1347:
302:, the main mechanisms by which horizontal cells inhibit cones probably do not involve the release of GABA by horizontal cells onto cones. Two mechanisms that are not mutually exclusive likely contribute to horizontal cell inhibition of glutamate release by cones. Both postulated mechanisms depend on the protected environment provided by the invaginating synapses that horizontal cells make onto cones. The first postulated mechanism is a very fast ephaptic mechanism that has no synaptic delay, making it one of the fastest inhibitory synapses known. The second postulated mechanism is relatively slow with a time constant of about 200 ms and depends on ATP release via Pannexin 1 channels located on horizontal cell dendrites invaginating the cone synaptic terminal. The ecto-ATPase NTPDase1 hydrolyses extracellular ATP to AMP, phosphate groups, and protons. The phosphate groups and protons form a pH buffer with a pKa of 7.2, which keeps the pH in the synaptic cleft relatively acidic. This inhibits the cone Ca channels and consequently reduces the glutamate release by the cones.
1346:
230:, measures the average level of illumination falling upon a region of the retinal surface, which horizontal cells then subtract a proportionate value from the output of photoreceptors to hold the signal input to the inner retinal circuitry within its operating range. Horizontal cells are also one of two groups of inhibitory interneurons that contribute to the surround of retinal ganglion cells:
26:
309:
of bipolar cells is thought to be inherited from cones. However, when recordings are made from parts of the cone that are distant from the cone terminals that synapse onto bipolar cells, center-surround antagonism seems to be less reliable in cones than in bipolar cells. As the invaginating synapses
1382:
1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and
221:
nearby photoreceptors. Conversely, in the light, a photoreceptor releases less glutamate, which hyperpolarizes the horizontal cell, leading to depolarization of nearby photoreceptors. Thus, horizontal cells provide
1383:
veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina
185:
Horizontal cells and other retinal interneuron cells are less likely to be near neighbours of the same subtype than would occur by chance, resulting in ‘exclusion zones’ that separate them.
171:
Horizontal cells span across photoreceptors and summate inputs before synapsing onto photoreceptor cells. Horizontal cells may also synapse onto bipolar cells, but this remains uncertain.
189:
arrangements provide a mechanism to distribute each cell type evenly across the retina, ensuring that all parts of the visual field have access to a full set of processing elements.
137:
conditions. Horizontal cells provide inhibitory feedback to rod and cone photoreceptors. They are thought to be important for the antagonistic center-surround property of the
182:, it is observed that A-type horizontal cells have a density of 225 cells/mm near the center of the retina and a density of 120 cells/mm in more peripheral retina.
292:
272:
252:
1366:
1363:
1626:
339:
1364:
1369:
310:
from horizontal cells are made onto cone terminals, the center-surround antagonism of cones is thought to be more reliably present in cone terminals.
1056:"Effects of pH buffering on horizontal and ganglion cell light responses in primate retina: evidence for the proton hypothesis of surround formation"
1378:
849:
Kamermans M, Fahrenfort I, Schultz K, Janssen-Bienhold U, Sjoerdsma T, Weiler R (May 2001). "Hemichannel-mediated inhibition in the outer retina".
714:
Verweij J, Kamermans M, Spekreijse H (December 1996). "Horizontal cells feed back to cones by shifting the cone calcium-current activation range".
1365:
1370:
1199:
1350:
298:
The exact mechanism by which depolarization of horizontal cells hyperpolarizes photoreceptors is uncertain. Although horizontal cells contain
1356:
1103:
Byzov AL, Shura-Bura TM (1986). "Electrical feedback mechanism in the processing of signals in the outer plexiform layer of the retina".
956:"pH changes in the invaginating synaptic cleft mediate feedback from horizontal cells to cone photoreceptors by modulating Ca2+ channels"
478:
Chaya, Taro; Matsumoto, Akihiro; Sugita, Yuko; Watanabe, Satoshi; Kuwahara, Ryusuke; Tachibana, Masao; Furukawa, Takahisa (2017-07-17).
