270:
helices contain positively-charged lysine or arginine residues in repeated motifs. In its resting state, half of each S4 helix is in contact with the cell cytosol. Upon depolarization, the positively-charged residues on the S4 domains move toward the exoplasmic surface of the membrane. It is thought that the first 4 arginines account for the gating current, moving toward the extracellular solvent upon channel activation in response to membrane depolarization. The movement of 10–12 of these protein-bound positive charges triggers a conformational change that opens the channel. The exact mechanism by which this movement occurs is not currently agreed upon, however the canonical, transporter, paddle, and twisted models are examples of current theories.
98:
24:
190:
316:
segment makes a scissor-like movement allowing ions to flow through, the S6 segment breaks into two segments allowing of passing of ions through the channel, or the S6 channel serving as the gate itself. The mechanism by which the movement of the S4 segment affects that of S6 is still unknown, however it is theorized that there is a S4-S5 linker whose movement allows the opening of S6.
389:
the duration and rate of action potential firing, which has a significant effect on electrical conduction along an axon as well as synaptic transmission. Potassium channels differ in structure from the other channels in that they contain four separate polypeptide subunits, while the other channels contain four homologous domain but on a single polypeptide unit.
324:. During inactivation, the chain folds in on itself and the ball blocks the flow of ions through the channel. Fast inactivation is directly linked to the activation caused by intramembrane movements of the S4 segments, though the mechanism linking movement of S4 and the engagement of the inactivation gate is unknown.
269:
In general, the voltage sensing portion of the ion channel is responsible for the detection of changes in transmembrane potential that trigger the opening or closing of the channel. The S1-4 alpha helices are generally thought to serve this role. In potassium and sodium channels, voltage-sensing S4
204:
The functionality of voltage-gated ion channels is attributed to its three main discrete units: the voltage sensor, the pore or conducting pathway, and the gate. Na, K, and Ca channels are composed of four transmembrane domains arranged around a central pore; these four domains are part of a single
388:
are the largest and most diverse class of voltage-gated channels, with over 100 encoding human genes. These types of channels differ significantly in their gating properties; some inactivating extremely slowly and others inactivating extremely quickly. This difference in activation time influences
315:
The gate is thought to be coupled to the voltage sensing regions of the channels and appears to contain a mechanical obstruction to ion flow. While the S6 domain has been agreed upon as the segment acting as this obstruction, its exact mechanism is unknown. Possible explanations include: the S6
247:
induces a conformational change in the potassium channel. The conformational change distorts the shape of the channel proteins sufficiently such that the cavity, or channel, opens to allow influx or efflux to occur across the membrane. This movement of ions down their concentration gradients
101:
Ions are depicted by the red circles. A gradient is represented by the different concentration of ions on either side of the membrane. The open conformation of the ion channel allows for the translocation of ions across the cell membrane, while the closed conformation does
297:" allows use of simple and inexpensive model systems to study the function of this region, its role in disease, and pharmaceutical control of its behavior rather than being limited to poorly characterized, expensive, and/or difficult to study preparations.
212:
with specialized functions. The fifth and sixth transmembrane segments (S5 and S6) and pore loop serve the principal role of ion conduction, comprising the gate and pore of the channel, while S1-S4 serve as the voltage-sensing region.
319:
Inactivation of ion channels occurs within milliseconds after opening. Inactivation is thought to be mediated by an intracellular gate that controls the opening of the pore on the inside of the cell. This gate is modeled as a
340:
have similar functional properties across many different cell types. While ten human genes encoding for sodium channels have been identified, their function is typically conserved between species and different cell types.
266:. One of these helices, S4, is the voltage sensing helix. The S4 segment contains many positive charges such that a high positive charge outside the cell repels the helix, keeping the channel in its closed state.
276:
The main functional part of the voltage-sensitive protein domain of these channels generally contains a region composed of S3b and S4 helices, known as the "paddle" due to its shape, which appears to be a
400:
are present in all types of neurons. With the chief responsibility of controlling excitability, chloride channels contribute to the maintenance of cell resting potential and help to regulate cell volume.
175:(Cl) ions have been identified. The opening and closing of the channels are triggered by changing ion concentration, and hence charge gradient, between the sides of the cell membrane.
459:
1499:
Chang, Abraham B.; Lin, Ron; Studley, W. Keith; Tran, Can V.; Saier, Milton H. Jr. (2004). "Phylogeny as a guide to structure and function of membrane transport proteins".
1331:
Cherny, V.V.; Markin, V.S.; DeCoursey, T.E. (1995), "The voltage-activated hydrogen ion conductance in rat alveolar epithelial cells is determined by the pH gradient",
197:
Voltage-gated ion channels are generally composed of several subunits arranged in such a way that there is a central pore through which ions can travel down their
2370:
3280:
513:
377:
nerve endings. In most cells, Ca channels regulate a wide variety of biochemical processes due to their role in controlling intracellular Ca concentrations.
220:
activity, which are located on the inner surface of the cell membrane and do not cross the membrane, and which are coassembled with the α-subunits in the
216:
The four subunits may be identical, or different from one another. In addition to the four central α-subunits, there are also regulatory β-subunits, with
28:
Each of the four homologous domains makes up one subunit of the ion channel. The S4 voltage-sensing segments (marked with + symbols) are shown as charged.
3015:
2267:
2200:
465:
273:
Movement of the voltage-sensor triggers a conformational change of the gate of the conducting pathway, controlling the flow of ions through the channel.
1134:
Webster SM, Del Camino D, Dekker JP, Yellen G (April 2004). "Intracellular gate opening in Shaker K channels defined by high-affinity metal bridges".
205:α-subunit in the case of most Na and Ca channels, whereas there are four α-subunits, each contributing one transmembrane domain, in most K channels.
943:
Long SB, Tao X, Campbell EB, MacKinnon R (November 2007). "Atomic structure of a voltage-dependent K channel in a lipid membrane-like environment".
835:
Murata Y, Iwasaki H, Sasaki M, Inaba K, Okamura Y (June 2005). "Phosphoinositide phosphatase activity coupled to an intrinsic voltage sensor".
84:
72:
1083:
Jiang Y, Lee A, Chen J, Cadene M, Chait BT, MacKinnon R (May 2002). "Crystal structure and mechanism of a calcium-gated potassium channel".
1603:
293:
into rat brain potassium channels results in a fully functional ion channel, as long as the whole intact paddle is replaced. This "
3211:
3194:
3189:
3184:
3179:
361:
With sixteen different identified genes for human calcium channels, this type of channel differs in function between cell types.
281:, interchangeable across a wide variety of cells and species. A similar voltage sensor paddle has also been found in a family of
125:
near the channel. The membrane potential alters the conformation of the channel proteins, regulating their opening and closing.
