Закономірності розвитку інтерфазної загибелі лімфоцитів селезінки щурів за участі Сa2+-залежних фосфоліпідних сигнальних систем після радіаційного ура

Ee?anueeee oi?aa?neoao ?iai? Oa?ana Oaa/aiea

IAOEOAANUeEA IEUeAA IAAE?AIA

OAeE: 577.125: 591: 144: 612.014

CAEIIII??IINO? ?ICAEOEO ?IOA?OACII? CAAEAAE? E?IOIOeEO?A NAEAC?IEE UO??A
CA O*ANO? Na2+ — CAEAAEIEO OINOIE?I?AeIEO NEAIAEUeIEO NENOAI I?NEss
?AAe?AOe?EIIAI O?AAEAIIss

03.00.04 — a?io?i?y

AAOI?AOA?AO

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaith

aeieoi?a a?ieia?/ieo iaoe

Ee?a — 1999

Aeena?oaoe??th ? ?oeiien

?iaioa aeeiiaia ia eaoaae?? a?io?i?? Ee?anueeiai oi?aa?neoaoo ?iai?
Oa?ana Oaa/aiea

Iaoeiaee eiinoeueoaio: /eai-ei?. IAI Oe?a?ie, i?ioani?

Eo/a?aiei Ieeiea ?aaeieeiiae, Ee?anueeee

oi?aa?neoao ?iai? Oa?ana Oaa/aiea, caa?aeoaa/

eaoaae?e a?io?i??

Io?oe?ei? iiiiaioe: aeieoi? a?ieia?/ieo iaoe, i?ioani?

Einoa??i Na?a?e Ieaen?eiae/, ?inoeooo

a?io?i?? ?i.I.A.Iaeeaae?ia IAI Oe?a?ie

caa?aeoaa/ a?aeae?eo a?io?i?? i(yc?a

aeieoi? a?ieia?/ieo iaoe

Aeieo?aiei Ieeiea Iao?iae/, ?inoeooo

aeia?a??ie ? oieneeieiaa?? ?i.E.?.Iaaeaaaey

IIC Oe?a?ie, caa?aeoaa/ eaai?aoi??? a?io?i??

aeieoi? a?ieia?/ieo iaoe

Ae?oaeei Ieeiea Ieaenaiae?iae/, ?inoeooo

aenia?eiaioaeueii? iaoieia??, iieieia?? ? ?aae?ia?ieia??

?i.?.?.Eaaaoeueeiai IAI Oe?a?ie, i?ia?aeiee

iaoeiaee ni?a?ia?oiee

I?ia?aeia onoaiiaa: Iaoe?iiaeueie iaaee/iee oi?aa?neoao ?iai?

I.I.Aiaiiieueoey, i.Ee?a

Caoeno a?aeaoaeaoueny 25 eenoiiaaea 1999 ?. i 14.00 aiaeei? ia
can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae 26.001.24 o Ee?anueeiio
oi?aa?neoao? ?i.O.Oaa/aiea ca aae?anith: i?ini.aeaaethAeooeiaa 2,
a?ieia?/iee oaeoeueoao, aoae.215.

Iiooiaa aae?ana: 01033, Ee?a — 33, aoe.Aieiaeeie?nueea, 64.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? Ee?anueeiai
oi?aa?neoaoo ?iai? Oa?ana Oaa/aiea ca aae?anith: aoe.Aieiaeeie?nueea,
58.

Aaoi?aoa?ao ?ic?neaie 25 aeiaoiy 1999 ?ieo.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee A?aeii I.A.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue ?iaioe. I?ioyaii inoaii?o ?ie?a iniaeeai? aeooaeueiino?
iaaoee aeine?aeaeaiiy a?io?i?/ieo iaoai?ci?a oi?ioaaiiy ee?oeiii?
a?aeiia?ae? ia ae?th aaiioiene/ieo oaeoi??a ??cii? i?e?iaee, o oiio
/ene? oaeiai, ye ?ii?coth/a ?aae?aoe?y. Iaeiei c aa??aio?a ee?oeiii?
a?aeiia?ae? aaaaoiee?oeiieo nenoai, iai?aaeaii? ia i?aeo?eiaiiy
ee?oeiiiai aiiainoaco oa ia caiia?aaiiy ia?a?iaeaeaiiy ee?oei aiane?aeie
ae?? iioeiaeaeoth/iai oaeoi?a ? aiiioic, aai i?ia?aiiaaia caaeaaeue
ee?oei o ?ioa?oac? (Kerr J.,1980.; Wyllie A.,1987).

Cia/iiai oni?oo o c’ynoaaii? a?io?i?/ieo caeiiii??iinoae ?icaeoeo
aiiioico aoei aeinyaiooi caaaeyee aeine?aeaeaiith ?aae?aoe?eii? caaeaae?
e?ioi?aeieo ee?oei. E?ioioeeoe — ?aeei? c ee?oei ?ioiii? nenoaie, ye?
aeiooue i?iii?oe?eii aei aeice ( o ae?aiacii? aei 3 A? ) aei anooio o
i?oic i?ioyaii ia?oi? aeiae i?ney ae?? ?aae?aoe??. Aenia?eiaioaeueia
iiaeaethaaiiy a?aeiia?ae? e?ioioeeo?a in vitro ia ae?th ?ii?coth/i?
?aae?aoe?? ca oiiae ii?ii?iaiiy ee?oeiii? noniaic?? oeiioeeo?a oa e?i?e
e?ioi?aeieo ee?oei aeicaieeei anoaiiaeoe i?e/eiiee ca’ycie i?ae oaeith
ii?oieia?/iith iciaeith caaeaae? ee?oei, ye i?eiic yaea? oa
i?aeioeeainiiiith aeaa?aaeaoe??th o?iiaoeio (Cohen J., 1992; Zhivotovsky
B. et al., 1981), aeyaeoe caeaaei?noue caaeaae? ee?oei a?ae aeni?an??
aai?a ( Hallahan D., 1992; Wilson L. et al.,1993) oa aeiaanoe, ui
?aae?aoe?eia caaeaaeue e?ioioeeo?a o ?ioa?oac? ? ??ciiaeaei?noth
aiiioico. Aoei aenoiooi a?iioaco, ca?aeii yei? aaciina?aaeiueith
i?oaiith ae?? ?aae?aoe?? ? AeIE, a ae??oaeueio ?ieue o ?i?oe?aoe??
aiiioico a?ae?a?a? ooeiaeaeaiiy aaiaoe/iiai iaoa??aeo, aiane?aeie /iai
aeaaeieo?oueny aaiaoe/ii aeaoa?i?iiaaia i?ia?aia caaeaae? (Duke R. et
al., 1983; Wyllie A., 1985). C ?ioiai aieo, aeine?aeaeaiiy oe?ieiai
niaeo?o ?iaeoeoi??a aiiioico aeyaeee caeaaei?noue ?ioa?oacii? caaeaae?
ee?oei a?ae aioo??oiueiee?oeiiiai aaeaino ?ii?a eaeueoe?y ( Nicotera
P.et al, 1994, Aaaae A. e ae?., 1990) oa a?ae nenoai ee?oeiii?
neaiae?caoe??, ye? caeaoi? iineethaaoe io?eiaiee neaiae oa ia?aaeaaaoe
eiai o?aine?eioe?eiei oaeoi?ai – ?aaoeyoi?ai aeni?an?? aai?a, ui
aeicaiey? aecia/aoe aiiioic ye caaeaaeue a?ae neaaeeo neaiaeueieo
aca?iiae?e (Uckun F. et al., 1993; Aeaeon E.,1997; Ei?unoia A. e ae?.,
1998).

Na?aae nenoai, ye? caaacia/othoue ?aoeaioe?th neaiaeo ia ??ai?
ieaciaoe/ii? iaia?aie e?ioioeeo?a oa eiai ia?aaea/o o
aioo??oiueiee?oeiiee i?ino??, ne?ae ianaiia?aae aeae?eeoe oae? Na2+-
caeaaei? oinoie?i?aei? o?ainaeoeoe?ei? nenoaie, ye iie?oinoi?iiceoeaeia
oa oeeee a?ao?aeiiiai? eeneioe. Eeth/ia? oa?iaioe oeeo nenoai —
neaiaeuei? oinoie?iace N oa A2 aeei?enoiaothoue iaia?aii? oinoie?i?aee
ye aeaea?aei aeey neioaco oi?aa?naeueieo aioo??oiueiee?oeiieo
iina?aaeiee?a – ?iiceoie-1,4,5-o?eoinoaoo, aeeaoeeeae?oea?ieo,
a?ao?aeiiiai? eeneioe oa ?? iaoaaie?o?a. Aioo??oiueiee?oeii? oeyoe
ia?aaea/? neaiaeo ca o/ano? oeeo iina?aaeiee?a, ui aeoeaothoueny o
e?ioioeeoao o aeiaaeeo ?iaeoeiaaii? ?aae?aoe??th ?ioa?oacii? caaeaae?,
cia/iith i??ith ni?aiaaeathoue c oeie, ye? eiio?ieththoue aeoeaaoe?th oa
aeeoa?aioe?aoe?th e?ioioeeo?a o ii?i?. A iaio aeiaaeeao iathoue i?noea
i?aeaeuaiiy eiioeaio?aoe?? eii?a eaeueoe?th o oeeoiieaci?, iineeaiiy
a?ae?ie?co iie?oinoi?iiceoeae?a, ?ioaineo?eaoe?y iai?io a?ao?aeiiiai?
eeneioe, aeni?an?y aai?a n-fos oa c-jun oa ?io.(Aoaeieoeeay A., 1986;
Uckun F. et al.,1992; Oaiioieeiaa A. e ae?.,1998). Caeeoa?oueny
a?aee?eoei ieoaiiy i?i oa, yeei /eiii aeoeaaoe?y o e?ioioeeoao
iaeiaeiaeo ?aaoeyoi?ieo iaoai?ci?a iiaea i?eaiaeeoe aei i?ioeeaaeieo
iane?aee?a. Aaaeeeaiai cia/aiiy aeey ae??oaiiy oe??? i?iaeaie iaaoaa?
iiooe iiaeeeaeo oeyo?a ia?ao?auaiiy ??cieo neaiaeueieo oeyo?a oa
aeoeaaoe?? iaoai?ci?a, ui ? niaoeeo?/ieie o?eaa?aie aiiioico, oaeeo,
ye aeoeaaoe?y no?iaii??e?iace, aeoeaaoe?y oeenoa?iiaeo i?ioa?iac
n?iaenoaa ?NA oa ?io. (Kolesnick R. & Kronke M, 1994; Kidd V., 1998).

Iaaeinoaoiuei aeine?aeaeai? iniaeeaino? ?ioa?oacii? caaeaae? e?ioioeeo?a
ca ii?ii?iaiiy oe?eiai i?aai?cio, oi/a naia oeae oei caaeaae? aecia/a?
oaeee niaoeeo?/iee i?iya i?iiaiaai? oai?iae nnaaoe?a, ye e?ioiiai?y
(ss?eeei A., 1988; Howie S. et al., 1994). Aaaaea?oueny anoaiiaeaiei, ui
i?eai?/aiiy ?ioiii? a?aeiia?ae? i?aai?cio i?ney ae?? ?aae?aoe??
coiiaeaia aea??eiaith ?ioa?oaciith caaeaaeeth e?ioioeeo?a aiane?aeie
ii?ooaiiyi ?aaoeyoi?ii? aca?iiae?? i?ae ??cieie noaiiioeyoe?yie
e?ioioeeo?a o i?aaiao e?ioiiiaco – e?noeiaiio iiceo, oeion? oa
naeac?ioe? (Ai?eciioia I. e ae?.,1983; Meyn R.et al., 1993). Oiio
aaaeeeao ?ieue o iaoai?ciao aaiioiene/ii? ae?? ?aae?aoe?? iaaeathoue
ia?ae?eiiei ianaiaeaea?ai –eaeeio???iai, oeeoie?i?iai, oaeoi?ai ?inoo,
ye? ?aaoeththoue o?eaaeo o /an? i?aeee?oeiio aca?iiae?th e?ioioeeo?a
??cieo noaiiioeyoe?e oa coiiaeththoue oiia?eueiaio e?iaoeeo ?iaeoeoe??
aai?a — ?aaoeyoi??a aiiioico ( Yeaeon E.1996; Wilson R. et al., 1993; Wu
X.et al., 1995).

O ca’yceo c oeei iniaeeai? aeooaeueiino? iaaoaa? aeine?aeaeaiiy o/ano?
??cieo aoi?eiieo iina?aaeiee?a oa aca?iiae?? neaiaeueieo nenoai ca
?icaeoeo ?iaeoeiaaii? a oiiaao in vivo ?aae?aoe?eii? ?ioa?oacii?
caaeaae? ee?oei o iiioeyoe?? e?ioioeeo?a oaeiai aaaeeeaiai
e?ioi?aeiiai i?aaio, ye naeac?iea.

Iaoa oa caaea/? aeine?aeaeaiiy. Iaoith ?iaioe aoei c’ynoaaoe ?ieue
eiiiiiaio?a Na2+ – caeaaeieo oinoie?i?aeieo o?ainaeoeoe?eieo nenoai o
?i?oe?aoe?? oa i?iaaaeaii? ?iaeoeiaaiiai ?ii?coth/ith ?aae?aoe??th
aiiioiaaiiiai neaiaeo o e?ioioeeoao naeac?iee uo??a.

Aoee iinoaaeai? oae? caaea/?:

— ia iniia? e?euee?niiai aiae?co oa ii??aiyiiy ii?oieia?/ieo ?
a?io?i?/ieo iieaciee?a noaio ee?oei o iiioeyoe?? e?ioioeeo?a naeac?iee
eiio?ieueieo oa ii?ii?iaieo ?aioaai?anueeeie i?iiaiyie uo??a aecia/eoe
aeico ?ii?coth/i? ?aae?aoe?? oa oa?i?i i?ney aieeao, ca yeeo
?i?oe?th?oueny ?ioa?oacia caaeaaeue e?ioioeeo?a;

— aecia/eoe eiioeaio?aoe?th a?eueiiai oeeoicieueiiai Na2+ o e?ioioeeoao
naeac?iee oa i?iieei?noue ieaciaoe/ii? iaia?aie aeey Na2+ ca ?icaeoeo
?ioa?oacii? caaeaae? ee?oei;

aeine?aeeoe ca’ycie i?ae no?oeoo?iei noaiii o?iiaoeio oa
eiioeaio?aoe??th oeeoiieaciaoe/iiai eaeueoe?th o e?ioioeeoao;

aeine?aeeoe aeoeai?noue oinoie?iace N oa ooai?aiiy ?iiceoieoinoao?a o
e?ioioeeoao naeac?iee o eiio?ie? oa i?ney ia?iaee ee?oei
iie?eeiiaeueieie i?oiaaiaie oa eaeueoe??aei ?iiioi?ii;

aeine?aeeoe aeoeai?noue oinoie?iace N oa ooai?aiiy ?iiceoieoinoao?a o
e?ioioeeoao naeac?iee o aeeiai?oe? oi?ioaaiiy a?aeiia?ae? ee?oei ia
ae?th ?aae?aoe??;

— aeine?aeeoe aeoeai?noue oinoie?iace A2 oa aea?eueiaiiy a?ao?aeiiiai?
eeneioe c iaia?aiieo oinoie?i?ae?a o e?ioioeeoao naeac?iee o eiio?ie? oa
i?ney ia?iaee ee?oei eaeueoe??aei ?iiioi?ii;

aeine?aeeoe aeoeai?noue oinoie?iace A2 oa aea?eueiaiiy a?ao?aeiiiai?
eeneioe c iaia?aiieo oinoie?i?ae?a e?ioioeeo?a naeac?iee ca ?icaeoeo
?ioa?oacii? caaeaae? ee?oei;

aeine?aeeoe aeoeai?noue e?iieneaaiace oa ai?no ia?aeiieo i?iaeoeo?a
e?iieneaaiaciiai ia?aoai?aiiy a?ao?aeiiiai? eeneioe o e?ioioeeoao
naeac?iee o aeeiai?oe? oi?ioaaiiy a?aeiia?ae? ee?oei ia ae?th
?aae?aoe??;

— aeine?aeeoe ca’ycie i?ae oa?iaioaoeaiei ia?aieeneaiiyi e?i?ae?a ii
e?iieneaaiaciiio oeyoo oa aeaa?aaeaoe??th AeIE ca ?icaeoeo ?ioa?oacii?
caaeaae? e?ioioeeo?a naeac?iee c aeei?enoaiiyi niaoeeo?/iiai ?ia?a?oi?a
e?iieneaaiace ;

?ic?iaeoe noaio oeyo?a ia?aaea/? oa iineeaiiy ?iaeoeiaaiiai
?aae?aoe??th aiiioiaaiiiai neaiaeo o e?ioioeeoao ca o/ano? Na2+-
caeaaeieo oinoie?i?aeieo o?ainaeoeoe?eieo nenoai .

Iaoeiaa iiaecia ? i?aeoe/ia cia/aiiy ?iaioe. O ?acoeueoao? i?iaaaeaieo
aeine?aeaeaiue aia?oa c’yniaaia ?ieue Na2+- caeaaeieo oinoie?i?aeieo
o?ainaeoeoe?eieo nenoai o ?icaeoeo ?ioa?oacii? caaeaae? e?ioioeeo?a
naeac?iee ca ?aae?aoe?eiiai o?aaeaiiy i?aai?cio. I?iaaaeaiee
ii??aiyeueiee aiae?c ia?aiao??a ooieoe?iioaaiiy iie?oinoi?iiceoeaeii?
nenoaie oa iaoaaie?cio a?ao?aeiiiai? eeneioe oa aeeiai?ee ?icaeoeo in
vivo ii?oieia?/ieo oa a?io?i?/ieo iciae ?icaeoeo ?ioa?oacii? caaeaae?
e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei.

Aia?oa aeyaeaii ?aii? c?inoaiiy eiioeaio?aoe?? a?eueiiai oeeoicieueiiai
Na2+ o e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei ca ?icaeoeo ?ioa?oacii?
caaeaae?. Iieacaii, ui i?aeaeuaiiy eiioeaio?aoe?? oeeoicieueiiai Na2+ o
ee?oeiao ia?aaeo? i?aeioeeainiii?e o?aaiaioaoe?? o?iiaoeio, a
iiaeiaaeaia oo?eiaiiy i?aeaeuaiiai ??aiy [Ca2+]i aiane?aeie c?inoaiiy
eaeueoe??ai? i?iieeiino? ieaciaoe/ii? iaia?aie i?enei?th? o?aaiaioaoe?th
AeIE.

