A.T. KamKnu W.C. Kucenesa Atnac NO OU3ZMONOTUM Tom 1 MocksaYALK 612(084.121) BBK 28.707.3461 Ks Hadano npu dunarcosoi noddepacxe Dedepanenozo deenmemea no nevamu u Maccostim KONMYKUKAUUAM 6 PaNKaX Pedeparsuoli uereeoti npoepanmes «Kyaemypa Poccuu>. Penewsenris: aaneayioMlinit KacbenpoHt¢puaHonOTHH WenoReKA H xIROTHX GHlONorHeCKOrO ¢pakynurera TOY BIO MEY wv. M.B. Jlomovocosa, nporp., -p Guon. Hayk Kamencxwii Anapeit Anexcanmponiy; gapeayioulni Kaceapoh dapmakoaorun TOY BIO PIMY Pocsapasa, npod., a-p Mea. Hayk Koanos Mean Tenpuxonus. Kama A. Knceneea H. K18 —Arnae no guovorori. B syn Tomax. [Tow I]: Ye6. noco6we] / A. Kawi, H.C. Kicenesa — M. : TOTAP- Meawa, 2010. —408.c.: un. ISBN 978-5-9704- 1596-2 (o6u1.) 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TEX MAM MHBIX cbyHKUNI Opranusnca, conposoxsaiores Taka W Hx onNcanheN "Arne npeavaahaven CTy2eHTaM MenHUMMCKHX W CHONOTHECKHX 9y200H bakyn4TETOB, nertonawaTenRM, acnpanTaM, ayn paGornunaw. On 6yner aXe NoneNeH BPaaM H OPAHHATOPAM, KeNaIOUHN NOBRCHT CAO KBAANUKALID. YAK 612(084.121) BBK 28.707.3861 Mpasa na dannoe usdanue npunadaexcam OOO Hsdamesocrax zpynna «I'9OTAP-Meduar. Bocnpoursedenue u pacnpocmpanenue « KaKoM 6s mo nu Oe.10 eude wacmu wu yeno20 UsdaHUR He Mozymi Guimb OcyuecmaxeHe Ge NucbMeNHoz0 paspewenus OOO Hsdameneckan epynna «FIOTAP-Meduar. © Kanewn A.T, Kntcenena H.C., 2008 © 000 Mianatensckas rpynna «P'30TAP-Meamay, 2010 ISBN 978-5-9704-1596-2 (o61.) © 000 Manatenscxan rpynna «30 TAP-Menuar, ISBN 978-5-9704-1290-9 (r. 1) opopmenite, 2010Copepykanne Camcox a65pesnaryp. Orasropos. Tams: fens fessor sostpmct reas Jinn >Xupae KuerOTEL Hefitpansinue HPA. octpornnepwas Sinrnss wen pan. Cémmrosnsamewentive nanins Creponas SMTA GHCAOR Mnewxnocrs nunino Benxi mewOpan Horrerpanunte wewpanniue Gens mone veMGpans Tpavenopr vepes wenGpany.. Nat-kanan, Tlaanowerpiveckan oprannauitn, 26 Nat-xanat, OGnemnne wonenst sarees ae Moen paSotw Na*-xanauna Ca?*-Kana. MnasoMerpveckas oprannsauns. Kantuwessie ToxW, o Monekynapras opraHwsauna kanbunesoro KaMama.....34 K’-kanan, Mpocteniee erpoenme ii Totenusazynpanisenme K*-xanasy (Ky)... 38 Cemetter¥0 Ky ncn sce dO K’-kanamt ¢ nays neTARMH 8 HOMCHE (tw0-P potassium channels — Kp) Ca?-axrusnpyenne K’-xaniani (Ke, Mexannani WoHHOM ceneKTHBHOCTH.. Cencop nanpaxeni ch-xanan Mexatioynpasniewsse Katia.ts Bonwit kann oriute nacoces $8 Co -AT@ana.. amar moi 0 Kammunesuit wacoc » muuuax 60 Kansuneauit Hacoc a naasmaTieckolt MeMGpAHE 60 PaGora Ca®*-AT@aau Ha MOneXY/IAPHOM YpOBHE .. Crema paGores Ca?*-AT@ ABB nee see Na/K-ATOasa 66 68 10 ‘Cena paGores Nat/K*-AT ast. OGwenHuKH, rpantcnoprepy Ko-rpaHcnopTepH SnekrpHeckute cnoicTAa KAETKHL Conporusnenne semGparn KneTKH.. Euxocts mem6pants Flocroninian spenenn mem6pans Moctostnas a1mmes MeMOpanet Tlorettunaa rioxos 4 ero MexaniHaMes Morenuuaay knerku, nosmuKaioune mpi ‘aenoxapisauitu ee MeMGpanui, a MpHHUUT NeKyecTBCHHOFO CMeMieHHs HOTeHUMAA Maccusnui siextpotommueck it notenunam, 90 Mexanitam naceusioro aexrporounueckoro Meroa gukeaumi Tox: Jloxanpitnii ovper. Tlorenunan aevicrsus ‘Tums 6wosnexrpueckoH axruBnoctH HEPBHBIX KACTOK esrrnne nnn 9K Zeticrsne sexrpuseckoro ToKa. 100 Mexativam noreHtnama 2eRCTBHR nnn VOD @asosise uaneHeHUA BOSBYAKMOETH nen 104 Meroa buxcauns norenunaaa... Patch-clamp, Cpasnenme meronos Monnwe T0Ku. Perucrpauws Wout ToKOS MeTosoM dbuKcauWH NOTEHUWAE - f ol Na*- K*- nposoausocte, 116 ‘Tox npx pasaux crynespxax norenumaa us Ocnosnute Gnoxatopss Na*- w K*-ToKos TTX u T9A... 120 Na*-TOKH sn iol 122 ‘Axrunautn w wnaxtueauiin Na"-TOKO®. 124 NQ*-KOHON osc ennmsninesnninnnnsennsnnnee 126 Tapawetpst Na*-ToKa.... sniseisD Mecta casauieani roxcnHon y Na’-Kanaa, 130 Uewtp comssinanicn | y Na*-Kavana, 132 Uerstp conssinaniin 2 y Na*-KaHama on 134 esp canssinanu 3 y Nat-xavana . 136 Uewrp constinani: wecrinix anecteritkos y Na*-Kanana.. aa 138 -TOK Ht €FO XApAKTEPHCTHKI. 42 wana L=THNA W NRHA snonnennnnns MA) janet R-vana w T-runa. 146 Baoxaropet m axtusaropti Ca’*-Kananos... 4s. Mwenenne Ca*-roxos non aeiicraue Ca” Moneats Ca?*-nopw.. rox w kana. Xapaxrepuctixu K’-10K08. K*-kaviamul sanepeannoro suinpawtenns. 158 Tpansuropmste K*-Kananes : 158 K,-Kanannt 160 Ca?*-axrusupyemue KT-Kavaet N62 Kanaa yreax — Kp. 164 Morenunanus aelicrons # TORN, Wx PopMapYOUINE MexaHoceHcuTipHaie noTeHuueants.. MexanocencrHpntie TOK Mutenusiusitposannoe 8on0xH0 Usaninosexas Knerxa Pacnpocrpanienue no axcony naccuaioro anekTpoTOHHMNeckoro noTeHuMaaa.. Pacnpocrpaienne noremusana aeRereHsMexanutam pacnpocrpavienns orenuitaa neHcraus ‘No HewHenHHu3HpoBAHHOMy BOTOKH Mexanitam pacnpoctpaHenus norenunana neficrBHs NO MueTHHIESHPOBAHHOMY BOROKHY. 182 Tiposenesine Box6yxxacxHA so nByM Tita BOROKON... 184 Buexnerowne norenumarts 186 3aKoMM MpODENEHMA BOIGYAIEHHA enn sone 188, 180 Caana 2, Bropwansie MeCCeMMEPE 191 oui npescransenin onyancutnansnon TPAMEAYKUMH on 192 Menpannie peusnropus, ennaaiinie ¢ G-Genxan. 194 Fereporprmepibie G-6enKH pasa 96) Monioeprine G-Genkit 198 Mexanusm paSora rereporpuwepnarx G-Geaxor.....200 Myr criaasMon rpancaykumn vepes rereporpumepunie G-6eaxn. snes 202 Tyr axtusaunu suytpuKnerounsix npoTennxntas... 208 WAMO-sanvcuvaa npoTenkunasa (nporewkinasa A) McemanKue c Hel curHanbnee nyTH 206 Foponaneias peryaauna axrupiocrn aneniaarunxaasss Se Mporenxnnasa Cw ceasanntie c Hell cHrHanbHBie ayn 210 @ocdhonnnasy: 4 camsaniiwe CHUM cHTHAABHBE /AYTH Ma IpHMEpe apaxHACHOBOM KHCHOTH nen 22 Kaspwoayaun: erpoenne u yHKu.. 24 Peuentopui ¢ co6cr8eMHoit depMeraTHsHOMt axruanocrovo (xaTansruveckwe pewentopi) 216 Peuciirop-cessanube THPOIMN-npoTeMHKNMASA ‘ua npusepe peuentopos muTepipepona 218 Ras-nionoGibte monomepnare G-Genxu ‘onlocpeaoBaH ble MMH TIYTM TPAHEAYKUWH ness 220 Perynauna rpalicKpnmunit GestkoM, ssanMoaeiicTEyIOUlNM cUAM@-aanucumbin anementoM JTHK (CREB)... 222 Tapa 3. Crnancs... 204 nexrpHveckoe ssanoeficreHte KAeTOK: ea aeKTpHYecktt CHHanC.. sree CCIPYRTYPA KOHHEKCOHOB. non nt Tipununn mexkaerounoro anextpoTonHyecxor ssanwomeficraHa. 