3216:
2314:
1586:
453:
305:
677:
548:
489:
3268:
2449:
3827:
3166:
2187:
1284:"Molecular dynamics of the sodium channel pore vary with gating: interactions between P-segment motions and inactivation"
2952:
483:
477:
259:
201:. The channels tend to be ion-specific, although similarly sized and charged ions may sometimes travel through them.
168:
471:
3545:
2544:
2322:
1638:
1409:
Ramsey, I. Scott; Mokrab, Younes; Carvacho, Ingrid; Sands, Zara A.; Sansom, Mark S.P.; Clapham, David E. (2010).
385:
164:
1037:
Perozo E, Cortes DM, Cuello LG (July 1999). "Structural rearrangements underlying K-channel activation gating".
3567:
3776:
1687:
350:
282:
1554:
3142:
2054:
495:
409:
262:
are made up of a single polypeptide with four homologous domains. Each domain contains 6 membrane spanning
566:"From ionic currents to molecular mechanisms: the structure and function of voltage-gated sodium channels"
3417:
445:
of voltage-gated sodium channels. Subsequent studies have shown that a variety of other ion channels and
321:
309:
3822:
1595:
1579:
184:
1981:
172:
3761:
2022:
1898:
1564:
1558:
1544:
198:
3766:
356:
97:
3797:
1789:
1776:
1763:
1743:
1730:
1695:
535:
Purves D, Augustine GJ, Fitzpatrick D, Katz LC, LaMantia AS, McNamara JO, Williams SM (2001).
3817:
1572:
255:
249:
232:
221:
160:
110:
1371:
DeCoursey, T.E. (2003), "Voltage-gated proton channels and other proton transfer pathways",
1670:
1233:
1143:
1092:
952:
899:
844:
799:
240:
8:
442:
286:
1237:
1147:
1096:
956:
903:
848:
803:
3137:
2013:
1660:
1524:
1435:
1410:
1349:
1308:
1299:
1283:
1167:
1116:
1062:
1019:
976:
920:
887:
868:
760:
735:
639:
433:
278:
122:
23:
1256:
1221:
711:
694:
582:
565:
133:, thus they must diffuse through the membrane through transmembrane protein channels.
2175:
1516:
1481:
1440:
1392:
1384:
1354:
1313:
1261:
1202:
1159:
1120:
1108:
1054:
1011:
968:
925:
860:
817:
765:
716:
673:
666:
631:
587:
544:
508:
236:
77:
1528:
1066:
2944:
1925:
1508:
1471:
1430:
1422:
1376:
1344:
1336:
1303:
1295:
1251:
1241:
1194:
1171:
1151:
1100:
1046:
1023:
1003:
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960:
915:
907:
872:
852:
807:
755:
747:
706:
643:
623:
577:
397:
374:
366:
193:
Conformation of the four homologous domains showing the formation of a central pore
1222:"Inhibition of inactivation of single sodium channels by a site-directed antibody"
1198:
89:
53:
3480:
1620:
1512:
751:
446:
362:
65:
1050:
3276:
1963:
1380:
1226:
Proceedings of the
National Academy of Sciences of the United States of America
661:
536:
337:
301:
290:
244:
217:
141:
118:
1458:
Koishi R, Xu H, Ren D, Navarro B, Spiller BW, Shi Q, Clapham DE (March 2004).
1007:
812:
787:
189:
159:
Voltage-gated ion-channels are usually ion-specific, and channels specific to
3811:
3230:
2903:
1388:
438:
126:
3263:
3161:
1246:
660:
Lodish H, Berk A, Zipursky SL, Matsudaira P, Baltimore D, Darnell J (2000).
627:
3583:
2974:
1520:
1485:
1476:
1459:
1444:
1396:
1317:
1163:
1112:
1058:
1015:
972:
929:
864:
821:
769:
720:
635:
591:
449:
are phylogenetically related to the voltage-gated ion channels, including:
413:
145:
1358:
1340:
1265:
1206:
994:
Yellen G (August 1998). "The moving parts of voltage-gated ion channels".
888:"Portability of paddle motif function and pharmacology in voltage sensors"
136:
Voltage-gated ion channels have a crucial role in excitable cells such as
3513:
3508:
3503:
3498:
3385:
3224:
1599:
1411:"An aqueous H permeation pathway in the voltage-gated proton channel Hv1"
263:
209:
156:, voltage-gated ion channels directionally propagate electrical signals.
114:
1549:
1155:
964:
911:
856:
300:
Although voltage-gated ion channels are typically activated by membrane
3493:
2859:
2227:
2195:
736:"Ion channel voltage sensors: structure, function, and pathophysiology"
370:
294:
208:
The membrane-spanning segments, designated S1-S6, all take the form of
1426:
441:
studies of proteins expressed in bacteria revealed the existence of a
3578:
3488:
1335:, vol. 105, no. 6 (published June 1995), pp. 861–896,
417:
1594:
1282:
Bénitah JP, Chen Z, Balser JR, Tomaselli GF, Marbán E (March 1999).
1104:
886:
Alabi AA, Bahamonde MI, Jung HJ, Kim JI, Swartz KJ (November 2007).
3591:
2939:
1918:
1913:
1908:
1903:
1835:
1830:
369:
similarly to Na channels in some neurons. They also play a role in
662:"Section 21.3, Molecular Properties of Voltage-Gated Ion Channels"
3718:
3475:
3311:
1812:
1807:
1802:
1185:
Armstrong CM (July 1981). "Sodium channels and gating currents".
153:
3395:
3390:
3124:
3119:
3065:
3060:
2876:
2864:
2854:
2849:
2837:
2832:
2827:
2714:
2709:
2704:
2596:
2532:
2527:
2522:
2517:
2512:
2507:
2502:
2437:
2432:
2422:
2417:
2412:
2363:
2358:
2348:
2343:
2220:
2215:
2210:
2205:
2111:
2106:
2101:
2091:
2086:
2076:
2071:
2001:
1996:
1991:
1986:
1889:
1884:
1879:
1874:
1862:
1857:
1852:
1847:
1756:
1723:
1718:
1713:
1708:
137:
534:
3629:
3560:
3555:
3550:
3467:
3454:
3449:
3444:
3439:
3434:
3429:
3380:
3373:
3368:
3363:
3358:
3353:
3348:
3343:
3338:
3326:
3321:
3316:
3306:
3301:
3296:
3291:
3206:
3150:
3107:
3102:
3097:
3092:
3087:
3082:
3077:
3055:
3050:
3045:
3040:
3035:
3030:
3025:
3006:
3001:
2996:
2991:
2979:
2896:
2891:
2886:
2871:
2809:
2804:
2799:
2790:
2785:
2771:
2766:
2757:
2752:
2747:
2738:
2733:
2724:
2719:
2686:
2681:
2676:
2671:
2662:
2653:
2648:
2643:
2634:
2629:
2624:
2619:
2610:
2605:
2591:
2586:
2581:
2576:
2571:
2566:
2561:
2497:
2492:
2487:
2482:
2477:
2472:
2467:
2462:
2427:
2397:
2392:
2387:
2382:
2353:
2302:
2297:
2292:
2287:
2277:
2272:
2252:
2237:
2232:
2147:
2142:
2137:
2132:
2116:
2096:
2081:
2042:
2037:
2032:
2027:
1935:
1930:
1653:
1648:
1643:
1133:
1460:"A superfamily of voltage-gated sodium channels in bacteria"
659:
424:-dependent manner. They function to remove acid from cells.
