
184:
173:
158:
151:
145:
134:
120:
104:
98:
49:
42:
21:
3251:
3250:
3246:
3245:
3244:
3242:
3241:
3240:
3221:
3220:
3217:
3212:
3200:
3160:
3139:
3123:
3092:
3017:
2956:
2905:
2898:ABC transporter
2884:
2800:
2795:
2760:
2724:(6746): 761–6.
2714:
2679:
2636:
2614:(12): 8269–81.
2601:
2558:
2523:
2486:
2456:
2418:
2383:
2339:Nature Genetics
2335:
2302:
2274:Nature Genetics
2267:
2233:Nature Genetics
2230:
2185:
2145:
2111:Nature Genetics
2107:
2079:Genome Research
2072:
2043:
2039:
2037:Further reading
2034:
1990:
1989:
1985:
1940:
1939:
1935:
1910:10.1038/nrg2933
1891:
1890:
1886:
1840:
1839:
1830:
1815:
1786:
1785:
1781:
1737:
1736:
1729:
1685:
1684:
1680:
1624:
1623:
1619:
1604:
1575:
1574:
1570:
1536:
1535:
1531:
1492:
1491:
1487:
1452:Nature Genetics
1448:
1447:
1443:
1434:
1433:
1426:
1413:
1412:
1408:
1395:
1394:
1390:
1378:
1369:
1357:
1348:
1344:
1332:
1314:combination of
1248:
1184:
1123:
1093:
1075:(ECD), and two
1011:View/Edit Mouse
1006:View/Edit Human
969:
962:
959:Location (UCSC)
945:
924:
898:
877:
790:ENSG00000198691
683:
674:lipid transport
627:
581:
567:ATPase activity
502:
471:
462:
457:
453:
449:
445:
441:
437:
433:
429:
425:
411:
400:
395:
393:skin of abdomen
391:
387:
383:
379:
375:
371:
367:
363:
349:
293:
280:
272:
262:
261:
260:
253:
233:
210:Gene location (
192:
179:
171:
161:
160:
159:
152:
130:
107:Gene location (
58:
45:
38:
17:
12:
11:
5:
3249:
3247:
3239:
3238:
3233:
3223:
3222:
3214:
3213:
3205:
3202:
3201:
3199:
3198:
3184:
3179:
3174:
3168:
3166:
3162:
3161:
3159:
3158:
3153:
3147:
3145:
3141:
3140:
3138:
3137:
3131:
3129:
3125:
3124:
3122:
3121:
3116:
3111:
3106:
3100:
3098:
3094:
3093:
3091:
3090:
3085:
3080:
3075:
3061:
3056:
3051:
3046:
3041:
3036:
3031:
3025:
3023:
3019:
3018:
3016:
3015:
3010:
3005:
3000:
2995:
2990:
2985:
2979:
2970:
2964:
2962:
2958:
2957:
2955:
2954:
2949:
2944:
2939:
2934:
2929:
2924:
2919:
2913:
2911:
2907:
2906:
2885:
2883:
2882:
2875:
2868:
2860:
2841:
2840:
2828:
2816:
2806:
2799:
2798:External links
2796:
2794:
2793:
2758:
2712:
2677:
2655:10.1086/302323
2649:(4): 1024–35.
2634:
2599:
2577:10.1086/302251
2556:
2521:
2495:(6): 699–705.
2489:Human Genetics
2484:
2454:
2422:Human Genetics
2416:
2381:
2333:
2300:
2265:
2228:
2183:
2143:
2105:
2085:(9): 791–806.
2070:
2040:
2038:
2035:
2033:
2032:
2003:(11): 934–42.