29 Fipwatunnts o6napyamehinn dleneRoro KOIFTAKTS sn 231 Pout weaevoro KonTaKra 8 npoBeseHHn BoxGyxxaeH WH aTKanin 231 Meron: vsy¥enns wienenbix KoNTaXToR, son 233 ‘Tpancnopr seutecrs 4epes uleneBon KoNTaKT. 235 Snexipuveckuit cnnanc 1 235 Pont wienevoro Konraxra e cepa 235 Siurawa-peuerrropHoe saanwoneficrene (ocuionntie onpeneness). 236 Ockostioil, wan optocrepuuecknil yaacrox cananinannn 237 Xanawweexuit ewnane. Conepmanne Axconamaatiit tpaycnopr Mexanivau abicaoGoxaeHnn TpaHcunrrepa, Tipeo6pasosanne Tpancmurtepa. “Tunas xuamueckux cunancos. ‘Tunei tpaneaurTepos H MexaHiramis Hx aeitersH ‘Tune aueriuxoantosHisx cHtancos onotponnitti winxorinoput NOMMHEPTHMECKMI CHHAMC. nner 254 Tayrasarauait eaane, 256 ®oedopuanposanne st sedecdopuanronane B FAYTAMATHOM CHHAMEE inns 258 MexanwaM aoarospemennoit notenumauin OATOBPENEHHOH CHMATITHYECKOH MENDECCH nn 260 Peuentropi NMDA ut AMPA onpezeastor paGory raytawaTHoro cnnanca... 1262 TAMK ax rpanemurtep. 264 anu xax tpancaurtep, 266 NO kax tpanenutrrep, 268 Booby xaaioulni nocrcnwantHvecxi noveHuMa (BIC) nesting ncn 210 Topsosnoft nocrennantusecknit noTeHuNan crnicn).. 270 Cumarriveckite HoMHKIe TOKH B yexOBUAX ‘bwxcaumm norenunasa m Buauaiioe annavne BIICT w TTICT 24 JIrania-ynpanasenie wonnwe Kanan. Mx rtoxn. 276 Paich-clamp an vayenin Kantanbnax TOxOE CHHAMER ones snes 218 Coeannenn c ennanriveckun sauanites.. 280 Taapa 4. Duswonorns momui. 283 O6uue npeacrasnenns o ctpoeni pasnwansx MBE 284 Tums wv aa iicaain 286 Ckenernive semis! 286 TRARKHE MELLIUB rnrnnnnnn 1 287 Mua cepaua. 288 uawonorvia CKEAETHLX MELLIL serennennnnennn 290 Moutexyta AK THM 292 Muosun. Capnonaaawarneckwt perMxyAys. SneKtpomexaitveckoe conpaxeHHe. Coxpantenite Met... Moneb CKONKIAMLHX HITE nnn Creu iveckne Genkn cxeneTHoH MaiuitBe Monexyna axtina H MMOSHHE Boamoneficrswe axruna u mwosHa. Mexannsm coxpauienus Ozwnoutoe coxpauieHMe MMU ‘Bynes seswewntsx coxpawennit Coornoutenue mexay uarpy3Kolt u cxopocriio YyRopovenus Coornouenne mexay naMHoft saMuiist wee nanpaxeuttem.. 310 310Conepwanne ynxunonatssas pons AT # npouecce coxpauienis ckeneTHOM MENU. ‘Tune ponokon exeneTHtsx Miu. Hanpaxenwe sesuiisa Onopio-awurarensiwit annapar.. Hepatto-Meiutenoe coeannennt ‘Bnekrpoexaniiecxoe conpaxeHne @yaMONOTMS! EMAAKHX MIE Yuwrraprive rraakwe Meaus Mynwriynirrapnise roankne MBiUiub. i Torenustants raaakHx MMI... z 326 Verownuxu nocrynnenia kansuiin 6 unTornasMy.... 328 CapkonnaawarnueckiA peTHKysyM FRQAKH MALU 5s 328 Coxpauienus enaakiex watt 330 AxriBaus| MonepeNHx MOCTHKOB... 332 Orem rostosHoro wosra Opranisauns cnunnoro mosra Crpoctite HeApOWa seen 342 Com sir : 342 Alenapurss AKcot. Opraneaau neiipona . 344 ‘Tunis weapouos. Buu Henenponiitux kneTOK.... Tunw neHipornun. ‘Crewa pacnpocrpanenin wuiopmauwt no wempony... $0 Bxyconute peuenropt... 354 ipoctpancrsennoe pacnpenenenne H MMHepBaWiEH BKYCORBIX TONEK. 354 Bkycouas now: : 356 Ueierpansibie axyconnte nyTH. 358 O6onsHHe: OsonsTeasuue peuentops UcrTpaasnise oGoustensnete nyTH Crpoene rnasa ‘Onriieckaa cuctesa rna3a, ‘Oprant cayxa, Hapyxtioe yxo Cpeanee yx0... Buyrpennee yxo. Koprutes opran. 7 MipeoGpanonaxine (rpaneayxuin) anya Tlepudbepusieckuit oraen secriGyanpHoit cucreMs... Ctpocame sectu6yaxpHoro naGupunTa. Munepsaums cencopuoro snurenns ectu6ynapHoro anmapara Mpeoparonaune ranenyxunn) neraSyaxpiaix curnanos. Tlonyxpyanste Kanan. Mutuesiise peuenrropet pacts xe1s Crpoenne 1 QyHxtnn smulewnoro neperena Mutmiewtie peurenropss pacraxenis Cyxoxnaseiit annapar Ponsa. ‘ynxunonuponanie mmulewnurx BepeTen. Muorarneckni peexe, 1a pecbaexe a pacraxenue. @aanvecxni pedpexe wa pacramenie. ‘Tonuueckui pedexe wa pacraxenie y-Morouenpoins u pecbnexcu na pactaxenue. OGparuast muorarHueckwA pedprexc... CruGatensnue pediaexces (Cuyenariseckwit oraen nerevaTHsKoh NepBHoi cuctenss i Tlapacumnarnveckuit oraen wererarusnas HepBHon cuctemes Bererarnanan HepsHas cucrema 360 360 32 364 370 372 32 372 4 316 378 380 382 388 388 400 402Cnucok a66pesuatyp AN@® — — anenosnnampocdar AKTD —aapenoxoptxorponusili ropMon AM® — anenosntmonodoctar AT@ ——anenosmnrpucpocdar AUX —aueraaxonun BIICT —sos6yxaaiouimh nocreumantHveckHit notenunan BIICT — sox6yxaaioutHh nocrenHantHYeckHA TOK TAMK —y-anmomacaanan kucaoTa Ti — —ryanosnnamocdar TT® —— ryanosnutpuboctar HK — —sesoxcnpnGonyknennosaa kucnoTa JIT —zoremrsnpytounit ropwor MAUX —MyckapHHosiit — aueTHAxoamHoBEsit (peuentop um cHtarc) MCK — Mexaitocericutnpttate nonitBe Karta WALX — HukoTHHOButi auleTHaXOnMHODEIT (peuerrtop mam cHManc) TIKA —nporenuxnnasa A TIKC —nporemtxitiasa C NKG —nporennxinasa G PHK — —puGonykaemsopas kueaora CIP —capxorinasmariuecknit perukyaym TNICK — topmosHoh noctenmantuyeckui notes una TTX ——tetponoroKcnn T9A —TeTpasTHTaMMoHHi CF — onauxynctamyanpyiounnsi ropmon uAM® — unkanyeckiit AMD ulM® — unknnvecksi FMD UHC —uentpansiias Hepsias cuctema SNP —skaonmasmatHyeckHit peTHKyAyM AC (aconitine) — akoHuTHK ANP (Atrial Natriuretic Peptide) npeacepanwii Harpuitypetawecksit nentHa, AMPA — a-amitHo-3-rnapoKen-5-merHn-4-130K- ca30N-MlponnoHonas KUCKOTA ATX ——ToKcHHb MopcKHx aHeMoH BIX (batrachotoxin) — GarpaxotoKcun CaM —creunduuecknii KanewonysnH CBP (CREB-binding protein) — CREB-censuivarounit nporenn CFTR (cystic fibrosis transmembrane conductance regulator) — pasHonnanocts —xopHoro kavasa CRE (CAMP response element) — yuactox LHK, ssaumoaeicrayrouni ¢ CREB CREB — 6exox, caastisaiouniticn c CRE CTX (ciguatoxin) ~ wryaroxcun DAG —anauuarnnuepon DPPX —annenTiana ammnonenrinasa-nono6H sit potent GAP — [Teasa-axrnaupyioune GenkH GNRF — ryannanyxaeorna06mennusii nporeus GNRP — ryanuu-HykacorHa-ocsoGoxnaioune Genk GRB2_— growth factor receptor—bound protein-2 GTX (grayanotoxin) — rpaaworoKcuH IP, nnositron-1.4,5-rpadocdar JAK — — Janus THposi-nporenHKHHasH LTD (long-term depression) — aonrospemenHas saenpecens MAP — suttoreH-aktusnpyiouunii nporent MAPK — xackaa MAP-kuHasii MAPKK (MEK) — kunasa MAP-KuHa36i NGF — qaxrop pocra nepsos NMDA — N-wetita-D-acnaprar PPI (phosphoprotein phosphatase 1) — chocdonporenrcbocrbarasa | PBTx (brevetoxin) — GpeneToKen Het RTK — —rupositu-nporenixnnasa SAC (stretch-activated channels) — Kanan, aKTHBMpY- JOULNECA MPH pacTAXKeHMH KIETKH — npeoGpasosarenk curHasa W akTHEATOpOB ‘TpanckpHMuMi STX (saxitaxin) — caxcuroKcu TGF — —¢axrop pocra onyxonn VER (veratridine) — neparpnaun STATOt apropos MpeniaraemMi naulemy BHUMAHiIO «ATIaC nO QwsHonOrHHe npeactasnaeT COBO! 