Iieacaii, ui ia ii/aoeia?e noaae?? ?icaeoeo ?ioa?oacii? caaeaae?
e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei a?aeaoaa?oueny aeoeaaoe?y
oinoie?iace N oa iiaeeo?eaoe?y aca?iiae?? eiiiiiaio?a o nenoai?
iiaa?oiaaee ?aoeaioi?-iaia?aiica’ycai? ?aaoeyoi?i? a?eee – oinoie?iaca
N. Aia?oa anoaiiaeaii, ui ia ?aiiueiio aoai? ?icaeoeo ?ioa?oacii?
caaeaae? o e?ioioeeoao naeac?iee iineeth?oueny neioac iniiaiiai
Na2+-iia?e?coth/iai iina?aaeieea — ?iiceoie-1,4,5-o?eoinoaoo.

Aeine?aeaeai? aeanoeaino? oinoie?iace A2 e?ioioeeo?a naeac?iee, ui
a?ae?ie?co? a?ao?aeiii?eai?ni? oinoie?i?aee oa ?aeaioeo?eiaai?
aiaeiaaii? noano?aoe oa?iaioo. Iieacaii, ui o i?ioean? ?iaeoeiaaii?
?aae?aoe??th ?ioa?oacii? caaeaae? e?ioioeeo?a naeac?iee a?aeaoaa?oueny
aea?eueiaiiy a?ao?aeiiiai? eeneioe c? neeaaeo iaia?aiieo oinoie?i?ae?a
oa iaeii/ania i?aeaeuaiiy o iaia?aii?e o?aeoe?? ee?oei aeoeaiino?
oinoie?iace A2, ui a?ae?ie?co? oinoaoeaeeeoie?i oa
oinoaoeaeeeaoaiieai?i.

Anoaiiaeaii, ui e?ioioeeoe naeac?iee i?iyaeythoue e?iieneaaiacio
aeoeai?noue oa ?aeaioeo?eiaaii aiaeiaaii? i?iaeoeoe e?iieneaaiaciiai
ia?aoai?aiiy a?ao?aeiiiai? eeneioe. Ciaeaeaii, ui o i?ioean? ?icaeoeo
?aae?aoe?eii? caaeaae? e?ioioeeo?a naeac?iee o ?ioa?oac? a?aeaoaa?oueny
?ioaineo?eaoe?y oa?iaioaoeaiiai ia?aieeneaiiy noano?ao?a e?iieneaaiacith
oa i?aeaeuaiiy ai?noo 15-a?ae?ieneaeeicaoao?a?iiai? eeneioe oa
eaeeio???io A4. Aeiaaaeaii, ui niaoeeo?/ia i?eai?/aiiy e?iieneaaiaciiai
oeyoo ia?aieeneaiiy a?ao?aeiiiai? eeneioe i?ecaiaeeoue aei cia/iiai
cieaeaiiy i?aeioeeainiiii? o?aaiaioaoe?? AeIE o e?ioioeeoao naeac?iee
ii?ii?iaieo oaa?ei.

Ia iniia? io?eiaieo aeaieo oa aeaieo e?oa?aoo?e ?ic?iaeaia noaia oeyo?a
ia?aaea/? oa iineeaiiy ?iaeoeiaaiiai ?aae?aoe??th aiiioiaaiiiai neaiaeo
o e?ioioeeoao naeac?iee ca o/ano? ?ii?a eaeueoe?th, ie?aieo eiiiiiaio?a
Na2+- caeaaeieo oinoie?i?aeieo o?ainaeoeoe?eieo nenoai oa oaeoi??a, ui ?
niaoeeo?/ieie o?eaa?aie aiiioico.

?acoeueoaoe i?iaaaeaieo aeine?aeaeaiue iathoue caaaeueiioai?aoe/ia
cia/aiiy aeey aaeaeaaoii? a?io?i?/ii? ioe?iee ieaeio?iiieo aoaeo?a
aeno?aiaeueieo iao?c?ieia?/ieo oa oeeoioiene/ieo oaeoi??a.
Aeine?aeaeaiiy oeyo?a ia?aaea/? neaiaeo aei ?icaeoeo ?ioa?oacii?
caaeaae? e?ioioeeo?a ? iaiao?aeiei aoaiii aeey anoaiiaeaiiy eiie?aoieo
a?io?i?/ieo iaoai?ci?a ?aae?caoe?? aiiioico ee?oei ??cieo oei?a.
?aeaioeo?eaoe?y ie?aieo neaiaeueieo oeyo?a aiiioico ia? aaaeeeaa
i?aeoe/ia cia/aiiy, ine?eueee caaacia/o? iiaeeea?noue ?ic?iaee
i?aeoiae?a aei eiio?ieth iaae oeei i?ioeanii, cie?aia, aei iai?aaeaiiai
iiooeo i?aia?ao?a, caeaoieo ?i?oe?thaaoe ?ioa?oacio caaeaaeue
ia?a?iaeaeaieo ceiye?nieo ee?oei aai ae iiia?aaeaeaoe caaeaaeue
ii?iaeueieo ee?oei o aeiaaeeo i?iiaiaai? oa?ai?? aai ae?? ?ioeo
iao?c?ieia?/ieo oaeoi??a.

Iniaenoee aianie aaoi?a. Aeena?oaioii iniaenoi cae?eniaiee
?ioi?iaoe?eiee iiooe oa ioe?iea e?oa?aoo?ieo aeaieo, ?ic?iaeaia noaia
oa iaa?oioiaaia iaoiaeieia?y iinoaiiaee aenia?eiaio?a, aeeiiai?
aenia?eiaioaeuei? aeine?aeaeaiiy, i?iaaaeaiee aiae?c io?eiaieo
?acoeueoao?a, i?aeaioiaeai? ae?oeiaai? i?aoe?. ?acoeueoaoe aeayeeo
i?ae?icae?e?a io?eiai? aaciina?aaeiuei aaoi?ii oa ca o/ano? ani??aio?a
eaoaae?e a?io?i?? (Ne?iaaoeue O.I. — i?ae?icae?e 3.2.1., Neaoa?inueei?
I.A.- i?ae?icae?e 3.3.3, Ianoooa A.I.- i?ae?icae?e 3.3.4., Nieiaeooei
A.I – i?ae?icae?e 3.1.1). Io?eiai? ?acoeueoaoe aeeeaaeai? o ni?eueieo
ioae?eaoe?yo. Aeaeo?iiii-i?e?ineii?/iee aiae?c i?aia?ao?a e?ioioeeo?a
naeac?iee cae?eniaiee o ni?a?ia?oieoeoa? c? n.i.n. ?inoeoooo a?io?i??
?i. I.A. Iaeeaae?ia *a?ieoiaei A.?.

Ca’ycie c iaoeiaith oaiaoeeith i?aai?caoe??. ?iaioa a?aeiia?aea? ieaio
iaoeiai-aeine?aeii? ?iaioe eaoaae?e a?io?i?? a?ieia?/iiai oaeoeueoaoo
Ee?anueeiai oi?aa?neoaoo ?iai? Oa?ana Oaa/aiea oa aeeiiaia o ?aieao oaie
N 97083 » A?io?i?/i? iaoai?cie ?icaeoeo i?iiaiaaiai o?aaeaiiy i?ney
ae?? ?ii?coth/i? ?aae?aoe?? » eiiieaenii? iaoeiai? i?ia?aie «Caei?ia’y
ethaeeie», a oaeiae oai 5.3/ 34 oa 5.4/483 I?i?noa?noaa Oe?a?ie o
ni?aaao iaoee ? oaoiieia??.

Ai?iaaoe?y ?iaioe. Iaoa??aee aeena?oaoe?? aoee i?aaenoaaeai? ia VI
(Ee?a, 1992) oa VII (Ee?a, 1997) Oe?a?inueeeo a?io?i?/ieo c’?caeao, II (
Ee?a, 1993) oa III (Iineaa, 1997) ?aae?ia?ieia?/ieo c’?caeao,
i?aeia?iaei?e iaoeia?e eiioa?aioe?? » Iaaeieeoi? na?aaeiaeua ? caei?ia’y
» ( *a?i?aoe?, 1993), I Oe?a?inueeiio c’?cae? a?io?ce/iiai oiaa?enoaa (
Ee?a, 1994), I i?aeia?iaei?e i?aeoe/i?e eiioa?aioe?? » Sustainable
Development: Environmental Pollution and Ecological Safety»(
Aei?i?iiao?ianuee, 1995), O I?aeia?iaeiiio eiia?an? c ?aae?aoe?eieo
aeine?aeaeaiue ( Ath?oeao?a, I?ia//eia, 1995), eiioa?aioe?? »
A?aeaeaeai? iane?aeee ii?ii?iaiiy a ?ioii?e nenoai? oa aaiiiiaoe/i?e
nenoaiao» ( Ee?a, 1996), eiioa?aioe?? » I?iaeaiu i?ioeaieo/aaie caueou»
( Iineaa, 1998).

Ioae?eaoe??. ?acoeueoaoe aeine?aeaeaiue i?aaenoaaeai? o 20 noaooyo oa
13 oacao, ye? iioae?eiaai? o i?io?eueieo a?o/eciyieo oa ca?oa?aeieo
aeo?iaeao, ca??ieeao iaoeiaeo i?aoeue oa iaoa??aeao c’?cae?a oa
eiioa?aioe?e.

No?oeoo?a aeena?oaoe??. Aeena?oaoe?eia ?iaioa aeeeaaeaia ia …noi??ieao
iaoeiiieniiai oaenoo ? neeaaea?oueny ?c anooio, iniiaii? /anoeie, ui
i?noeoue iaeyae e?oa?aoo?e ( 4 ?icae?ee) oa aenia?eiaioaeueio /anoeio (3
?icae?ee), caeeth/ii? /anoeie (1 ?icae?e), aeniiae?a oa oeeoiaaii?
e?oa?aoo?e ( …. aeaea?ae). ?iaioa i?noeoue ….?enoie?a , …. oaaeeoeue
oa 1 noaio.

IAEssAe E?OA?AOO?E

A iaeyae? e?oa?aoo?e aeieeaaeii ?icaeyioo? ieoaiiy a?io?i?/ii?
?aaoeyoe?? i?io?eaiiy i?ioean?a aeoeaaoe??, aeeoa?aioe?aoe?? oa
aiiioe/ii? caaeaae? e?ioi?aeieo ee?oei. I?iaaaeaiee aeaoaeueiee aiae?c
no/anieo oyaeaiue i?i iieaeoey?i? iaoai?cie oa ooieoe?iiaeueia cia/aiiy
oaeeo ?aaoeyoi?ieo nenoai, ye Na2+-iia?e?coth/a iie?oinoi?iiceoeaeia oa
oeeee a?ao?aeiiiai? eeneioe. Aena?oeaii a?io?i?/i? aniaeoe ?icaeoeo
?ioa?oacii? caaeaae? ee?oei, aeeeeeaii? ?ii?coth/ith ?aae?aoe??th.

IAOA??AEE OA IAOIAeE AeINE?AeAEAIUe

O ?iaio? aoee aeei?enoai? oae? ?aaaaioe oa iaoa??aee: Ficoll-Paque (
Pharmacia, Oaaoe?y), ii?aeea?ae?iaoaea?aoeeiaa eeneioa (Serva,
I?ia//eia), ea?ia-2 aoeaoieneiaoeeiaee ao??(Amersham, Aaeeea A?eoai?y),
IA?AS, ?iiioi? A23187, o?oiaaiaaethoei?i oa e?iiiie?naoa?eae E.
coli, eoa?ie ?O, e?iieaaa eeneioa, o?eoii O-100 (Ferak, I?ia//eia),
14N-a?ao?aeiiiaa eeneioa, 14N-2- a?ao?aeiii?eoinoie?i?aee, 14N-2-
e?iiea?eoinoaoeaeeeaoaiieai?i, oinoaoeaeee-2-3I-?iiceoie,
3I-2-i?i?iiceoie, (Amersham, Aiae?y), iaai?e aeey aecia/aiiy ai?noo
15-IAOA ( Advanced Magnetic Inc, COA), eaeeio???io A4,
i?inoaaeaiaeei?a F2( oa E2 ( Amersham, Aiae?y); ieanoeie aeey
oiieioa?iai? o?iiaoia?ao??, n?e?eaaaeue RH 40/5, o?eueo?e «Neiii?» N6
(Oaiaiie, *ao?y), eieiiee Particil 10 SAX aeey ai?iiiiai?iii?
o?iiaoia?ao?? ?iiceoieoinoao?a (Whatman, Aiae?y); oinoaoeaeeena?ei,
oinoaoeaeeeoie?i, oinoaoeaeeeaoaiieai?i, oinoaoeaeee?iiceo,
oinoaoeaeeeae?oea?ie (caaiae aaeoa??aeueieo i?aia?ao?a, Oa?e?a);
45NaCl2 , 32?-I3?I4 – i?aei?e?inoai «?cioii», Ee?a.

Aeine?aee i?iaiaeeee ia uo?ao e?i?? A?noa? iaio noaoae ianith 130 –150
a. Oaa?ei i?aeaeaaaee oioaeueiiio iaeii?aciaiio ii?ii?iaiith ia
?aioaai?anuee?e onoaiiaoe? ?OI-17 ca oaeeo aeniiceoe?eieo aeic – 1,29(
10-2 Ee/ea ( iiaeeiooa aeica – 0,5 A?) oa 2,58( 10-2 Ee/ea ( iiaeeiooa
aeica – 1 A?). Oiiae ii?ii?iaiiy: iiooaei?noue aeice– 0, 2 A?/oa,
o?eueo?e 0,5 ii ( Cu + Al), iai?oaa 180 eA, neea no?oio 5 iA, oieonia
a?aenoaiue – 50 ni.

E?ioioeeoe io?eioaaee iaoiaeii a?aae??ioiiai oeaio?eooaoaaiiy noniaic??
ee?oei naeac?iee o ?ic/ei? Ficoll-Paque aai o?eie-aa?ia?ao?io ( ( =
1,077) (Boyum A., 1968). I?ney ?oeioaaiiy ee?oei o a?iiiniioe/iiio
aooa?? oa oeaio?eooaoaaiiy io?eioaaee o?aeoe?? aac’yaea?iiai aiiiaaiaoo
( noia?iaoaio 1000g, 5 oa), oeeoicieueio oa iaia?aiio ( 100000g, 60 oa).

Eiioeaio?aoe?th a?eueiiai Na2+ o ee?oeiao aecia/aee c canoinoaaiiyi
oeoi?anoeaioiiai ciiaeo ea?i-2 (Tsien R.et al.,1982). Oeoi?anoeaioe?th
noniaic?? ee?oei aei??thaaee ia niaeo?ioethi?eiao?? Shimadzu RF-510
( ssiii?y), ( caoaeaeaiiy — 339 ii, ( ai?n?? – 490 ii.

Ai?no iaeiiiaiai aeeaeueaeaa?aeo aecia/aee ca ?aaeoe??th c
o?iaa?a?oo?iaith eeneioith ( Noaeueiay E. e Aa?eoaeee O., 1977).
Aioeieenethaaeueio aeoeai?noue aecia/aee ca eiinoaioith i?eai?/aiiy
?aaeoe?? ieeneaiiy 2,4- aeeoei?oaiie?iaeioaiieo a?ieia?/iei iaoa??aeii
(Naiaiia A.,1985).

Ioe?ieo o?aaiaioaoe?? AeIE i?iaiaeeee i?ney e?ceno ee?oei ?ic/eiii c 2
iI AAeOA oa 0,5 % o?eoiiii O-100. I?iae oeaio?eooaoaaee ( 15000 g, 20
oa) aeey ?icae?eaiiy ?ioaeoiiai o?iiaoeio (inaae) a?ae a?aeuaieaieo
o?aaiaio?a AeIE (noia?iaoaio). Ai?no AeIE o c?aceao aecia/aee ?c
canoinoaaiiyi aeeoai?eai?iiaiai ?aaeoeao ( Burton K.,1956).

AeIE aeno?aaoaaee noi?ooth oei?ioi?i : ?ciai?eiaee nie?o (24:1) i?ney
aeai?ioa?i?caoe?? i?ia (Skalka M. et al., 1976). Aeaeo?ioi?ac AeIE
i?iaiaeeee o 1% aaa?iciiio aae? ca iai?oae iiey 2 iA/nae i?ioyaii 15
aiae (200 N). O yeino? ia?ea?ieo aeei?enoiaoaaee o?aaiaioe AeIE oaaa (.,
ooai?ai? ?ano?eeoacith Pst I. Aae? oa?aoaaee ?ic/eiii a?ii?aeo aoeae?th
(1 iea/ie) ? i?ney caoaeaeaiiy oeoi?anoeaioe?? oeueo?ao?ieaoii
oioia?aooaaee ia ie?aeo I?e?ao-300. Iaaaoeae aeaineoiiao?oaaee ia
i?e?iaeaineoiiao?? MD-100 ( I?ia//eia).

Ao?ae eaeueoe?th o e?ioioeeoe aecia/aee ca ?ieoaoaaiiy noniaic??
e?ioioeeo?a o noaiaea?oiiio nieueiaiio na?aaeiaeu?, ui i?noeei 1 ieE?
45NaCl2. ?aaeoe?th coieiyee oaeaeeith o?eueo?aoe??th i?ia /a?ac
o?eueo?e Neiii? N 6.

Ioe?ieo ooai?aiiy ?iiceoieoinoao?a i?iaiaeeee c aeei?enoaiiyi
ee?oei, ?ieoaoaaieo i?ioyaii 1 aiae o na?aaeiaeu? 199, ui i?noeei 5
ieE? /ie 3H-2-i?i-?iiceoieo. O na?aaeiaeua ?ieoaaoe?? iiia?aaeiuei
i?/aieo ee?oei aeiaeaaaee 0,1% ANA oa 10 iI LiCl aeey i?eai?/aiiy
aeoeaiino? oinoiiiiianoa?ac. ?aaeoe?th coieiyee aeiaeaaaiiyi noi?o?
oei?ioi?i:iaoaiie (1:2). Aiaeio oaco aeei?enoiaoaaee aeey
?icae?eo ?iiceoieoinoao?a ia eieiieao Partisil 10 SAX. Aethoe?th
?iiceoie-1-oinoaoo, ?iiceoie-1,4-aeeoinoaoo oa ?iiceoie-1,4,5-o?eoinoaoo
i?iaiaeeee iine?aeiaii, aeei?enoiaoth/e ii 2 ie 0,2I, 0,4I oa 0,8I
EI2 ?I4 (?I 3,35) a?aeiia?aeii (Sugiya I. et al., 1987).

