IAOeIIIAEUeIA AEAAeAIIss IAOE OE?A?IE

IINOEOOO AIIOIII? ii.
I.A.IAEEAAeIIA

EAIEss

IEAENAIAe?
AIAe?IEIAE*

OAeE 577.152.361

CO?OEOO?II-OOIEOeIIIAEUeII AEANOEAINOI

ICIOI?I EAOAEIOE*II? NOAIAeEIEOeI
Na+,E+-AO?-ace

I?E IAIA?AIIO?IIIEO AIEEAAO

03.00.04 — aiioiiiy

AAOI?AOA?AO

aeena?oaoei? ia caeiaoooy
iaoeiaiai nooiaiy

aeieoi?a
aiieiai/ieo iaoe

EE?A — 1999

Aeena?oaoei?th ? ?oeiien.

?iaioa aeeiiaia a eaai?aoi?i? o?ainii?oieo AOOac Iinoeoooo
aiioiii? ii.I.A.Iaeeaaeiia Iaoeiiiaeueii? aeaaeaii? iaoe Oe?a?ie

Ioioeieii iiiiaioe: aeieoi? aiieiai/ieo iaoe, noa?oee iaoeiaee
niia?iaioiee Iaeeoaaa

Ia?aa?eoa Einoyioeiiaia (caaiaeoaa/ aiaeaeieo iae?ioiii?
Iinoeoooo

o?c?ieia?? ?i. I.I.Aiaiiieueoey IAI Oe?a?ie);

aeieoi? aiieiai/ieo iaoe, noa?oee iaoeiaee niia?iaioiee Eioioaiei

Ieaenaiae? sseiae/ (ea?iaiee eaai?aoi?i? iieaeoey?ii? aaiaoeee

?inoeoooo aa?iioieiai? AII Oe?a?ie);

aeieoi? aiieiai/ieo iaoe, i?ioani? Onaothe Iao?i Aieiaeeie?iae/

(i?ioani? eaoaae?e aiioiii? i aiioaoiieiai? Iaoeiiiaeueiiai
aa?a?iiai

oiiaa?neoaoo Eaaiiaoo Iiiino?ia Oe?a?ie).

I?iaiaeia onoaiiaa — Ee?anueeee iaoeiiiaeueiee oiiaa?neoao ii.Oa?ana
Oaa/aiea,eao.a?io?i??

Caoeno aiaeaoaeaoueny «27 «aa?aniy 1999 ?ieo i 14 aiaeeii ia caniaeaiii
niaoeiaeiciaaii? a/aii? ?aaee Ae 26.240.01 a Iinoeoooi aiioiii? ii.
I.A.Iaeeaaeiia IAI Oe?a?ie ca aae?anith: 252601, i.Ee?a-30, aoe.
Eaiioiae/a,9.

C aeena?oaoei?th iiaeia iciaeiieoeny a aiaeiioaoei Iinoeoooo aiioiii?
?i. I.A. Iaeeaaeiia Iaoeiiiaeueii? aeaaeaii? iaoe Oe?a?ie.

Aaoi?aoa?ao ?icineaiee «26»na?iiy 1999 ?ieo.

A/aiee nae?aoa?

niaoeiaeiciaaii? a/aii? ?aaee
I.A.Ei?naiei

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeueiinoue oaie. Oa?iaio Na+,E+-AO?-aca (AO?-oinoiaiae?ieaca, EO
3.6.1.37), ?ioaa?aeueiee a?eie ieaciaoe/ieo iaia?ai ee?oei aoea??io,
cae?enith? ni?yaeaiee c a?ae?ie?cii AO? i?ioeiai?aaeaiee o?ainiaia?aiiee
ia?aiin Na+ oa E+ ? oei naiei caaacia/o? i?aeo?eiaiiy aeaeo?io?i?/iiai ?
iniioe/iiai a?aae??io?a iaeiiaaeaioieo ?ii?a, iaiao?aeieo aeey
ii?iaeueiiai ooieoe?iioaaiiy ee?oei.Oa?iaio a?ae?a?a? aieiaio ?ieue a
?aae?caoe?? aaaaoi/enaeueieo ee?oeiieo ooieoe?e oa i?ioean?a, ye?
caeaaeaoue a?ae ?nioaaiiy ?iiieo a?aae??io?a [Aieaeu?aa A.A.,Iaeueaoiia
A.E.,1985].

Ia a?aei?io a?ae ?ioeo AO?-ac ?-oeio Na+,E+-AO?-aca ii?aeio? a ?aeei?e
iieaeoe? ii?yae c o?ainii?oii-a?ae?ie?oe/iith ? ?aoeaioi?io ooieoe?th
[Anner B.M.,1985], niaoeeo?/ii aca?iiae?th/e c aeciaaiieie ?ia?a?oi?aie
?ineeiiiai iioiaeaeaiiy — na?oeaaeie ae?eiceaeaie, aai ?o aiaeiaaiieie
aiaeiaaie [Doris P.A.,1994; Martinka E.,1995; McDonough A.A.et
al.,1995].

Ia oeae /an oe?ieee ?icaeoie iaaoee aeine?aeaeaiiy ii c’ynoaaiith
no?oeoo?ii-ooieoe?iiaeueieo iniaeeainoae ?cioi?i Na+,E+-AO?-ace,
yei,i/aaeaeii, caaacia/othoue niaoeeo?/i? iio?aae ee?oei a ?aaoeyoe??
?iiiiai o?ainii?oo [Sweadner K.J.,1989; Vasilets L.A.,Schwarz
W.,1993;Pressley T.A.,1996].Iacaaaeath/e ia anoaiiaeaio aiioiii/io i a
?yaei aeiaaeeia ooieoeiiiaeueio niaoeeoi/iinoue icioi?i, ia oeae /an
caeeoa?oueny aiaee?eoei ieoaiiy i?i oa, yei iniaeeainoi no?oeoo?ii?
i?aaiicaoei? a iaia?aii aai ai aca?iiaei? c eiiiaeiei ioi/aiiyi
iaoiiaeththoue ?o oeaieiiniaoeeoi/io ooieoeiiiaeueio niaoeiaeicaoeith.

A ca’yceo c ?ioaa?aeueiith no?oeoo?ith ? ooiaeaiaioaeueiith ooieoe??th
oa?iaioo Na+,E+-AO?-aca caeo/aia aei ?icaeoeo aaaaoueio, a oiio /enei i
iae?iiaeueieo, ee?oeiieo iaoieia?e, ye? noi?iaiaeaeothoueny co?oeoo?iith
ia?aaoaeiaith iaia?ai [Aeai?aoeeee A.E. e ae?.,1990;Aieaeu?aa A.,1992;
Aieaeu?aa A.A.,Eoeeae I.E.,1996; Aieeia A.E.,1996].

Iaeei c oi?aa?naeueieo oeyoia a?iiioeiaeaeaiiy iaia?ai ?aae?co?oueny
/a?ac iiaeeoieaoeith ?o eiiiaeiiai eiiiiiaioa. Oa?iaio Na+,E+-AO?-aca ye
eiioi?iaoeieii-eaaieueiee iioaa?aeueiee aieie oa?aeoa?eco?oueny aenieith
?aaoeyoi?iith caeaaeiinoth aiae noaio eiiiaeii? oace iaia?aie [Aea?aoiia
A.Ae. e ae?.,1984;Robinson J.D.,Pratap P.R., 1993]. Ie?aii aeaii
naiae/aoue i?i caeaaeiinoue ooieoeiiiaeueieo aeanoeainoae icioi?i
Na+,E+-AO?-ace aiae oiceei-oiii/ieo iniaeeainoae ?o eiiiaeiiai ioi/aiiy
a iaia?aii [Matsuda T.,Iwata H.,1986;1988; Iwata H.et al.,1988;Sweadner
K.J.,1989].

A ca’yceo c oeei iaaoaa? aeooaeueiinoi aea/aiiy iiaeeaiaeoaeueii?
/ooeeainoi icioi?i Na+,E+-AO?-ace aei ii?ooaiue no?oeoo?iiai noaio
iaia?aie. Oaei aeineiaeaeaiiy aaaeeeai aeey c’ynoaaiiy
caeiiiii?iinoae,yei iaoiiaeththoue caeaaeiinoue ooieoeiiiaeueieo
aeanoeainoae icioi?i aiae ?aaoeyoi?iiai i aeano?oeoeaiiai aieeaia ia
iaia?aio aiaeiaaiieo aoaeoi?ia oa aeno?aiaeueieo oaeoi?ia. Aieueo
oiai,aiie ? iniiaiiieiaeieie i?e aea/aiii iaoaiiciia, yei caaacia/othoue
aaeaioeaio noieeinoue icioi?i Na+,E+-AO?-ace aei ciii oiceei-oiii/ieo
aeanoeainoae ieaciaoe/ii? (cie?aia iae?iiaeueii?) iaia?aie i?e
oiciieiai/ieo oa iaoieiai/ieo noaiao i?aaiicio.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. ?iaioa aeeiiaia
a eaai?aoi?i? o?ainii?oieo AOOac o aiaeiiaiaeiinoi c ieaiaie
iaoeiai-aeineiaeieo ?iaio Iinoeoooo aiioiii? ii.I.A.Iaeeaaeiia IAI
Oe?a?ie.

Iaoa i caaea/i aeineiaeaeaiiy. Iaoa ?iaioe iieyaaea a aeineiaeaeaiii
/ooeeainoi ?icieo ?aoeaioi?ieo oeiia icioi?i eaoaeioe/ii? noaiaeeieoei
Na+,E+-AO?-ace iiceo aei iaia?aiio?iiieo aieeaia in vitro aeey
c’ynoaaiiy caeiiiii?iinoae ?o ooieoeiiiaeueieo ii?ooaiue a iaia?aii.

Aeey aeinyaiaiiy iaoe aoee iinoaaeaii oaei caaea/i:

1. Iaeaioeoieoaaoe icioi?ie (-noaiaeeieoei Na+,E+-AO?-ace a iaia?aiieo
i?aia?aoao oa?iaioo iiceo ia iiaenoaai aeaoaeoei? eaoaeioe/ieo
oinoiiioa?iaaeiaoia oa?iaioo. Ca aeiiiiiaith iiaiaioi?iiai aiaeico
aeeoa?aioeithaaoe oa?iaioaoeaio aeoeaiinoue icioi?i a i?aia?aoao,
aeineiaeeoe aeaeiao niaoeeoi/iinoue ?aoeaioi?ieo oeiia oa?iaioo,
ioa?aeoa?ecoaaoe ?o aeayei no?oeoo?ii-ooieoeiiiaeueii aeanoeainoi oa
aeoeaiinoue a iinoiaoaeueiiio iioiaaiaci uo?ia.

2. Aea/eoe aeaeo?ioi?aoe/ii aeanoeainoi aiae?ioiaieo aieeia icioi?i
eaoaeioe/ieo iieiiaioeaeia Na+,E+-AO?-ace, yei aeyaeythoueny
oa?iiia?iaeith i?aia?aoia a i?enooiinoi DS-Na.

3. Aea/eoe caeiiiii?iinoi oa?iiiiaeoeaaoeii icioi?i eaoaeioe/ii?
noaiaeeieoei Na+,E+-AO?-ace iiceo oa ie?ie.

4. Aea/eoe /ooeeainoue icioi?i eaoaeioe/iie noaiaeeieoei Na+,E+-AO?-ace
iiceo aei iaia?aiio?iiii? aei? oinoieiiace A2 c io?ooe
na?aaeiueiaciaonueei? eia?e.

5. Aea/eoe /ooeeainoue icioi?i eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace
aei iaia?aiio?iiii? aei? aiiiiiiai aeaoa?aaioo DS-Na a oiiaao ciiie
oiceei-oiii/ieo aeanoeainoae ieaciaoe/ieo iaia?ai eeioei aieiaiiai iiceo
in vitro oa a iinoiaoaeueiee ia?iiae o uo?ia.

6. Aea/eoe /ooeeainoue icioi?i eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace
c ?iciith SH-caeaaeiinoth aei iaoa?iaioaoeaii? ieenii? iiaeeoieaoei?
iaia?ai.

7. Aeyaeoe caeiiiii?iinoi, yei aecia/athoue niaoeeoieo /ooeeainoi
icioi?i eaoaeeoe/ii? noaiaeeieoei iaia?aiica’ycaii? Na+,E+-AO?-ace iiceo
aei iaia?aiio?iiieo aieeaia.

Iaoeiaa iiaecia iaea?aeaieo ?acoeueoaoia. A ?acoeueoaoi i?iaaaeaieo
aeineiaeaeaiue anoaiiaeaii, ui icioi?ie eaoaeioe/ii? noaiaeeieoei
iaia?aiica’ycaii? Na+,E+-AO?-ace iiceo aeyaeythoue ?icio noieeinoue aei
aeanoaaieicoth/eo iaia?aio aieeaia in vitro.

Iniaeeainoi aeaeo?ioi?aoe/ieo aeanoeainoae a iieiae?eeaiiaeiiio aaei a
i?enooiinoi DS-Na aiae?ioiaieo aieeia icioi?i (-noaiaeeieoei
Na+,E+-AO?-ace iiceo iiney niethaieicaoei? i?aia?aoia i?e oa?iiia?iaoei
(1000C) iiaeoaa?aeaeothoue inioaaiiy aiaeiiiiinoae aieeiai-aeaoa?aaioieo
aca?iiaeie aeey icioi?i.

Anoaiiaeaii, ui (1-icioi?ia Na+,E+-AO?-ace iiceo ia? aieueo aenieo
oa?iinoaaieueiinoue (i?e 50-550N) o ii?iaiyiii c (+. Niaoeeoiea
oa?ii/ooeeainoi icioi?i ia aecia/a?oueny eiioi?iaoeieiei noaiii oa?iaioo
(Na+-oa E+-eiioi?iaoei?). Aeaii naiae/aoue i?i aiaeiiiiinoi no?oeoo?ii?
noieeinoi a iaia?aii icioi?i Na+,E+-AO?-ace iiceo oa ie?ie.

