ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ, 2009, ÚÓÏ 26, ‹ 1, Ò.

64–74

ìÑä 577.3

çÂÒÂÎÂÍÚË‚Ì˚ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ ͇̇Î˚

‚Ó ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II
© 2009 „. ê. Ä. êÓχÌÓ‚, ç. Ç. 䇷‡ÌÓ‚‡, ë. ã. å‡ÎÍËÌ, ë. ë. äÓÎÂÒÌËÍÓ‚
àÌÒÚËÚÛÚ ·ËÓÙËÁËÍË ÍÎÂÚÍË êÄç, 142290, åÓÒÍÓ‚Ò͇fl Ó·Î., èÛ˘ËÌÓ, àÌÒÚËÚÛÚÒ͇fl, 3;
Ù‡ÍÒ: 8-(4967)-33-0509; ˝ÎÂÍÚÓÌ̇fl ÔÓ˜Ú‡: cheglok@rambler.ru
èÓÒÚÛÔË· ‚ ‰‡ÍˆË˛ 06.10.2008 „.

Ç Ò‡Ï˚ı ‡ÁÌÓÓ·‡ÁÌ˚ı ÍÎÂÚ͇ı ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ (èá) ‚˚ıÓ‰fl˘Ë ÚÓÍË ÔÂËÏÛ˘ÂÒÚ‚ÂÌÌÓ ÔÂ-

ÂÌÓÒflÚÒfl ËÓ̇ÏË ä+. Ç ÚÓ Ê ‚ÂÏfl ‚Ó ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II èá ‚˚ıÓ‰fl˘Ë ÚÓÍË ÒÓı‡Ìfl˛ÚÒfl
‚ ÛÒÎÓ‚Ëflı, ‚ ÍÓÚÓ˚ı K+-ÒÂÎÂÍÚË‚Ì˚ ͇̇Î˚ ËÌ„Ë·ËÓ‚‡Ì˚. Ç ‰‡ÌÌÓÈ ‡·ÓÚ Ï˚ ËÒÒΉӂ‡ÎË
ËÓÌÌ˚ ͇̇Î˚, Ó·ÂÒÔ˜˂‡˛˘Ë èá ‚˚ıÓ‰fl˘Ë ÚÓÍË Ò ÌÂÓ·˚˜Ì˚ÏË Ò‚ÓÈÒÚ‚‡ÏË, ‚Ó ‚ÍÛÒÓ‚˚ı
ÍÎÂÚ͇ı ÚËÔ‡ II. Å˚ÎÓ ÛÒÚ‡ÌÓ‚ÎÂÌÓ, ˜ÚÓ ˝ÚË Í‡Ì‡Î˚ ı‡‡ÍÚÂËÁÛ˛ÚÒfl ÌËÁÍÓÈ ÒÂÎÂÍÚË‚ÌÓÒÚ¸˛ Ë ÔÓ-
Ìˈ‡ÂÏ˚ ‰Îfl ÍÛÔÌ˚ı ÏÓÎÂÍÛÎ, Ú‡ÍËı Í‡Í NMDG, „βÍÓÌ‡Ú Ë Äíê. ëӄ·ÒÌÓ ÓˆÂÌ͇Ï, ҉·ÌÌ˚Ï
Ò ÔÓÏÓ˘¸˛ ÌÂÒÚ‡ˆËÓ̇ÌÓ„Ó ÙÎÛÍÚÛ‡ˆËÓÌÌÓ„Ó ‡Ì‡ÎËÁ‡, ÔÓ‚Ó‰ËÏÓÒÚ¸ Ó‰ËÌÓ˜ÌÓ„Ó Í‡Ì‡Î‡ ÒÓÒÚ‡‚-
ÎflÂÚ ÔÓfl‰Í‡ 200 ÔëÏ. èÓÎÛ˜ÂÌÌ˚ ‰‡ÌÌ˚ ҂ˉÂÚÂθÒÚ‚Û˛Ú Ó ÚÓÏ, ˜ÚÓ ÔÓ Ò‚ÓËÏ Ò‚ÓÈÒÚ‚‡Ï ËÒÒÎÂ-
‰Ó‚‡‚¯ËÂÒfl ËÓÌÌ˚ ͇̇Î˚ ·ÓΠ‚ÒÂ„Ó ÒıÓ‰Ì˚ Ò ÔÓÎÛÔÓ‡ÏË, ÙÓÏËÛÂÏ˚ÏË ‚ ̇ÛÊÌÓÈ ÏÂÏ·‡-
Ì ÍÎÂÚÓÍ ÔÓÁ‚ÓÌÓ˜Ì˚ı ·ÂÎ͇ÏË ˘Â΂˚ı ÍÓÌÚ‡ÍÚÓ‚ – ÍÓÌÌÂÍÒË̇ÏË.

èÓÔÛÎflˆËfl ÍÎÂÚÓÍ ‚ÍÛÒÓ‚ÓÈ ÔÓ˜ÍË ÏÎÂÍÓÔËÚ‡-
˛˘Ëı „ÂÚÂÓ„ÂÌ̇ Ë ‚Íβ˜‡ÂÚ ˜ÂÚ˚ ÚËÔ‡ ÍÎÂ-
ÚÓÍ, ˉÂÌÚËÙˈËÓ‚‡ÌÌ˚ı ̇ ÓÒÌÓ‚Â ÏÓÙÓÎÓ„Ë-
˜ÂÒÍËı ÍËÚÂË‚. íË ÚËÔ‡ ÍÎÂÚÓÍ – ÚËÔ I, ÚËÔ II Ë
ÚËÔ III – ËÏÂ˛Ú ‚ÂÂÚÂÌÓÓ·‡ÁÌÛ˛ ÙÓÏÛ Ë ‚˚ÔÓÎ-
Ìfl˛Ú ˆÂÔÚÓÌÛ˛, ÔÓ‰‰ÂÊË‚‡˛˘Û˛ Ë ÒÂÍÂÚÓ-
ÌÛ˛ ÙÛÌ͈ËË [1–3]. éÍÛ„Î˚ ÍÎÂÚÍË ÚËÔ‡ IV, Ú‡Í
̇Á˚‚‡ÂÏ˚ ·‡Á‡Î¸Ì˚ ÍÎÂÚÍË, ‡ÒÒχÚË‚‡˛ÚÒfl
‚ ͇˜ÂÒÚ‚Â ÍÎÂÚÓÍ-Ô‰¯ÂÒÚ‚ÂÌÌËÍÓ‚, ‰ËÙÙÂÂÌ-
ˆËÛ˛˘ËıÒfl ‚Ó ‚ÍÛÒÓ‚˚ ÍÎÂÚÍË ÔÂ‚˚ı ÚÂı ÚË-
ÔÓ‚ [3]. ÇÂÏfl ÊËÁÌË ÍÎÂÚÓÍ ÚËÔ‡ IV ‚Ó ‚ÍÛÒÓ‚ÓÈ
ÔӘ͠‚ Ò‰ÌÂÏ ÒÓÒÚ‡‚ÎflÂÚ ÔËÏÂÌÓ 10 ‰ÌÂÈ [4,
5]. èÓÔÛÎflˆËfl ‚ÂÂÚÂÌÓÓ·‡ÁÌ˚ı ‚ÍÛÒÓ‚˚ı ÍÎÂÚÓÍ
„ÂÚÂÓ„ÂÌ̇ Ì ÚÓθÍÓ ÏÓÙÓÎӄ˘ÂÒÍË – ‚ ÌÂÈ
ÏÓÊÌÓ ‚˚‰ÂÎËÚ¸ ÚË ‡Á΢Ì˚ „ÛÔÔ˚ (ÚËÔ˚ A,
B, C) ÍÎÂÚÓÍ Ì‡ ÓÒÌÓ‚Â Ëı ˝ÎÂÍÚÓÙËÁËÓÎӄ˘Â-
ÒÍËı ı‡‡ÍÚÂËÒÚËÍ [6]. àÒÔÓθÁÛfl ÚÓÚ Ù‡ÍÚ, ˜ÚÓ
˝ÎÂÍÚÓÙËÁËÓÎӄ˘ÂÒÍË ҂ÓÈÒÚ‚‡ ‚ÍÛÒÓ‚˚ı ÍÎÂ-
ÚÓÍ ÒÚÓ„Ó ÍÓÂÎËÛ˛Ú Ò ÔÓÙËÎÂÏ ˝ÍÒÔÂÒÒËË
ÓÔ‰ÂÎÂÌÌ˚ı χÍÂÌ˚ı ·ÂÎÍÓ‚ [7, 8], ̇ÏË ·˚ÎÓ
ÛÒÚ‡ÌÓ‚ÎÂÌÓ ÒÎÂ‰Û˛˘Â ÒÓÓÚ‚ÂÚÒÚ‚ËÂ: ÚËÔ Ä = ÚËÔ II,
ÚËÔ B = ÚËÔ III, ÚËÔ ë = ÚËÔ I [8].
ÇÓ ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II Ë ÚËÔ‡ III ÙÛÌ͈Ë-
ÓÌËÛ˛Ú ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ (èá) Na+-͇̇Î˚
[6–8], Ë ÔÓ˝ÚÓÏÛ ÓÌË ÒÔÓÒÓ·Ì˚ „ÂÌÂËÓ‚‡Ú¸ ÔÓ-
ÚÂ̈ˇÎ˚ ‰ÂÈÒÚ‚Ëfl [8–10]. é·˚˜ÌÓ ÂÔÓÎflËÁ‡ˆËfl
˝ÎÂÍÚ˘ÂÒÍË ‚ÓÁ·Û‰ËÏ˚ı ÍÎÂÚÓÍ ÔÓËÒıÓ‰ËÚ Á‡
Ò˜ÂÚ ‡ÍÚË‚‡ˆËË èá K+-͇̇ÎÓ‚ ÌÂÒÍÓθÍËı ÚËÔÓ‚,
Ë ÔÓ˝ÚÓÏÛ ‚ Ò‡Ï˚ı ‡ÁÌÓÓ·‡ÁÌ˚ı ÍÎÂÚ͇ı èá ‚˚-
ıÓ‰fl˘Ë ÚÓÍË ÔÂËÏÛ˘ÂÒÚ‚ÂÌÌÓ Ó·ÂÒÔ˜˂‡˛ÚÒfl
ËÓ̇ÏË ä+. Ç ÚÓ Ê ‚ÂÏfl ‰Îfl ÍÎÂÚÓÍ ÚËÔ‡ II ı‡‡Í-
ÚÂÌÓ Ì‡Î˘Ë èá ‚˚ıÓ‰fl˘Ëı ÚÓÍÓ‚ ‰Ó ÌÂÒÍÓθ-
ÍËı ̇ÌÓ‡ÏÔÂ, ÍÓÚÓ˚ ÒÓı‡Ìfl˛ÚÒfl ‚ ÛÒÎÓ‚Ëflı, ‚
ÍÓÚÓ˚ı K+-ÚÓÍË Ó·˚˜ÌÓ ËÌ„Ë·ËÓ‚‡Ì˚, ̇ÔË-
ÏÂ, ÔË ‰Ë‡ÎËÁ ÍÎÂÚÓÍ ‡ÒÚ‚ÓÓÏ, ÒÓ‰Âʇ˘ËÏ
CsCl ‚ÏÂÒÚÓ KCl. èÓ˝ÚÓÏÛ ˆÂθ˛ ‰‡ÌÌÓÈ ‡·ÓÚ˚
·˚ÎÓ Óı‡‡ÍÚÂËÁÓ‚‡Ú¸ ËÓÌÌ˚ ͇̇Î˚, ÍÓÚÓ˚Â
‚Ó ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II Ú‡ÌÒÔÓÚËÛ˛Ú èá
‚˚ıÓ‰fl˘Ë ÚÓÍË Ò ÌÂÓ·˚˜Ì˚ÏË Ò‚ÓÈÒÚ‚‡ÏË.
ùäëèÖêàåÖçíÄãúçÄü óÄëíú
ùÍÒÔÂËÏÂÌÚ˚ ÔÓ‚Ó‰ËÎË Ì‡ Ó‰ËÌÓ˜Ì˚ı ‚ÍÛÒÓ-
‚˚ı ÍÎÂÚ͇ı, ËÁÓÎËÓ‚‡ÌÌ˚ı ËÁ ÊÂÎÓ·Ó‚‡ÚÓ„Ó Ë
ÎËÒÚӂˉÌÓ„Ó ÒÓÒÓ˜ÍÓ‚ flÁ˚͇ Ï˚¯Ë ‚ ‚ÓÁ‡ÒÚ ÓÚ
6 ‰Ó 12 ̉Âθ. ÇÍÛÒÓ‚˚ ÍÎÂÚÍË ‚˚‰ÂÎflÎË ËÁ ‚ÍÛ-
ÒÓ‚˚ı ÒÓÒÓ˜ÍÓ‚, Í‡Í ÓÔËÒ‡ÌÓ ‡Ì [11].
ùÎÂÍÚ˘ÂÒ͇fl ‡ÍÚË‚ÌÓÒÚ¸ ‚ÍÛÒÓ‚˚ı ÍÎÂÚÓÍ Â-
„ËÒÚËÓ‚‡Î‡Ò¸ ÏÂÚÓ‰ÓÏ patch clamp ‚ ÍÓÌÙË„Û‡-
ˆËË whole-cell Ë perforated patch clamp Ò ËÒÔÓθÁÓ‚‡-
ÌËÂÏ ÛÒËÎËÚÂÎfl Axopatch 200 B, ñÄè-Äñè ÍÓÌ-
‚ÂÚÂ‡ DigiData 1322A Ë Ô‡ÍÂÚ‡ ÎˈÂÌÁËÓÌÌ˚ı
ÔÓ„‡ÏÏ pClamp8.0 (‚Ò “Axon Instruments”,
ëòÄ). ëËÒÚÂχ ÔÂÙÛÁËË Ó·ÂÒÔ˜˂‡Î‡ ÒÏÂÌÛ
‚ÌÂÍÎÂÚÓ˜ÌÓ„Ó ‡ÒÚ‚Ó‡ ÒÓ ÒÍÓÓÒÚ¸˛ 0.1–1 ÏÎ/Ò
[12]. ê„ËÒÚËÛ˛˘Ë ÔËÔÂÚÍË Á‡ÔÓÎÌflÎË ‡ÒÚ‚Ó-
‡ÏË, ÒÓ‰Âʇ˘ËÏË (‚ Ïå): 140 KCl/CsCl, 1 MgCl2,
0.5 EGTA, 10 HEPES-KOH (CsOH), pH 7.4. Ç ÒÎÛ˜‡Â
ËÒÔÓθÁÓ‚‡ÌËfl ÏÂÚÓ‰‡ perforated patch clamp Í ÔËÔÂ-
ÚÓ˜ÌÓÏÛ ‡ÒÚ‚ÓÛ ‰Ó·‡‚ÎflÎË 400 ÏÍ„/ÏÎ ‡ÏÙÓÚÂ-
ˈË̇ Å. èË ËÒÔÓθÁÓ‚‡ÌËË ÍÓÌÙË„Û‡ˆËË
whole-cell, Á‡ ËÒÍβ˜ÂÌËÂÏ ÒÔˆˇθÌ˚ı ˝ÍÒÔÂË-
ÏÂÌÚÓ‚, ÔËÔÂÚÓ˜Ì˚È ‡ÒÚ‚Ó ‚ÏÂÒÚÓ 1 Ïå MgCl2
ÒÓ‰ÂʇΠ2 Ïå MgATP. ŇÁÓ‚˚È ‚ÌÂÍÎÂÚÓ˜Ì˚È
‡ÒÚ‚Ó ÒÓ‰ÂʇΠ‚ Ïå: 140 NaCl, 2.5 KCl, 1 CaCl2,
1 MgCl2, 10 HEPES-NaOH, pH 7.4. èÓ ÏÂ ÌÂÓ·ıÓ-
‰ËÏÓÒÚË ‡ÒÚ‚Ó˚ ÏÓ‰ËÙˈËÓ‚‡ÎË ÔÛÚÂÏ ˝Í‚Ë-
ÏÓÎflÌÓÈ Á‡ÏÂÌ˚ ËÓÌÓ‚ Na+ Ë/ËÎË K+ ‰Û„ËÏË Í‡-
ÚËÓ̇ÏË Ë/ËÎË ËÓÌÓ‚ Cl– – ‰Û„ËÏË ‡ÌËÓ̇ÏË. ê‡Ò-
Ú‚Ó 50 Ïå Mg2ATP ÔÓÎÛ˜‡ÎË ÒÎÂ‰Û˛˘ËÏ
Ó·‡ÁÓÏ. ÇÓ‰Ì˚È ‡ÒÚ‚Ó MgATP ‚ ÍÓ̈ÂÌÚ‡ˆËË