Aeoeai?noue oinoie?iace N aecia/aee o na?aaeiaeu?, ui i?noeei 0,02 ieE?
oinoaoeaeee-2- 3H- ?iiceoieo ( 19,9 E?/iiieue) (Carter I. et al., 1987).
?aaeoe?th coieiyee aeiaeaaaiiyi noi?o? oei?ioi?i : iaoaiie : HCl ( 100
: 200 : 0,3 ), 0,6 ie oei?ioi?io oa 0,6 ie aiaee. E?euee?noue
a?aeuaieaiiai ?iiceoieo aecia/aee ca ?aae?iaeoeai?noth aa?oiuei? oace.

Aeey aeeth/aiiy a?ao?aeiiiai? eeneioe e?ioioeeoe ?ieoaoaaee o
na?aaeiaeu? 199, ui i?noeei 0,5 ieE? 14N- a?ao?aeiiiai? eeneioe ( 56
iE?/iiieue) i?ioyaii 1 aiae i?e 370N. E?i?aee aeno?aaoaaee noi?ooth
oei?ioi?i : iaoaiie : ICl (1 : 2 : 0,06). Oiieioa?iao o?iiaoia?ao?th
e?i?ae?a i?iaiaeeee ia ieanoeiao “Oaiaiie” o nenoai? oei?ioi?i : iaoaiie
: ai?ae ( 68 : 28 : 4) ( Hirata F. et al.,1978). ?iciiae?e 14N-
a?ao?aeiiaoo iii?ae oinoie?i?ae?a ioe?ithaaee ca ?aae?iaeoeai?noth ieyi.

Aeey ioe?iee aea?eueiaiiy a?ao?aeiiiai? eeneioe iiia?aaeiuei i?/ai? ca
14N — a?ao?aeiiaoii e?ioioeeoe ?ieoaoaaee o noaiaea?oiiio nieueiaiio
?ic/ei?, ui i?noea aeae?i?aeeciaaiee 0,2 % ANA aeey ca’ycoaaiiy
aea?eueiaii? aee?ii? eeneioe. ?aaeoe?th coieiyee, aeiaeath/e oieiaeiee
?ic/ei 10 iI EI2?I4 (?I 7,4). I?ney oeaio?eooaoaaiiy i?ia ( 1000g, 5 oa)
aecia/aee ?aae?aeoeai?noue iaaeinaaeiai? ??aeeie.

Aeoeai?noue oinoie?iace A2 aecia/aee o na?aaeiaeu?, ui i?noeei 0,05
ieE? 1-noaa?i?e-2-14N-a?ao?aeiii?e-OO aai 1-aoeee-2-14N-a?ao?aeiii?e –
OA (56 iE?/iiieue) (Katsumata M. et al.,1986). ?aaeoe?th coieiyee,
aeiaeath/e 5% ?ic/ei o?eoiio — O-100 oa 40 iI AAeOA. A?ao?aeiiiao
eeneioo aeno?aaoaaee, aeiaeath/e 150 ia noeueoaoo iao??th oa 2,5 ie
aaenaio. ??aaiue aea?eueiaiiai a?ao?aeiiaoo ?ic?aoiaoaaee ca
?aae?iaeoeai?noth ae?eaioe aaenaio i?ney ?icae?eaiiy oac.

Aeoeai?noue e?iieneaaiace aecia/aee niaeo?ioioiiao?e/iei niiniaii c
aeei?enoaiiyi e?iieaai? eeneioe (Aooiae/ E. e Eooa?ue A.,1989).
?aaeoe?th i?iaiaeeee o oa?iinoaoiaaieo ethaaoao niaeo?ioioiiao?a Specord
M-40 i?e 35 0N, ?a?no?oth/e i?e??no iioe/iiai iiaeeiaiiy i?e 234ii,
ca’ycaiiai c ooai?aiiyi a?ae?iia?ieneaeieo iio?aeieo e?iieaai? eeneioe.

Aecia/aiiy ai?noo iaoaaie?o?a a?ao?aeiiiai? eeneioe o ee?oeiao
i?iaiaeeee ?aae?i?ioiiei iaoiaeii i?ney aeno?aaoaaiiy aeeicaii?ae?a
oieiaeiei aoeeaoeaoaoii c aeei?enoaiiyi iaai??a aeey aecia/aiiy
eaeeio???io A4 (Amersham, Aiae?y) oa 15 -IAOA (Advanced Magnetic Inc,
NOA) ca?aeii ?aeiiaiaeaoe?e o??ie – ae?iaieea.

Noaoenoe/io ia?iaeo aeaieo i?iaiaeeee caaaeueiii?eeiyoeie iaoiaeaie
aa??aoe?eii? noaoenoeee (Ieioeineee I.,1981) ?ic?aooiee oa iiaoaeiaa
a?ao?e?a i?iaiaeeeenue ia IBM PC-486 c aeei?enoaiiyi i?eeeaaeieo
i?ia?ai.

?ACOEUeOAOE OA IAAIAI?AIIss

Ii?oieia?/ia oa a?io?i?/ia ioe?iea noaio e?ioioeeo?a naeac?iee i?ney
?aioaai?anueeiai ii?ii?iaiiy oaa?ei. Ia?oi/a?aiaei ioieoii
aeine?aeaeaiiy oi?ioaaiiy in vivo a?aeiia?ae? e?ioioeeo?a naeac?iee ia
ae?th ?ii?coth/i? ?aae?aoe?? aoei io?eiaiiy iiioeyoe?? ?aae?i/ooeeaeo
ee?oei naeac?iee oa ii?oieia?/ia ioe?iea ?o noaio o aeeiai?oe? i?ney
?aioaai?anueeiai ii?ii?iaiiy oaa?ei. Ca?aeii aeaieo
aeaeo?iiii-i?e?ineii?/iiai aiae?co, io?eioaaiee i?aia?ao e?ioioeeo?a
naeac?iee i?aaenoaaeaiee iaeeie oa na?aaei?ie e?ioioeeoaie c
??aiii??iei ?ici?uaiiyi aoo?iiaoeiiaeo oa aaoa?io?iiaoeiiaeo cii o
yae??. Oea ee?oeie, ui ia?aaoaathoue o noaae?yo G0 oa G1 ee?oeiiiai
oeeeeo ? oa?aeoa?ecothoueny ianoaa?euei?noth aaiiio oa cia/iith
/ooeea?noth aei ae?? ?aae?aoe??.

Aeey c’ynoaaiiy caeaaeiino? o?aaeaiiy e?ioioeeo?a naeac?iee uo??a
a?ae aeice ?aae?aoe?? oaa?ei i?aeaeaaaee oioaeueiiio ?aioaai?anueeiio
ii?ii?iaiith o aeicao 0,5 oa 1 A? oa aecia/aee ai?no ee?oei o i?aia?ao?
e?ioioeeo?a naeac?iee o oa?i?i aei 6 ae?a. Ca(ycie, yeee
niinoa??aa?oueny i?ae aeicith ii?ii?iaiiy oaa?ei, e?euee?noth caaeaeeo
ee?oei oa ii/aoeii ?o ?aaaia?aoe?? o naeac?ioe? aeicaieea aeei?enoaoe
aaee/eio caaaeueii? e?eueeino? e?ioioeeo?a o iaeeieoe? ia(?io oeaieie
ye e?euee?niee e?eoa??e ioe?iee ?o i?iiaiaaiai o?aaeaiiy. I?ney
ii?ii?iaiiy oaa?ei o aeic? 0,5 A? e?euee?noue e?ioioeeo?a o
aeine?aeaeoaai?e iiioeyoe?? ia ci?ithaaeanue i?ioyaii 2 aiae,
cieaeoaaeanue /a?ac 1 aeiao ia 27,8%, a /a?ac 3 aeiae iiai?noth
a?aeiiaethaaeanue. I?ney ii?ii?iaiiy oaa?ei o aeic? 1 A? cieaeaiiy
e?eueeino? e?ioioeeo?a naeac?iee niinoa??aaeinue o a?eueo ?aii?e oa?i?i
? aoei a?eueo cia/iei – ia 31,9 % /a?ac 12 aiae ? ia 53,2 % /a?ac 1
aeiao, /anoeiaa a?aeiiaeaiiy iaei i?noea /a?ac 6 ae?a. Ioaea, oioaeueia
?aioaai?anueea ii?ii?iaiiy oaa?ei o aeicao 0,5 oa 1 A? i?ecaiaeeoue aei
??ciiai ca oaeaee?noth oa aaee/eiith cieaeaiiy e?eueeino? iaeeo oa
na?aaei?o e?ioioeeo?a naeac?iee i?ioyaii ia?oi? aeiae. I?ioean
nionoioaiiy iiioeyoe?? e?ioioeeo?a i?ney ae?? ?aae?aoe?? aecia/a?oueny
aeaiia iieacieeaie: ?ioa?oaciith caaeaaeeth ee?oei o ?aii?e ia??iae
i?ney ae?? /eiieea oa iiaeaeueoith ?aae?caoe??th o?aaeaiiy o oiae?
?ai?iaeoeoeaii? caaeaae? ee?oei ( Aaeioniaa I. e ae?., 1979; ss?eeei A.
e ae?., 1982). Aeey aecia/aiiy oa?aeoa?o ?aae?aoe?eii? caaeaae? ee?oei
aeine?aeaeoaaee ii?oieia?/i? oa a?io?i?/i? iieacieee noaio e?ioioeeo?a
o oa?i?i, yeee ia?aaeo? iaeneiaeueiiio cieaeaiith ?o e?eueeino? o
iiioeyoe??.

O i?aia?aoao e?ioioeeo?a, io?eiaieo /a?ac 3 aiae i?ney ii?ii?iaiiy
oaa?ei o aeic? 0,5 A?, ii?oieia?/ieo a?aei?iiinoae ii??aiyii c
eiio?ieai ia aeyaeaii. *a?ac 12 aiae o iiioeyoe?? e?ioioeeo?a
aeyaeythoueny ie?ai? ee?oeie c ia??aiii??iei ?ici?uaiiyi
o?iiaoeio oa ciiith ia?eioeeaa?iiai i?ina?oeaiiy, ui ?icoe?iththoue ye
iciaeo aeoeaaoe?? e?ioioeeo?a oa i?ioiaeaeaiiy S – G2 oac ee?oeiiiai
oeeeeo ( Iao?ia ?. e ae?., 1983).

A?eueo cia/i? ii?oieia?/i? ci?ie aeyaeythoueny o i?aia?aoao e?ioioeeo?a
naeac?iee oaa?ei, ii?ii?iaieo o aeic? 1 A? – /a?ac 3 aiae o iiioeyoe??
e?ioioeeo?a i?enooi? ee?oeie c iniio?eueieie neoi/aiiyie a?aioey?iiai
aaoa?io?iiaoeio, cina?aaeaeaieie a?ey iaeiiai c iiethn?a yaea?ii?
iaia?aie, a oaeiae ee?oeie c yae?ii ciaioaieo ?ici???a, caiiaiaiei
eiiaeainiaaiei aaoa?io?iiaoeiii. Oae? ii?oieia?/i? ci?ie
a?aeiia?aeathoue ?aii?i iciaeai ?icaeoeo ?ioa?oacii? caaeaae? ee?oei,
eiee ia? i?noea ooai?aiiy e?oiieo (50 – 300 oeny/ ia? iniia) o?aaiaio?a
AeIE (Sun X. & Cohen G., 1994). *a?ac 12 aiae i?ney ii?ii?iaiiy oaa?ei
o aeic? 1 A? o iiioeyoe?? e?ioioeeo?a naeac?iee, ie??i ee?oei c
i?eiioe/iei yae?ii, aeyaeaii aaeoie?ciaai? ee?oeie oa ioi/ai? iaia?aiith
o?aaiaioe, ui i?noyoue o?iiaoei oa ?ioaeoi? i?aaiaee. Oae? iciaee ?
iieacieeii i?ciuei? noaae?? aiiioico oa ?icaeoeo iae?ioe/ieo i?ioean?a
o ee?oeiao (Columbano A., 1995; Kroemer G. et al., 1998).

Aeey a?io?i?/ii? ioe?iee no?oeoo?iiai noaio o?iiaoeio o e?ioioeeoao
aeine?aeaeoaaee ai?no i?iaeoeo?a aeaa?aaeaoe?? o?iiaoeio –
iecueeiiieaeoey?ieo o?aaiaio?a AeIE iie?aeaceene?eaiioeeaioeae?a (IAeI),
ye? aeno?aaothoueny c yaea? o ?ic/ei c iecueeith ?iiiith neeith
(?en.1). O e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei aeyaeaii
i?aeaeuaiiy /anoee aeaa?aaeiaaii? AeIE, yea caeaaeeoue a?ae aeice
?aae?aoe??. I?ney ii?ii?iaiiy oaa?ei o aeic? 0,5 A? i?aeaeuaiiy ai?noo
IAeI o ee?oeiao aeyaey?oueny /a?ac 12 aiae ? ? iacia/iei – aei 16% a?ae
caaaeueiiai ai?noo AeIE.

I?ney ae?? ?aae?aoe?? o aeic? 1 A? i?aeaeuaiiy ai?noo IAeI o
ee?oeiao aeyaey?oueny /a?ac 6 aiae, /a?ac 12 aiae aiii noaiiaeoue 31 %
a?ae caaaeueiiai ai?noo AeIE. sse iieacaei aeaeo?ioi?aoe/ia ?icae?eaiiy
AeIE o aaa?iciiio aae?, o?aaiaioe AeIE e?ioioeeo?a naeac?iee
ii?ii?iaieo oaa?ei ?iciiae?eyeenue o aeaeyae? nioa, ?ioaineai?noue yeeo
aoea cia/ii a?eueoith o aeiaaeeo ?icae?eaiiy AeIE e?ioioeeo?a oaa?ei,
ii?ii?iaieo o aeic? 1 A? (?en. 2). Ca?aeii ia?ea?iiai aiae?co, aeiaaeeia
o?aaiaio?a o nioaao neeaaeaea 400, 600 oa 800 ia? ioeeaioeae?a, oiaoi
aoea e?aoiith aeiaaeei? AeIE ioeeainii. Ioaea, o e?ioioeeoao naeac?iee
oaa?ei, ii?ii?iaieo o aeic? 1 A?, a?aeaoaa?oueny cia/ia aii?yaeeiaaia
i?aeioeeainiiia ?icuaieaiiy o?iiaoeio ca o/ano? oa?iaio?a, ye?
aoaeothoue e?iea?i? ae?eyiee AeIE. Iaeiei c oaeeo oa?iaio?a ? Mg2+, Ca2+
— caeaaeia aiaeiioeeaaca, ui eiinoeoooeaii aeni?ano?oueny o e?ioioeeoao
oa aeoeao?oueny ca ?icaeoeo ?ioa?oacii? caaeaae? ee?oei (Zhivotovsky B.
et al., 1993; Ieeiiiaa E. e ae?., 1998).

Ii?ooaiiy o nenoai? i?aeo?eiaiiy noaa?eueiino? o?iiaoeio cia/iith i??ith
aecia/a?oueny ci?iith oaeeo ia?aiao??a iaoaaie?/iiai noaio ee?oeie, ye
ieenii-aioeieneaeaioia ??aiiaaaa oa eaeueoe??aee aiiainoac. sse
iieacaei aeine?aeaeaiiy aioeieneaeaioii? aeoeaiino? oa ai?noo iaeiiiaiai
aeeaeueaeaa?aeo, oi?iaeiaae 6 aiae i?ney ii?ii?iaiiy oaa?ei o aeic? 0,5
A? i?ioeane a?eueii?aaeeeaeueiiai ieeneaiiy o e?ioioeeoao naeac?iee
oo?eiothoueny ia i?i?iaeueiiio noaoe?iia?iiio ??ai?. I?ney ae??
?aae?aoe?? o aeic? 1 A? niinoa??aa?oueny i?aeaeuaiiy ai?noo IAeA
ii??aiyii c eiio?ieai o oa?i?i 3 — 6 aiae oa cieaeaiiy aioeieneaeaioii?
aeoeaiino? /a?ac 12 aiae. Ioaea, ii?ii?iaiiy oaa?ei o aeic? 1 A?
i?ecaiaeeoue aei ?aiiueiai cnoao ieenii-aioeieneaeaioii? ??aiiaaae o
e?ioioeeoao o a?e iaaei??ii? ?ioaineo?eaoe?? ?aaeoe?e
a?eueii?aaeeeaeueiiai ia?aieeneaiiy. Ia iaoo aeoieo, aeoeaaoe?y
i?ioean?a ia?aeeniiai ieeneaiiy iaia?aiieo e?i?ae?a, yea cia/iith i??ith
iicia/a?oueny ia no?oeoo?iiio noai? iaia?ai, iiaea i?ecaiaeeoe oaeiae
aei ii?ooaiiy eiiia?oiaioae?caoe?? oaeiai aaaeeeaiai aoi?eiiiai
ianaiaeaea?a, ye eaeueoe?e.

Ca?aeii ?acoeueoao?a, io?eiaieo iaie o aenia?eiaioao c aeei?enoaiiyi
oethi?anoeaioiiai ciiaeo ea?i-2, ii?ii?iaiiy oaa?ei o aeicao 0,5 oa 1 A?
i?ecaiaeeoue aei ii?ooaiiy eaeueoe??aiai aiiainoaco o e?ioioeeoao
naeac?iee (oaae.1). Eiioeaio?aoe?y a?eueiiai oeeoicieueiiai Ca2+ o
e?ioioeeoao eiio?ieueieo oaa?ei noaiiaeea 117 ( 10 iI. I?ney ii?ii?iaiiy
oaa?ei o aeic? 0,5 A? cia/aiiy [Na2+]i o e?ioioeeoao ia
ci?ith?oueny i?ioyaii 3-6 aiae ? c?inoa? /a?ac 12 aiae. I?ney
ii?ii?iaiiy oaa?ei o aeic? 1 A? eiioeaio?aoe?y eao?iio o oeeoicie?
ee?oei c?inoa? /a?ac 3 aiae ? oo?eio?oueny ia i?aeaeuaiiio ??ai?
oi?iaeiaae 12 aiae.

Oaaeeoey 1

Eiioeaio?aoe?y a?eueiiai oeeoicieueiiai Na2+ ( iI) o e?ioioeeoao
naeac?iee i?ney ii?ii?iaiiy oaa?ei (n = 6-8)

Aeica ii?ii?iaiiy 3 aiae 6 aiae 12 aiae

0,5 A? 112 ( 18 110 ( 15 238 ( 31 *

1 A? 167 ( 21 * 245 ( 32 * 195 ( 25 *

* ? ( 0,05 ii??aiyii c eiio?ieai

E?eoa??yie ii?ooaiiy eaeueoe??aiai aiiainoaco o ee?oeiao ? ia eeoa
i?aeaeuaiiy eiioeaio?aoe?? Na2+ o oeeoiieaci?, aea e ci?ia iaia?aiii?
i?iieeiino? aeey eao?iio. Aeey ioe?iee eaeueoe??ai? i?iieeiino?
ieaciaoe/ii? iaia?aie e?ioioeeo?a aeine?aeaeoaaee ao?ae Na2+ o ee?oeie
(?en.3) oa ?ic?aoiaoaaee e?iaoe/i? ia?aiao?e i?ioeano – ii/aoeiao
oaeaee?noue iaaeoiaeaeaiiy (Vi) oa iaeneiaeueiee ao?ae (?max) eao?iia.
Aiae?c e?iaoe/ieo ia?aiao??a aeicaieea i?iaiae?coaaoe iiaeeea? i?e/eie
aeyaeaiiai iaie i?aeaeuaiiy eiioeaio?aoe?? eii?a a?eueiiai
oeeoicieueiiai eaeueoe?y o e?ioioeeoao ii?ii?iaieo oaa?ei.

