Особливості пошкодження ендотелію при розвитку експериментального артеріосклерозу менкебергівського типу: Автореф. дис… канд. мед. наук / О.В. Голдо

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IAOe?IIAEUeIA AEAAeAI?ss IAOE OE?A?IE

?INOEOOO O?C?IEIA?? ?IAI? I.I. AIAIIIEUeOess

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 Aieaeia?ia Ieaia Aieiaeeie??aia

OAeE: 616.018.74-001:546.41:612.017

INIAEEAINO? IIOEIAeAEAIIss AIAeIOAE?TH 

I?E ?ICAEOEO

AENIA?EIAIOAEUeIIAI  A?OA??INEEA?ICO

IAIEAAA?A?ANUeEIAI OEIO

14.03.04. — iaoieia?/ia o?c?ieia?y

Aaoi?aoa?ao

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa iaaee/ieo iaoe

EE?A   1999

Aeena?oaoe??th ? ?oeiien

?iaioa aeeiiaia ia eaoaae?? iaoieia?/ii? o?c?ieia??

Iaoe?iiaeueiiai iaaee/iiai oi?aa?neoaoo  ?i. I.I. Aiaiiieueoey (i.
Ee?a)

Iaoeia? ea??aieee: aeieoi? iaaee/ieo iaoe, i?ioani?

Aeoeue TH??e A?eoi?iae/,

IIO ?i. I.I. Aiaiiieueoey,

caa?aeoaa/ eaoaae?e iaoio?c?ieia??

aeieoi? a?ieia?/ieo iaoe, i?ioani?

Noa/aiei Ethaeieea Ieaenaiae??aia,

IIO ?i. I.I.Aiaiiieueoey, IAeEOe,

caa?aeoaa/ eaai?aoi???
aeaeo?iiii? i?e?ineii??

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

A?aoonue Aieiaeeie? Aaneeueiae/,

IAe? ?i.aeaae. I.Ae. No?aaeanei IIC
Oe?a?ie,

caa?aeoaa/ eaai?aoi???
iaoio?c?ieia??

aeieoi? iaaee/ieo iaoe, i?ioani?

Oaaeioia Aaaono Ieeeiiae/,

EIAII, i?ioani? eaoaae?e
iaoieia?/ii? aiaoii??

I?ia?aeia onoaiiaa: ?inoeooo aa?iioieia?? AII Oe?a?ie,

eaai?aoi??y iaoio?c?ieia?? ?
?ioiieia??

Caoeno a?aeaoaeaoueny “ 24 ” ethoiai 1999 ?. i “ 14 ” aiae.
ia can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae 26.198.01 i?e
?inoeooo? o?c?ieia?? ?i. I.I. Aiaiiieueoey IAI Oe?a?ie ca aae?anith:
252024, i. Ee?a, aoe. Aiaiiieueoey, 4.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? ?inoeoooo
o?c?ieia??

?i. I.I. Aiaiiieueoey IAI Oe?a?ie ca aae?anith: 252024, i. Ee?a, aoe.
Aiaiiieueoey, 4.

Aaoi?aoa?ao ?ic?neaiee “ 22 ” n?/iy 1999 ?.

A/aiee nae?aoa? niaoe?ae?ciaaii? a/aii? ?aaee

aeieoi? a?ieia?/ieo iaoe
Ni?ie?ia-Ia??ia C.I.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue i?iaeaie. A?oa??ineea?ic iaieaaa?a?anueeiai oeio,
oa?aeoa?iei i?iyaii yeiai ? iaae?aeaeueoeeiic, cono??/a?oueny oae naii
/anoi, ye aoa?ineea?ic, aea e aein? eeoa?oueny iaeiaea/aiith oi?iith
(Schoen F.J., 1994; Nieieiaa ?.E., 1996). Caaaeueiia?aeiii, ui
aoa?ineea?ic ?icaeaa?oueny a ae?eyieao ia?aeiiiai iioeiaeaeaiiy
aiaeioae?th (Ross R., Glomset J.A., 1976; Shimokava H., 1998). A oie aea
/an ?icaeoie iaae?aeaeueoeeiico a a?oa??aeuei?e no?ioe? aecia/a?oueny
ia?aaeon?i iae?icii ?? aeaaeaiueeiiyciaeo ee?oei (AIE). Noai ? iiaeeeaa
?ieue aiaeioae?th i?e oeueiio ? i?aeoe/ii ia aea/aieie.

Caycie i?ae eae?a?ii a?oa???, nooiaiai ?? iaaaiiaiiy oa e?euee?noth
i?iaiaeioa-e?aeueieo eiioaeo?a oa?aeoa?eco?oueny caeaaei?noth: /ei iaioa
e?euee?noue AIE a a?oa???, oei a?eueo ae?aaeaia ?aaoeyoe?y ?o
oiiono aiaeioae??i ? oei a?eueoa nooi?iue iaaaiiaiiy a?oa??? i?e
iaae?aeaeueoeeiic? (Naiioei I.A., ?iiaiaiei A.A., Naeieia I.N., 1990).
Oaeiae a?aeiii, ui ao?aoa iaaii? e?eueeino? aiaeioae?ioeeo?a
noi?iaiaeaeo?oueny ?iceaaeaie aiaeioae?ecaeaaeii? ?aaoeyoe?? oiiono
noaeeiieo iyc?a, ia?o ca ana ?c-ca a?aenooiino? aiaeioae?aeueiiai
eiiiiiaioo ?aaeoe?? ?o ?icneaaeaiiy (Iieaaiei I.I., Naaa/ A.O., Oaiiaae
E.I. ? ni?a., 1997; Griffith T.M. et al., 1984; Inagami T., Naruse M.,
Hoover R., 1995). ?c neacaiiai aeieeaa?, ui ?a/iaeie, ye? iaeii/anii
ni?e/eiththoue niaci AIE ? iioeiaeaeaiiy aiaeioae?th, iiaeooue
?iaeoeoaaoe ia?anei?i/aiiy AIE oa iaae?aiae?ic. Aeaiei aeiiaai
?aeaaeueii a?aeiia?aeathoue eaoaoieai?ie, ye? aia?oa aoee aeei?enoai?
aeey iiaeaethaaiiy a?oa??ineea?ico Iaieaaa?aa (Oe?ieiai?ia E.E., 1907).
Ana oea aea? i?aenoaae aeey i?eiouaiiy, ui iioeiaeaeaiiy aiaeioae?th
iiaea a?ae?a?aaaoe iaaio ?ieue a iaoiaaiac? a?oa??aeueiiai
iaae?aeaeueoe?iico.

?acii c oei, aeey ?a/iaei, ye? ia ? aacieiino?eeoi?aie, aea oaeiae
caeaoi? ?i?oe?thaaoe iaae?aneea?iciaaiac (iaoaaie?/i? yaee, a?oai?i Ae3
oa ?i.), ne?ae ia?aaeaa/aoe ?io? iaoai?cie iaoiaaiii? ae?? ia noaeeiio
no?ieo. Cie?aia, ni?eyoe ?? iioeiaeaeaiith iiaea a?ia?eaeueoe??i?y ca
oiia iaaeinoaoiino? iaoai?ci?a ?aaoeyoe?? eiioeaio?aoe??
aioo??oiueiee?oeiiiai eaeueoe?th ([Na2+]i) a AIE. Ia ei?enoue oeueiai
i?eiouaiiy na?ae/eoue ? oa, ui eaoaoieai?ie noeioeththoue
ooieoe?iiaeueio aeoeai?noue ia?aoe?ioeeo?a (Hargis G. et al., 1980;
Fischer J.A. et al., 1982). Iaaeinoaoiy e?euee?noue ?ioi?iaoe?? i?i
ci?ie, ye? aeieeathoue a nenoai? ?aaoeyoe?? eaeueoe??i?? i?e ?icaeoeo
a?oa??aeueiiai eaeueoeeiico, aeicaiey? eeoa a caaaeueieo ?enao
iaaiai?thaaoe o/anoue iaoanoaoe/ii? eaeueoeeo?eaoe?? a eiai iaoiaaiac?.
Aeiaeaoeia? a?aoiaioe ia ei?enoue aeaiiai iaoai?cio eaeueoeeo?eaoe??
iiaeia io?eiaoe i?e ia?aeaeueiiio aecia/aii? ai?noo caaaeueiiai
eaeueoe?th, aeoeaiino? eoaeii? oinoaoace (EO) ? -aeooai?eo?ainoa?ace
(-AO) a ne?iaaooe? e?ia? i?aeaeine?aeieo oaa?ei o ?aiiueiio ia??iae?
?icaeoeo iaae?aeaeueoeeiico ai?oe. Anoaiiaeaii, ui ne?iaaoeiaa EO ?
noi?ooth i?eaeecii ??aii? e?eueeino? e?noeiaeo ? ia/?ieiaeo ?cioa?iaio?a
(Wolf P.L., 1979). Ine?eueee i?aeaeuaiiy aeoeaiino? -AO a ne?iaaooe?
a?aeia?aaea? ca?eueoaiiy aeoeaiino? ia/?ieiaeo ?cioa?iaio?a EO, c??no
aaee/eie a?aeiioaiiy “ne?iaaoeiaa EO/ -AT” na?ae/eoue ia ei?enoue ?inoo
aeie? e?noeiaeo ?cioa?iaio?a EO ca ?aooiie ca?eueoaiiy ooieoe??
inoaiaeano?a i?ae ae??th ia?aoai?iiio (Eoean I., Aoaeoaae Aeae., 1985;
Oaiia?iai Aeae., Oaiia?iai O., 1989; Oeoa? I.I., 1992; Posen S., Doherty
E., 1981).

Naiino?eiith i?iaeaiith ? ??cia /ooeea?noue a?oa??e ? aai aei ?icaeoeo
neea?i-oe/ieo ci?i. Anoaiiaeaii, ui iaeiei c iaoai?ci?a aeniei?
?acenoaioiino? AIE aai, ii??aiyeueii c AIE a?oa??e, aei ae?? iaoiaaieo
oaeoi??a ? a?eueo aeniea caeaoi?noue ia?oeo aei a?aeiiaeaiiy ii?iaeueii?
[Ca2+]i ca ?aooiie a?eueoi? iiooaeiino? iaoai?ci?a ?aaoeyoe?? (Aeoaeei
I.I., Aeoeue TH.A., 1994). Ia aeeeth/aii, ui naaiaioa?ia
iaiaeii??aei?noue aiaeioae?th (Aonaa N.A. e niaao., 1988; ?na?/e?ia
?.I., Oa?aiaiei A.I., 1990; Naaa/ A.O., Oea/aiei I.I., Aaceethe I.A.,
1992; Iton H. et. al., 1994) oaeiae a?ae?a?a? a oeueiio iaaio ?ieue.

Oaeei /eiii, iiaeia aaaaeaoe, ui aea/aiiy a ?aiiueiio ia??iae? ?icaeoeo
aenia?eiaioaeueiiai iaae?aeaeueoeeiico ai?oe oeueo?ano?oeoo?e
aiaeioae?th noaeei ??cieo oei?a c iaoith anoaiiaeaiiy oa?aeoa?ieo aeey
ai?oe iniaeeainoae iioeiaeaeaiy aiaeioae?th, a oaeiae ai?noo caaaeueiiai
eaeueoe?th, aeoeaiino? EO ? -AO a ne?iaaooe? e?ia? uo??a aiana iaaio
yni?noue a cacia/aio i?iaeaio.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. ?iaioo aeeiiaii
a ?aieao iaoeiai? ?iaioe eaoaae?e iaoieia?/ii? o?c?ieia?? IIO ?i. I.I.
Aiaiiieueoey: “Aea/aiiy ?ie? eaeueoe?th a iaoai?ciao iioeiaeaeaiiy
iyc?a ??cieo oei?a”.

Iaoa aeine?aeaeaiiy: aecia/aiiy iniaeeainoae oeueo?ano?oeoo?ii?
i?aai?caoe?? aiaei-oae?th, ai?noo caaaeueiiai eaeueoe?th a ne?iaaooe? ?
iiaeeeaeo ci?i a nenoai? eiai ?aaoey-oe?? o uo??a a ?aiiueiio ia??iae?
?icaeoeo aenia?eiaioaeueiiai iaae?aeaeueoeeiico ai?oe.