1983:
1933:
1884:
1828:
1813:
1779:
1727:
1678:
1617:
1602:
1568:
1529:
1496:Human Genetics
1485:
1441:
1424:
1406:
1388:
1367:
1345:
1343:
1340:
1339:
1338:
1331:
1328:
1275:-retinylidene-
1267:-retinylidene-
1247:
1244:
1226:-retinylidene-
1196:-retinylidene-
1182:
1122:
1119:
1092:
1089:
1071:extracellular
1018:
1017:
1014:
1013:
1008:
998:
997:
991:
990:
987:
986:
984:
982:
975:
974:
967:
960:
956:
955:
952:
951:
941:
940:
934:
931:
930:
920:
919:
913:
909:
908:
905:
904:
894:
893:
887:
884:
883:
873:
872:
866:
862:
861:
858:
857:
849:
848:
842:
839:
838:
830:
829:
823:
817:
816:
813:
812:
804:
803:
797:
794:
793:
785:
784:
778:
772:
771:
768:
767:
759:
758:
752:
749:
748:
740:
739:
733:
727:
726:
721:
716:
712:
711:
701:
700:
697:
696:
685:
684:
682:
681:
676:
671:
666:
661:
656:
651:
646:
641:
635:
633:
629:
628:
626:
625:
620:
615:
610:
605:
600:
595:
589:
587:
583:
582:
580:
579:
574:
569:
564:
559:
554:
549:
544:
539:
533:
531:
527:
526:
516:
515:
512:
511:
508:
507:
499:
498:
487:
481:
480:
477:
476:
468:
467:
464:
463:
461:
460:
459:tail of embryo
456:
455:corneal stroma
452:
448:
444:
440:
436:
432:
428:
424:
420:
417:
416:
405:
402:
401:
399:
398:
397:jejunal mucosa
394:
390:
386:
382:
378:
374:
370:
366:
362:
358:
355:
354:
342:
341:
333:
322:
316:
315:
312:RNA expression
304:
303:
300:
299:
291:
287:
286:
278:
275:
270:
264:
263:
254:
247:
241:
237:
236:
231:
225:
224:
216:
215:
203:
202:
199:
198:
190:
186:
185:
177:
174:
169:
163:
162:
153:
146:
140:
136:
135:
128:
122:
121:
113:
112:
100:
99:
56:
52:
51:
43:
35:
34:
30:
29:
26:
25:
15:
13:
10:
9:
6:
4:
3:
2:
3248:
3237:
3234:
3232:
3229:
3228:
3226:
3219:
3211:
3210:
3203:
3196:
3192:
3188:
3185:
3183:
3180:
3178:
3175:
3173:
3170:
3169:
3167:
3163:
3157:
3154:
3152:
3149:
3148:
3146:
3142:
3136:
3133:
3132:
3130:
3126:
3120:
3117:
3115:
3112:
3110:
3107:
3105:
3102:
3101:
3099:
3095:
3089:
3086:
3084:
3081:
3079:
3076:
3073:
3069:
3065:
3062:
3060:
3057:
3055:
3052:
3050:
3047:
3045:
3042:
3040:
3037:
3035:
3032:
3030:
3027:
3026:
3024:
3020:
3014:
3011:
3009:
3006:
3004:
3001:
2999:
2996:
2994:
2991:
2989:
2986:
2983:
2980:
2978:
2974:
2971:
2969:
2966:
2965:
2963:
2959:
2953:
2950:
2948:
2945:
2943:
2940:
2938:
2935:
2933:
2930:
2928:
2925:
2923:
2920:
2918:
2915:
2914:
2912:
2908:
2903:
2899:
2896:
2892:
2888:
2881:
2876:
2874:
2869:
2867:
2862:
2861:
2858:
2854:
2853:
2851:
2850:public domain
2847:
2838:
2834:
2833:
2829:
2826:
2822:
2821:
2817:
2814:
2810:
2807:
2805:
2802:
2801:
2797:
2790:
2786:
2781:
2776:
2772:
2768:
2764:
2759:
2755:
2751:
2747:
2743:
2739:
2738:10.1038/23468
2735:
2731:
2727:
2723:
2719:
2713:
2709:
2705:
2700:
2695:
2692:(4): 504–10.
2691:
2687:
2683:
2678:
2674:
2670:
2665:
2660:
2656:
2652:
2648:
2644:
2640:
2635:
2631:
2627:
2622:
2617:
2613:
2609:
2605:
2600:
2596:
2592:
2587:
2582:
2578:
2574:
2571:(2): 422–34.
2570:
2566:
2562:
2557:
2553:
2549:
2544:
2539:
2535:
2531:
2527:
2522:
2518:
2514:
2510:
2506:
2502:
2498:
2494:
2490:
2485:
2481:
2477:
2473:
2469:
2466:(1): 139–42.
2465:
2461:
2455:
2451:
2447:
2443:
2439:
2435:
2431:
2427:
2423:
2417:
2413:
2409:
2404:
2399:
2396:(3): 355–62.