*a?ac 3-6 aiae i?ney ii?ii?iaiiy oaa?ei o aeic? 0,5 A? e?iaoe/i?
ia?aiao?e aoiaeo Na2+ o e?ioioeeoe ia ci?iththoueny. Iineeaiiy aoiaeo
eao?iio aiane?aeie c?inoaiiy ii/aoeiai? oaeaeeino? eiai iaaeoiaeaeaiiy o
ee?oeie aeyaey?oueny /a?ac 12 aiae ( ?en. 3A). Ioaea, i?aeaeuaiiy ??aiy
[Ca2+]i o e?ioioeeoao i?ney ae?? ?aae?aoe?? o aeic? 0,5 A? ni?aiaaea? c
i?aeaeuaiiyi eaeueoe??ai? i?iieeiino? ieaciaoe/ii? iaia?aie ee?oei ?
coiiaeaia iineeaiiyi aoiaeo eao?iio ?c iicaee?oeiiiai na?aaeiaeua.

*a?ac 3 aiae i?ney ii?ii?iaiiy oaa?ei o aeic? 1 A? e?iaoe/i?
ia?aiao?e aoiaeo Na2+ o e?ioioeeoe oaeiae ia ci?iththoueny ( ?en. 3A).
Oea icia/a?, ui i?aeaeuaiiy ??aiy [Ca2+]i ia ?aiiueiio aoai?
?i?oe?aoe?? ?ioa?oacii? caaeaae? ee?oei a?aeaoaa?oueny aiane?aeie
aeoiaeo eao?iio c aioo??oiueiee?oeiieo aeaii. *a?ac 6 aiae a?aei?/aii
aeaie?aoia i?aeaeuaiiy ii/aoeiai? oaeaeeino? iaaeoiaeaeaiiy Na2+ o
e?ioioeeoe, a /a?ac 12 aiae ia? i?noea c?inoaiiy ia eeoa Vi, aea e
?ioaneaiino? iioieo eaeueoe?th /a?ac ieaciaoe/io iaia?aio, i?i ui
na?ae/eoue i?aeaeuaiiy aaee/eie ?max. Oaeei /eiii, ?aii? i?aeaeuaiiy
??aiy a?eueiiai Na2+ o oeeoicie? e?ioioeeo?a oa iiaeiaaeaia eiai
oo?eiaiiy i?ney ii?ii?iaiiy oaa?ei o aeic? 1 A? ? iieacieeii
iaaeeeoeiaiai iaeiie/aiiy Na2+ o e?ioioeeoao aiane?aeie
iacaaeainiaaiino? i?ioean?a aooa?oaaiiy eao?iia o aioo??oiueiee?oeiieo
aeaii oa eiai aoiaeo /a?ac ieaciaoe/io iaia?aio.

Aeyaeaia iaie i?aeaeuaiiy eaeueoe??ai? i?iieeiino? ieaciaoe/ii? iaia?aie
e?ioioeeo?a o oa?i?i 3-6 aiae i?ney ii?ii?iaiiy oaa?ei o aeic? 1 A?
aeicaieeei aeine?aeeoe aieea Na2+ ia i?ioean i?aeioeeainiiii?
aeaa?aaeaoe?? o?iiaoeio o ee?oeiao. Aeey oeueiai aea/aee aeeiai?eo
iaeiie/aiiy o?aaiaio?a AeIE (IAeI) o eiio?ieueieo e?ioioeeoao oa
e?ioioeeoao, aeae?eaieo /a?ac 3 aiae i?ney ae?? ?aae?aoe?? i?ioyaii
90-oaeeeiii? ?ieoaaoe?? ee?oei o na?aaeiaeu? aac aeiaeaaaiiy eaeueoe?th
oa o na?aaeiaeu?, ui i?noeei 2 iI NaCl2 (?en.4).

?ieoaaoe?y ee?oei, aeae?eaieo c eiio?ieueieo oaa?ei, noi?iaiaeaeo?oueny
iacia/iei, iacaeaaeiei a?ae iayaiino? eaeueoe?th o na?aaeiaeu?
?ieoaaoe??, c?inoaiiyi ai?noo IAeI. Ia ii/aoeiaiio aoai? ?ieoaaoe??
e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei o na?aaeiaeu? aac aeiaeaaaiiy
eaeueoe?th ia? i?noea i?aeaeuaiiy oaeaeeino? o?aaiaioaoe?? AeIE
ii??aiyii c eiio?ieai, iaeiae, ia a?eueo i?ci?o aoaiao ?ieoaaoe??
oaeaee?noue iaeiie/aiiy IAeI o ee?oeiao cieaeo?oueny. Aeiaeaaaiiy
eaeueoe?th o na?aaeiaeua ?ieoaaoe?? i?ecaiaeeoue aei cia/iiai c?inoaiiy
oaeaeeino? ?icuaieaiiy AeIE oa aei i?aeaeuaiiy ai?noo IAeI, iaeiie/aieo
i?ioyaii 90 oa. Ioaea, iaeiie/aiiy Na2+ o e?ioioeeoao naeac?iee
aiane?aeie i?aeaeuaiiy eaeueoe??ai? i?iieeiino? ieaciaoe/ii? iaia?aie
iiaea aooe iaei??th c i?e/ei iiina?aaeeiaaieo /a?ac eaeueoe?e
iacai?ioi?o ii?ooaiue no?oeoo?e oa oe?e?niino? o?iiaoeio ca ?icaeoeo
?ioa?oacii? caaeaae? e?ioioeeo?a naeac?iee i?ney ii?ii?iaiiy oaa?ei o
aeic? 1 A?.

I?iaaaeaiee iaie aiae?c ii?oieia?/ieo oa a?io?i?/ieo iieaciee?a noaio
ee?oei o iiioeyoe?? iaeeo oa na?aaei?o e?ioioeeo?a naeac?iee na?ae/eoue,
ui oi?ioaaiiy a?aeiia?ae? ee?oei in vivo ia ae?th ?ii?coth/i? ?aae?aoe??
o aeicao 0,5 oa 1 A? a?aeaoaa?oueny ??cieie oeyoaie. Iayai?noue
ii?oieia?/ieo iciae aeoeaaoe?? ee?oei, iacia/ia e?euee?noue caaeaeeo
ee?oei, i?aeaeuaiiy [Ca]i oa ai?noo IAeA o ee?oeiao o oa?i?i,
a?aeaeaeaiee a?ae ae?? i?iiaiaaiai oaeoi?a aeacothoue, ui ii?ii?iaiiy
oaa?ei o aeic? 0,5 A? ia aeeeeea? iaoaaie?/ieo ii?ooaiue, aeinoaoi?o
aeey aeoeaaoe?? o?eaa?iiai iaoai?cio ?icaeoeo o?aaeaiiy e?ioioeeo?a ? o
iiioeyoe?? e?ioioeeo?a naeac?iee ia?aaaaeathoue i?ioeane aaeaioeaii?
i?ie?oa?aoe??, o oiae? yei? aeiooue iaea?eueo ?aae?i/ooeea? ee?oeie.

Ii?ii?iaiiy oaa?ei o aeic? 1 A? noi?iaiaeaeo?oueny cia/iei cieaeaiiyi
e?eueeino? ee?oei o iiioeyoe?? iaeeo oa na?aaei?o e?ioioeeo?a naeac?iee
oa i?iyaeaiiyi ii?oieia?/ieo oa a?io?i?/ieo iciae ?ioa?oacii? caaeaae?
ee?oei. Oiia?eueiaia aeeiai?ea ?icaeoeo ?aae?aoe?eii? caaeaae?
e?ioioeeo?a, a oaeiae oie oaeo, ui c?inoaiiy eiioeaio?aoe?? a?eueiiai
oeeoicieueiiai Ca2+ o e?ioioeeoao ia?aaeo? ?icaeoeo i?aeioeeainiiii?
aeaa?aaeaoe?? o?iiaoeio aeicaieythoue i?eionoeoe aaaeeeao ?ieue
Na2+-caeaaeieo iaia?aiieo nenoai ee?oeiii? neaiae?caoe?? o iineeaii?
aeeeeeaiiai ae??th ?ii?coth/i? ?aae?aoe?? aiiioiaaiiiai neaiaeo.

Oaeei /eiii, iiioeyoe?y iaeeo oa na?aaei?o e?ioioeeo?a naeac?iee
ii?ii?iaieo uo??a iiaea aooe aeei?enoaia ye iiaeaeue aeey
aeine?aeaeaiiy ?icaeoeo in vivo ?iaeoeiaaii? ?aae?aoe??th ?ioa?oacii?
caaeaae? e?ioioeeo?a, a ii??aiyeueia aeine?aeaeaiiy ooieoe?iioaaiiy
Na2+-caeaaeieo ?aaoeyoi?ieo nenoai a?ae?ie?co iaia?aiieo oinoie?i?ae?a
o ee?oeiao i?ney ae?? in vivo ?ii?coth/i? ?aae?aoe?? o aeicao 0,5 oa 1
A? – ye aenia?eiaioaeueiee i?eeii aeey c’ynoaaiiy ioneiaeo iaoai?ci?a
?ioa?oacii? caaeaae? e?ioioeeo?a.

Aeine?aeaeaiiy aeoeaiino? iie?oinoi?iiceoeaeii? ianaiaeaea?ii? nenoaie o
e?ioioeeoao naeac?iee o ?aii?e ia??iae i?ney ii?ii?iaiiy oaa?ei.
Oinoie?iaca N, eeth/iaee oa?iaio iie?oinoi?iiceoeaeii? nenoaie,
cae?enith? a?ae?ie?c oinoiao??iiai ca’yceo on?o o?ueio oei?a
oinoi?iiceoeae?a c ooai?aiiyi aoi?eiieo ianaiaeaea??a — aiane?aeie
?icuaieaiiy oinoaoeaeee?iiceoieo oa oinoaoeaeee?iiceoie-4-oinoaoo o
ee?oeiao i?aeaeuo?oueny ai?no aeeaoeeeae?oea?iea, a aiane?aeie
?icuaieaiiy oinoaoeaeee?iiceoie-4,5-aeeoinoaoo – aeeaoeeeae?oea?iea oa
?iiceoie-1,4,5-o?eoinoaoa (Rhee S. et al, 1989; Cockroft S, 1992).
Aea/aiiy aeoeaiino? oinoie?iace N o e?ioioeeoao c aeei?enoaiiyi
aeciaaiiiai noano?aoo oinoaoeaeee?iiceoieo iieacaei, ui oa?iaioaoeaia
aeoeai?noue aeyaey?oueny ye o iaia?aii?e, oae ? oeeoicieuei?e o?aeoe?yo,
aeinyaa? iioeiaeueiiai ??aiy o iaeano? ?I 6,5 oa ca 80ieI
eiioeaio?aoe?? noano?aoo. Aeinoaoiuei aenieee ??aaiue oa?iaioaoeaii?
aeoeaiino? iaio o?aeoe?e i?iyaey?oueny o na?aaeiaeu? aac aeiaeaaaiiy
eaeueoe?th, iaeiae iaeneiaeueia oaeaee?noue oa?iaioaoeaiiai ?icuaieaiiy
oinoaoeaeee?iiceoieo, ye ? ?ioeo aeciaaiieo oinoie?i?ae?a ( Carter H. &
Smith A., 1987 ) ?icaeaa?oueny o i?enooiino? eaeueoe?th o
noai?e?iiey?i?e eiioeaio?aoe?? o na?aaeiaeu? ?ieoaaoe??

Aeey ioe?iee aeoeaiino? oinoie?iace N o iaaeeaeaieo aei in vivo oiiaao,
oiaoi ca o?c?ieia?/ii? eiioeaio?aoe?? noano?aoa o ee?oeii?e iaia?ai? oa
eii?a eaeueoe?th o oeeoicie? ee?oei, aoee aeei?enoai? e?ioioeeoe,
iiia?aaeiuei i?/ai? ca 3I-?iiceoieii. Ine?eueee oinoie?iaca N caaacia/o?
o?ainiaia?aiiee aoai i?iaaaeaiiy aeoeaaoe?eiiai neaiaeo o
aioo??oiueiee?oeiiee i?ino??, ianaiia?aae aeine?aeaeoaaee oa?iaioaoeaio
aeoeai?noue e?ioioeeo?a ca oiiae aeoeaaoe?? aioeaainiaoeeo?/ieo
?aoeaioi?ieo eiiieaen?a ieaciaoe/ii? iaia?aie. Aeey oeueiai
aeei?enoiaoaaee iie?eeiiaeuei? i?oiaaii? e?aaiaee o?oiaaiaaethoei?i
(OAA) oa e?iiiie?naoa?eae E. coli (EIN), ye? caeaoi? ia?ao?anii
coeaaoeny c iiaa?oiaaeie ?aoeaioi?aie ee?oei. Io?eiai? e?ea? caeaaeiino?
aoaeoo e?aaiaea a?ae /ano ?ieoaaoe?? c e?ioioeeoaie a?aeia?aaeathoue
aoaeoeai?noue ?o ca’ycoaaiiy c ee?oeiiith iaia?aiith, iaiao?aeiiai aeey
?iaeoeoe?? aeoeaiino? oinoie?iace N (?en.5). O?iiaoia?ao?/ia ?icae?eaiiy
noia?ii? o?aeoe?? ooai?aieo o oiae? ?ieoaaoe?? ee?oei ?iiceoieoinoao?a
iieacaei, ui ye EIN, oae ? OAA iineeththoue neioac on?o o?ueio oei?a
?iiceoieoinoao?a, iaeiae iaea?eueoith i??ith —
?iiceoie-1,4,5-o?eoinoaoo (?O3). Ioaea, o aeine?aeaeoaai?e iaie
iiaeaeuei?e nenoai? ooieoe?iio? iaoai?ci ia?aaea/? neaiaeo a?ae
?aoeaioi?a ciai?oiuei? iaia?aie ana?aaeeio ee?oeie ca o/ano?
iie?oinoi?iiceoeaeii? nenoaie. Aieea OAA oa EIN ia aeoeai?noue
oinoie?iace N ii??aithaaee c ae??th eaeueoe??aiai ?iiioi?a A23187,
oaeiae caeaoiiai i?iyaeyoe i?oiaaiiee aoaeo. Aeiaeaaaiiy ?iiioi?a o
na?aaeiaeua ?ieoaaoe?? ee?oei aeeeeeaei iacia/ia i?aeaeuaiiy
aeeth/aiiy i?oee o ?iiceoieoinoao (?O) oa ?iiceoie-1,4-aeeoinoao (?O2)
? ia aieeaaei ia ooai?aiiy ?O3 ( ?en. 5). Oea ocaiaeaeo?oueny c
e?oa?aoo?ieie aeaieie, ca?aeii yeeo neioac ?iiceoieoinoao?a
a?aeaoaa?oueny ca o?c?ieia?/ii? eiioeaio?aoe?? ?ii?a eaeueoe?th o
oeeoicie? (Nakanishi H. et al., 1989; Conti A.et al., 1993). Ioaea,
aca?iiae?y oinoie?iace N c iaia?aiieie oinoi?iiceoeaeaie aecia/a?oueny,
ia?o ca ana, noaiii ?aoeaioi?iiai eiiieaeno o neeaae? ieaciaoe/ii?
iaia?aie e?ioioeeo?a.

Aeine?aeaeaiiy aeoeaiino? oinoie?iace N o e?ioioeeoao naeac?iee
i?iaiaeeee /a?ac 1, 3, 6 oa 12 aiae i?ney ii?ii?iaiiy oaa?ei,
aeei?enoiaoth/e aac’yaea?iee aiiiaaiao ee?oei oa i?/aiee noano?ao
oinoaoeaeee?iiceoie. Ca?aeii i?aaenoaaeaieo ia ?en. 6 aeaieo, o ?aii?e
ia??iae i?ney ae?? ?ii?coth/i? ?aae?aoe?? a?aeaoaa?oueny i?aeaeuaiiy
oa?iaioaoeaii? aeoeaiino? e?ioioeeo?a naeac?iee, ??aaiue yeiai
caeaaeeoue a?ae aeice ?aae?aoe??. I?ney ii?ii?iaiiy oaa?ei o aeic? 0,5
A? oa?iaioaoeaia aeoeai?noue ee?oei c?inoa? /a?ac 3 aiae. Ii?ii?iaiiy
oaa?ei o aeic? 1 A? i?ecaiaeeoue aei a?eueo cia/ii? iiaeeo?eaoe??
aeoeaiino? oinoie?iace N — oa?iaioaoeaia aeoeai?noue i?aeaeuo?oueny
/a?ac 1 aiae, aeinyaa? iaeneiaeueiiai ??aiy /a?ac 3 aiae ?
i?eai?/o?oueny ii??aiyii c eiio?ieai /a?ac 12 aiae.

Aiae?c oa?iaioaoeaii? aeoeaiino? noaee?oeiieo o?aeoe?e e?ioioeeo?a
iieacaa, ui /a?ac 3 aiae i?ney ii?ii?iaiiy oaa?ei o aeic? 0,5 A?
oinoie?iacia aeoeai?noue iaia?aiii? o?aeoe?? c?inoa? ia 33,5 % ii??aiyii
c eiio?ieai, a oeeoicieueii? – ia ci?ith?oueny. I?ney ii?ii?iaiiy
oaa?ei o aeic? 1 A? oa?iaioaoeaia aeoeai?noue iaia?aiii? oa
oeeoicieueii? o?aeoe?e i?aeaeuo?oueny ia 120 oa 73 % ii??aiyii c
eiio?ieai a?aeiia?aeii. Io?eiai? aeai? aeicaieythoue i?eionoeoe, ui
iaei??th c i?oaiae ae?? ?ii?coth/i? ?aae?aoe?? o aeic? 1 A? ? eiiiiiaioe
iie?oinoi?iiceoeaeii? nenoaie, ca’ycai? c ieaciaoe/iith iaia?aiith
e?ioioeeo?a naeac?iee.