Caaea/? aeine?aeaeaiiy: 1.  Aeine?aeeoe oeueo?ano?oeoo?io i?aai?caoe?th
aiaeioae?th noaeei ??cieo oei?a, iaiaeiaeiai /ooeeaeo aei ?icaeoeo
neea?ioe/ieo ci?i (ai?oa, naeac?ieiaa a?oa??y, caaeiy ii?iaeienoa aaia)
ia ?aii?o aoaiao ?icaeoeo a?oa??ineea?ico iaieaaa?a?anueeiai oeio i?e
??cieo niiniaao eiai iiaeaethaaiiy.

2. I?iaanoe ii?oiiao?e/iee ? ii??aiyeueiee aiae?c oeueo?ano?oeoo?ieo
ci?i aiaeioae?th noaeei ??cieo oei?a o i?aeaeine?aeieo oaa?ei.

3. Aecia/eoe ci?ie ai?noo caaaeueiiai eaeueoe?th,aeoeaiino? eoaeii?
oinoaoace ? -aeooa-i?eo?ainoa?ace a ne?iaaooe? e?ia? uo??a a oiiaao
aenia?eiaioaeueiiai iiaeaethaaiiy iaae?aeaeueoeeiico ai?oe.

Iaoeiaa iiaecia iaea?aeaieo ?acoeueoao?a.  A aeena?oaoe?ei?e ?iaio?
aia?oa i?e aeine?aeaeaii? a?oa??ineea?ico iaieaaa?a?anueeiai oeio
nooo?aa oaaaa i?eae?ey?oueny aiaeioae?th. Anoaiiaeaii, ui iioeiaeaeaiiy
aiaeioae?th ??ciiai nooiaiy oyaeeino? ia? i?noea a ai?o?, naeac?ieia?e
a?oa??? ? caaei?e ii?iaeieno?e aai? aaea a ?aiiueiio ia??iae? ?icaeoeo
aenia?eiaioaeueiiai iaae?aeaeueoeeiico ai?oe (a?oa??ineea?ico
iaieaaa?a?a-nueeiai oeio). Ii??aiyeueiee aiae?c ci?i oeueo?ano?oeoo?e
aiaeioae?th noaeei, /ooeeaeo ? no?eeeo aei ?icaeoeo iaae?aeaeueoeeiico,
na?ae/eoue i?i oa, ui a ia?oo aeiao aenia?eiaioaeueiiai aieeao (aino?a
a?ia?aae?aiae?iai?y) ianeaia iioeiaeaeaiiy aiaeioae?th (noaoioaeueia
aeaioaeaoe?y, aeano?oeoe?y aacaeueii? iaia?aie) ia? i?noea o?eueee a
ai?o?.

Aia?oa c?iaeaia ni?iaa ioe?ieoe ?ieue iioeiaeaeaiiy aiaeioae?th a
iaoiaaiac? a?oa??aeueiiai iaae?aeaeueoeeiico.

Oai?aoe/ia ? i?aeoe/ia cia/aiey iaea?aeaieo ?acoeueoao?a.  ?acoeueoaoe
aeine?aeaeaiue aeicaieeee aecia/eoe oa?aeoa? ? iniaeeaino? ci?i
oeueo?ano?oeoo?ii? i?aai?caoe?? aiaeioae?th, ye? aeanoea? ?aiiueiio
ia??iaeo ?icaeoeo aenia?eiaioaeueiiai iaae?aeaeue-oeeiico ai?oe.
Io?eiaii iia? aeai? i?i ?nioaaiiy a iaoiaaiac? “aae?aiae?iiaiai”
iaae?aeaeueoeeiico ai?oe oaei? aaaeeeai? eaiee, ye aino?a ianeaia
iioeiaeaeaiiy ?? aiaeioae?th. Oea aeicaiey? c?iaeoe i?eioeeiiaa
aeiiiaiaiiy a ?nioth/? oyaeaiiy uiaei iaoiaaiaco a?oa??ineea?ico
iaieaaa?a?anueeiai oeio. Iioeiaeaeaiiy aiaeioae?th iiaea aooe i?e/eiith
ii?ooaiiy ?aeaenaoe??, ia?anei?i/aiiy ? iae?ioe/ieo ci?i AIE c
iiaeaeueoei ?icaeoeii iaae?aeaeueoeeiico a?oa??e. I?e IEA-?ioieneeaoe??
ye iaoai?ci iaae?aiae?ico a?eueo a??ia?aeia i?yia ae?y IEA
aaciina?aaeiuei ia AIE na?aaeiuei? iaieiiee ai?oe. Iioeiaeaeaiiy
aiaeioae?th i?e oeueiio iiaea aenooiaoe eeoa a ?ie? oiiae, yea ni?ey?
?icaeoeo iaae?aeaeueoeeiico.

Aeiaeaoeiao iaoiaaiaoe/io ?ieue a ?aiiueiio ia??iae? ?icaeoeo
a?oa??ineea?ico Iaieaaa?aa iiaeooue iaoe aeyaeai? ci?ie a nenoai?
?aaoeyoe?? eaeueoe??i??.

Io?eiai? ?acoeueoaoe iaaeathoue iiaeeea?noue ?ic?iaee iaoiaaiaoe/ii
iaa?oioiaaieo caoiae?a i?io?eaeoeee eaeueoeeiico a?oa??e.

Iniaenoee aianie caeiaoaa/a. Caeiaoaa/ai i?iaaaeaii aiae?c e?oa?aoo?ieo
aeaea?ae, aeeiiaii iaiao?aei? aenia?eiaioaeuei? aeine?aeaeaiiy,
noaoenoe/ii ia?iaeaii, i?iaiae?-ciaaii oa ocaaaeueiaii io?eiai? aeai?.
I?eaiooaaiiy oeueo?aoiieeo c??c?a noaeei, ye? aeei?enoaii o ?iaio?, ?o
ia?aaeyae a aeaeo?iiiiio i?e?ineii? i?iaaaeaii ca o/anoth ni?a?ia?oiee?a
eaai?aoi??? aeaeo?iiii? i?e?ineii?? IAeEOe IIO ?i. I.I. Aiaiiieueoey.

Ai?iaaoe?y ?iaioe. Iniiai? iieiaeaiiy ?iaioe aoee i?aaenoaaeai? ia
Ieaioi? iaoeiai-iaaee/iiai oiaa?enoaa iaoio?c?ieia?a Oe?a?ie (i. Euea?a,
23-25 aa?aniy 1993), II eiia?an? iaoio?c?ieia?a Oe?a?ie (i. Ee?a, 1996),
can?aeaii? Ee?anueeiai oiaa?enoaa iaoio?c?ieia?a (i. Ee?a, 9 /a?aiy
1998), can?aeaii? ai?iaaoe?eii? ?aaee c i?iaeai «Oai?aoe/ia iaaeeoeeia»
IIO ?i. I.I. Aiaiiieueoey  (i. Ee?a, 11 /a?aiy 1998).

Ioae?eaoe??. Ca ?acoeueoaoaie ?iaioe iioae?eiaaii 8 iaoeiaeo ?ia?o, c
ieo 5 — aeo?iaeuei? noaoo?.

No?oeoo?a oa ianya aeena?oaoe??. Aeena?oaoe?y neeaaea?oueny c anooio,
iaeyaeo e?oa?aoo?e, iieno iaoa??ae?a oa iaoiae?a aeine?aeaeaiiy,
aeeeaaeaiiy ?acoeueoao?a oa ?o iaaiai?aiiy, aeniiae?a oa nieneo 217
aeei?enoaieo aeaea?ae. ?iaioa aeeeaaeaia ia 120 noi??ieao, i?noeoue 10
oaaeeoeue oa ?ethno?iaaia 20 iaethieaie.

INIIAIEE CI?NO

Iaoa??aee oa iaoiaee aeine?aeaeaiue

Aeine?aee aeeiiai? ia 83 aacii?iaeieo a?eeo uo?ao-naioeyo, ye?
oo?eioaaaaeenue ia noaiaea?oiiio ?aoe?ii? a?aa??th. Iaoa??aeii aeey
aeine?aeaeaiiy aoee oiaoi/ee noaeei oa ne?iaaoea e?ia? ?ioaeoieo ?
i?aeaeine?aeieo uo??a.

Iiaeaethaaiiy iaae?aeaeueoeeiico ai?oe i?iaaaeaii a oiiaao “aino?iai” ?
“i?aeaino?iai” aieea?a aae?aiae?io ? iiiieiaeaoeaoaoo (IEA). ?ic/ei
aae?aiae?io a?ae?ioei??aeo c ?ic?aooieo 50 iea/ea aaiaeeee iaeii?aciai
i?e «aino?iio» aieea? ? uiaeaiii i?ioyaii 6 aei?a — i?e «i?aeaino?iio».
A oiiaao “aino?iai” aieeao ?ic/ei IEA aaiaeeee a aeic? 25 ia/ea.
«I?aeaino?o» IEA-?ioieneeaoe?th a?aeoai?thaaee ca noaiith: 1-e aeaiue —
25ia/ea, ianooii? 5 aei?a — ii 10 ia/ea uiaeiaiai. E?ioeaaiai cia/aiiy
?I ?ic/eio IEA (7,4+0,02) aeinyaaee ca aeiiiiiaith iane/aiiai ?ic/eio
NaHCO3 a o?c?ic/ei?. An? ?ic/eie aeaioiaeyee ? aaiaeeee a /a?aaio
ii?iaeieio c aeio?eiaiiyi i?aaee anaioeee.

Aeoiaey/e c oiai, ui iaeia?o? ii?oieia?/i? i?iyae a?oa??ineea?ico
Iaieaaa?-aa niinoa??aathoueny /a?ac aeiao i?ney aaaaeaiiy a?aeiia?aeiiai
i?aia?aoo, iaoa??ae a?aee /a?ac aiaeeio ? aeiao i?ney aino?iai oa ia 7-o
aeiao i?aeaino?iai aieeao. I?ney aeaeai?oaoe?? oaa?ei cae?aee e?ia aeey
io?eiaiiy ne?iaaoee (Niieyieoeeee A.ss., 1987). Oaeaeei aeae?eyee
oiaoi/ee noaeei, ye? ia?iaeyee caaaeueiii?eeiyoeie iaoiaeaie aeey
iiaeaeueoiai aeaeo?iiiii?e?ineii?/iiai aiae?co (Ea?oio A.ss.,
1984).Ia?eoe caeeaaee a aeaeaoeiia? eainoee ? iie?ia?ecoaaee i?e +60i N.
C aeie?a aeaioiaeyee iai?aoiie? c??ce, ye? oa?aoaaee oieo?aeeiiaei nei?i
c iaoith ?aeaioeo?eaoe?? noaeei ia na?oeiiioe/iiio ??ai?. Ia
oeueo?ai?e?ioii? LKB-3 aeaioiaeyee oeueo?aoiie? c??ce, ye? eiio?anooaaee
o?ai?eaoeaoaoii ? oeeo?aoii naeioeth (Watson, 1958; Reynolds, 1963),
ia?aaeyaeaee a aeaeo?iiiiio i?e?ineii? AIA-100A?, oioia?aooaaee. C
iaoith e?euee?nii? ioe?iee oeueo?ano?oeoo? aiaeioae?aeueieo ee?oei (AE)
aoa canoiniaaiee iaoiae ii?oiiao?e/iiai aiae?co (Aaoaiaeeeia A.A.,
Iaaci?ia A.I., Iaaci?iaa I.O., 1984). Aeei?enoiaoaaee aecia/aiiy
ianooiieo noa?aiiao?e/ieo ia?aiao??a i?oioiiae??e oa ca?ienoi?
aiaeiieaciaoe/ii? n?oee (CAN): /anoea ia?io i?aaiaee a oeeoiieaci? ( %);
e?euee?noue i?aaiae a iaeeieoe? ia?io oeeoiieacie (/enaeueia
u?euei?noue, 1/iei3); ieiua (iei2) oa oaeoi? oi?ie (oiia.iae.).
Iaiao?aei? aei??thaaiiy i?iaiaeeee ca aeiiiiiaith i?ia?aie «I?aaiaea».
Aeine?aeaeaiiy cae?enithaaeenue a IAeEOe IIO ?i.I.I. Aiaiiieueoey.