3709:
3704:
3699:
3689:
3684:
3679:
3669:
3664:
3659:
3654:
3649:
3644:
3639:
3624:
3619:
3614:
3609:
3604:
3599:
3538:
3533:
3528:
3523:
3518:
3424:
3412:
3407:
3402:
3333:
3286:
3250:
3245:
3240:
3235:
2962:
2780:
2338:
1675:
1665:
149:
60:
47:
1281:
460:
ryanodine-inositol 1,4,5-triphosphate receptor Ca channels
2282:
2242:
1408:
695:"Structure of a voltage-dependent K channel beta subunit"
130:
1550:
The IUPHAR Compendium of
Voltage-gated Ion Channels 2005
1219:
942:
834:
614:
Bezanilla F (March 2005). "Voltage-gated ion channels".
289:
of the paddle region from a species of volcano-dwelling
421:
543:(2nd ed.). Sunderland, Mass: Sinauer Associates.
140:
and muscle tissues, allowing a rapid and co-ordinated
1330:
885:
672:(4th ed.). New York: Scientific American Books.
514:
Catecholaminergic polymorphic ventricular tachycardia
1220:
Vassilev P, Scheuer T, Catterall WA (October 1989).
1498:
785:
692:
530:
528:
1457:
1082:
1036:
665:
3809:
3752:
609:
607:
605:
603:
601:
525:
243:is introduced over the membrane, the associated
1277:
1275:
1375:, vol. 83, no. 2, pp. 475–579,
1580:
1078:
1076:
598:
1545:IUPHAR-DB Voltage-gated ion channel subunits
1272:
936:
781:
779:
693:Gulbis JM, Mann S, MacKinnon R (June 1999).
490:monovalent cation:proton antiporters, type 1
252:sufficient to depolarize the cell membrane.
879:
727:
420:, and are activated by depolarization in a
1587:
1573:
1073:
655:
653:
392:
380:
344:
22:
2171:
1616:
1557:at the U.S. National Library of Medicine
1475:
1434:
1415:Nature Structural & Molecular Biology
1370:
1348:
1307:
1255:
1245:
1184:
919:
811:
776:
759:
733:
710:
613:
581:
563:
16:Type of ion channel transmembrane protein
466:transient receptor potential Ca channels
332:
188:
96:
3740:
1959:
786:Sands Z, Grottesi A, Sansom MS (2005).
650:
404:
306:inward-rectifier potassium ion channels
3810:
993:
3739:
2924:
2170:
1958:
1615:
1568:
2953:Calcium-activated chloride channels
1464:The Journal of Biological Chemistry
616:IEEE Transactions on NanoBioscience
484:calcium-dependent chloride channels
248:subsequently generates an electric
117:that are activated by changes in a
13:
2023:Amiloride-sensitive cation channel
1300:10.1523/JNEUROSCI.19-05-01577.1999
327:
14:
3839:
2925:
1538:
322:ball tethered to a flexible chain
1555:Voltage-Dependent+Anion+Channels
427:
3546:Voltage-dependent anion channel
1639:Inositol trisphosphate receptor
1492:
1451:
1402:
1364:
1324:
1213:
1178:
1127:
1030:
996:Quarterly Reviews of Biophysics
987:
3568:General bacterial porin family
828:
686:
557:
454:inwardly rectifying K channels
283:voltage sensitive phosphatases
1:
1333:Journal of General Physiology
1199:10.1152/physrev.1981.61.3.644
712:10.1016/s0092-8674(00)80805-3
583:10.1016/S0896-6273(00)81133-2
519:
410:Voltage-gated proton channels
351:Voltage-gated calcium channel
256:Voltage-gated sodium channels
129:are generally impermeable to
1513:10.1080/09687680410001720830
788:"Voltage-gated ion channels"
752:10.1016/j.neuron.2010.08.021
537:"Voltage-Gated Ion Channels"
478:glutamate-gated ion channels
227:
178:
7:
1288:The Journal of Neuroscience
1051:10.1126/science.285.5424.73
564:Catterall WA (April 2000).
502:
412:carry currents mediated by
308:, are activated instead by
10:
3844:
3828:Integral membrane proteins
2987:Chloride Channel Accessory
1596:Membrane transport protein
1381:10.1152/physrev.00028.2002
472:polycystin cation channels
431:
354:
348:
185:Cation channel superfamily
182:
144:in response to triggering
107:Voltage-gated ion channels
3793:
3748:
3735:
3576:
3465:
3261:
3159:
3135:
2937:
2933:
2920:
2543:
2448:
2313:
2186:
2182:
2166:
2053:
2012:
1982:Epithelial sodium channel
1974:
1970:
1954:
1686:
1631:
1627:
1611:
1008:10.1017/s0033583598003448
813:10.1016/j.cub.2004.12.050
304:, some channels, such as
199:electrochemical gradients
83:
71:
59:
46:
38:
33:
21:
1899:Cation channels of sperm
1559:Medical Subject Headings
371:neurotransmitter release
239:have shown that, when a
235:structural studies of a
1247:10.1073/pnas.86.20.8147
628:10.1109/tnb.2004.842463
357:Calcium channel blocker
1477:10.1074/jbc.M313100200
668:Molecular Cell Biology
496:potassium transporters
393:Chloride (Cl) channels
381:Potassium (K) channels
194:
111:transmembrane proteins
103:
1975:Constitutively active
1373:Physiological Reviews
1341:10.1085/jgp.105.6.861
1187:Physiological Reviews
734:Catterall WA (2010).
432:Further information:
345:Calcium (Ca) channels
222:endoplasmic reticulum
192:
183:Further information:
100:
333:Sodium (Na) channels
285:in various species.