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Lpyruwit cnoBaMH, OCHO- poll 110A HX NOCTPOCHHA ABARETER He KAMUEPMH, a ramucpodociopnan kucnora. Eon B nonoxcHitit 3 y PMtUepHHa HaxXomHTeH OcTaTOK ochopHO! KHC- OTH, TO COOTETCTBYIOUIE COCMHEHME HasHBAI0T hoctbarnawnonoit xHcHOTON. Kpome toro, x 3Tomy dociaty ® pane cnyvaes MpHcoeAMNAeTeH HeGoNbiIaR NoARpHAA HONK pORAHHaA asoTCoNepxaUIas MoeKyaa, HanpHMep, kak y ocdaruanacepnna 1 docdbarnanaxoaMna (pu. 1-25). Oru rpynnet cocraszmor nonapyyie (rnapodnawnyt) oGnacte Ha OnHOM KoHe doc- qhonunuaa. lee apyrue rwapoxcumsnee rpynmst ranuepuna (rpynne | 42) mpencrannawt coboit OcTaTKM XHPHBIX KHCAOT H OGecreuHAaIoT OGpar0- BAHHe HenOAApHOrO (rHspomoGHOro) yuacTka NpOTHBONOTOKHOM KOHILe MONEKY.IH. TakwM o6pasom, MoneKynul ocex ocoranuepn- AoB conepxaT NOARPHYIO ToAOBY H ABA HerOARPHEX yenenonopoansix xBoca. MosToMy OHM HaaKIBaKOTCs aMibunaTiecKHMi, T. €. COBMELaIOMIHMY CBOFICTBA H rapodwsHOCTH, H rHapodo6HocTH. 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HaHECTH KANLAIO PACTBOPEHHEIX B Kakom-1H60 siety¥em pacrropuene ocipomni0n MAM XHPHEX KHeAOT, TO MOCAe pacpencscHMs. HX Momekys1 10 RoAMOH NoBEPXHOCTH M MeMapeHWs pac- ‘TpopiTeaa OGpasyerca MOHOMOseKYARpHAR TneH Ka, Kak ycranosita JIatrwiop, mpi nonHoM HactiuieHiti NOBEPXHOCTHOTO cs10R aNCOPGHPOBAKHLe MoneKY- AbL AMIHLIOB pacnonarawTeR nepreNANKyARPHO K NOBEPXHOCTH ROAM TAKUM O6pa3OM, 4TO B BORY Norpyxacten rvapodwasHan nonapHan rpynna, a HenoaspHas yraesonopontas uen’ HanpaBne- Ha pepTHKaNbHO BRepx. Tako opHeHTHponaHNDlit CHO MoeKyn HastiBa\OT «4acTOKOIOM JISHTMIOpa» (pwc. 1-21). Ecai creknaiyto MMacTMHKy ONYCTHTTE B BOY, Ha OBEPXHOCTH KOTOPOM HaXOAHTCR MOHOMONEKYASPHA uIeHKa AMMH08, TO ITY MMCHKY MOKHO MepeHecTHt Ha ovepxHocts maacTanki. [pH NoBropHENX norpyxe- HMMAX Ha TMACTHHKe POSHHKBIOT GuMoneKYARpHUC naenu (pue. 1-2). - Sochoannnant x mpuxyunax hoprnpowanna nex6partes. A, B— docbonunmaw men6pax npenmywiecrsenno npencrasnens Gocbornnyepunann. B pochormmepnnax ona vs neppwsanax rapoNcnasarx rpynn tanyepana (rpynna 3) 23 iT. €. Yor OcHOBA Me TMMepHH, a PmiEposocopHaA KH a OAMOM KOH (oc Cobo ocrarw usuponannan asorconepmamian MoReKyRa, cocrasnaer nonapiyw (ruaposunsiyw) o6nacr pyri rniyepuna (rpynma 1 M2) npencrasns puduyNpovada He xUPHOR, a HochopHO; ora. K bochary npucoenwaserca nonspxan jeparvaunaranonanusa. 31a rpynna nunuza. Zee ApyrHe THAPOKCHRMSHE woGecnesnsaior neonmpxoro (-yapo)o6Horo) ywacrKa. B— o6osavenwe dochommnwsa. T— nonocnol gocbonumusoe 2 ‘one. A — 6ucnoi docconumunosO6mjan usKonorna BosbyAMMaIx TKAHEH A Poccberuqnnaranonanant Sravonamnn p4ososx sprusX FP) Tunpobositie nuniansbie xe0CTbI Tunpocpunesbie ronoeKnJivnuapi Mem6pan B HavGonsuiem Konuuectse B opranitaMe enoneKa HW XMBOTHEIX BTpeYaIOTeR TaKHE hochormuuepHAL (pue. 1-24), Kak ocdarnanazTanonamnn, Baonon- Henne, pucytcrayioT ocdarnaKacepan, hocdarn- AMAMHOSHTOR H SocarwaKrxoAHH (pHe. 1-3 A). Ont MeraGonHveckit cBsixaMB! 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TIpH rMaponie cébuTo- aMTHI08 OBpasyioTeA OAH MoneKy7a XUPHOMi KHC- JOTEL, OANA MONEKY.1a HEHACHULeHHOTO AMMHOCTIMPTA cqhiitrosiiHa HIM ero HaCKIUIeNHOTO atlazora AMrHA- poctbunrosiia, onka MoneKyza ocdbopHont KHCAOTE ona monexyna cnutpra (X—OH). Favtuepina ean Foamnundi He conepxat. Han6onee pacnpoctparen counroaunia cobunro- meant (pic. 1-3 B), conepxaunit B KavecrBe X-rpyn- nua xonitn, Dns cebwtironenmna, Tak xe KaK WANS BCX ApYFHx cibUHTONHTIAOR, XapaKTepHO TO, ¥TO OAHIM 13 FO YieBONOpOANBIX XBOCTOB CHYAKHT AMMHHAR anit charivieckan Uen’ cibuHrosMHa, @ ApyrHM — sTepH- dbuunponannas KUpHaA KucHIOTA. TakHM oGpas0m, KOH(OpMaLIMH Cc>HHTONHTHAOB ONEHE CXOKH ¢ oc- oranuepiaamn, Tak Kak HX MONeKyA Takxe COAep- >Xar NORSPHYIO FOAOBY HABA HenoAAPHBIX KBOCTA. Yrawxonunnazos rososy womeKynm o6pasy- jor MonspHMe THAPODALHIE yrneAONHNe rpyn- nui, atte scero D-ranaxtosa. OnHaxo 8 ormHuMe oT cobukronuninos, riWKonHnAaB He comepxaT Tnapa 1 Hochopuon Kucroti. Knpocretunm ranKon- TIMAaM OTHOCATCA TAMKOSHTAMAUMATAMUCpHHE OOWapYXCHHBIC B PACTEHHAX M MHKPOOpraHHaMax. Tpyraa rpynna, uepeGposnati, moxer GurTt oTHece- Ha Kak k PAMKOAMTINZaM, TAK HK cHroauTuAaN, MOCKOAbKY 3TH CoeAMHeHHR conepKAT MH caxap, 1 coiroaun. Mostomy ob HastiBaiores Lepe6po3H- Abi, WU PMKOCMHTOTUTHAM. OCOGEKHO BEAHKO conepxaHne uepe6posnnon 8 MeMGpaHax HepBHbIX KACTOK, B MACTHOCTH B MHEAMHOBOR OGOROUKE. OGiapyxenninie B uepeSposumax KUPHHE KHCTOTH weobwunel, Tak Kak conepxaT 24 aToMa yraepona. Yauie pcero scTpeaioTeH HepBOHOBAR, UepeGPOHO- bas M AMTHOUEpHHOBARKKEAOTEL. KMPHOKMCHOTH EH 2bMp cmnrosmHa, He conepxaULMit caxapa, Hasti- sajor epamia, Creponapt Crepowaui pesko oraHyaioren no crpyktype oT MoneKya Apyrnx monknaccos sKTHLAOR. Crepowast — TO MpOMSBORHME neprHapoUMKAoneHTAH-*eHaHT- PeHOBOTO supa, Conepxaulero TPH KOHAEHCHPOBAH- HBX UHKAOTEKCAHOBEIX KOLA, COPAMHEHHLX APYT apyrom. 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Buisi0 npeanonoxeHo, “To s1ekTpHNecKoe noxe nefic- TayeT Ha CeHCOP HANPAxEHHA, KOTOPHIA onpene- AgeT TpaticMeNGpaHHesit NoTeHtMaT. 3aTem ceHCOp HanpsxxeH na HomDKeH NepenaTb sty MHKopMaLIO Ha caMY KAHAABHSWO MOMEKYAY 11181 €€ KOH>OPMALLMOH!