Aeyaeaii iaeiiiai?aaeaiinoue niaoeeoiee iaia?aiio?iiieo aieeaia
oinoieiiace A2 c io?ooe na?aaeiueiaciaonueei? eia?e oa DS-Na ia
aeoeaiinoue icioi?i a iaia?aiieo i?aia?aoao Na+,E+-AO?-ace ei?e
aieiaiiai iiceo uo?a. A oiiaao iiii?ii? iiaeoeaaoei? Na+,E+-AO?-ace
oinoieiiacith A2 aoaeo cieaeaiiy /ooeeainoi oa?iaioo aei oaaa?io
onoaa?oueny ia?iaeith i?aia?aoia ne?iaaoeiaei aeueaoiiiii aeey aeeo/aiiy
aieueieo aee?ieo eeneio oa a oiiaao iiaiaoaaiiy, iioeiaeueieo aeey
ca’ycoaaiiy aeieiceaea. I?e iiaeaeueoie iiaeoeaaoei? Na+,E+-AO?-ace a
iaia?aii (1-icioi?ia iiaeoeao?oueny oaeaeoa, iiae (+. Aia?oa
anoaiiaeaii, ui iaiai?ioia iiaeoeaaoeiy Na+,E+-AO?-ace DS-Na, ui
i?aaaeth? aeey (1-icioi?ie, noi?iaiaeaeo?oueny aieueo aoaeoeaiei ??
aeeo/aiiyi c iaia?ai a i?aia?aoao iie?inii ni?i? ?a/iaeie iiceo uo?a oa
aeea. Aiaeiai/ia niaoeeoiea iiaeoeaaoei? icioi?i aeaoa?aaioii oa?aeoa?ia
oaeiae aeey Na+,E+-AO?-ace iici/ea, ii ia aeyaey?oueny o i?aia?aoao c
aeiaaanoiai iiceo. Aiaeiiiiinoi a /ooeeainoi icioi?i aei DS-Na
onoaathoueny a oiiaao oaiia?aoo?ii? iiaeeoieaoei? iaia?aiiiai iao?ieno
(200C(370C) oa i?e caeeneaiii na?aaeiaeua (pH 7,5(6,2).

Aia?oa anoaiiaeaii, ui ieenia iiaiaoaaiiy icioi?i Na+,E+-AO?ace ni?i?
?a/iaeie iiceo uo?a a nenoaii «Fe2++anei?aao» aeua aeey (+ o ii?iaiyiii
c (1, ui aiaeiiaiaea? SH-caeaaeiinoi oa?iaioaoeaii? aeoeaiinoi icioi?i.
Niaoeeoiea iiaeoeaaoei? icioi?i ia caeaaeeoue aiae ia?ieneaeii?
aeano?oeoei? eiiiaeiiai eiiiiiaioa iaia?aie. Aeaii naiae/aoue ia
ei?enoue oiai, ui aieeiaa iieaeoea iae?iiaeueii? Na+,E+-AO?-ace ?
aaciina?aaeiueith iioaiith icioi?ii-niaoeeoi/ii? aei? ieneaeaioia.

Noi?ioeueiaaia aiiioaca i?i inioaaiiy no?oeoo?ieo aiaeiiiiinoae
aieeiai-eiiiaeieo eiiieaenia icioi?i eaoaeioe/ii? noaiaeeieoei
Na+,E+-AO?-ace iae?iiaeueieo iaia?ai, iaoiiaeaieo iniaeeainoyie
no?oeoo?ii? i?aaiicaoei? ?o eiiiaeiiai ioi/aiiy.

I?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoaoia. ?acoeueoaoe aeineiaeaeaiue
iathoue caaaeueiioai?aoe/ia aeey aiioiii? cia/aiiy,noai?ththoue iiai
iaoiaee/ii iiaeoiaee aeey aaeaeaaoii? ioeiiee iaoioiciieiai/ieo oa
oa?iaeieiai/ieo iaia?aiio?iiieo aoaeoia i iiaeooue aooe iniiaith aeey
ooiaeaiaioaeueieo oa i?eeeaaeieo aeineiaeaeaiue a aaeoci aiioiii/ii?
iaia?aiieiai? oa oa?iaeieiai? i?e aea/aiii oa ei?aeoei? iae?iiaeueiiai
iaoaaieicio i?e iaia?aiio?iiieo aieeaao. Iieiaeaiiy aeena?oaoei?
iiaeooue aooe aeei?enoaii a o/aiaiio i?ioeani i?e aea/aiii
iaia?aiieiai?, aiioiii? oa oiciieiai?.

Iniaenoee aianie aeena?oaioa. Aeena?oaioii iniaenoi iaa?oioiaaia iaeay
oa iaoiaeieiaiy ?iaioe, iiaeia?aii i ia?iaeaii aeaii eioa?aoo?e,
aeeiiaii aenia?eiaioaeueii aeineiaeaeaiiy, i?iaaaeaiee iaoeiaee aiaeic
io?eiaiiai iaoa?iaeo, noi?ioeueiaaii iniiaii iieiaeaiiy i aeniiaee,
iiaeaioiaeaii iniiaii ae?oeiaaii i?aoei. Aenieii/euaii iaia?aiii
i?aia?aoe Na+,K+-AO?-ace ie?ie naeii etha’ycii iaaeaii eaiae.aiie.iaoe
E?aaoeiaith A.A. (eaai?aoi?iy o?ainii?oieo AOOac Iinoeoooo aiioiii?
ii.I.A.Iaeeaaeiia IAI Oe?a?ie).

Ai?iaaoeiy ?acoeueoaoia aeena?oaoei?. Iaoa?iaee aeena?oaoei?
aeiiiaiaeaeenue ia Iiaeia?iaeiiio neiiicioii ii o?ainii?oo iiiia
(Oaieini,1989), VI e VII Oe?a?inueeeo aiioiii/ieo c’?caeao
(Ee?a,1992,1997), 3-ie eiioa?aioei? aiioiiieia Ocaaeenoaio
(Oaoeaio,1996), 4th European conference on engineering and medicine
(Warsaw,1997), II c’?caei Oe?a?inueeiai aiioice/iiai oiaa?enoaa
(Oa?ueeia,1998), ia caniaeaiiyo A/aii? ?aaee Iinoeoooo aiioiii?
ii.I.A.Iaeeaaeiia IAI Oe?a?ie (1996-1999?.).

Ioaeieaoei?. Iniiaiee ciino ?iaioe aeeeaaeaii o 24 iaoeiaeo i?aoeyo,c
ieo:1 iiiia?aoi? (iaeiiiniaia), 17 noaooyo o aio/eciyieo i ca?oaiaeieo
i?ioieueieo aeo?iaeao (3 iaeiiiniaii), iaoa?iaeao oa oacao 6
?anioaeieainueeeo oa iiaeia?iaeieo iaoeiaeo c’?caeia oa eiioa?aioeie.

No?oeoo?a oa ianya ?iaioe. Aeena?oaoeiy neeaaea?oueny c oaeeo ?icaeieia:
«Anooi», «Iaeyae eioa?aoo?e» (3 iiae?icaeiee), «Iaoa?iaee oa iaoiaee
aeineiaeaeaiue», «?acoeueoaoe aeineiaeaeaiue oa ?o iaaiai?aiiy» (6
iiae?icaeieia), «Caeeth/aiiy», «Aeniiaee», «Nienie eioa?aoo?e (378
aeaea?ae). Aeena?oaoeith aeeeaaeaii ia 265 noi?iieao iaoeiiieno, aiia
iinoeoue 49 iaethieia oa 10 oaaeeoeue.

IAEssAe EIOA?AOO?E

A iaeyaei eioa?aoo?e aeieeaaeii ?icaeyiooi ieoaiiy
no?oeoo?ii-ooieoeiiiaeueii? ?aeiinoi iieaeoee Na+,K+-AO?-ace i
eiiiaeiiai ioi/aiiy a ieaciaoe/iie iaia?aii. I?iaaaeaii aeaoaeueiee
aiaeic no/anieo oyaeaiue i?i ooieoeiiiaeueio ?ieue icioi?i i
?aoeaioi?ieo oeiia Na+,K+-AO?-ace, cie?aia a oeaieiao iiceo, ?o
aaiaoe/io aeaoa?iiiiaaiinoue. Aaciina?aaeith oaaao i?eaeieaii
iaa?oiooaaiith ia?aaeoiia inioaaiiy aca?iica’yceo ooieoeiiiaeueieo
aeanoeainoae icioi?i Na+,K+-AO?-ace c eiiiaeiei ioi/aiiyi a iaia?aii.

IAOA?IAEE I IAOIAeE
AeINEIAeAEAIUe

Ia’?eoii iniiaieo aeineiaeaeaiue aoa aieiaiee iicie uo?ia (ni?a
?a/iaeia). A ii?iaiyeueieo aeineiaeao aeei?enoiaoaaee oaeiae iici/ie i
aeiaaanoee iicie uo?a, iicie aeea i e?iey (ni?a ?a/iaeia) oa ie?ee
(ciaiioiy iiceiaa ?a/iaeia) uo?ia,e?iey,aeea,naeii.

I?aia?aoe Na+,K+-AO?-ace a iaia?aiica’ycaiie oi?ii io?eioaaee ?c
o?aeoeii iie?inii (50000g) ca aeiiiiiaith «i’yei?» aeno?aeoe?? DS-Na ca
iaoiaeii Ei?aainaia-Naiaeia? [Jorgensen P.L.,1974;Sweadner K.J.,1978] c
ianooiiei oeueo?aoeaio?eooaoaaiiyi(105000g, 4 aiaeeie) a nooiaiaaiio
a?aaei?ioi u?eueiino? naoa?ice (15,25 i 30%,aaaa/ia’?i) ca
iiaeeo?eiaaiei iaoiaeii [Matsuda T.et al.,1984]. Na?aaeiaeua ia?iaee
iie?inii aeaoa?aaioii i?noeei: 50 iI ?i?aeacie (?I 7,5), 1 iI AAeOA, 3
iI o?in-AO?, 0,16 I naoa?ico, 5 ia/ie a?eea iie?inii ? 1,2 /e 1,4 ia/ie
DS-Na (aeey oeaiei iiceo oa ie?ie aiaeiiaiaeii). *an ia?iaee — 30 oa i?e
200N. Aieeiai o?aeoei? a iaaeao 15/25% (ciia 1,iicie) i 25/30% (ie?ee)
naoa?ice (a ?yaei aenia?aiaioia o 15% naoa?ici,ciia 2,iicie)
inaaeaeoaaee oeaio?eooaoaaiiyi (105000g,1 aiaeeia). I?aia?aoe oa?iaioo
noniaicoaaee a na?aaeiaeu?, ui i?noeoue 10 iI iiiaeacie (?I 7,5), 0,16I
naoa?ico, 0,1-0,5iI AAeOA, caa??aaee a ??aeeiio acio?, ?icii?iaeoaaee
iaeii?aciai,aeei?enoiaoaaee ye iaoeaiee iaia?aiica’ycaiee i?aia?ao
oa?iaioo. Eiioeaio?aoeith aieea aecia/aee ca iiaeeoieiaaiei iaoiaeii
Eio?i i niiaaao. [Cadman E.et al.,1979].

Aeaeo?ioi?ac i?aia?aoia [Weber K., Osborn M.,1969; Laemmli U.K.,1970]
i?iaiaeeee o 5-6%-iiio IAAA. Iieiaeaiiy eaoaeioe/ieo noaiaeeieoeue
icioi?i Na+,K+-AO?-ace eiio?iethaaee aooi?aaeiia?aoi/ii ca
Na+-caeaaeiei, K+- oa oaaa?i/ooeeaei oinoi?eethaaiiyi [(-32P]AO?
[Sweadner K.J.,1979]. Aea/aiiy oa?iiiiaeoeiaaieo aeaeo?ioi?aoe/ieo
aeanoeainoae (-noaiaeeieoeue icioi?i Na+,K+-AO?-ace i?iaiaeeee o
aiaeiiaiaeiinoi c ?iaioith [Ohta T. et al.,1982].

Caaaeueio AO?-acio aeoeai?noue aecia/aee a na?aaeiaeu?: 30 iI o?in-HCl
(?I 7,4 i?e 370N), 130 iI NaCl, 20 iI KCl, 3 iI MgCl2, 3 iI o??n-AO?,
0,2 iI AAeOA, 1,25 iI aeeo?io?a?oie. Mg2+-AO?-acio aeoeai?noue aecia/aee
a i?enooiino? aeiaeaoeiai 3(10-3oaaa?ia. Na+,K+AO?-acio aeoeai?noue
?ic?aoiaoaaee ca ??cieoeath i?ae caaaeueiith AO?-aciith oa
Mg2+-AO?-aciith aeoeaiinoyie. O aaceeoeueiaaiie o?aeoei? iie?inii
Na+,K+-AO?-acio aeoeaiinoue aecia/aee iiney aeaianeoaaiiy eaoaioii?
aeoeaiinoi a i?aia?aoao (2ia aieea/ie) [DS-Na]opt (0,2-0,3 ia/ia aieea)
a na?aaeiaeui caiaeii Ei?aainaio-Naiaeia?. Ieoiia aeoeaiinoue
Na+,K+-AOP-ace (ieiieae Pi/ia aieea ca 1 aiaeeio) noaiiaeea aeey
iie?inii: 130-200 (aeiaaanoee iicie, ie?ee uo?a), 90-120 (ei?a aieiaiiai
iiceo, iici/ie uo?a), 50-60 (ei?a aieiaiiai iiceo aeea);aeey
iaia?aiica’ycaiiai oa?iaioo: 380-540 (aeiaaanoee iicie), 320-520 oa
210-260 (ei?a aieiaiiai iiceo, ciia 1 oa ciia 2 aiaeiiaiaeii), 350-720
(ie?ee uo?a); 210-350 (oeaieie aeea); 160-290 (oeaieie e?iey); 720-1200
(ie?ee naeii).