Uia aeine?aeeoe aieea ?ii?coth/i? ?aae?aoe?? ia ooieoe?iiaeueiee noai
?aoeaioi??a ieaciaoe/ii? iaia?aie e?ioioeeo?a, ye? ni?yaeai? c
oinoie?iacith N oa aeoeaothoueny i?ae ae??th e?aaiae?a, caeaoieo
aeeeeeaoe ?o aa?aaaoe?th, aea/aee aieea e?iiiie?naoa?eaea ia
aeoeai?noue oinoie?iace N o e?ioioeeoao o eiio?ie? oa /a?ac 3 aiae i?ney
ii?ii?iaiiy oaa?ei. I?a?ieoaaoe?y i?ioyaii 5 oa o i?enooiino?
e?iiiie?naoa?eaeo e?ioioeeo?a naeac?iee eiio?ieueieo oaa?ei aeeeeea?
i?aeaeuaiiy aeoeaiino? oinoie?iace N o aac’yaea?iiio aiiiaaiao? ee?oei (
oaae. 2), ui ocaiaeaeo?oueny c aeaieie, io?eiaieie iaie i?e
aeine?aeaeaii? aieeao EIN ia ooai?aiiy ?iiceoieoinoao?a o e?ioioeeoao.
Aeoeaoth/ee aoaeo i?iyaey?oueny oaeiae ca i?a?ieoaaoe?? c i?oiaaiii
e?ioioeeo?a naeac?iee oaa?ei, ii?ii?iaieo o aeic? 0,5 A?. Iaeiae,
aeiaeaaaiiy EIN o na?aaeiaeua i?a?ieoaaoe?? e?ioioeeo?a naeac?iee
oaa?ei, ii?ii?iaieo o aeic? 1 A?, ia aieeaa? ia aeoeai?noue oinoie?iacio
aeoeai?noue ee?oei.

Oaaeeoey 2

Aieea e?iiiie?naoa?eaeo ia aeoeai?noue oinoie?iace N e?ioioeeo?a
naeac?iee o eiio?ie? oa /a?ac 3 aiae i?ney ii?ii?iaiiy oaa?ei (n = 4)

Oiiae aenia?eiaioa Aeoeai?noue,

iiieue ?iiceoieoinoaoo ( ia a?eea (oa)-1

Oiiae i?a?ieoaaoe??

aac aeiaaaee + EIN

eiio?ieue 6.1 ( 0,6 19,4 ( 1,2**

ii?ii?iaiiy 0,5 A? 8,6 ( 0,8 12,3 ( 1,1**

ii?ii?iaiiy 1 A? 10,2 ( 0,9* 9,8 ( 0,7

* ?( 0,05 ii??aiyii c eiio?ieai

** ?( 0,05 ii??aiyii c i?a?ieoaaoe??th aac aeiaaaee

I/aaeaeii, ii?ii?iaiiy oaa?ei o aeic? 1 A? i?ecaiaeeoue aei no?oeoo?ieo
aai ooieoe?iiaeueieo ci?i o noai? ieaciaoe/ii? iaia?aie e?ioioeeo?a
naeac?iee, ye? iiaeeo?eothoue aca?iiae?th oinoie?iace N c
iaia?aiica’ycaieie a?eeaie-?aaoeyoi?aie oa aeeeeeathoue aeoeaaoe?th
oa?iaioa, yea ia eiio?ieth?oueny /a?ac noeioeyoe?th
aioeaainiaoeeo?/ieo ?aoeaioi??a.

Io?eiai? aeai? na?ae/aoue i?i aeoeaaoe?th o e?ioioeeoao ii?ii?iaieo
oaa?ei eeth/iaiai eiiiiiaioa iie?oinoi?iiceoeaeii? nenoaie – oinoie?iace
N, iaeiae ia aeathoue oyaeaiiy i?i i?e?iaeo ooai?thaaieo i?iaeoeo?a
a?ae?ie?co iie?oinoi?iiceoeae?a. Neioac ?iiceoieoinoao?a aeine?aeaeoaaee
ca ?ieoaaoe?? e?ioioeeo?a naeac?iee, aeae?eaieo /a?ac 1, 3 oa 6 aiae
i?ney ii?ii?iaiiy oaa?ei oa iiia?aaeiuei i?/aieo ca 3I-?iiceoieii.
Ai?no i?oee o noia?i?e o?aeoe?? ?iiceoieoinoao?a i?ney 5-oaeeeiii?
?ieoaaoe?? eiio?ieueieo ee?oei neeaaeaa 490 ( 45 ?ii/oa ia 107 ee?oei.
a?aei?/aia I?aeaeuaiiy ii??aiyii c eiio?ieai aeeth/aiiy i?oee o noia?io
o?aeoe?th ?iiceoieoinoao?a o e?ioioeeoao aeyaeaia /a?ac 1 aiae i?ney
ii?ii?iaiiy oaa?ei o aeic? 0,5 A?, oa /a?ac 1 ? 3 aiae i?ney
ii?ii?iaiiy oaa?ei o aeic? 1 A?. Oe? aeai? i?aeoaa?aeaeothoue oaeo
?aiiuei? aeoeaaoe?? oinoie?iace N o e?ioioeeoao ii?ii?iaieo oaa?ei.

O?iiaoia?ao?/ia ?icae?eaiiy noia?ii? o?aeoe?? ?iiceoieoinoao?a
aeicaieeei i?iaanoe ii??aiyeueiee aiae?c ooai?aiiy ??cieo oei?a
?iiceoieoinoao?a o e?ioioeeoao eiio?ieueieo oa ii?ii?iaieo oaa?ei. O
e?ioioeeoao eiio?ieueieo oaa?ei aoa aeyaeaiee oaeee ?iciiae?e i?oee: ?O
– 240 ( 35, ?O2 –140 ( 23 , ?O3 – 120 ( 25 ?ii /oa ia 107 ee?oei.
Iaie ciaeaeaii, ui iineeaiiy neioaco ?iiceoieoinoao?a o e?ioioeeoao
/a?ac 1 aiae i?ney ii?ii?iaiiy oaa?ei o aeic? 0,5 A? coiiaeaia
ooai?aiiyi ?iiceoie-1-oinoaoo, ai?no yeiai, ia a?aei?io a?ae ?ioeo
?iiceoieoinoao?a, i?aeaeuo?oueny ii??aiyii c eiio?ieai. Ia iniia? oeeo
aeaieo c?iaeaiee aeniiaie, ui iniiaiei ianaiaeaea?ii, yeee ooai?th?oueny
iie?oinoi?iiceoeaeiith nenoaiith e?ioioeeo?a naeac?iee o ?aii?e oa?i?i
i?ney ae?? ?aae?aoe?? o aeic? 0,5 A? ? aeeaoeeeae?oea?ie. I/aaeaeii, o
oeueiio aeiaaeeo a?aeaoaa?oueny iai?aaeaiiy neaiaeo o a?e
iiina?aaeeiaaii? /a?ac aca?iiae?th c aeeaoeeeae?oea?ieii aeoeaaoe??
Na2+, oinoie?i?ae- caeaaeii? i?ioa?ie?iace oa eaneaaeo i?ioa?ie?iac, ui
oinoi?eeththoue ??ci? aoaeoi?i? a?eee. Oaea iinoo?aineyoe?eia
iiaeeo?eaoe?y a?ee?a eaaeeoue a iniia? oi?ioaaiiy aaeaioeaii?
a?aeiia?ae? ee?oei /a?ac noeioeyoe?th i?ie?oa?aoe?? (Meyn R. et al.,
1993, Prasad A.et al., 1994).

Niaeo? ?iiceoieoinoao?a, ooai?thaaieo o e?ioioeeoao naeac?iee oaa?ei,
ii?ii?iaieo o aeic? 1 A?, ?ioee ( ?en.7). *a?ac 1 aiae i?ney ae??
?aae?aoe?? i?aeaeuo?oueny ai?no i?oee o neeaae? neioaciaaiiai o
ee?oeiao ?iiceoie-1-oinoaoo, a /a?ac 3 aiae – o neeaae? ?iiceoie
–1,4-aeeoinoaoo. Iaeiae iaea?eueoith i??ith ii??aiyii c ?ioeie
?iiceoieoinoaoaie o ia??iae 1-3 aiae i?ney ae?? ?aae?aoe?? iineeth?oueny
ooai?aiiy oaeiai aoi?eiiiai ianaiaeaea?a, ye ?iiceoie-1,4,5-o?eoinoao.
Ioaea, ?ioaineo?eaoe?y oa?iaioaoeaiiai a?ae?ie?co
iie?oinoi?iiceoeae?a o e?ioioeeoao naeac?iee oaa?ei, ii?ii?iaieo o aeic?
1 A?, noi?iaiaeaeo?oueny aeoeaaoe??th ie?aiiai neaiaeueiiai eaioethaa,
ui eiio?ieth? ??aaiue oeeoiieaciaoe/iiai Na2+ o ee?oeiao. Ine?eueee
?icuaieaiiy oinoaoeaeee?iiceoie-4,5-aeeoinoaoo o neeaae? ee?oeiieo
iaia?ai oa ooai?aiiy ?iiceoie-1,4,5-o?eoinoaoo – oea aieiaiee iaoai?ci
iia?e?caoe?? eaeueoe?th c aiaeiieaciaoe/iiai ?aoeeoeoia e?ioioeeo?a
(Corado I., 1990;Conti A.et al., 1993), ? i?aenoaae aaaaeaoe, ui
aeyaeaia iaie ?aii? i?aeaeuaiiy eiioeaio?aoe?? Na2+ o oeeoicie?
e?ioioeeo?a naeac?iee oaa?ei, ii?ii?iaieo o aeic? 1 A?, coiiaeaia
aea?eueiaiiyi Na2+ naia c oeueiai aioo??oiueiee?oeiiiai eiiia?oiaioo.

A?aoiao/e, ui ?icaeoie ?ioa?oacii? caaeaae? e?ioioeeo?a naeac?iee i?ney
ii?ii?iaiiy oaa?ei a?aeaoaa?oueny ca oiiae iaeii/aniiai i?aeaeuaiiy
eiioeaio?aoe?? oeeoicieueiiai Na2+ o ee?oeiao oa iineeaiiy i?ioean?a
ia?aeeniiai ieeneaiiy e?i?ae?a, aeine?aeaeoaaee ?ieue o oi?ioaaii?
iino?aae?aoe?eieo ?aaeoe?e e?ioioeeo?a oa?iaio?a, aeoeai?noue yeeo
cia/iith i??ith aecia/a?oueny ooai?aiiyi i?iaeoeo?a ?aaeoe?e
a?eueii?aaeeeaeueiiai ia?aieeneaiiy oa aioo??oiueiee?oeiiith
eiioeaio?aoe??th eaeueoe?y — oinoie?iace A2, ui a?ae?ie?co?
a?ao?aeiii?eai?ni? oinoie?i?aee, oa e?iieneaaiace.

Aeine?aeaeaiiy aeoeaiino? oinoie?iace A2 oa Na2+-caeaaeiiai iaoaaie?cio
a?ao?aeiiiai? eeneioe o e?ioioeeoao naeac?iee i?ney ii?ii?iaiiy
oaa?ei.

Aeine?aeaeaiiy aea?eueiaiiy a?ao?aeiiiai? eeneioe c? neeaaeo iaia?aiieo
oinoie?i?ae?a e?ioioeeo?a naeac?iee i?iaiaeeee ia ee?oeiao, ui aeeth/eee
14N-a?ao?aeiiao i?ioyaii 90-oaeeeiii? ?ieoaaoe?? c i?oeith. Ine?eueee
iia? i?noey aeey aeeth/aiiy aee?ii? eeneioe aei neeaaeo oinoie?i?ae?a
c’yaeythoueny eeoa i?ney ae?? oinoie?iace A2, ioe?iea ?iciiae?eo i?oee
i?ae iniiaieie o?aeoe?yie e?i?ae?a o i?/aieo ca 14N-a?ao?aeiiaoii
e?ioioeeoao aeicaiey? ?aeaioeo?eoaaoe aiaeiaaii? noano?aoe oinoie?iace
A2. Ca?aeii ?acoeueoao?a o?iiaoia?ao?/iiai ?icae?eaiiy e?i?aeii?
o?aeoe?? i?oea ?iciiae?ey?oueny oaeei /eiii ( % a?ae caaaeueii?
e?eueeino? aeeth/aii? i?oee) – aoeeeae?oea?eie — 11,6;
e?cioinoaoeaeeeoie?i — 5,7; oinoaoeaeeena?ei + oinoaoeaeee?iiceoie —
9,5; oinoaoeaeeeoie?i – 21,8; oinoaoeaeeaoaiieai?i – 47,3;
oinoaoeaeeeae?oea?ie – 4,1. Ioaea, o e?ioioeeoao ?niothoue aaoa?iaaii?
ioee a?ao?aeiiaoo, aeinooii?noue yeeo aeey oinoie?iace A2 iiaea aooe
??ciith. Oe? iniaeeaino? noai?ththoue oiiae aeey eiio?ieueiaaiiai
aea?eueiaiiy a?ao?aeiiiai? eeneioe i?ae aieeaii ??ciiiai?oieo
?aaoeyoi?ieo oaeoi??a oa ?? ooieoe?iioaaiiy o yeino? eieaeueiiai
aoi?eiiiai iina?aaeieea.

E?euee?noue i?oee, ui aea?eueiy?oueny i?ioyaii ?ieoaaoe?? e?ioioeeo?a
naeac?iee eiio?ieueieo oaa?ei o na?aaeiaeua, yea i?noeei
aeae?i?aeeciaaiee ANA aeey ca’ycoaaiiy aee?ii? eeneioe, ? iacia/iith (
?en. 8A), ui na?ae/eoue i?i iecueeo ?ioaineai?noue iai?io a?ao?aeiiiai?
eeneioe o ianoeioeueiaaieo e?ioioeeoao. Aeiaeaaaiiy o na?aaeiaeua
?ieoaaoe?? e?ioioeeo?a 0,5 ieI eaeueoe??aiai ?iiioi?o A23187 cia/ii
noeioeth? aea?eueiaiiy a?ao?aeiiiai? eeneioe. Aeiaeaaaiiy
e?iiiie?naoa?eaeo oaeiae iineeth? aea?eueiaiiy a?ao?aeiiiai? eeneioe o
na?aaeiaeua ?ieoaaoe?? ee?oei, oi/a aoaeo ? a?eueo oiia?eueiaiei ? iaio
aeyaeaiei ii??aiyii c A23187. Oe? aeai? aeacothoue, ui aaaeeeaei
oaeoi?ii, yeee eiio?ieth? i?ioean aea?eueiaiiy a?ao?aeiiiai? eeneioe c
iaia?aiieo oinoie?i?ae?a o e?ioioeeoao naeac?iee, ? eiioeaio?aoe?y
aioo??oiueiee?oeiiiai Na2+, a iia?e?caoe?y a?ao?aeiiiai? eeneioe
aiane?aeie aeoeaaoe?? ?aoeaioi?a i?oiaaiiei e?aaiaeii
iiina?aaeeiao?oueny /a?ac aeiiii?aei? ?aaoeyoi?i? iaoai?cie, na?aae yeeo
iniiaiee – oinoaoeaeee?iiceoieiaee oeeee (Errasfa et al., 1989).

Ii??aiyeueiee aiae?c e?iaoe/ieo ia?aiao??a aeeth/aiiy 14N-a?ao?aeiiaoo
o e?ioioeeoe naeac?iee eiio?ieueieo oa ii?ii?iaieo oaa?ei iieacaa, ui
i?ney ae?? ?ii?coth/i? ?aae?aoe?? iai?i a?ao?aeiiiai? eeneioe
i?enei?th?oueny. I?ney ii?ii?iaiiy oaa?ei o aeic? 0,5 A? ci?ie
aeyaeythoueny /a?ac 12 aiae, eiee ia? i?noea c?inoaiiy ii/aoeiai?
oaeaeeino? V0 aeeth/aiiy i?oee o e?ioioeeoe. *a?ac 6 aiae i?ney
ii?ii?iaiiy oaa?ei o aeic? 1 A? aeyaey?oueny i?aeaeuaiiy cia/aiiy ia
eeoa V0, aea e O05 — /ano iai?aiaeneiaeueiiai aeeth/aiiy
14N-a?ao?aeiiaoo o e?ioioeeoe.

Ca?aeii ?acoeueoao?a, i?aaenoaaeaieo ia ?en.8A,A ?ioaineo?eaoe?y iai?io
a?ao?aeiiiai? eeneioe o e?ioioeeoao naeac?iee i?ney ae?? ?aae?aoe??
iia’ycaia c iineeaiiyi ?? aea?eueiaiiy c? neeaaeo iaia?aiieo
oinoie?i?ae?a. Oae, ca ?ieoaaoe?? e?ioioeeo?a, aeae?eaieo /a?ac 12 aiae
i?ney ii?ii?iaiiy oaa?ei o aeic? 0,5 A?, e?euee?noue 14C-a?ao?aeiiiai?
eeneioe o na?aaeiaeu? ?ieoaaoe?? i?aeaeuo?oueny ii??aiyii c eiio?ieai.
Ca ?ieoaaoe?? e?ioioeeo?a naeac?iee oaa?ei, ii?ii?iaieo o aeic? 1 A?,
cia/ii c?inoa? ia eeoa e?euee?noue a?ao?aeiiiai? eeneioe o na?aaeiaeu?
?ieoaaoe??, aea e oaeaee?noue ?? aea?eueiaiiy. Caa?oa? ia naaa oaaao
iiae?ai?noue o oa?aeoa?? aea?eueiaiiy a?ao?aeiiaoo c iaia?aiieo
oinoie?i?ae?a o e?ioioeeoao naeac?iee oaa?ei i?ney ii?ii?iaiiy o aeic? 1
A? oa ca oiiae i?a?ieoaaoe?? eiio?ieueieo e?ioioeeo?a o i?enooiino?
eaeueoe??aiai ?iiioi?a A23187. Oaea iiae?ai?noue na?ae/eoue, ui
i?aeaeuaiiy ??aiy aioo??oiueiee?oeiiiai Na2+aieeaa? ia aea?eueiaiiy
a?ao?aeiiiai? eeneioe c iaia?aiieo oinoie?i?ae?a ee?oei iacaeaaeii a?ae
oiai, aeeeeeaia aiii eaeueoe??aei ?iiioi?ii /e i?aeaeuaiiyi eaeueoe??ai?
i?iieeiino? ieaciaoe/ii? iaia?aie e?ioioeeo?a aiane?aeie ae??
?aae?aoe??. Eaeueoe?e, oaeei /eiii, ? iaeiei c o?eaa?ieo oaeoi??a
aeoeaaoe?? aea?eueiaiiy a?ao?aeiiiai? eeneioe o e?ioioeeoao naeac?iee,
ui caciathoue ?ioa?oacii? caaeaae?.