194:
1361:
1354:
1373:
1372:
1226:
1353:
168:
Depending on the species, there are typically one or two classes of horizontal cells, with a third type sometimes proposed.
1352:
1872:
1360:
1371:
1362:
1359:
93:
1368:
1367:
1351:
1374:
905:"Extracellular ATP hydrolysis inhibits synaptic transmission by increasing ph buffering in the synaptic cleft"
217:
from photoreceptors, which happens in the absence of light. Depolarization of a horizontal cell causes it to
1375:
1817:
1785:
1622:
1770:
1739:
1731:
1411:
1003:
Vessey JP, Stratis AK, Daniels BA, Da Silva N, Jonz MG, Lalonde MR, Baldridge WH, Barnes S (April 2005).
218:
1923:
1566:
1556:
1377:
1005:"Proton-mediated feedback inhibition of presynaptic calcium channels at the cone photoreceptor synapse"
324:
306:
1358:
1219:
129:. Among their functions, horizontal cells are believed to be responsible for increasing contrast via
1376:
1379:
1248:
197:
transmembrane proteins have critical roles in the formation of the mosaics by horizontal cells and
68:
1357:
1561:
1517:
659:"MEGF10 and MEGF11 mediate homotypic interactions required for mosaic spacing of retinal neurons"
299:
198:
903:
Vroman R, Klaassen LJ, Howlett MH, Cenedese V, Klooster J, Sjoerdsma T, Kamermans M (May 2014).
1701:
1325:
1355:
1877:
1682:
1606:
1544:
1532:
1487:
334:
153:
142:
1928:
1867:
1212:
858:
802:"Center-surround antagonism mediated by proton signaling at the cone photoreceptor synapse"
670:
615:
491:
277:
257:
237:
226:
to photoreceptors. The moderately wide lateral spread and coupling of horizontal cells by
8:
1892:
1598:
1549:
1537:
1527:
1482:
1416:
1330:
1277:
446:
114:
1204:
862:
674:
619:
606:
Wässle H, Riemann HJ (March 1978). "The mosaic of nerve cells in the mammalian retina".
495:
1862:
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658:
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522:
479:
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430:
399:
374:
319:
149:
130:
126:
727:
1902:
1618:
1522:
1439:
1310:
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1120:
1116:
1085:
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527:
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404:
344:
223:
1132:
886:
643:
1933:
1887:
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1315:
1267:
1171:
1163:
1112:
1075:
1071:
1067:
1026:
1021:
1016:
975:
967:
926:
916:
866:
821:
813:
772:
762:
723:
686:
678:
623:
575:
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517:
499:
450:
442:
394:
386:
1800:
1759:
1755:
1677:
1167:
921:
390:
138:
73:
1805:
1795:
1780:
1775:
1472:
1464:
1444:
1320:
1295:
504:
329:
210:
186:
175:
157:
86:
1917:
1830:
1765:
1602:
513:
214:
46:
870:
1790:
1614:
1449:
1431:
1185:
1089:
1040:
989:
940:
878:
835:
786:
700:
627:
589:
531:
464:
408:
227:
41:
1124:
971:
848:
817:
735:
1882:
1751:
1708:
1646:
1610:
682:
635:
99:
1696:
1454:
480:"Versatile functional roles of horizontal cells in the retinal circuit"
122:
1672:
1667:
1586:
1240:
178:
of horizontal cells towards the central region of the retina. In the
113:
are the laterally interconnecting neurons having cell bodies in the
1590:
80:
1842:
1743:
1687:
1403:
1837:
1747:
1501:
1287:
1259:
190:
125:
eyes. They help integrate and regulate the input from multiple
118:
56:
1477:
902:
134:
1002:
477:
1391:
713:
25:
1234:
179:
748:
1053:
549:
547:
545:
543:
541:
749:
Verweij J, Hornstein EP, Schnapf JL (November 2003).
280:
260:
240:
1054:
Davenport CM, Detwiler PB, Dacey DM (January 2008).