Iiaiaoaaiiy Na+,K+-AOP-ace oaaa?iii aea/aee a i?ioeanni ?icaeoeo
AO?-acii? ?aaeoei? [Matsuda T.et al.,1984] /e iiney iiia?aaeiuei?
iieoaaoei? i?aia?aoia c iiaiaioi?ii 10-15 oa i?e 370N o aiaenooiinoi
ENl. AO?-acio ?aaeoeith caioneaee aeiaeaaaiiyi 20 iI ENl [Urayama
O.,Nakao M.,1979]. Ia?aiao?e iiaiaoaaiiy oaaa?iii Na+,K+-AO?-acii?
aeoeaiinoi icioi?i oa?iaioo ((+ e (1) a i?aia?aoao c iiceo uo?a
?ic?aoiaoaaee caiaeii c iiaeaei aeey aeaio iacaeaaeieo oeaio?ia
ca’ycoaaiiy aeieiceaea. Niaoeeoi/io aeith aaaioia ia icioi?ie
eaoaeioi/ii? noaiaeeieoei Na+,E+-AO?-ace ioeiithaaee ia iniiai aiaeico
ciii aeaiooacii? e?eai? caeaaeiinoi oa?iaioaoeaii? aeoeaiinoi io
eiioeaio?aoei? oaaa?io [Sweadner K.J.,1989]]. Eiioeaaa ?icaaaeaiiy
na?aaeiaeua ia?iaee i?e aecia/aiii AO?-acii? aeoeaiinoi noaiiaeoue
100-200 ?acia. A noaoeiiia?ieo oiiaao iiaiaoaaiiy aeoeaiino? ie?aieo
?cioi?i ?ic?aoiaoaaee ca ??cieoeath i?ae aeoeaiinoyie aac ?ia?a?oi?o ? a
i?enooiino? 5(10-6I oaaa?io ((+: noia?ia aeoeai?noue oaaa?i/ooeeaeo
?cioi?i (2 ? (3 ic noiaeeie aiioiii/ieie aeanoeainoyii,aeoeaiinoi yeeo
ia aeeoa?aioeiththoueny eiiaoe/ii) aai 5(10-6I ? 3(10-3 I oaaa?io
(oaaa?i?acenoaioia (1-?cioi?ia).

Ia?iaeo oa?iaioo iiceo 1 iI N-aoeeiaeaiiiaeii (N-AI) oa o?eineiii (a
i?enooiinoi 60iI ENl) i?iaiaeeee o aiaeiiaiaeiinoi c ?iaioaie [Sweadner
K.J.,1979; Matsuda T.et al.,1984]. Aea/aiiy oa?iiiiaeoeaaoei?
Na+,K+-AO?-ace i?iaiaeeee i?e 500N aai 550N a na?aaeiaeui: 10 iI
iiiaeacie (?I 7,5), 2,5 iI aeeoiio?a?oie, 0,2 iI EAeOA, 5% naoa?ica, 40
iea/ie aieea i?aia?aoo. Ia?iaeo i?aia?aoia Na+,K+-AO?-ace iiceo
oinoieiiacith (OE) A2 c io?ooe Naja naja oxiana (icioa?iaio c I.i.=13
eAea) i?iaiaeeee 30 oa i?e 370N a na?aaeiaeui, yea iinoeei 25 iI
iiiaeacie (?I 7,5), 1iI NaNl2. ?aaeoeith coieiyee 10-e?aoiei
?icaaaeaiiyi ?ic/eiii c AAOA (eiioeaaa eiioeaio?aoeiy 5iI).

Ieenio iiaeeo?eaoe?th iaia?aiieo i?aia?aoia Na+,K+-AO?-ace (0,5 ia/ie)
i?iaiaeeee a 30 iI o??n-HCl aooa?? (pH 7,4 i?e 370N), ui i?noeoue 10 ieI
FeSO4 ? 0,2 iI anei?a?iiao eeneioo. ?aaeoe?th coieiyee 10-e?aoiei
?icaaaeaiiyi oieiaeiei 10iI o??n-HCl aooa?ii, ui i?noea 0,5 iI AAeOA
(eiioeaaa eiioeaio?aoeiy). Aecia/aee aiino iaeiiiaiai aeiaeueaeaaiaeo
(IAeA) e eieueeinoue aieueieo SH-a?oi ca aeiiiiiaith
5,5′-aeioii-ain(2-iio?iaaicieii? eeneioe) (AeOIAE) [Aieaeu?aa A.A. e
ae?.,1992; Eo?aeea A.A. e ae?.,1996].

?acoeueoaoe ia?iaeaii noaoenoe/ii [Ieioeineee A.I.,1981].

?ACOEUeOAOE OA ?O IAAIAI?AIIss

Oa?aeoa?enoeea iaia?aiieo i?aia?aoia Na+,K+-AO?-ace. A i?aia?aoao
Na+,E+-AO?-ace ei?e aieiaiiai iiceo ia aiaeiiio aiae ie?ie ((1)
iaeaioeoieiaaii aeaeo?ioi?aoe/ii aeene?aoii aieeiai ciie
[(-32P]oinoiiioa?iaaeiaoia icioi?i (-noaiaeeieoeue ((1 oa (+).
I?iaeaiiino?iaaii niaoeeoieo aei? o?eineio ia (1-icioi?io Na+,E+-AO?-ace
a A2-eiioi?iaoei? oa?iaioo oa N-AI ia (+-oi?io ye e?eoa?ith aeey ?o
aeaoaeoei? a aaoa?iaaiieo ii icioi?iiiio neeaaeo i?aia?aoao oa?iaioo
iiceo.

A i?aia?aoao c iiceo uo?a iaeaioeoieiaaii aeoeaiinoi aeieiceae/ooeeaiai
oa aeieiceae?acenoaioiiai oeiia icioi?i Na+,E+-AO?-ace ((+ oa (1) o
aiaeiiaiaeiinoi c oaaa?icaeaaei?noth ?o oinoiiioa?iaaeiaoia oa
niaoeeoieith aei? o?eineio, N-EM, oa?iiia?iaee. Oei iaoiaee/ii iiaeoiaee
canoiniaaii aeey iai?aaeaii? iiaeoeaaoeii icioi?i a i?aia?aoao iiceo
aeea. Aeyaeaii ?o aenieo noiaeo /ooeeainoue aei na?oeaaiai aeieiceaeo.
Aeey ?icieo aeaeia oaa?ei iieacaii, ui aiaeiiiiinoi nii?iaeiaiinoi aei
oaaa?io oa?aeoa?ii aeey (1-icioi?ie Na+,E+-AO?-ace. Oa?aeoa?enoeee
iiaiaoaaiiy Na+,E+-AO?-ace iiceo uo?a oa aeea a aeaio aenia?eiaioaeueieo
oiiaao iaaaaeaii a oaae.1,2.

Oaaeeoey 1

Ia?aiao?e ?ia?aoaaiiy oaaa?iii Na+,K+-ATP-acii? aeoeaiino? a iaia?aiieo
i?aia?aoao c ei?e aieiaiiai iiceo uo?a oa aeea (?ia?aoaaiiy oa?iaioo a
i?ioean? AO?-acii? ?aaeoe??, M(m,n=3-6).

Iicie Ie?ee

Ia?aiao?e

Ia?iaea

eiio?ieue o?einei N-AI 500N eiio?ieue

Uo?

A., % * 100 26,2 ( 2,5 # # 9,6 ( 1,6 # # 19,0 ( 0,8 #
# 100

Ei,1,ieI 0,16 ( 0,02 0,27 ( 0,06 – –

(+ ** 71,2 ( 1,0 # 96,6 ( 0,4 # # – –

Ei,2, ieI 76,3 ( 2,8 – 146 ( 12 79,1 ( 7,2 3,4 (
2,4

(1 ** 31,6 ( 1,5 – 82,8 ( 3,7 # # 95,3 ( 2,5 # # 100,4
( 2,6

Aee

A., % * 100 29,3 ( 5,1 # # 24,2 ( 1,3 # # 25,4 ( 6,9 # #
100

Ei, ieI 0,25 ( 0,02 0,20 ( 0,01 0,27 ( 0,01 0,23 ( 0,00 0,38
( 0,02

I?ei?oea: * — A — oa?iaioaoeaia aeoeai?noue i?aia?aoo; **- iaaaaeaii
Vmax (%) aeniei (1)- oa iecueeiao?iiiai (2) eiiiiiaio?a ?ia?aoaaiiy
Na+,K+-ATP-acii? aeoeaiino? oaaa?iii ((+ oa (1-?cioi?ie); # p(0,001 a
ii??aiyii? c Vmax aeey (1; # #- p(0,001 a ii??aiyii? c eiio?ieai.

Oaaeeoey 2

Ia?aiao?e ?ia?aoaaiiy oa?iaioo a i?ioean? iiia?aaeiuei? ?ieoaaoe??
c Mg2+, ATP, Na+, oaaa?iii

Ia?aiao?e Iicie, eiio?ieue Ie?ee,
eiio?ieue

Uo?

Ei,1, iI 24,2 ( 3,5

(+ ** 65,0 ( 2,4 #

Ei,2, ieI 66,1 ( 8,0
80,3 ( 8,9

(1 ** 33,5 ( 2,0
100

Aee

Ei, iI 14,7 ( 2,2
16,3 ( 1,8

I?ei?oea: iicia/aiiy aeea. a oaae. 1

Aeey aeaianeiaaii? Na+,E+-AO?-ace iie?inii aeiaaanoiai iiceo (~90%
aeoeaiinoi (+ a i?aia?aoao) aeyaeaii aieueo aenieo o ii?iaiyiii c
(1-icioi?iith oa?iaioo ie?ie oyaio nii?iaeiaiinoue aei Na+ oa noano?aoo
Mg(AO?2- i iaeiaeiao /ooeeainoue aei aeoeaaoei? E+ oa iiaiaoaaiiy Na2+.

Iieacaii, ui a ieaciaoe/ieo iaia?aiao c ni?i? ?a/iaeie iiceo uo?a a
iinoiaoaeueiee ia?iiae noi?iiaaiee noaaieueiee icioi?iiee eiiieaen ic
noiaeeie ?aoeaioi?ieie aeanoeainoyie. Oa?iaioaoeaia aeoeaiinoue icioi?i
aeinyaa? iaeneioio ia 30-e aeaiue iinoaia?iiiaeueiiai ?icaeoeo – /ano
noaaieicaoei? aeaeo?ioiciieiai/ieo oa?aeoa?enoee iiceo [Venadakis
A.,Woodbury D.M.,1962;Abdel-Latif A.A.et al.,1967].

Oaeei /eiii, ca icioi?iiei neeaaeii i no?oeoo?ii-ooieoeiiiaeueieie
oa?aeoa?enoeeaie iaia?aiii i?aia?aoe Na+,E+-AO?-ace aiaeiiaiaeathoue
oei,ui aeei?enoiaothoueny i?e aea/aiii no?oeoo?ii-ooieoeiiiaeueieo
aca?iica’yceia oei?? oa?iaioii? nenoaie a iaia?aii [Matsuda T., Iwata
H.,1986;1988; Iwata H.et al.,1988; Sweadner K.J.,1989].

Aeaeo?ioi?aoe/ia aaoa?iaaiiinoue aieeiai-aeaoa?aaioieo eiiieaenia
eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace, ?iaeoeiaaia oa?iia?iaeith.
I?iaaaeaii aea/aiiy aeaeo?ioi?aoe/iiai ?iciiaeieo a IAAA
aieeiai-aeaoa?aaioieo eiiieaen?a aiae?ioiaieo aieeia icioi?i
(-noaiaeeieoeue Na+,E+-AO? -ace iiney oa?iiia?iaee i?aia?aoia a
i?enooiinoi 1% DS-Na i 1% 2-ia?eaioiaoaiieo.

Iieacaii,/oi a oiiaao aeaeo?ioi?aco o aenieiii?enoiio iieiae?eeaiiaeiiio
aaei aeyaey?oueny oa?iiiiaeoeiaaia (i?e t0>700C) ?icaeai?iiy aiiiaaiii?
aieeiai? ciie (1-noaiaeeieoei (~90 eAea) Na+,E+-AO?-acu ie?ie
(naeii,uo?a,aeea) c ooai?aiiyi aeiaeaoeiai? aieeiai? ciie c aieueoith
aeaeo?ioi?aoe/iith ?ooiiinoth (II (~85 eAea) (o aiaeiiaiaeiinoi c
aeineiaeaeaiiyie ia oa?iaioi c ie?ie niaaee [Ohta T.,1982]).

Iiney iioaineaii? oa?iiia?iaee (1000C,(1oa) i?iaeaiiino?iaaii iiyao i?e
aeaeo?ioi?aci ii Eaaiei (Laemli K.,1970) oaeiae aeeoocii? aieeiai? ciie
a iaeanoi oyaieo iieaeoey?ieo ian aeueo, iiae o caeeoeiai? aieeiai?
iieine (I (?en.1). Oea ia ? ianeiaeeii ooai?aiiy aa?aaaoia, ai iaaea
aeeoocii? ciie (aiaeii/aia () ia ia?aaeuo? 105 eAea. Iieacaii, ui
i?ioean oa?iiiiaeoeiaaii? o?ainoi?iaoei? aieeiai? ciie (-noaiaeeieoei
/ooeeaee aei oiia aiaeiiaeaiiy S-S-ca’yceia. C /anii ia?iaee c?inoa?
iioaineaiinoue aeeoocii? aieeiai? ciie ia oiii ciaioaiiy iioaineaiinoi
iniiaieo aieeiaeo cii (I e (II. E?ii oiai,iiaeaeuaiiy eiioeaio?aoei?
2-ia?eaioiaoaiieo c 1iI aei 5iI (aea ia DS-Na) ni?eyei ia?aaaaeiiio
ooai?aiith ciie «(«.

Aeyaeaiee oaiiiai ? oa?aeoa?iei aeey (1-noaiaeeieoei Na+,E+-AO?-ace
ie?ie onio aeineiaeaeaieo aeaeia oaa?ei. Oea naiae/eoue i?i inioaaiiy
caaaeueieo no?oeoo?ieo caeiiiii?iinoae i?e oa?iiiiaeoeiaaiiio
ia?aoai?aiii (-noaiaeeieoei, ia caeaaeieo aiae no?oeoo?ieo iniaeeainoae
?aoeaioi?ieo oeiia oa?iaioo.

I?e/eia aiiiaeueii? iiaaaeiiee aieeiaeo cii eaoaeioe/ii? noaiaeeieoei
iieyaa? o inioaaiii iacae/aeieo aieeiai-aeaoa?aaioieo
aca?iiaeie,iaiaeiaeiaeo aeey aiae?ioiaieo aieeia icioi?i Na+,E+-AO?-ace
[Cortas N. et al.,1991]. Oea iiaea iaoiiaethaaoe niaoeeoi/io caeaoiinoue
aiae?ioiaieo aieeia (-noaiaeeieoeue iaaoaaoe iaiiaiinoth ?icai?iaii?
eiioi?iaoei? (aieueo aeiaoey?ii?) c aeuith aeaeo?ioi?aoe/iith
?ooiiinoth,iiae iaaioue o oeieeii ?icai?iaii? eiioi?iaoei? c
iaeneiaeueiith eieueeinoth ca’ycaiiai aeaoa?aaioo, a a oiiaao
oa?iiia?iaee ni?e/eiy? ooai?aiith eiioi?iaoe?eieo “inaaaei?cioi?i”
[Sweadner K.J.,1990]. A oeueiio aeiaaeeo aeeoocia ciia (() a?aeiia?aea?
?icai?iai?e eiioi?iaoe?? oa?iaioo, ui ooai?th?oueny a oiiaao
?ioaineaiiai a?aeiiaeaiiy aeenoeueo?aeieo ca’yce?a, c oyaiith
iieaeoey?iith ianith, yea iaaeeaea?oueny aei ni?aaaeiuei? iieaeoey?ii?
iane eaoae?oe/ii? noaiaeeieoe?.