241:potential difference
3742:By gating mechanism
1238:1989PNAS...86.8147V
1156:10.1038/nature02468
1148:2004Natur.428..864W
1097:2002Natur.417..515J
965:10.1038/nature06265
957:2007Natur.450..376L
912:10.1038/nature06266
904:2007Natur.450..370A
857:10.1038/nature03650
849:2005Natur.435.1239M
804:2005CBio...15..R44S
405:Proton (H) channels
287:Genetic engineering
3269:TRP cation channel
3167:CNG cation channel
2450:Tandem pore domain
1661:Ryanodine receptor
434:Ion channel family
386:Potassium channels
279:conserved sequence
195:
148:. Found along the
123:membrane potential
104:
3823:Electrophysiology
3805:
3804:
3789:
3788:
3785:
3784:
3777:Stretch-activated
3754:Ion channel class
3731:
3730:
3727:
3726:
2916:
2915:
2912:
2911:
2188:Calcium-activated
2176:Potassium channel
2162:
2161:
2158:
2157:
1950:
1949:
1946:
1945:
1427:10.1038/nsmb.1826
843:(7046): 1239–43.
679:978-0-7167-3136-8
550:978-0-87893-742-4
509:Potassium channel
398:Chloride channels
367:action potentials
310:hyperpolarization
237:potassium channel
95:
94:
3835:
3750:
3749:
3737:
3736:
2945:Chloride channel
2935:
2934:
2922:
2921:
2315:Inward-rectifier
2184:
2183:
2168:
2167:
1972:
1971:
1956:
1955:
1926:Two-pore channel
1629:
1628:
1613:
1612:
1589:
1582:
1575:
1566:
1565:
1533:
1532:
1496:
1490:
1489:
1479:
1455:
1449:
1448:
1438:
1406:
1400:
1399:
1368:
1362:
1361:
1352:
1328:
1322:
1321:
1311:
1279:
1270:
1269:
1259:
1249:
1217:
1211:
1210:
1182:
1176:
1175:
1131:
1125:
1124:
1091:(6888): 515–22.
1080:
1071:
1070:
1034:
1028:
1027:
991:
985:
984:
951:(7168): 376–82.
940:
934:
933:
923:
883:
877:
876:
832:
826:
825:
815:
783:
774:
773:
763:
731:
725:
724:
714:
690:
684:
683:
671:
657:
648:
647:
611:
596:
595:
585:
561:
555:
554:
532:
260:calcium channels
233:Crystallographic
26:
19:
18:
3843:
3842:
3838:
3837:
3836:
3834:
3833:
3832:
3808:
3807:
3806:
3801:
3781:
3744:
3723:
3572:
3471:
3461:
3257:
3155:
3131:
2929:
2908:
2822:
2699:
2556:
2539:
2444:
2408:
2378:
2334:
2326:
2309:
2263:
2178:
2154:
2127:
2066:
2049:
2008:
1966:
1942:
1825:
1797:
1784:
1771:
1751:
1738:
1703:
1682:
1623:
1621:Calcium channel
1607:
1593:
1541:
1536:
1497:
1493:
1456:
1452:
1407:
1403:
1369:
1365:
1329:
1325:
1280:
1273:
1232:(20): 8147–51.
1218:
1214:
1183:
1179:
1142:(6985): 864–8.
1132:
1128:
1105:10.1038/417515a
1081:
1074:
1035:
1031:
992:
988:
941:
937:
898:(7168): 370–5.
884:
880:
833:
829:
792:Current Biology
784:
777:
732:
728:
691:
687:
680:
658:
651:
612:
599:
562:
558:
551:
533:
526:
522:
505:
436:
430:
416:in the form of
407:
395:
383:
359:
353:
347:
338:Sodium channels
335:
330:
328:Different types
230:
187:
181:
109:are a class of
73:OPM superfamily
29:
17:
12:
11:
5:
3841:
3831:
3830:
3825:
3820:
3803:
3802:
3794:
3791:
3790:
3787:
3786:
3783:
3782:
3780:
3779:
3774:
3769:
3764:
3758:
3756:
3746:
3745:
3733:
3732:
3729:
3728:
3725:
3724:
3722:
3721:
3716:
3715:
3714:
3713:
3712:
3707:
3702:
3694:
3693:
3692:
3687:
3682:
3674:
3673:
3672:
3667:
3662:
3657:
3652:
3647:
3642:
3634:
3633:
3632:
3627:
3622:
3617:
3612:
3607:
3602:
3588:
3586:
3574:
3573:
3571:
3570:
3565:
3564:
3563:
3558:
3553:
3543:
3542:
3541:
3536:
3531:
3526:
3521:
3516:
3511:
3506:
3501:
3496:
3485:
3483:
3469:
3463:
3462:
3460:
3459:
3458:
3457:
3452:
3447:
3442:
3437:
3432:
3422:
3421:
3420:
3415:
3405:
3400:
3399:
3398:
3393:
3388:
3378:
3377:
3376:
3371:
3366:
3361:
3356:
3351:
3346:
3341:
3331:
3330:
3329:
3324:
3319:
3314:
3309:
3304:
3299:
3294:
3284:
3273:
3271:
3259:
3258:
3256:
3255:
3254:
3253:
3248:
3243:
3238:
3233:
3221:
3220:
3219:
3214:
3209:
3199:
3198:
3197:
3192:
3187:
3182:
3171:
3169:
3157:
3156:
3154:
3153:
3147:
3145:
3143:Proton channel
3133:
3132:
3130:
3129:
3128:
3127:
3122:
3112:
3111:
3110:
3105:
3100:
3095:
3090:
3085:
3080:
3070:
3069:
3068:
3063:
3058:
3053:
3048:
3043:
3038:
3033:
3028:
3018:
3013:
3012:
3011:
3010:
3009:
3004:
2999:
2994:
2984:
2983:
2982:
2977:
2967:
2966:
2965:
2949:
2947:
2931:
2930:
2918:
2917:
2914:
2913:
2910:
2909:
2907:
2906:
2901:
2900:
2899:
2894:
2889:
2879:
2874:
2869:
2868:
2867:
2862:
2857:
2852:
2842:
2841:
2840:
2835:
2830:
2820:
2815:
2814:
2813:
2812:
2807:
2802:
2795:
2794:
2793:
2788:
2783:
2776:
2775:
2774:
2769:
2762:
2761:
2760:
2755:
2750:
2743:
2742:
2741:
2736:
2729:
2728:
2727:
2722:
2717:
2712:
2707:
2697:
2692:
2691:
2690:
2689:
2684:
2679:
2674:
2667:
2666:
2665:
2658:
2657:
2656:
2651:
2646:
2639:
2638:
2637:
2632:
2627:
2622:
2615:
2614:
2613:
2608:
2601:
2600:
2599:
2594:
2589:
2584:
2579:
2574:
2569:
2564:
2554:
2549:
2547:
2541:
2540:
2538:
2537:
2536:
2535:
2530:
2525:
2520:
2515:
2510:
2505:
2500:
2495:
2490:
2485:
2480:
2475:
2470:
2465:
2454:
2452:
2446:
2445:
2443:
2442:
2441:
2440:
2435:
2430:
2425:
2420:
2415:
2406:
2402:
2401:
2400:
2395:
2390:
2385:
2376:
2368:
2367:
2366:
2361:
2356:
2351:
2346:
2341:
2332:
2328:
2324:
2319:
2317:
2311:
2310:
2308:
2307:
2306:
2305:
2300:
2295:
2290:
2285:
2280:
2275:
2270:
2261:
2257:
2256:
2255:
2247:
2246:
2245:
2240:
2235:
2225:
2224:
2223:
2218:
2213:
2208:
2203:
2192:
2190:
2180:
2179:
2164:
2163:
2160:
2159:
2156:
2155:
2153:
2152:
2151:
2150:
2145:
2140:
2135:
2125:
2121:
2120:
2119:
2114:
2109:
2104:
2099:
2094:
2089:
2084:
2079:
2074:
2064:
2059:
2057:
2051:
2050:
2048:
2047:
2046:
2045:
2040:
2035:
2030:
2019:
2017:
2010:
2009:
2007:
2006:
2005:
2004:
1999:
1994:
1989:
1978:
1976:
1968:
1967:
1964:Sodium channel
1952:
1951:
1948:
1947:
1944:
1943:
1941:
1940:
1939:
1938:
1933:
1923:
1922:
1921:
1916:
1911:
1906:
1895:
1894:
1893:
1892:
1887:
1882:
1877:
1867:
1866:
1865:
1860:
1855:
1850:
1840:
1839:
1838:
1833:
1823:
1818:
1817:
1816:
1815:
1810:
1805:
1795:
1787:
1782:
1774:
1769:
1761:
1760:
1759:
1749:
1741:
1736:
1728:
1727:
1726:
1721:
1716:
1711:
1701:
1692:
1690:
1684:
1683:
1681:
1680:
1679:
1678:
1673:
1668:
1658:
1657:
1656:
1651:
1646:
1635:
1633:
1625:
1624:
1609:
1608:
1592:
1591:
1584:
1577:
1569:
1563:
1562:
1552:
1547:
1540:
1539:External links
1537:
1535:
1534:
1507:(3): 171–181.
1501:Mol Membr Biol
1491:
1470:(10): 9532–8.
1450:
1421:(7): 869–875.
1401:
1363:
1323:
1294:(5): 1577–85.
1271:
1212:
1177:
1126:
1072:
1045:(5424): 73–8.
1029:
986:
935:
878:
827:
775:
726:
685:
678:
649:
597:
556:
549:
523:
521:
518:
517:
516:
511:
504:
501:
500:
499:
493:
487:
481:
475:
469:
463:
457:
429:
426:
406:
403:
394:
391:
382:
379:
349:Main article:
346:
343:
334:
331:
329:
326:
302:depolarization
291:archaebacteria
245:electric field
229:
226:
218:oxidoreductase
180:
177:
146:voltage change
142:depolarization
127:Cell membranes
93:
92:
87:
81:
80:
75:
69:
68:
63:
57:
56:
51:
44:
43:
40:
36:
35:
31:
30:
27:
15:
9:
6:
4:
3:
2:
3840:
3829:
3826:
3824:
3821:
3819:
3816:
3815:
3813:
3800:
3799:
3792:
3778:
3775:
3773:
3772:Voltage-gated
3770:
3768:
3765:
3763:
3760:
3759:
3757:
3755:
3751:
3747:
3743:
3738:
3734:
3720:
3717:
3711:
3708:
3706:
3703:
3701:
3698:
3697:
3695:
3691:
3688:
3686:
3683:
3681:
3678:
3677:
3675:
3671:
3668:
3666:
3663:
3661:
3658:
3656:
3653:
3651:
3648:
3646:
3643:
3641:
3638:
3637:
3635:
3631:
3628:
3626:
3623:
3621:
3618:
3616:
3613:
3611:
3608:
3606:
3603:
3601:
3598:
3597:
3595:
3594:
3593:
3590:
3589:
3587:
3585:
3582:
3580:
3575:
3569:
3566:
3562:
3559:
3557:
3554:
3552:
3549:
3548:
3547:
3544:
3540:
3537:
3535:
3532:
3530:
3527:
3525:
3522:
3520:
3517:
3515:
3512:
3510:
3507:
3505:
3502:
3500:
3497:
3495:
3492:
3491:
3490:
3487:
3486:
3484:
3482:
3479:
3477:
3473:
3464:
3456:
3453:
3451:
3448:
3446:
3443:
3441:
3438:
3436:
3433:
3431:
3428:
3427:
3426:
3423:
3419:
3416:
3414:
3411:
3410:
3409:
3406:
3404:
3401:
3397:
3394:
3392:
3389:
3387:
3384:
3383:
3382:
3379:
3375:
3372:
3370:
3367:
3365:
3362:
3360:
3357:
3355:
3352:
3350:
3347:
3345:
3342:
3340:
3337:
3336:
3335:
3332:
3328:
3325:
3323:
3320:
3318:
3315:
3313:
3310:
3308:
3305:
3303:
3300:
3298:
3295:
3293:
3290:
3289:
3288:
3285:
3282:
3278:
3275:
3274:
3272:
3270:
3267:
3265:
3260:
3252:
3249:
3247:
3244:
3242:
3239:
3237:
3234:
3232:
3229:
3228:
3227:
3226:
3222:
3218:
3215:
3213:
3210:
3208:
3205:
3204:
3203:
3200:
3196:
3193:
3191:
3188:
3186:
3183:
3181:
3178:
3177:
3176:
3173:
3172:
3170:
3168:
3165:
3163:
3158:
3152:
3149:
3148:
3146:
3144:
3141:
3139:
3134:
3126:
3123:
3121:
3118:
3117:
3116:
3113:
3109:
3106:
3104:
3101:
3099:
3096:
3094:
3091:
3089:
3086:
3084:
3081:
3079:
3076:
3075:
3074:
3071:
3067:
3064:
3062:
3059:
3057:
3054:
3052:
3049:
3047:
3044:
3042:
3039:
3037:
3034:
3032:
3029:
3027:
3024:
3023:
3022:
3019:
3017:
3014:
3008:
3005:
3003:
3000:
2998:
2995:
2993:
2990:
2989:
2988:
2985:
2981:
2978:
2976:
2973:
2972:
2971:
2968:
2964:
2961:
2960:
2959:
2956:
2955:
2954:
2951:
2950:
2948:
2946:
2943:
2941:
2936:
2932:
2928:
2927:Miscellaneous
2923:
2919:
2905:
2904:Shaker (gene)
2902:
2898:
2895:
2893:
2890:
2888:
2885:
2884:
2883:
2880:
2878:
2877:minK/ISK-like
2875:
2873:
2870:
2866:
2863:
2861:
2858:
2856:
2853:
2851:
2848:
2847:
2846:
2843:
2839:
2836:
2834:
2831:
2829:
2826:
2825:
2824:
2817:
2816:
2811:
2808:
2806:
2803:
2801:
2798:
2797:
2796:
2792:
2789:
2787:
2784:
2782:
2779:
2778:
2777:
2773:
2770:
2768:
2765:
2764:
2763:
2759:
2756:
2754:
2751:
2749:
2746:
2745:
2744:
2740:
2737:
2735:
2732:
2731:
2730:
2726:
2723:
2721:
2718:
2716:
2713:
2711:
2708:
2706:
2703:
2702:
2701:
2694:
2693:
2688:
2685:
2683:
2680:
2678:
2675:
2673:
2670:
2669:
2668:
2664:
2661:
2660:
2659:
2655:
2652:
2650:
2647:
2645:
2642:
2641:
2640:
2636:
2633:
2631:
2628:
2626:
2623:
2621:
2618:
2617:
2616:
2612:
2609:
2607:
2604:
2603:
2602:
2598:
2595:
2593:
2590:
2588:
2585:
2583:
2580:
2578:
2575:
2573:
2570:
2568:
2565:
2563:
2560:
2559:
2558:
2551:
2550:
2548:
2546:
2545:Voltage-gated
2542:
2534:
2531:
2529:
2526:
2524:
2521:
2519:
2516:
2514:
2511:
2509:
2506:
2504:
2501:
2499:
2496:
2494:
2491:
2489:
2486:
2484:
2481:
2479:
2476:
2474:
2471:
2469:
2466:
2464:
2461:
2460:
2459:
2456:
2455:
2453:
2451:
2447:
2439:
2436:
2434:
2431:
2429:
2426:
2424:
2421:
2419:
2416:
2414:
2411:
2410:
2409:
2403:
2399:
2396:
2394:
2391:
2389:
2386:
2384:
2381:
2380:
2379:
2372:
2369:
2365:
2362:
2360:
2357:
2355:
2352:
2350:
2347:
2345:
2342:
2340:
2337:
2336:
2335:
2329:
2327:
2321:
2320:
2318:
2316:
2312:
2304:
2301:
2299:
2296:
2294:
2291:
2289:
2286:
2284:
2281:
2279:
2276:
2274:
2271:
2269:
2266:
2265:
2264:
2258:
2254:
2251:
2250:
2248:
2244:
2241:
2239:
2236:
2234:
2231:
2230:
2229:
2226:
2222:
2219:
2217:
2214:
2212:
2209:
2207:
2204:
2202:
2199:
2198:
2197:
2194:
2193:
2191:
2189:
2185:
2181:
2177:
2174:
2169:
2165:
2149:
2146:
2144:
2141:
2139:
2136:
2134:
2131:
2130:
2129:
2122:
2118:
2115:
2113:
2110:
2108:
2105:
2103:
2100:
2098:
2095:
2093:
2090:
2088:
2085:
2083:
2080:
2078:
2075:
2073:
2070:
2069:
2068:
2061:
2060:
2058:
2056:
2055:Voltage-gated
2052:
2044:
2041:
2039:
2036:
2034:
2031:
2029:
2026:
2025:
2024:
2021:
2020:
2018:
2015:
2011:
2003:
2000:
1998:
1995:
1993:
1990:
1988:
1985:
1984:
1983:
1980:
1979:
1977:
1973:
1969:
1965:
1962:
1957:
1953:
1937:
1934:
1932:
1929:
1928:
1927:
1924:
1920:
1917:
1915:
1912:
1910:
1907:
1905:
1902:
1901:
1900:
1897:
1896:
1891:
1888:
1886:
1883:
1881:
1878:
1876:
1873:
1872:
1871:
1868:
1864:
1861:
1859:
1856:
1854:
1851:
1849:
1846:
1845:
1844:
1841:
1837:
1834:
1832:
1829:
1828:
1827:
1820:
1819:
1814:
1811:
1809:
1806:
1804:
1801:
1800:
1799:
1791:
1788:
1786:
1778:
1775:
1773:
1765:
1762:
1758:
1755:
1754:
1753:
1745:
1742:
1740:
1732:
1729:
1725:
1722:
1720:
1717:
1715:
1712:
1710:
1707:
1706:
1705:
1697:
1694:
1693:
1691:
1689:
1688:Voltage-gated
1685:
1677:
1674:
1672:
1669:
1667:
1664:
1663:
1662:
1659:
1655:
1652:
1650:
1647:
1645:
1642:
1641:
1640:
1637:
1636:
1634:
1630:
1626:
1622:
1619:
1614:
1610:
1605:
1601:
1597:
1590:
1585:
1583:
1578:
1576:
1571:
1570:
1567:
1560:
1556:
1553:
1551:
1548:
1546:
1543:
1542:
1530:
1526:
1522:
1518:
1514:
1510:
1506:
1502:
1495:
1487:
1483:
1478:
1473:
1469:
1465:
1461:
1454:
1446:
1442:
1437:
1432:
1428:
1424:
1420:
1416:
1412:
1405:
1398:
1394:
1390:
1386:
1382:
1378:
1374:
1367:
1360:
1356:
1351:
1346:
1342:
1338:
1334:
1327:
1319:
1315:
1310:
1305:
1301:
1297:
1293:
1289:
1285:
1278:
1276:
1267:
1263:
1258:
1253:
1248:
1243:
1239:
1235:
1231:
1227:
1223:
1216:
1208:
1204:
1200:
1196:
1193:(3): 644–83.
1192:
1188:
1181:
1173:
1169:
1165:
1161:
1157:
1153:
1149:
1145:
1141:
1137:
1130:
1122:
1118:
1114:
1110:
1106:
1102:
1098:
1094:
1090:
1086:
1079:
1077:
1068:
1064:
1060:
1056:
1052:
1048:
1044:
1040:
1033:
1025:
1021:
1017:
1013:
1009:
1005:
1002:(3): 239–95.