64
 çÂÒÂÎÂÍÚË‚Ì˚ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ ͇̇Î˚ 65

‡ ·
50 ÏÇ
–70 ÏÇ 3

àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ
–100 ÏÇ
ÍÓÌÚÓθ 10 Ïå íÖÄ 2

50 ÏÒ 0
KClin /NaClout
1 ÌÄ
–120 –60 0 60
åÂÏ·‡ÌÌ˚È ÔÓÚÂ̈ˇÎ, ÏÇ

‚ „
50 ÏÇ
–70 ÏÇ 2

àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ
–100 ÏÇ
CsClin /NaClout 1

0
25 ÏÒ
–90 –60 –30 0 30 60
1 ÌÄ åÂÏ·‡ÌÌ˚È ÔÓÚÂ̈ˇÎ, ÏÇ

êËÒ. 1. àÌÚ„‡Î¸Ì˚ ÚÓÍË ‚Ó ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II. ‡ – ëÂÏÂÈÒÚ‚Ó ÚÓÍÓ‚, „ËÒÚËÛÂÏ˚ı ÓÚ ‚ÍÛÒÓ‚ÓÈ ÍÎÂÚÍË ‚ ÓÚ-
‚ÂÚ Ì‡ 100 ÏÒ ÔÓÎflËÁ‡ˆË˛ (‚ÂıÌË ԇÌÂÎË) ÓÚ –100 ‰Ó 50 ÏÇ ‚ ÔÒ‚‰ÓÙËÁËÓÎӄ˘ÂÒÍËı ÛÒÎÓ‚Ëflı ‚ ÍÓÌÚÓΠ(΂‡fl ÌËÊ-
Ìflfl Ô‡ÌÂθ) Ë ‚ ÔËÒÛÚÒÚ‚ËË ‚ ̇ÛÊÌÓÏ ‡ÒÚ‚Ó 10 Ïå ÚÂÚ‡˝ÚË·ÏÏÓÌËfl (TEA) (Ô‡‚‡fl ÌËÊÌflfl Ô‡ÌÂθ). ëËÏ‚ÓÎ˚ ̇‰
ÍË‚˚ÏË ÚÓÍÓ‚ Û͇Á˚‚‡˛Ú ÏÓÏÂÌÚ˚ ÒÌflÚËfl ‚ÓθÚ-‡ÏÔÂÌ˚ı (I–V) ı‡‡ÍÚÂËÒÚËÍ. · – ÇÓθÚ-‡ÏÔÂÌ˚ ı‡‡ÍÚÂËÒÚËÍË
ÒÚ‡ˆËÓ̇Ì˚ı ÚÓÍÓ‚, Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ‡. ‚ – ëÂÏÂÈÒÚ‚Ó ÚÓÍÓ‚, „ËÒÚËÛÂÏ˚ı ÓÚ ‚ÍÛÒÓ‚ÓÈ ÍÎÂÚÍË ‚ ÛÒÎÓ‚Ëflı ‰Ë‡ÎËÁ‡
‡ÒÚ‚ÓÓÏ, ÒÓ‰Âʇ˘ËÏ 140 Ïå CsCl. „ – ÇÓθÚ-‡ÏÔÂÌ˚ ı‡‡ÍÚÂËÒÚËÍË ÚÓÍÓ‚, Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ‚. èÛÌÍÚËÌ˚ ÎË-
ÌËË ‚ · Ë „ ÒÓÓÚ‚ÂÚÒÚ‚Û˛Ú ÎËÌËË ÌÛÎÂ‚Ó„Ó ÚÓ͇. ÇÓ ‚ÒÂı ÒÎÛ˜‡flı ËÒÔÓθÁÓ‚‡ÎÒfl ÏÂÚÓ‰ perforated patch.

‚˚¯Â 50 Ïå ÚËÚÓ‚‡ÎË Mg(OH)2 ‰Ó pH 7.4, ÔÓÒÎÂ
˜Â„Ó ‡ÒÚ‚Ó ÙËθÚÓ‚‡ÎË ‰Îfl Û‰‡ÎÂÌËfl ÌÂ‡ÒÚ‚Ó-
ÂÌÌÓ„Ó Mg(OH)2 Ë ‡Á·‡‚ÎflÎË ‚Ó‰ÓÈ Ú‡Í, ˜ÚÓ·˚ ÍÓ-
̘̇fl ÍÓ̈ÂÌÚ‡ˆËfl Mg2ATP ÒÓÒÚ‡‚Îfl· 50 Ïå.
ÇÒ ÒÓÎË, ˘ÂÎÓ˜Ë, EGTA, EDTA, Na2ATP,
MgATP Ë ÙÎÛÓÂÒˆÂËÌ ·˚ÎË ÍÛÔÎÂÌ˚ ‚ “Sigma-Al-
drich” (ëòÄ). ÇÒ ÓÒڇθÌ˚ ‡„ÂÌÚ˚ (1,2-‰ËÓÍ-
Ú‡ÌÓËÎ-sn-„ÎˈÂËÌ (DOG), ‡ÏÙÓÚÂˈËÌ Å,
U73122, SITS, DIDS, ÌËÙÎÛÏÓ‚‡fl ÍËÒÎÓÚ‡, NPPB,
9AC) – ‚ “Calbiochem” (ëòÄ). èÂÔÚˉ 43GAP26
·˚Î ÒËÌÚÂÁËÓ‚‡Ì ‚ ㇷÓ‡ÚÓËË ıËÏËË ÔÂÔÚˉӂ,
îàÅï êÄç, „. èÛ˘ËÌÓ.
êÖáìãúíÄíõ
Ç ÔÒ‚‰ÓÙËÁËÓÎӄ˘ÂÒÍËı ÛÒÎÓ‚Ëflı ([KCl]in =
= 140 Ïå, [NaCl]out = 140 Ïå) ÍÎÂÚÍË ÚËÔ‡ II „ÂÌÂ-
ËÓ‚‡ÎË ‚ ÓÚ‚ÂÚ Ì‡ ‰ÂÔÓÎflËÁ‡ˆË˛ ·˚ÒÚ˚ ‚ıÓ-
‰fl˘Ë ÚÓÍË, ·ÎÓÍËÛÂÏ˚ TTX, Ú.Â. Ó·ÛÒÎÓ‚ÎÂÌ-
Ì˚ ‡ÍÚË‚ÌÓÒÚ¸˛ èá Na+-͇̇ÎÓ‚, ‡ Ú‡ÍÊ ÓÚÌÓÒË-
ÚÂθÌÓ Ï‰ÎÂÌÌÓ ‡ÍÚË‚ËÛ˛˘ËÂÒfl, ÌÂË̇ÍÚË‚Ë-
Û˛˘ËÂÒfl ‚˚ıÓ‰fl˘Ë ÚÓÍË. àÁ‚ÂÒÚÌÓ, ˜ÚÓ Û ·Óθ-
¯ËÌÒÚ‚‡ ÍÎÂÚÓÍ èá ‚˚ıÓ‰fl˘Ë ÚÓÍË Ó·ÂÒÔ˜˂‡-
˛ÚÒfl K+-͇̇·ÏË. é‰Ì‡ÍÓ ˜‡ÒÚ˘̇fl Á‡ÏÂ̇ ‚ÌÂ-
ÍÎÂÚÓ˜ÌÓ„Ó NaCl ıÎÓˉÓÏ ÚÂÚ‡˝ÚË·ÏÏÓÌËfl
(TEACl) – ·ÎÓ͇ÚÓÓÏ Í‡ÎË‚˚ı ͇̇ÎÓ‚ – Ò··Ó
·ÎÓÍËÓ‚‡Î‡ ‚˚ıÓ‰fl˘Ë ÚÓÍË (ËÒ. 1‡, ·). ùÚÓ Û͇-
Á˚‚‡ÂÚ Ì‡ ÌÂÁ̇˜ËÚÂθÌ˚È ‚Í·‰ èá K+-͇̇ÎÓ‚,
ÔÓ Í‡ÈÌÂÈ ÏÂ TEA-˜Û‚ÒÚ‚ËÚÂθÌ˚ı. êfl‰ ‰Û„Ëı
Ù‡ÍÚÓ‚ Ú‡ÍÊ ҂ˉÂÚÂθÒÚ‚Ó‚‡Î Ó ÚÓÏ, ˜ÚÓ Óθ
K+-͇̇ÎÓ‚ ‚ „ÂÌÂ‡ˆËË èá ‚˚ıÓ‰fl˘Ëı ÚÓÍÓ‚ ‚
ÍÎÂÚ͇ı ÚËÔ‡ II ‚ ˆÂÎÓÏ Ï‡Î‡. Ç ˜‡ÒÚÌÓÒÚË, ÂÔÓÎfl-
ËÁ‡ˆËfl ÍÎÂÚÓÍ ÚËÔ‡ II ‰Ó ÔÓÚÂ̈ˇ· Vh = –70 ÏÇ
‚˚Á˚‚‡Î‡ ‚ıÓ‰fl˘Ë ı‚ÓÒÚÓ‚˚ ÚÓÍË (ËÒ. 1‡), ıÓÚfl
ÂÒÎË ·˚ ËÓÌ˚ K+ ·˚ÎË ÌÓÒËÚÂÎflÏË ˝ÚËı ÚÓÍÓ‚, ÚÓ
ÓÌË ‰ÓÎÊÌ˚ ·˚ÎË ·˚ ·˚Ú¸ ‚˚ıÓ‰fl˘ËÏË. ÑÂÈÒÚ‚Ë-
ÚÂθÌÓ, Ï„ÌÓ‚ÂÌÌ˚È ı‚ÓÒÚÓ‚ÓÈ (tail) ÚÓÍ IT ÏÓÊÌÓ
Á‡ÔËÒ‡Ú¸ ‚ ‚ˉÂ:
I T = G ( V ) ( V h – V r ), (1)
„‰Â G(V) – ËÌÚ„‡Î¸Ì‡fl ÔÓ‚Ó‰ËÏÓÒÚ¸ ÔË ‰‡ÌÌÓÏ
ÏÂÏ·‡ÌÌÓÏ ÔÓÚÂ̈ˇÎ V, Vh – ÔÓ‰‰ÂÊË‚‡ÂÏ˚È