- Hot MepecrpoliKH HM COOTSETCTBYIOUIeTO HaMEHEHHA actor OTKPHTHS H 3aKpbrTuM KAKA B 1992. B. Xware npeanonoxia, 4TO HaMeHeHHs KOHMOPMALIKM MPONCKOLAT B pesyabTaTe OGuiero Tnapal epepactpenenesus 3apsiaa B MaKpowoneKyae. o6pa- 3yWOULelt Kaliaa, M BLIpUXaWOTCR B BME OTKPHITHA iam 3axpeTHa opoT KaHaza (pwc. I-10 A). Ipyruan CHOBANH, BOIMOAHOCT OTKPHITHA MH JaKpHTHA Bopor Kanasia KONTpoANpyerca CeHCopom Hanpx- xeHia. Beayae notenunanynpasaaemux Kana- JOB CEHCOP JONKCH BKINOYATL MHOTO sapaxCHHEX Apynn, KOTOpHe aBHrawTcs Mon meficTBMEM 21eKT- puvieckoro non. Heo6xonHMo 3aMerHTb, 470 paGora CeNcopa HaNpAxCHKS Mt BOpOT KAHANa, NOKAZAH- wan Ha pic. I-I0A, npenctapnara co6oi pa6ouyio runoresy B. Xwane, BuICKaSAHHY!O Ha OCHORAHNH SAEKTPOSHIHONOTHYECKOFO HayMEHHI NPOROAKMOE- Tw KaHaa, Paccmarpiipas KaHan ¢ nosMUMiL HAMM Ton Ko ABYX — AKTHBALIMOMHBIX HM MHAKTHBALHOHMEX ~ BOPOT, MOXHO NpencraBHT cilenyioUtyro nocmenoR- TenpnocTe coGuTHi (pic. 1-106). B oTeeT Ha .tero- ASpMBALLKIO MeMBpaNts NPONCXOART HaMEHEHIL KOH- POpMaUNH KaHaIBHOM MONeKYAM: omHORpeMcHO NaNWHalOT CMCULATECR EKTHBALMOKHBIE 1 MHAKTHBA LMOHHBIe BopOTA, HO C paaHOR CKOPOCTHO M B Pa MHBIX HaNpaBAeHMAX: AKTHBALMOMHBIE BOPOTA CTPEMAITCR OTKPAITS KAHAN, @ MHAKTHBALMOMHMe — 3aKPBITS. Ckopocts cMewienia 4acritu, oGpasyroulnx arnault ‘ontibIe BopOTA, Go:TbWe, H 3TO MIPHBOANT K OTKPHITIO Kanama (Nepexon M3 COCTOAKHA OKOR B COcToRHHe aKTHBatLIN). B9T0 BpeMs Hepes KAHAN MIPOXOAAT HOH Harpns. Cnenyioutee 3a 3THM KaMeHeHMe TpaHeMeMG- PaltHoro noreHumata ripHBOAHT K 3aKPBITHO HHaKTH- alLHORHBIX BOPOT. ITO MepeBOAMT KAHAN M3 AKTHBH- POBaHHOTO COCTORHHA B HHAKTHEHPOBAHHOE. 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FloBBIUICHHE IutTosORBHOM KOHIICH- rpatinw Ca?” puisbiBaer pasaMuHste KieTOUHEE peak- eH, HANPHMeP COKPaUIeHHE, ceKpeUMtIO, BEIGPOC Meawaropa, npoueccht TpaNcKPH NUM. Ca?*-Kanasb npenctannsiT coGoi KonteKc Ges- Kop, oBpasoanntsit 13 a,-cyOueanNHEe H BCMOMO- Fareabntix a,8-, B- u y-cySveannnu. Cy6venunnua a hopuupyer npovonsiuyw nopy, ofa conepxnT ceHcop HanpAXeHHA HM annapat BopoT KaHana. Tleraa mexay TpaHcmemOpanniimn cerentantn SS 1H S6 xaxOM MOMeHe OMpenenseT CeneKTHBHOCTE M npovonumocts Kanara. CenexTHBHbiit banetp Ca?*-Kanasa 202keH yaHaTb HOH Ca?* yoke Ha BXOIE 8 Katia. 9TH COBRITHA OCTaTONHO pesKH NO cpas: HeHINO ¢ BxONAMM HOHOB Na*, KOAMMECTHO KOTOPHIX nO BHleK.neTOYHON cpeme NPKMEPHO B 100 pas Goss ue. Xora Horm Ca’ 4 Na" aMeior wneHTHsHeH awamerp (2 Ao), KaHan Moxer BbiGHpaTs Ca?” B Gonsuied crenenit, vem Na*, BcooTHOUIENMH 1000:1 Hitkakoe cnTo He MoxeT TaK s¢bbeKTHBHO aNdbe- PeHulMpovarh Mone HAeHTHNHOFO pasMepa. Ckopec cero, Mopa conepxNT crenuuYecKoe MecTO, Tnasa 1 Koropoe cnocoGo cexstiBare Ca” mpi ero vel HIGKNX KOHWEHTPALMAX 8 pactaope (10 °M), a BCE ApyrHe dSHONOTHNECKME HOHE CBAIMNACT B Hamnoro Gonee BSICOKHX KOHUEHTpALLHSX Ca*-Katrants Gem KsIaccHM>HLIMpORAHEA Ha OCHO ne anys npuntunnon: © xuMuveckiti cHNBON raaBHOrO Hoa, 12 KOTO- oro OH NpOHHUaeMBI (B MAHHOM cya HOHE a?*y; © npmitusin dbyanonornuccKoii peryasiut puGors KaHaNa, HMER B BAY. NTO 9TH KAMA NlOTEHU- anynpasanembie (voltage gated calcium channels) Ca?*-Kanaiel, TO MoIOKTO ABECTH OGoINAYe- Hue CaV (pute. 1-11 B). Tlepnas umbpa nocne oGosHavenna cooT- percrayet B Cayo, cyGeeAnHMue HOMepy reHHO- Fo noncemeficrsa (or | 203 8 HacToRuIee BpEMs), 4 BTopan uMMhpa, KOTOpaN cTABHTEA ocne TONKH, coorBercTByeT NopaAky OTKPHITHA OpHTHHanb: HOR WIOhopMN a, CYOLERMHMLM BKYTPH >TOFO noacemeitetsa (or | 0m). CoorseretsenHo ato Homenkaatype, Cay! nonceneftetso (oT Cayl.1 10 Cayl.4) wkalowaeT KaHate, COREPKAUME Cys. cies ip M Gp, AK KOTOPKIX XapaKTepen L-THMT KaMb- unesbix ToKos. Moacemelictso Cay2 (oT Cay2.1 no Cay2.3) BkMONaeT KaHATM, CONepKAMINe Ui). yg M Gye, ANA KOTOpHX xapakTepen P/Q-THN, N-rni R-Titn KasbulneBeix TOKOS COOTBETCTBCHHO. Mloacemertctso Ca,3 (or Ca,3.1 10 Ca,3.3) BknvoNaeT KaHanbl, COMEPAALINE Cic,. Oy, H Gy, AAR KOTOPIIX xapakrepen T-THM KambitHenbix TOKOB Pxe. 1-11, Tnawomerpuwecnan Monens KoneKynapuok opransayyn norexynanynpasanenoro Ca*'-Kawana A— ocvonnan a,-cy6rennxna, Sopnupyioman nopy. OcwoREMe CXpyXTYpH c,-cyBRERNKHIK NoKazaHN Kak pancmen6pavuile RAAMMApH, MpeRcTARAADUME COBOH c-cnypansuste cermeuTus, B— Mogens euinonvewa Mt ocHose 2xcnepHMeNTaMbHLIX NaNNUX KAYNEHNR CTPYKTYpM C22-vaKANOB, COCTORMME M3 MATH Cy6BERKHML Baonoamenne x c,-cyGsennunue MpencraeneNa AovaaWsouaKKan BxyTpHKAeTONNO f-cy6 © NOKeNWaR BNEKNETONNO c,-CYONERMNMNA, COPRANENHAR RUCYALGURKOM CRRIND C f-cyGRERMNUNEH ex Macuira6 qnuna muni MpwEnnsuTenbxo cooTeercreyer AAKHe Monunentwsnesx cermentos. Ywacrk ean © dy-cyOnenmonue MeMAY NexorOpEHN Ca’-raKanaMN MENSPAK sev pacrONOMeKN B NH,-»2%°-80 ‘acti » swapopo6wx cermenrax DIS6 w DIVS3 v » nnwxepe (upronnasnarwecxox CHRaY OWEN YMACTXe) owenam IIL. AID (ay-interaction domain) — nepswsnan oGnacrs censuisann nna scex Caya,-°yoe Ca,p-cySremoonann, Haseannan ax a,-ssanNoneicrayouit ROMeN. CuMBorOm «Po O60NaNen MECTa (> Gopuanponannun paavannn mporenatknsasann. B— nocnenosarensHoe CKORETBO cH ‘mpasngennx Ca*-xaKanoB.TponeMoNCTpApoeano NnOreMeTHNECKOE REPEEO MEPEHHNK "0 CaP -waxanop. B peaynotate cpanneua nap nocnezoparenxocreh werv0 AnbdepeNtOBaNi TPM cevcl-92 Nencenelicreenitunn nocnexosarensioctann, wRextHgnunposaritint Goree we 80% (Cay, Ca,2, C33). ne ‘oRC780 NOCREAOEATENBHOCTH GHNO ONPEAENENO ABR KANROFO CEMENCTEA, H 3TH TPM MOCHERORATEMSHOCEA panes oma ‘enero MOeMeROBareRAMOCTAMM, wReNTH>MIUAPORANNIN Ka 52% 3 no epanieninn €Ca1 nn e C2) pyro 2) wa 28% ( no cpankexoOfmyan d>usxonornn Bos6yqunnx rKanen A LVA ® Hassanne kanana [—— Cat Cal2 ro aus Licata capt ri caz2 L_caz3 cast cas2 L_ casa 2 © 40 60 80 = 100 NpouerT mpenTu4HOCTH aMMHOKMCNOT U Ue Crapoe Tuntoxa~Homep agBaHne reva kanana, y wenoeeKa ais L CACNAIS, ic re CACNAIC 1D L CACNAID o1F L CACNAIF ota P,Q CACNAIA o1B N CACNAIB ote R CACNATE aG T CACNAIG 1H T CACNATH old T CACNAII y enosena 1931-32 12p13.33 spi4.3 Xp11.23 19913 9934 1925-31 17922 16p13.2 2eqie—Ze: Kanbynespie ToKK Ca" -Kanaasi Gut oGHapyxcHE npaxTHecKH 0 BCex KeTKaX. Ca?*-TOKM, 3aperncTpHpopaNHe 3 pASMMAHBIX THNIAX KiIETOK, MMCIOT onpeneneHHe isHoTOrMMeCKHe H dapwaKonorHYeckMe cBOiic- ‘pa, VaHavanbHas GykneHHan HOMCHKAaTypa Obina Npeatoxena HcxonA M3 KUHETHK Ca?