1001:
997:
990:
982:
978:
974:
970:
966:
962:
958:
954:
950:
946:
939:
931:
927:
922:
917:
913:
909:
905:
901:
897:
893:
889:
882:
874:
870:
866:
862:
858:
854:
850:
846:
842:
838:
831:
823:
819:
814:
809:
805:
801:
797:
793:
789:
782:
780:
771:
767:
762:
757:
753:
749:
746:(6): 915–28.
745:
741:
737:
730:
722:
718:
713:
708:
705:(7): 943–52.
704:
700:
696:
689:
681:
675:
670:
669:
663:
656:
654:
645:
641:
637:
633:
629:
625:
621:
617:
610:
608:
606:
604:
602:
593:
589:
584:
579:
575:
571:
567:
560:
552:
546:
542:
538:
531:
529:
524:
515:
512:
510:
507:
506:
497:
494:
491:
488:
485:
482:
479:
476:
473:
470:
467:
464:
461:
458:
455:
452:
451:
450:
448:
444:
440:
435:
428:Phylogenetics
425:
423:
419:
415:
414:hydrogen ions
411:
402:
399:
390:
387:
378:
376:
372:
368:
364:
358:
352:
342:
339:
325:
323:
317:
313:
311:
307:
303:
298:
296:
292:
288:
284:
280:
274:
271:
267:
265:
264:alpha helices
261:
257:
253:
251:
246:
242:
238:
234:
225:
223:
219:
214:
211:
210:alpha helices
206:
202:
200:
191:
186:
176:
174:
170:
166:
162:
157:
155:
151:
147:
143:
139:
134:
132:
128:
124:
120:
116:
112:
108:
99:
91:
88:
86:
82:
79:
76:
74:
70:
67:
64:
62:
58:
55:
52:
49:
45:
41:
37:
32:
25:
20:
3818:Ion channels
3795:
3771:
3762:Ligand-gated
3753:
3741:
3584:Gap junction
3577:
3466:
3262:
3223:
3201:
3174:
3160:
3136:
3114:
3072:
3020:
2986:
2969:
2957:
2938:
2926:
2881:
2844:
2818:
2695:
2552:
2457:
2404:
2374:
2330:
2259:
2172:
2123:
2062:
1960:
1869:
1842:
1821:
1793:
1780:
1767:
1747:
1734:
1699:
1632:Ligand-gated
1617:
1600:ion channels
1504:
1500:
1494:
1467:
1463:
1453:
1418:
1414:
1404:
1372:
1366:
1332:
1326:
1291:
1287:
1229:
1225:
1215:
1190:
1186:
1180:
1139:
1135:
1129:
1088:
1084:
1042:
1038:
1032:
999:
995:
989:
948:
944:
938:
895:
891:
881:
840:
836:
830:
798:(2): R44–7.
795:
791:
743:
739:
729:
702:
698:
688:
667:
622:(1): 34–48.
619:
615:
576:(1): 13–25.
573:
569:
559:
541:Neuroscience
540:
447:transporters
439:Phylogenetic
437:
408:
396:
384:
375:pre-synaptic
360:
336:
318:
314:
299:
275:
272:
268:
254:
231:
215:
207:
203:
196:
158:
135:
115:ion channels
106:
105:
3767:Light-gated
2249:IK channel
443:superfamily
363:Ca channels
152:and at the
121:electrical
85:OPM protein
34:Identifiers
3812:Categories
2970:Bestrophin
2228:SK channel
2196:BK channel
1870:γ-subunits
1843:β-subunits
1826:δ-subunits
520:References
355:See also:
295:modularity
171:(Ca), and
113:that form
3798:disorders
3796:see also
3579:Cytoplasm
3489:Aquaporin
2958:Anoctamin
2781:11.1/hERG
1389:205658168
1121:205029269
418:hydronium
228:Mechanism
179:Structure
165:potassium
3592:Connexin
2872:minK/ISK
1529:45284885
1521:15204625
1486:14665618
1445:20543828
1397:12663866
1318:10024345
1164:15103379
1113:12037559
1067:26775433
1059:10390363
1016:10384687
973:18004376
930:18004375
865:15902207
822:15668152
770:20869590
721:10399921
636:15816170
592:10798388
503:See also
365:produce
173:chloride
138:neuronal
3719:Innexin
3476:solutes
1436:4035905
1359:7561747
1350:2216954
1309:6782169
1266:2554301
1234:Bibcode
1207:6265962
1172:1329210
1144:Bibcode
1093:Bibcode
1039:Science
1024:2605660
981:4320272
953:Bibcode
921:2709416
900:Bibcode
873:4427755
845:Bibcode
800:Bibcode
761:2950829
644:8212388
250:current
169:calcium
154:synapse
2016:-gated
2014:Proton
1790:T-type
1777:R-type
1764:Q-type
1744:P-type
1731:N-type
1696:L-type
1561:(MeSH)
1527:
1519:
1484:
1443:
1433:
1395:
1387:
1357:
1347:
1316:
1306:
1264:
1257:298232
1254:
1205:
1170:
1162:
1136:Nature
1119:
1111:
1085:Nature
1065:
1057:
1022:
1014:
979:
971:
945:Nature
928:
918:
892:Nature
871:
863:
837:Nature
820:
768:
758:
740:Neuron
719:
676:
642:
634:
590:
570:Neuron
547:
163:(Na),
161:sodium
119:cell's
54:CL0030
39:Symbol
3630:GJA10
3481:Porin
3381:TRPML
3151:HVCN1
2845:KCNIP
2700:α7-12
1604:TC 1A
1525:S2CID
1168:S2CID
1117:S2CID
1063:S2CID
1020:S2CID
977:S2CID
869:S2CID
640:S2CID
492:, and
167:(K),
66:1.A.1
3710:GJD4
3705:GJD3
3700:GJD2
3690:GJC3
3685:GJC2
3680:GJC1
3670:GJB7
3665:GJB6
3660:GJB5
3655:GJB4
3650:GJB3
3645:GJB2
3640:GJB1
3625:GJA9
3620:GJA8
3615:GJA5
3610:GJA4
3605:GJA3
3600:GJA1
3425:TRPV
3408:TRPP
3403:TRPN
3334:TRPM
3287:TRPC
3277:TRPA
3115:CLNS
3073:CLIC
3021:CLCN
3016:CFTR
2963:ANO1
2882:MiRP
2810:12.3
2805:12.2
2800:12.1
2791:11.3
2786:11.2
2772:10.2
2767:10.1
2557:α1-6
2371:GIRK
1785:α2.3
1772:α2.1
1739:α2.2
1517:PMID
1482:PMID
1441:PMID
1393:PMID
1385:OCLC
1355:PMID
1314:PMID
1262:PMID
1203:PMID
1160:PMID
1109:PMID
1055:PMID
1012:PMID
969:PMID
926:PMID
861:PMID
818:PMID
766:PMID
717:PMID
699:Cell
674:ISBN
632:PMID
588:PMID
545:ISBN
258:and
150:axon
131:ions
102:not.