Ai?no caaaeueiiai eaeueoe?th a ne?iaaooe? e?ia? aecia/aee ca
eieuei?iaith ?aaeoe??th c iaoeeoeiieiaei aeaeeoiei (Gindler, King,
1972). Aecia/aiiy aeoeaiino? eoaeii? oinoaoace i?iaiaeeee ca
?ioaineai?noth a?ae?ie?co n-i?o?ioai?eoinoaoo (Bessey, Lowry, Brock,
1940). Aeoeai?noue -aeooai?eo?ainoa?ace aecia/aee ca oi?o?eiaaiith
iaoiaeeeith c noano?aoii  L—aeooai?e-4-i?o?iai?e?aeii  ?
ae?oe?eae?oeeiiaei aooa?ii (Nielsen, 1978). B ?iaio? aeei?enoai? iaai?e
?aaeoea?a “Roche” (I?ia//eia) oa a?aeiia?aei? ?ino?oeoe?? ii ?o
aeei?enoaiith. Oeeo?ia? aeai?, io?eiai? a ?acoeueoao? aeine?aeaeaiue,
ia?iaeyee iaoiaeii aa??aoe?eii? noaoenoeee c aeei?enoaiiyi e?eoa??th
«t» Nouethaeaioa ? oaaeeoeue a??ia?aeiino? «?» O?oa?a. An? ?ic?aooiee
ii noaoenoe/i?e ia?iaoe? aeaieo cae?eniaii ia i?e?ieaeueeoeyoi?? «IE-61»
ca a?aeiia?aeieie i?ia?aiaie (Eaaiia TH.E., Iiai?aethe I.I., 1990).

?acoeueoaoe Aeine?aeaeaiue oa ?o iaaiai?aiiy

A ?acoeueoao? i?iaaaeaieo aeine?aeaeaiue aoei anoaiiaeaii, ui
iioeiaeaeaiiy ??ciiai nooiaiy oyaeeino?, caaeaaeue ? ianooiia
aeaneaaiaoe?y aiaeioae?th a ai?o?, naeac?i-eia?e a?oa??? ? caaei?e
ii?iaeieno?e aai?, a oaeiae ci?ie ai?noo caaaeueiiai eaeueoe?th a
ne?iaaooe? e?ia? i?aeaeine?aeieo oaa?ei aeyaeythoueny aaea a ?aiiueiio
ia??iae? ?icaeoeo aenia?eiaioaeueiiai iaae?aeaeueoeeiico ai?oe
(a?oa??ineea?ico iaieaaa?a?anueeiai oeio). Oea ?noioii aeiiiaith?
?nioth/? oyaeaiiy i?i oa, ui aae?aiae?i a oiene/ieo aeicao aea??eiai
a?aaea? ee?oeie na?aaeiuei? iaieiiee e?oiieo a?oa??e c ianooiiei
?icaeoeii oiene/iiai a?oa??ineea?ico (No?oeia A.E., Na?ia A.A., 1979).

Io?eiai? iaie aeai? aeicaieythoue oaeiae noaa?aeaeoaaoe, ui /ooeea?noue
aiaeioae?th ai?oe, naeac?ieiai? a?oa??? ? ii?iaeienoi? aaie uo??a aei
ae?? iiiieiaeaoeaoaoo ? aae?aiae?io ??cia. Aeaia caeeth/aiiy aeo?ea? ?c
?acoeueoao?a aiae?co ci?i ?yaeo ye?nieo ? e?euee?nieo e?eoa???a,
noeoii?noue yeeo aeicaieth? ioe?ieoe oa?aeoa? oa oyaee?noue no?oeoo?ieo
ci?i aiaeioae?th, a oaeiae eiai ?aia?aoeaieo iiaeeeainoae. Iaaecae/aeii
ae?aaeaia i?aaenoaaieoeoai i?aaiae neioaoe/iiai oeio a AE naeac?ieiai?
a?oa??? ? ii?iaeienoi? aaie a ie?ai? ia??iaee niinoa?aaeaiiy aea?
i?aenoaao noaa?aeaeoaaoe i?i ?aia?aoeaio ?aaaia?aoe?th aiaeioae?th ye
aeineoue iiooaeiee iaoai?ci eiai a?aeiiaeaiiy oa oi?ioaaiiy aiane?aeie
oeueiai aeniei? no?eeino? ie?aieo noaeeiieo ?aa?ii?a aei ae?? iaoiaaieo
oaeoi??a.

Oeueo?ano?oeoo?a aiaeioae?th noaeei a oiiaao “ aae?aiae?iiai?
iiaeae?” iaae?aeaeueoeeiico ai?oe (a?oa??ineea?ico
iaieaaa?a?anueeiai oeio)

sse iieacaee iao? aeine?aeaeaiiy, ia ?aii?o noaae?yo ?icaeoeo
aenia?eiaioaeueiiai a?oa??ineea?ico iaieaaa?a?anueeiai oeio a aiaeioae??
ai?oe, naeac?ieiai? a?oa??? ? caaeiuei? ii?iaeienoi? aaie uo??a
a?aeaoaathoueny aeano?oeoeai? ci?ie oeueo?ano?oeoo? oeeoieaie,
oeeoiieacie, yae?a ? i?aaiae AE, ?o o?aaiaioaoe?y, iae?ic, aeaneaaiaoe?y
oa aeiaeaoeia? eiioaeoe c AIE ? ee?oeiaie e?ia?. Oea iiaiaeaeo?oueny c
aeaieie e?oa?aoo?e uiaei iaeiioeiiino? oeueo?ano?oeoo?ieo ci?i
aiaeioae?th i?e ae?? ia inoaii?e ??cieo iioeiaeaeoth/eo oaeoi??a
(Aieii-ssnaiaoeeay O.A. e niaao., 1988; Noaieia I.E. e niaao., 1989). A
oie aea /an io?eiai? aeai? na?ae/aoue i?i oa, ui ae?aaeai?noue
iioeiaeaeaiiy aiaeioae?th iaiaeiaeiaa a noaeeiao, /ooeeaeo ? no?eeeo
aei ?icaeoeo iaae?aeaeueoeeiico.

Ca iaoeie aeaieie, aaea /a?ac aiaeeio i?ney aaaaeaiiy aae?aiae?io a
ie?aieo ae?eyieao ai?oe a?aeaoaa?oueny aiaieuaaa aeaneaaiaoe?y
aiaeioae?th. Aeaye? AE ciaoiaeyoueny a noai? nei?i/aiiy, ?io? iathoue
i?ina?oeaio oeeoiieacio ? iioeiaeaeai? i?oioiiae???. Ca ?acoeueoaoaie
ii?oiiao?e/ieo aeine?aeaeaiue, ca?eueoaiiy /eneiai? u?eueiino?
i?oioiiae??e i?iii?oe?eii ciaioaiith ?o ieiu? (oaae. 1). Oea aeicaiey?
i?eionoeoe iiae??aiaiiy i?aaiae. CAN iaaoaa? a?eueoi? iie?ii?oiino?. I?i
oea na?ae/eoue c?inoaiiy iieaciee?a /eneiai? oa ia?iii? u?eueiino?
eaiaeueoe?a ia oii? caa?aaeaiiy ?o ii/aoeiaeo ?ici???a (oaae. 1).
A?aeaoaa?oueny ?icioooaaiiy ? /anoeiaa o?aaiaioaoe?y aacaeueii? iaia?aie
aiaeioae?th, iaa?ye noaaiaeioae?aeueiiai i?inoi?o.

Ii??aiyeueiee aiae?c ci?i oeueo?ano?oeoo?e aiaeioae?th a ai?o?,
naeac?ieia?e a?oa??? ? caaei?e ii?iaeieno?e aai? uo??a /a?ac aeiao i?ney
aaaaeaiiy aae?aiae?io aea? aaaeeeao ?ioi?iaoe?th uiaei a?aei?iiinoae
ci?i aiaeioae?th a noaeeiao ??cieo oei?a.

A i?ina?o? ai?oe aeoaea /anoi niinoa??aathoueny caeeoee AE, a ae?eyieao
aeaioaeaoe?? — aa?aaaoe?y ? aaeaac?y o?iiaioeeo?a. Ia aaaaoueio c??cao
AE i?aeianai? iaae iiaa?oiath noaaiaeioae?th iaa?yeiaith ??aeeiith. A
AE, ui caa?aaee caycie c noaaiaeioae??i, aeyaeythoueny /eneaii? aai
aaeee? aaeoie?, c?inoa? (aei naaiaioaoe??) caeaeno?noue eiioo??a yaea?.
?o o?aaiaioe /anoi cnoioo? ia ia?eoa??th a?aaionueeeie aaeoieyie. A AE
ia?aaaaeathoue i?oioiiae??? c ooeiaeaeaieie ciai?oi?ie iaia?aiaie,
i?ina?oeaiei iao?eenii, ?aaeoeiaaieie e??noaie. E?euee?noue i?oioiiae??e
ii??aiyii c iiia?aaei?i oa?i?iii niinoa?aaeaiiy ciaioaia. A oie aea /an
ca ?aooiie ae?aaeaiiai iaa?yeo i?oioiiae??e ca?eueoaiee ia?i
oeeoiieacie, yeee aiie caeiathoue a AE (oaae. 1). Ciaioaiiy caaaeueii?
e?eueeino? i?oioiiae??e i?e oeueiio iiaeia iiynieoe ?o iniie?cii. Aiae?c
ci?i ii?oiiao?e/ieo iieaciee?a ? ye?nieo iniaeeainoae CAN aeicaiey?
c?iaeoe aeniiaie, ui ?aaeoe?y inoaiiuei? a?ae??ciy?oueny a?ae ?aaeoe??
i?oioiiae??e (aeano?oeoeai? ci?ie a inoaii?o ae?aaeai? a a?eueo?e i???).
Ii?ooaia no?oeoo?a aacaeueii? iaia?aie aiaeioae?th. Ianeaia
iioeiaeaeaiiy aiaeioae?th ni?e/eith? noaoioaeueio aeaioaeaoe?th no?iee
ai?oe ? iaa?ye, yeee ?iciianthaeaeo?oueny ia aioo??oi? oa?e na?aaeiuei?
iaieiiee.

A naeac?ieia?e a?oa??? ? caaei?e ii?iaeieno?e aai? aeeno?io?/i? ci?ie AE
? ?o i?aaiae ia iaaoaathoue oaeiai nooiaiy ae?aaeaiino?, ye a ai?o?.
Ci?ie aiaeioae?th a a?eueoino? aeiaaee?a iaiaaeai? iaa?yeii ee?oei oa ?o
i?aaiae, ?, ye i?aaeei, ia noi?iaiaeaeothoueny aeaioaeaoe??th noaeeiii?
no?iee. Oae, a AE aaie ca?eueoaia na?aaeiy ieiua i?oioiiae??e —
i?iii?oe?eii ca?eueoaiith ia?io, yeee aiie caeiathoue a oeeoi-ieaci? AE.
Iaci?ii?noue iieacieea /eneiai? u?eueiino? i?e oeueiio (oaae. 2) iiaea
aooe iciaeith ca?eueoaiiy ?ici???a i?oioiiae??e ye aiane?aeie ?o
iaa?yeo, oae ? a?ia?o?io??. Oio?einue ae i?aee?aneeoe, ui aeeno?io?/i?
ci?ie i?oioiiae??e a AE aaie ae?aaeai? cia/ii iaioa, i?ae a ai?o?, a
cayceo c /ei ia niinoa??aathoueny iniie?c ? ciaioaiiy e?eueeino?
i?oioiiae??e. Aiae?c noaio CAN (ca?eueoaiiy /enaeueii? ? ia?iii?
u?eueiinoae eaiaeueoe?a, oaae. 2) aeicaiey? i?eionoeoe, ui a AE ia?
i?noea ?? a?ia?o?io?y. Aacaeueia iaia?aia aiaeioae?th naeac?ieiai?
a?oa??? ? caaeiuei? ii?iaeienoi? aaie a aeaiee ia??iae niinoa?aaeaiiy
caa??aa? naith cae/aeio oeueo?ano?oeoo?io i?aai?caoe?th.