Ine?eueee i?aeaeuaiiy ai?noo a?eueii? a?ao?aeiiiai? eeneioe o ee?oeiao
a?aeaoaa?oueny aiane?aeie aeoeaaoe?? aiaeiaaiiiai oinoie?i?aeiiai
a?ae?ie?co, aea/aee ?ieue o oeueiio i?ioean? oinoie?iace A2.
Aeine?aeaeaiiy aeoeaiino? oinoie?iace A2 o e?ioioeeoao naeac?iee
i?iaiaeeee, aeei?enoiaoth/e aeciaaii? i?/ai? noano?aoe
14N-a?ao?aeiii?eoinoaoeaeeeaoaiieai?i oa
14N-a?ao?aeiii?eoinoaoeaeeeoie?i. Aeoeai?noue oinoie?iace A2 o
e?ioioeeoao naeac?iee aoea aeuith ca iayaiino? caeeoeo a?ao?aeiiiai?
eeneioe ii??aiyii c iea?iiaith o sn-2 iieiaeaii? ia?aoai?thaaiiai
oinoie?i?aea, aeinyaaea iaeneiaeueiiai ??aiy o iaeano? eoaeieo cia/aiue
?I, i?eai?/oaaeanue o i?enooiino? 2 iI AAOA, i?iyaeyeanue ca
o?c?ieia?/ii? eiioeaio?aoe?? eaeueoe?y o na?aaeiaeu? ?ieoaaoe?? oa
c?inoaea ca ?? i?aeaeuaiiy o ae?aiacii? noai?e?iiey?ii? eiioeaio?aoe??
eao?iio, ui a oe?eiio a?aeiia?aea? aeanoeainoyi oeeoicieueii?
oinoie?iace A2 oeio IV (Lesllie C.et al., 1990; Dennis E., 1994).
?iciiae?e oinoie?iacii? aeoeaiino? i?ae iaia?aiiith oa oeeoicieueiith
o?aeoe?yie e?ioioeeo?a, caeaaei?noue oa?iaioaoeaiiai a?ae?ie?co a?ae
eiioeaio?aoe?? eii?a eaeueoe?th oa eiai iaeneiaeueia oaeaee?noue aoee
??cieie aeey aeaio noano?ao?a. Oae, o iaia?aii?e o?aeoe?? e?ioioeeo?a
naeac?iee aeyaey?oueny 24,4% oa 41,9 % a?ae caaaeueii? oa?iaioaoeaii?
aeoeaiino? ca aeei?enoaiiy noano?ao?a oinoaoeaeeeoie?io oa
oinoaoeaeeeaoaiieai?io a?aeiia?aeii, cia/aiiy ENa2+ — eiinoaioe
iai?aiaeneiaeueii? aeoeaaoe?? eaeueoe??i ?aaeoe?? oa?iaioaoeaiiai
a?ae?ie?co aeaio noano?ao?a noaiiaeee 0,13 ( 0,01 oa 0,55 ( 0,07 iI
a?aeiia?aeii, iaeneiaeueia oaeaee?noue oinoie?i?aeiiai a?ae?ie?co aoea
aeuith ca aeei?enoaiiy noano?aoo oinoaoeaeeeaoaiiai?io.

Iaie aeyaeaii i?aeaeuaiiy oinoie?iacii? aeoeaiino? iaia?aiii? o?aeoe??
e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei, yea a?aeaoaa?oueny o ??ciee
oa?i?i i?ney ae?? ?aae?aoe?? caeaaeii a?ae aeice ii?ii?iaiiy oa oeio
noano?aoo ( oaae. 3). Oae, /a?ac 12 aiae i?ney ii?ii?iaiiy oaa?ei o
aeic? 0,5 A? o iaia?aii?e o?aeoe?? ee?oei i?aeaeuo?oueny a?ae?ie?oe/ia
aeoeai?noue oinoie?iace A2 noiniaii oinoaoeaeeeoie?io, a /a?ac 6 aiae
i?ney ii?ii?iaiiy oaa?ei o aeic? 1 A? iineeth?oueny oa?iaioaoeaia
?icuaieaiiy ye oinoaoeaeeeoie?io, oae ? oinoaoeaeeeaoaiieai?io.

Oaaeeoey 3

Aeoeai?noue oinoie?iace A2 (iiie? a?ao?aeiiaoo/ oa ( ia a?eea) o
iaia?aii?e o?aeoe?? e?ioioeeo?a naeac?iee i?ney ii?ii?iaiiy oaa?ei( n =
4-6)

oiiae aenia?eiaioo noano?ao

14N-2-a?ao?aeiii?e-

oinoaoeaeeeoie?i 14N-2-a?ao?aeiii?e

oinoaoeaeeeaoaiieai?i

eiio?ieue 0,40 ( 0,03 1,50 ( 0,16

0,5 A? , 6 aiae 0,53 ( 0,06 2,02 ( 0,25

12 aiae 0,70 ( 0,08* 2,04 ( 0,29

1 A? 6 aiae 0,58 ( 0,05* 2,51 ( 0,22*

12 aiae 0,42 ( 0,03 1,91 ( 0,30*

*? ( 0,05

I?aeaeuaiiy oinoie?iacii? aeoeaiino? iaia?aii?e o?aeoe?? e?ioioeeo?a
naeac?iee oaa?ei, ii?ii?iaieo o aeic? 1 A?, ni?aiaaea? o /an? c
aeyaeaiith iaie ?ioaineo?eaoe??th i?ioean?a ia?aeeniiai ieeneaiiy
iaia?aiieo e?i?ae?a o ee?oeiao. No?oeoo?iee noai iaia?ai ? iaeiei c
oaeoi??a, ui aecia/a? ?ioaineai?noue oinoie?i?aeiiai a?ae?ie?co o
ee?oeiao- aiane?aeie iineeaiiy ?aaeoe?e a?eueii?aaeeeaeueiiai ieeneaiiy
aei ??aiy, yeee ? e?eoe/iei aeey ?i?oe?aoe?? ii?ooaiiy no?oeoo?e
iaia?aie, a?aeaoaa?oueny eieaeueia i?aeaeuaiiy aeoeaiino? oinoie?iace A2
(Eaieei A., 1984; Burak W.et al., 1995). Io?eiai? iaie aeai? na?ae/aoue
i?i oa, ui aea?eueiaiiy a?ao?aeiiiai? eeneioe c iaia?aiieo oinoie?i?ae?a
e?ioioeeo?a naeac?iee ca ?icaeoeo ?ioa?oacii? caaeaae? a?aeaoaa?oueny
aiane?aeie aeoeaaoe?? oinoie?iace A2. Noano?aoii aeoeaiaaiiai oa?iaioo
iiaeooue aooe ??ci? oinoie?i?aee, eieae?ciaai? o iaaieo ae?eyieao
iaia?aie, aeinooii?noue yeeo aeey oa?iaioo caeaaeeoue a?ae no?oeoo?iiai
noaio iaia?aie.

?aaoeyoe?y eaoae?oe/ii? aeoeaiino? oinoie?iace A2 aaeeeith i??ith
a?aeaoaa?oueny /a?ac aca?iiae?th oeueiai oa?iaioa c ?iiaie eaeueoe?th —
i?aeaeuaiiy eiioeaio?aoe?? Na2+ o noai?e?iiiey?iiio ae?aiacii?
i?ecaiaeeoue aei ca’ycoaaiiy eao?iia oinoie?iacith A2 oa aei ia?aoiaeo
oa?iaioa o oi?io c a?eueo aenieith nii??aeiai?noth aei noano?aoa, a ca
i?aeaeuaiiy Na2+ o ae?aiacii? 10 –100ieI a?aeaoaa?oueny ca’ycoaaiiy
oa?iaioa c iaia?aiith oa eiai aeoeaaoe?y ( Lin L.et al., 1993;
Nalefski E. et al., 1994). Ca?aeii ?acoeueoao?a, i?aaenoaaeaieo ia ?en.
9, i?a?ieoaaoe?y e?ioioeeo?a naeac?iee eiio?ieueieo oaa?ei c
eaeueoe??aei ?iiioi?ii A23187 aeeeeea? i?aeaeuaiiy aeoeaiino?
oinoie?iace A2 o o?aeoe?? aac’yaea?iiai aiiiaaiaoo noiniaii
14N-2-a?ao?aeiii?eoinoaoeaeeeaoaiieai?io. *a?ac 6 aiae i?ney ii?ii?iaiiy
oaa?ei o aeic? 0,5 A? oinoie?iacia aeoeai?noue e?ioioeeo?a
i?aeaeuo?oueny ia 21% ? c?inoa? ca i?a?ieoaaoe?? ee?oei c ?iiioi?ii
oae naii, ye ? o e?ioioeeoao eiio?ieueieo oaa?ei. Oinoie?iacia
aeoeai?noue e?ioioeeo?a naeac?iee oaa?ei, ii?ii?iaieo o aeic? 1 A?,
c?inoa? ia 75% ii??aiyii c eiio?ieai, ia?iaea ee?oei eaeueoe??aei
?iiioi?ii ia i?ecaiaeeoue aei i?aeaeuaiiy aeoeaiino? oinoie?iace A2. Oe?
aeai? aeacothoue, ui i?ney ii?ii?iaiiy oaa?ei o aeic? 1 A?
eiioeaio?aoe?y Na2+ o oeeoicie? e?ioioeeo?a naeac?iee i?aeaeuo?oueny
aei ??aiy, yeee ? aeinoaoi?i aeey aeoeaaoe?? oa?iaioaoeaiiai a?ae?ie?co
a?ao?aeiii?eai?nieo oinoie?i?ae?a oa aea?eueiaiiy cia/ii? e?eueeino?
a?ao?aeiiiai? eeneioe.

Ooee?caoe?y iaaeeeoeo aea?eueiaii? c iaia?aiieo oinoie?i?ae?a
a?ao?aeiiiai? eeneioe caaacia/o?oueny /a?ac Na2+- caeaaeio ?iaeoeoe?th
e?iieneaaiacii? aeoeaiino? o ee?oeiao (Eaie?i A., 1984; Keppler
D.,1992). O i?iaaaeaieo iaie aenia?eiaioao c aeei?enoaiiyi noano?aoo
e?iieaai? eeneioe oa noia?iaoaioo 18000g, io?eiaiiai i?ney
oeaio?eooaoaaiiy aac’yaea?iiai aiiiaaiaoo ee?oei, ye oa?iaioiiai
i?aia?aoo, iieacaii, ui o e?ioioeeoao naeac?iee a?aeaoaa?oueny
oa?iaioaoeaia ia?aieeneaiiy aeciaaiii? iie?iaiane/aii? eeneioe c
ooai?aiiyi a?ae?iia?aeenieo ?aaeeeae?a. E?iieneaaiacia aeoeai?noue
aeinyaaea iioeiaeueiiai ??aiy ca ?I 6,4 oa 0,1ieI eiioeaio?aoe??
noano?aoo, i?iyaeyeanue ca o?c?ieia?/ii? eiioeaio?aoe?? ?ii?a Na2+,
c?inoaea ca i?aeaeuaiiy eiioeaio?aoe?? NaCl2 aei 0,1 iI oa
i?eai?/oaaeanue o i?enooiino? 0,2 iI AAOA o na?aaeiaeu? ?ieoaaoe??.
?aae?i?ioiiei iaoiaeii o e?ioioeeoao naeac?iee aoee ?aeaioeo?eiaai?
aiaeiaaii? i?iaeoeoe e?iieneaaiaciiai ia?aoai?aiiy a?ao?aeiiiai? eeneioe
– 15- a?ae?ieneaeeicaoao?a?iiaa eeneioa (15-IAOA) oa eaeeio???i A4.

I?ney ?aioaai?anueeiai ii?ii?iaiiy oaa?ei e?iieneaaiacia aeoeai?noue
e?ioioeeo?a naeac?iee ci?ith?oueny. I?ney ae?? ?aae?aoe?? o aeic? 0,5
A? i?aeaeuaiiy oa?iaioaoeaii? aeoeaiino? aeyaeaia /a?ac 12 aiae (?en.
10). Ii?ii?iaiiy oaa?ei o aeic? 1 A? aeeeeea? a?eueo ?aii? ?
o?eaaea i?aeaeuaiiy e?iieneaaiacii? aeoeaiino? e?ioioeeo?a – aiii
aeyaey?oueny /a?ac 3 aiae ? oo?eio?oueny oi?iaeiaae 6 aiae, iaeiae
/a?ac 24 aiae oa?iaioaoeaia aeoeai?noue i?eai?/o?oueny ii??aiyii c
eiio?ieai ia 25%. Aeyaeaiee iaie oa?aeoa? ci?ie e?iieneaaiacii?
aeoeaiino? e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei ocaiaeaeo?oueny c
iniaeeainoyie ?aaoeyoe?? oa?iaioo – yeui i?iaeoeoe iaoa?iaioaoeaiiai
ia?aeeniiai ieeneaiiy e?i?ae?a o iecueeeo eiioeaio?aoe?yo aeoeaothoue
e?iieneaaiaco, oi ooai?aiiy cia/ii? e?eueeino? i?iaeoeo?a
oa?iaioaoeaiiai IIE i?ecaiaeeoue aei i?eai?/aiiyi aeoeaiino? oa?iaioo (
Lands W. et al., 1985).

Iineeaiiy e?iieneaaiacii? aeoeaiino? e?ioioeeo?a naeac?iee ii?ii?iaieo
oaa?ei noi?iaiaeaeo?oueny iaeii/aniei i?aeaeuaiiyi ai?noo aiaeiaaiieo
i?iaeoeo?a e?iieneaaiaciiai ia?aoai?aiiy a?ao?aeiiiai? eeneioe o
ee?oeiao ( oaae.4). Oae, /a?ac 12 aiae i?ney ii?ii?iaiiy oaa?ei o aeic?
0,5 A? o e?ioioeeoao naeac?iee aeyaey?oueny i?aeaeuaiiy ai?noo
ia?aeiiiai i?iaeoeoo e?iieneaaiacii? ?aaeoe?? 15-IAOA. O e?ioioeeoao
naeac?iee oaa?ei, ii?ii?iaieo o aeic? 1 A?, e?euee?noue neioaciaaii?
15-IAOA cia/ii c?inoa? /a?ac 3 – 6 aiae. Oaea naia caeaaei?noue a?ae
aeice ?aae?aoe?? aeyaeaia ? i?e aeine?aeaeaii? ai?noo eaeeio???io A4 o
e?ioioeeoao naeac?iee i?ney ii?ii?iaiiy..

Oaaeeoey 4

Ai?no 15-IAOA oa EO A4 (ia/107 ee?oei) o e?ioioeeoao naeac?iee

i?ney ii?ii?iaiiy oaa?ei

Oiiae aenia?eiaioa Oa?i?i IAOA, EO A4, ia/107 ee?oei

eiio?ieue

57.8 ( 3.7 19,3 ( 2,2

ii?ii?iaiiy, 0,5 A? 3 aiae 63,4 ( 7.4 —

6 aiae 70.2 ( 9.4 28,6 ( 4,9

12 aiae 221.8 ( 25.5* 43,1 ( 5,3*

ii?ii?iaiiy, 1 A? 3 aiae 122.6 ( 14.2* 42,3 ( 5,1*

6 aiae 185.8 ( 18.6* 78,1 ( 8,2*

12 aiae 49.1 ( 5.8 39,2 ( 6,2*

* ? ( 0,05 ii??aiyii c eiio?ieai

Aeyaeaia iaie i?aeaeuaiiy e?iieneaaiacii? aeoeaiino? e?ioioeeo?a
naeac?iee i?ney ae?? ?aae?aoe?? iiaeia iiynieoe ye ci?io
aaeaioaoe?eiiai oa?aeoa?o, iai?aaeaio ia cieaeaiiy o ii?ii?iaiee ee?oei?
??aiy aeoeaieo oi?i eenith aiane?aeie ?o ia?aianaiiy e?iieneaaiacith ia
iie?iaiane/ai? aee?i? eeneioe. Iaeiae ooai?aiiy o iaaei??ieo e?eueeinoyo
oaeeo i?iaeoeo?a ?aaeeeaeueii? i?e?iaee, ye ia?aeii? iaoaaie?oe
e?iieneaaiaciiai ia?aoai?aiiy a?ao?aeiiiai? eeneioe, iiaea i?ecaiaeeoe
aei ii?ooaiiy ieenii-aioeieneaeaioii? ??aiiaaae o ee?oeiao oa aei ci?ie
aeni?an?? aai?a (Aoaeieoeeay A., 1986; Tebbey P.et al.,1993).

Aeey c’ynoaaiiy iiaeeeai? ?ie? ?ioaineo?eaoe?? e?iieneaaiaciiai
ieeneaiiy a?ao?aeiiiai? eeneioe o ?icaeoeo ?iaeoeiaaii? ?aae?aoe??th
?ioa?oacii? caaeaae? e?ioioeeo?a aea/aee aeeiai?eo iaeiie/aiiy
o?aaiaio?a AeIE (IAeI) o eiio?ieueieo e?ioioeeoao oa e?ioioeeoao,
aeae?eaieo /a?ac 3 aiae i?ney ae?? ?aae?aoe?? i?ioyaii 60-oaeeeiii?
?ieoaaoe?? ee?oei o na?aaeiaeu? aac aeiaaaie oa o i?enooiino?
ii?aeea?ae?iaoay?aoeeiai? eeneioe — ?ia?a?oi?a ieeniiai iaoaaie?cio
iie?iaiane/aieo aee?ieo eeneio, yeee naeaeoeaii i?eai?/o? aeoeai?noue
e?iieneaaiace (Sircar J. et al, 1983; Eoee A.,1988). Aeiaeaaaiiy 20 ieI
IAeAE o na?aaeiaeua ?ieoaaoe?? i?eaiaeeoue aei cia/iiai i?eai?/aiiy
?icuaieaiiy AeIE o e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei, iniaeeai
ia ii/aoeiaeo aoaiao ?ieoaaoe?? ee?oei (?en.10). Io?eiai? aeai?
aeicaieythoue ae?eoe aeniiaeo, ui e?iieneaaiaci? iaoaaie?oe
a?ao?aeiiiai? eeneioe, ye? iineeaii ooai?ththoueny o e?ioioeeoao ?aii?e
ia??iae i?ney ii?ii?iaiiy oaa?ei o aeic? 1 A?, aeeiiothoue ?ieue
aiaeiaaiieo iaae?aoi??a ?ioa?oacii? caaeaae? e?ioioeeo?a.