90:2a79
61:TCDB
50:clan
48:Pfam
3474:(+
3312:4AP
3225:HCN
2758:9.3
2753:9.2
2748:9.1
2739:8.2
2734:8.1
2725:7.5
2720:7.4
2715:7.3
2710:7.2
2705:7.1
2687:6.4
2682:6.3
2677:6.2
2672:6.1
2663:5.1
2654:4.3
2649:4.2
2644:4.1
2635:3.4
2630:3.3
2625:3.2
2620:3.1
2611:2.2
2606:2.1
2597:1.8
2592:1.7
2587:1.6
2582:1.5
2577:1.4
2572:1.3
2567:1.2
2562:1.1
2458:K2P
2438:7.1
2433:6.2
2428:6.1
2423:5.1
2418:4.2
2413:4.1
2398:3.4
2393:3.3
2388:3.2
2383:3.1
2364:2.6
2359:2.4
2354:2.3
2349:2.2
2344:2.1
2339:1.1
2325:ATP
2303:5.1
2298:4.2
2293:4.1
2288:3.1
2283:2.3
2278:2.2
2273:2.1
2268:1.1
2253:IK1
2243:SK3
2238:SK2
2233:SK1
2112:1.9
2107:1.8
2102:1.7
2097:1.6
2092:1.5
2087:1.4
2082:1.3
2077:1.2
2072:1.1
1961:Na:
1813:3.3
1808:3.2
1803:3.1
1757:2.1
1724:1.4
1719:1.3
1714:1.2
1709:1.1
1618:Ca:
1509:doi
1472:doi
1468:279
1431:PMC
1423:doi
1377:doi
1345:PMC
1337:doi
1304:PMC
1296:doi
1252:PMC
1242:doi
1195:doi
1152:doi
1140:428
1101:doi
1089:417
1047:doi
1043:285
1004:doi
961:doi
949:450
916:PMC
908:doi
896:450
853:doi
841:435
808:doi
756:PMC
748:doi
707:doi
624:doi
578:doi
373:in
42:VIC
3814::
3696:D
3676:C
3636:B
3596:A
3478:):
3231:FC
3125:1B
3120:1A
3108:L1
3066:KB
3061:KA
2940:Cl
2533:18
2528:17
2523:16
2518:15
2513:13
2508:12
2503:10
2407:ir
2377:ir
2333:ir
2262:Ca
2221:β4
2216:β3
2211:β2
2206:β1
2201:α1
2173:K:
2124:Na
2117:7A
2063:Na
1890:γ4
1885:γ3
1880:γ2
1875:γ1
1863:β4
1858:β3
1853:β2
1848:β1
1794:Ca
1781:Ca
1768:Ca
1748:Ca
1735:Ca
1700:Ca
1598::
1523:.
1515:.
1505:21
1503:.
1480:.
1466:.
1462:.
1439:.
1429:.
1419:17
1417:.
1413:.
1391:,
1383:,
1353:,
1343:,
1312:.
1302:.
1292:19
1290:.
1286:.
1274:^
1260:.
1250:.
1240:.
1230:86
1228:.
1224:.
1201:.
1191:61
1189:.
1166:.
1158:.
1150:.
1138:.
1115:.
1107:.
1099:.
1087:.
1075:^
1061:.
1053:.
1041:.
1018:.
1010:.
1000:31
998:.
975:.
967:.
959:.
947:.
924:.
914:.
906:.
894:.
890:.
867:.
859:.
851:.
839:.
816:.
806:.
796:15
794:.
790:.
778:^
764:.
754:.
744:67
742:.
738:.
715:.
703:97
701:.
697:.
664:.
652:^
638:.
630:.
618:.
600:^
586:.
574:26
572:.
568:.
539:.
527:^
422:pH
312:.
224:.
3581::
3561:3
3556:2
3551:1
3539:9
3534:8
3529:7
3524:6
3519:5
3514:4
3509:3
3504:2
3499:1
3494:0
3472:O
3470:2
3468:H
3455:6
3450:5
3445:4
3440:3
3435:2
3430:1
3418:2
3413:1
3396:3
3391:2
3386:1
3374:8
3369:7
3364:6
3359:5
3354:4
3349:3
3344:2
3339:1
3327:7
3322:6
3317:5
3307:4
3302:3
3297:2
3292:1
3283:)
3281:1
3279:(
3266::
3264:M
3251:4
3246:3
3241:2
3236:1
3217:3
3212:2
3207:1
3202:β
3195:4
3190:3
3185:2
3180:1
3175:α
3164::
3162:M
3140::
3138:H
3103:6
3098:5
3093:4
3088:3
3083:2
3078:1
3056:7
3051:6
3046:5
3041:4
3036:3
3031:2
3026:1
3007:4
3002:3
2997:2
2992:1
2980:2
2975:1
2942::
2897:3
2892:2
2887:1
2865:4
2860:3
2855:2
2850:1
2838:3
2833:2
2828:1
2823:β
2821:v
2819:K
2698:v
2696:K
2555:v
2553:K
2498:9
2493:7
2488:6
2483:5
2478:4
2473:3
2468:2
2463:1
2405:K
2375:K
2373:/
2331:K
2323:K
2260:K
2148:4
2143:3
2138:2
2133:1
2128:β
2126:v
2067:α
2065:v
2043:4
2038:3
2033:2
2028:1
2002:δ
1997:γ
1992:β
1987:α
1936:2
1931:1
1919:4
1914:3
1909:2
1904:1
1836:2
1831:1
1824:2
1822:α
1798:α
1796:v
1792:/
1783:v
1779:/
1770:v
1766:/
1752:α
1750:v
1746:/
1737:v
1733:/
1704:α
1702:v
1698:/
1676:3
1671:2
1666:1
1654:3
1649:2
1644:1
1606:)
1602:(
1588:e
1581:t
1574:v
1531:.
1511::
1488:.
1474::
1447:.
1425::
1379::
1339::
1320:.
1298::
1268:.
1244::
1236::
1209:.
1197::
1174:.
1154::
1146::
1123:.
1103::
1095::
1069:.
1049::
1026:.
1006::
983:.
963::
955::
932:.
910::
902::
875:.
855::
847::
824:.
810::
802::
772:.
750::
723:.
709::
682:.
646:.
626::
620:4
594:.
580::
553:.
498:.
486:,
480:,
474:,
468:,
462:,
456:,
78:8
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