898:
896:
751:"Surround antagonism in macaque cone photoreceptors"
538:
340:
List of distinct cell types in the adult human body
424:
422:
420:
418:
286:
266:
246:
893:
1915:
1349:
1102:
415:
953:
553:
368:
366:
364:
362:
360:
1220:
605:
1047:
996:
947:
842:
1096:
742:
656:
357:
1227:
1213:
707:
556:"Lateral interactions in the outer retina"
428:
24:
1175:
1152:"The neuronal organization of the retina"
1079:
1030:
1020:
979:
930:
920:
825:
776:
766:
690:
579:
554:Thoreson WB, Mangel SC (September 2012).
521:
503:
454:
398:
375:"The neuronal organization of the retina"
799:
1149:
372:
254:Center photoreceptor hyperpolarization
1916:
954:Hirasawa H, Kaneko A (December 2003).
601:
599:
1208:
294:Surround photoreceptor depolarization
657:Kay, JN; Chu, MW; Sanes, JR (2012).
447:10.1146/annurev-vision-082114-035334
431:"Functional Circuitry of the Retina"
429:Demb JB, Singer JH (November 2015).
133:and adapting both to bright and dim
596:
13:
1345:
768:10.1523/JNEUROSCI.23-32-10249.2003
274:Horizontal cell hyperpolarization
14:
1945:
1193:
572:10.1016/j.preteyeres.2012.04.003
94:Anatomical terms of neuroanatomy
1143:
608:Proc. R. Soc. Lond. B Biol. Sci
1072:10.1523/JNEUROSCI.2735-07.2008
1022:10.1523/JNEUROSCI.5253-04.2005
793:
650:
471:
281:
261:
241:
160:, and retinal ganglion cells.
148:Other retinal neurons include
1:
728:10.1016/S0042-6989(96)00142-3
350:
1873:Optical coherence tomography
1627:Photosensitive ganglion cell
1168:10.1016/j.neuron.2012.10.002
1117:10.1016/0042-6989(86)90069-6
922:10.1371/journal.pbio.1001864
391:10.1016/j.neuron.2012.10.002
163:
7:
1623:Giant retina ganglion cells
1412:Capillary lamina of choroid
313:
204:
10:
1950:
1567:Retinal pigment epithelium
1557:External limiting membrane
1200:Webvision: Horizontal Cell
505:10.1038/s41598-017-05543-2
307:center-surround antagonism
1855:
1816:
1730:
1721:
1655:
1575:
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1500:
1463:
1430:
1402:
1390:
1343:
1286:
1258:
1247:
92:
79:
67:
62:
52:
40:
35:
23:
18:
199:starburst amacrine cells
30:Plan of retinal neurons.
1562:Layer of rods and cones
1518:Inner limiting membrane
871:10.1126/science.1060101
1384:
628:10.1098/rspb.1978.0026
288:
268:
248:
143:retinal ganglion cells
1878:Eye care professional
1683:Foveal avascular zone
1545:Outer plexiform layer
1533:Inner plexiform layer
1488:Iris sphincter muscle
1381:
972:10.1085/jgp.200308863
818:10.1085/jgp.200308947
800:Barnes S (Dec 2003).
289:
269:
249:
209:Horizontal cells are
1898:Physiological Optics
1868:Ocular immune system
1607:Retina ganglion cell
1150:Masland, RH (2012).
373:Masland, RH (2012).