*-ToKoB. L-THn (or +long-lasting» — noaro naawnitea) Ca?*-roKoB TpeGyeT CHABHOM AenoAApHIAUNM AR AKTHBALHH, of ONTO ANAMUMMCA H GnoKMpyeres opranHueckH- Mu attTaronmcramn L-THna Ca?*-KaHaston, BKTO4aR AMTMApOMMpHANHE, eniaTKHsTAMMHD H GeH30- Twasenuite. Ca?*-toxn L-Tuna spasiores rat HIN B MBILILIAX MSHMOKPHHHBIX KMETKAX, THe on HuMuMHpyIoT coKpauieHHe H cexpeuMto. N-THn (ot sneither long nor transients — nn L, xm T), P/Q-tun m R-THN KAABUHEBLIX TOKOB TAKE TPEOY- JOT cHIbHOMI AeNOAAPHZAMMH LAR aKTHBAUMN. OHH OTHOCHTENBHO HeYYACTBHTENBHY K AHTATORMCTAMH L-twna Ca?*-Kanatos, Ho GaoKMpyioTea creundt- HECKHMM TOTHTENTHAHKIMH TOKCHHAMH M3 S108 YAWTKH HAH nayKa, STH TOKH B OcHOBHOM BEIPAKe- Hb B HeipoHax, re OHM HHMUMMPy!oT HeHpoTpan- eMHCCHWO JU GonbinMHcTHa ObICTPKIX CHHANCOR ‘Takxe onocpeaywor Bron Ca®* B KjeTONHELE Tena H aenapuTti. T-THn (oT «transiente— npexonsuiuit) Ca?*-ToKoB axTHAMpyeTCR cna6o!t AEnOnApHSaLtH- efi, MOTH TOK MAMOZeTHBe (npexomsutne). OHH HeYyBcTBHTesbHM K OpraHMECKHM arTarONHCTAN WTOKCHHAM 3Mefi H TayKOB, KOTOPHIE HeMOAIBAyIOTCH aaa onpenenenua N- w P/Q Ca?*-roKos. Ca?*-ToKH T-rnna supaxents & GonbilloM crexTpe KeTONHHIX TUMOB, THe OHM BOBAEYEHE 8 PASBTHE NOTEHUMANA AeliCTONA H BAXNB B KACTKAX M TKANAX, OGAaaI0- LHX pHETMHeCKOM AKTHBHOCTEN. Monekynapnan oprannsayua Kanbuvesoro KaHana Ca?*-Kananbi, KoTOpHe GEITH oXapaKTepHs0BaKE GuOXUMMYECKHMH MeTOnAMH, NpeACTABAAOT COBO! KoMMexe Gerkos, o6pazonansinifl M3 NeTMpeX WAH NATH ONpeTeAeHHEX cyGBEMKHHL, KOAMpyIOUINXCs, Gomusinm ceneficT#OM reHoE. Tnapa 1 Ha wonexynaprom yposnie Ca?*-Kattan cocraanen va opMMpyioulell Nopy TpaHtcMeMGpaHHo! c-cy6e- eAMHMUBI H BeOMOTATETBHEIX 0,8, BH Y--VO8e1 8 (pic, 1-12). Tip 3t0M cy-cySveantinua Hiaxonirres Ha eHeulHeR cropoHe MeMbpantl, B-cyBeeaunnua— Ha BHYTPeHHeR CTopoHe MeMOpartel, a 5-H 7-CyO- eAHHMUKL MpemcTaBAAIOT COGO!L TpareMeMGpanHHe ctpyxtypei. Crpykrypa Ca**-kaniana cepnesHon FaanKoH MBILULIB! — 3T0 CTPYKTYpa C HETHIpENA CYOL- eaMHMUAMH Kanata, H OWA HaniowMaeT Ca?*-KaHa- aM HeApona. BenoMoratenwiwe a,5- m B-cyGvens- HMUB YBeNM4MBAIOT TpaHcnopT MOHOB Ca”* Nepes nopy, o6pasoBantriyi a,-cy6veanHHueA, Ht MoRy- AMpyIOT MOTeMUMaTIABHCHMYIO KHNETHKY KaHaAa, BuytpnkneTownas f-cyGeennkmua H TpaHCMeMGpaH- MBIA aNcymbeua-cBasaHHtAM KoMNteKc a,5-cyGeea- MMU SAAIOTCA KOMTOMEHTAMM GOABIIMACTRA THTOR Ca?*-kananos. Batononvenne K pont B-cy6xens- MulUM B TpaHCHopTe HOHOB MepeS KaHa, OHA TAKA MIpaeT MPHHUNTHAABHYW poms B aTeKTpOMEXAHH- MecKOM CONPAXEHHH B MBILICYHLIX BONOKHAX cke- sneribtx most, Bsansonesicrans cyGReaMyints a,8- B-cy6veanHnusi B HacTomuice BpemA XOpOUO H3y4C- vos. Hamnine cyGveannuuss y Obi10 TaKxe OOHAPY- xeno » Ca?*-Kananax ckeneTHLIX MBILILL, B Kapsit- OMMOUHTAX H HeRPOHAX ronoBHOTO Mosra. XoTH 3TH noGaRowHbIe cYGREAMNHUM MORYAUPYIOT cHofiCTHA MoHHoro KaHana, apMaKonorM4eckHe H eK TPOdH- awonoruveckHe ocoGexHoctH Ca?*-KaHasi08 8 OCHO HOM cBsizaHihtc cYULeCTBOAHHEM « -cyBReMMH IL, B-cyGreannuus: Ca?*-Kanaso8 croco6ws ¢bopmn- ovate rereporennuse KoMmiekcti in vivo Min viro P-cyGeeamHMuM n a,-cyOsennHnua oGnanawT KOH CepBaTHBHBIMH OBNIACTAMH BIAMMONERCTBHS, “TO CHo- coGcrayeT*bopwnpovanimioreTeporeHHMXKOMIINEKCOB Kaviana, o,5-cyOvenmniuist Monyaupy.oT pasnHunie oy cyGbenKHMUL fn vitro, M_BOSMOKHO, 4TO 37a reTe- PoreHHocTs pacnpocrpaliaeTca Ha cHTYAMLKH in vivo. Taxoe pashoo6pasne ssanmoneiictena cyOvennnntt NOTEHUMATDHO YBEANYHLAO Gbt YHCTO BOSMOKHEIX BapHaHTOB KaHaT08 ¢ OTANAIOULHMHEA GHOdUSHEC- KHMG Ht DHSHOTOTHYECKHMH CBOIiCTBAMM, HTO MOKET obecrewpare pasHoobpaame K:eTONNEIX OTBETOB. @usnonornyeckaa Moness MoTeHuvastynpaBaseMo- ro Na*-xanaza noxasana wa pcyHke (pitc. I-12 B) Puc. 1-12. O6nemnan mogen» ssanMonencrsnit cyGveqnnn norexinanynpasnaeMore Ca*-nanana. A— y-cyGnemmamya ws06paxena xax TpancnenGpaxnii Genow C YeTMDAMR CermeNTaMK ¢ BKYTDIKRETON N-C-xonstan, Nepean ronoswna y-cyGrenMnwun Bianmoneicrayer c a,-cyBrexvnnnen. llepean sexnerowIian TeTnA ConepyM? 3apAMeNNME OCTATKH M YHACTKH FnNKosMAMPoBA;THA, Onpegenciis e2annonencray‘) c 7H a,5- W B-cySveaunnua (B-Boannoneficrsyounk nome) ¢ o,-cybsenuinuen. B— MaKOROTHYECKAR KOR jovenumanyrpasnaemoro Na*-rawana, Toxazaiia oCHOBNAR «,-cYGEeRMKMUa OPMMPYIWAR NOPY M B ‘eneumte a, :, Yu BecyorenvexnogheOswjan dbusnonorns sos6yqumerx rKaHen A o,-cySvennnnua 5 Buexnetoutas cpena (out) Tpacmemopannnin wHTerpanbHbii GenoK, Popmupyiouywn Kanan a, -cyObenMHnua ,-cySvenviiua Cenextuerii cnn S-cySvenmnnua yeyOvenmmaya BeeyOnennanya Benok \ Vinaxrueaynonneie umTockenera Bopora Arrapaynonntie eopora BryTpukneTounasl KMAKOCTE (in)K*-kanan. Mpocteiwee crpoeHue TpwHatan 8 Hacromttee BpeMa KraccMbuKaNMA aenut ace K*-Kaitazbi ra: ‘© notenunanynpannsembie K*-KaHtanes (Ky); = Ca?*- axrionpyemste K*-Kawtartt (Ke); © K*-kanaml anoManbHoro abinpaMAEHMA c ToKOM sxoaMuero HanpannertnNs (inward rectifier K-channels) (K,,); * K’-kaaant ¢ fleyMa netasMH 8 noMeHE (two-P K’-channels) (Kp). Onucatine MozekyaspHoit oprannsaunn K*-Kana- OB Mb HABE ¢ HaNGoee MpOCTOH dope, Mpes- crapenHof, HanpuMep, 8 GaKreput Streptomyces lividans, Moaen Monekyaapnoi oprantusainn TaKOTO HanGoutee npoctoro K*-Kanana Gaxtepin Streptomyces lividans nponemonetpuposatia wa pic. 1-13. Hloxasannetit wa pac. 1-13 A K'-Kawtaa Gaxrepii CcoctonT H3 YeTbIpex oAHHaKoBLX cyOvEAKHHU, O6b- emMHeHHBIX B eANHOE UeZOE Tes10 KAHAN. Ha nianerax B, Bm F noKasaHo, To P-netas 10K: ANQOBAHA OKONO BHEKMETONHO! NOBEPXHOCTH MeMG- pane! HW oGwenMseT a-cnupann OGoHx cermerton SS WS6. Ona cocrouT Ha HecnpanbHolt YacTH, bopMix- Pyloutelt BepxHto10 acre Nope, KOPOTKOMl a-crMpamH W yanMHmoULelt MeTAH, KOTOpAR BBLAAeTCA B Cy>Ka- foulyiocs acts nope H Gopmipyer CeneKTHBHEI (ussrp. BTOT qMALTP nosBoALeT NpoxoAHTE Hepes Kaitaa Tanbxo woHaM K*, Ho He apyrHM MOHAN. Hixe cbbrpa HaXoMMTCH eHTpAbMad NacTe NopEl, O6pa- sosanHas a-cnupanbio cermettta S6. BTeTpamepHoM K*-kanate Gaxtepun Streptomyces lividans cermentta SS 11 $6 oGosiiayentt Tak, NOCKOAEKY OHM NpeAcTABAROT coGoii cTpyKTypHte 3kBHBATEHTH CerMeHTO SS it S6 B nloTenunaaynpassxemom K*-Kantaze (pic. 1-13). Puc. 1-13. Mogens monexynapwom oprannsauym nanGonee A— stor K’-wanan npencrasnser c ve cxopiite tpaxcmen6pakkwe a-cnupan, 06031 nopy. B etna P (posoeas). B wa ceepry rerpamepkan opranusainn HMGAUMA ORHOH va WeTNIpeX cyEveR Tnasa 1 Ha pucynixe 1-13 B nokasatiet crpyktypHiie eT an kanaaa. NH,-koweu cyOveanniusi Hanne HapyxxHoft crpasisio (opanxenax), por iieit MeMGpany oT uNTOMAASMaTHHECKOH croc JO HapyxHOH nosepxHocTH. 