?en.1. Aeaeo?ioi?aa?aia (ca Laemmli; 5,6% IAAA) i?aia?aoia
Na+,K+-ATP-ace ie?ie oa iiceo

i?ney niethaieicaoei? i?e 1000N. A: 1,2–20iea, 3–10iea; A: 1,2–25iea;
3–15iea; A: 35iea a?eea/o?ae.

I?e ii??aiyii? aeaeo?ioi?aoe/ieo aeanoeainoae a?eeiai-aeaoa?aaioieo
eiiieaen?a ?cioi?i eaoae?oe/ii? noaiaeeieoe? Na+,E+-AO?-ace iiceo
anoaiiaeaii, ui a?eeiaa ciia (+ a?eueo no?eea aei i?ioeano oa?iiia?iaee,
i?ae (1 (?en.1,A). Aeyaeai? caeiiii??iino? aeaeo?ioi?aoe/ii? iiaaae?iee
a?eeiai-aeaoa?aaioieo eiiieaen?a (+ ? (1 a oiiaao aeaeo?ioi?aco a
iie?ae?eeai?aeiiio aae? a i?enooiino? DS-Na naiae/aoue i?i iniaeeainoi
niethaieicaoei? aiae?ioiaieo aieeia icioi?i (-iieiiaioeaeia
Na+,E+-AO?-ace o aiaeiiaiaeiinoi c anoaiiaeaiith aeey ieo niaoeeoieith
aiae?ioiaieo aieeiai-aeaoa?aaioieo aca?iiaeie [Cortas N.et al.,1991].

Oa?ii/ooeeainoue Na+,K+-AO?-acu iiceo oa ie?ie. Aeey ioeiiee no?oeoo?ii?
noieeinoi ooieoeiiiaeueii-aeoeaii? iaoeaii? eiioi?iaoei? aieeiai?
aeiaoee Na+,E+-AO?-ace a iaia?aii aoei aeei?enoaii iaoiae
oa?iiiiaeoeaaoei? (Eiceia TH.I. e ae?.,1983).

I?e aeineiaeaeaiii eiiaoeee oa?iiiiaeoeaaoei? oa?iaioo c iiceo oa ie?ie
?yaeo aeaeia oaa?ei aeyaeaii oaeo iineiaeiaiinoue a oa?ii/ooeeainoi
Na+,E+-AO?-ace a i?aia?aoao (o aiaeiiaiaeiinoi c aiinoii (+-oi?ie):
aeiaaanoee iicie > ni?a ?a/iaeia (iici/ie) > ie?ee (?en.2,a). Noaey/e ic
ciaioaiiy aenieiaoiiiiai eiiiiiaioa iiaiaoaaiiy oaaa?iii aeoeaiinoi
Na+,E+-AO?-ace iiceo uo?a ia oiii caa?aaeaiiy oyaii? nii?iaeiaiinoi aei
oaaa?io /anoeiai iiaeoeaiaaii? (1-icioi?ie (?en.2,a;oaae.1) i ii?ooaiiy
ooai?aiiy oinoiiioa?iaaeiaoo (+ (?en.2,a), anoaiiaeaii, ui (+-oi?ia
oa?aeoa?eco?oueny aenieith oa?iieaaieueiinoth. Iieacaii, ui oaea
iioa?i?aoaoeiy i?ioeano oa?iiiiaeoeaaoei? iiaea aooe canoiniaaia i aeey
iioeo i?aia?aoia c ei?e iiceo aeea oa e?iey, yei iinoyoue (+-oi?io
oa?iaioo. Na+,E+-AO?-aca ie?ie ((1-icioi?ia) oa?aeoa?eco?oueny cia/iith
oa?iinoieeinoth.

Oa?ii/ooeeainoue Na+,E+-AO?-ace ye c oeaiei iiceo, oae i ie?ie inoioii
caeaaeeoue aiae noaaieicaoei? oa?iaioo a eiie?aoieo eiioi?iaoeieieo
noaiao, iiaeoeiaaieo iiiaie iao?ith oa eaeith (Na+- oa K+-eiioi?iaoeiy),
ui aiaeiiaiaea? aeuie aiae?ioiaiinoi iaia?aiiiai ioi/aiiy eaoaeioe/ieo
iieiiaioeaeia Na+,E+-AO?-ace a eiioi?iaoei? A2, iiae A1 [Jorgensen
P.L.,Andersen J.B.,1986]. I?ioa aiaeiiiiinoi a oa?iieaaieueiinoi icioi?i
ia aecia/athoueny eiioi?iaoeieiei noaiii oa?iaioo ye ?acoeueoao
aiaeiiiiinoae a eiioi?iaoeieiie ?iaiiaaci, yei aeyaeaii aeey icioi?i
Na+,E+-AO?-ace [Matsuda T.,Iwata H.,1988].

I?iaaaeaii aeineiaeaeaiiy naiae/aoue i?i aieueo aenieo no?oeoo?io
noaaieueiinoue a iaia?aii (1-icioi?ie Na+,E+-AO?-ace iiceo oa ie?ie o
ii?iaiyiii c (+. Aeniea oa?iinoaaieueiinoue (1-icioi?ie Na+,E+-AO?-ace ?
?? caaaeueiith no?oeoo?iith oa?aeoa?enoeeith i ia iaoiiaeaia
no?oeoo?ieie iniaeeainoyie aeieiceae?acenoaioiiai oeio oa?iaioo uo?a.

?en.2. Aeoeaiinoue (a), /ooeeainoue aei oaaa?io (a) oa eaoaeioe/ia
oinoi?eethaaiiy (a,a) Na+,K+-ATP-ace iiceo oa ie?ie a oiiaao
oa?iiiiaeoeaaoei? i?e 500N; a: 1–4-ie?ee naeii, aeea, uo?a, e?iey
aiaeiiaiaeii; 5–7 – ei?a aieiaiiai iiceo aeea, e?iey, uo?a aiaeiiaiaeii;
8 – iici/ie uo?a; 9,10 – aeiaaanoee iicie e?iey oa uo?a aiaeiiaiaeii;
n=2 – 4; a: caeeoeiaa aeoeaiinoue oa?iaioo ei?e iiceo uo?a,% aei
eiio?ieth (1): 62% (2), 28% (3), 18% (4), oa?iaio ie?ie aac oa?iiia?iaee
(5). Aeaeo?ioi?ac ca Laemmli (a) oa aooi?aaeiia?aia aaeth (a):oa?iaio
iiceo uo?a oinoi?eethaaee [(-32?]AO? a i?enooiinoi 130iI NaCl (1,2) oa
130iI NaCl+20iI KNl (3,4) aac (1,3) oa iiney oa?iiia?iaee (2,4),
caeeoeiaa aeoeaiinoue-16%; 35iea aieea/o?ae.

A i?iaaaeaieo aeineiaeaeaiyo aeyaeaii aiaeiiiiinoi a oa?iieaaieueiinoi
iiae aenieiaiiieiai/ieie ca aiiiieeneioiith iineiaeiaiinoth (1-oi?iaie
oa?iaioo a i?aia?aoao ie?ie ?icieo aeaeia (?en.2,a), a oaeiae iiae
iaeaioe/ieie (1-icioi?iaie Na+,E+-AO?-ace ?icieo oeaiei iaeiiai aeaeo:
iiceo oa ie?ie uo?a. Oea naiae/eoue,ui oa?ii/ooeeainoue Na+,E+-AO?-ace
aecia/athoue oaeiae oeaieiii iniaeeainoi i?aaiicaoei? oa?iaioo a
iaia?aii.

*ooeeainoue icioi?i Na+,K+-ATP-ace aei oinoieiiace A2. Aeey c’ynoaaiiy
aianeo eiiiaeiiai eiiiiiaioa a noaaieicaoeith ooieoeiiiaeueii-aeoeaii?
eiioi?iaoei? icioi?i Na+,E+-AO?-ace a iaia?aii i?iaaaeaii aeineiaeaeaiiy
c aeei?enoaiiyi OE A2 c io?ooe Naja naja oxiana.

Iieacaii, ui iiii?ia iiaeoeaaoeiy iaia?aiica’ycaii? Na+,E+-AO?-ace ei?e
aieiaiiai iiceo uo?a iiae aieeaii OE A2 (aeecueei 40% caeeoeiai?
oa?iaioaoeaii? aeoeaiinoi a i?aia?aoi) noi?iaiaeaeo?oueny iiiioiei
ciaioaiiyi /ooeeainoi aei oaaa?io (?en.3,a). Oeae aoaeo ia ?
?acoeueoaoii ciiie niiaaiaeiioaiiy aeoeaiinoae ie?aieo icioi?i a
i?aia?aoi aianeiaeie ?o ?icii? /ooeeainoi aei iiaeeoieoth/iai iaia?aio
aieeao OE A2, aea iiaea aooe iiyniaiee niiaieueiaiiyi ca’coaaiiy
iiaiaioi?o. Aianeiaeie oeueiai ca noaiaea?oiee /an iieoaaoei? c oaaa?iii
ia aeinyaa?oueny noaoeiiia?iee ?iaaiue iiaiaoaaiiy iiaeeoieiaaii?
Na+,E+-AO?-ace ia aiaeiiio aiae iaoeaiiai oa?iaioo. Iiaeaeuaiiy /ano
iieoaaoei? i?eaiaeeoue aei aiaeiiaeaiiy oa?aeoa?o caeaaeiinoi aeoeaiinoi
Na+,E+-AO?-ace aiae eiioeaio?aoei? oaaa?io,oeiiaiai aeey iaoeaiiai
oa?iaioo.

E?ii oiai, iiney iiia?aaeiuei? iieoaaoei? c oaaa?iii oeo naieo
i?aia?aoia a i?enooiinoi eiaaiaeia (AO?, Na+, Mg2+),yei noaaieicothoue
E2-P-eiioi?iaoeith Na+,E+-AO?-ace,c yeith oaaa?i ca’yco?oueny c
iaeaieueoith oaeaeeinoth [Aieaeu?aa A.A.,Iaeueaoiia
A.E.,1985],cieeathoue ?icaiaeiinoi a oa?aeoa?i iiaiaoaaiiy eiio?ieueieo
i iiii?ii iiaeoeaiaaieo OE A2 i?aia?aoia (?en.3,a),ui iieyaathoue a
ciaioaiii /ooeeainoi oa?iaioo aei aeieiceaeo a oiiaao iiaiaoaaiiy a
i?ioeani ?icaeoeo AO?-acii? ?aaeoei?. Ia?iaea i?aia?aoia Na+,E+-AO?-ace
ne?iaaoeiaei aeueaoiiiii (NA) aeey aeeo/aiiy aieueieo aee?ieo eeneio –
i?iaeoeoia aiae?ieico oinoieiiiaeia, oaeiae aiaeiiaeth? /ooeeainoue
Na+,E+- AO?-ace aei oaaa?io i?e iiii?iie ?? iiaeoeaaoei? iiae aei?th OE
A2 aei ?iaiy iaoeaiiai oa?iaioo (?en.3,a). Oea aeaco? ia iiaeeo?eoth/ee
aieea aee?ieo eeneio ia oa?iaio. I?iieaea aac NA ia aoaeoeaia.

?en.3. Caeaaeiinoue aeoeaiinoi iaia?aiica’ycaii? Na+,K+-AO?-ace ei?e
aieiaiiai iiceo uo?a (a,a,a) oa aeea (a) aiae eiioeaio?aoei? oaaa?io a
oiiaao iiaeeoieaoei? i?aia?aoia oinoieiiacith A2 aac (a,a,a) oa iiney
ia?iaee NA (a).[OE A2],iea/ie:0(1,2);1(3,4);5(5,6);8(7);
a,a,aI-iiaiaoaaiiy oaaa?iii a i?ioeani AO?-acii? ?aaeoei?,30oa(1,3,5,7)
oa 60oa(2,4,6) iieoaaoei?; a,aII-iiaiaoaaiiy oaaa?iii a oiiaao
iiia?aaeiuei? iieoaaoei? c AO?,Na+,Mg2+,15 oa. Caeeoeiaa aeoeaiinoue
Na+,K+-AO?-ace a i?aia?aoao: a:1,2-100%;3,4-40%;5,6-20%;7-8%; a:
1-100%;3-56%;5-8%;a:1,2-100%,3,4-39%;5,6-6%.