5 ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009

66 ÓχÌÓ‚ Ë ‰.
ÔÓÚÂ̈ˇÎ, Vr – ÔÓÚÂ̈ˇΠ‚ÂÒËË. àÁ Û‡‚ÌÂÌËfl
(1) ÒΉÛÂÚ, ˜ÚÓ ‚ ÔÒ‚‰ÓÙËÁËÓÎӄ˘ÂÒÍËı ËÓÌÌ˚ı
ÛÒÎÓ‚Ëflı ÔË Vh = –70 ÏÇ Í‡ÎË‚˚ ͇̇Î˚ ‰ÓÎÊ-
Ì˚ ÔÂÂÌÓÒËÚ¸ ‚˚ıÓ‰fl˘Ë ı‚ÓÒÚÓ‚˚ ÚÓÍË, Ú‡Í
Í‡Í ‚ ÛÒÎÓ‚Ëflı [K+]in = 140 Ïå/[K+]out = 2.5 Ïå ÔÓ-
ÚÂ̈ˇΠ‚ÂÒËË ‰Îfl ä+-ÒÂÎÂÍÚË‚Ì˚ı ͇̇ÎÓ‚ ÒÓ-
ÒÚ‡‚ÎflÂÚ ÓÍÓÎÓ –100 ÏÇ. ëΉӂ‡ÚÂθÌÓ, ‚ıÓ‰fl˘ËÂ
ı‚ÓÒÚÓ‚˚ ÚÓÍË Ì ÏÓ„ÛÚ ÔÂÂÌÓÒËÚ¸Òfl ËÓ̇ÏË ä+,
ÌÓ Ó·ÂÒÔ˜˂‡˛ÚÒfl ÎË·Ó Í‡Ì‡Î‡ÏË, ÔÓÌˈ‡ÂÏ˚-
ÏË ‰Îfl ‰Û„Ëı ÙËÁËÓÎӄ˘ÂÒÍË ‚‡ÊÌ˚ı ËÓÌÓ‚, Ú‡-
ÍËı Í‡Í Na+, Ca2+ Ë Cl–, ÎË·Ó ÌÂÒÂÎÂÍÚË‚Ì˚ÏË ËÓÌ-
Ì˚ÏË Í‡Ì‡Î‡ÏË. àÓÌÌ˚ ÚÓÍË, ÚÂÍÛ˘Ë ˜ÂÂÁ ˝ÚË
͇̇Î˚, ÔË ÙËÁËÓÎӄ˘ÂÒÍËı ËÓÌÌ˚ı „‡‰ËÂÌÚ‡ı
ı‡‡ÍÚÂËÁÛ˛ÚÒfl ÔÓÎÓÊËÚÂθÌ˚ÏË ËÎË ·ÎËÁÍËÏË
Í ÌÛβ ÔÓÚÂ̈ˇ·ÏË ‚ÂÒËË; ÒΉӂ‡ÚÂθÌÓ,
ÔË Vh = –70 ÏÇ ı‚ÓÒÚÓ‚˚ ÚÓÍË ˜ÂÂÁ ˝ÚË Í‡Ì‡Î˚
‰ÓÎÊÌ˚ ·˚Ú¸ ‚ıÓ‰fl˘ËÏË.
Ç ˝ÍÒÔÂËÏÂÌÚ‡ı, ‚ ÍÓÚÓ˚ı Ô˝Ú˜-ÔËÔÂÚÍË ÒÓ-
‰ÂʇÎË 140 ÏM CsCl, Û ÍÎÂÚÓÍ II ÚËÔ‡ ÒÓı‡ÌflÎËÒ¸
èá ωÎÂÌÌÓ ‡ÍÚË‚ËÛ˛˘ËÂÒfl, ÌÂË̇ÍÚË‚ËÛ˛-
˘ËÂÒfl ÚÓÍË (‚ ‰‡Î¸ÌÂȯÂÏ SA-ÚÓÍË, slowly activat-
ing) Ë ‚ıÓ‰fl˘Ë ı‚ÓÒÚÓ‚˚ ÚÓÍË (ËÒ. 1‚). ÄÏÔÎË-
ÚÛ‰‡, ÍËÌÂÚË͇ Ë ÔÓÚÂ̈ˇΠ‚ÂÒËË ˝ÚËı ÚÓÍÓ‚
·˚ÎË ·ÎËÁÍË Í ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘ËÏ Ô‡‡ÏÂÚ‡Ï, ̇-
·Î˛‰‡ÂÏ˚Ï ‚ ÛÒÎÓ‚Ëflı ‰Ë‡ÎËÁ‡ ÍÎÂÚÓÍ ‡ÒÚ‚ÓÓÏ
Ò 140 Ïå KCl (ËÒ. 1‡, ·). ùÚÓ Ò‚Ë‰ÂÚÂθÒÚ‚ÛÂÚ Ó
ÚÓÏ, ˜ÚÓ SA-ÚÓÍË Ó·ÂÒÔ˜˂‡˛ÚÒfl ËÓÌÌ˚ÏË Í‡Ì‡-
·ÏË ÔÂËÏÛ˘ÂÒÚ‚ÂÌÌÓ Ó‰ÌÓ„Ó ÚËÔ‡, ÍÓÚÓ˚ ÏÓ-
„ÛÚ ·˚Ú¸ ÔÓÌˈ‡ÂÏ˚ ‰Îfl ËÓÌÓ‚ Na+, K+, Cs+ Ë/ËÎË
Cl–. ç‡ ˝ÚÓ Ê Û͇Á˚‚‡ÂÚ Ë ÚÓÚ Ù‡ÍÚ, ˜ÚÓ ÔÓ‰ ‰ÂÈ-
ÒÚ‚ËÂÏ ·ÎÓ͇ÚÓÓ‚ Ë ÏÂÌfl˛˘ËıÒfl ËÓÌÌ˚ı ÛÒÎÓ‚ËÈ
‚ÓθÚ-‡ÏÔÂÌ˚ (I–V) ı‡‡ÍÚÂËÒÚËÍË SA-ÚÓÍÓ‚
ÎË·Ó Ò‰‚Ë„‡ÎËÒ¸ ÔÓ ÓÒË ÔÓÚÂ̈ˇÎÓ‚, ÎË·Ó Ï‡Ò¯Ú‡-
·ËÓ‚‡ÎËÒ¸ ÔÓ ÓÒË ÚÓÍÓ‚, Ú.Â. Ù‡ÍÚ˘ÂÒÍË ÔÓ‰‚Â-
„‡ÎËÒ¸ ÎËÌÂÈÌ˚Ï ÔÂÓ·‡ÁÓ‚‡ÌËflÏ ·ÂÁ ÒÛ˘ÂÒÚ‚ÂÌ-
ÌÓ„Ó ËÁÏÂÌÂÌËfl Ëı ÙÓÏ˚ (ÒÏ. ÌËÊÂ). Ç ‰‡Î¸ÌÂȯËı
˝ÍÒÔÂËÏÂÌÚ‡ı Ò‚ÓÈÒÚ‚‡ SA-ÚÓÍÓ‚ ËÒÒΉӂ‡ÎËÒ¸ ‚
ÛÒÎÓ‚Ëflı ‰Ë‡ÎËÁ‡ ÍÎÂÚÓÍ Cs+ ËÎË ‰Û„ËÏË Í‡ÚËÓ̇-
ÏË, Ì ÔÓÌË͇˛˘ËÏË ˜ÂÂÁ K+-ÒÂÎÂÍÚË‚Ì˚ ͇̇-
Î˚, ˜ÚÓ·˚ ÔÓÎÌÓÒÚ¸˛ ËÒÍβ˜ËÚ¸ ‚Í·‰ ÔÓÒΉÌËı
‚ èá ËÌÚ„‡Î¸Ì˚ ÚÓÍË.
ÑÎfl SA-ÚÓÍÓ‚ ·˚ÎÓ ı‡‡ÍÚÂÌÓ ‚˚‡ÊÂÌÌÓÂ
‚˚ÔflÏÎÂÌË ÔÓ ‚˚ıÓ‰fl˘ÂÏÛ ÚÓÍÛ. ç‡ I–V ı‡‡Í-
ÚÂËÒÚËÍ ÚÓ͇ ‚ ӷ·ÒÚË Ì·Óθ¯Ëı ÓÚˈ‡ÚÂθ-
Ì˚ı ÔÓÚÂ̈ˇÎÓ‚ ΂ ÔÓÚÂ̈ˇ· ‚ÂÒËË SA-ÚÓ-
ÍÓ‚ Ó·˚˜ÌÓ Ì‡·Î˛‰‡Î‡Ò¸ Ì·Óθ¯‡fl ‚ıÓ‰fl˘‡fl
ÍÓÏÔÓÌÂÌÚ‡ (ËÒ. 1·, „), ‚Â΢Ë̇ ÍÓÚÓÓÈ ‚‡¸Ë-
Ó‚‡Î‡ ÓÚ ˝ÍÒÔÂËÏÂÌÚ‡ Í ˝ÍÒÔÂËÏÂÌÚÛ. ùÚÓ ·˚ÎÓ
Ò‚flÁ‡ÌÓ Ò ÚÂÏ, ˜ÚÓ ÔÓÎÓÊÂÌË ‡ÍÚË‚‡ˆËÓÌÌÓÈ ÍË-
‚ÓÈ SA-͇̇ÎÓ‚ ̇ ÓÒË ÔÓÚÂ̈ˇÎÓ‚ Ë ÔÓÚÂ̈ˇ·
‚ÂÒËË SA-ÚÓÍÓ‚ ‚‡¸ËÓ‚‡ÎË ÓÚ ÍÎÂÚÍË Í ÍÎÂÚ-
ÍÂ. èÓÚÂ̈ˇΠ‚ÂÒËË ÓÔ‰ÂÎflÎÒfl ÔÛÚÂÏ ‡Ì‡ÎË-
Á‡ ı‚ÓÒÚÓ‚˚ı ÚÓÍÓ‚, ÔÓÒÍÓθÍÛ SA-ÚÓÍË ‚ÂÒË-
Ó‚‡ÎËÒ¸ ‚ÓÁΠÌÛÎÂ‚Ó„Ó ÔÓÚÂ̈ˇ·, ÔË ÍÓÚÓÓÏ
Ëı ‡ÏÔÎËÚÛ‰‡ ·˚· χ·, ˜ÚÓ Ì ÔÓÁ‚ÓÎflÂÚ Ò ‰ÓÒÚ‡-
ÚÓ˜ÌÓÈ ÚÓ˜ÌÓÒÚ¸˛ ÓˆÂÌËÚ¸ Vr.
àÌÚ„‡Î¸Ì˚È ÒÚ‡ˆËÓ̇Ì˚È ÚÓÍ IS, ËÁÏÂflÂ-
Ï˚È ‚ ÍÎÂÚ͇ı, ‰Ë‡ÎËÁÛÂÏ˚ı 140 Ïå CsCl, ÏÓÊÌÓ
ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009
Ô‰ÒÚ‡‚ËÚ¸ ‚ ‚ˉ ÒÛÏÏ˚ ‰‚Ûı ÌÂÁ‡‚ËÒËÏ˚ı ÍÓÏ-
ÔÓÌÂÌÚ, ‡ ËÏÂÌÌÓ – ÒÚ‡ˆËÓ̇ÌÓ„Ó SA-ÚÓ͇ ISA Ë ÚÓ-
͇ ÛÚ˜ÍË Il:
I S = I SA + I l . (2)
èË ·Óθ¯Ëı ÍÓχ̉Ì˚ı ÔÓÚÂ̈ˇ·ı SA-ÚÓÍË ‰Ó-
ÒÚË„‡ÎË 2 ÌÄ Ë ·ÓÎÂÂ, Ë ÔÓ˝ÚÓÏÛ ÓÚÌÓÒËÚÂθÌ˚È
‚Í·‰ ÚÓ͇ ÛÚ˜ÍË ‚ ÒÛÏχÌ˚È ÚÓÍ ·˚ΠχÎ. äÓ-
Ï ÚÓ„Ó, ÚÓÍ ÛÚ˜ÍË, ÍÓÚÓ˚È Ò˜ËÚ‡ÎÒfl ÎËÌÂÈÌÓ Á‡-
‚ËÒËÏ˚Ï ÓÚ ÏÂÏ·‡ÌÌÓ„Ó ÔÓÚÂ̈ˇ·, ‡‚ÚÓχÚË-
˜ÂÒÍË ÍÓÏÔÂÌÒËÓ‚‡ÎÒfl ÔË ÔÓÏÓ˘Ë ÔÓ„‡ÏÏ˚
Clamfit 8.2.0. èÓ˝ÚÓÏÛ ‚ ӷ·ÒÚË ÔÓÎÓÊËÚÂθÌ˚ı
ÏÂÏ·‡ÌÌ˚ı ÔÓÚÂ̈ˇÎÓ‚ Ò ıÓÓ¯ÂÈ ÚÓ˜ÌÓÒÚ¸˛
I S = I SA = G SA ( V ) ( V – V r ), (3)
„‰Â GSA(V) – SA-ÔÓ‚Ó‰ËÏÓÒÚ¸ ÔË ‰‡ÌÌÓÏ ÏÂÏ-
·‡ÌÌÓÏ ÔÓÚÂ̈ˇÎ V. àÁ Û‡‚ÌÂÌËÈ (1) Ë (3) ÒÎÂ-
‰ÛÂÚ:
I
----S = a ( V – V r ), (4)
IT
1
„‰Â a = ----------------- = const.
Vh – Vr
ç‡ ËÒ. 2‡ (ÍÛÊÓ˜ÍË) Ô‰ÒÚ‡‚ÎÂÌÓ ÛÒ‰ÌÂÌ-
ÌÓ ÓÚÌÓ¯ÂÌË IS/IT Í‡Í ÙÛÌ͈Ëfl ÍÓχ̉ÌÓ„Ó ÔÓ-
ÚÂ̈ˇ·, ÔÓÎÛ˜ÂÌÌÓ ÔÓ ËÁÏÂÂÌËflÏ Ì‡ 10 ÍÎÂ-
Ú͇ı. ãËÌÂÈ̇fl „ÂÒÒËfl ˝ÍÒÔÂËÏÂÌڇθÌ˚ı ‰‡Ì-
Ì˚ı Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ Û‡‚ÌÂÌËfl (4) (ËÒ. 2‡,
ÎËÌËfl) ÔÓÁ‚ÓÎË· ÓÔ‰ÂÎËÚ¸ Á̇˜ÂÌË ÍÓ˝ÙÙˈË-
ÂÌÚ‡ a Ë ÚÂÏ Ò‡Ï˚Ï ÓˆÂÌËÚ¸ Ò‰Ì Á̇˜ÂÌË ÔÓ-
ÚÂ̈ˇ· ‚ÂÒËË, ÍÓÚÓ˚È Ó͇Á‡ÎÒfl Vr = –5.5 ÏÇ.
ç‡ ËÒ. 2· Ô‰ÒÚ‡‚ÎÂÌ˚ I–V ı‡‡ÍÚÂËÒÚËÍË
SA-ÚÓÍÓ‚ (ÔÓÎÛ˜ÂÌ˚ ‚˚˜ËÚ‡ÌËÂÏ ÚÓÍÓ‚ ÛÚ˜ÍË)
‰Îfl ‰‚Ûı ‡ÁÌ˚ı ÍÎÂÚÓÍ, ‰Îfl Ó‰ÌÓÈ ËÁ ÍÓÚÓ˚ı
‚ıӉ̇fl ÍÓÏÔÓÌÂÌÚ‡ SA-ÚÓ͇ ıÓÓ¯Ó ‚˚‡ÊÂ̇
(
), ‡ ‰Îfl ‰Û„ÓÈ – Ó̇ Ô‡ÍÚ˘ÂÒÍË ÓÚÒÛÚÒÚ‚ÛÂÚ
(). àÒÔÓθÁÛfl ÓÔËÒ‡ÌÌÛ˛ ‚˚¯Â ÏÂÚÓ‰ËÍÛ ÓÔ‰Â-
ÎÂÌËfl ÔÓÚÂ̈ˇ· ‚ÂÒËË ÔÓ ı‚ÓÒÚÓ‚˚Ï ÚÓ͇Ï,
Ï˚ ÓˆÂÌËÎË Vr ‚ ͇ʉÓÏ ËÁ ˝ÚËı ÒÎÛ˜‡Â‚ Í‡Í –0.5
(
) Ë –8.5 ÏÇ (). ë ËÒÔÓθÁÓ‚‡ÌËÂÏ ˝ÚËı ‰‡ÌÌ˚ı Ë
Û‡‚ÌÂÌËfl (3), ·˚ÎË ÔÓ‰Ò˜ËÚ‡Ì˚ ÔÓ‚Ó‰ËÏÓÒÚË GSA
‰Îfl ͇ʉÓÈ ÍÎÂÚÍË Í‡Í ÙÛÌ͈ËË ÏÂÏ·‡ÌÌÓ„Ó ÔÓ-
ÚÂ̈ˇ·, ÍÓÚÓ˚ ‚ ÌÓÏËÓ‚‡ÌÌÓÏ ‚ˉ Ô‰-
ÒÚ‡‚ÎÂÌ˚ ̇ ËÒ. 2‚. èÓ‚Ó‰ËÏÓÒÚË ‡ÔÔÓÍÒËÏËÓ-
‚‡ÎËÒ¸ Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ Û‡‚ÌÂÌËfl ÅÓθˆÏ‡Ì‡ [13]:
G SA ( V )/G max = 1/ ( 1 + exp ( – ( V – V g )/V g0 ) ). (5)
á‰ÂÒ¸ Gmax – χÍÒËχθ̇fl ÔÓ‚Ó‰ËÏÓÒÚ¸, Vg – ÔÓ-
ÚÂ̈ˇΠÔÓÎÛ‡ÍÚË‚‡ˆËË, Vg0 = RT/nF, „‰Â n – ˝Í‚Ë‚‡-
ÎÂÌÚÌÓ ˜ËÒÎÓ ‚ÓÓÚÌ˚ı Á‡fl‰Ó‚, ‚˚‡ÊÂÌÌ˚ı ‚
˝ÎÂÏÂÌÚ‡Ì˚ı Á‡fl‰‡ı; R, T Ë F – „‡ÁÓ‚‡fl ÔÓÒÚÓflÌ-
̇fl, ‡·ÒÓβÚ̇fl ÚÂÏÔÂ‡ÚÛ‡ Ë ÍÓÌÒÚ‡ÌÚ‡ î‡‡‰Âfl
ÒÓÓÚ‚ÂÚÒÚ‚ÂÌÌÓ. Ä̇ÎËÁ Ô˂‰ÂÌÌ˚ı ˝ÍÒÔÂËÏÂÌ-
ڇθÌ˚ı ‰‡ÌÌ˚ı ÔÓÁ‚ÓÎËÎ ÔÓÎÛ˜ËÚ¸ ÒÎÂ‰Û˛˘ËÂ
ÓˆÂÌÍË: Vg = –15 ÏÇ, Vg0 = 6 ÏÇ (
) Ë Vg = 4 ÏÇ, Vg0 =
= 12 ÏÇ (). í‡ÍËÏ Ó·‡ÁÓÏ, ‡ÍÚË‚‡ˆËÓÌ̇fl ÍË-
‚‡fl SA-͇̇ÎÓ‚, ÙÛÌ͈ËÓÌËÛ˛˘Ëı ‚ ÍÎÂÚÍÂ, ‚ ÍÓ-
 çÂÒÂÎÂÍÚË‚Ì˚ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ ͇̇Î˚ 67

IS /IT Inorm Gnorm

‡ 1.0 · 1.0 ‚

–0.4
0.5
–0.8 0.5

0
–1.2
0
20 40 60 –50 0 50 –50 0 50
åÂÏ·‡ÌÌ˚È ÔÓÚÂ̈ˇÎ, ÏÇ

êËÒ. 2. ï‡‡ÍÚÂËÒÚËÍË SA-ÚÓÍÓ‚ ‚ ÍÎÂÚ͇ı, ‰Ë‡ÎËÁÛÂÏ˚ı ‡ÒÚ‚ÓÓÏ Ò 140 Ïå CsCl. ‡ – éÚÌÓ¯ÂÌË ‡ÏÔÎËÚÛ‰˚ ÒÚ‡ˆËÓ-
̇ÌÓ„Ó ÚÓ͇ ÔË ‰‡ÌÌÓÏ ÍÓχ̉ÌÓÏ ÔÓÚÂ̈ˇÎÂ Í ‡ÏÔÎËÚۉ ı‚ÓÒÚÓ‚˚ı ÚÓÍÓ‚. ëËÏ‚ÓÎ˚ ÓÚ‡Ê‡˛Ú ˝ÍÒÔÂËÏÂÌڇθÌ˚Â
‰‡ÌÌ˚Â, Ô‰ÒÚ‡‚ÎÂÌÌ˚Â Í‡Í Ò‰Ì ± Òڇ̉‡Ú̇fl ӯ˷͇. ãËÌËfl ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ Û‡‚ÌÂÌ˲ (4) ÔË Vr = –5.5 ÏÇ Ë Vh =
= –70 ÏÇ. · – ëÚ‡ˆËÓ̇Ì˚È ÚÓÍ, ÌÓÏËÓ‚‡ÌÌ˚È Ì‡ χÍÒËχθÌÓ Á̇˜ÂÌËÂ, Í‡Í ÙÛÌ͈Ëfl ÏÂÏ·‡ÌÌÓ„Ó ÔÓÚÂ̈ˇ·, ÔÓ-
ÎÛ˜ÂÌ̇fl ‰Îfl ‰‚Ûı ‡ÁÌ˚ı ‚ÍÛÒÓ‚˚ı ÍÎÂÚÓÍ ÚËÔ‡ II. ‚ – éÚÌÓÒËÚÂθ̇fl ËÌÚ„‡Î¸Ì‡fl ÔÓ‚Ó‰ËÏÓÒÚ¸ ÍÎÂÚÓÍ, I–V ı‡‡ÍÚÂ-
ËÒÚËÍË ÍÓÚÓ˚ı Ô‰ÒÚ‡‚ÎÂÌ˚ ‚ ·. ëËÏ‚ÓÎ˚ ÒÓÓÚ‚ÂÚÒÚ‚Û˛Ú ˝ÍÒÔÂËÏÂÌڇθÌ˚Ï ‰‡ÌÌ˚Ï, ÍË‚˚ ÔÓÎÛ˜ÂÌ˚ ‚˚˜ËÒÎÂ-
ÌËÂÏ Ò ÔÓÏÓ˘¸˛ Û‡‚ÌÂÌËfl (5) (ÔÓ‰Ó·ÌÓÒÚË ‚ ÚÂÍÒÚÂ). ÇÓ ‚ÒÂı ÒÎÛ˜‡flı ËÒÔÓθÁÓ‚‡ÎÒfl ÏÂÚÓ‰ perforated patch.