3a HapyXHO! © pansio caeayeT KopoTkas cnupans (kpacnas vanpasjteHHas 8 nopy. 3areM cnesyeT BAYTpE crMpanb, KoTOpaR BosBpAULaeTCA K UMTOTIANA: ueckoii cTopone (opanxesas). CoeaMHsioutas tes Mexay Hapy KHOM H KOPOTKOMl CMPAAAMH Op CTPYKTYPY. GOPMUPYIOUIY! HapyKHOE OTBEpL ops. Ira WacTs COMEPAHT LEHTP CBAIKIBAN HS HeceneKTHBHoro G1oxaTopa K*-KaHanon rer atuzamMonus (TOA). YeTsipe TaKHX MeTAH Ko of cyGveanHnuE PopMUPYIOT BHEUIHHE CTEM)! TOPs, a MMEHHO TOM ee HACTH, KOTOpas OTHE THeHHA 38 CENEKTHBHOCTS KaHasia — CeneKTHDH: cbuawrtp. Vs6nparensocth Kanata nocTHraeTe# Kak pas MepoM ero nlopss, Tak M MosIeKy:1PHOM OpraHMsaLiH elt ceneKTHBHOTO dbuaBTpa. JLMaMeTp CeeKTHRHO uasrpa pave npumepHo 0.3 HM. AmMHOKH i ef CTeHKM OpHeNTHpORAHM Tak, TO 1 TeAbHUte KObUA, OGpazoBAH HIE NETLIPLMA CHNbHEIMH rpynnaxH (NO OMMOH OF KAXZOH eaMHituBt), OBpautenbi BHYTPS Nopes. JwaMeTp AOCTATONEH M14 NPOXOXIEHNS woHia K’, MMetoutero anaMeTp 0,27 HM. ATOMB! i Jopoa B CTeHKe NopEs 3aMeHRIOT FHAPATALIO V0 Hons Menbitiero paswepa, Hanpumep Na’, ¢ 0 PON 0,19 HM, He MoryT npoxonHTe Hepes KT=\.) NOCKOABKY OHM HE MoryT cchopmupoBaTe CBA3b OHOBPEMEHHO CO BCEMIK HETMPEMS! AMM CTEHKH Mopbi. ITOT MEXARHAM B MeTAARX "pe! crapnen Ha pic. 1-19, mpocroro K’-kaxana Gaxrepuu Streptomyces lividans. i Terpamep ua MAENTHNHTTK CYBRERMHUN, KAMAAR 1 KOTOPHK CONepMUT awaemate xax $5 1 SB, 4 Xo} 0 u 55 $6 Ky P-nerne, GopHy| wpann (x pieer 4yvanynpaBanemple K*-KaHanbt Ky K, kananet o6pasonanet GenkoBKinn cyOueamin- aM aByx THNOB: a-cyGveaHHALANM, GOPMHPyIO- LIM Nlopy, H HcrioMoraressHtiMn B-cySBeaMiMa- Mu. Cy6ueaninuts a opmupyior TerpaMep, "robb oGpasowaTb dbyHKUMOHAABHBIM KaHan, npoBonAMIN” wont K*. Onna cySnemmuuua Ky-Katiana — Genox ¢ wecTaI0 TpaHcmemGpankinn cermenTamm (S) a-cnupann (SI-S6) 1 pacnonoxeHHeimn 8 uMTORAAMe N- 4 C-repmunaannin, HanGomee Hay 4eHbI BHEKCTONHaR netas P mexaay 5S u 6S (bopmupyer K*-cenexTHituti euneTp mM nopy Kawlana) HW 4S, KoTOpHIM NoTOKH- TenbHO JapRxeH M SOpMHPYET CeHCOP HANpAKCHHA Geaka (pc. I-14 A). Kpome Toro, mHoro dynkuni o6siuo npuTicerBaan Gonbusm N- w C-KoHttEBEIM neTaam. OHH BKAIOYAIOT KOHTpOA 3a HHAKTHBALME! KaHaton, GesloK-Genkosoe pxaMmostelicraMe (no THNY TeTpaMepHsaunn a-cySveamnt, BsaMMoeltcTBHE C uitronzasmaTH4eckHMM B-cyGuennniuann W coxpa- HeHMe nocaesoBarenbHocTeit). Ky a-cyGvenninust woryt opmposare wnK Fomorerpameptt (homotetramers), naw reTepoTeTpa- mepei (helerotetramers), neportHo, orpanuseniinie B TOM xe caMOM mlonceMelicTBe. OrpoMHoe pasHo- oGpasme Ky-ToKo in vivo ects caecTBHe cyutec- TeOBAHWA CHAaMCHHTOBEX BapHanTos (H30>opM Tetla, KOnMpyIOULMX OAHY M TY XE a-cyOBEAHHMUY), cGopkit TeTpaMepa c-cyOLenMHMLL ¢ pasaMuHenn Tnaza 1 BcHOMOraTeABHEIMH cyGEenMAMUAMH M reTepoMmepK- sauna a-cy6vennniu. Kak wspectHo. Hons: Kt HeakBHBareHTHO pac- Tpenenensi MEXity BHEKMETONHOM cpesoit H LMTON- na3Mof, M 9TO cosMacT MBIKYUIYIO CHAY ATA suixona HoHOS K* 8 Gonbwx cbusMonornuecKH npenenax Mem6pakitioro notenumana. Ky-kaHans OTKPHITH! IPH slenopHaLIMH TpaHCMeMGpaHHoro HoTeHuMama, TO crlocoGcrByeT BUXOAY HOHOB K’. Jicnonspysauns moxet Gute npeacrasneHa KaK AKKYMYARUMA MOSHTHBHBIX 3apA20B Ha BHEUIHER cTOpoHe MeMGpanti. OnHaKo Bbixoa nosHTHBHO JapRAXCHHBIX HOHOB, TAKHX, KAK HOHE K’, cMMTaeT MeMOpanituil noTeHuNan HiKKE NOTEHUMATa NOKOR. Taxi oGpasom, Ky-KaHanus 4yBcraytot jenonspi- 3aUMWo H, B CBO! ONepeRb, eHICTEYIOT TAK, HTOBHI Ce yetpanurs. Ontia m3 HanGonee oxapaKTepHsoBal- Hbtx pore Ky-Katanon 3aknW0¥aeTCR B TOM, “TOG 3AKOHMITE CHAbHYIO AeMOAAPHIALMO, BUIBAHHYD AKTHBALMEM NOTEHUMANIABHCHMOTO BXOTA KATHO- Hos (coamecTHo oBpasytoutnx noTeHtiMan neftcTBHA). ‘Takum o6pa30M, MOxHO MOCTYAMpoRATS, HTO MOBWE yeennuenua Ky-kaHarbHoit axTHBHOcTH Gysyt NpuBonHTh K Goxee abexTHBHOMy OKOHYAHM AENOARPHAUMK (OGMHHO YKOpANKBaR HAM yOH- pas noreHumat aciicrena). Sta yHKunonannsan cnocoGHocTs Ky-KaHan08 TaKKE TEKNT B OcHOBE opmuposanina naxewvolt axTuBHocTH (B3peiBHO!). ‘Taku o6pasoM, akTHBHocTe Ky-KaHasoe Hrpaer, HAMPHMep, Pont B KOAMPOBAHMM MIKpOPMALMK B HepsHofi cucreme. Puc. 1-14. TnaMomerpwyeckan Monen> MOReKYRAPMOK opraNusauMM noTeMuManympasnaeMoro K’-Kawana (Ky-vamana). A— onwa cy6sennnuna Ky-nanana mpegcrasnaer co6oi a-crupans ¢ mrecrhio TpaNcMeMOpanMusmn CerMenTaM (61-86) pacronomeximn » mironnaane N- « C-x $5 u $6 dopxnpyer cenexrusich gumrp axx nowos K° x nop\ ‘4 gopmupyer cexcop sianipamevis Cenka. Zammnue N- ¥ C-Tepmuxannitae *e anamu, Brexnerouian P-netna nemay cermenrann awanov v Genox-Genkonce Bsannoneficrene, 6 — accouwauma NeTAspex cy6remuny # Ky-KawanOsman dusMonorua BoaGyqumerx TKANenTI Cemeicteo K, 2lo HacTosutero BpemeHi 8 reHOMe YesIOBeKA On cane 38 revios, KoaMpyIOUINX pastintEtC KantaTbs cymepceneficrsa Ky. Ky-KaHasst BkuOaIOT KstaC- enveckne KareropHn Katiatos: K'-Katiani samep- >XxaHHoro BbinpaMEHHs (delayed rectifier K*-channels) i GticTpHe TpaHsMTOpHBIe K'-KaHaNe BHIXORAULEO TToxa (fast transient K*-channels: fast transient K*-current I, uau transient outward current 1) Haswanne «delayed rectifier” K-channels» 610 neppotaansHo nano moTeHuHanynpanaaemem K'- KaWianaM THraltTcKoro akcona KanbMapa 3-34 HX oTcpoveHHolt aKTHBAUKH (no CpaBHeHKIo ¢ GuicTpO axTHensHpytounimuca Na*-Kavatamn). Went Bcex K, moncemestcrs (ekno¥as Kyl—4, EAG « KCNQ) Moryt (bopmuposart delayed rectifier K’-channels Fast transient K*-channels (hopmupytouine fast transient K-current: I,, WaW. transient outward current 1.) — Suictpte TpansiTopHete K*-KaHanb BuIXORA: lero ToKa npeacTasnaOT