Iiaeaeueoa cieaeaiiy aeoeaiino? Na+,E+-AO?-ace i?ae aieeaii OE A2
noi?iaiaeaeo?oueny i?ia?anoth/ei ciaioaiiyi eiiiiiaioa c iecueeith
nii??aeiai?noth aei oaaa?io a aeaio aenia?eiaioaeueieo oiiaao
?ia?aoaaiiy (?en.3,a,a). ?acoeueoaoe ii aecia/aiith ai?noo aeoeaiino?
(1-?cioi?ie a i?aia?aoao i?e aeinyaiaii? noaoe?iia?ieo ??ai?a
?ia?aoaaiiy oa?iaioo, iaaaaeai? ia ?en. 4, na?ae/aoue i?i ?? i?enei?aio
?iaeoeaaoe?th o ii??aiyii? c (+. Oae, aeoeai?noue (1-?cioi?ie
Na+,E+-AO?-ace i?ney ia?iaee i?aia?ao?a OE A2 (5 iea/ie) cieaeoaaeanue
aei 18,6(2,4% a?ae noia?ii? Na+,E+-AO?-acii? aeoeaiino? i?aia?aoo o
ii??aiyii? c ?? ??aiai a eiio?ie? – 31,0(0,6% (M(m,n=4-5,p<0,05), Ei,1 =0,44 ieI, Ei,2=39,4 ieI. A oeeo oiiaao niaoeeo?ea /ooeeaino? aeoeaiino? ?cioi?i Na+,E+-AO?-ace aei OE A2 ia cacia? ye?nieo ci?i i?ney ia?iaee NA (?en.3,a). ?en.4. Aeoeaiinoue (1-icioi?ie (1,2) a i?aia?aoao Na+,K+-AO?-ace ei?e aieiaiiai iiceo uo?a a oiiaao iiaeoeaaoei? oa?iaioo oinoieiiacith A2 (3): 1-iiaiaoaaiiy oaaa?iii a i?ioeani AO?-acii? ?aaeoei? (30-60oa);2-a oiiaao iiia?aaeiuei? iieoaaoei? c AO?,Mg2+,Na+, 15oa; 100%-aeoeaiinoue noia?ii? Na+,K+-AO?-ace i?e [OE A2]i (1,2) /e aac ia?iaee i?aia?aoia OE A2 (3). Aeey oaaa?i/ooeeai? Na+,E+-AO?-ace ei?e aieiaiiai iiceo aeea oaeiae i?iaeaiiino?iaaii cieaeaiiy oyaii? nii??aeiaiino? iiaeeo?eiaaiiai oa?iaioo aei na?oeaaiai ae?eiceaeo (Ei~0,5ieI) o ii??aiyii? c eiio?ieai (Ei~0,2ieI) a oiiaao ?ia?aoaaiiy a i?ioean? AO?-oinoia?ae?ieaciiai oeeeeo (?en.3,a). Oe? a?aei?iiino? i?aaeththoueny i?e iiia?aaei?e ?ieoaaoe?? oa?iaioo a oiiaao oinoi?eethaaiiy (Ei~15iI). Aeniea /ooeea?noue (1-?cioi?ie Na+,E+-AO?-ace iiceo aei ?iaeoeaaoe?? ia ? iane?aeeii ?? aea??iiai i?ioaie?co, i?i ui na?ae/eoue caa?aaeaiiy a aeine?aeaeaiiyo aeaeo?ioi?aoe/iiai i?io?eth ?cioi?i eaoaeioe/ii? noaiaeeieoei a oiiaao ia?iaee i?aia?aoia oa?iaioo OE A2 oa noieeinoue Na+,E+-AO?-ace iiceo aei aiaeiaaiieo i?ioaac [Aeaaeeie?iaa I.I.e ae?.,1995]. Iiaeia noaa?aeaeoaaoe, ui aeyaeaii caeiiiii?iinoi aei? OE A2 ia icioi?ie iaoiiaeaii aeaa?aaeaoei?th ?o eiiiaeiiai ioi/aiiy, a ?o oa?aeoa? aecia/a?oueny nooiaiai iiaeoeaaoei? oa?iaioo. Iaaaaeaii aeaii naiae/aoue i?i inioaaiiy niaoeeoiee no?oeoo?ii? i?aaiicaoei? aieeiai-eiiiaeieo eiiieaenia icioi?i eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace iae?iiaeueieo iaia?ai. Aeniiaie i?i aiaeiiiiinoi a eiiiaeiie /ooeeainoi icioi?i iiceo oaeiae aeoiaeeoue c aeineiaeaeaiue c aeei?enoaiiyi iioeo oinoieiiac [Matsuda T.,Iwata H.,1986;Bougis P.E.et al.,1989]. *ooeeainoue icioi?i Na+,K+-AO?-ace aei DS-Na. Ia?iaea iie?inii c ni?i? ?a/iaeie iiceo uo?a DS-Na a eiioeaio?aoeiyo, aeua iioeiaeueieo,ui aeei?enoiaothoueny aeey aeaianeoaaiiy oa?iaioaoeaii? aeoeaiinoi o aaceeoeao, noi?iaiaeaeo?oueny ia?aaaaeath/ith iiaeoeaaoei?th (1-ecioi?ie oa?iaioo o ii?iaiyiii c (+. Oea aeyaey?oueny o iineiaeiaiiio ciaioaiii a i?aia?aoao aiaeiiaiaeiiai eiiiiiaioa Na+,E+-AO?-acii? aeoeaiinoi c iecueeith nii?iaeiaiinoth aei oaaa?io i?e niiaaiaeiioaiii aeaoa?aaio/aieie aeua aaee/eie 0,4 ia DS-Na/ia aieea (?en.5). Oyaia nii?iaeiaiinoue aei oaaa?io icioi?i Na+,E+-AO?-ace a i?ioeani iiaeoeaaoei? aeaoa?aaioii ia ciiith?oueny, ui aea? iiaeeeainoue ioeiieoe aieea aeaoa?aaioo ia ?o aeoeaiinoue. ?en.5. Caeaaeiinoue aeoeaiinoi noia?ii? Na+,K+-ATP-ace ei?e aieiaiiai iiceo uo?a aiae eiioeaioaoei? oaaa?io (a) oa aeoeaiinoue (1-icioi?ie (a,1) a i?aia?aoao iie?inii iiney ?o ia?iaee DS-Na; a: ia DS-Na/ia aieea: 1-0,2; 2-0,6; 3-0,8; a: 100%-aeoeaiinoue noia?ii? Na+,K+-ATP-ace i?e [DS-Na]i; IN ia?iaeyee i?e [DS-Na]opt, caeeoeiao eieueeinoue aeaoa?aaioo aiineee a na?aaeiaeua AO?-acii? ?aaeoei? (a,2). Aeey c'ynoaaiiy i?e/ei niaoeeoiee iiaeoeaaoei? icioi?i Na+,E+-AO?-ace aeaoa?aaioii iie?iniie,ia?iaeaii DS-Na a oiiaao iaiai?ioii? iiaeoeaaoei?, oiaoi i?e niiaaiaeiioaiiyo aei aieea aeua iioeiaeueieo aeey i/enoee iaoeaieo iaia?aiica'ycaieo i?aia?aoia oa?iaioo ca iaoiaeii Ei?aainaia-Naiaeia?, oeaio?eooaoaaee a a?aaei?ioi uieueiinoi naoa?ice. sse ?acoeueoao iaiaeii?iaeiinoi oiceei-oiii/ieo aeanoeainoae iiioeyoeie iaia?aiieo o?aaiaioia iie?inii a a?aaei?ioi uieueiinoi naoa?ice aeyaeythoueny aeai aieeiai ciie: ia iaaei 15/25% naoa?ice (ciia 1) e a 15% naoa?ici (ciia 2), yei aiae?iciythoueny ca /ooeeainoth Na+,E+-AO?-ac aei DS-Na, aea, ye i o aeiaaeeo iie?inii,oa?aeoa?ecothoueny aieueoith /ooeeainoth (1-icioi?ie oa?iaioo o ii?iaiyiii c (+ aei iaiai?ioii? iiaeoeaaoei? aeaoa?aaioii (?en.6,a). Ca aeiiiiiaith aeaeo?ioi?aco a IAAA a i?enooiinoi DS-Na i?aia?ao?a iaia?aiica’ycaii? Na+,E+-AO?-ace ei?e aieiaiiai iiceo uo?a iieacaii, ui iaiai?ioia ?iaeoeaaoe?y ?cioi?i aeaoa?aaioii noi?iaiaeaeo?oueny aeaeaeaiiyi ?o eaoae?oe/ieo noaiaeeieoeue c iaia?ai, yea i?aaaeth? aeey (1-?cioi?ie (?en.6,a). A oiiaao ??aiiaaeeeeo ni?aa?aeiioaiue aeaoa?aaio/a?eie oeae aoaeo aeyaeaii i?e eiioeaio?aoe?yo DS-Na, ye? iii??ii ia?aaeuothoue aaee/eio e?eoe/ii? eiioeaio?aoe?? i?oeaeiooai?aiiy, oae ? ieueo ca ia?. Oea na?ae/eoue i?i oa, ui ca’ycoaaiiy iiiiia?ii? oi?ie DS-Na, a ia i?oeaey?ii?, iaoiiaeth? niaoeeo?/iee aei ?cioi?i iaia?aiio?iiiee aoaeo aeaoa?aaioo. Oae? caeiiii??iino? ?iaeoeaoth/i? ? nietha?e?coth/i? ae?? DS-Na aeey ?cioi?i Na+,E+-AO?-ace aeyaeaii oaeiae a i?aia?aoao ei?e aieiaiiai iiceo aeea (?en.6,a,a), iici/eo uo?a, aea ia aeiaaanoiai iiceo, aea /ooeea?noue ?cioi?i aei aeaoa?aaioo iaeiaeiaa. ?en.6. Caeaaeiinoue aeoeaiinoi iaia?aiica'ycaii? Na+,K+-ATP-ace aiae eiioeaio?aoei? oaaa?io (a,a) oa aeaeo?ioi?aa?aia (a,a) i?aia?aoia c ei?e aieiaiiai iiceo uo?a (a,a) oa aeea (a,a); ia DS-Na/ia aieea i?e aeno?aeoei? iie?inii (5ia/ie): 1-0,5; 2-0,6; 3-0,8 (a); 1,2-0,4; 3,4-0,6; 5-0,7 (a); aieeiai ciie a a?aaei?ioi uieueiinoi naoa?ice: 15/25% (ionoi neiaiee), 15% (iiaii); aeaeo?ioi?ac ca Laemmli (ciia eaoaeioe/ii? noaiaeeieoei): oinoi?ieaca b, 92,5eAea (a1,a,1); 30 iea a?eea/o?ae ca aeiyoeii a: 5,6-50 iea. Io?eiai? aeai? i?i a?aei?iiino? o noaa?eueiino? ?cioi?i Na+,E+-AO?-ace iiceo aei aeac?ioaa?oth/i? iaia?aio ae?? DS-Na aiaeiiaiaeathoue c?iaeaiiio aeua aeniiaeo i?i niaoeeoieo i?aaiicaoei? a ieaciaoe/ieo iaia?aiao eeioei ei?e aieiaiiai iiceo aieeiai-eiiiaeieo eiiieaenia icioi?i eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace. Iieacaii, ui aeyaeaiee aoaeo DS-Na ia ? ianeiaeeii aeai?iiai i?ioaieico eaoaeioe/ii? noaiaeeieoei a i?ioeani ia?iaee iie?inii aeaoa?aaioii. Niaoeeoi/ia aei (1-icioi?ie Na+,E+-AO?-ace iaia?aiio?iiia aeiy DS-Na iacaeaaeii aiae oeio ?? /ooeeainoi aei na?oeaaeo aeieiceaeia (uo?,aee) naiae/eoue i?i oiiaa?naeueii iniaeeainoi ?? no?oeoo?ii? i?aaiicaoei? a ieaciaoe/iie iaia?aii eeioei ni?i? ?a/iaeie, a ia aiei? ?a/iaeie (aeiaaanoiai iiceo). I/aaeaeii, ui niaoeeoiea aei? aeaoa?aaioo iaoiiaeaia oiceei-oiii/ieie aeanoeainoyie iaia?aie, a naia ?? eiiiaeiiai eiiiiiaoa, ai iathoue iinoea iniaeeainoi a /ooeeainoi aei DS-Na iaeiaeiaeo icioi?i eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace a ?icieo noaiiioeyoeiyo iaia?aiieo o?aaiaioia ei?e aieiaiiai iiceo, a oaeiae iniaeeainoi iiaeoeaaoei? icioi?i a i?aia?aoao c ?icieo oeaiei. Caieueoaiiy /ano iiia?aaeiuei? iieoaaoei? iie?inii c aeaoa?aaioii iiaeaeuo? aoaeoeaiinoue iiaeoeaaoei? Na+,E+-AO?-ace, ii ia onoaa? ?icieoei a /ooeeainoi icioi?i aei DS-Na. Oaeaeeinoue iiaeoeaaoei? aeaoa?aaioii oaeiae aeua aeey (1-icioi?ie (?en.7). Aiaeii/aii aeua aiaeiiiiinoi a /ooeeainoi aei iiaeoeaaoei? aeaoa?aaioii icioi?i Na+,E+-AO?-ace iiceo aei?ineeo uo?ia aeyaeythoueny oaeiae a iie?iniiao iiceo oaa?ei iieiaeiai aieo i ia caeaaeaoue aiae ciiie oiceei-oiii/ieo aeanoeainoae ieaciaoe/ieo iaia?ai a iinoiaoaeueiiio iioiaaiaci. Oea naiae/eoue, ui iniaeeainoi aieeiai-eiiiaeieo eiiieaenia icioi?i noi?iiaaii a iinoaia?iiiaeueiee ia?iiae. C iaoith aea/aiiy aieeao aeaeo?inoaoe/ieo aca?iiaeie a iaoaiicii iiaeoeaaoei? Na+,E+-AO?-ace aeaoa?aaioii aeineiaeaeaiiy i?iaaaeaii a oiiaao caeeneaiiy na?aaeiaeua. Iiii?ia iiaeaeuaiiy eeneioiinoi na?aaeiaeua ia?iaee iie?inii DS-Na a aeiaiaciii ?I 7,5-6,2 noi?iaiaeaeo?oueny ?iceei c?inoaiiyi /ooeeainoi Na+,E+-AO?-ace iiceo uo?a aei aeaoa?aaioo, ui aeaco? ia ?ieue iiaa?oiaaiai ca?yaeo a caaacia/aiii eiai iiaeoeaoth/i? aei? ia oa?iaio (?en.8). ?en.7. Eiiaoeea iiaeoeaaoei? icioi?i eaoaeioe/ii? noaiaeeieoei Na+,K+-AO?-ace DS-Na (0,6ia/ia aieea) a i?aia?aoao iie?inii ei?e aieiaiiai iiceo uo?a; a: 1-(+; 2-(1; 100%-iaeneiaeueia aeoeaiinoue aiaeiiaiaeii? icioi?ie i?e [DS-Na]opt=0,2ia/ia aieea; a: 1-noia?ia Na+,K+-AO?-acia aeoeaiinoue; 2-i?ioeaioiee aiino aeoeaiinoi (1-icioi?ie a i?aia?aoi. ?en.8. *ooeeainoue Na+,K+-AO?-ace iie?inii ei?e aieiaiiai iiceo uo?a aei DS-Na i?e ?icieo cia/aiiyo ?I na?aaeiaeua; a: noia?ia Na+,K+-AO?-acia aeoeaiinoue i?aia?aoo; 1,2-?I=7,5; 3,4-?I=6,5; 5,6-?I=6,2; 1,3,5-3iI AO?; 2,4,6-aac AO?; a: aeoeaiinoue icioi?i; 1,3,5- (+; 2,4,6-(1; 1,2-?I=7,5; 3-6-?I=6,2; 1,2,5,6-aac AO?; 3,4-c AO?. Aiaeic nooiaiy iiiicaoei? ie?aieo a?oi aieeia oa eiiiaeia a oeeo oiiaao naiae/eoue i?i iiaeeeainoue aeieeiaiiy aeiaeaoeiaeo iiceoeaieo ca?yaeia ca ?aooiie aiiiieeneioieo caeeoeia ainoeaeeia,yei ianooue neaaieoaeio a?oio. A oeueiio aeiaaeeo iiyaa aeiaeaoeiaeo iiceoeaieo ca?yaeia iiaea ni?eyoe ?iceiio iiaeaeuaiith ca'ycoaaiiy aeaoa?aaioo c aeniiiiaaieie aeiiaiaie iieaeoee oa?iaioo, aeaci?aaiicaoeiy yeeo ia? nooo?aa cia/aiiy a iiaeoeaaoei? oa?iaioo. sse aeaeii c ?en.8, caoeniee aoaeo AO? a aieueoee ii?i aeyaey?oueny a oiiaao iaiai?ioii? iiaeoeaaoei? oa?iaioo. I?e ?I 7,5 niaoeeoiea aei? DS-Na ia icioi?ie eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace inoioii ia caeaaeeoue aiae iayaiinoi AO? a na?aaeiaeui ia?iaee iie?inii aeaoa?aaioii. I?ioa i?e caeeneaiii na?aaeiaeua aeyaeaii caeiiiii?iinoi a iiaeoeaaoei? icioi?i DS-Na caa?iaathoueny eeoa a i?enooiinoi ioeeaioeaeo. Oea aeaco? ia ae?aioth/ee aoaeo AO?, a i?eeiath/e aei oaaae inioaaiiy eiioaeoo /anoeie AO?