ÚÓÓÈ ‚ıÓ‰fl˘‡fl ÍÓÏÔÓÌÂÌÚ‡ SA-ÚÓÍÓ‚ ·˚· ÏËÌË-
χθ̇ (ËÒ. 2‚, ), ·˚· Ò‰‚ËÌÛÚ‡ ‚ ÒÚÓÓÌÛ ÔÓÎÓ-
ÊËÚÂθÌ˚ı ÔÓÚÂ̈ˇÎÓ‚ ÔËÏÂÌÓ Ì‡ 20 ÏÇ ÔÓ
Ò‡‚ÌÂÌ˲ Ò ‡ÍÚË‚‡ˆËÓÌÌÓÈ ÍË‚ÓÈ, ÔÓÎÛ˜ÂÌÌÓÈ
‰Îfl ÍÎÂÚÍË Ò ‚˚‡ÊÂÌÌÓÈ ‚ıÓ‰fl˘ÂÈ ÍÓÏÔÓÌÂÌÚÓÈ
(ËÒ. 2‚,
).
èÓÒÍÓθÍÛ ÔË ËÒÔÓθÁÛÂÏÓÏ ËÓÌÌÓÏ „‡‰ËÂÌ-
Ú ([CsCl]in = 140 Ïå/[NaCl]out = 140 Ïå) ÔÓÚÂ̈ˇÎ
‚ÂÒËË SA-ÚÓ͇ ·˚Î ·ÎËÁÓÍ Í 0 ÏÇ, ÚÓ ‚ ÒÎÛ˜‡Â,
ÂÒÎË ÓÌ Ó·ÂÒÔ˜˂‡ÂÚÒfl ËÓÌÌ˚ÏË Í‡Ì‡Î‡ÏË Ó‰ÌÓ„Ó
ÚËÔ‡, ˝ÚË Í‡Ì‡Î˚ ÎË·Ó ‚ ‡‚ÌÓÈ ÒÚÂÔÂÌË ÔÓÌˈ‡Â-
Ï˚ ‰Îfl Cs+ Ë Na+, ÎË·Ó ÒÂÎÂÍÚË‚Ì˚ ‰Îfl ËÓÌÓ‚ ıÎÓ-
‡, ÎË·Ó ı‡‡ÍÚÂËÁÛ˛ÚÒfl Ò··ÓÈ Í‡ÚËÓÌ/‡ÌËÓÌ-
ÌÓÈ ÒÂÎÂÍÚË‚ÌÓÒÚ¸˛. äÓ„‰‡ ‚ÌÂÍÎÂÚÓ˜Ì˚È Cl– Á‡-
ÏÂÌflÎÒfl ÍÛÔÌ˚Ï ‡ÌËÓÌÓÏ „βÍÓ̇ÚÓÏ, ÍÓÚÓ˚È
Ì ÔÓÌË͇ÂÚ ˜ÂÂÁ Cl–-͇̇Î˚, ÔÓÚÂ̈ˇΠ‚Â-
ÒËË SA-ÚÓ͇ ÒÏ¢‡ÎÒfl ‚ ÒÚÓÓÌÛ ÔÓÎÓÊËÚÂθÌ˚ı
ÔÓÚÂ̈ˇÎÓ‚ [8] (ËÒ. 3‡, ·) ̇ 7.7 ± 2.2 ÏÇ. ùÚÓ Ò‚Ë-
‰ÂÚÂθÒÚ‚ÛÂÚ Ó ÚÓÏ, ˜ÚÓ SA-ÚÓÍ ˜‡ÒÚ˘ÌÓ ÔÂÂÌÓ-
ÒËÚÒfl ËÓ̇ÏË Cl–. åÓÊÌÓ Ô‰ÔÓÎÓÊËÚ¸, ˜ÚÓ SA-
ÚÓÍ ˜‡ÒÚ˘ÌÓ Ó·ÂÒÔ˜˂‡ÂÚÒfl Cl–-͇̇·ÏË, ‡ ˜‡-
ÒÚ˘ÌÓ – ͇ÚËÓÌÌ˚ÏË Í‡Ì‡Î‡ÏË. å˚ ÔÓ‚ÂflÎË
˝ÚÛ ‚ÓÁÏÓÊÌÓÒÚ¸, ËÒÒΉÛfl ˜Û‚ÒÚ‚ËÚÂθÌÓÒÚ¸ SA-ÚÓ-
͇ Í fl‰Û ·ÎÓ͇ÚÓÓ‚, ‚Íβ˜‡fl NPPB, 9-AC, ÌËÙ-
ÎÛÏÓ‚Û˛ ÍËÒÎÓÚÛ, SITS, DIDS, ÍÓÚÓ˚ ËÒÔÓθÁÓ-
‚‡ÎËÒ¸ ‚ ÍÓ̈ÂÌÚ‡ˆËflı, ÔË ÍÓÚÓ˚ı ÓÌË Ó·˚˜ÌÓ
‚˚Á˚‚‡˛Ú ÔÓ˜ÚË ÔÓÎÌÛ˛ ·ÎÓ͇‰Û Cl–-͇̇ÎÓ‚ ÒÓ-
ÓÚ‚ÂÚÒÚ‚Û˛˘Ëı ÔÓ‰ÚËÔÓ‚. é͇Á‡ÎÓÒ¸, ˜ÚÓ ÌË Ó‰ËÌ
ËÁ ÌËı Ì Ó͇Á˚‚‡Î ÒÛ˘ÂÒÚ‚ÂÌÌÓ„Ó ‚ÎËflÌËfl ̇ ‡Ï-
ÔÎËÚÛ‰Û SA-ÚÓ͇ (ËÒ. 3‚). ë‡Ï˚È ˝ÙÙÂÍÚË‚Ì˚È ËÁ
·ÎÓ͇ÚÓÓ‚, DIDS, ÒÌËʇΠ‡ÏÔÎËÚÛ‰Û ÚÓ͇ ‚ Ò‰-
ÌÂÏ Ì‡ 22.8 ± 5.4% (ËÒ. 3‚, „). í‡ÍËÏ Ó·‡ÁÓÏ, ‚ÓÁ-
ÏÓÊÌ˚È ‚Í·‰ DIDS-˜Û‚ÒÚ‚ËÚÂθÌ˚ı Cl–-͇̇ÎÓ‚
‚ SA-ÚÓÍ Ì ÏÓ„ Ô‚˚¯‡Ú¸ 23%, Ë, ÒΉӂ‡ÚÂθÌÓ,
77% ÚÓ͇ ‰ÓÎÊÌ˚ ·˚ÎË ·˚ Ó·ÂÒÔ˜˂‡Ú¸Òfl ͇ÚË-
ÓÌÌ˚ÏË Í‡Ì‡Î‡ÏË. ÄθÚÂ̇ÚË‚ÌÓ ӷ˙flÒÌÂÌËÂ
̇·Î˛‰‡‚¯ËıÒfl fl‚ÎÂÌËÈ ÒÓÒÚÓflÎÓ ‚ ÚÓÏ, ˜ÚÓ SA-
ÚÓÍ Ó·ÛÒÎÓ‚ÎÂÌ ‡ÍÚË‚ÌÓÒÚ¸˛ ͇̇ÎÓ‚ Ó‰ÌÓ„Ó ÚËÔ‡,
ÔÓÌˈ‡ÂÏ˚ÏË Í‡Í ‰Îfl ͇ÚËÓÌÓ‚, Ú‡Í Ë ‰Îfl ‡ÌËÓ-
ÌÓ‚, Ë ˜Û‚ÒÚ‚ËÚÂθÌ˚ÏË Í DIDS. Ç ˝ÚÓÏ ÒÎÛ˜‡Â ÒÚ‡-
ÌÓ‚ËÚÒfl ÔÓÌflÚÌ˚Ï, ÔÓ˜ÂÏÛ ÔÓÏËÏÓ ÛÏÂ̸¯ÂÌËfl
‡ÏÔÎËÚÛ‰˚ SA-ÚÓ͇ DIDS (Ë ‚ ÏÂ̸¯ÂÈ ÒÚÂÔÂÌË –
SITS) Û‚Â΢˂‡Î Â„Ó ÙÎÛÍÚÛ‡ˆËË Ë Á‡Ï‰ÎflÎ Â„Ó ‰Â-
‡ÍÚË‚‡ˆË˛ ÔË ÂÔÓÎflËÁ‡ˆËË ÏÂÏ·‡Ì˚ (ËÒ.
3„,‰). Ç ÚÓ Ê ‚ÂÏfl ˝ÚË Ù‡ÍÚ˚ Ì‚ÓÁÏÓÊÌÓ ‡ˆË-
Ó̇θÌÓ Ó·˙flÒÌËÚ¸, ÂÒÎË SA-ÚÓÍ Ô‰ÒÚ‡‚ÎflÂÚ ÒÓ-
·ÓÈ ÒÛÏÏÛ ÚÓÍÓ‚ ‚ ÒÓÓÚÌÓ¯ÂÌËË 23 Í 77, ÍÓÚÓ˚Â
ÌÂÁ‡‚ËÒËÏÓ ÔÂÂÌÓÒflÚÒfl ˜ÂÂÁ DIDS-˜Û‚ÒÚ‚ËÚÂθ-
Ì˚ Cl–-͇̇Î˚ Ë DIDS-̘ۂÒÚ‚ËÚÂθÌ˚ ͇ÚËÓÌ-
Ì˚ ͇̇Î˚.
äÓ„‰‡ ÍÎÂÚÍË ‰Ë‡ÎËÁÓ‚‡ÎËÒ¸ ‡ÒÚ‚ÓÓÏ, ÒÓ‰Â-
ʇ˘ËÏ 140 Ïå NMDG-Cl, ‡ ‚ÌÂÍÎÂÚÓ˜Ì˚È ‡ÒÚ‚Ó
ÒÓ‰ÂʇΠ140 Ïå NaCl (n = 9), ‡ÏÔÎËÚÛ‰‡ ‚˚ıÓ‰fl-
˘Ëı ÚÓÍÓ‚ Ë ÔÓÚÂ̈ˇΠ‚ÂÒËË (Vr = –4.7 ÏÇ) ‚
Ò‰ÌÂÏ ·˚ÎË ‚ ÚÓÏ Ê ‰Ë‡Ô‡ÁÓÌÂ, ˜ÚÓ Ë ‚ ÒÎÛ˜‡Â
‰Ë‡ÎËÁ‡ ÍÎÂÚÓÍ ‚ ÔËÒÛÚÒÚ‚ËË ‚ÌÛÚËÍÎÂÚÓ˜ÌÓ„Ó
140 Ïå CsCl (n = 28) (ËÒ. 3‡; ΂‡fl Ô‡ÌÂθ; ËÒ. 4‡,
΂‡fl Ô‡ÌÂθ; ËÒ. 4·). ùÚÓ ÓÁ̇˜‡ÎÓ, ˜ÚÓ ‚˚ıÓ‰fl-
˘ËÈ ÚÓÍ ÔË ÔÓÎÓÊËÚÂθÌ˚ı ÏÂÏ·‡ÌÌ˚ı ÔÓÚÂÌ-
ˆË‡Î‡ı ÏÓ„ ·˚Ú¸ Ó·ÛÒÎÓ‚ÎÂÌ ‚˚ıÓ‰fl˘ËÏ ÔÓÚÓÍÓÏ
NMDG+ Ë/ËÎË ‚ıÓ‰fl˘ËÏ ÔÓÚÓÍÓÏ Cl–. èË Á‡ÏÂÌÂ
‚ÌÂÍÎÂÚÓ˜ÌÓ„Ó Cl– „βÍÓ̇ÚÓÏ ÔÓËÒıÓ‰ËÎÓ Û‚Â-
΢ÂÌË ‚ıÓ‰fl˘ÂÈ ÍÓÏÔÓÌÂÌÚ˚ ËÌÚ„‡Î¸ÌÓ„Ó ÚÓ-
͇, ÒÏ¢ÂÌË ÔÓÚÂ̈ˇ· ‚ÂÒËË ‰Ó 14 ± 4 ÏÇ Ë
ÔÓ‰‡‚ÎÂÌË ‚˚ıÓ‰fl˘Â„Ó ÚÓ͇, ·ÓΠÒËθÌÓÂ, ÌÂ-
ÊÂÎË ÔË ‰Ë‡ÎËÁ ÍÎÂÚÓÍ CsCl (66.0 ± 12.5% ÔÓÚË‚
26.9 ± 5.0%) (ËÒ. 4‚). ùÚË Ù‡ÍÚ˚ ҂ˉÂÚÂθÒÚ‚Û˛Ú
Ó ÚÓÏ, ˜ÚÓ NMDG+ ÔÓÌË͇ÂÚ ˜ÂÂÁ SA-͇̇Î˚,
ÍÓÚÓ˚ ڇÍÊ ÔÓÌˈ‡ÂÏ˚ ‰Îfl ‡ÌËÓÌÓ‚ „βÍÓ-
̇ڇ, ıÓÚfl Á‡ÏÂÚÌÓ ıÛÊÂ, ˜ÂÏ ‰Îfl ËÓÌÓ‚ Cl–.
àÚ‡Í, ÓÔËÒ‡ÌÌ˚ ‚˚¯Â ÂÁÛθڇÚ˚ ÚÛ‰ÌÓ Ó·˙-
flÒÌËÚ¸, ÂÒÎË Ì Ô‰ÔÓÎÓÊËÚ¸, ˜ÚÓ SA-ÚÓÍ ‚Ó ‚ÍÛ-
ÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II Ó·ÂÒÔ˜˂‡ÂÚÒfl ÌÂÒÂÎÂÍÚË‚-
Ì˚ÏË ËÓÌÌ˚ÏË Í‡Ì‡Î‡ÏË, ÍÓÚÓ˚ ÔÓÌˈ‡ÂÏ˚
‰Îfl ͇ÚËÓÌÓ‚ Ë ‡ÌËÓÌÓ‚, ‚Íβ˜‡fl Cs+, Na+, Cl–,
NMDG+ Ë „βÍÓ̇Ú. í‡Í‡fl ÌÂÒÔˆËÙ˘ÌÓÒÚ¸ ͇-

ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009 5*

68 ÓχÌÓ‚ Ë ‰.