Oaaeeoey 1. Ii?oiiao?e/i? ia?aiao?e i?oioiiae??e ? ca?ienoi?
aiaeiieaciaoe/ii? n?oee aiaeioae?th /a?aaii? ai?oe uo??a i?e
a?ia?aae?aiae?iai?? (I + m)

Ia??iae niinoa?aaeaiiyIa?iia u?euei?noue (%)*enaeueia u?eue-i?noue
(10-2/iei3)Ieiua i?io?-e?a (10-2iei2)Oaeoi? oi?ie

i?aaiaeeI?OIOIIAe???, Eiio?ieue2,310,482,380,3693,138,710,900,011
aiaeeia i?ney aino?i? a?ia?aae?aiae?iai??2,570,76 27,14+5,63

*12,931,94

* 0,880,051 aeiaa i?ney aino?i? a?ia?aae?aiae?iai??7,341,47

* ** 11,28+2,48

* ** 46,094,20

* ** 0,910,01

**7-e aeaiue i?aeaino?i? a?ia?aae?aiae?iai??3,081,27

**12,982,53

* 24,053,73

* **0,830,04

* **CAN,

Eiio?ieue1,12 0,1633,54 8,067,76 0,600,51 0,021 aiaeeia i?ney
aino?i? a?ia?aae?aiae?iai??6,78 1,12

*123,5339,98

*9,42 1,800,83 0,021 aeiaa i?ney aino?i? a?ia?aae?aiae?iai??1,95
0,27

* **51,2410,167,34 0,630,71 0,027-e aeaiue i?aeaino?i?
a?ia?aae?aiae?iai??0,83 0,16