-ca'ycoth/i? aeieyiee c iaia?aiith [Lutsenko S.,Kaplan J.H.,1993], i noaaieicoth/ee aieea ia oa?iaio. Ia?iaea iie?inii aeaoa?aaioii (na?aaeiaeua aac AO?) a i?enooiinoi iiiia iaaiith (eaeueoeith),ui aiineee aeey iaeo?aeicaoei? aiiiiieo a?oi eiiiiiaioia iaia?aie,noi?iaiaeaeo?oueny cia/iei iiaeaeuaiiyi /ooeeainoi oa?iaioo aei DS-Na. A oeueiio aeiaaeeo ia onoaa?oueny niaoeeoiea aei? aeaoa?aaioo ia icioi?ie (?en.9,a). Aeey c'ynoaaiiy caeaaeiinoi aoaeoo aeaoa?aaioo aiae ?iaeeiieo aeanoeainoae eiiiaeii? oace iaia?aie aieea DS-Na ia aeoeaiinoue icioi?i Na+,E+-AO?-ace aea/eee a aeiaiaciii oaiia?aoo? 20-370N. Iaiai?ioia iiaeoeaaoeiy Na+,E+-AO?-ace DS-Na iiaeneeth?oueny c oaiia?aoo?ith. I?e aeno?aeoei? iie?inii aeaoa?aaioii i?e 370N onoaathoueny ye aiaeiiiiinoi a /ooeeainoi aei DS-Na Na+,E+-AO?-ace a iaia?aiieo i?aia?aoao ?icieo oeaiei,oae i ie?aieo icioi?i a iie?iniiao c ei?e aieiaiiai iiceo uo?a (?en.9,a). ?en.9. *ooeeainoue Na+,K+-AO?-ace ei?e aieiaiiai iiceo uo?a aei DS-Na i?e ?icieo oiiaao ia?iaee iie?inii aeaoa?aaioii; a:na?aaeiaeua aac AO?; 1,2- eiio?ieue; 3,4-5iI MgCl2; 1,3-(+, 2,4-(1; a: ia?iaea a i?enooiinoi AO?: 1,2-250N; 3,4-300N; 5,6-370N;1,3,5-(+;2,4,6-(1. ?acoeueoaoe aeineiaeaeaiue aiaeiiaiaeathoue eiioeaioei? eieaeueii? a'yceio?iiii? ?aaoeyoei? ooieoeiiioaaiiy iaia?aiieo oa?iaioia, yea aecia/a?oueny oice/iei noaiii iia?aie/ieo eiiiaeia [Aea?aoiia A.Ae. e ae?.,1984]. Caaaeath/e ia iaoaiici iaia?aiio?iiii? aei? DS-Na ia iioaa?aeueii aieee [Ialenius A., Simons K.,1975;Lichtenberg D.et al.,1983;E?aaoeia A.A.,Aeaenaaiei E.?.,1993], niaoeeoiea iiaeoeaoth/iai i niethaieicoth/iai aieeao aeaoa?aaioo ia icioi?ie eaoaeioe/ii? noaiaeeieoei Na+,E+-AO?-ace iiynith?oueny inioaaiiyi iniaeeainoae o aeinooiiinoi aiae?ioiaieo aeieyiie ia aieeo aeey ca'ycoaaiiy aeaoa?aaioo,iaoiiaeaieo aiaeiiiiinoyie a aoaeoeaiinoi aieie-eiiiaeieo aca?iiaeie.A oeueiio aeiaaeeo nooiiiue iiiiaieicaoei? aoeeeueieo eaioethaia i?oeii ca’ycaieo oinoie?i?ae?a iiaea oa?aeoa?ecoaaoe aoaeoeai?noue a?ae?ioiaieo a?eeiai-e?i?aeieo aca?iiae?e, ui aecia/a? ?o no?ee?noue aei cai?ie ia a?eie-aeaoa?aaioi? aca?iiae??. Ci?ia i?e?ia’yceino? iia?aie/iiai e?i?aeo i?e oa?iio?iiiiio oaciaiio ia?aoiae? noi?iaiaeaeo?oueny onoiaiiyi e?eoe/ieo a?aei?iiinoae a aoaeoeaiino? a?eie-e?i?aeieo aca?iiae?e aeey ?cioi?i Na+,K+-ATP-ace ia ca ?aooiie ineaaeaiiy ?iiieo aca?iiae?e, a a ?acoeueoao? ci?ie u?eueiino? iaeoaaiiy iia?aie/ieo e?i?ae?a. Oea i?eaiaeeoue aei eiioi?iaoe?eii? ia?aaoaeiae oa?iaioo, i?aeaeuaiiy aeinooiiino? aeey DS-Na a?ae?ioiaieo ae?eyiie ia a?eeo, aeaiaoo?aoe?? oa nietha?e?caoe?? oa?iaioo. Aeoeai?noue ?cioi?i Na+,K+-ATP-ace iiceo i?e ia?ieneaei?e iiaeeo?eaoe?? iaia?ai. Ia?iaea i?aia?ao?a iaia?aiica’ycaii? Na+,K+-ATP-ace ei?e aieiaiiai iiceo uo??a a nenoai? "Fe2+/anei?aao" i?eaiaeeea aei iaiai?ioiiai ?ia?aoaaiiy oa?iaioaoeaii? aeoeaiino? (?en.10,a), yea noi?iaiaeaeoaaeinue ia?aeaeueiei ooai?aiiyi IAeA (?en.11) i?e ia?ieneaeiiio ieeneaiii eiiiaeia (IIE) oa ciaioaiiyi eieueeinoi aieueieo SH-a?oi a aieeiaeo eiiiiiaioao i?aia?aoia oa?iaioo (75,53±3,26 oa 31,92±2,65 iiieae/ia aieea a eiio?iei oa iiney 60 oa ia?iaee aiaeiiaiaeii,M±m,n=3-5,p<0,05). AAeOA onoaa? aoaeo iiaeoeaaoei? Na+,K+-ATP-ace, ui aeaco? ia ca'ycie iiaiaoaaiiy oa?iaioo c iaoaiiciii iiioeiaoei? ieenieo i?ioeania iiiaie ia?aoiaeieo iaoaeia (Fe). ?en.10. Eiiaoeea iiaeoeaaoei? iaia?aiica'ycaii? Na+,K+-AO?-ace ni?i? ?a/iaeie iiceo uo?a a nenoaii "Fe2+/anei?aao" (M±m,n=3-9); a: noia?ia Na+,K+-AO?-acia aeoeaiinoue i?aia?aoo: 1-aac aeiaaaie, 2-150iI oieiioei?eae, 3-150iI ENl, 4-150iI NaCl, 5-30iI oiina/iaeia, 6-1iI AAeOA; p<0,05 a ii?iaiyiii c eiio?ieai aac aeiaaaie (1); a: aeoeaiinoue (1-icioi?ie (% aei noia?ii? oa?iaioaoeaii? aeoeaiinoi i?aia?aoo): 1-aac aeiaaaie, 2-150iI ENl, 3-30iI oiina/iaeia; ?<0,05 a ii?iaiyiii c ii/aoeiaei aiinoii (1-icioi?ie a i?aia?aoi. Eiiieaeniinoue oa ia?aeaeueiinoue i?ioeania ia?ieneaeii? iiaeeoieaoei? eiiiaeiiai oa aieeiaiai eiiiiiaioia iaia?aie ia aeicaiey? aeeoa?aioeithaaoe ?o aieea i?e ii?ooaiii ooieoeiiioaaiiy Na+,K+-ATP-ace ? aeieaciai noaa?aeaeoaaoe i?i ia?aii?e/eio ?? ieenii? ?iaeoeaaoe?? a iae?iiaeuei?e iaia?ai?. Oiio aeine?aeaeaiiy aoee ni?yiiaai? ia aea/aiiy aianeo ieenii? iiaeeo?eaoe?? a?eeiai? iieaeoee oa?iaioo oa e?i?aeiiai iao?eeno iaia?aie a iaoai?ci ?iaeoeaaoe?? Na+,K+-ATP-ace. ?en.11. Eiiaoeea iaeiie/aiiy IAeA i?e Fe2+/anei?aao-caeaaeiie iiaeoeoei? IIE a i?aia?aoao iaia?aiica'ycaii? Na+,K+-AO?-ace ni?i? ?a/iaeie iiceo uo?ia (M±m,n=3-5): 1-aac aeiaaaie, 2-30 iI oiina/iaeia; p < 0,05 a ii?iaiyiii c eiio?ieai aac iieoaaoei? c ieneaeaioaie. O?ina/iaeia (ON), “ianoea” a?ae?ienee-?aaeeeae?a [Ohta A. et al.,1989], aoaeoeaii aeieo? i?ioean iaeiie/aiiy IAeA (?en.11) ? cia/ii niia?eueith? ?ia?aoaaiiy Na+,K+-AO?-ace (?en.10,a). ?acoeueoaoe aeine?aeaeaiue na?ae/aoue i?i iaoiiaeai?noue ?ia?aoaaiiy Na+,K+-AO?-ace i?yiith iiaeeo?eaoe??th iieaeoee oa?iaioo, a ia eiai e?i?aeiiai ioi/aiiy, i?ae aieeaii aeoeaieo oi?i eenith ia ?aii?o aoaiao ieeniiai ooeiaeaeaiiy iaia?ai, a oiiaao, ye? ia noi?iaiaeaeothoueny ieeniith aeano?oeoe??th e?i?aeiiai iao?eeno. Caaaeath/e ia ci?io ni?aa?aeiioaiiy aeoeaiinoae ?cioi?i Na+,K+-AO?-ace a i?aia?aoao, ??aaiue ?iaeoeaaoe?? (+ ia?aaeuo? oaeee aeey (1 (?en.10,a). A i?enooiino? ON niaoeeo?ea ?iaeoeaaoe?? ?cioi?i ia ci?ith?oueny, aea ?aae?co?oueny i?e iaioeo ??aiyo ?iaeoeaaoe?? oa?iaioo. I?e aeine?aeaeaii? aieeao ?iiieo oiia ia Fe2+/anei?aao-caeaaeio ?iaeoeaaoe?th Na+,K+-AO?-ace anoaiiaeaii, ui ieenia ia?iaea a i?enooiinoi ENl ( NaCl ) ia aiaeiiio aiae oieiioei?eaeo onoaaea niaoeeoieo a iiaeoeaaoei? icioi?i Na+,K+-AO?-ace (?en.10). A na?aaeiaeui c KCl ?iaaiue ooai?aiiy IAeA i?aeoe/ii ia ciiithaaaany, a eieueeinoue ieeneaieo SH-a?oi ciaiooaaeanue. Iaaii, oaea iiiia ioi/aiiy iii'yeoo? ia?ieneaeio iiaeoeaaoeith Na+,K+-AO?-ace, noaaieicoth/e iaia?aiio no?oeoo?o oa?iaioo. Aiaeiii, ui icioi?ie oa?iaioo aiae?iciythoueny ca eieueeinoth, ooieoeiiiaeueiei cia/aiiyi oa aeniiiiaaiinoth aeniei/ooeeaeo aei ieeneaiiy SH-a?oi [Iwata H. et al.,1988; Sweadner K.J., 1989; Vasilets L., Schwarz W., 1993], yei aiaeia?athoue cia/io ?ieue a noaaieicaoei? o?aoeiii? no?oeoo?e oa?iaioo a iaia?aii [Eo?aeea A.A. e ae?.,1996]. Io?eiaii aeaii ciaoiaeyoueny o aiaeiiaiaeiinoi c ?acoeueoaoaie iioeo aeineiaeaeaiue, a yeeo aeyaeaii aiaeiiiiinoi a /ooeeainoi aei ieneaeaioia Na+,K+-AO?-ace iiceo oa ie?ie, (3-icioi?ie a na?eieaii iiiea?aeo ooi?a oa (2-icioi?ie, aeni?aniaaii? a eeioeiao eiiao eiii? Sf-9, uiaei (1-icioi?ie [Huang W. et al.,1994;Xie Z. et al., 1995; Aieaeu?aa A.A., 1995, 1996]. I?ioa a i?iaaaeaieo aeineiaeao aia?oa i?iaeaiiino?iaaii aeuo ieneaeaioio /ooeeainoue SH-caeaaeieo icioi?i (+ ((2 oa (3) o ii?iaiyiii c (1 a i?aia?aoao Na+,K+-AO?-ace ni?i? ?a/iaeie iiceo a oiiaao aeeoa?aioeiaoei? aianeo ieeniiai ooeiaeaeaiiy aieeiaiai oa eiiiaeiiai eiiiiiaioia iaia?aie. Iiaeii c?iaeoe aeniiaie, ui ieenia iiaeoeaaoeiy Na+,K+-AO?-ace ? eiiieaeniei i?ioeanii. Iiaeneeaiiy iiaeoeaaoei? oa?iaioo i?e iioe?aiii ieeniiai i?ioeano i aeoeaaoei? IIE aeaco? ia o/anoue ?icieo oeyoia ieeniiai ooeiaeaeaiiy Na+,K+-AO?-ace. Iaeiae, niaoeeoiea ieeniiai aoaeoo iaoiiaeth?oueny no?oeoo?ieie iniaeeainoyie naia aieeiaeo iieaeoe icioi?i oa?iaioo o aiaeiiaiaeiinoi c SH-caeaaeiinoth ?o ooieoeiiiaeueii? aeoeaiinoi i ia ca'ycaia c iioaineaiinoth ooeiaeaeaiiy eiiiaeiiai iao?eeno iaia?aie. CAEETH*AIIss Oaeei /eiii, ?acoeueoaoe aeineiaeaeaiue aeacothoue ia inioaaiiy iniaeeainoae no?oeoo?ii? i?aaiicaoei? aieeiai-eiiiaeieo eiiieaenia icioi?i eaoaeioe/ii? noaiaeeieoei Na+,K+-AO?-ace a ieaciaoe/ieo iaia?aiao eeioei ei?e aieiaiiai iiceo, yei iaoiiaeaii iniaeeainoyie no?oeoo?ii? i?aaiicaoei? ?o iia?aie/ieo eiiiaeia. Iiaenoiiaoth/e io?eiaii ?acoeueoaoe neiae aiaeiioeoe, ui ia?aaeoiiaith inioaaiiy aiaeiiiiinoae eiiiaeiiai aioethna icioi?i Na+,K+-AO?-ace ? ?o ?icia eieaeicaoeiy a eeioeiao iae?iiia, a oaeiae a ?icieo oeiao eeioei ei?e aieiaiiai iiceo [Sweadner K.J.,1989;1992]. Ia aeeeth/aia iiaeeeainoue inioaaiiy aaoa?iaaiiinoi eiiiaeiiai ioi/aiiy icioi?i a eiie?aoiie aeieyioei iae?iiaeueii? iaia?aie. I?eionea?oueny, ui (1-icioi?ia caeienith? ?aaoeyoeith caaaeueii? eiioeaio?aoei? iiiia Na+ a oeeoiieacii iae?iiia, a (2- oa (3-icioi?ie — a iaiaaeaiiio i?inoi?i iiae ieaciaoe/iith iaia?aiith oa ?aoeeoeoiii [Juhaszova M., Blaustein M.P., 1997; Aeaaeeie?iaa I.I. e ae?.,1998]. Inioaaiiy oiceei-oiii/ieo iniaeeainoae eiiiaeiiai ioi/aiiy icioi?i iiaea caaacia/eoe niaoeeoi/io iiaeoeyoeith eiioi?iaoeieii? ?ooiiinoi /e iioeo eaoaeioe/ieo aeanoeainoae icioi?i aai niaoeeoieo ?o ooieoeiiiaeueii? noieeinoi i?e aeeiaii/ieo ciiiao a iaia?aii caiaeii aiaeiiiio iaoaiicio a'yceio?iiii? ?aaoeyoei? ooieoeiiioaaiiy iioaa?aeueieo iaia?aiieo oa?iaioia, a naia Na+,K+-AO?-ace [Aea?aoiia A.Ae. e ae?.,1984; ?ost R.L.,Klodos I.,1996]. A oie aea /an aeniea ieneaeaioia /ooeeainoue niaoeiaeiciaaieo icioi?i (2 oa (3 iiaea aooe iaeiei c oaeoi?ia, ui aecia/a? eaoano?ioi/iinoue i?ioeania ieeniiai no?ano o ?icaeoeo iae?iiaeueieo iaoieiaie [Aieaeu?aa A.A.,Eoeeae I.E.,1996]. ?acoeueoaoe aeineiaeaeaiue naiae/aoue i?i inioaaiiy aiaeiiiiinoae ooieoeiiiaeueieo ii?ooaiue ooiaeaiaioaeueii? (1- oa niaoeiaeiciaaieo (2- oa (3-icioi?i Na+,K+-AO?-ace i?e iiaeeoieaoei? eiiiaeii? oace iaia?aie /e ieeniie iiaeoeaaoei?, ui iiaea aooe ia?aaeoiiaith iniaeeainoae aaeaioeaii? noieeinoi icioi?i i?e iaaieo oiciieiai/ieo oa iaoieiai/ieo noaiao i?aaiicio. AENIIAEE 1. Aeyaeaii aiaeiiiiinoi a noieeinoi aei iaia?aiio?iiieo aieeaia in vitro icioi?i eaoaeioe/ii? noaiaeeieoei iaia?aiica'ycaii? Na+,K+-AO?-ace iiceo. 2. Aeey aiae?ioiaieo aieeia (-noaiaeeieoei Na+,K+-AO?-ace iiceo oa ie?ie (uo?,aee,naeiy) anoaiiaeaii noiaeinoue aeaeo?ioi?aoe/ieo aeanoeainoae iiney niethaieicaoei? i?aia?aoia a oiiaao oa?iiia?iaee (>700C) a
i?enooiinoi DS-Na oa 2-ia?eaioiaoaiieo. Aeey Na+,K+-AO?-ace ni?i?
?a/iaeie iiceo aeyaeaii aieueo aenieo noieeinoue aei oa?iiia?iaee (i?e
1000N) aieeiai? ciie (+ o ii?iaiyiii c (1.