‡ ·
50 ÏÇ

àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ
–70 ÏÇ 4
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ
3 100 ÏÇ 3
2
1
0 0
CsClin /NaClout CsClin / –1
Na-„βÍÓ̇Úout –50 0 50
–3 åÂÏ·‡ÌÌ˚È
ÔÓÚÂ̈ˇÎ, ÏÇ
‚

0.2 Ïå NPPB
ÌËÙÎÛÏÓ‚‡fl

1 Ïå DIDS
1 Ïå 9-AC
1 Ïå SITS
ÍËÒÎÓÚ‡
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

1 Ïå

4
3
2
1
0
0 3 6 9 12 15
ÇÂÏfl, ÏËÌ
„ ‰
50 ÏÇ
–70 ÏÇ äÓÌÚÓθ 0.5 Ïå DIDS
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

100 ÏÇ
3

0
25 ÏÒ
äÓÌÚÓθ 0.5 Ïå DIDS
–3

êËÒ. 3. ÑÂÈÒÚ‚Ë ·ÎÓ͇ÚÓÓ‚ Cl–-͇̇ÎÓ‚ ̇ SA-ÚÓÍË. ‡ – SA-ÚÓÍË, „ÂÌÂËÛÂÏ˚ ‚ ÓÚ‚ÂÚ Ì‡ 100 ÏÒ ÔÓÎflËÁ‡ˆË˛ (‚ÂıÌflfl
Ô‡ÌÂθ), ‚ ÍÓÌÚÓΠ(ÌËÊÌflfl ΂‡fl Ô‡ÌÂθ) Ë ÔÓÒΠÁ‡ÏÂÌ˚ 140 Ïå NaCl ‚ ̇ÛÊÌÓÏ ‡ÒÚ‚Ó ̇ 140 Ïå Na-„βÍÓ̇Ú
(ÌËÊÌflfl Ô‡‚‡fl ̇ÌÂθ). ëËÏ‚ÓÎ˚ ̇‰ ÍË‚˚ÏË ÚÓÍÓ‚ Û͇Á˚‚‡˛Ú ÏÓÏÂÌÚ˚ ÒÌflÚËfl I–V ı‡‡ÍÚÂËÒÚËÍ. · – ÇÓθÚ-‡ÏÔÂ-
Ì˚ ı‡‡ÍÚÂËÒÚËÍË ÚÓÍÓ‚, Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ‡. ‚ – ÑÂÈÒÚ‚Ë ‡ÌËÓÌÌ˚ı ·ÎÓ͇ÚÓÓ‚ ̇ ‡ÏÔÎËÚÛ‰Û SA-ÚÓ͇, „ËÒÚËÛ-
ÂÏÓ„Ó ÔË ‰ÂÔÓÎflËÁ‡ˆËË ÍÎÂÚÍË ÓÚ –70 ‰Ó 50 ÏÇ. „ – çËÊÌË ԇÌÂÎË: SA-ÚÓÍË ‚ ÍÓÌÚÓΠ(΂‡fl Ô‡ÌÂθ) Ë ‚ ÔËÒÛÚÒÚ‚ËË
·ÎÓ͇ÚÓ‡ Cl–-͇̇ÎÓ‚ DIDS (Ô‡‚‡fl Ô‡ÌÂθ). ‰ – DIDS Û‚Â΢˂‡ÂÚ ‰ËÒÔÂÒ˲ SA-ÚÓ͇ Ë Á‡Ï‰ÎflÂÚ ÍËÌÂÚËÍÛ Â„Ó ‰Â‡Í-
ÚË‚‡ˆËË. ÇÂıÌË ԇÌÂÎË: ÙÎÛÍÚÛ‡ˆËË ÒÚ‡ˆËÓ̇ÌÓ„Ó SA-ÚÓ͇ ÔË 50 ÏÇ ‚ ÍÓÌÚÓΠ(΂‡fl Ô‡ÌÂθ) Ë ‚ ÔËÒÛÚÒÚ‚ËË 0.5 Ïå
DIDS (Ô‡‚‡fl Ô‡ÌÂθ). çËÊÌË ԇÌÂÎË: ı‚ÓÒÚÓ‚˚ ÚÓÍË ‚ ÓÚ‚ÂÚ Ì‡ ÂÔÓÎflËÁ‡ˆË˛ ÏÂÏ·‡Ì˚ ÓÚ 50 ‰Ó –70 ÏÇ ‚ ÍÓÌÚÓÎÂ
(΂‡fl Ô‡ÌÂθ) Ë ‚ ÔËÒÛÚÒÚ‚ËË DIDS (Ô‡‚‡fl Ô‡ÌÂθ) чÌÌ˚ ‚ ‰ ÌÓÏËÓ‚‡Ì˚ ̇ ‚Â΢ËÌÛ Ï‡ÍÒËχθÌÓ„Ó ÚÓ͇. ÇÓ ‚ÒÂı
ÒÎÛ˜‡flı ËÒÔÓθÁÓ‚‡ÎÒfl ÏÂÚÓ‰ whole-cell, Ô˝Ú˜-ÔËÔÂÚ͇ ÒÓ‰Âʇ· 140 Ïå CsCl.

̇θÌÓÈ ÔÓ˚ Ô‰ÔÓ·„‡ÂÚ Â ‚˚ÒÓÍÛ˛ ÔÓ‚Ó‰Ë- „‰Â D Ë I – ‰ËÒÔÂÒËfl Ë Ò‰Ìflfl ‚Â΢Ë̇ ËÌÚ„‡Î¸-
ÏÓÒÚ¸. àÒÔÓθÁÛfl ÌÂÒÚ‡ˆËÓ̇Ì˚È ÙÎÛÍÚÛ‡ˆËÓÌ- ÌÓ„Ó ÚÓ͇ ÒÓÓÚ‚ÂÚÒÚ‚ÂÌÌÓ, ÔÓÎÛ˜ÂÌÌ˚ ÛÒ‰ÌÂÌË-
Ì˚È ‡Ì‡ÎËÁ [14] ı‚ÓÒÚÓ‚˚ı ÚÓÍÓ‚, „ËÒÚËÛÂÏ˚ı ÂÏ ÔÓ ‡ÌÒ‡Ï·Î˛; i Ë N – ÚÓÍ ˜ÂÂÁ Ó‰ËÌÓ˜Ì˚È Í‡Ì‡Î
‚ „‡‰ËÂÌÚ Csin/Naout, Ï˚ ÔÓÔ˚Ú‡ÎËÒ¸ ÓˆÂÌËÚ¸ ÔÓ- Ë ˜ËÒÎÓ Ú‡ÍËı ͇̇ÎÓ‚. Ç ÂÔÂÁÂÌÚ‡ÚË‚ÌÓÏ ˝ÍÒÔÂ-
‚Ó‰ËÏÓÒÚ¸ Ó‰ËÌÓ˜ÌÓ„Ó SA-͇̇· ̇ ÓÒÌÓ‚Â ÒΉÛ- ËÏÂÌÚÂ, Ô‰ÒÚ‡‚ÎÂÌÌÓÏ Ì‡ ËÒ. 4„, ‡ÔÔÓÍÒËχ-
˛˘Â„Ó ÒÓÓÚÌÓ¯ÂÌËfl: ˆËfl ‰ËÒÔÂÒËË SA-ÚÓ͇ Í‡Í ÙÛÌ͈ËË Ò‰ÌÂ„Ó SA-ÚÓ-
͇ ÔÓ ÏÂÚÓ‰Û Ì‡ËÏÂ̸¯Ëı Í‚‡‰‡ÚÓ‚ Ò ËÒÔÓθÁÓ‚‡ÌË-
2
D = iI – I N, (6) ÂÏ Û‡‚ÌÂÌËfl (6) ‰‡ÎÓ I = 10.8 ÔÄ ÔË ˝ÎÂÍÚÓ-

ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009

 çÂÒÂÎÂÍÚË‚Ì˚ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ ͇̇Î˚ 69

‡ ·
50 ÏÇ
–70 ÏÇ

àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ
3 –100 ÏÇ 2
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

1
0

NMDGClin / NMDGClin /
NaClout Na-„βÍÓ̇Úout 0
CsClin NMDGClin
–3

‚ „
3 5000
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

ÑËÒÔÂÒËfl ÚÓ͇, ÔÄ2

2

1 2500

–1 0
–100 –50 0 50 0 500 1000 1500
åÂÏ·‡ÌÌ˚È ÔÓÚÂ̈ˇÎ, ÏÇ ë‰ÌËÈ ÚÓÍ, ÔÄ

êËÒ. 4. SA-ÚÓÍË ‚ ‡ÁÌ˚ı ÛÒÎÓ‚Ëflı ‰Ë‡ÎËÁ‡. ‡ – SA-ÚÓÍË, „ËÒÚËÛÂÏ˚ ÓÚ Ó‰ÌÓÈ Ë ÚÓÈ Ê ÍÎÂÚÍË ÔË ËÓÌÌ˚ı „‡‰ËÂÌ-
Ú‡ı 140 Ïå NMDG-Clin/140 Ïå NaClout (΂‡fl Ô‡ÌÂθ) Ë ‚ 140 Ïå NMDG-Clin/140 Ïå Na-„βÍÓ̇Úout (Ô‡‚‡fl Ô‡ÌÂθ). · –
ë‰ÌË ‡ÏÔÎËÚÛ‰˚ ÒÚ‡ˆËÓ̇Ì˚ı SA-ÚÓÍÓ‚ ÔË 50 ÏÇ ‚ ÛÒÎÓ‚Ëflı 140 Ïå CsClin/140 Ïå NaClout (΂˚È ÒÚÓηˆ, 28 ÍÎÂ-
ÚÓÍ) Ë 140 Ïå NMDGClin/140 Ïå NaClout (Ô‡‚˚È ÒÚÓηˆ, 9 ÍÎÂÚÓÍ). чÌÌ˚ Ô‰ÒÚ‡‚ÎÂÌ˚ Í‡Í Ò‰Ì ± Òڇ̉‡Ú̇fl
ӯ˷͇. ‚ – ÇÓθÚ-‡ÏÔÂÌ˚ ı‡‡ÍÚÂËÒÚËÍË SA-ÚÓÍÓ‚, Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ‡. „ – ùÍÒÔÂËÏÂÌڇθ̇fl Á‡‚ËÒËÏÓÒÚ¸ ‰ËÒÔÂ-
ÒËË SA-ÚÓ͇ () ÓÚ Ò‰ÌÂÈ ‚Â΢ËÌ˚ ËÌÚ„‡Î¸ÌÓ„Ó ÚÓ͇, ÓˆÂÌÂÌ̇fl ÔÓ ı‚ÓÒÚÓ‚˚Ï ÚÓ͇Ï, „ÂÌÂËÛÂÏ˚Ï ‚ ÓÚ‚ÂÚ Ì‡ Â-
ÔÓÎflËÁ‡ˆË˛ ÍÎÂÚÓ˜ÌÓÈ ÏÂÏ·‡Ì˚ ÓÚ 30 ‰Ó –70 ÏÇ. çÂÔÂ˚‚̇fl ÍË‚‡fl ÒÓÓÚ‚ÂÚÒÚ‚ÛÂÚ Û‡‚ÌÂÌ˲ (6) ÔË N = 151 Ë i =
= 10.8 ÔÄ. Ç ‡–· ÚÓÍË „ËÒÚËÓ‚‡ÎËÒ¸ ÏÂÚÓ‰ÓÏ whole-cell. Ç „ ËÒÔÓθÁÓ‚‡ÎÒfl ÏÂÚÓ‰ perforated patch ÔË 140 Ïå CsCl ‚
Ô˝Ú˜-ÔËÔÂÚÍÂ Ë 140 Ïå NaCl ‚ ̇ÛÊÌÓÏ ‡ÒÚ‚ÓÂ.

‰‚ËÊÛ˘ÂÈ ÒËΠ(Vh – Vr) = 62.8 ÏÇ, ˜ÚÓ ˝Í‚Ë‚‡ÎÂÌÚ-
ÌÓ ÔÓ‚Ó‰ËÏÓÒÚË 171 ÔëÏ. àÒÔÓθÁÛfl ‰‡ÌÌ˚È ÔÓ‰-
ıÓ‰ ‰Îfl ‡Ì‡ÎËÁ‡ ı‚ÓÒÚÓ‚˚ı SA-ÚÓÍÓ‚ ‚ ¯ÂÒÚË ÍÎÂÚ-
͇ı, Ï˚ ÔÓÎÛ˜ËÎË, ˜ÚÓ ‚ Ò‰ÌÂÏ ÔÓ‚Ó‰ËÏÓÒÚ¸
Ó‰ËÌÓ˜ÌÓ„Ó SA-͇̇· ÒÓÒÚ‡‚ÎflÂÚ 178 ± 21 ÔëÏ.
í‡Í‡fl ‚Â΢Ë̇ ÔÓ‚Ó‰ËÏÓÒÚË ÔÓ˚ Ë Â Ò··‡fl ÒÂ-
ÎÂÍÚË‚ÌÓÒÚ¸ ‰Îfl ͇ÚËÓÌÓ‚ Ë ‡ÌËÓÌÓ‚ ı‡‡ÍÚÂÌ˚
‰Îfl ËÓÌÌ˚ı ͇̇ÎÓ‚, ÙÓÏËÛÂÏ˚ı ÍÓÌÌÂÍÒË̇ÏË
‚ Ô·ÁχÚ˘ÂÒÍÓÈ ÏÂÏ·‡Ì ÍÎÂÚÓÍ ÔÓÁ‚ÓÌÓ˜Ì˚ı
[15, 16]. Ç ˝ÚÓÈ Ò‚flÁË Ï˚ ËÒÒΉӂ‡ÎË ‰ÂÈÒÚ‚Ë fl-
‰‡ ‚¢ÂÒÚ‚, ÍÓÚÓ˚ ËÌ„Ë·ËÛ˛Ú ÍÓÌÌÂÍÒËÌÓ‚˚Â
ÔÓÎÛÔÓ˚.
à‰ÂÌÚËÙË͇ˆËfl ËÓÌÌ˚ı ͇̇ÎÓ‚, ÙÓÏËÛÂ-
Ï˚ı ÍÓÌÌÂÍÒË̇ÏË, ÍÓÚÓ˚Â Û ÏÎÂÍÓÔËÚ‡˛˘Ëı
ÍÓ‰ËÛ˛ÚÒfl ÔËÏÂÌÓ 20 „Â̇ÏË [17], ÓÒÎÓÊÌflÂÚÒfl
ÚÂÏ, ˜ÚÓ Ì ÒÛ˘ÂÒÚ‚ÛÂÚ Ëı ÒÔˆËÙ˘ÂÒÍËı Ë ÒÚÓ„Ó
ÒÂÎÂÍÚË‚Ì˚ı ·ÎÓ͇ÚÓÓ‚. ëÂ‰Ë ‚¢ÂÒÚ‚, ‚˚Á˚‚‡-
˛˘Ëı ·ÎÓ͇‰Û ÍÓÌÌÂÍÒËÌÓ‚˚ı ͇̇ÎÓ‚, Ï˚ ËÒÔÓθ-
ÁÓ‚‡ÎË ÌÂÍÓÚÓ˚ ÏËÏÂÚ˘ÂÒÍË ÔÂÔÚˉ˚ [18], ͇-
·ÂÌÓÍÒÓÎÓÌ Ë ÓÍÚ‡ÌÓÎ. é͇Á‡ÎÓÒ¸, ˜ÚÓ Î˯¸ ÓÍÚ‡-
ÌÓÎ Ë ÔÂÔÚˉ 43GAP26 ‚˚Á˚‚‡˛Ú ÓÚÌÓÒËÚÂθÌÓ
ÒÔˆËÙ˘ÂÒÍÓÂ Ë ÒÛ˘ÂÒÚ‚ÂÌÌÓ ·ÎÓÍËÓ‚‡ÌË SA-
ÚÓÍÓ‚ (ËÒ. 5‡, ·). àÌ„Ë·ËÛ˛˘Â ‰ÂÈÒÚ‚Ë ˝ÚËı ‚Â-
˘ÂÒÚ‚ ҂ˉÂÚÂθÒÚ‚ÛÂÚ ‚ ÔÓθÁÛ ÍÓÌÌÂÍÒËÌÓ‚ÓÈ
ÔËÓ‰˚ SA-͇̇ÎÓ‚, ÙÛÌ͈ËÓÌËÛ˛˘Ëı ‚Ó ‚ÍÛÒÓ-
‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II. àÌÚÂÂÒÌÓ ÓÚÏÂÚËÚ¸, ˜ÚÓ ‚ÌÂ-
ÍÎÂÚӘ̇fl ‡ÔÔÎË͇ˆËfl ÒËÌÚÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎÓ„‡ ‰Ë-
‡ˆË΄ÎˈÂÓ· DOG ËÎË ‰Ë‡ÎËÁ ÍÎÂÚÓÍ ‡ÒÚ‚Ó-
ÓÏ, Ì ÒÓ‰Âʇ˘ËÏ ATP, ‚˚Á˚‚‡Î Ô‡ÍÚ˘ÂÒÍË
ÔÓÎÌÓÂ Ë ÌÂÓ·‡ÚËÏÓ ËÌ„Ë·ËÓ‚‡ÌË SA-ÚÓÍÓ‚ ‚
Ú˜ÂÌË 5–10 ÏËÌ (ËÒ. 5‚). ùÚÓ Ò‚Ë‰ÂÚÂθÒÚ‚Ó‚‡ÎÓ
Ó ÚÓÏ, ˜ÚÓ Í‡Ì‡Î˚, Ó·ÂÒÔ˜˂‡˛˘Ë SA-ÚÓÍ (SA-͇-

ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009

70 ÓχÌÓ‚ Ë ‰.