* **162,0256,38

* **1,88 0,19

* **0,47 0,02Ooo ? a oaae. 2 — 6 * —
a??ia?aeii o ni?anoaaeaii? c eiio?ieai

** — a??ia?aeii a?aeiinii iiia?aaeiueiai oa?i?io niinoa?aaeaiiy (p
 <  0,05) Oaaeeoey 2. Ii?oiiao?e/i? ia?aiao?e i?oioiiae??e ? ca?ienoi? aiaeiieaciaoe/ii? n?oee aiaeioae?th caaeiuei? ii?iaeienoi? aaie uo??a i?e a?ia?aae?aiae?iai?? (I + m) Ia??iae niinoa?aaeaiiyIa?iia u?euei?noue (%)*enaeueia u?eue-i?noue (10-2/iei3)Ieiua i?io?e?a (10-2 iei2) Oaeoi? oi?ieI?OIOIIAe??? Eiio?ieue 4,300,52 14,683,82 18,792,31 0,890,021 aeiaa i?ney aino?i? a?ia?aae?aiae?iai?? 8,121,72*15,651,8936,632,29*0,900,027-e aeaiue i?aeaino?i? a?ia?aae?aiae?iai??7,541,887,792,16**56,578,99* **0,810,05AIN, Eiio?ieue1,900,43135,8818,123,060,370,450,031 aeiaa i?ney aino?i? a?ia?aae?aiae?iai??3,410,41*278,9148,64*3,230,240,660,027-e aeaiue i?aeaino?i? a?ia?aae?aiae?iai?? 6,611,36* **40,411,05* **8,021,13 * **0,770,03Aeine?aeaeaiiy ? ii??aiyeueiee aiae?c ci?i oeueo?ano?oeoo?e aiaeioae?th a noaeeiao ??cieo oei?a o uo??a i?e i?aeaino??e a?ia?aae?aiae?iai?? oaeiae aeyaeee ?aa?iiaeuei? a?aei?iiino? ci?i aiaeioae?th. A ai?o? iciaee a?oaeo ii?ooaiue no?oeoo?e aiaeioae?th i?aeoe/ii a?aenooi?, a oie /an ye a aiaeioae?? naeac?ieiai? a?oa??? ? caaeiuei? ii?iaeienoi? aaie iaee i?noea aeano?oeoeai? ci?ie. I?e oeueiio ?o oyaee?noue aoea a?eueoith, i?ae i?e aino??e a?ia?aae-?aiae?iai??. sse iieacaee iao? aeine?aeaeaiiy, ia 7-o aeiao i?aeaino?i? a?ia?aae?aiae?iai?? a aiaeioae?? ai?oe ii?iae?co?oueny no?oeoo?a i?oioiiae??e, a?aeiiaeth?oueny aaee/eia iieacieea ?o ia?iii? u?eueiino?. Na?aaeiy ieiua i?oioiiae??e ciaioo?oueny a 3,9 ?ac?a a /enaeueia ?o u?euei?noue ia?aa?eueoo? eiio?ieueia cia/aiiy a 5,4 ?ace (oaae. 1). Ca?eue-oaiiy e?eueeino? aeia?aaeaea? ciaioaiiy ieiu? i?aaiae ? aeicaiey? i?eioneaoe, ui a?aeiiaeaiiy noaio i?oioiiae??iio a AE a?aeaoaa?oueny ca ?aooiie iiaiooai?aiiy i?oioiiae??e. Aiaeia?/i? (a?ia?ieanoe/i?) ci?ie a?aeaoaathoueny ? a CAN (oaae. 1). Ia oii? iienaiiai aeua ianeaiiai iioeiaeaeaiiy aiaeioae?th ai?oe i?e aino??e a?ia?aae?aiae?iai?? oea ? aeaui ianiiae?aaiei. O cayceo c oeei iaoo oaaao i?eaa?ioee no?oeoo?e na?aaeiuei? iaieiiee ai?oe, a ye?e, ca aeaieie e?oa?aoo?e, a aeaiee oa?i?i iathoue i?noea yaeua iaae?aeaeueoeeiico. Ae?enii, a na?aaei?e iaieiioe? niinoa??aaeenue aeeno?io?/ii ci?iai?, iaa?oue c yaeuaie iae?ico, AIE ? ee?oeie-"iaaoee", ye? aoee noenioo? ?icoe?aiei ? ae?ae ou?eueiaiei aeanoe/iei ea?eanii ai?oe. Ia i?aenoaa? io?eiaieo aeaieo iiaeia aaaaeaoe, ui a?aeiinia caa?aaeai?noue aiaeioae?th ai?oe i?ney 6-aeaiiiai aaaaeaiiy aae?aiae?io ? iane?aeeii ci?ie /ooeeaino? AE aei eiai iioeiaeaeoth/i? ae?? oa aoaeoeaii? ?aia?aoe?? iiia?aaeiueiai iioeiaeaeaiiy. A?aeiii, ui iioeiaeaeaiiy aiaeioae?th a?oa??e eaoaoieai?iaie iiaycaii c aeoeaaoe??th aae?aii?aoeaioi??a AE (Eiiaeaeia E.E. e niaao., 1989). Ca aeaieie Iton H. et al. (1994), na?aae -aae?aii?aoeaioi??a a a?oa??? ia?aaaaea? 1- , a a aai? - 2- noaoei. Iiaeia, oaeei /eiii, aeoiaoe, ui aeniea /ooeea?noue aiaeioae?th ai?oe aei iioeiaeaeoth/i? ae?? aino?i? a?ia?aae?aiae?iai?? iaoiiaeaia iaeneiaeueiei (ii??aiyiii c noaeeiaie aoaeue-yeiai ?ioiai oeio) i?aaenoaaieoeoaii 1-aae?aii?aoeaioi??a. sse a?aeiii, na?aae iane?aee?a i?neyae?? aaii?no?a c 1-aae?aii?aoeaioi?aie, e??i ca?eueoaiiy [Ca2+]i oa iioeiaeaeaiiy ee?oeie, iiaeeeaee aoaeo aeoeaaoe?? (Na+/I+)-iai?iieea (/a?ac ae?aoeeeae?oea?ie ? Ca2+-caeaaeio C-e?iaco) (Aeaa?on A., 1994). Eiia??ii, ui iaoai?ci eiio?ieth ?Ii ? i?ie?oa?aoe??, yeee caionea?oueny a aiaeioae?? i?e aeoeaaoe?? 1-aae?aii?aoeaioi??a, ia? a?aeiioaiiy aei eiai ?aia?aoeaii? ?aaaia?aoe?? ? aaeaioaoe?? aei iioeiaeaeoth/i? ae??  eaoaoieai?i?a. ?c oeueiai aeo?ea?, ui naia ni?aa?aeiioaiiy ie?aieo noaoei?a aae?aii?aoeaioi??a ia AE aecia/a? ni?aa?aeiioaiiy “?aoeaioi?-caeaaeia iioeiaeaeaiiy / ?aoeaioi?-iiina?aaeeiaaia aeoeaaoe?y i?ie?oa?aoe??" oa, a cia/i?e i???, e?ioeaaee ?acoeueoao aca?iiae??. A?aenooi?noue e?eoa???a ioe?iee aaoa?iaaiiino? aiaeioae?th aaeeeeo noaeei (Aonaa N.A. e niaao., 1988) ia aeicaiey? cae?enieoe ia?eoeaiee aiae?c oaiiiaio aaeaioaoe?? aiaeioae?th ai?oe aei ae?? a?ia?aae?aiae?iai??. Caeiaaaeii eiai iiaeia iiynieoe niaoeeo?/i?noth i?aaenoaaieoeoaa -aae?aii?aoeaioi??a ia AE, a?ae yei? caeaaeeoue e?iaoeue i?ioeano aeanaineo?caoe??. Ia a?aei?io a?ae ai?oe, a naeac?ieia?e a?oa??? ? c?aei?e ii?iaeieno?e aai? ca oiia i?aeainoi? a?ia?aae?aiae?iai?? niinoa??aa?oueny aiaieuaaa aeaneaaiaoe?y aiaeioae?th, a eiai aacaeueia iaia?aia a oe?eiio caa??aa? naith cae/aeio no?oeoo?io i?aai?caoe?th. Aeiaie? /anoi iaa?yee? AE i?noyoue aaaaoi i?e?ii?iioeeoicieo aaceeoe, yae?a AE aeeaiei iie?ayi?, a oeeoiieaci? cono??/athoueny iioeiaeaeai? i?oioiiae???. Ca iaoeie aeaieie, a aiaeioae?? naeac?ieiai? a?oa??? i?e oeueiio a?aeaoaa?oueny ca?eueoaiiy aaniethoii? e?eueeino? i?oioiiae??e ? aeaiaio?a CAN. Oae, /eneiaa ? ia?iia u?euei?nue i?oioiiae??e a AE ca?eueoai? ii??aiyii c eiio?ieai (oaae. 3). Caa?oa? ia naaa oaaao 6-e?aoia ca?eueoaiiy /enaeueii?, 7-e?aoia - ia?iii? u?eueiinoae eaiaeueoe?a CAN (oaae. 3). Oaeei /eiii, ii?oieia?/i? aea?aaeaioe ?aia?aoeaii? ?aaaia?aoe?? aiaeioae?th naeac?ieiai? a?oa??? aeoaea ae?aaeai?. A caaei?e ii?iaeieno?e aai? ca?eueoaiiy ?ici???a i?oioiiae??e a?aeaoaa?oueny ia oii? ciaioaiiy ?o e?eueeino? a iaeeieoe? ia?io oeeoiieacie AE ? ia noi?iaiaeaeo?oueny ?inoii iieacieea a?aeiiniiai ia?io i?aaiae (oaae. 2). Aeyaeai? ci?ie iiaeooue aooe ?acoeueoaoii caaeaae? /anoeie i?oioiiae??e ia oii? iaa?yeo aai a?ia?o?io?? ?ioeo. A AE a?aeaoaa?oueny ca?eueoaiiy ia?iii? u?eueiino? ? na?aaeiuei? ieiu? eaiaeueoe?a CAN, ciaioo?oueny /enaeueia u?euei?noue ?? eaiaeueoe?a (oaae. 2). Aeai? ci?ie iathoue i?noea ? i?e iaa?yeo eaiaeueoe?a aiaeiieaciaoe/ii? n?oee. Oaaeeoey 3. Ii?oiiao?e/i? ia?aiao?e i?oioiiae??e ? ca?ienoi? aiaeiieaciaoe/ii? n?oee aiaeioae?th naeac?ieiai? a?oa??? uo??a i?e i?aeaino??e a?ia?aae?aiae?iai?? (I + m) I?aaiaeaI?oioiiae???Ca?ienoa aiaeiieaciaoe/ia n?oeaIa?aiao?Eiio?ieueAeine?aeEiio?ieueAeine?aeIeiua i?io?e?a (10-2 iei2)36,478,1532,963,565,260,494,650,30Ia?iia u?euei?noue (%)2,670,876,071,14*0,690,224,880,92**enaeueia u?euei?noue (10-2 / iei3)3,041,2414,793,55*25,147,41154,1632,99*Oaeoi? oi?ie0,860,040,880,030,350,040,720,02* Oeueo?ano?oeoo?a aiaeioae?th noaeei a oiiaao “ IEA- iiaeae?” iaae?aeaeueoeeiico ai?oe (a?oa??ineea?ico iaieaaa?a?anueeiai oeio) A oiiaao IEA-?ioieneeaoe?? aeine?aeaeaiiy oeueo?ano?oeoo?e aiaeioae?th ii?iae-ienoi? aaie ? /a?aaii? ai?oe i?iaiaeeeiny ye aeey aecia/aiiy oa?aeoa?ieo aeey iaae?aeaeueoeeiico ai?oe iniaeeainoae iioeiaeaeaiiy aiaeioae?th, oae ? aiae?co iioeiaeaeaiiy aiaeioae?th ai?oe a ??cieo iiaeaeyo ?? iaae?aeaeueoeeiico c iaoith aeine?aeaeaiiy ?ie? oeueiai iioeiaeaeaiiy a iaoiaaiac? iaae?aeaeueoeeiico Iaieaaa?aa. Iaie anoaiiaeaii, ui iioeiaeaeaiiy aiaeioae?th a oiiaao IEA-?ioieneeaoe?? ia? i?noea a /a?aai?e ai?o? ? caaei?e ii?iaeieno?e aai? uo??a. Oae, /a?ac aiaeeio i?ney aaaaeaiiy IEA a ai?o? niinoa??aa?oueny aeaneaaiaoe?y ie?aieo aiaeioae?aeueieo ee?oei c ianooiiith ae??aiiaiaieuaaith aeaioaeaoe??th. sse i?aaeei, EE, ye? i?noyoue o?eueoey Aaeaaey-Iaeaaea, ?icoaoiaai? ia aacaeuei?e iaia?ai?, ?iiae? i?aeianai? iaae ?? iiaa?oiath iaa?yeiaith ??aeeiith. I?aeaiaeioae?aeuei? eiioaeoe i?aaenoaaeai? i?inoeie i?aeee?oei-ieie c?aeiaiiyie. A iiiioa?? aeyaeythoueny "oaii?" EE c ae?aaeaiei i?aaenoaa-ieoeoaii iie?nii ? ?eainii. Ca?eueoaii oiaooaaiiy yaea? AE. I?oioiiae??? i?noyoue iao?een na?aaeiuei? aeaeo?iiii? u?eueiino? ? iaeaea ne??cue caa??aathoue naith i?aai?caoe?th. ?aaeoe?y i?oioiiae??iio ia IEA-?ioieneeaoe?th oioiaeia ?aaeoe?? ia a?ia?aae?aiae?iai?th /a?ac 1 aiae. niinoa?aaeaiiy (oaae. 1, 4) ? ia ia? niaoeeo?/ieo iciae. Iieacieee, ye? oa?aeoa?ecothoue ia?aiao?e CAN a AE ai?oe, ia a?ae??ciythoueny a?ae eiio?ieueieo (oaae. 4). Aacaeueia iaia?aia AE caa??aa? cae/aeio no?oeoo?io i?aai?caoe?th. Noaaiaeioae?aeueiee i?ino?? ?icoe?aiee, i?ina?oeaiee, i?noeoue aaeoiey?i? ooai?aiiy. *a?ac aeiao i?ney aaaaeaiiy IEA ae?eyiee aeaioaeiaaii? iiaa?oi? a ai?o? ia aeyaeythoueny, aea AE iathoue iciaee aeeno?io?/ieo ci?i. A oe?eiio, aeey a?eueoino? EE ia oa?aeoa?i? aeano?oeoeai? ci?ie i?aaiae, eeoa a ie?aieo ee?oeiao niinoa??aathoueny i?oioiiae??? c i?ina?oeaiei iao?eenii oa ?aaeoeiaaieie e??noaie. Na?aaeiy ieiua i?oioiiae??e ca?eueoaia ii a?aeiioaiith aei iiia?aaeiueiai oa?i?io niinoa?aaeaiiy, aea iaioa, i?ae a eiio?ie?. Ca?eueoaiiy ieiu? i?oioiiae??e noi?iaiaeaeo?oueny c?inoaiiyi iieacieea ?o a?aeiiniiai ia?io a oeeoiieaci? EE. Ieiua eaiaeueoe?a CAN oaeiae ca?eueoo?oueny a?aeiinii iiia?aaeiueiai oa?ieio niinoa?aaeaiiy. Ia?iia u?euei?noue eaiaeueoe?a i?aeaeuaia a?aeiinii ye eiio?ieth, oae ? iiia?aaeiueiai oa?i?io niinoa-?aaeaiiy (oaae. 4). Aiae?c CAN aeaco? ia a?ia?o?io?th ?? eaiaeueoe?a ?, oaeei /eiii, ia ca?eueoaiiy ooieoe?iiaeueii? aeoeaiino? n?oee. Aacaeueia iaia?aia aiaeioae?th caeaenoa ? ?icoe?aia. A noaaiaeioae?aeueiiio i?inoi?? ae?aaeaiee iaa?ye. Ia a?aei?io a?ae ai?oe, aiaeioae?e aaie /a?ac aiaeeio i?ney aaaaeaiiy IEA iaeaea ne??cue caa??aa? caycie c noaaiaeioae?ai. A EE niinoa??aa?oueny noaiaea?oiee iaa?? i?aa-iae ? aeeth/aiue, ?iiae? iayai? aeeno?io?