2395:
2391:
2387:
2382:
2378:
2374:
2370:
2366:
2361:
2356:
2352:
2348:
2344:
2340:
2334:
2330:
2326:
2322:
2318:
2314:
2310:
2306:
2301:
2297:
2293:
2288:
2283:
2279:
2275:
2271:
2266:
2262:
2258:
2254:
2250:
2246:
2242:
2238:
2234:
2229:
2225:
2221:
2217:
2213:
2209:
2205:
2201:
2197:
2194:(2): 247–52.
2193:
2189:
2184:
2180:
2176:
2171:
2166:
2162:
2158:
2154:
2150:
2144:
2140:
2136:
2132:
2128:
2124:
2120:
2117:(3): 236–46.
2116:
2112:
2106:
2102:
2098:
2093:
2088:
2084:
2080:
2076:
2071:
2067:
2063:
2059:
2055:
2052:(3): 288–92.
2051:
2047:
2042:
2041:
2036:
2028:
2024:
2019:
2014:
2010:
2006:
2002:
1998:
1994:
1987:
1984:
1979:
1975:
1970:
1965:
1961:
1957:
1954:(6): 469–78.
1953:
1949:
1945:
1937:
1934:
1929:
1925:
1920:
1915:
1911:
1907:
1904:(3): 167–78.
1903:
1899:
1895:
1888:
1885:
1880:
1876:
1871:
1866:
1861:
1856:
1852:
1848:
1844:
1837:
1835:
1833:
1829:
1824:
1820:
1816:
1810:
1806:
1802:
1798:
1793:
1792:
1783:
1780:
1775:
1771:
1767:
1763:
1758:
1753:
1749:
1745:
1741:
1734:
1732:
1728:
1723:
1719:
1714:
1709:
1705:
1701:
1697:
1693:
1689:
1682:
1679:
1674:
1670:
1666:
1662:
1657:
1652:
1648:
1644:
1640:
1636:
1633:(4): 1171–7.
1632:
1628:
1621:
1618:
1613:
1609:
1605:
1599:
1595:
1591:
1587:
1583:
1579:
1572:
1569:
1564:
1560:
1556:
1552:
1548:
1544:
1540:
1533:
1530:
1525:
1521:
1517:
1513:
1509:
1505:
1501:
1497:
1489:
1486:
1481:
1477:
1473:
1469:
1465:
1461:
1458:(3): 236–46.
1457:
1453:
1445:
1442:
1437:
1431:
1429:
1425:
1420:
1416:
1410:
1407:
1402:
1398:
1392:
1389:
1385:
1381:
1376:
1374:
1372:
1368:
1364:
1360:
1355:
1353:
1351:
1347:
1341:
1337:
1334:
1333:
1329:
1327:
1325:
1321:
1317:
1312:
1308:
1303:
1301:
1297:
1293:
1289:
1285:
1280:
1278:
1274:
1270:
1266:
1262:
1257:
1253:
1250:Mutations in
1245:
1243:
1241:
1237:
1236:
1231:
1229:
1225:
1220:
1215:
1207:
1206:
1201:
1199:
1195:
1190:
1186:
1180:
1176:
1171:
1167:
1163:
1162:retinaldehyde
1159:
1155:
1151:
1150:knockout mice
1147:
1143:
1139:
1135:
1131:
1127:
1120:
1118:
1115:
1111:
1108:forms into a
1107:
1103:
1099:
1090:
1088:
1086:
1082:
1078:
1074:
1070:
1066:
1062:
1058:
1054:
1050:
1045:
1043:
1040:
1036:
1032:
1028:
1024:
1012:
1007:
1003:
999:
996:
992:
985:
983:
980:
976:
972:
968:
965:
961:
957:
950:
948:
942:
938:
935:
929:
927:
921:
917:
914:
910:
903:
901:
895:
891:
888:
882:
880:
874:
870:
867:
865:RefSeq (mRNA)
863:
856:
855:
850:
846:
843:
837:
836:
831:
827:
824:
822:
818:
811:
810:
805:
801:
798:
792:
791:
786:
782:
779:
777:
773:
766:
765:
760:
756:
753:
747:
746:
741:
737:
734:
732:
728:
725:
722:
720:
717:
713:
710:
706:
702:
695:
691:
686:
680:
677:
675:
672:
670:
667:
665:
662:
660:
657:
655:
652:
650:
647:
645:
642:
640:
637:
636:
634:
631:
630:
624:
621:
619:
616:
614:
611:
609:
606:
604:
601:
599:
596:
594:
591:
590:
588:
585:
584:
578:
575:
573:
570:
568:
565:
563:
560:
558:
555:
553:
550:
548:
545:
543:
540:
538:
535:
534:
532:
529:
528:
525:
524:Gene ontology
521:
517:
505:
500:
496:
491:
488:
486:
482:
474:
469:
458:
454:
450:
446:
442:
438:
434:
430:
426:
422:
421:
418:
414:
409:
406:
396:
392:
388:
384:
380:
376:
372:
368:
364:
360:
359:
356:
352:
347:
344:
343:
340:
338:
334:
332:
331:
327:
326:
323:
321:
317:
313:
309:
305:
297:
292:
288:
284:
279:
269:
265:
258:
251:
245:
238:
230:
226:
222:
217:
213:
208:
204:
196:
191:
187:
183:
178:
168:
164:
157:
150:
144:
137:
133:
127:
123:
119:
114:
110:
105:
101:
97:
93:
89:
85:
81:
77:
73:
69:
65:
61:
53:
48:
41:
36:
31:
27:
22:
19:
3218:
3206:
2931:
2843:
2842:
2831:
2819:
2773:(6): 720–4.