3. Aeyaeaii oaeo iineiaeiaiinoue a oa?ii/ooeeainoi (i?e 500N)
Na+,K+-AO?-ace c oeaiei iiceo oa ie?ie, yea aiaeiiaiaea? aiinoo (+-oi?ie
a i?aia?aoao: aeiaaanoee iicie>ni?a ?a/iaeia (iici/ie)>ie?ee. Aeey
icioi?i Na+,K+-AO?-ace iiceo aeyaeaii aieueo aenieo oa?iinoaaieueiinoue
(1 o ii?iaiyiii c (+. Niaoeeoiea oa?iiiiaeoeaaoei? icioi?i ia caeaaeeoue
aiae ?aoeaioi?iiai oeio oa?iaioo oa eiai eiioi?iaoeieiiai noaio (Na+- /e
K+-oi?ie). Aeyaeaii aiaeiiiiinoi a oa?iieaaieueiinoi (1-icioi?ie a
i?aia?aoao oa?iaioo c ni?ith ?a/iaeie iiceo oa ie?ie.

4. Aia?oa anoaiiaeaii, ui a i?aia?aoao iaia?aiica’ycaii? Na+,K+-AO?-ace
c ni?i? ?a/iaeie iiceo uo?a (1-icioi?ia oa?iaioo aieueo /ooeeaa o
ii?iaiyiii c (+ aei iiaeoeaoth/i? aei? oinoieiiace A2 c io?ooe Naja
naja oxiana.

5. Aia?oa anoaiiaeaii, ui a i?aia?aoao iie?inii c ni?i? ?a/iaeie i
iici/ea uo?a ia aiaeiiio aiae iaia?ai c aeiaaanoiai iiceo (1-icioi?ia
Na+,K+-AO?-ace oa?aeoa?eco?oueny aeuith o ii?iaiyiii c (+ /ooeeainoth
aei iiaeoeaaoei? DS-Na. Iieacaii, ui iaiai?ioia iiaeoeaaoeiy
aeaoa?aaioii Na+,K+-AO?-ace a i?aia?aoao c ei?e aieiaiiai iiceo uo?a oa
aeea noi?iaiaeaeo?oueny aeeo/aiiyi (-coaiaeeieoeue c iaia?aie, aieueo
aoaeoeaiei aeey (1-icioi?ie.

6. Anoaiiaeaii, ui iiaeaeuaiiy iiaeoeaoth/iai aoaeoo DS-Na (i?e
caieueoaiii o?eaaeinoi ia?iaee iie?inii aeaoa?aaioii, iiaeaeuaiii
oaiia?aoo?e, eeneioiinoi na?aaeiaeua, a i?enooiinoi MgCl2, a iie?iniiao
iiceo iieiaeeo uo?ia) noi?iaiaeaeo?oueny onoiaiiyi aiaeiiiiinoae a
noieeinoi icioi?i Na+,K+-AO?-ace iiceo aei aeaoa?aaioo eeoa a oiiaao
oa?iio?iiii? iiaeeoieaoei? oice/iiai noaio eiiiaeiiai iao?eeno
(200N(370N) oa caeeneaiii na?aaeiaeua (?I 7,5(6,2).

7. Iieacaii, ui (1-icioi?ia Na+,K+-AO?-ace iiceo oa ie?ie aeea
aiaeiineoueny aei aeieiceae/ooeeaiai oeio oa?iaioo. A oiiaao /anoeiai?
oa?iiiiaeoeaaoei? oa aei? DS-Na oyaia nii?iaeiaiinoue aei oaaaiio
icioi?i Na+,K+-AO?-ace ei?e aieiaiiai iiceo uo?a oa aeea ia
ciiith?oueny. Iiii?ia iiaeoeaaoeiy Na+,K+-AO?-ace iiceo oinoieiiacith A2
noi?iaiaeaeo?oueny cieaeaiiyi ?? /ooeeainoi aei oaaa?io o ii?iaiyiii c
iaoeaiei oa?iaioii a oiiaao iiaiaoaaiiy a i?ioeani
AO?-oinoiaiae?ieaciiai oeeeeo. Oeae aoaeo onoaa?oueny ia?iaeith
i?aia?aoia ne?iaaoeiaei aeueaoiiiii aeey aeeo/aiiy aieueieo aee?ieo
eeneio oa ia aeyaey?oueny i?e caieueoaiii o?eaaeinoi iiaiaoaaiiy oa
iiney iiia?aaeiuei? iieoaaoei? oa?iaioo c oaaa?iii a i?enooiinoi
AO?,Na+,Mg2+.

8. Aia?oa iieacaii,ui a oiiaao Fe2+/anei?aao-caeaaeiiai ia?ieneaeiiai
ieeneaiiy eiiiaeia,yea noi?iaiaeaeo?oueny iaiai?ioiith iiaeoeaaoei?th
oa?iaioo i ieeneaiiyi SH-a?oi aieeia a i?aia?aoao iaia?aiica’ycaii?
Na+,E+-AO?-ace ei?e aieiaiiai iiceo uo?a, icioi?ie oa?iaioo (+ a
aieueoie ii?i o ii?iaiyiii c (1 /ooeeai aei ieeniiai iiaiaoaaiiy, ui
aiaeiiaiaea? aeyaeaiei aiaeiiiiinoyi a SH-caeaaeiinoi ?o oa?iaioaoeaii?
aeoeaiinoi.

9. Oiina/iaeia aeieo? ooai?aiiy iaeiiiaiai aeiaeueaeaaiaeo i
niiaieueith? iiaiaoaaiiy oa?iaioo a nenoaii «Fe2+/anei?aao». Niaoeeoiea
/ooeeainoi icioi?i aei ieeneaiiy ia ciiith?oueny i ia caeaaeeoue aiae
iaeiie/aiiy iaeiiiaiai aeiaeueaeaaiaeo. Iaea?aeaii aeaii naiae/aoue i?i
iiaeeeainoue aaciina?aaeiueiai ieeniiai ooeiaeaeaiiy aieeiai? iieaeoee
iae?iiaeueii? Na+,E+-AO?-ace o aiaeiiaiaeiinoi c iiaeeaiaeoaeueiith
ieneaeaioiith /ooeeainoth icioi?i.

10. Cai?iiiiiaaia aiiioaca i?i inioaaiiy aiaeiiiiinoae no?oeoo?ii?
i?aaiicaoei? aieeiai-eiiiaeieo eiiieaenia icioi?i eaoaeioe/ii?
noaiaeeieoei Na+,E+-AO?-ace iiceo, iaoiiaeaieo no?oeoo?ieie
iniaeeainoyie eiiiaeiiai ioi/aiiy a iae?iiaeueiie iaia?aii.

NIENIE IIOAEIEIAAIEO I?AOeUe CA OAIITH AeENA?OAOeI?

1. Eaiey A.A. (Iiiia?aoey). No?oeoo?iay i?aaiecaoeey e ooieoeeiiaeueiay
?ieue ecioa?iaioia Na+,K+-ATP-acu.- Eeaa: Ecae-ai “Eeaaneee
oieaa?neoao”,1998.-162n.

2. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A., Ai?eoiye A.I. Ecioi?iu
eaoaeeoe/aneie noauaaeeieoeu Na+,E+-AO?-acu iicaa e eo
/oanoaeoaeueiinoue e oaaaeio//Iae?ioeiey.-1989.-8,N2.-N.216-223.

3. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A., Ai?eoiye A.I.
Yeaeo?ioi?aoe/aneay aaoa?iaaiiinoue eaoaeeoe/aneie noauaaeeieoeu
Na+,K+-ATP-acu, eiaeooee?oaiay oa?iiia?aaioeie//Aeiee.AI
ONN?.Na?.A.-1989.-N6.-N.59-61.

4. Eaiey A.A., E?aaoeia A.A. Ecioi?iu eaoaeeoe/aneie noauaaeeieoeu
Na+,E+-AO?-acu a oeaiyo aeeaioiuo//Oe?.aeioei.aeo?i.-1990.-62,
N3.-N.17-30.

5. Eaiey A.A.,E?aaoeia A.A. Oa?iieaaeeueiinoue (+-ecioi?iu
eaoaeeoe/aneie noauaaeeieoeu Na+,E+-AO?-acu
iicaa//Oe?.aeioei.aeo?i.-1990.-62,N4.- N.39-44.

6. Eaiey A.A., E?aaoeia A.A. *oanoaeoaeueiinoue Na+,K+-ATP-acu
iae?iiaeueiuo iaia?ai e oaaaeio//Oe?.aeioei.aeo?i.-1992.-64,N4.N.38-43.

7. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A. Oa?ii/oanoaeoaeueiinoue
ecioa?iaioia Na+,E+-AO?-acu iicaa e ii/ae//Oe?.aeioei.aeo?i.-
1994.-66,N6. — N.58-66.

8. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A. *oanoaeoaeueiinoue ecioi?i
eaoaeeoe/aneie noauaaeeieoeu Na+,E+-AO?acu iicaa e?unu e Ds-Na//
Aeioeiey.-1995.-60,N6.-N.970-975.