287:{\displaystyle \to }
278:
267:{\displaystyle \to }
258:
247:{\displaystyle \to }
238:
1722:Anatomical regions
1583:Photoreceptor cells
1550:Outer nuclear layer
1538:Inner nuclear layer
1528:Ganglion cell layer
1483:Iris dilator muscle
1278:Trabecular meshwork
863:2001Sci...292.1178K
683:10.1038/nature10877
675:2012Natur.483..465K
620:1978RSPSB.200..441W
496:2017NatSR...7.5540C
320:Photoreceptor cells
174:There is a greater
150:photoreceptor cells
127:photoreceptor cells
115:inner nuclear layer
1385:
560:Prog Retin Eye Res
484:Scientific Reports
284:
264:
244:
213:by the release of
131:lateral inhibition
1924:Human eye anatomy
1911:
1910:
1903:Visual perception
1851:
1850:
1818:Posterior segment
1786:Posterior chamber
1717:
1716:
1619:Bistratified cell
1523:Nerve fiber layer
1496:
1495:
1440:Ciliary processes
1341:
1340:
857:(5519): 1178–80.
345:Gunnar Svaetichin
224:negative feedback
141:of many types of
108:
107:
103:
1941:
1888:Refractive error
1826:Vitreous chamber
1771:Anterior chamber
1732:Anterior segment
1728:
1727:
1508:
1507:
1417:Bruch's membrane
1400:
1399:
1392:Uvea / vascular
1348:
1268:Episcleral layer
1256:
1255:
1229:
1222:
1215:
1206:
1205:
1189:
1179:
1137:
1136:
1100:
1094:
1093:
1083:
1051:
1045:
1044:
1034:
1024:
1000:
994:
993:
983:
951:
945:
944:
934:
924:
900:
891:
890:
846:
840:
839:
829:
797:
791:
790:
780:
770:
761:(32): 10249–57.
746:
740:
739:
711:
705:
704:
694:
654:
648:
647:
614:(1141): 441–61.
603:
594:
593:
583:
551:
536:
535:
525:
507:
475:
469:
468:
458:
435:Annu Rev Vis Sci
426:
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402:
370:
293:
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273:
271:
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139:receptive fields
111:Horizontal cells
100:edit on Wikidata
97:
28:
16:
15:
1949:
1948:
1944:
1943:
1942:
1940:
1939:
1938:
1914:
1913:
1912:
1907:
1847:
1812:
1801:Capsule of lens
1756:Lacrimal system
1723:
1713:
1673:Parafoveal area
1668:Perifoveal area
1651:
1595:Horizontal cell
1571:
1492:
1459:
1426:
1422:Sattler's layer
1393:
1386:
1380:
1337:
1282:
1273:Schlemm's canal
1251:
1243:
1235:Anatomy of the
1233:
1196:
1146:
1141:
1140:
1101:
1097:
1052:
1048:
1015:(16): 4108–17.
1001:
997:
960:J. Gen. Physiol
952:
948:
915:(5): e1001864.
901:
894:
847:
843:
798:
794:
747:
743:
722:(24): 3943–53.
712:
708:
669:(7390): 465–9.
655:
651:
604:
597:
552:
539:
476:
472:
427:
416:
371:
358:
353:
316:
279:
276:
275:
259:
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255:
239:
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207:
166:
104:
31:
19:Horizontal cell
12:
11:
5:
1947:
1937:
1936:
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1909:
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1895:
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1845:
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1835:
1834:
1833:
1822:
1820:
1814:
1813:
1811:
1810:
1809:
1808:
1806:Zonule of Zinn
1803:
1793:
1788:
1783:
1778:
1776:Aqueous humour
1773:
1768:
1763:
1736:
1734:
1725:
1719:
1718:
1715:
1714:
1712:
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1706:
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1514:
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1490:
1485:
1480:
1475:
1469:
1467:
1461:
1460:
1458:
1457:
1452:
1447:
1445:Ciliary muscle
1442:
1436:
1434:
1428:
1427:
1425:
1424:
1419:
1414:
1408:
1406:
1397:
1388:
1387:
1344:
1342:
1339:
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1336:
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1333:
1328:
1323:
1318:
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1224:
1217:
1209:
1203:
1202:
1195:
1194:External links
1192:
1191:
1190:
1162:(2): 266–280.
1145:
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1095:
1046:
995:
946:
892:
841:
792:
741:
706:
649:
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537:
470:
414:
385:(2): 266–280.