2770:
2766:
2721:
2717:
2689:
2685:
2646:
2642:
2611:
2607:
2568:
2564:
2536:(3): 291–5.
2533:
2529:
2492:
2488:
2463:
2459:
2425:
2421:
2393:
2389:
2342:
2338:
2312:
2308:
2277:
2273:
2236:
2232:
2191:
2188:FEBS Letters
2187:
2155:(1): 142–6.
2152:
2148:
2114:
2110:
2082:
2078:
2049:
2045:
2000:
1996:
1986:
1951:
1947:
1936:
1901:
1897:
1887:
1850:
1846:
1790:
1782:
1750:(1): 13–23.
1747:
1743:
1698:(5): 602–3.
1695:
1691:
1681:
1630:
1627:FEBS Letters
1626:
1620:
1581:
1571:
1546:
1542:
1532:
1499:
1495:
1488:
1455:
1451:
1444:
1418:
1409:
1400:
1391:
1311:cleft palate
1304:
1283:
1281:
1276:
1272:
1268:
1264:
1251:
1249:
1234:
1233:
1227:
1223:
1222:
1218:
1213:
1211:
1204:
1203:
1197:
1193:
1192:
1165:
1145:
1137:
1125:
1124:
1105:
1094:
1069:glycosylated
1064:
1046:
1038:
1030:
1026:
1022:
1021:
944:
923:
897:
876:
852:
833:
807:
788:
762:
743:
723:
718:
451:pineal gland
443:ciliary body
335:
328:
294:121,973,772
281:121,838,092
55:External IDs
18:
2428:(1): 21–6.
2360:10261/39477
2345:(1): 11–2.
2239:(1): 15–6.
2170:10261/39369
1502:(1): 21–6.
1110:heterodimer
557:ATP binding
193:94,121,148
180:93,992,834
33:Identifiers
3225:Categories
2280:(1): 122.
1656:1874/19996
1386:, May 2017
1365:, May 2017
1342:References
1077:nucleotide
1053:eukaryotes
339:(ortholog)
76:HomoloGene
3207:see also
1307:cleft lip
1179:phosphate
1142:rhodopsin
1102:phosphate
1091:Structure
947:NP_031404
926:NP_000341
900:NM_007378
879:NM_000350
709:Orthologs
84:GeneCards
3187:Sterolin
2789:10612508
2746:10466724
2708:10206579
2673:10090887
2630:10075733
2517:34452470
2460:Genomics
2450:22070963
2149:Genomics
2139:31677978
2066:15947798
2027:23008150
1978:21618603
1928:21331089
1879:18658157
1823:17249610
1774:18605680
1766:10412977
1722:19306869
1673:27946190
1665:16376334
1612:10736747
1524:22070963
1480:31677978
1382:–
1361:–
1330:See also
1324:ARHGAP29
1286:include
1214:ABCR -/-
1154:flippase
1121:Function
1114:retinoid
995:Wikidata
688:Sources:
603:membrane
369:testicle
2754:4394997
2726:Bibcode
2664:1377826
2595:9973280
2586:1377752
2552:9781034
2509:9703434
2480:9503029
2442:9490294
2412:9466990
2369:9425888
2329:9295268
2309:Science
2296:9288113
2253:9288089
2224:7701478
2216:9202155
2196:Bibcode
2179:9070931
2131:9054934
2101:8889548
2018:3501616
1969:3180858
1919:3086810
1870:2546559
1713:3371273
1635:Bibcode
1563:9295268
1543:Science
1516:9490294
1472:9054934
1384:Ensembl
1363:Ensembl
1309:and/or
1073:domains
1035:protein
1033:, is a
821:UniProt
776:Ensembl
715:Species