Caeeth/ia /anoeia.

sse iiaeaeue ?icaeoeo in vivo ?ioa?oacii? caaeaae? e?ioioeeo?a iaie
aeei?enoaia iiioeyoe?y iaeeo oa na?aaei?o e?ioioeeo?a naeac?iee uo??a,
e?euee?noue ee?oei o ye?e cia/ii cieaeo?oueny o ?aii?e ia??iae
?aae?aoe?eiiai o?aaeaiiy i?ney ?aioaai?anueeiai ii?ii?iaiiy oaa?ei o
aeic? 1 A?. Ia iniia? ii??aiyeueiiai aeine?aeaeaiiy aeeiai?ee ?icaeoeo
ii?oieia?/ieo oa a?io?i?/ieo iciae ?ioa?oacii? caaeaae? ee?oei o
iiaeaeuei?e nenoai? oa ia?aiao??a ooieoe?iioaaiiy Na2+-caeaaeieo
oinoie?i?aeieo o?ainaeoeoe?eieo nenoai ?ic?iaeaia noaia oi?ioaaiiy
?iaeoeiaaiiai ?aae?aoe??th aiiioiaaiiiai neaiaeo o e?ioioeeoao
naeac?iee ( ?en.11).

Ca?aeii io?eiaieo iaie ?acoeueoao?a, ia?oi/a?aiao ?ieue o ?icaeoeo
?ioa?oacii? caaeaae? e?ioioeeo?a naeac?iee i?ney ?aioaai?anueeiai
ii?ii?iaiiy oaa?ei a?ae?a?athoue iaia?aii? eiiiiiaioe neaiaeueieo
nenoai. ?i?oe?aoe?y ?icaeoeo ii?oieia?/ieo iciae ?ioa?oacii? caaeaae?
e?ioioeeo?a o aeine?aeaeoaai?e iiioeyoe?? e?ioioeeo?a naeac?iee
a?aeaoaa?oueny iaeii/anii c ?ioaineo?eaoe??th i?ioean?a ia?aeeniiai
ieeneaiiy, i?aeaeuaiiyi aeoeaiino? oinoie?iace N, iineeaiiyi neioaco
?iiceoie-1,4,5-o?eoinoaoo o ee?oeiao oa c?inoaiiyi eiioeaio?aoe??
a?eueiiai oeeoicieueiiai Na2+. Ao?aoa /ooeeaino? oinoie?iace N aei ae??
iie?eeiiaeueiiai i?oiaaia EIN na?ae/eoue, ui aeoeaaoe?y oa?iaioo oa
i?iaaaeaiiy ?iaeoeiaaiiai ?aae?aoe??th neaiaeo o e?ioioeeoao
a?aeaoaathoueny /a?ac iaoai?cie, ye? ia iio?aaothoue ca’ycoaaiiy
e?aaiaea c? ni?yaeaiei n oinoie?iacith N iiaa?oiaaei ?aoeaioi?ii.
Iaea?eueo a??ia?aeieie c oaeeo iaoai?ci?a ? aa?aaaoe?y ?aoeaioi??a
ieaciaoe/ii? iaia?aie aiane?aeie coeaie, ooai?aieo i?e ?aeiia?iaoe??
?aaeeeae?a a?ee?a ( Koteles G., 1982, Sosiynsky V., 1991) oa aeoeaaoe?y
/ooeeaeo aei eeniaai?nieo ?aaeeeae?a oe?iceiiaeo i?ioa?ie?iac fyn oa
lck, ye? caaacia/othoue ca’ycoaaiiy oinoie?iace N c iaia?aiith oa
aeoeaaoe?th oa?iaioa (Chalupny N. et al.; 1991Rao A., 1994).

Aeyaeaia iaie i?aeaeuaiiy aeoeaiino? oinoie?iace N o iaia?aii?e o?aeoe??
e?ioioeeo?a naeac?iee oa iineeaiiy ooai?aiiy ?O3 o e?ioioeeoao
naeac?iee ii?ii?iaieo oaa?ei a?aeaoaa?oueny o ?aii?e i?ney?aae?aoe?eiee
ia??iae, eiee i?iieei?noue ieaciaoe/ii? iaia?aie aeey ?ii?a eaeueoe?th
ia ci?ith?oueny. Oea na?ae/eoue, ui i?e/eiith ?aiiueiai i?aeaeuaiiy
eiioeaio?aoe?? oeeoicieueiiai Na2+ o e?ioioeeoao, ui aeiooue o
?ioa?oac?, ? iia?e?caoe?y eao?iio c aiaeiieaciaoe/iiai ?aoeeoeoio /a?ac
?aoeaioi?e-eaiaeaie, nii??aeiai?noue yeeo aei ?O3 c?inoa? i?e
iacia/iiio i?aeaeuaii? eiioeaio?aoe?? ianaiaeaea?a o iaiaiiey?iiio
ae?aiacii? (Berridge M., 1984; Chow S.& Jondal M., 1990; Scharff O. et
al, 1993). Aea?eueiaiiy eao?iio c aiaeiieaciaoe/iiai ?aoeeoeoia iiaea
aooe ioneiaei iaoai?ciii ii?ooaiiy aioo??oiueiee?oeiii?
eiiia?oiaioae?caoe?? eao?iio a ii?ii?iaieo ee?oeiao oa iiaeiaaeaiiai
oo?eiaiiy eiioeaio?aoe?? a?eueiiai oeeoicieueiiai Na2+ ia i?aeaeuaiiio
??ai?, ine?eueee i?aeaeuaiiy eaeueoe??ai? i?iieeiino? ieaciaoe/ii?
iaia?aie e?ioioeeo?a o a?eueo a?aeaeaeaiee oa?i?i i?ney ae?? ?aae?aoe??
… iacaaeainiaaiee ao?ae eao?iio o aioo??oiueiee?oeiiee i?ino??.

Iane?aeeii ii?ooaiiy eaeueoe??aiai aiiainoaco oa iaeiie/aiiy Na2+ o
oeeoiieaci? ee?oei iiaea aooe iio?aieyiiy eao?iio o yae?i oa iineeaiiy
Na2+-caeaaeii? aeni?an?? aai?a n?iaenoaa gadd – iiceoeaieo
?aaoeyoi??a aiiioico (Bartlett L. et al., 1992; Smeyne R. et al.,
1993). Iaaei??iiai iiiiaiaiiy yaea?iiai eaeueoe??aiai ioeo iiaea aooe
iaei??th c i?e/ei ci?ie eiioi?iaoe?? o?iiaoeio oa aeoeaaoe?? Na2+ –
caeaaeiiai aiaeiioeeaie?co AeIE.

Iaei??th c ?aaeoe?e ee?oei ia ae?th aenoa?iaeueieo oaeoi??a oa
ieneaeaoeaiiai no?ano ? ?iaeoeoe?y neioaco oinoie?iace A2 oa
e?iieneaaiace (Schewe T. & Kunh H.,1991; Chen X. et al., 1996). O?eaaea
oo?eiaiiy i?aeaeuaii? eiioeaio?aoe?? Na2+ o oeeoiieaci? e?ioioeeo?a
naeac?iee ii?ii?iaieo oaa?ei i?aeaeuo? a??ia?aei?noue iane/aiiy eao?iiii
iaio Na2+-ca’ycoth/eo oa?iaio?a, iaoai?ciii aeoeaaoe?? yeeo ? Na2+
-caeaaeia o?aineieaoe?y aei iaia?aie (Keppler D., 1992; Lin L. et al.,
1993). Aeoeaaoe?y oa?iaio?a o oaeee niin?a a?aeaoaa?oueny i?ney iaaiiai
eaa-ia??iaeo ? caaacia/o? iineeaiiy ?iaeoeiaaiiai ?aae?aoe??th
eaeueoe??aiai neaiaeo oa ?icaeoie o e?ioioeeoao naeac?iee ii?ii?iaieo
oaa?ei a?eueo iia?eueieo ci?i iaoaaie?cio, ui ?aaoeththoueny
ooai?thaaieie aoi?eiieie ianaiaeaea?aie. Oae, a?ao?aeiiiaa eeneioa, /an
?nioaaiiy yei? o ee?oeiao aeiaoee ii??aiyii c ?O3, ? ?aaoeyoi?ii
eaeueoe??ai? i?iieeiino? ieaciaoe/ii? iaia?aie e?ioioeeo?a, caeaoia
iia?e?coaaoe Na2+ c ooieoe?iiaeueieo ??cieo aioo??oiueiee?oeiieo
eaeueoe??aeo aeaii ( Aoeianeay A. e ae?., 1989; Liskovitch M., 1992;
Tsunoda et al., 1993) oa ? aeoeaaoi?ii no?iaii??e?iace – oa?iaioa,
a?aeiia?aeaeueiiai ca neioac oea?ai?aea – o?eaa?a aiiioico, aeoeaaoi?a
i?ioa?iac n?iaenoaa ?NA ( Cifone M. et al., 1995). Aiane?aeie ?icaeoeo
eaioethaiaeo ?aaeoe?e ia?aieeneaiiy e?i?ae?a oa iieaaoaiiy aeinooia
ca’ycaii? c iaia?aiaie aeoeaiaaii? oinoie?iace A2 aei noano?aoo o
e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei a?aeaoaa?oueny aea?eueiaiiy
a?ao?aeiiiai? eeneioe c? neeaaeo iaia?aiieo oinoie?i?ae?a, ui aoee
iaaeinooiieie aeey oa?iaioa o eiio?ie?. Na?aae iaoaaie?o?a, ui
ooai?ththoueny i?e ooee?caoe?? e?iieneaaiacith iaaeeeoeo a?ao?aeiiiai?
eeneioe o e?ioioeeoao naeac?iee, 15-I?AOA ? iaea?eueo aoaeoeaiei
?iaeoeoi?ii aeni?an?? c-fos – iaeiiai c aai?a ?aiiuei? a?aeiia?ae?,
?aaoeyoi?a ee?oeiiiai oeeeeo. Iineeaiiy aeni?an?? oeueiai aaio
iacaeaaeii a?ae noaae?? ee?oeiiiai oeeeeo ? iaei??th c oiia ?icaeoeo
?ioa?oacii? caaeaae? ee?oei ( Manome Y. et al.,1993; Wilson R. et al.,
1993). Ooai?thaai? eaeeio???ie iiaeooue ca’ycoaaoenue ?aoeaioi?aie ia
iiaa?oi? ee?oei oa aeeiioaaoe ?ieue aooi- oa ia?ae?eiieo iaae?aoi??a
aiiioico, ui ocaiaeaeo?oueny c oiia?eueiaiith aeeiai?eith ?ioa?oacii?
caaeaae? ee?oei o aeine?aeaeoaai?e iiioeyoe?? iaeeo oa na?aaei?o
e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei.

Oaeei /eiii, ?icaeoie in vivo ?aae?aoe?eii? caaeaae? e?ioioeeo?a o
?ioa?oac? a?aeaoaa?oueny aiane?aeie ?iaeoeiaaii? ?aae?aoe??th
ieaeio?iiieo ci?i o ooieoe?iioaaii? iaia?aiieo neaiaeueieo nenoai oa
iiaeeo?eaoe?? i?inoi?iai-/aniai? i?aai?caoe?? ?o aca?iiae?? —
iaeii/anii? aeoeaaoe?? aeae?eueeio iaia?aiieo Na2+-caeaaeieo
oinoie?i?aeieo o?ainaeoeoe?eieo oeyo?a, ii?ooaiiy iine?aeiaiino?
ooai?aiiy oa o?eaaeino? neaiae?a, ye? ?aaoeththoue i?io?eaiiy
iaoaaie?/ieo i?ioean?a ye ia iaia?aiiiio, oae ? aaiiiiiio ??aiyo.

Aeniiaee

1. Anoaiiaeaii, ui i?ney oioaeueiiai ?aioaai?anueeiai ii?ii?iaiiy uo??a
o aeic? 1 A? o iiioeyoe?? iaeeo oa na?aaei?o e?ioioeeo?a naeac?iee
a?aeaoaa?oueny cieaeaiiy e?eueeino? ee?oei aiane?aeie ?o caaeaae? o
?ioa?oac?. ?ioa?oaciee oa?aeoa? caaeaae? e?ioioeeo?a aeiaaaeaiee ia
iniia? ye ii?oieia?/ieo (eiiaeainaoe?y o?iiaoeio, i?eiioecaoe?y yae?a),
oae ? a?io?i?/ieo ( iaeiie/aiiy o ee?oeiao o?aaiaio?a AeIE aeiaaeeiith
400, 600 oa 800 ia? iniia, ii?ooaiiy aioo??oiueiee?oeiiiai eaeueoe??aiai
aiiainoaco) iciae. Aeyaeaia oiia?eueiaia aeeiai?ea ?icaeoeo in vivo
?iaeoeiaaii? ?aae?aoe??th ?ioa?oacii? caaeaae? e?ioioeeo?a naeac?iee,
yea na?ae/eoue i?i o?ainoi?iaoe?th o iiioeyoe?? ee?oei ?aae?aoe?eiiai
neaiaeo ca o/ano? iaia?aiieo neaiaeueieo nenoai.

2. Iieacaii, ui o aeine?aeaeoaai?e iiioeyoe?? e?ioioeeo?a naeac?iee
ooieoe?iiothoue iaoai?cie i?iaaaeaiiy aeoeaaoe?eiiai neaiaeo a?ae
ieaciaoe/ii? iaia?aie o aioo??oiueiee?oeiiee i?ino?? ca o/ano?
neaiaeueieo oinoie?iac N oa A2 – coeaaiiy iiaa?oiaaeo ?aoeaioi??a
e?ioioeeo?a iie?eeiiaeueieie i?oiaaiaie EIN oa OAA i?ecaiaeeoue aei
iineeaiiy a?ae?ie?co iie?oinoi?iiceoeae?a oa ooai?aiiy ?iiceoieoinoao?a,
a ia?iaea ee?oei eaeueoe??aei ?iiioi?ii aeeeeea? iineeaiiy aea?eueiaiiy
a?ao?aeiiiai? eeneioe c? neeaaeo a?ao?aeiii?eai?nieo oinoie?i?ae?a.

3. Anoaiiaeaii, ui iaei??th c ?aii?o ?aaeoe?e ca ?i?oe?aoe?? ?ioa?oacii?
caaeaae? e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei ? aeoeaaoe?y
oinoi?iiceoeae-niaoeeo?/ii? oinoie?iace N o iaia?aii?e o?aeoe?? ee?oei
oa iiaeeo?eaoe?y aca?iiae?? eiiiiiaio?a o nenoai? iiaa?oiaaee ?aoeaioi?
e?ioioeeoa – iaia?aiica’ycai? a?eee –oinoie?iaca N, aiane?aeie yei?
i?aeaeuaiiy aeoeaiino? oinoie?iace N a?aeaoaa?oueny /a?ac iaoai?cie, ui
ia iio?aaothoue ca’ycoaaiiy e?aaiaea c? ni?yaeaiei c oinoie?iacith N
?aoeaioi?ii.

4. Iieacaii, ui ia ?aiiueiio aoai? ?icaeoeo ?ioa?oacii? caaeaae?
e?ioioeeo?a naeac?iee ii?ii?iaieo oaa?ei a?aeaoaa?oueny iineeaiiy
neioaco ?iiceoieoinoao?a oa ci?ia ?o niaeo?o o ee?oeiao. Iaoiaeii
o?iiaoia?ao?/iiai ?icae?eaiiy ?iiceoieoinoao?a aeiaaaeaii, ui
iaea?eueoith i??ith ii??aiyii c ?O oa ?O2 o e?ioioeeoao naeac?iee i?ney
ii?ii?iaiiy oaa?ei o aeic? 1 A? iineeth?oueny ooai?aiiy
Na2+-iia?e?coth/iai ianaiaeaea?a — ?iiceoie-1,4,5-o?eoinoaoo.

3. I?ney ii?ii?iaiiy oaa?ei o aeic? 1 A? o e?ioioeeoao naeac?iee
a?aeaoaa?oueny i?aeaeuaiiy eiioeaio?aoe?y a?eueiiai oeeoicieueiiai Na2+,
aei??yii? ca aeiiiiiaith eaeueoe??aiai oeoi?anoeaioiiai ciiaeo ea?i-2.
Anoaiiaeaii, ui i?aeaeuaiiy ??aiy Na2+ o oeeoicie? e?ioioeeo?a o ?aii?e
oa?i?i i?ney ae?? ?aae?aoe?? ia?aaeo? iaeiie/aiith i?iaeoeo?a
i?aeioeeainiiii? o?aaiaioaoe?? AeIE ? a?aeaoaa?oueny ca a?aenooiino?
ci?i e?iaoe/ieo ia?aiao??a aoiaeo eaeueoe?y o ee?oeie. Oe? aeai?
aeacothoue, ui ioneiaei iaoai?ciii ii?ooaiiy eaeueoe??aiai aiiainoaco o
e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei ? iiina?aaeeiaaia /a?ac
?iiceoie-1,4,5-o?eoinoao aea?eueiaiiy eao?iio c aiaeiieaciaoe/iiai
?aoeeoeoio.

4. Iieacaii, ui o?eaaea oo?eiaiiy eiioeaio?aoe?? oeeoicieueiiai
eaeueoe?th ia i?aeaeuaiiio ??ai? o e?ioioeeoao naeac?iee ia a?eueo
i?ci?o aoaiao ?icaeoeo ?aae?aoe?eii? caaeaae? ee?oei coiiaeaia
c?inoaiiyi ii/aoeiai? oaeaeeino? iaaeoiaeaeaiiy 45Na2+ o ee?oeie oa
ca?eueoaiiyi ?ioaineaiino? o?ainiaia?aiiiai iioieo eao?iia. I?aeaeuaiiy
eaeueoe??ai? i?iieeiino? ieaciaoe/ii? iaia?aie e?ioioeeo?a naeac?iee
ni?aiaaea? o /an? c iineeaiiyi i?ioean?a ia?aeeniiai ieeneaiiy e?i?ae?a,
cieaeaiiyi aioeieneaeaioii? aeoeaiino? oa i?enei?aiiyi i?aeioeeainiiii?
aeaa?aaeaoe?? AeIE.

5. Anoaiiaeaii, ui iniiaieie aiaeiaaiieie noano?aoaie oinoie?iace A2 o
i?/aieo ca 14N-a?ao?aeiiaoii e?ioioeeoao naeac?iee ?
oinoaoeaeeeaoaiieai?i oa oinoaoeaeeeoie?i. O i?ioean? ?icaeoeo
?ioa?oacii? caaeaae? e?ioioeeo?a c?inoa? oaeaee?noue aea?eueiaiiy
a?ao?aeiiiai? eeneioe c? neeaaeo iaia?aiieo oinoie?i?ae?a e?ioioeeo?a oa
e?euee?noue aea?eueiaii? o na?aaeiaeua ?ieoaaoe?? a?ao?aeiiiai? eeneioe.
Iaeii/anii o iaia?aii?e o?aeoe?? ee?oei i?aeaeuo?oueny aeoeai?noue
oinoie?iace A2, yea a?ae?ie?co? a?ao?aeiii?e-OA oa a?ao?aeiii?e-OO. Oe?
aeai? aeacothoue, ui iineeaiiy aea?eueiaiiy a?ao?aeiiiai? eeneioe o
e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei a?aeaoaa?oueny aiane?aeie
aeoeaaoe?? oinoie?iace A2, ui cae?enith? a?ae?ie?c ??cieo ioe?a
a?ao?aeiii?eai?nieo oinoie?i?ae?a.