355:
354:
352:
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342:
337:
335:Ganglion cells
332:
330:Amacrine cells
327:
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158:amacrine cells
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1893:Accommodation
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1831:Vitreous body
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1766:Fibrous tunic
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1738:
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1608:
1604:
1603:Amacrine cell
1600:
1596:
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1580:
1578:
1574:
1568:
1565:
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1560:
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1519:
1516:
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1509:
1506:
1503:
1499:
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1389:
1332:
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1291:
1289:
1285:
1279:
1276:
1274:
1271:
1269:
1266:
1265:
1263:
1261:
1257:
1254:
1250:
1249:Fibrous tunic
1246:
1242:
1238:
1230:
1225:
1223:
1218:
1216:
1211:
1210:
1207:
1201:
1198:
1197:
1187:
1183:
1178:
1173:
1169:
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1161:
1157:
1153:
1148:
1147:
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1122:
1118:
1114:
1110:
1106:
1099:
1091:
1087:
1082:
1077:
1073:
1069:
1066:(2): 456–64.
1065:
1061:
1057:
1050:
1042:
1038:
1033:
1028:
1023:
1018:
1014:
1010:
1006:
999:
991:
987:
982:
977:
973:
969:
966:(6): 657–71.
965:
961:
957:
950:
942:
938:
933:
928:
923:
918:
914:
910:
906:
899:
897:
888:
884:
880:
876:
872:
868:
864:
860:
856:
852:
845:
837:
833:
828:
823:
819:
815:
811:
807:
806:J Gen Physiol
803:
796:
788:
784:
779:
774:
769:
764:
760:
756:
752:
745:
737:
733:
729:
725:
721:
717:
710:
702:
698:
693:
688:
684:
680:
676:
672:
668:
664:
660:
653:
645:
641:
637:
633:
629:
625:
621:
617:
613:
609:
602:
600:
591:
587:
582:
577:
573:
569:
566:(5): 407–41.
565:
561:
557:
550:
548:
546:
544:
542:
533:
529:
524:
519:
515:
511:
506:
501:
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489:
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457:
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419:
410:
406:
401:
396:
392:
388:
384:
380:
376:
369:
367:
365:
363:
361:
356:
346:
343:
341:
338:
336:
333:
331:
328:
326:
325:Bipolar cells
323:
321:
318:
317:
311:
308:
303:
301:
296:
295:
234:Illumination
231:
229:
228:gap junctions
225:
220:
219:hyperpolarize
216:
212:
202:
200:
196:
192:
188:
183:
181:
177:
172:
169:
161:
159:
155:
154:bipolar cells
151:
146:
144:
140:
136:
132:
128:
124:
120:
116:
112:
101:
95:
91:
88:
85:
82:
78:
75:
72:
70:
66:
61:
58:
55:
51:
48:
47:Visual system
45:
43:
39:
34:
27:
22:
17:
1897:
1791:Ciliary body
1631:Diencephalon
1630:
1615:Parasol cell
1599:Bipolar cell
1594:
1450:Pars plicata
1432:Ciliary body
1300:
1159:
1155:
1144:Bibliography
1111:(1): 33–44.
1108:
1104:
1098:
1063:
1059:
1049:
1012:
1008:
998:
963:
959:
949:
912:
908:
854:
850:
844:
812:(6): 653–6.
809:
805:
795:
758:
754:
744:
719:
715:
709:
666:
662:
652:
611:
607:
563:
559:
487:
483:
473:
438:
434:
382:
378:
304:
297:
233:
232:
208:
184:
173:
170:
167:
147:
110:
109:
1929:Human cells
1883:Eye disease
1863:Keratocytes
1752:Conjunctiva
1709:Ora serrata
1647:Muller glia
1611:Midget cell
1331:Endothelium
1321:Dua's layer
1060:J. Neurosci
1009:J. Neurosci
755:J. Neurosci
490:(1): 5540.
441:: 263–289.