COGOR KaHambi, aKTHBIH- PyeMble HHSKHM noTeHLMaTOM (low vollage-activated), OkH Guictpo uHaKTHEMPyIoUIMEcs (rpaKaHTOpHLIe, MHave — BpemeHHbie, muMoneTHNe K*-KaHatt). Kavanu sroro Tana o6u4HO OGpaxonaHt 13 4eHOD Pe. 1-15, Tnasal Kyl # KA noaceneiicrs 1 scnomoratenstoit B~y0 ENKEI, 4ACTO HEOBXORUMOM ALI eHOMeHa O1iC ofl HHaKTHBALHM, HeaBHo Ost1a OGHApY ENA Hs Bcnomorarensian cyGeeammituia — CD26, peoicrver Has OUMENTHAMA aMMHONeNTHAasaNOLOGHOMY p< tenny (Dipeptidyl Aminopeptidase-noaoGuisit Gea. DPPX), # 3T0 nosBonMAO MpHCBONTE Katty He Ipc Hop xapaxtepnctuky K4-KaHias08 Punorenerueckue aepenbs Ky-Kananon npose- MoncTpHpopatiit Ha pic. I-15. Ha Hem npH TeHETHMECKOM PeKOHCTPYKUMH, OCHOBAHHOM Va CXORHOM AMHHOKHCTOTHOM NocneROBATeAbHOCTH, Nokasanbl noreHunaaynpasraemue (voltagega') K*-xanants cemeficts Kyl—K\6, Ky7 a K\8—K,9. Cpean 9roit rpynmsi TombKo KyI.8 B Hactos- lilee BPENR MenBITBIBZeT HeMOCTATOK B HasBaNHM HGNC. Mate wnenos rpynnm Ky cemefictua Ky7 (KCNQI-KCNQS) 8 Hactosuiee Bpems (C2005 1.) MOryT GHITS BHECEHIL B COWS C APYTHMM GeaKa- Mit Katiana Ky, # MosToMy He MoKasaKB! # OTAe- AIbHOM ILTOTeHETHECKOM epeRE, KAK ITO Mpes- cTapaaaoct patie. Tp ocTapuinec ceMeiicTea Ky — Ky10, Ky Il # Ky12 — nocrarouno 6ansKo cas- 3aHM pyr ¢ ApYrOM, 4TOGH GAT NOKASaHHLMH B ommentitom busioreneTHYecKoM nepene. mnoreneTHweckne nepesea ana cemencrua Ky, A— dunorenernvecnoe nepeno ana cenelicrea Ky1-K,9-xananos. Amnouucnorawe nocnenoBarenbxocrK K,ramana exmovaior Kyi-Ky6 « K,B-K,9 cemeiic Moctw K,71-K,75, K6-4 « K,B.2 Gn nobaei rugpogobrie apa xaana (S1-S6) uenonss08ann: K,10-K,12-nawanoe “Tua, KoTOpwe Geiny awecen 10 2005 r. B — nocnenosarens HL X cymecrayIMeNy GunorenerwvecKoKy nepeny. Tonbko ce na axiannaa. B— dnorenerwiecnoe Repeno Ana cenedicraO6mjan @usnonorun woabyaumnrx rKaKel A ;— K5.1 [KCNF1, 2p25) E421 IKON, 20013] K,2.2 [KCNB2, 8q12] \\ 8.1 [KCNG1, 20913), K,6.2 [KONG2, 18922] '— k 6.3 [KCNGS, 2421] [-K.8.1 IKCNBS, 8922) | -— KQ.3 [KCNSS, 2p24) [—K.9.2 [KCNS2, 8922) '— k 9.1 [KCNS1, 20912] K34 [KCNC4,1p21) K,3.2 IKCNC2, 19913) K,3.1 [KCNC1,11p14) 23 [KCNCS, 19913) K,4.1 [KCND1, Xp11) K,4.2 [KCND2, 7431) K,43 [KCND3,1p13) —k,1.7 IKCNA7, 19413) K,1.4 [KCNM,11p14) Kv1.6 [KCNA6, 12p13) roc: 5 [KCNAS, 12p13) K,1.8 [KY1.10, 19913) K,1.2 [KCNA2, 1p13) “Teich teen s2pt9) K(1.3 [KCNA3, 1p21) —— K 5.1 [KCNF1, 2p25) a KE! CNB, 20913] K 2.2 [KCNB2, 8q13} [—K.6.3 IKCNGS, 2p21) K.6.1 [KCNGH, 20413] K 6.2 [KCNG2, 18422} _ k 6.4 (KCNG4, 16924) [47 IVT, 1 4p15) K7.3 [KCNQS, 8924] U K7.2 [KCNQ2, 2013} UP: 4 [KCNQS, 1p34] K,7.5 [KCNQS, 6914] [— K,8.1 [KCNV1,8q22] (__k8.2 (KCNV2, 9p24] [—K 9.3 [KCNSS, 2p24) [- K,9.2 IKCNS2, 8q22] L— k 9.1 [KCNS1, 20912] [- K3-4 [KCNC4,1p21] K,3.2 [KCNC2,19q13] ‘Lp Kes [KCNC1, 11p14] K,33 [KCNC3, 19913] K,4.1 [KCND1, Xp11] rLpKee [KCND2, 7431] 4.3 [KCNDS, 1p13] [— K,1.7[KCNAZ, 19913} K 1.4[KCNA4, 11p14] K,1.6 [KCNA6, 12p13] [—K,1.5 [KCNAS, 12p13] L—k 1.8 [Kv1,10, 19913] K,1.2 IKCNA2, 1p13] —K1.1 [KCNAI, 12p13] '~K_1.3 [KONAS, 1p21] [— K,12.2 [EIk-2, KCNHG, 1213] [—K,12.1 [EIk-1, KCNH8, 3p24) L123 [EIk-3, KCNH14, 17421] t—K,10.1 [eag-1, KCNH1, 1432] 1K '10.2 [eag-2, KCNHS, 14423] -—K,11.1 [eag-1, KCNH2, 7435] _k (11.2 [eag-2, KCNH6, 17435) 1k .11.3 [eag-2, KCNH7, 2424)Tpynnt K,, Carey tout katace 25-Genkos pkmiovaer TpH rpynt- nus KaHat08 inward rectifiers channels (K,,), KOTOPBIE MOxkHO HasBaTb, XOTA H ONEHE AAHHKO, Kak «K*-Katia- 2161 ANOMAAABHOTO BBITIPAMZIEH A ¢ TOKOM BxOARHLETO wanpasnenuse, N- 4 C-repmiiiant 9THXx KaHa0B ‘Takxe pacrianoxenst B LuTOMTaIMe, P-neTnA MeXAY ABYMS: TpavicMeNGpannesMst aoMeHamu ¢opsupyer HOpy, H DYHKUMOHHPYLOUNNT Kaan npeacrasaser coGoH Terpamep 3THx 2S/1P-cyGvennHMt Knonnponarine nepnuix invard rectifiers K*-chan- nels— K, 1.1 (ROMK1) 4 K,2.1 UIRK1) — Gus0 ocyuieeTBeHo 8 1993, M MocmenoBaTenbHocT® HOBLIX “ieHOB 2TOrO ceNeiicTBA Guia WZEHTH- quunposana, ekmovan K-Kanan, cesgaHnei ¢ G-Genkom, (K,,3) # AT@-syscrenrenbausil K-xawan (K,,6). DTH KaHaasl MrpawT BaxHyIO ditsHOTOTN- NeCKyto pomb BAYHKUMM MHOFHX opraHton, BKmI0vaA MOST, CepmNte, MONKM, IHOKPHHHME KAETKM, CAY- XOBBIC KTETKH H KAETKH ceTHaTKH. MoaeKyaapHan opranusauna THX KaHatoB mpHeeaena Ha pic, 1-16A. @urorenerHeckoe s1epeno, noKasaH- Hoe Ha put. I-16 B, mamocTpHpyeT pomcTBEHHBIe CBH3H, OCHOBANHIME Ha MocKenOBATETBHOCTH aMit- HOKHCAOTHDIX OCTATKOB, MEXIY CEMbIO OCHOBHBIN [nasa 1 noncemefictsamu K,,. BHacronuice BpeMa HOBHE neHEt 3TOTO ceMefiCTRA He GLITM HAeHTHDMLLKPORA- ist, Tockonsxy 3T0 éitaorenerHeckoe mepeBo nos- BuaOCS TOKO 8 2002 r. 8 NanaHM [UPHAR, mato- BEPORTHO, TO KaKHe-7NGO apyrHe HoBLie “eH GuiaH oGtapyxcHits 34 9TO BpeMA BnocaemHee npema meTon peHtrenoctpyKtyp- Hof KpHcranzorpadun aan BosMoxHocT onN- cars CTPyKTYpy UNTONAAIMaTHYeCKOTO. perHoHs K,3.1, noanyo crpyktypy Gaxrepnanbsix K,,1.L Ht uitronnasMatneckyw oGzacte K,,2.1. StH manne AeMOHCTpUpyIoT, TO inward rectifiers K*-channels MMEIOT ATMHHY!O Mopy, YXOMAUy!O 8 UNTORAASMy, H NOAYEPKMBAIOT BAXHOCTh OTPHLIATEABHBIX SApAOE AMMHOKHCKOTHEX OCTaTKOB Ha CTeHKE WHTONAAs- MATHMECKOA ACTH Opti. KOTOPAR HrpaeT BAKHYK Poab aioManbHoro BLinpAMaeHuA. ITH HecAeno- aHHs OGeCTeNMAM TONMNAHHE MexXaHMaMOB pery- AMpoBaHitA BOpOT KaHaTA NOcpeACTBOM AHTAHAO® ‘uepes G-Genkn mchocdbatnananHosnTon 4,5-Guiboc- ar. Machopmauns, ronysientias 6naronaps avamsy KDHCTasMNeCKOH CTPYKTYPHI, !pesBLIAHIHO MEH Tak Kak MO3BOMET OLIEHUTe CTpyKTYPHO-cbyHk- UMoHarbHue oTHOUHHA. Takxe npHMeNaTenbit HaAAHHS, NOCBSIIEHHHIE HIYNEHMIO AMHAM HECK acekTos cbyHKUuIM KaHa.ta. Puc. 1-16. Monexynapwan oprawnsaunn inwardly rectifying potassium channels— K'-xananov anonansoro ‘BIpAnnewR ¢ TOKOM BKORAIIETO ManpaBAeHNA (K,,). A— onma cyGvenunnna K,-ranana npencrasnset (85 w S6), pacronomeKKUMa B uwronasme Re w C co6of a-cnupant ¢ aByMA xpanicmenGpaxnnnn cermenrant pRunannin, M KOPOTKyO P-TeTIO, PopRUPY ONY B — accoimanus seruper cybseawinal 8 K,-vavan. B— unorenernveckoe nepeno 21s K,KawanonO6man cbusmonorua Bos6yAMMBIx TKaHeEn A K