9. Eaiey A.A., E?aaoeiaa A.A., E?aaoeia A.A. Noaaeeueiinoue ?aoeaioi?iie
ooieoeee ecioa?iaioia Na+,K+-ATP-acu iicaa a oneiaeyo iiaeeoeeaoeee eo
oa?iaioaoeaiie aeoeaiinoe// Oe?.aeioei.aeo?i.-1995.-67, N5.-N.43-48.

10. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A. Aeoeaiinoue ecioa?iaioia
Na+,E+-AO?-acu ei?u aieiaiiai iicaa e eo /oanoaeoaeueiinoue e
aeiaeaoeeenoeueoaoo iao?ey a iinoiaoaeueiii iioiaaiaca e?un//Oe?.
aeioei.aeo?i.- 1996.-68,N1.-N.32-39.

11. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A., Aa?on I.A.
Oa?iieiaeooee?oaiay o?ainoi?iaoeey aaeeiai-aeaoa?aaioiuo eiiieaenia
eaoaeeoe/aneeo noauaaeeieoe Na+,K+-ATP-acu ii/ae e iicaa i?e
yeaeo?ioi?aca a IAAA//Oe?.aeioei.aeo?i.-1996.-68,N2.- N.51-57.

12. Eaiey A.A., E?aaoeiaa A.A., E?aaoeia A.A. Aeeyiea oinoieeiacu A2 ec
yaea Naja naja oxiana ia aeoeaiinoue ecioa?iaioia Na+,E+- AO?acu iicaa
e?unu//Aeioeiey.-1996.-61,N6.-N.998-1005.

13. Eaiey A.A.,E?aaoeiaa A.A., E?aaoeia A.A. Eiaeoeaaoeey ecioi?i
eaoaeeoe/aneie noauaaeeieoeu Na+,E+-AO?-acu iicaa aeiaeaoeeenoeueoaoii
iao?ey//Aeie.iaia?.-1997.- 14,N1.- C.73-82.

14. Eaiey A.A., E?aaoeia A.A. Eiaeoeaaoeey Na+,K+-AO?-acu iicaa e?un
aeiaeaoeeenoeueoaoii iao?ey: aeeyiea ?I, eiiia iaaiey, oaiia?aoo?u//
Oe?. aeioei. aeo?i.-1997.-69,N4.-C.3-8.

15. Eaiey A.A. No?oeoo?iay i?aaiecaoeey ecioa?iaioia Na+,K+-ATP-acu a
ieaciaoe/aneie iaia?aia//Oe?.aeioei.aeo?i.-1997.-69, N5-6.-C.12-24.

16. Eaiey A.A. Oeceieiae/aneay e aaeaioaoeeiiiay cia/eiinoue
ecioa?iaioia Na+,K+-ATP-acu//Oe?.aeioei.aeo?i.-1998.-70, N3.-N.3-11.

17. Eaiey A.A. *oanoaeoaeueiinoue ecioi?i eaoaeeoe/aneie noauaaeeieoeu
Na+,K+-ATP-acu iicaa e?unu e ia?ieneaeiie iiaeeoeeaoeee
iaia?ai//Oe?.aeioei.aeo?i.-1998.-70,N4.-C.118-122.

18. Eaiey A.A., E?aaoeia A.A. Iieaeoey?iua iaoaieciu ?aoeaioeee
na?aea/iuo aeeeiceaeia Na+,K+-ATP-acie// Oniaoe nia?. aeie.-1999.-119,
N1.- N.84-94.

19. Eaiey A.A., E?aaoeia A.A., E?aaoeiaa A.A., Ai?eoiye A.I. Ecioi?iu
eaoaeeoe/aneie noauaaeeieoeu Na+,K+-ATOacu a oeaie iicaa//
Iaaeaeoia?iaeiue neiiiceoi ii o?ainii?oo eiiia.Oacenu aeieeaaeia.-
Oaeeene,1989.-N.31.

20. Eaiey I.A., E?aaoeia I.A. Icioi?ie eaoaeioe/ii? noaiaeeieoei
Na+,K+-ATP-ace:/ooeeainoue aei i?ioaieico,
N-aoeeiaeaiiiaeo,oa?iiiiaeoeaaoei?,oaaa?io// VI Oe?.aiioii.c’?cae. Oace
aeiiiaiaeae, /ano.I.- Ee?a, 1992.-N.76.

21. Eaiey A.A., E?aaoeia A.A. Iaia?aiiua naienoaa ecioa?iaioia
Na+,K+-ATPacu iicaa e?unu: ?acee/ey a /oanoaeoaeueiinoe e oinoieeiaca A2
e eiaeoeaaoeee SDS// 3 Eiioa?aioeey aeioeieeia Ocaaeenoaia. Oacenu
aeieeaaeia.-Oaoeaio.-1996.-N.35.

22. Eaiey I.A., E?aaoeia I.A. Iniaeeainoi no?oeoo?ii? i?aaiicaoei?
aieeiai-eiiiaeieo eiiieaenia icioi?i eaoaeioe/ii? noaiaeeieoei
Na+,K+-ATPace// VII Oe?a?inueeee aiioiii/iee c’icae.Oace
aeiiiaiaeae,/ano.I-Ee?a.-1997.-C.41.

23. Kaplya A.A., Kravtsov A.V. Membrane-dependent properties of brain
Na+,K+-ATPase isoenzymes// 4th European conference on engineering and
medicine.Book of abstracts.-Warsaw.-1997.-P.395.

24. Eaiey I.A., E?aaoeia I.A. Iiaeoeaaoeiy icioi?i eaoaeioe/ii?
noaiaeeieoei Na+,K+-ATP-ace iiceo i?e iaia?aiio?iiieo aieeaao in vitro//
II c’?cae oe?a?inueeiai aiioice/iiai oiaa?enoaa.Oace
aeiiiaiaeae.-Oa?eia.-1998.-N.105.

Eaiey I.A. No?oeoo?ii-ooieoeiiiaeueii aeanoeainoi icioi?i eaoaeioe/ii?
noaiaeeieoei Na+,K+-ATP-ace i?e iaia?aiio?iiieo aieeaao. – ?oeiien.

Aeena?oaoeiy ia caeiaoooy iaoeiaiai nooiaiy aeieoi?a aiieiai/ieo iaoe ca
niaoeiaeueiinoth 03.00.04 – aiioiiiy. – Iinoeooo aiioiii?
ii.I.A.Iaeeaaeiia IAI Oe?a?ie,Ee?a,1999?.

Aeena?oaoeith i?enay/aii aeineiaeaeaiith iniaeeainoae ooieoeiiiaeueii?
noieeinoi icioi?i eaoaeioe/ii? noaiaeeieoei Na+,K+-ATP-ace iiceo aei
iaia?aiio?iiieo aieeaia in vitro aeey c’ynoaaiiy caeiiiii?iinoae ?o
ooieoeiiiaeueieo ii?ooaiue a ieaciaoe/iie iaia?aii. Aia?oa anoaiiaeaii,
ui (1-icioi?ia iaia?aiica’ycaii? Na+,K+-ATP-ace ni?i? ?a/iaeie iiceo (o
ii?iaiyiii c (+) ia? aeuo /ooeeainoue aei iiaeoeaaoei? oinoieiiacith A2
c io?ooe Naja naja oxiana, iiaeoeaaoei? oa niethaieicaoei? DS-Na,
oa?iinoaaieueiinoue oa noieeinoue aei ieeniiai iiaiaoaaiiy a nenoaii
«Fe2+/anei?aao. Cai?iiiiiaaia aiiioaca i?i inioaaiiy aiaeiiiiinoae
no?oeoo?ii? i?aaiicaoei? aieeiai-eiiiaeieo eiiieaenia icioi?i
Na+,K+-ATP-ace iiceo, iaoiiaeaieo no?oeoo?ieie iniaeeainoyie eiiiaeiiai
ioi/aiiy a iaia?aii. Aeaii naiae/aoue,ui iae?iiaeueia Na+,K+-ATP-aca ?
aaciina?aaeiueith iioaiith ieneaeaioii? aei?, niaoeeoi/ii? aei
SH-caeaaeieo oaaa?i/ooeeaeo icioi?i ((+), iaaioue a oiiaao, yei ia
noi?iaiaeaeothoueny ia?ieneaeiith aeano?oeoei?th eiiiaeia.

Eeth/iai neiaa: Na+,K+-ATP-aca,icioi?ie eaoaeioe/ii? noaiaeeieoei,
iiaeeo?eaoe?y iaia?ai.

Eaiey A.A. No?oeoo?ii-ooieoeeiiaeueiua naienoaa ecioi?i eaoaeeoe/aneie
noauaaeeieoeu Na+,K+-ATP-acu i?e iaia?aiio?iiiuo aicaeaenoaeyo. –
?oeiienue.

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie aeieoi?a aeieiae/aneeo iaoe ii
niaoeeaeueiinoe 03.00.04 – aeioeiey. – Einoeooo aeioeiee
ei.A.A.Iaeeaaeeia IAI Oe?aeiu,Eeaa,1999a.

Aeenna?oaoeey iinayuaia enneaaeiaaieth iniaaiiinoae ooieoeeiiaeueiie
onoie/eainoe ecioi?i eaoaeeoe/aneie noauaaeeieoeu Na+,K+-ATP-acu iicaa e
iaia?aiio?iiiui aicaeaenoaeyi in vitro aeey auyniaiey caeiiiia?iinoae eo
ooieoeeiiaeueiuo ia?ooaiee a ieaciaoe/aneie iaia?aia. Aia?aua
onoaiiaeaii, /oi (1-ecioi?ia iaia?aiinaycaiiie Na+,K+-ATP-acu na?iai
aauanoaa iicaa (ii n?aaiaieth n (+) iaeaaeaao aieaa aunieie
/oanoaeoaeueiinoueth e eiaeoeaaoeee oinoieeiacie A2 ec yaea Naja naja
oxiana, eiaeoeaaoeee e niethaeeecaoeee Ds-Na, oa?iinoaaeeueiinoueth e
onoie/eainoueth e ieeneeoaeueiiio eiaeae?iaaieth a nenoaia
«Fe2+/anei?aao». I?aaeeiaeaia aeiioaca i nouanoaiaaiee ?acee/ee
no?oeoo?iie i?aaiecaoeee aaeeiai-eeieaeiuo eiiieaenia ecioi?i
Na+,K+-ATP-acu iicaa,iaoneiaeaiiuo no?oeoo?iuie iniaaiiinoyie eeieaeiiai
ie?oaeaiey a iaia?aia. Aeaiiua naeaeaoaeuenoaotho,/oi iae?iiaeueiay
Na+,K+-ATP-aca yaeyaony iaiin?aaenoaaiiie ieoaiueth ieneaeaioiiai
aeaenoaey,niaoeeoe/iiai e SH-caaeneiui oaaaei/oanoaeoaeueiui ecioi?iai
((+), aeaaea a oneiaeyo, ia nii?iaiaeaeathueony ia?ieneaeiie
aeano?oeoeeae eeieaeia.

Eeth/aaua neiaa: Na+,K+-ATP-aca, ecioi?iu eaoaeeoe/aneie noauaaeeieoeu,
iiaeeoeeaoeey iaia?ai.

Kaplya A.A. Structural and functional properties of Na+,K+-ATP-ase
catalytic subunit isoforms under membrane-disturbing influences. –
Manuscript.

Thesis for doctor’s degree on speciality 03.00.04 – biochemistry.
O.V.Palladin Biochemistry Institute of National Academy of Sciences of
Ukraine,Kyiv,1999.

The dissertation is devoted to investigation of the functional
persistence of the receptor types of brain Na+,K+-ATP-ase catalytic
subunit isoforms under membrane-disturbing influences in vitro for
elucidation of the peculiarities of their functional disorders in plasma
membrane.

The differenses in the electrophoretic properties in SDS of the
hydrophobic proteins of the Na+,K+-ATP-ase (-subunit isoforms after
thermal solubilization of the cerebral cortex preparations prove that
isoforms differ in their protein-detergent interactions.

The different susceptibility to thermal inactivation at 50-550C of brain
Na+,K+-ATP-ase isoforms (higher for (+) and (1-isoform of brain and
kidney (higher for brain) reflects the differences in their structural
stability in membrane.

The higher sensitivity of (1-isoform of membrane-bound Na+,K+-ATP-ase
from cerebral cortex (in comparison to (+) to phospholipase A2 from Naja
naja oxiana venom was ascertained revealing the peculiarities of the
isoform lipid dependence. Na+,K+-ATP-ase partial inactivation is
accompanied by decrease of their apparent affinity to ouabain. This
phospholipase effect was abolished after treatment of the preparations
by serum albumin to remove fatty acids or after incubation of the enzyme
at the optimal conditions for inhibitor binding.

In contrast to enzyme preparations from medulla oblongata the higher
sensitivity of the Na+,K+-ATP-ase (1-isoform to SDS inactivation
accompanied by its, at least partial, removal from the membrane, was
also found in preparations from cerebral cortex. The differences in the
sensitivity to SDS was eliminated after extraction of microsomes with
the detergent at 370C under conditions of termotropic phase transition
of the membrane lipids. The interpretation of the results is based on
the assumed structural differences of isoform protein-lipid complexes
dependent on the differences in structural organization of the boundary
lipids of the neuronal Na+,K+-ATP-ase catalytic subunit isoforms,thus
determining the differences in the sensitivity of Na+,K+-ATP-ase
isozymes in viscotropic regulation of their functional activity.

The lower oxidant sensitivity of the (1-isoform in «Fe2+/ascorbate»
system was revealed. Thiourea, the hydroxyl radical scavenger,
effectively blocked malonic dialdehyde formation and reduced the rate of
the Na+,K+-ATP-ase inactivation. The relative isoform sensitivity to
oxidation is not changed. It is concluded that Na+,K+-ATP-ase is the
direct target of the oxidative membrane damage specific to SH-dependent
ouabain-sensitive isoforms ((+) even in absence of peroxide lipid
destruction.

The differences of the functional disorders of the «house-keeping»
(1-isoform and specialized (2- and (3-isoforms of the neuronal
Na+,K+-ATP -ase at some brain diseases and their specific sensitivity to
membrane-disturbing agents are supposed.

Key words: Na+,K+-ATP-ase, catalytic subunit isoforms, membrane
perturbation.

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