/i? ci?ie i?oioiiae??e. I?e ni?anoaaeaii? c eiio?ieai, a EE cieaeaia /eneiaa u?euei?noue oa ca?eueoaia na?aaeiy ieiua i?io?e?a i?oioiiae??e. Iieaciee a?aeiiniiai ia?io, yeee i?oioiiae??? caeiathoue a EE, ia a?ae??ciy?oueny a?ae eiio?ieth (oaae. 5). C oeueiai iiaeia c?iaeoe aeniiaie, ui ciaioaiiy e?eueeino? i?oioiiae??e ? iane?aeeii ceeooy ie?aieo i?aaiae. Oaaeeoey 4. Ii?oiiao?e/i? ia?aiao?e i?oioiiae??e ? ca?ienoi? aiaeiieaciaoe/ii? n?oee aiaeioae?th /a?aaii? ai?oe uo??a i?e aino??e IEA-?ioieneeaoe?? (I + m) Ia??iae niinoa?aaeaiiyIa?iia u?euei?noue (%)*enaeueia u?eue-i?noue (10-2/iei3)Ieiua i?io?-e?a (10-2 iei2)Oaeoi? oi?ie i?aaiaeeI?OIOIIAe???, Eiio?ieue2,310,482,380,3693,138,710,900,011 aiaeeia i?ney aino?i? IEA- ?ioieneeaoe??2,82 0,77 4,22 0,66* 48,16 4,53* 0,880,021 aeiaa i?ney aino?i? IEA- ?ioieneeaoe??5,79 1,29* **3,54 0,70 69,72 5,97* **0,837 0,02CAN, Eiio?ieue1,12 0,1633,54 8,067,76 0,600,513 0,021 aiaeeia i?ney aino?i? IEA- ?ioieneeaoe??1,08 0,2333,72 8,926,23 0,420,417 0,021 aeiaa i?ney aino?i? IEA- ?ioieneeaoe??1,93 0,25* **38,48 6,939,30 0,64**0,444 0,01 Oaaeeoey 5. Ii?oiiao?e/i? ia?aiao?e i?oioiiae??e ? ca?ienoi? aiaeiieaciaoe/ii? n?o-ee aiaeioae?th caaeiuei? ii?iaeienoi? aaie uo??a i?e aino??e IEA-?ioieneeaoe?? (I+m) Ia??iae niinoa?aaeaiiyIa?iia u?euei?noue (%)*enaeueia u?eue-i?noue (10-2/iei3)Ieiua i?io?-e?a (10-2 iei2)Oaeoi? oi?ie i?aaiaeeI?OIOIIAe???, Eiio?ieue4,30 0,52 14,68 3,8218,79 2,310,89 0,021 aiaeeia i?ney aino?i? IEA- ?ioieneeaoe??2,61 1,245,36 2,37*43,67 10,44*0,87 0,031 aeiaa i?ney aino?i? IEA- ?ioieneeaoe??5,81 1,243,90 0,81*78,22 6,47* **0,85 0,02CAN, Eiio?ieue1,90 0,43135,88 18,123,06 0,370,45 0,031 aiaeeia i?ney aino?i? IEA- ?ioieneeaoe??1,43 0,5129,57 9,46*8,29 1,00*0,49 0,021 aeiaa i?ney aino?i? IEA- ?ioieneeaoe??2,44 0,4934,47 7,80*8,73 0,52*0,439 0,01 Ca aeaieie e?oa?aoo?e, i?oioiiae??? a aeeaeo ee?oeiao ? iaaecae/aeii ieanoe/ieie ? ?ooiieie i?aaiaeaie, ye? iino?eii ci?iththoue naith oi?io, caeaoi? aei ceeooy iaeia c iaeiith oa ?ic?aeiaiiy (Aeaa?on A., 1994). Ia iaoo aeoieo, oaeiai oeio ci?ie c aieo i?oioiiae??e, ye? ie ? niinoa??aaee a aiaeioae?? /a?ac 1 aiaeeio i?ney aaaaeaiiy aae?aiae?io ? IEA, iiaeooue aooe iciaeith a?aeiinii eaaeiai iioeiaeaeaiiy i?oioiiae??iio ? ee?oeie a oe?eiio. A aeaiee oa?i?i aeine?aeaeaiiy a AE aaie ia? i?noea ca?eueoaiiy i?aaenoaaieoeoaa eaiaeueoe?a aeaaeaiueei? ? ca?ienoi? aiaeiieaciaoe/ii? n?oee. Ciaioaiiy /eneiai? u?eueiino? ? ca?eueoaiiy ieiu? eaiaeueoe?a a?aeaoaa?oueny a CAN ia oii? caa?aaeaiiy ?? a?aeiiniiai ia?io a oeeoiieaci? EE (oaae. 5). Ae?aaeai? aeeno?io?/i? ci?ie a CAN i?e oeueiio a?aenooi?. Aacaeueia iaia?aia aiaeioae?th caa??aa? naith cae/aeio no?oeoo?o. *a?ac aeiao i?ney aaaaeaiiy IEA a  aiaeioae?? aaie na?aaeiy ieiua i?oioiiae??e c?inoa?, a ?o e?euee?noue a iaeeieoe? ia?io oeeoiieacie AE ia ci?ith?oueny. I?e oeueiio ia?iia u?euei?noue i?oioiiae??e a AE, ye ? a iiia?aaei?e no?ie, ia a?ae??ciy?oueny a?ae eiio?ieth (oaae. 5). I?e/eiith oaeeo ci?i c aieo i?oioiiae??iio iiaea aooe iaa?ye iaeieo i?oioiiae??e ? a?ia?o?io?y ?ioeo. Aea neeaaea?oueny a?aaeaiiy, ui aeeno?io?/i? ci?ie i?aaiae ia ? ia?aaaaeath/eie. A AE aaie ae?aaeaia i?aaenoaaieoeoai CAN ? eiiieaeno Aieueaeae?, aeaui /ano?oa cae/aeiiai cono??/athoueny e?ciniie. Aacaeueia iaia?aia aiaeioae?th caa??aa? naith i?aai?caoe?th. *anoi AE eiioaeoothoue c e?ioioeeoaie e?ia?. A oiiaao i?aeaino?i? IEA- ?ioieneeaoe?? a?aeaoaa?oueny noaoioaeueia aeaioaeaoe?y ai?oe. “Ino??aoe?” EE, ye? caa?aaee caycie c noaaiaeioae??i, niinoa??aaeenue ia/anoi, oiio ii?oiiao?e/i? aeine?aeaeaiiy iaie ia i?iaiaeeeenue. Iaaiaee, a i?ina?o? ai?oe iino?eii cono??/aeenue o?aaiaioe iae?ioe/ii ci?iaieo EE. A?oa? aeano?oeoeai? ci?ie oaeiae oa?aeoa?i? aeey aacaeueii? iaia?aie aiaeioae?th, yea aoea i?aeoe/ii a?aenooiy a ie?aieo ae?eyieao ai?oe. A na?aaei?e iaieiioe? ai?oe aoee iayai? ci?ie, ui neeaaeathoue ii?oieia?/io iniiao a?oa??ineea?ico Iaieaaa?aa. A caaei?e ii?iaeieno?e  aai? iaie niinoa??aaeenue ae?eyiee aeaioaeiaaii? iiaa?oi?, ye? iaaeothoue c ino??aoeyie a?aeiinii caa?aaeaiiai aiaeioae?th, ui ia ao?aoea caycie c i?aeeaaeeie aacaeueiith iaia?aiith ? noaaiaeioae??i. EE iathoue ?ic?eae oeeoieaie, iaa?yeeo oeeoiieacio. A ?o yae?ao cia/ii ca?eueoaia caeaeno?noue eiioo??a, ui a aeayeeo aeiaaeeao coiiaeth? naaiaioaoe?th yae?a. A i?aaiaeao ae?aaeai? aeeno?io?/i? ci?ie, ye i?aaeei, ciaioaia caaaeueia e?euee?noue i?aaiae. A ciiao aeaioaeaoe?? niinoa??aathoueny eiioaeoe ee?oei e?ia? c aaiiciith no?ieith. Ia a?aei?io a?ae ai?oe, i?e i?aeaino?iio aieea? IEA aacaeueia iaia?aia aiaeioae?th a aai? a oe?eiio caa?aaeaia ? o?eueee i?noeyie ia?a?ea/anoa, ? aeaui ?icoa?iaaia. Ii??aithth/e noai aiaeioae?th a ai?o? ? aai? a oiiaao aieeao IEA, iaiao?aeii a?aei?oeoe: 1) iioeiaeaeaiiy ? aeano?oeoeai? ci?ie aacaeueii? iaia?aie aiaeioae?th cia/ii a?eueoa ae?aaeai? a ai?o?, 2) i?e aino??e IEA-?ioieneeaoe?? iioeiaeaeaiiy AE ia? i?noea ? a ai?o?, ? a aai?, 3) i?e i?aeaino??e IEA-?ioieneeaoe?? iioeiaeaeaiiy aiaeioae?th iineeth?oueny ? a ai?o?, ? a aai?. Ia i?aenoaa? oeueiai iiaeia aaaaeaoe, ui iioeiaeaeaiiy aiaeioae?th i?yii i?iii?oe?eii ?ioaineaiino? ae?? IEA. sse ? a oiiaao aino?i? a?ia?aae?aiae?iai??, aei iioeiaeaeoth/i? ae?? oaeoi?a a?eueo /ooeeaee aiaeioae?e ai?oe. Ai?no caaaeueiiai eaeueoe?th, aeoeai?noue EO ? - AO a ne?iaaooe? e?ia? uo??a ia ?aii?o noaae?yo aenia?eiaioaeueiiai a?oa??ineea?ico Iaieaaa?aa Eieeaaiiy ai?noo caaaeueiiai eaeueoe?th, ci?ie aeoeaiino? EO ? -AO a ne?iaaooe? uo??a niinoa??aathoueny aaea ia ii/aoeiaiio noai? ?icaeoeo aenia?eiaioaeueiiai a?oa??ineea?ico Iaieaaa?aa (oaae. 6). Oaaeeoey 6. Ai?no caaaeueiiai eaeueoe?th, aeoeai?noue eoaeii? oinoaoace ? - aeooai?eo?ainoa?ace a ne?iaaooe? uo??a i?e ?icaeoeo aenia?eiaioaeueiiai a?oa??ineea?ico Iaieaaa?aa (I + m) Oiiae aenia?eiaioocaaaeueiee eaeue-oe?e (iiieue/e)eoaeia oinoaoaca (oiia.iae.)-aeooai?eo?ain-oa?aca(oiia.iae.) “Aae?aiae?iiaa” iiaeaeueEiio?ieue (n = 10)2,09+0,19899,9+53,62,30+0, 30I?ney a?ia?aae?aiae?iai??, /a?ac1 aiaeeio (n = 10)1,51+0,20*798,5+50,33,00+0,521 aeiao (n = 10)2,38+0,10**790,0+51,88,30+2,18*,**7 ae?a (n = 8)1,58+0,06*,**741,3+76,30,75+0,25*,**“Iiiieiaeaoeaoaoia” iiaeaeueEiio?ieue (n = 10)2,03+0,07562,6+29,51,25+0,23I?ney IEA- ?ioieneeaoe??, /a?ac1 aeiao ( n = 8 )1,80+0,08442,0+40,8*5,50+0,82*7 ae?a ( n = 9)2,45+0,07*,**845,0+54,7*,**1,88+0,48** Ciaioaiiy ai?noo caaaeueiiai eaeueoe?th a ne?iaaooe? uo??a ia? i?noea ye /a?ac aiaeeio, oae ? i?ney 6-aeaiiiai aaaaeaiiy aae?aiae?io. ??no a?aeiioaiiy “aeoeai?noue EO / aeoeai?noue -AO” a oiiaao i?aeaino?i? a?ia?aae?aiae?iai?? (oaae. 6) aeicaiey? i?eioneaoe ca?eueoaiiy e?noeiaeo ?cioa?iaio?a EO a ne?iaaooe? uo??a ?, a?aeiia?aeii, ooieoe?iioaaiiy ia?aoai?iiia. Na?aae aaaaoueio i?e/ei a?iieaeueoe??i?? ia inoaiith ?ieue iiaea a?ae?a?aaaoe iaaei??ia iaaeoiaeaeaiiy eaeueoe?th a ee?oeie noaeeiii? no?iee. *a?ac aeiao i?ney aaaaeaiiy aae?aiae?io ai?no caaaeueiiai eaeueoe?th o ne?iaaooe? a?aeiiaeth?oueny. I?e??no eaeueoe?th a e?ia? (aiane?aeie aeeth/aiiy ?aaoeyoi?ieo iaoai?ci?a) iiaea aooe oaeoi?ii, yeee ni?ey? iai?oaeaiith ? aeaeiiiainaoe?? iaoai?ci?a ?aaoeyoe?? [Na2+]i a AE, AIE noaeei c iaea?eueoei i?aaenoaaieoeoaii 1-aae?aii?aoeaioi??a. Oe?eeii a??ia?aeii, ui ca?eueoaiiy aoiaeo iicaee?oeiiiai Na2+ o EK, AIE a?oa??e i?e ae?? aaii?no?a (eaoaoieai?ie), ye? iia?e?cothoue Na2+ c aioo??oiueiee?oeiieo aeaii, ni?ey? cieaeaiith ooieoe?iiaeueiiai ?aca?ao, ia?aiaaai-oaaeaiith ? a?eueo oaeaeeiio ?icaeoeo aeaeiiiainaoe?? a nenoaiao a?aeiiaeaiiy [Na2+]i a AIE a?oa??e aeanoe/iiai oeio ii??aiyii c AIE a?oa??e ?ioeo oei?a ? aai. I?e i?aeaino?iio aieea? IEA anoaiiaeaii ca?eueoaiiy ai?noo caaaeueiiai eaeueoe?th a ne?iaaooe? (a?aeiinii eiio?ieth ? iiia?aaeiueiai oa?i?io niinoa?aaeaiiy), a oaeiae i?aeaeuaiiy a?aeiioaiiy “aeoeai?noue EO/aeoeai?noue -AO” (oaae. 6) ye iai?yiee aeieac iineeaiiy ooieoe?iioaaiiy IOA. A?aeiii, ui i?e aoeeaeic?, yeee o i?aeaeine?aeieo uo??a ?iaeoeiaaiee aaaeeith oeaieiiith a?iien??th i?ae aieeaii IEA, a?ia?eaeueoe??i?y ? iane?aeeii aeeth/aiiy iaoai?ci?a ei?aeoe?? iaaeeeoea eeneioe o iicaee?oeii?e ??aeei? (O?onea E., 1987). I?aeaeuaiiy ooieoe?iioaaiiy IOA ca aeaieo oiia aeaui iac?icoi?ei. Iiaeeeai, aiii iiaycaii ?c ci?iith /ooeeaino? ia?aoe?ioeeo?a aei neaiaeo oa/aai ii?ooaiiyi aca?iiae?? IOA c ?aoeaioi?aie ia ee?oeiao-i?oaiyo. O oaeiio ?ac? iaa?oue ca?eueoaia [Na2+]e iiaea ni?eeiaoeny ia?aoe?ioeeoaie ye iaaeinoaoiy, a ci?ie a nenoai? eaeueoe??i?? iiaeooue iaoe cia/aiiy a iaoiaaiac? iaae?aeaeueoeeiico ai?oe. Ia cae?i/aiiy aaaeeeai a?aei?oeoe, ui: Oa?aeoa?enoeee ? aeeiai?ea ?icaeoeo iioeiaeaeaiiy aiaeioae?th aecia/athoueny iaoai?ciii iioeiaeaeoth/i? ae?? iaoiaaiiiai oaeoi?a (?aoeaioi?