8. E?ioioeeoe naeac?iee i?iyaeythoue e?iieneaaiacio aeoeai?noue,
ia?aoai?thth/e aeciaaiio e?iieaao eeneioo c ooai?aiiyi
a?ae?ieneia?ieneaeieo iio?aeieo. ?aeaioeo?eiaai? ia?aeii? aiaeiaaii?
i?iaeoeoe e?iieneaaiaciiai ia?aoai?aiiy a?ao?aeiiiai? eeneioe o ee?oeiao
– 15-a?ae?ieneaeeicaoao?a?iiaa eeneioa oa eaeeio???i A4. Anoaiiaeaii,
ui ca ?icaeoeo ?ioa?oacii? caaeaae? e?ioioeeo?a naeac?iee ii?ii?iaieo
oaa?ei e?iieneaaiacia aeoeai?noue i?aeaeuo?oueny, a ai?no 15-IAOA oa EO
A4 c?inoa?.

9. Anoaiiaeaii, ui niaoeeo?/iee ?ia?a?oi? e?iieneaaiace
ii?aeea?ae?iaoaea?aoeeiaa eeneioa i?eai?/o? i?aeioeeainiiio
aeaa?aaeaoe?th AeIE o e?ioioeeoao naeac?iee ii?ii?iaieo oaa?ei. Oe?
aeai? aeacothoue ia ?ieue ia?aeiieo i?iaeoeo?a e?iieneaaiaciiai
ia?aieeneaiiy a?ao?aeiiaoo ye iaae?aoi??a ?icaeoeo ?ioa?oacii? caaeaae?
e?ioioeeo?a.

10. Cai?iiiiiaaia noaia i?iaaaeaiiy ?iaeoeiaaiiai ?aae?aoe??th neaiaeo
oa eiai o?ainoi?iaoe?? ia iaia?aiiiio oa aaiiiiiio ??aiyo o aiiioiaaiiee
neaiae, yea a?aoiao? ?ieue eaeueoe?y o ii?ooaii? i?inoi?iai-/aniai?
aca?iiae?? ??cieo eiiiiiaio?a Na2+-caeaaeieo oinoie?i?aeieo
o?ainaeoeoe?eieo nenoai o e?ioioeeoao i?ney ii?ii?iaiiy oaa?ei.

Nienie iioae?eiaaieo ca oaiith aeena?oaoe?? ?ia?o

1.?o/ei I.A., Neaoaeineay A.A., Iaouoaaneay I.I., Eo/a?aiei I.A.
Oaaee/aiea eiioeaio?aoeee naiaiaeiiai Na2+, ii?aaeaeaiiie n iiiiuueth
eaeia –2 a eeioioeeoao naeacaiee iaeo/aiiuo e?un// Oe?. aeioei. aeo?i. –
1990. – o.62, N 4. – N. 110 –113.

2. Eo/a?aiei I.A., Iaouoaaneay I.I., Inoai/aiei E.E., Ia?oiiaoe O.E.,
?o/ei I.A., Neaoaeineay A.A. Niaea?aeaiea oeeeee/aneeo ioeeaioeaeia,
naiaiaeiiai oeeoiieaciaoe/aneiai Na2+ e iaeiiiaiai aeeaeueaeaaeaea a
eeioioeeoao naeacaiee e oeiioeeoao i?e aeaenoaee iaaunieeo aeic
?aaeeaoeee// ?aaeeiaeieiaey. – 1991. – O.31, N 5. – N. 739 – 742.

3. Neaoaeineay A.A., Iaouoaaneay I.I., Eo/a?aiei I.A. Oinoieeiaca A2 a
eeioioeeoao naeacaiee e?un, aeae?ieecothuay a?aoeaeiiieeoinoieeieaeu//
Oe?. aeioei. aeo?i. – 1991. – O. 63, N 6. – N. 48-52.

4. Iico? A.E., Iaouoaaneay I.I., Inoai/aiei E.E., Eo/a?aiei I.A.
Eiaeoeoeey aaeoa?eaeueiuie iaioeaeiaeeeaiii e eeiiiieenaoa?eaeii ?aiieo
e i?iiaaeooi/iuo i?eciaeia aeoeaaoeee eeioieaeiuo eeaoie
yenia?eiaioaeueiuo aeeaioiuo// Eiioiieiaey. – 1993. – N1. – N. 25-27

5. Ianooo A.I., Iaoeoaanueea I.I., Eo/a?aiei I.?. Aea/aiiy aeoeaiino?
5-e?iieneaaiace a e?ioioeeoao naeac?iee oa oeion? uo??a// A?niee
Ee?anueeiai oi?aa?neoaoo. A?ieia?y. – 1993. – N25. – N.14 – 17.

6. Ianooo A.I., Iaoeoaanueea I.I., Eo/a?aiei I.?. Aea/aiiy oiia
i?io?eaiiy 5-e?iieneaaiacii? ?aaeoe?? a e?ioioeeoao naeac?iee uo??a//
Aeiiia?ae? AI OE?a?ie. – 1993. – N6. – N.131- 134.

7. Iaouoaaneay I.I., Neaoaeineay A.A., Eo/a?aiei I.A. Oinoieeiaca A2 a
aeioeie/aneeo ?aaeoeeyo eeioioeeoia naeacaiee ia ?aioaaiianeia
iaeo/aiea// Aii?inu iaaeeoeeineie oeiee. – 1994. – O.40, N 2. – N. 28 –
31.

8. Iaouoaaneay I.I., Neaoaeineay A.A., Eo/a?aiei I.A. Aunaiaiaeaeaiea
a?aoeaeiiaoa ec eeioioeeoa naeacaiee e?un iinea iaeo/aiey // Oe?.
aeioei. aeo?i. – 1994. – O. 66, N 3. – N. 60- 66.

9. Neaoa?inueea I.A., Iaoeoaanueea I.I., Eo/a?aiei I.?. Aieea
?aioaai?anueeiai ii?ii?iaiiy ia aeoeai?noue oinoe?iace A2 o
noaee?oeiieo o?aeoe?yo e?ioioeeo?a naeac?iee oa oeiono uo??a// A?niee
Ee?anueeiai oi?aa?neoaoo. A?ieia?y. – 1995. – N26. – N.99 – 104.

10. Nieiaeooei A.I., Iaoeoaanueea I.I., Eo/a?aiei I.?. Aiaeiioeeaie?c
o?iiaoeio o e?ioioeeoao naeac?iee oa oeiono uo??a i?ney ii?ii?iaiiy //
Oe?. a?io?i. aeo?i. – 1997.- O.69, N 4. – N. 88 – 94.

11. Nieiaeooei A.I., Iaoeoaanueea I.I., Eo/a?aiei I.?. Na, Mg-caeaaeia
aiaeiioeeaacia aeoeai?noue oeiioeeo?a uo??a i?ney ii?ii?iaiiy //
Aeiiia?ae? IAI Oe?a?ie. – 1996. – N10. – N. 147 – 150

12. Iaoeoaanueea I.I. Aiiioic – cai?ia?aiiaaia oi?ia ee?oeiii? caaeaae?
// A?niee Ee?anueeiai oi?aa?neoaoo. A?ieia?y. – 1998. — N 27. – N.3 – 7.

13. Iaouoaaneay I.I. ?ieue eiiceoieo?eoinoaoa a oi?ie?iaaiee
eaeueoeeaaiai neaiaea a O-eeioioeeoao // Oe?. aeioei. aeo?i. – 1998. –
O.70, N4. – N.3 – 15.

14. Ne?iaaoeue O.A., Ia?oiiaoeue O.?., Iaoeoaanueea I.I., Eo/a?aiei I.?.
Aieea ?aioaai?anueeiai ii?ii?iaiiy ia aeoeai?noue oinoie?iace N, ui
a?ae?ie?co? oinoaoeaeee?iiceoie o e?ioioeeoao naeac?iee uo??a // A?niee
Ee?anueeiai oi?aa?neoaoo. A?ieia?y. – 1998. — N 28. – N.12 – 14

15. Iaoeoaanueea I.I. Ooai?aiiy ?iiceoieo?eoinoaoo a e?ioioeeoao
naeac?iee uo??a ia ?aiiueiio aoai? i?ney ?aioaai?anueeiai ii?ii?iaiiy //
Oe?. aeioei. aeo?i.- 1998. – O. 70, N 4. – N. 94 – 100.

16. Iaoeoaanueea I.I. Iine?aeiai?noue aeeth/aiiy neaiaeueieo iaoai?ci?a
/a?ac oinoie?iaco N/ ?iiceoieo?eoinoao oa oinoie?iaco A2 / a?ao?aeiiao
o e?ioioeeoao naeac?iee uo??a// A?niee Ee?anueeiai oi?aa?neoaoo.
A?ieia?y. – 1998. — N 28. – N.14-16.

17. Iaouoaaneay I.I., Ianooo A.I., Nieiaeooei A.A. Eeiieneaaiaciay
aeoeaiinoue e niaea?aeaiea 15-IAOA a eeioioeeoao naaeacaiee e?un i?e
aeaenoaee eiiece?othuae ?aaeeaoeee // Oe?. aeioei. aeo?i.- 1998. – O.
70, N 5. – N. 91 –96

18. Nieiaeooei A.I., A?eithe ?.?., Ai?enia N.?., Iaoeoaanueea I.I.
Caeaaei?noue a?ae eaeueoe?th ?iaeoeiaaiiai ?aae?aoe??th aiaeiioeeaie?co
o?iiaoeio o e?ioi?aeieo ee?oeiao // A?niee Ee?anueeiai oi?aa?neoaoo.
A?ieia?y. – 1998. — N 28. – N.3 – 7.

19. Iaouoaaneay I.I. Aeioeie/aneea aniaeou aucaaiiiai ?aaeeaoeeae
aiiioica // Oe?. aeioei. aeo?i.- 1998. – O. 70, N 5. – N. 15 – 29.

20. Iaouoaaneay I.I., Ianooo A.I., Nieiaeooei A.A. Eiaeae?iaaiea
eeiieneaaiaciie aeoeaiinoe nieaeaao eiaeooee?iaaiioth ?aaeeaoeeae
o?aaiaioaoeeth AeIE eeioioeeoia // ?aaeeaoeeiiiay aeieiaey.
?aaeeiyeieiaey. – 1999. – O.39, N 2-3. – N. 282 – 286.

Iaoeoaanueea I.I. Caeiiii??iino? ?icaeoeo ?ioa?oacii? caaeaae?
e?ioioeeo?a naeac?iee uo??a ca o/ano? Na2+-caeaaeieo oinoie?i?aeieo
neaiaeueieo nenoai i?ney ?aae?aoe?eiiai o?aaeaiiy. – ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy aeieoi?a a?ieia?/ieo iaoe ca
niaoe?aeuei?noth 03.00.04 – a?io?i?y. – Ee?anueeee oi?aa?neoao ?iai?
Oa?ana Oaa/aiea, Ee?a, 1999.

Aeena?oaoe?th i?enay/aii aeine?aeaeaiith o/ano? iie?oinoi?iiceoeaeii?
nenoaie oa Na2+-caeaaeiiai iaoaaie?cio a?ao?aeiiiai? eeneioe o ?icaeoeo
?ioa?oacii? caaeaae? ee?oei o iiioeyoe?? ?aae?i/ooeeaeo e?ioioeeo?a
naeac?iee i?ney oioaeueiiai ?aioaai?aneiai ii?ii?iaiiy oaa?ei.
Anoaiiaeaii, ui ?i?oe?aoe?y ?icaeoeo ii?oieia?/ieo iciae aiiioico
e?ioioeeo?a i?ney ae?? ?aae?aoe?? o aeic? 1 A? a?aeaoaa?ouey iaeii/anii
c i?aeaeuaiiyi aeoeaiino? oinoie?iace N o iaia?aii?e o?aeoe??
e?ioioeeo?a, iineeaiiyi neioaco ?iiceoie-1,4,5-o?eoinoaoo oa c?inoaiiyi
??aiy oeeoicieueiiai Na2+ o ee?oeiao. Iiaeiaaeaia oo?eiaiiy i?aeaeuaii?
eiioeaio?aoe?? oeeoicieueiiai Na2+ aiane?aeie c?inoaiiy eaeueoe??ai?
i?iieeiino? ieaciaoe/ii? iaia?aie ia a?eueo i?ciueiio aoai? ?icaeoeo
?ioa?oacii? caaeaae? e?ioioeeo?a noi?iaiaeaeo?oueny aeoeaaoe??th
oinoie?iace A2, aea?eueiaiiyi a?ao?aeiiiai? eeneioe c? neeaaeo
iaia?aiieo oinoie?i?ae?a, i?aeaeuaiiyi ai?noo ia?aeiieo i?iaeoeo?a
e?iieneaaiaciiai ia?aoai?aiiy a?ao?aeiiiai? eeneioe oa i?enei?aiiyi
i?aeioeeainiiii? aeaa?aaeaoe?? o?iiaoeio. ?ic?iaeaii noaio i?iaaaeaiiy
?iaeoeiaaiiai ?aae?aoe??th neaiaeo oa eiai o?ainoi?iaoe?? o
aiiioiaaiiee, yea a?aoiao? ?ieue eaeueoe?y o ii?ooaii?
i?inoi?iai-/aniai? aca?iiae?? ??cieo eiiiiiaio?a Na2+-caeaaeieo
oinoie?i?aeieo o?ainaeoeoe?eieo nenoai o e?ioioeeoao.

Eeth/ia? neiaa: aiiioic e?ioioeeo?a, ?aae?aoe?y,
?iiceoie-1,4,5-o?eoinoao, a?ao?aeiiiaa eeneioa, e?iieneaaiaca,
eaeueoe?e, o?aaiaioaoe?y AeIE.

Iaouoaaneay I.I. Caeiiiia?iinoe ?acaeoey eioa?oaciie aeaaee eeioioeeoia
naeacaiee e?un i?e o/anoee Na2+-caaeneiuo oinoieeieaeiuo neaiaeueiuo
nenoai iinea ?aaeeaoeeiiiiai aicaeaenoaey. – ?oeiienue.

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie aeieoi?a aeieiae/aneeo iaoe ii
niaoeeaeueiinoe 03.00.04 – aeioeiey. – Eeaaneee oieaa?neoao eiaie Oa?ana
Oaa/aiei, Eeaa, 1999.

Aeenna?oaoeey iinayuaia enneaaeiaaieth ?iee iieeoinoieiiceoeaeiie
nenoaiu e Na2+-caaeneiiai iaoaaieecia a?aoeaeiiiaie eeneiou a ?acaeoee
eioa?oaciie aeaaee eeaoie a iiioeyoeee ?aaeei/oanoaeoaeueiuo eeioioeeoia
naeacaiee iinea oioaeueiiai ?aioaaiianeiai iaeo/aiey aeeaioiuo.
Onoaiiaeaii, /oi eieoeeaoeey ?acaeoey ii?oieiae/aneeo i?eciaeia iinea
aeaenoaey ?aaeeaoeee a aeica 1 A? i?ienoiaeeo iaeiia?aiaiii n
aeoeaaoeeae oinoieeiacu N, oneeaieai neioaca eiiceoie-1,4,5-o?eoinoaoa e
iiauoaieai o?iaiy oeeoicieueiiai Na2+ a eeaoeao. Aeeeoaeueiia
iiaeaea?aeaiea iiauoaiiie eiioeaio?aoeee oeeoicieueiiai Na2+ aneaaenoaea
oaaee/aiey eaeueoeeaaie i?iieoeaaiinoe ieaciaoe/aneie iaia?aiu ia aieaa
iicaeiai yoaia ?acaeoey eioa?oaciie aeaaee eeioioeeoia nii?iaiaeaeaaony
aeoeaaoeeae oinoieeiacu A2, inaiaiaeaeaieai a?aoeaeiiiaie eeneiou ec
ninoaaa iaia?aiiuo oinoieeieaeia, oaaee/aieai niaea?aeaiey ia?ae/iuo
i?iaeoeoia eeiieneaaiaciiai i?aa?auaiey a?aoeaeiiaoa e oneeaieai
iaaeioeeainiiiie o?aaiaioaoeee AeIE. ?ac?aaioaia noaia i?iaaaeaiey
eiaeooee?iaaiiiai ?aaeeaoeeae neaiaea e aai o?ainoi?iaoeee a
aiiioiaaiiue, o/eouaathuay ?ieue eaeueoeey a ia?ooaiee
i?ino?ainoaaiii-a?aiaiiiai acaeiiaeaenoaey ?acee/iuo eiiiiiaioia
Na2+-caaeneiuo oinoieeieaeiuo o?ainaeoeoeeiiiuo nenoai eeioioeeoia.

Eeth/aaua neiaa: aiiioic eeioioeeoia, ?aaeeaoeey,
eiiceoie-1,4,5-o?eoinoao, a?aoeaeiiiaay eeneioa, eeiieneaaiaca,
eaeueoeee, o?aaiaioaoeey AeIE.

Matyshevska O.P. Mechanisms of rat spleen lymphocytes interphase death
involving Ca2+-dependent phospholipid signalling systems after radiation
injury. – Manuscript.

Thesis for a doctor’s degree by speciality 03.00.04 – biochemistry. –
Kyiv Taras Shevchenko University, Kyiv, 1999.

The dissertation is devoted to investigation of the role of
polyphosphoinositide system and Ca2+-dependent metabolism of arachidonic
acid in interphase cells death in the population of radioisensitive
spleen lymphocytes after total X-ray irradiation of rats in a dose 1 Gy.
Appearance of the early morphologic features of apoptosis is shown
to be induced concurrently with phospholipase C activation,
inositol-1,4,5-triphosphate synthesis stimulation and elevation of free
cytosolic Ca2+ concentration. Increase in calcium permeability of the
lymphocytes plasma membrane leading to more sustained cytosolic Ca2+
elevation closely correlates with phospholipase A2 activation,
arachidonic acid release from membrane phospholipids, increase of the
content of initial products of arachidonic acid lypooxydation and DNA
internucleosomal fragmentation. The model of radiation induced signal
transduction and transformation into apoptotic signal considering the
role of calcium in disturbance of spatiotemporal interacion of
Ca2+-dependent phospholipid signalling systems was elaborated.

Eey words: lymphocyte apoptosis, radiation, inositol-
1,4,5-triphpsphate, arachidonic acid, lypoxygenase, calcium, DNA
fragmentation.

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