‡ · ‚
50 ÏÇ 50 ÏÇ
–70 ÏÇ –70 ÏÇ 1.0

çÓÏËÓ‚‡ÌÌ˚È ÚÓÍ
äÓÌÚÓθ
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

2 äÓÌÚÓθ 4

0.5
éÍÚ‡ÌÓÎ
0 43GAP26
0

0
–2 –4 0 5 10 15
ÇÂÏfl, ÏËÌ

„ ‰
50 ÏÇ
200

àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ
–50 ÏÇ
–100 ÏÇ
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

200 100

100
0
0

–100 –100
50 Ïå Mg2Äíê 12.5 Ïå Mg2Äíê –80 –40 0 40
–200 åÂÏ·‡ÌÌ˚È ÔÓÚÂ̈ˇÎ, ÏÇ

êËÒ. 5. SA-ÚÓÍË Ú‡ÌÒÔÓÚËÛ˛ÚÒfl ÍÓÌÌÂÍÒËÌÓÔÓ‰Ó·Ì˚ÏË Í‡Ì‡Î‡ÏË. ‡ – å‰ÎÂÌÌÓ (10 ÏËÌ) ËÌ„Ë·ËÛ˛˘Â ‰ÂÈÒÚ‚ËÂ
ÔÂÔÚˉ‡ 43GAP26 (0.5 Ïå) ̇ SA-ÚÓÍ, „ËÒÚËÛÂÏ˚È ‚ ÓÚ‚ÂÚ Ì‡ 100 ÏÒ ‰ÂÔÓÎflËÁ‡ˆË˛ ÍÎÂÚÍË ‰Ó 50 ÏÇ. · – àÌ„Ë·ËÓ-
‚‡ÌË SA-ÚÓ͇, „ËÒÚËÛÂÏÓ„Ó ÔË 50 ÏÇ, ‚ ÔËÒÛÚÒÚ‚ËË 3 Ïå ÓÍÚ‡ÌÓ·, ·ÎÓÍËÛ˛˘Â ‰ÂÈÒÚ‚Ë ÍÓÚÓÓ„Ó ‡Á‚Ë‚‡ÂÚÒfl
ÏÂÌ ˜ÂÏ Á‡ 1 ÏËÌ. íÓÍË ‚ · „ËÒÚËÓ‚‡ÎËÒ¸ ‚ ÓÚ‚ÂÚ Ì‡ ‰ÂÔÓÎflËÁ‡ˆË˛ ÍÎÂÚÍË ‰Ó 50 ÏÇ. Ç ‡ Ë · ËÒÔÓθÁÓ‚‡ÎËÒ¸ ‡ÁÌ˚Â
ÍÎÂÚÍË. ‚ – ÇÂÏÂÌÌÓÈ ıÓ‰ ËÌ„Ë·ËÓ‚‡ÌËfl SA-ÚÓÍÓ‚ ‚ ÔËÒÛÚÒÚ‚ËË ÒËÌÚÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎÓ„‡ ‰Ë‡ˆË΄ÎˈÂË̇ DOG () Ë
‚ ÛÒÎÓ‚Ëflı ‰Ë‡ÎËÁ‡ ‚ÍÛÒÓ‚˚ı ÍÎÂÚÓÍ ‡ÒÚ‚ÓÓÏ, Ì ÒÓ‰Âʇ˘ËÏ ATP (). „ – SA-ÚÓÍË, „ËÒÚËÛÂÏ˚ ÓÚ ‚ÍÛÒÓ‚ÓÈ ÍÎÂÚ-
ÍË ‚ ÛÒÎÓ‚Ëflı 50 Ïå Mg2ATPout/50 Ïå Mg2ATPin (΂‡fl Ô‡ÌÂθ) Ë 12.5 Ïå Mg2ATPout/50 Ïå Mg2ATPin (Ô‡‚‡fl Ô‡ÌÂθ).
‰ – ÇÓθÚ-‡ÏÔÂÌ˚ ı‡‡ÍÚÂËÒÚËÍË ÚÓÍÓ‚, Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ „, ÒËÏ‚ÓÎ˚ ÒÓÓÚ‚ÂÚÒÚ‚Û˛Ú ÏÓÏÂÌÚÛ ËÁÏÂÂÌËfl ÒÚ‡ˆËÓ̇-
ÌÓ„Ó SA-ÚÓ͇. Ç ‡, · Ë ‚ () ÚÓÍË „ËÒÚËÓ‚‡ÎËÒ¸ ÏÂÚÓ‰ÓÏ perforated patch ÔË 140 Ïå CsCl ‚ Ô˝Ú˜-ÔËÔÂÚÍÂ Ë 140 Ïå
NaCl ‚ ̇ÛÊÌÓÏ ‡ÒÚ‚ÓÂ. Ç „ Ë ‚ () ËÒÔÓθÁÓ‚‡ÎÒfl ÏÂÚÓ‰ whole-cell.

̇Î˚), ̇ıÓ‰flÚÒfl ÔÓ‰ ÊÂÒÚÍËÏ ÏÂÚ‡·Ó΢ÂÒÍËÏ
ÍÓÌÚÓÎÂÏ.
ÑÎfl ˘Â΂˚ı ÍÓÌÚ‡ÍÚÓ‚ Ë ÔÓÎÛÔÓ, ÙÓÏËÛÂ-
Ï˚ı ÍÓÌÌÂÍÒË̇ÏË, ı‡‡ÍÚÂÌÓ ÚÓ, ˜ÚÓ ÓÌË ÔÓÌË-
ˆ‡ÂÏ˚ ‰Îfl ÏÓÎÂÍÛÎ ‚ÂÒÓÏ ‰Ó 1 Íч, ‚Íβ˜‡fl Ú‡ÍËÂ,
Í‡Í Äíê, ÒÄåê, IP3, Lucifer Yellow Ë fl‰ ‰Û„Ëı
[15, 16]. èÓ˝ÚÓÏÛ ‚ fl‰Â ˝ÍÒÔÂËÏÂÌÚÓ‚ Ï˚ ‚˚flÒ-
ÌflÎË, ÔÓÌˈ‡ÂÏ˚ ÎË SA-͇̇Î˚ ‰Îfl ÍÛÔÌ˚ı ÏÓ-
ÎÂÍÛÎ, ‚ ˜‡ÒÚÌÓÒÚË ATP. Ç ÛÒÎÓ‚Ëflı, ÍÓ„‰‡ ‚ÌÛÚË-
ÍÎÂÚÓ˜Ì˚È Ë ‚ÌÂÍÎÂÚÓ˜Ì˚È ‡ÒÚ‚Ó˚ ÒÓ‰ÂʇÎË
Î˯¸ 50 Ïå MgATP + 50 Ïå Mg(OH)2, ÔË ‰ÂÔÓÎfl-
ËÁ‡ˆËË „ËÒÚËÓ‚‡ÎËÒ¸ ‚˚ıÓ‰fl˘Ë SA-ÚÓÍË ‚Â-
΢ËÌÓÈ ‰Ó ÌÂÒÍÓθÍËı ÒÓÚÂÌ ÔËÍÓ‡ÏÔÂ (ËÒ. 5„,
΂‡fl Ô‡ÌÂθ). ùÚÓ Ò‚Ë‰ÂÚÂθÒÚ‚Ó‚‡ÎÓ Ó ÚÓÏ, ˜ÚÓ
SA-͇̇Î˚ ÔÓÌˈ‡ÂÏ˚ ‰Îfl Mg2+ Ë/ËÎË MgATP2–,
Ú‡Í Í‡Í ÔË ÌÂÈÚ‡Î¸Ì˚ı Á̇˜ÂÌËflı pH ‰Û„Ë Á‡-
flÊÂÌÌ˚ ÙÓÏ˚ Äíê ÔËÒÛÚÒÚ‚Û˛Ú Î˯¸ ‚ ÒÎÂ-
‰Ó‚˚ı ÍÓ΢ÂÒÚ‚‡ı. í‡Í, Òӄ·ÒÌÓ Ì‡¯ËÏ ‡Ò˜Â-
Ú‡Ï, ‚ ‡ÒÚ‚ÓÂ, ÒÓ‰Âʇ˘ÂÏ 50 Ïå MgATP + 50 Ïå
Mg(OH)2, ÔË pH 7.4 Ë 25°C ÔËÒÛÚÒÚ‚Û˛Ú 21.0 Ïå
MgATP2–, 21.1 Ïå Mg2+ Ë 28.9 Ïå Mg2ATP. ê‡Á·‡‚-
ÎÂÌË ‚ÌÂÍÎÂÚÓ˜ÌÓ„Ó ‡ÒÚ‚Ó‡ ‚ 4 ‡Á‡ ‰Ó 12.5 Ïå
MgATP + 12.5 Ïå Mg(OH)2 (ÒÓÓÚ‚ÂÚÒÚ‚ÂÌÌÓ ÍÓ̈ÂÌ-
Ú‡ˆËË ‡Á΢Ì˚ı ÙÓÏ: 7.04 Ïå MgATP2–, 7.1 Ïå
Mg2+ Ë 5.42 Ïå Mg2ATP) ÔË‚Ó‰ËÎÓ Í ÒÌËÊÂÌ˲
‡ÏÔÎËÚÛ‰˚ ‚˚ıÓ‰fl˘Â„Ó ÚÓ͇ Ë ÒÏ¢ÂÌ˲ ̇ „‡-
ÙË͇ı I–V ı‡‡ÍÚÂËÒÚËÍ ÔÓÚÂ̈ˇ· ‚ÂÒËË
‚Ô‡‚Ó Ì‡ 3.8 ± 0.8 (n = 4) (ËÒ. 5„, ‰). ùÚÓ Û͇Á˚‚‡ÂÚ
̇ ÚÓ, ˜ÚÓ SA-͇̇Î˚ ÔËÏÂÌÓ ‚ ‡‚ÌÓÈ ÒÚÂÔÂÌË
ÔÓÌˈ‡ÂÏ˚ ‰Îfl Mg2+ Ë MgATP2–.
Ç fl‰Â ÒÎÛ˜‡Â‚ Ï˚ ÓˆÂÌË‚‡ÎË ÔÓÌˈ‡ÂÏÓÒÚ¸
SA-͇̇ÎÓ‚ ‰Îfl ÌÂÍÓÚÓ˚ı ÙÎÛÓÂÒˆÂÌÚÌ˚ı Í‡-
ÒËÚÂÎÂÈ, Ú‡ÌÒÔÓÚËÛÂÏ˚ı ÍÓÌÌÂÍÒËÌÓ‚˚ÏË Í‡-
̇·ÏË. ç‡ ËÒ. 6 Ô‰ÒÚ‡‚ÎÂÌ Ó‰ËÌ ËÁ Ú‡ÍËı ˝ÍÒÔÂ-
ËÏÂÌÚÓ‚ (n = 6). àÌÍÛ·‡ˆËfl (‚ÔÎÓÚ¸ ‰Ó 10 ÏËÌ)
ÍÎÂÚÍË ÚËÔ‡ II (ËÒ. 6, ËÁÓ·‡ÊÂÌË 1) ‚ ‡ÒÚ‚Ó Ò
ÓÚˈ‡ÚÂθÌÓ Á‡flÊÂÌÌ˚Ï Í‡ÒËÚÂÎÂÏ ÙÎÛÓÂÒ-
ˆÂËÌÓÏ ÔË ÔÓ‰‰ÂÊË‚‡ÂÏÓÏ ÔÓÚÂ̈ˇÎ Vh = –70 ÏÇ

ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009

 çÂÒÂÎÂÍÚË‚Ì˚ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ ͇̇Î˚ 71

‡ 50 ÏÇ
èÓÚÂ̈ˇΠ–70 ÏÇ
10 c
íÓÍ 2 ÌÄ

0.05 Ï„/ÏÎ
îÎÛÓÂÒˆÂËÌ 0 Ï„/ÏÎ
1 2 3
·

1 2 3

êËÒ. 6. ÑÂÔÓÎflËÁ‡ˆËfl ‚ÍÛÒÓ‚˚ı ÍÎÂÚÓÍ ÚËÔ‡ II ÒÚËÏÛÎËÛÂÚ Ì‡ÍÓÔÎÂÌË ÙÎÛÓÂÒˆÂË̇. ‡ – ëıÂχ ˝ÍÒÔÂËÏÂÌÚ‡ ÔÓ Á‡-
„ÛÁÍ ‚ ÍÎÂÚÍÛ ÙÎÛÓÂÒˆÂË̇. ÇÂıÌflfl Ô‡ÌÂθ – ‚ÂÏÂÌÌÓÈ ıÓ‰ ÔÓÚÂ̈ˇ·, ÔÓ‰‰ÂÊË‚‡ÂÏÓ„Ó Ì‡ ÏÂÏ·‡Ì ‚ÍÛÒÓ‚ÓÈ
ÍÎÂÚÍË. ë‰Ìflfl Ô‡ÌÂθ – „ËÒÚËÛÂÏ˚È ËÌÚ„‡Î¸Ì˚È ÚÓÍ ‚ÍÛÒÓ‚ÓÈ ÍÎÂÚÍË. çËÊÌflfl Ô‡ÌÂθ – ËÁÏÂÌÂÌË ÍÓ̈ÂÌÚ‡-
ˆËË ÙÎÛÓÂÒˆÂË̇ ‚Ó ‚ÌÂÍÎÂÚÓ˜ÌÓÏ ‡ÒÚ‚ÓÂ; ˆËÙ˚ ‚ÌËÁÛ ÒÓÓÚ‚ÂÚÒÚ‚Û˛Ú ÏÓÏÂÌÚ‡Ï ‚ÂÏÂÌË ÔÓÎÛ˜ÂÌËfl ËÁÓ·‡ÊÂÌËÈ
ËÒÒΉÛÂÏÓÈ ÍÎÂÚÍË, Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ·. · – àÁÓ·‡ÊÂÌËfl ÍÎÂÚÍË ‚ ÔÓıÓ‰fl˘ÂÏ (1) Ë ÙÎÛÓÂÒˆÂÌÚÌÓÏ (2, 3) Ò‚ÂÚÂ. ë‚Â-
˜ÂÌË ÙÎÛÓÂÒˆÂË̇ ‚ÓÁ·Ûʉ‡ÎÓÒ¸ Ò‚ÂÚÓÏ ‚ ‰Ë‡Ô‡ÁÓÌ 480 ± 5 ÌÏ, ˝ÏËÒÒËfl „ËÒÚËÓ‚‡Î‡Ò¸ Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ ÓÔÚ˘Â-
ÒÍÓ„Ó ÙËθÚ‡ 515 ± 10 ÌÏ.