-iiina?aaeeiaaia aai ?aoeaioi?-iacaeaaeia), a oaeiae ?aa?iiaeueieie a?aei?iaie /ooeeaino? aiaeioae?th. Na?aae oaeoi??a, ye? aecia/athoue /ooeea?noue aiaeioae?th, iiaeooue aooe: a) ooieoe?iiaeuei? iiaeeeaino? nenoai, ui iiia?aaeaeothoue iioeiaeaeaiiy, a) iiaeeeaino? nenoai caaacia/aiiy ?aia?aoe?? iioeiaeaeaiiy, ?, o aeiaaeeo ?aoeaioi?-iiina?aaeeiaaiiai iioeiaeaeaiiy, a) iniaeeaino? i?aaenoaaieoeoaa ?aoeaioi??a. Iaoiaaiaoe/ia cia/aiiy a aaiac? iaae?aeaeueoeeiico ai?oe ia? aino?a ianeaia iioeiaeaeaiiy aiaeioae?th. Ca iaaieo oiia (“aae?aiae?iiaa iiaeaeue”) aiii iiaea aenoo-iaoe a ?ie? oi?iiooai?thth/iai (iaae?aeaeueoeeiic) oaeoi?a. Ii?ooaiiy ??aiiaaae a nenoai? ?aaoeyoe?? oiiono AIE aiane?aeie iaaeinoaoiino? aiaeioae?aeueiiai eiiiiiaioo a oiiaao eaoaoieai?iiai? aa?an?? iiaea aooe i?e/eiith a?ia?nei?i/aiiy AIE, aeoeaaoe?? Na2+-iaoai?ci?a iioeiaeaeaiiy ? iae?ico AIE ai?oe. Ianeaia iioeiaeaeaiiy aiaeioae?th ni?e/eith? ii?ooaiiy o?ainaiaeioae?aeueiiai o?ainii?oo ? ?iceaaee o?io?/iiai caaacia/aiiy AIE na?aaeiuei? iaieiiee aaeeeeo a?oa??e. Ca aeaieo oiia eiioaeo no?oeoo?ieo aeaiaio?a iaieaii? noaeeiii? no?iee c aa?aneaieie ?a/iaeiaie (? ee?oeiaie) e?ia?, aeoeaaoe?y i?ioean?a e?oe/iiai oeio (i?ioaie?c, aeanoie?c) c on?ia ?o iiaeeeaeie iane?aeeaie oaeiae iiaeooue ni?e/eithaaoe iioeiaeaeaiiy AIE . I?e IEA-?ioieneeaoe?? a ee?oeiao noaeeiii? no?iee ?icaeaa?oueny oyaeea oeaieiia a?iien?y, yea iiaea aooe i?e/eiith iaa?aeiia?aeiino? iiooaeiino? iaoai?ci?a cieaeaiiy [Ca2+ ]i ?ioaineaiino? eiai iaaeoiaeaeaiiy a oeeoicieue. sse iaoai?ci iaae?aiae?ico a?eueo a??ia?aeia i?yia iioeiaeaeoth/a ae?y IEA aaciina?aaeiuei ia AIE na?aaeiuei? iaieiiee a?oa???. Iioeiaeaeaiiy aiaeioae?th i?e oeueiio iiaea aenooiaoe eeoa a ?ie? oiiae, yea ni?ey? ?icaeoeo iaae?aiae?ico ? iaae?aeaeueoeeiico. Aeyaeai? ci?ie ai?noo caaaeueiiai eaeueoe?th oa aeoeaiino? EO ? - AO a ne?iaaooe? e?ia? i?aeaeine?aeieo uo??a aeicaieythoue i?eioneaoe, ui eieeaaiiy a nenoai? ?aaoeyoe?? [Ca2+]e, ni?yiiaai? ia ?? a?aeiiaeaiiy, a oiiaao ii?ooaiiy ?aaoeyoe?? [Ca2+]i a AIE (i?e iioeiaeaeaii?) iiaeooue aooe oaeoi?ii, yeee aeeth/a? iaoanoaoe/io ? iineeth? aeeno?io?/io eaeueoeeo?eaoe?th no?oeoo? na?aaeiuei? iaieiiee ai?oe i?e ?icaeoeo a?oa??ineea?ico iaieaaa?a?anueeiai oeio. AENIIAEE 1. A?oa??ineea?ic ? aaaeeeaith iaaeeei-nioe?aeueiith i?iaeaiith ye iniiaia i?e/eia ?iaae?aeiino? oa nia?oiino? ianaeaiiy a a?eueoino? e?a?i na?oo. Ieoaiiy ao?ieia?? oa iaoiaaiaco iaae?aeaeueoeeiico a?oa??e aeanoe/iiai ? aeanoe/ii-iyciaiai oeio, cie?aia ?ieue aiaeioae?th a ?icaeoeo oe??? oi?ie a?oa??ineea?ico, caeeoathoueny i?aeoe/ii ia aea/aieie. 2. I?e iiaeaethaaii? iaae?aeaeueoeeiico ai?oe ca aeiiiiiaith a?ia?aae?aiae?iai?? ? IEA- ?ioieneeaoe?? a /a?aai?e ai?o?, naeac?ieia?e a?oa??? ? caaei?e ii?iaeieno?e aai? i?aeaeine?aeieo uo??a aoee aeyaeai? ci?ie oeueo?ano?oeoo?ii? i?aai?caoe?? aiaeioae?th. Inoaii? iaee oa?aeoa? aeeno?io?/ieo, aeano?oeoeaieo aai na?ae/eee i?i iineeaiiy neioaoe/ieo ? ?aia?aoeaieo i?ioean?a. Ni?aa?aeiioaiiy aeano?oeoeaieo ? ?aia?aoeaieo ci?i AE ??cia a noaeeiao /ooeeaeo (ai?oa, naeac?ieiaa a?oa??y) oa no?eeeo (ii?iaeienoa aaia) aei ?icaeoeo iaae?aeaeueoeeiico. 3. *a?ac aiaeeio i?ney aino?i? a?ia?aae?aiae?iai?? ? IEA-?ioieneeaoe?? aeeno?io?/i? ci?ie oeueo?ano?oeoo?e AE ? ia?aaaaeath/eie. A ai?o? aiie a?eueo ae?aaeai?, i?ae a naeac?ieia?e a?oa??? ? caaei?e ii?iaeieno?e aai?. *a?ac aeiao a?ae ii/aoeo ae?? aeei?en-oaieo iaoiaaiieo oaeoi??a a?aei?/a?oueny ?ica?aei?noue ci?i aiaeioae?th: a) aino?a a?ia?aae?aiae?iai?y noi?iaiaeaeo?ony ca?eueoaiiyi nooiaith oyaeeinoe aeano?oeoeaieo ci?i AE ai?oe, ia?aaaaeath/e oaeo a naeac?ieia?e a?oa??? ? caaei?e ii?iaeieno?e aai?, a) aino?a IEA-?ioieneeaoe?y noi?iaiaeaeo?ony ca?eueoaiiyi nooiaith oyaeeinoe aeeno?i-o?/ieo ci?i AE naeac?ieiai? a?oa??? ? caaeiuei? ii?iaeienoi? aaie. I?e oeueiio aeano?oeoeai? ci?ie a AE ii?aeiothoueny c ii?oieia?/ieie aea?aaeaioaie ?aia?aoeaii? ?aaaia?aoe??, ia ui iaeeaaea? a?aeaeoie naaiaioa?ia iaiaeii??aei?noue aiaeioae?th. A?ia?ieanoe/i? ? a?ia?o?io?/i? ci?ie i?aaiae neioaoe/iiai oeio a?eueo i?aaenoaaeai? a caaei?e ii?iaeieno?e aai? ? naeac?ieia?e a?oa???, i?ae a ai?o?. 4.  *ooeea?noue aiaeioae?th a?oa??e c ??ciith e?euee?noth aeaaeaiueeiiyciaeo aeaiaio?a a na?aaei?e iaieiioe? ? aaie uo??a aei iioeiaeaeoth/i? ae?? aae?aiae?io ? IEA ia iaeiaeiaa. ?c ca?eueoaiiyi e?eueeino? AIE a no?ioe? a?oa??? /ooeea?noue ?? aiaeioae?th aei ae?? aino?i? a?ia?aae?aiae?iai?? ? IEA- ?ioieneeaoe?? ciaioo?oueny, a a aai? i?i?iaeueia. 5. A oiiaao i?aeaino?i? (6-aeaiii?) a?ia?aae?aiae?iai?? a ai?o? a?aenooi? ae?aaeai? aeano?oeoeai? ci?ie ?? aiaeioae?th, a oie /an ye a naeac?ieia?e a?oa??? oa caaei?e ii?iaeieno?e aai? aiie ia?inoathoue ? noi?iaiaeaeothoueny aeaioaeaoe??th noaeeiii? no?iee. Ioaea, o aeiaaeeo ae?? ia aiaeioae?e oaeoi?a, iioeiaeaeoth/a ae?y yeiai iiaycaia c aeoeaa-oe??th ?aoeaioi??a (eaoaoieai?ie), iiaeeeaa ci?ia /ooeeaino? aiaeioae?th aei eiai ae??. A?aenooi?noue nooo?aeo ci?i oeueo?ano?oeoo?e aiaeioae?th ai?oe i?ney 6-aeaiiiai aaaaeaiiy aae?aiae?io iiaea aooe i?iyaii aaeaioaoe?? aiaeioae?th aei eiai iioeiaeaeoth/i? ae?? ca ?aooiie cieaeaiiy /ooeeaino? aae?aii?aaeoeaieo no?oeoo?. 6. I?aeaino?a IEA-?ioieneeaoe?y /eieoue ae?aaeaio iioeiaeaeoth/o ae?th ia aiaeioae?e ai?oe ? caaeiuei? ii?iaeienoi? aaie uo??a. Oyaee?noue aeano?oeoeaieo ci?i aiaeioae?th a ai?o? i?e oeueiio ia?aaeuo? oaeo a caaei?e ii?iaeieno?e aai?. Iioeiaeaeaiiy aiaeioae?th iaa?no?aeueieo a?oa??e oa aaie uo??a i?e ae?? ia noaeeiio no?ieo IEA i?yii i?iii?oe?eii e?euee?ni?e oa?aeoa?enoeoe? eiai ae??. I?e oeueiio caa??aa?oueny caei-iii??i?noue, ui oa?aeoa?ia aeey iaeii?aciaiai aino?iai aieeao iaoiaaiiiai oaeoi?a c ?aoeaioi?-iiina?aaeeiaaiei iaoai?ciii iioeiaeaeaiiy: aiaeioae?e a?oa??? c i?i?iaeue-iith e?euee?noth AIE (aeanoe/iiai oeio) a?eueo /ooeeaee aei ae?? iaoiaaiiiai oaeoi?a. 7. Aino?a ianeaia iioeiaeaeaiiy aiaeioae?th ai?oe c yaeuaie noaoioaeueii? aeaioaeaoe?? ? iioeiaeaeaiiyi aacaeueii? iaia?aie a ?aiiueiio ia??iae? ?icaeoeo aenia?eiaioaeueiiai a?oa??ineea?ico iaieaaa?a?anueeiai oeio (ia?oa aeiaa aino?i? a?ia?aae?aiae?iai??) ? aaaeeeaith eaieith iaoiaaiaco "aae?aiae?iiaiai" iaae?aiae?ica oa iaae?aeaeueoe?iica. 8. Aeayea  iaoiaaiaoe/ia cia/aiiy a ?aiiueiio ia??iae? ?icaeoeo a?oa??ineea?ico iaieaaa?a?anueeiai oeio iiaeooue iaoe aeyaeai? ci?ie ai?noo caaaeueiiai eaeueoe?th a ne?iaaooe? e?ia? oa cnoae a nenoai? eiai ?aaoeyoe??. Nienie iioae?eiaaieo i?aoeue ca oaiith aeena?oaoe?? 1. Aieaeiaeia A.A., Noa/aiei E.A., Auoeue TH.A. Oeueo?ano?oeoo?iua eciaiaiey yiaeioa-eey iaaeno?aeueiuo ninoaeia i?e ino?ie e iiaeino?ie aeia?aae?aiaeeiaiee // Aeiiia?ae? IAI Oe?a?ie. - 1996.- ? 9.- N.147-152. 2. Aieaeiaeia A.A., Auoeue TH.A. Nii?yaeaiea noeioe-nae?aoeey a yiaeioaeee e ?aaoeyoeey oiiona aeaaeeeo ninoaeenouo iuooe // Iae?iaoii?aeueiua iaoaieciu iaoieiae/aneeo i?ioeannia: Na. iao/. o?. - O.: Oaeo, 1996. - 130n. 3. Aieaeia?ia I.A., Aeoeue TH.A., Noa/aiei E.I. Aino?a ianeaia iioeiaeaeaiiy aiaeioae?th ye eaiea iaoiaaiaco a?oa??ineea?ico iaieaaa?a?anueeiai oeio // Aoeiaeinueeee iaaee/iee a?niee.- 1998.- O.2, ?2.- N. 41-47. 4. Aieaeia?ia I.A. Ci?ie ai?noo caaaeueiiai eaeueoe?th, aeoeaiino? eoaeii? oinoaoace ? -aeooai?eo?ainoa?ace a ne?iaaooe? e?ia? uo??a i?e aenia?eiaioaeueiiio a?oa??i-neea?ic? Iaieaaa?aa // O?c?ieia?/iee aeo?iae. - 1998.- O.44, ? 1-2. - N.71-76. 5. Aieaeia?ia I.A. Oeueo?ano?oeoo?i? ci?ie aiaeioae?th a ii/aoeiaiio ia??iae? ?icaeoeo aenia?eiaioaeueiiai a?oa??ineea?ico iaieaaa?a?anueeiai oeio // Oe?. Iaoeiai-iaaee/iee iieiae?aeiee aeo?iae.- 1998.- ? 2-3.- N. 12-15. 6. Aeoeue TH.A., Aieaeia?ia I.A., Aeoaeei I.I. Aeine?aeaeaiiy iaoai?ci?a ??cii? ?acenoaio-iino? a?oa??e oa aai aei iioeiaeaeaiiy //Iaoa??aee ?? eiia?ano iaoio?c?ieia?a Oe?a?ie i?ena. 100-??//th a?ae aeiy ia?iaeaeaiiy aeaae. I.I. Ne?ioei?ia.- O?c?ieia?/iee aeo?iae. - 1996. - O. 42, ? 3-4.- N.57. 7. Aieaeiaeia A.A. Eaoaoieaieiiaia iia?aaeaeaiea yiaeioaeey a?oa?ee e aai // I?iaeaiu yenia?eiaioaeueiie eaai/ii-na?aea/iie iaaeinoaoi/iinoe. Oacenu aeieeaaeia iaaeaocianeiai neiiiceoia, iina. 30-eaoeth IEEOe EIE.