694:QuickGO
314:pattern
40:Aliases
2902:TC 3A1
2815:(MeSH)
2787:
2752:
2744:
2718:Nature
2706:
2671:
2661:
2628:
2593:
2583:
2550:
2515:
2507:
2478:
2448:
2440:
2410:
2377:125620
2375:
2367:
2327:
2294:
2261:759924
2259:
2251:
2222:
2214:
2177:
2137:
2129:
2099:
2064:
2025:
2015:
1976:
1966:
1926:
1916:
1877:
1867:
1821:
1811:
1797:465–70
1772:
1764:
1720:
1710:
1671:
1663:
1610:
1600:
1586:879–97
1561:
1522:
1514:
1478:
1470:
1298:, and
1081:retina
1063:. The
981:search
979:PubMed
854:O35600
835:P78363
731:Entrez
485:BioGPS
172:1p22.1
72:109424
64:601691
2832:ABCA4
2820:ABCA4
2750:S2CID
2513:S2CID
2446:S2CID
2373:S2CID
2257:S2CID
2220:S2CID
2135:S2CID
1770:S2CID
1669:S2CID
1520:S2CID
1476:S2CID
1284:ABCA4
1261:opsin
1252:ABCA4
1146:ABCA4
1134:cones
1106:ABCA4
1065:ABCA4
1039:ABCA4
1027:ABCA4
764:11304
724:Mouse
719:Human
690:Amigo
365:gonad
337:Mouse
330:Human
277:Start
212:Mouse
176:Start
109:Human
88:ABCA4
47:ABCA4
24:ABCA4
3064:C8-9
2973:B2-3
2785:PMID
2742:PMID
2704:PMID
2669:PMID
2626:PMID
2591:PMID
2548:PMID
2505:PMID
2476:PMID
2438:PMID
2408:PMID
2365:PMID
2325:PMID
2292:PMID
2249:PMID
2212:PMID
2175:PMID
2127:PMID
2097:PMID
2062:PMID
2023:PMID
1974:PMID
1924:PMID
1875:PMID
1819:PMID
1809:ISBN
1762:PMID
1744:Cell
1718:PMID
1661:PMID
1608:PMID
1598:ISBN
1559:PMID
1512:PMID
1468:PMID
1320:IRF6
1212:The
1138:ABCR
1132:and
1130:rods
1126:ABCR
1042:gene
1031:ABCR
447:iris
320:Bgee
268:Band
229:Chr.
167:Band
126:Chr.
60:OMIM
3088:C13
3083:C11
3078:C10
3013:B11
2952:A13
2947:A12
2775:doi
2771:128
2734:doi
2722:400
2694:doi
2690:117
2659:PMC
2651:doi
2616:doi
2612:274
2581:PMC
2573:doi
2538:doi
2497:doi
2493:102
2468:doi
2430:doi
2426:102
2398:doi
2355:hdl
2347:doi
2317:doi
2313:277
2282:doi
2241:doi
2204:doi
2192:409
2165:hdl
2157:doi
2119:doi
2087:doi
2054:doi
2013:PMC
2005:doi
1964:PMC
1956:doi
1914:PMC
1906:doi
1865:PMC
1855:doi
1851:283
1801:doi
1752:doi
1708:PMC
1700:doi
1651:hdl
1643:doi
1631:580
1590:doi
1551:doi
1547:277
1504:doi
1500:102
1460:doi
1235:A2E
1219:bis
1205:A2E
1175:ADP
1170:ATP
1166:cis
1029:or
290:End
189:End
92:OMA
80:298
68:MGI
3227::
3195:G8
3193:,
3191:G5
3182:G4
3177:G2
3172:G1
3156:F2
3151:F1
3135:E1
3119:D4
3114:D3
3109:D2
3104:D1
3072:C9
3070:,
3068:C8
3059:C7
3054:C6
3049:C5
3044:C4
3039:C3
3034:C2
3029:C1
3008:B9
3003:B7
2998:B6
2993:B5
2988:B4
2982:B3
2977:B2
2968:B1
2942:A8
2937:A7
2932:A4
2927:A3
2922:A2
2917:A1
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1198:N
1194:N
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