Ì Ô˂Ӊ˷ Í ÒÍÓθÍÓ-ÌË·Û‰¸ Á‡ÏÂÚÌÓÏÛ ËÁÏÂÌÂ- ΠÓÚˈ‡ÚÂθÌ˚ı ÔÓÚÂ̈ˇÎÓ‚, Û‚Â΢˂‡Î‡Ò¸

Ì˲ ÙÎÛÓÂÒˆÂ̈ËË ÍÎÂÚÍË (ËÒ. 6·, ËÁÓ·‡ÊÂÌË 2).
Ç ÚÓ Ê ‚ÂÏfl Í‡ÚÍÓ‚ÂÏÂÌ̇fl ‰ÂÔÓÎflËÁ‡ˆËfl
ÍÎÂÚÍË, ÒÚËÏÛÎËÛ˛˘‡fl SA-ÚÓÍË (ËÒ. 6‡), ‚˚Á˚-
‚‡Î‡ ·˚ÒÚÛ˛ ‡ÍÍÛÏÛÎflˆË˛ ‚ÌÂÍÎÂÚÓ˜ÌÓ„Ó ÙÎÛÓ-
ÂÒˆÂË̇ ‚ ÍÎÂÚÍÂ, ˜ÚÓ ‚˚‡Ê‡ÎÓÒ¸ ‚ Á̇˜ËÚÂθ-
ÌÓÏ Û‚Â΢ÂÌËË Â ÙÎÛÓÂÒˆÂ̈ËË (ËÒ. 6·, ËÁÓ·-
‡ÊÂÌË 3). èÓ ‚ÒÂÈ ‚ˉËÏÓÒÚË, ÙÎÛÓÂÒˆÂËÌ
ÔÓÌË͇Π‚ ÍÎÂÚÍÛ ˜ÂÂÁ SA-͇̇Î˚, ‡ Ì ˜ÂÂÁ ͇-
ÍËÂ-ÎË·Ó ËÌ˚ Ú‡ÌÒÔÓÚÌ˚ ÒÚÛÍÚÛ˚, ÔÓÒÍÓθ-
ÍÛ Ì‡ÍÓÔÎÂÌË Í‡ÒËÚÂÎfl Ô‡ÍÚ˘ÂÒÍË Ì ̇·Î˛-
‰‡ÎÓÒ¸ ‚ ÍÓÌÚÓθÌ˚ı ÍÎÂÚ͇ı, ‚ ÍÓÚÓ˚ı SA-͇-
̇Î˚ ·˚ÎË Á‡·ÎÓÍËÓ‚‡Ì˚ ÔÓÒΠËÌÍÛ·‡ˆËË ‚
‡ÒÚ‚ÓÂ, ÒÓ‰Âʇ˘ÂÏ DOG (n = 3).
֢ ӉÌÓÈ ı‡‡ÍÚÂÌÓÈ ÓÒÓ·ÂÌÌÓÒÚ¸˛ ÍÓÌÌÂÍ-
ÒËÌÓ‚˚ı ͇̇ÎÓ‚ fl‚ÎflÂÚÒfl Ëı ˜Û‚ÒÚ‚ËÚÂθÌÓÒÚ¸ Í
ë‡2+ Ë ‰Û„ËÏ ‰‚Ûı‚‡ÎÂÌÚÌ˚Ï Í‡ÚËÓÌ‡Ï [19, 20]. Ç
˜‡ÒÚÌÓÒÚË, ÒÌËÊÂÌË ‚ÌÂÍÎÂÚÓ˜ÌÓÈ ÍÓ̈ÂÌÚ‡ˆËË
ë‡2+ Ó·˚˜ÌÓ ‡ÍÚË‚ËÛÂÚ ÍÓÌÌÂÍÒËÌÓ‚˚ ͇̇Î˚, Ë
˝ÚÓÚ ˝ÙÙÂÍÚ ˜‡ÒÚÓ ËÒÔÓθÁÛÂÚÒfl ‚ ÙËÁËÓÎӄ˘Â-
ÒÍËı ˝ÍÒÔÂËÏÂÌÚ‡ı ÔÓ ËÁÛ˜ÂÌ˲ ÓÎË ÍÓÌÌÂÍÒË-
ÌÓ‚ ‚ ÙÛÌ͈ËÓÌËÓ‚‡ÌËË ÍÎÂÚÓÍ [19, 20]. Ç ˝ÚÓÈ
Ò‚flÁË Ï˚ ËÒÒΉӂ‡ÎË ‚ÎËflÌË ‚ÌÂÍÎÂÚÓ˜Ì˚ı ‰‚Ûı-
‚‡ÎÂÌÚÌ˚ı ͇ÚËÓÌÓ‚ ̇ SA-ÚÓÍË.
àÁÏÂÌÂÌË ÍÓ̈ÂÌÚ‡ˆËË ‚ÌÂÍÎÂÚÓ˜Ì˚ı Mg2+ Ë
Ca2+ ÏÓ‰ËÙˈËÓ‚‡ÎÓ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏÓÒÚ¸
ÔÓ‚Ó‰ËÏÓÒÚË SA-͇̇ÎÓ‚. èË ÔÓÌËÊÂÌËË ÍÓÌ-
ˆÂÌÚ‡ˆËË Mg2+ Ë Ca2+ ‚Ó ‚ÌÂÍÎÂÚÓ˜ÌÓÏ ‡ÒÚ‚ÓÂ
ÔÓÓ„ ‡ÍÚË‚‡ˆËË SA-ÚÓÍÓ‚ ÒÏ¢‡ÎÒfl ‚ ӷ·ÒÚ¸ ·Ó-
χÍÒËχθ̇fl ÔÓ‚Ó‰ËÏÓÒÚ¸, ‡ÍÚË‚‡ˆËfl ÚÓÍÓ‚
ÛÒÍÓfl·Ҹ, ‡ ‰Â‡ÍÚË‚‡ˆËfl ÚÓÍÓ‚ ÔË ÂÔÓÎflËÁ‡-
ˆËË, ̇ÔÓÚË‚, Á‡Ï‰Îfl·Ҹ (ËÒ. 7‡,·). èÓ‚˚¯Â-
ÌË ‚ÌÂÍÎÂÚÓ˜ÌÓÈ ÍÓ̈ÂÌÚ‡ˆËË Ca2+ ‰Ó 10 Ïå
Ó͇Á˚‚‡ÎÓ ÔÓÚË‚ÓÔÓÎÓÊÌÓ ‰ÂÈÒÚ‚ËÂ: Á‡‚ËÒË-
ÏÓÒÚ¸ ÔÓ‚Ó‰ËÏÓÒÚË ÓÚ ÔÓÚÂ̈ˇ· ÒÛ˘ÂÒÚ‚ÂÌÌÓ
ÒÏ¢‡Î‡Ò¸ ‚Ô‡‚Ó, ‡ÍÚË‚‡ˆËfl ÚÓ͇ Á‡Ï‰Îfl·Ҹ, ‡
‰Â‡ÍÚË‚‡ˆËfl ÚÓÍÓ‚ ÔË ÂÔÓÎflËÁ‡ˆËË ÛÒÍÓfl·Ҹ
(ËÒ. 7‚,„). ÄÔÔÎË͇ˆËfl 10 Ïå Ba2+ Á̇˜ËÚÂθÌÓ
ÔÓ‰‡‚Îfl· ‚˚ıÓ‰fl˘Ë SA-ÚÓÍË (ËÒ. 7‰) Á‡ Ò˜ÂÚ ÒÛ-
˘ÂÒÚ‚ÂÌÌÓ„Ó Ò‰‚Ë„‡ Ëı ‡ÍÚË‚‡ˆËÓÌÌÓÈ ÍË‚ÓÈ ‚
ÒÚÓÓÌÛ ÔÓÎÓÊËÚÂθÌ˚ı ÔÓÚÂ̈ˇÎÓ‚ (ËÒ. 7Â).
éÚÏÂÚËÏ, ˜ÚÓ Í‡ÚÍÓ‚ÂÏÂÌ̇fl ËÌÍÛ·‡ˆËfl ÍÎÂÚÓÍ
‚ ÔËÒÛÚÒÚ‚ËË 10 Ïå Ba2+ ‚˚Á˚‚‡Î‡ Ó·‡ÚËÏÓÂ
ÛÏÂ̸¯ÂÌË SA-ÚÓÍÓ‚, ÍÓÚÓ˚ ËÌ„Ë·ËÓ‚‡ÎËÒ¸
ÌÂÓ·‡ÚËÏÓ ÔË ‡ÔÔÎË͇ˆËflı Ba2+ ‰ÎËÚÂθÌÓÒÚ¸˛
5 ÏËÌ Ë ·ÓÎÂÂ.
ùÙÙÂÍÚ˚, ÓÔËÒ‡ÌÌ˚ ‚˚¯Â (ËÒ. 7), ÏÓ„ÎË
·˚Ú¸ ‚ Á̇˜ËÚÂθÌÓÈ ÒÚÂÔÂÌË Ó·ÛÒÎÓ‚ÎÂÌ˚ ËÁÏÂ-
ÌÂÌËflÏË ÔÓ‚ÂıÌÓÒÚÌÓ„Ó ÔÓÚÂ̈ˇ· ÏÂÏ·‡Ì˚,
ËÌˈËËÛÂÏ˚ÏË ËÁÏÂÌÂÌËÂÏ ‚ÌÂÍÎÂÚÓ˜ÌÓÈ ÍÓÌ-
ˆÂÌÚ‡ˆËË ‰‚Ûı‚‡ÎÂÌÚÌ˚ı ͇ÚËÓÌÓ‚. ç‡ÔËÏÂ,
ÔË Û‚Â΢ÂÌËË ÍÓ̈ÂÌÚ‡ˆËË ë‡2+ ‚ ̇ÛÊÌÓÈ
Ò‰ Ú‡ÌÒÏÂÏ·‡ÌÌÓ ˝ÎÂÍÚ˘ÂÒÍÓ ÔÓΠۂÂ-
΢˂‡ÂÚÒfl, ˜ÚÓ ˝Í‚Ë‚‡ÎÂÌÚÌÓ „ËÔÂÔÓÎflËÁ‡ˆËË
ÍÎÂÚÍË. ëÓÓÚ‚ÂÚÒÚ‚ÂÌÌÓ, ‡ÍÚË‚‡ˆËÓÌÌ˚ ÍË‚˚Â
ÚÓ͇ Ë Â„Ó ÔÓÚÂ̈ˇΠ‚ÂÒËË Ì‡ ÓÒË ÍÓχ̉Ì˚ı
ÔÓÚÂ̈ˇÎÓ‚ ‰ÓÎÊÌ˚ ·˚Ú¸ Ò‰‚ËÌÛÚ˚ ‚Ô‡‚Ó. Ç

ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009

72 ÓχÌÓ‚ Ë ‰.

‡ ·
50 ÏÇ
50
–70 ÏÇ
–100 ÏÇ
3 25

0
1 Ïå Ca2+ 100 Ìå Ca2+ 0
1 Ïå Mg2+ 0 Mg2+ –50 0 50
–3

‚ „ 40

èÓ‚Ó‰ËÏÓÒÚ¸, ÌëÏ
3
àÌÚ„‡Î¸Ì˚È ÚÓÍ, ÌÄ

20

0
1 Ïå Ca2+ 10 Ïå Ca2+
1 Ïå Mg2+ 1 Ïå Mg2+ 0
–50 0 50
–3

50
‰ Â

3
25

äÓÌÚÓθ + 10 Ïå Ça2+
0
–3 –50 0 50
åÂÏ·‡ÌÌ˚È ÔÓÚÂ̈ˇÎ, ÏÇ

êËÒ. 7. ᇂËÒËÏÓÒÚ¸ SA-ÚÓÍÓ‚ ÓÚ ÒÓ‰ÂʇÌËfl ‰‚Ûı‚‡ÎÂÌÚÌ˚ı ͇ÚËÓÌÓ‚ ‚ ̇ÛÊÌÓÏ ‡ÒÚ‚ÓÂ. ‡ – êÂÔÂÁÂÌÚ‡ÚË‚ÌÓ ÒÂ-
ÏÂÈÒÚ‚Ó ËÌÚ„‡Î¸Ì˚ı ÚÓÍÓ‚, „ËÒÚËÛÂÏ˚ı ‚ ÓÚ‚ÂÚ Ì‡ 100 ÏÒ ÔÓÎflËÁ‡ˆË˛ ÓÚ –100 ‰Ó 50 ÏÇ ÔË ÙËÁËÓÎӄ˘ÂÒÍËı
ÍÓ̈ÂÌÚ‡ˆËflı (1 Ïå) Ca2+ Ë Mg2+ (΂‡fl Ô‡ÌÂθ) Ë ÔË ÔÓÌËÊÂÌÌÓÈ ÍÓ̈ÂÌÚ‡ˆËË ‚ÌÂÍÎÂÚÓ˜Ì˚ı ‰‚Ûı‚‡ÎÂÌÚÌ˚ı ͇ÚË-
ÓÌÓ‚ (100 Ìå Ca2+ + 0 Ïå Mg2+) (Ô‡‚‡fl Ô‡ÌÂθ). · – ëÚ‡ˆËÓ̇̇fl ÔÓ‚Ó‰ËÏÓÒÚ¸ ÚÓÈ Ê ҇ÏÓÈ ÍÎÂÚÍË, ˜ÚÓ Ë ‚ ‡, ‚˚˜ËÒ-
ÎÂÌ̇fl Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ Û‡‚ÌÂÌËfl (1) ‚ ‡ÁÌ˚ı ÛÒÎÓ‚Ëflı: 1 Ïå Ca2+ + 1 Ïå Mg2+ (), 1 Ïå Ca2+ + 0.1 Ïå Mg2+ (0), 100 Ìå
Ca2+ + 0 Ïå Mg2+ (). ‚ – ùÙÙÂÍÚ Û‚Â΢ÂÌËfl ‚ÌÂÍÎÂÚÓ˜ÌÓÈ ÍÓ̈ÂÌÚ‡ˆËË Ca2+ ‰Ó 10 Ïå ̇ ËÌÚ„‡Î¸Ì˚ ÚÓÍË ÔË ÌÂ-
ËÁÏÂÌÌÓÈ ÍÓ̈ÂÌÚ‡ˆËË Ì‡ÛÊÌÓ„Ó Mg2+ (1 Ïå). „ – èÓ‚Ó‰ËÏÓÒÚ¸ ÚÓÈ Ê ҇ÏÓÈ ÍÎÂÚÍË, ˜ÚÓ Ë ‚ ‚, ‚ ÍÓÌÚÓΠ(1 Ïå
Ca2+, ) Ë ‚ ÔËÒÛÚÒÚ‚ËË 10 Ïå Ca2+ (). ‰ – àÌÚ„‡Î¸Ì˚ ÚÓÍË, „ËÒÚËÛÂÏ˚ ‚ ÍÓÌÚÓΠ(1 Ïå Ca2+ + 1 Ïå Mg2+)
(΂‡fl Ô‡ÌÂθ) Ë ‚ ÔËÒÛÚÒÚ‚ËË 10 Ïå Ba2+ ‚ ·‡ÁÓ‚ÓÏ ‚ÌÂÍÎÂÚÓ˜ÌÓÏ ‡ÒÚ‚Ó (Ô‡‚‡fl Ô‡ÌÂθ).  – ëÚ‡ˆËÓ̇̇fl ÔÓ‚Ó-
‰ËÏÓÒÚ¸ ÚÓÈ Ê ҇ÏÓÈ ÍÎÂÚÍË, ˜ÚÓ Ë ‚ ‰, ‚ ÍÓÌÚÓΠ() Ë ‚ ÔËÒÛÚÒÚ‚ËË 10 Ïå Ba2+ ().