Hassanne Homep kanana Kat K74 KA2 Kaa K 5.1 Ka k24 — K22 L— kos K34 -— 33 Homep eva xpomocome! ysenosexa y YenoseKa KNIT 1724 KCNII3 2997 KCNJIS 21922 KCNJI0 1422 KON16 17423 KONJ2 17423 KCNJ14 19413 KCNI2—17p11 KCN4 ——-22q13 KCN3 ——_2q24 KcN9 tga KONIS ——11q24 KCNIs 2122 KCB 12p11 KONI 11p15K*-kawanbi c ABYMA NeTAAMM B ROMeHE (two-P potassium channels — K,») Menine 4em 3a necaTHAeTHe, HaNHaR ¢ OTKDU- ‘rita Ky»-KaHaOR, Hx HBHEHHE MOKA3A0, ¥TO 6430- Bag yTewKa HoHOB K* Yepes cneuHaaHaHponanHtie nyTH mpeactaBaseT coGOH sqxpexTHBHO peryrMpy- embiit MexaninaM KOHTpOAR BOSGYANMOCTH KMCTOK. Tlyru yreukit MoHoB K*, akTHBHEie B MOKOE, cTaGH- ANSMpYIOT MeMGpaHtiuif MoTeHUMAN Hike Mopora TeHepaLitH B3pEIBHOM (naYeuHo!) akTHBHOCTH yexopaior penoaspysaunw. Xora cyulectsosaKne TOKOR yreuku Gui70 npeaoAOxCHO # 1952 r. Hodgkin W Huxley, on ocrasanics GHodHaMNeckuM Kypbe~ 30M BTeNeHHE NTH AecATHNETHH. MneHTHd>MKALMM NepBUX MOREKYNSPHLX MexAHH3MOB TOKA yTeY- KW Motos Kt cnocoGcrsosano KroHHPoRaHMe K*- KaHaon ¢ aByMa nopodopMupyiownMH meraAN B KaXi0A CYOVERMHHUe H YeTHIPMs TPAHCMeNG- pANHUIMM cerMeHTaMit (HAH BOCEMbIO TpaHeMeMO- pantuimi cermentamn) 8 Saccharomyces cerevisiae w Caenorhabditis elegans. Nlocae 3t0r0 Kp 681 430- AMpowaH M3 HepBHO-MENeYHO! TKAHH Drosophila ‘melanogaster. Buodwanveckne xapakrepucTHKHt TloKasaai, 4TO Kp HO1XCH GMT K*-ceneKTHBHWM kaHasoM ¢ onpenereHHuiMH MapaMeTpaMH MpoBO- AMMOCTM, Te. ¢ OTEHUMAAHesaBHICHMBIM BOPOTHBM MeXaHHGMOM. HM ripenckasaHiim Goldman, Hodgkin, Katz sbinpawsiennem. Kora KOnueHTpauns HoHOB K! cumerpitina c anyx cropon MeMOpanni, Kyp-TOK Taaza 1 HM ReMoHeTpHpyioT aAMHeliHYIO saRMEHMOCTH ov norenumaaa. TIpn dH3HOAOTHYECKHX ycAoNi"»s (@bicoKas KOHLIEHTPALIMA HOHOB K* BHYTDH H 1 Kaa cHapyxn) Kp mpononut Goree sHauHTen rit TOK 13 KACTKM, HEM aHYTPH KAETKH, Ha6nON€ TOTOK HOHOB, HanpaaeHHEsi 8 Gorbuen cree Hapyxy, SEM BHYTDb KETKH. Nopasurenbiax yepta Kp-kananos — opr aii uns wx cyGneamHnusi. Kaxaan ueer ape P ner ertipe S cermentra (pic. I-17 A). ra oranuarouias Tonoaornsi (2P/4S) Moxer 6tir» Haiiaena B Ganee °% 70 npeackasaHMbix roMonOrax 8 Ga3e AAHHBIX, 110" MeHHBIX MPH HByHEHHM reHOMa. MaTHamuAT® re io» MaeKOMTaIOULHKX B CeMeficTHRE OBO3HaNeHBI Kak KCNK, kommpyioune K»-Kanianut (pic. 1-17 B a 8 Tipu axcnpecems nerko oOnapyxwBaeTcs YH woHHoro KaHana. Kak oxiaa0cb, 278 pery ‘BosGYAMMOCTH Kyp-KaHalIbs HAXOANTEH MOM CHT! KOHTPONEM MHOKECTBA XHMHMECKHX H cbHSHMS: cTHMynon, eKMO¥aR HaNpAXxeHHE KHeROpOa, Pp SMM, MEXAHHINECKHE PACTSXEHHS, HC!lpOT)ak CaMTTepAt HM peuenTopsl, cBAIAHHBIE ¢ G-GenKo%! Kananu Taxxe cayxat MOneKYsPHBIMI 9/10/0850 DUI HeKOTOPLIX NETYYHX M_MECTHEX aiiec Peryasuus Kzp-KaHanos CBAIAHA C 1M! B MX cpoficTBAX, HanpMMep BepORTHOCTH OTKP! Ti! Kanaros, Tem He Meee HEKOTOPHIE peri ODN Mk waMeHeHua, Hanpumep doccopenuponanne K, OGecrewnBaeT MOABAEHHE BLITIPAMARIOWHX CBOM CHYBCTAHTeTBHOCTHIO K NoTeHUMAAY. Topol Puc. 1-17. Monexynapwan opraxwaauna ana two-P potassium channels — K’-waManon c AByMA MeTRAMM B AOMEKE (Ka) ‘A —ogna cyGseanmnua K.y-namana mpencrasnsi nn, pacnonoxewuuumn w untonnaame N- u C-repwinva qwaima nByx cy6wenvin 8 K,,-KaHan. B— ob aR Kyy-nananoe O6parure wxunanne, 470 3TOF KAMAN uaiTIOMNAeT YHKUMO TAK MasbIBaeMOrO KaNANA YTEERH. Got a-cimpanb c Yerapbin tpantewenGpantitantn cerventn mn, u ABYKA P-retnANn, GOPMHYIOWKMK TOPY e waobpanenne K,,-nanana. T— punorenerwectoe nepe'O6man dpusnonorua Boabyqumerx TKaHen Pt Hageanne kanana Kp kK 10.1 Kg2t KI74 K, 16.1 K 5.4 K,,6.1 Ky7A Kept ————- k,.9.1 -——— k,31 1 15.1 —— K131 Kpt24 Homep rena y 4enopeKa KCNK4 KCNK10 KONK2 KCNK17 KCNK16 KCNKS KCNK6 KCNK7 KCNK1 KCNKS KCNK3 KCNK15, KCNK13 KCNKI2 xpomocome! ysenosexa 11q13 14931 1941 6p21 6p2t 6p2t 19q13 11q13 1942 8q 24 2p 23 20q 12 14924 2p22=o Ca?*-axTueupyempie K*-KaHanet (K,,) MonexyaspHas oprannsauns K,,-kaHaios npea- crasnena Ha pute. I-18, B. Duzorenetuveckoe DepesO, MpencraweHHOE Ha PHCYHKE, MANWCTpH- pyer akr, 470 3TH KaHanu opMupyior aBe xopo- WO OnpeneAEHHbie, HO TONEKO OTAATEHHO CBAIAH- hie rpynmis. Oana Ha 9THX rpynn akatoYaer TpH Ke,-KaHaa ¢ HM3KOM nponoanMoctbIO (Ke,2.1. Kei2.2 # Ke,2.3), a Take Kava Ke,3.1 ¢ MpOMeXy- ‘TOYO MPoROANMOCTDIO. ITH KaHa.TE He MOTH AIYNPABAXEMM M AKTHBUSMPYIOTCR HHGKMMIL KOH- heTpauHAM BHyTPHKACTOHOTO Ca™* (THM KAMbMORYMHOM MpHBOMAT K KOH- opMALHOHHLIM H3MCHEHHAM, KOTOPEIC, B CBOW ONepens, OTHETCTBEHHE 3a BOPOTHEIH MeXAHHM, xanana, @uoreneruveckoe aepewo, MoKasaHHOe Ha puc. 1-18 B, uaniocrpupyer 3asucMMocts nocaeno- naTersHocreit rpynm! Ke,-KalanoB, KOTOpAR BKMIO- waer Ke, 1.1 (Slo nan Slol), Ke,4.1 (Slack win Slo2.2), Ko42 Glick stan $102.1) # Ke,5.1 (Slo3), Kanianet Kill soywanr 8 KaeTkaX Mo3ra, YOMTKM HM Mbt UAX, M HaBeCTHO, YTO epenyioulMecH crnaiicHn- Tw wx MPHK orsereTseHHnt 3a ux sHaNTenbHBe OyHKUMOHATbHHE pasnnns, Botmtane or Ke,2- Ke,3-Kana7on, cBasbiBanne noHoB Ca?*-KaanoM Puc, 1-18. 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OnHako B MacrOsuliee HpeMs nospuacce Gonbwie MiopMaLHH o dyHKUKOHAAS- Bix cBOHcTRAX “eHOB 9TOFO CeMeNICTBA TeHOR. eM Gvin0 MaveCTHO HeCKOADKO eT Ha3aa, KOFI OTH HaMMeHonaHHs GBIAM NpeaTOxeHD!, H 9TO rIpeR- craRnser BOIMOKHYIO JaMaLy JL CHeUM>UKALH, OcHoBAHHO!, CKOpee, Ha YHKUMOHAsbHOM Cxonc- ‘Tee, tem Ha cTpyktypHom. Borawine oT Haiinen- Huax sieHon Ke,L.l, ak THYECKH AKTHEHPYIOULXCR BHYTpHKeTONNMM Ca”. HH OMA 13 pyrHX WIeHOR TOM FPYANBI, NO-BHANMOMY, He MOKET GuTb TAKE aktusnposan Ca. @axraveckn, riasHeim OGpa- 30M, 9TH TPH HENYACTRHTENBHLIX K BHYTPHKAIETON womy Ca kaiiata K,4.2 4 Ke,4.1, axtnenpyiorcn Bhytpukserountmu Honann Na” 4 Ch, a KSI AKTHBHGMPYETCA BHYTPHKACTONHDIM SauleNasitBa- went (caBMFOM pH B ulenoNHy!o CTOpoRy). Takin 06pasoM, xors OHM CTPYKTYpHO CAAA ¢ Ke, OTH TPH KaHasia He MOryT GWT ONNCAHE MpaBiibHo Kak ¢kaMeDbie KAMAIH1, AKTHBHPYEMME KA,TBILNEN® a octone dy HKUMOHaNbHBIX KpHTEpHEB. ITO MOXET OnITs TeMoMt 19 OGcyxaeHHA cpeaM NecneMOBATeNE! aanNoit OGnacTH. 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