211:depolarized
63:Identifiers
1918:Categories
1724:of the eye
1697:Optic disc
1455:Pars plana
1326:Descemet's
1306:Epithelium
1105:Vision Res
716:Vision Res
351:References
123:vertebrate
1702:Optic cup
1587:Cone cell
1241:human eye
909:PLOS Biol
514:2045-2322
282:→
262:→
242:→
215:glutamate
201:in mice.
164:Structure
87:nifext_40
1591:Rod cell
1396:(middle)
1311:Bowman's
1186:23083731
1133:21785150
1090:18184788
1041:15843613
990:14610018
941:24844296
887:20660565
879:11349152
836:14610023
787:14614083
701:22407321
644:28724457
590:22580106
532:28717219
465:28532365
409:23083731
314:See also
205:Function
117:of the
81:NeuroLex
53:Location
1934:Neurons
1843:Choroid
1744:Eyebrow
1688:Foveola
1504:(inner)
1404:Choroid
1252:(outer)
1239:of the
1177:3714606
1125:3012877
1081:3057190
1032:6724943
981:2229595
932:4028192
859:Bibcode
851:Science
827:2229589
778:6741006
736:9068848
692:3310952
671:Bibcode
616:Bibcode
581:3401171
523:5514144
492:Bibcode
456:5749398
400:3714606
176:density
74:D051248
36:Details
1838:Retina
1748:Eyelid
1740:Adnexa
1663:Macula
1643:K cell
1639:M cell
1635:P cell
1511:Layers
1502:Retina
1473:Stroma
1316:Stroma
1301:layers
1296:Limbus
1288:Cornea
1260:Sclera
1184:
1174:
1156:Neuron
1131:
1123:
1088:
1078:
1039:
1029:
988:
978:
939:
929:
885:
877:
834:
824:
785:
775:
734:
699:
689:
663:Nature
642:
634:
588:
578:
530:
520:
512:
463:
453:
407:
397:
379:Neuron
195:MEGF11
191:MEGF10
187:Mosaic
119:retina
57:Retina
42:System
1856:Other
1760:Orbit
1678:Fovea
1656:Other
1593:) → (
1576:Cells
1478:Pupil
1394:tunic
1237:globe
1129:S2CID
883:S2CID
640:S2CID
636:26058
135:light
98:[
1796:Lens
1781:Iris
1629:) →
1605:) →
1597:) →
1465:Iris
1182:PMID
1121:PMID
1086:PMID
1037:PMID
986:PMID
937:PMID
875:PMID
832:PMID
783:PMID
732:PMID
697:PMID
632:PMID
586:PMID
528:PMID
510:ISSN
461:PMID
405:PMID
305:The
300:GABA
193:and
69:MeSH
1601:→ (
1172:PMC
1164:doi
1113:doi
1076:PMC
1068:doi
1027:PMC
1017:doi
976:PMC
968:doi
964:122
927:PMC
917:doi
867:doi
855:292
822:PMC
814:doi
810:122
773:PMC
763:doi
724:doi
687:PMC
679:doi
667:483
624:doi
612:200
576:PMC
568:doi
518:PMC
500:doi
451:PMC
443:doi
395:PMC
387:doi
180:cat
145:.
121:of
1920::
1758:,
1754:,
1750:,
1746:,
1645:,
1641:,
1637:,
1633::
1625:,
1621:,
1617:,
1613:,
1589:,
1180:.
1170:.
1160:76
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1127:.
1119:.
1109:26
1107:.
1084:.
1074:.
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1062:.
1058:.
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1025:.
1013:25
1011:.
1007:.
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974:.
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925:.
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907:.
895:^
881:.
873:.
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853:.
830:.
820:.
808:.
804:.
781:.
771:.
759:23
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753:.
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720:36
718:.
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677:.
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661:.
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622:.
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598:^
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564:31
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383:76
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359:^
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83:ID
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869::
861::
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765::
738:.
726::
703:.
681::
673::
646:.
626::
618::
592:.
570::
534:.
502::
494::
488:7
467:.
445::
439:1
411:.
389::
102:]
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