- 8-10 ai?aey 1992.-N. 67-68. 8. Aeoeue TH.A., Aeoaeei I.I., Aieaeia?ia I.A. Ee?oeii? oa iieaeoey?i? iaoai?cie ?icaeoeo iaoieia?/ieo i?ioean?a. Oace aeiiia?aeae Ieaioio iaoeiai-iaaee/iiai oiaa?enoaa iaoio?c?ieia?a Oe?a?ie (23-25 aa?aniy 1993 ?., i. Euea?a).- N.8.  Aieaeia?ia I. A. Iniaeeaino? iioeiaeaeaiiy aiaeioae?th i?e ?icaeoeo aenia?eiaioaeueiiai a?oa??ineea?ico iaieaaa?a?anueeiai oeio. - ?oeiien. Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa iaaee/ieo iaoe ca niaoe?aeuei?noth 14.03.04 - iaoieia?/ia o?c?ieia?y. - ?inoeooo o?c?ieia?? ?i. I.I. Aiaiiieueoey IAI Oe?a?ie, Ee?a, 1999. Aeena?oaoe?th i?enay/aii aea/aiith iniaeeainoae iioeiaeaeaiiy aiaeioae?th ? eaeueoe??i??, ye? oa?aeoa?i? aeey a?oa??ineea?ico Iaieaaa?aa. Ca aeiiiiiaith iaoiaeo o?aini?n?eii? aeaeo?iiii? i?e?ineii?? aeine?aeaeaii ci?ie no?oeoo?e aiaeioae?th noaeei, /ooeeaeo ? no?eeeo aei ?icaeoeo iaae?aeaeueoeeiico. Ca aeiiiiiaith a?io?i?/ieo iaoiae?a aeine?aeaeaii ai?no caaaeueiiai eaeueoe?th, aeoeai?noue eoaeii? oinoaoace ? -aeooai?eo?ain-oa?ace a ne?iaaooe? e?ia? uo??a a ?aiiueiio ia??iae? ?icaeoeo aenia?eiaioaeueiiai iaae?aeaeueoeeiico ai?oe. Anoaiiaeaii, ui aino?a ianeaia iioeiaeaeaiiy aiaeioae?th a ai?o? a?aeaoaa?oueny ai?iaeiaae ia?oeo aiaeei ? aeiae i?ney a?ia?aae?aiae?iai??. Ioaea, aeno?aiaeueia iioeiaeaeaiiy aiaeioae?th ye ia?aaeoiiao ia?anei?i/aiiy ? iae?ico AIE ai?oe i?e eaoaoieai?iia?e aa?an?? ne?ae aaaaeaoe aaaeeeaei iaoai?ciii a iaoiaaiac? "aae?aiae?iiaiai" iaae?aeaeueoe?iico ai?oe. I?e IEA-?ioieneeaoe?? ye iaoai?ci iaae?a-iae?ico a?eueo a??ia?aeia i?yia ae?y IEA aaciina?aaeiuei ia AIE na?aaeiuei? iaieiiee ai?oe. Iioeiaeaeaiiy aiaeioae?th i?e oeueiio iiaea aenooiaoe eeoa a ?ie? oiiae, yea ni?ey? ?icaeoeo iaae?aeaeueoeeiico. Io?eiai? aeai? i?i eieeaaiiy ai?noo caaaeueiiai eaeueoe?th ? i?aeaeuaiiy aeoeaiino? e?noeiaeo ?cioa?iaio?a EO a ne?iaaooe? aeine?aeieo iaoethe?a ne?ae ioe?ithaaoe ye aeac?aeo ia oa, ui ci?ie a nenoai? eaeueoe?ai?? iiaeooue iaoe iaoiaaiaoe/ia cia/aiiy a ?aiiueiio ia??iae? a?oa??ineea?ico Iaieaaa?aa. Eeth/ia? neiaa: a?ia?aae?aiae?iai?y, iiiieiaeaoeaoaoia ?ioieneeaoe?y, a?oa??ineea?ic Iaieaaa?aa, iaae?aeaeueoeeiic, aino?a ianeaia iioeiaeaeaiiy aiaeioae?th, ia?anei?i/aiiy AIE, caaaeueiee eaeueoe?e, ne?iaaoeiaa eoaeia oinoaoaca, - aeooai?e-o?ainoa?aca. Goldobina O.V. The endothelial injury characteristic features during the development of experimental Monckebergs medial calcific sclerosis. - Manuscript. Thesis for a candidates degree by speciality 14.03.04 - pathological physiology.- The O.O. Bogomolets Institute of Physiology, National Academy of Sciences of Ukraine, Kyiv, 1999. The dissertation is devoted to investigation of the endothelial injury characteristics and calciemia in modelling Monckebergs arteriosclerosis using either single dose or multidose administration adrenaline and monoiodacetatic acid (MIA). The transmission electron microscopy, the changes in endothelial vascular ultrastructure which can be either sensitive or innsensitive to medial calcific sclerosis are explored. It was established that acute massive injury of endothelium had been present in aorta on the first day after single-action adrenalinemia. Therefore, acute massive endothelial injury is a precondition of the hyperconstriction and necrosis of aortal SMCs in the condition of hyperadrenalinemia and may be considered as an important mechanism in pathogenesis of “adrenalinal” aorta mediacalcinosis. Under the conditions of MIA- intoxication, direct MIA effects on SMCs may be considered as a major mechanism of medianecrosis in the aorta but the endothelial injury may play a contributive role in that. By means of the biochemical methods, the common calcium level, alkaline phosphotase and -glutamiltranspheras activities in blood serum in the early period of the aortal medial calcific sclerosis are explored. Probably, the changes in calciemia regulatory system may play pathogenetical role in early period Monckebergs arteriosclerosogenesis. Key words: hyperadrenalinemia, monoiodacetatic acid intoxication, Monckebergs medial calcific sclerosis, acute massive endothelial injury, hyperconstriction of SMCs, common calcium level, alkaline phosphotase and -glutamiltranspherase activities. Aieaeiaeia A.A. Iniaaiiinoe iia?aaeaeaiey yiaeioaeey i?e ?acaeoee yenia?eiaioaeueiiai a?oa?eineea?ica iaieaaa?aianeiai oeia. - ?oeiienue. Aeenna?oaoeey ia nieneaiea o/aiie noaiaie eaiaeeaeaoa iaaeeoeeineeo iaoe ii niaoeeaeueiinoe 14.03.04 - iaoieiae/aneay oeceieiaey. - Einoeooo oeceieiaee ei. I.I. Aiaiiieueoea IAI Oe?aeiu, Eeaa, 1999. Aeenna?oaoeey iinayuaia ii?aaeaeaieth a ?aiiai ia?eiaea ?acaeoey yenia?eiaioaeueiiai iaaeeaeaeueoeeiica ai?ou oa?aeoa?iuo aeey a?oa?eineea?ica iaieaaa?aianeiai oeia iniaaiiinoae iia?aaeaeaiey yiaeioaeey, a oaeaea eciaiaiee niaea?aeaiey iauaai eaeueoeey, aeoeaiinoe uaei/iie oinoaoacu e - aeooaieeo?ainoa?acu a nuai?ioea e?iae. Iaaeeaeaeueoeeiic ai?ou o e?un iiaeaee?iaaee n iiiiuueth aaaaeaiey aae?aiaeeia aeae?ioei?eaea e iiiieiaeaoeaoaoa (IEA). Enoiaey ec oiai, /oi ia?aua ii?oiei-ae/aneea i?iyaeaiey iaaeeaeaeueoeeiica auyaeythony a ai?oa /a?ac nooee iinea aaaaeaiey niioaaonoaothuaai i?aia?aoa, iieacaoaee, oa?aeoa?ecothuea ninoiyiea yiaeioaeey, eco/aeenue /a?ac /an e nooee iinea iaeiie?aoiiai aaaaeaiey aauanoaa (“ino?ia” aicaeaenoaea) e ia 7-a nooee iinea 6-aeiaaiiai aaaaeaiey (“iiaeino?ia” aicaeaenoaea). N iiiiuueth iaoiaea o?ainienneiiiie yeaeo?iiiie iee?ineiiee enneaaeiaaia oeueo?ano?oeoo?a yiaeioaeey a?thoiie ai?ou, naeacaii/iie a?oa?ee e caaeiae iieie aaiu. N oeaeueth eiee/anoaaiiie ioeaiee i?aaiaee yiaeioaeeaeueiuo eeaoie (ieoioiiae?ee, ca?ienoay yiaeiieaciaoe/aneay naoue) aue i?eiaiai iaoiae ii?oiiao?e/aneiai aiaeeca. Eniieueciaaee ii?aaeaeaiea neaaeothueo noa?aiiao?e-/aneeo ia?aiao?ia: aeiey iauaia i?aaiaeeu a oeeoiieacia (%), /enei i?aaiaee a aaeeieoea iauaia (1/iei3), ieiuaaeue (iei2 ) e oaeoi? oi?iu i?aaiaee. Eae iieacaee enneaaeiaaiey, a ?aiiai ia?eiaea ?acaeoey yenia?eiaioaeueiiai iaaeeaeaeueoeeiica ai?ou eciaiaiey yiaeioaeey iaeiioeiiu e i?aaenoaaeaiu aeano?oeoeaiuie e ?aia?aoeaiuie eciaiaieyie oeueo?ano?oeoo? YE. N?aaieoaeueiue aiaeec eciaiaiee yiaeioaeey ninoaeia iieacae, /oi oyaeanoue iia?aaeaeaiey yiaeioaeey e au?aaeaiiinoue ii?oieiae/aneeo yeaeaaeaioia ?aia?aoeaiie ?aaaia?aoeee ?acee/iu a ninoaeao, /onoaeoaeueiuo e onoie/eauo e ?acaeoeth iaaeeaeaeueoeeiica. Oa?aoa?iie iniaaiiinoueth yiaeioaeey ai?ou yaeyaony aai aunieay /oanoaeoaeueiinoue e iia?aaeaeathuaio aeaenoaeth ino?ie aeia?aae?aiaeeiaiee e IEA-eioieneeaoeee. Onoaiiaeaii, /oi ianneaiia iia?aaeaeaiea yiaeioaeey eiaao ianoi a ai?oa a ia?aua nooee iinea ino?ie aeia?aae?aiaeeiaiee. A oneiaeyo eaoaoieaieiiaie aa?annee ino?ia ianneaiia iia?aaeaeaiea yiaeioaeey a?oa?ee n iaeneiaeueiui i?aaenoaaeoaeuenoaii 1-aae?aii?aoeaioi?ia (yeanoe/aneiai oeia - ai?oa) e neaaeothuaa ca iei ia?ooaiea ?aaoeyoeee oiiona AIE aneaaenoaea iaaeinoaoi/iinoe aa yiaeioaeeaeueiiai eiiiiiaioa iiaeao auoue i?e/eiie aeia?nie?auaiey AIE, aeoeaaoeee eaeueoeeaauo iaoaieciia iia?aaeaeaiey e iae?ioe/aneeo eciaiaiee AIE n?aaeiae iaiei/ee. A iienaiiie neooaoeee au?aaeaiiua ?anno?ienoaa o?ioe/aneiai iaania/aiey AIE n?aaeiae iaiei/ee e?oiiuo a?oa?ee ?acaeaathony aneaaenoaea eae niacia v.vasorum, oae e ia?ooaiee o?ainyiaeioaeeaeueiiai o?ainii?oa aauanoa. Aeoeaiue eiioaeo no?oeoo?iuo yeaiaioia iaieaiiie ninoaeenoie noaiee n aa?anneaiuie aauanoaaie (e eeaoeaie) e?iae, aeoeaaoeey i?ioeannia eeoe/aneiai oeia (i?ioaieec, yeanoieec) ni anaie eo aiciiaeiuie iineaaenoaeyie oaeaea iiaoo auoue i?e/eiie iia?aaeaeaiey AIE. Iiyoiio ino?ia ianneaiia iia?aaeaeaiea yiaeioaeey, eae aiciiaeioth i?e/eio iae?ioe/aneeo eciaiaiee AIE, neaaeoao n/eoaoue aaaeiui iaoaieciii iaoiaaiaca "aae?aiaeeiiaiai" iaaeeaeaeueoeeiica ai?ou. N iiiiuueth aeioeie/aneeo iaoiaeia eco/aii niaea?aeaiea iauaai eaeueoeey, aeoeaiinoue uaei/iie oinoaoacu e -aeooaieeo?ainoa?acu a nuai?ioea e?iae iiaeiiuoiuo e?un. Iieo/aiiua a ?aaioa iai?yiua naeaeaoaeuenoaa iiauoaiey aeoeaiinoe einoiuo ecioa?iaioia UO a nuai?ioea iiaeiiuoiuo e?un neaaeoao ioeaieaaoue eae oeacaiea ia oi, /oi eciaiaiey a nenoaia ?aaoeyoeee eaeueoeeaiee iiaoo eiaoue iaoiaaiaoe/aneia cia/aiea a ?aiiai ia?eiaea a?oa?eineea?ica Iaieaaa?aa. Eeth/aaua neiaa: aeia?aae?aiaeeiaiey, iiiieiaeaoeaoaoiay eioieneeaoeey, a?oa?eineea?ic Iaieaaa?aa, iaaeeaeaeueoeeiic, ino?ia ianneaiia iia?aaeaeaiea yiaeioaeey, ia?anie?auaiea AIE, iauee eaeueoeee, nuai?ioi/iay uaei/iay oinoaoaca, -aeooaieeo?ainoa?aca.

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