˜‡ÒÚÌÓÒÚË, Û‚Â΢ÂÌË ÍÓ̈ÂÌÚ‡ˆËË ë‡2+ Ò 1 ‰Ó 5 ÏÇ ‚ÎÂ‚Ó ÔË Û‚Â΢ÂÌËË ÍÓ̈ÂÌÚ‡ˆËË Ì‡ÛÊ-

10 Ïå ÒÏ¢‡ÎÓ I–V ı‡‡ÍÚÂËÒÚËÍÛ èá Na+-ÚÓ-
ÍÓ‚, „ËÒÚËÛÂÏ˚ı ‚ ÍÎÂÚ͇ı ÚËÔ‡ II, Ë ËÁÏÂflÂ-
Ï˚È ÔÓÚÂ̈ˇΠÌÛÎÂ‚Ó„Ó ÚÓ͇ ‚Ô‡‚Ó Ì‡ 10–15 ÏÇ
(Ì ÔÓ͇Á‡ÌÓ). é‰Ì‡ÍÓ ‚ ÒÎÛ˜‡Â SA-ÚÓÍÓ‚ ̇·Î˛-
‰‡ÎÒfl Ì·Óθ¯ÓÈ Ò‰‚Ë„ ÔÓÚÂ̈ˇ· ‚ÂÒËË Ì‡ 3–
ÌÓ„Ó Ca2+ ËÎË Ba2+. ùÚÓ Ò‚Ë‰ÂÚÂθÒÚ‚ÛÂÚ Ó ÚÓÏ, ˜ÚÓ
˝ÙÙÂÍÚ˚ ‰‚Ûı‚‡ÎÂÌÚÌ˚ı ͇ÚËÓÌÓ‚ ̇ SA-ÚÓÍ ÌÂ
Ò‚Ó‰flÚÒfl Í ËÁÏÂÌÂÌ˲ ÔÓ‚ÂıÌÓÒÚÌÓ„Ó ÔÓÚÂ̈ˇ·
Ë ˜ÚÓ ÒÂÎÂÍÚË‚ÌÓÒÚ¸ SA-͇̇ÎÓ‚ ÏÓÊÂÚ ÏÓ‰ÛÎËÓ-
‚‡Ú¸Òfl ‚ÌÂÍÎÂÚÓ˜Ì˚Ï ë‡2+.

ÅàéãéÉàóÖëäàÖ åÖåÅêÄçõ      ÚÓÏ 26      ‹ 1      2009

 çÂÒÂÎÂÍÚË‚Ì˚ ÔÓÚÂ̈ˇÎ-Á‡‚ËÒËÏ˚ ͇̇Î˚
éÅëìÜÑÖçàÖ êÖáìãúíÄíéÇ
чÌÌ˚Â, ËÁÎÓÊÂÌÌ˚ ‚˚¯Â, ҂ˉÂÚÂθÒÚ‚Û˛Ú Ó
ÚÓÏ, ˜ÚÓ SA-ÚÓÍË ‚Ó ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II ÔÂ-
ÂÌÓÒflÚÒfl ÌÂÒÂÎÂÍÚË‚Ì˚ÏË Í‡Ì‡Î‡ÏË Ó‰ÌÓ„Ó ÚËÔ‡,
ÍÓÚÓ˚ ÔÓÌˈ‡ÂÏ˚ Í‡Í ‰Îfl ͇ÚËÓÌÓ‚, Ú‡Í Ë ‰Îfl
‡ÌËÓÌÓ‚, ‚Íβ˜‡fl Ú‡ÍË ÍÛÔÌ˚ ÏÓÎÂÍÛÎ˚, ͇Í
ATP. èÓ Ò‚ÓËÏ ·ËÓÙËÁ˘ÂÒÍËÏ Ò‚ÓÈÒÚ‚‡Ï – ÌËÁ͇fl
ÒÂÎÂÍÚË‚ÌÓÒÚ¸, ‚˚ÒÓ͇fl ÔÓ‚Ó‰ËÏÓÒÚ¸ Ó‰ËÌÓ˜ÌÓ„Ó
͇̇·, ‚˚ÔflÏÎÂÌË ÔÓ ‚˚ıÓ‰fl˘ÂÏÛ ÚÓÍÛ, ω-
ÎÂÌ̇fl ÍËÌÂÚË͇ ‡ÍÚË‚‡ˆËË-‰Â‡ÍÚË‚‡ˆËË, Ò··‡fl
˜Û‚ÒÚ‚ËÚÂθÌÓÒÚ¸ Í ‡ÌËÓÌÌ˚Ï ·ÎÓ͇ÚÓ‡Ï Ë Á‡-
ÏÂÚ̇fl ˜Û‚ÒÚ‚ËÚÂθÌÓÒÚ¸ Í ‚ÌÂÍÎÂÚÓ˜ÌÓÈ ÍÓ̈ÂÌ-
Ú‡ˆËË ‰‚Ûı‚‡ÎÂÌÚÌ˚ı ͇ÚËÓÌÓ‚ – ˝ÚË Í‡Ì‡Î˚, ̇-
Á‚‡ÌÌ˚ ̇ÏË ‰Îfl Û‰Ó·ÒÚ‚‡ SA-͇̇·ÏË, ·ÓÎÂÂ
‚ÒÂ„Ó ÒıÓ‰Ì˚ Ò Í‡Ì‡Î‡ÏË, Ó·‡ÁÓ‚‡ÌÌ˚ÏË ÍÓÌÌÂÍ-
ÒË̇ÏË.
Ç ÔÓθÁÛ ˝ÚÓ„Ó Ô‰ÔÓÎÓÊÂÌËfl „Ó‚ÓËÚ Ë ÚÓÚ
Ù‡ÍÚ, ˜ÚÓ SA-͇̇Î˚ ËÌ„Ë·ËÓ‚‡ÎËÒ¸ ÏËÏÂÚ˘Â-
ÒÍËÏ ÔÂÔÚˉÓÏ 43GAP26, ˝ÙÙÂÍÚ ÍÓÚÓÓ„Ó ·˚Î
‚ÔÓÎÌ ÒÔˆËÙ˘ÂÌ: ÔÂÔÚˉ˚ GAP32 [8] Ë 10Px1
(ÌÂÓÔÛ·ÎËÍÓ‚‡ÌÌ˚ ‰‡ÌÌ˚Â), ËÌ„Ë·ËÛ˛˘Ë ÍÓÌ-
ÌÂÍÒËÌ ëı32 Ë Ô‡ÌÌÂÍÒËÌ Panx1, ÒÓÓÚ‚ÂÚÒÚ‚ÂÌÌÓ, ÌÂ
Ó͇Á˚‚‡ÎË ·ÎÓÍËÛ˛˘Â„Ó ‰ÂÈÒÚ‚Ëfl ̇ SA-ÚÓÍË.
èÂÔÚˉ 43GAP26 ·˚Î ÒÍÓÌÒÚÛËÓ‚‡Ì ̇ ÓÒÌÓ‚Â
‡ÏËÌÓÍËÒÎÓÚÌÓÈ ÔÓÒΉӂ‡ÚÂθÌÓÒÚË ‚ÚÓÓÈ ˝ÍÒ-
Ú‡ÍÎÂÚÓ˜ÌÓÈ ÔÂÚÎË ÍÓÌÌÂÍÒË̇ ëı43 [20]. ë˜ËÚ‡-
ÂÚÒfl, ˜ÚÓ ËÌ„Ë·ËÛ˛˘Â ‰ÂÈÒÚ‚Ë ÔÂÔÚˉ‡ 43GAP26
ÓÔÓÒ‰ÛÂÚÒfl ‰ÓÏÂÌÓÏ SHVR ‚ ‡ÏËÌÓÍËÒÎÓÚÌÓÈ
ÔÓÒΉӂ‡ÚÂθÌÓÒÚË ÍÓÌÌÂÍÒË̇ ëı43 [20]. é‰Ì‡ÍÓ
SHVR-ÏÓÚË‚ ËÏÂÂÚÒfl Ú‡ÍÊÂ Û ÍÓÌÌÂÍÒËÌÓ‚ ëı30.3,
ëı31.1, ëı32, ëı33, ëı45, ëı47, ÍÓÚÓ˚Â ÔÓ˝ÚÓÏÛ
Ú‡ÍÊ ÏÓ„ÛÚ ËÌ„Ë·ËÓ‚‡Ú¸Òfl ÔÂÔÚˉÓÏ 43GAP26,
ıÓÚfl, ̇ÒÍÓθÍÓ Ì‡Ï ËÁ‚ÂÒÚÌÓ, ˜Û‚ÒÚ‚ËÚÂθÌÓÒÚ¸
‚˚¯ÂÔÂ˜ËÒÎÂÌÌ˚ı ÍÓÌÌÂÍÒËÌÓ‚ Í ÔÂÔÚˉÛ
43GAP26 Ì ËÒÒΉӂ‡Î‡Ò¸. ÇÓ ‚ÍÛÒÓ‚ÓÈ Ú͇ÌË Ó·-
̇ÛÊÂÌ˚ Ô‡ÌÌÂÍÒËÌ 1 (Panx1) Ë fl‰ ÍÓÌÌÂÍÒËÌÓ‚
(ëx26, ëı30.3, ëı31.1, ëı33, ëı43) [8, 21–23], ÌÂÍÓ-
ÚÓ˚ ËÁ ÍÓÚÓ˚ı ÏÓ„ÛÚ ÙÓÏËÓ‚‡Ú¸ ÙÛÌ͈ËÓ-
̇θÌÓ ‡ÍÚË‚Ì˚ ATP-ÔÓÌˈ‡ÂÏ˚ ËÓÌÌ˚ ͇̇-
Î˚. ëÂ‰Ë ·ÂÎÍÓ‚ ˘Â΂˚ı ÍÓÌÚ‡ÍÚÓ‚, ˉÂÌÚËÙË-
ˆËÓ‚‡ÌÌ˚ı ‚Ó ‚ÍÛÒÓ‚ÓÈ Ú͇ÌË, Cx26 Ë Cx43, ‡
Ú‡ÍÊ Panx1, ÒÔÓÒÓ·Ì˚ Ó·‡ÁÓ‚˚‚‡Ú¸ ÙÛÌ͈ËÓ-
̇θÌ˚ ÔÓÎÛÔÓ˚, ÔÓÌˈ‡ÂÏ˚ ‰Îfl ATP [20, 24].
Ç ÚÓ Ê ‚ÂÏfl ÔÓÌˈ‡ÂÏÓÒÚ¸ Cx30.3, Cx31.1 Ë
Cx33 ‰Îfl Äíê Ì ·˚· ËÒÒΉӂ‡Ì‡ ‚ ÙËÁËÓÎӄ˘Â-
ÒÍËı ˝ÍÒÔÂËÏÂÌÚ‡ı.
֢ ӉÌËÏ ‡„ÛÏÂÌÚÓÏ ‚ ÔÓθÁÛ ÚÓ„Ó, ˜ÚÓ ÍÓÌ-
ÌÂÍÒËÌÓ‚˚ ͇̇Î˚ ‰ÂÈÒÚ‚ËÚÂθÌÓ ÙÛÌ͈ËÓÌËÛ-
˛Ú ‚Ó ‚ÍÛÒÓ‚˚ı ÍÎÂÚ͇ı ÚËÔ‡ II, fl‚ÎflÂÚÒfl ÚÓÚ Ù‡ÍÚ,
˜ÚÓ ˝ÚË ÍÎÂÚÍË ÒÔÓÒÓ·Ì˚ ‡ÍÍÛÏÛÎËÓ‚‡Ú¸ ÓÚˈ‡-
ÚÂθÌÓ Á‡flÊÂÌÌ˚ ÙÎÛÓÂÒˆÂÌÚÌ˚ Í‡ÒËÚÂÎË
[8], ‚Íβ˜‡fl ÙÎÛÓÂÒˆÂËÌ (ËÒ. 6). éÚÏÂÚËÏ ‰Îfl
Ò‡‚ÌÂÌËfl, ˜ÚÓ ÙÎÛÓÂÒˆÂËÌ Ì ÔÓÌË͇ÂÚ ˜ÂÂÁ
͇̇Î˚, Ó·‡ÁÓ‚‡ÌÌ˚ Panx1 [23].
í‡ÍËÏ Ó·‡ÁÓÏ, ‚ ÓÚ΢ˠÓÚ ·Óθ¯ËÌÒÚ‚‡ ËÒ-
ÒΉӂ‡ÌÌ˚ı ÍÎÂÚÓÍ, ‚Íβ˜‡fl ‚ÍÛÒÓ‚˚ ÍÎÂÚÍË
ÚËÔ‡ I Ë ÚËÔ‡ III, ‚˚ıÓ‰fl˘Ë èá ÚÓÍË ‚Ó ‚ÍÛÒÓ‚˚ı
ÍÎÂÚ͇ı ÚËÔ‡ II ÔÂËÏÛ˘ÂÒÚ‚ÂÌÌÓ Ó·ÛÒÎÓ‚ÎÂÌ˚ ‡Í-
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73
ÚË‚ÌÓÒÚ¸˛ Ì ä+-͇̇ÎÓ‚, ‡ ÌÂÒÂÎÂÍÚË‚Ì˚ı ͇̇-
ÎÓ‚, ÒÍÓ ‚ÒÂ„Ó ÙÓÏËÛÂÏ˚ı ͇̇θÌ˚ÏË ·ÂÎ-
͇ÏË ÍÓÌÌÂÍÒË̇ÏË. è˘ËÌ˚ ËÒÔÓθÁÓ‚‡ÌËfl ÌÂ-
ÒÂÎÂÍÚË‚Ì˚ı ͇̇ÎÓ‚ Ë Ëı Óθ ‚ ÙËÁËÓÎÓ„ËË
‚ÍÛÒÓ‚˚ı ÍÎÂÚÓÍ ÚËÔ‡ II Ô‰ÒÚÓËÚ ‚˚flÒÌËÚ¸ ‚
‰‡Î¸ÌÂȯËı ˝ÍÒÔÂËÏÂÌÚ‡ı.
ꇷÓÚ‡ ‚˚ÔÓÎÌÂ̇ ÔË ÔÓ‰‰ÂÊÍ èÓ„‡ÏÏ˚
èÂÁˉËÛχ êÄç “åÓÎÂÍÛÎfl̇fl Ë ÍÎÂÚӘ̇fl
·ËÓÎÓ„Ëfl”, „‡ÌÚ‡ êÓÒÒËÈÒÍÓ„Ó ÙÓ̉‡ ÙÛ̉‡ÏÂÌ-
ڇθÌ˚ı ËÒÒΉӂ‡ÌËÈ (‹ 08-04-00033-‡), èÓ-
„‡ÏÏ˚ èÂÁˉÂÌÚ‡ êî („‡ÌÚ˚ åä-1332.2008.4)
Ë îÓ̉‡ ÒÓ‰ÂÈÒÚ‚Ëfl ÓÚ˜ÂÒÚ‚ÂÌÌÓÈ Ì‡ÛÍÂ.
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Non-Selective Voltage-Gated Channels Operating in Type II Taste Cells

R. A. Romanov, N. V. Kabanova, S. L. Malkin, S. S. Kolesnikov
Institute of Cell Biophysics, Russian Academy of Sciences, 142290, Pushchino, Moscow oblast,
e-mail: cheglok@rambler.ru

In a variety of different cells, voltage-gated (VG) outward currents are largely carried by K+ ions. Surprisingly,
robust VG outward currents are still observed in taste cells of Type II under the conditions when K+-selective
channels are generally inhibited. Here we studied ion channels responsible for atypical VG outward currents in
Type II taste cells. As was found, these channels are weakly selective and are permeable for large molecules
such as NMDG, gluconate, and ATP. The estimate of nearly 200 pS was obtained for the single channel con-
ductance using the non-stationary fluctuation analysis. Altogether, the data obtained indicate that by their prop-
erties, the channels studied are mostly reminiscent of hemichannels that are formed by junctional proteins con-
nexins in the plasma membrane of vertebrate cells.

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