?inoeooo aiaeie?eiieia?? oa iai?io
?a/iaei

?i. A.I.Eii?na?aiea AII Oe?a?ie

OAeEOAeE
[616.379 — 008.64 + 616.8] :

612.015.31.63

EINOTHE Ieaia Ieaoii?aia

Ci?ie eaeueoe??aiai aiiainoaco

a ia?aiaeo ee?oeiao i?e ?icaeoeo

ae?aaaoe/ieo iae?iiao?e

14.01.14 aiaeie?eiieia?y

Aaoi?aoa?ao

aeena?oaoe?? ia caeiaoooy iaoeiaiai
nooiaiy

aeieoi?a iaaee/ieo iaoe

Ee?a, 1999

Aeena?oaoe?y ? ?oeiienii

?iaioa aeeiiaia a ?inoeooo? aiaeie?eiieia?? oa iai?io ?a/iaei
?i.A.I.Eii?na?aiea AII Oe?a?ie oa ?inoeooo? o?c?ieia?? ?i.
I.I.Aiaiiieueoey IAI Oe?a?ie

Iaoeia? eiinoeueoaioe:

aeieoi? iaaee/ieo iaoe, i?ioani?, aeaaeai?e IAI oa AII Oe?a?ie ?o?iia
Aiae??e Naiaiiae/, caa?aeoaa/ ae?aaaoieiae/iiai a?aeae?eo ?inoeoooo
aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A.I.Eii?na?aiea AII Oe?a?ie

aeieoi? iaaee/ieo iaoe, i?ioani?, eaeaaeai?e IAI oa AII Oe?a?ie
O?ieuee?n Aieiaeeie? Aai?ai?iiae/, caa?aeoaa/ eaai?aoi??? o?c?ieia??
?inoeoooo aa?iioieia?? AII Oe?a?ie

Io?oe?ei? iiiiaioe:

aeieoi? iaaee/ieo iaoe, i?ioani? Aiaeia? Iao?i Ieeieaeiae/, caa?aeoaa/
eaoaae?e aiaeie?eiieia?? Iaoe?iiaeueiiai iaaee/iiai oi?aa?neoaoo ?i.
I.I.Aiaiiieueoey

aeieoi? a?ieia?/ieo iaoe E?oioaiei Ieaenaiae? sseiae/, caa?aeoaa/
eaai?aoi??? iieaeoey?ii? aaiaoeee ?inoeoooo aa?iioieia?? AII Oe?a?ie

aeieoi? iaaee/ieo iaoe, i?ioani?, /eai-ei?aniiiaeaio IAI Oe?a?ie Naaa/
Aaaeei Oaaei?iae/, caa?aeoaay a?aeae?eo o?c?ieia?? e?iaiia?ao ?inoeoooo
o?c?ieia?? ?i. I.I.Aiaiiieueoey IAI Oe?a?ie.

I?ia?aeia onoaiiaa:

?aaii-O?aie?anueea Aea?aeaaia iaaee/ia aeaaeai?y, eaoaae?a eao

Caoeno a?aeaoaeaoueny 15 /a?aiy… 1999 ?ieo i 13e aiaeei? ia
can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae 26.558.01 c aiaeie?eiieia??
a ?inoeooo? aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A.I.Eii?na?aiea AII
Oe?a?ie (254114, Ee?a-114, aoe.Aeoai?iaenueea 69).

C aeena?oaoe??th iiaeia iciaeiieoenue a a?ae?ioaoe? ?inoeoooo
aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A.I.Eii?na?aiea AII Oe?a?ie
(Ee?a-114, aoe Aeoai?iaenueea 69).

Aaoi?aoa?ao ?ic?neaii 12 o?aaiy 1999?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee

aeieoi? a?ieia?/ieo iaoe
E.I.Eaeeinueea

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue oaie. Ai?aeai?ieia?/i? aeine?aeaeaiiy, i?iaaaeai? a ?yae?
e?a?i na?oo ca inoaii? aeanyoe??//y, iieacaee, ui oeoe?iaee ae?aaao ia
nueiaiaeiyoi?e aeaiue noaiiaeoue iaae?ei-nioe?aeueio i?iaeaio, a ii
nai?e /eiiino? caeia? o?ao? i?noea i?ney na?oeaai-noaeeiieo oa
iieieia?/ieo caoai?thaaiue (Aaeaaiee?i, 1994, ?o?iia oa Ne?iaiinueea,
1998). A?aei?/aii c?inoaiiy i?ioeaioiiai ni?aa?aeiioaiiy oai?eo ye ia
?inoe?icaeaaeiee (?COeAe), oae ? ia ?inoe?iiacaeaaeiee (?IOeAe)
oeoe?iaee ae?aaao na?aae ianaeaiiy ?a?iiaenueeeo e?a?i, ui na?aae ?ioeo
oaeoi??a iiynith?oueny i?aeaeuaiiyi a iiioeyoe?yo iin??a aai?a,
ni?eyoeeaeo aei ?icaeoeo oeoe?iaiai ae?aaaoo, ca?eueoaiiyi aeyaeaiiy
oeoe?iaiai ae?aaaoo (ca ?aooiie iie?auaiiy ae?aaiinoeee ia?aaaaeii
?IOeAe) oa nooo?aiai ca?eueoaiiy o?eaaeino? aeeooy oai?eo ia oeoe?iaee
ae?aaao, ca ?aooiie iie?auaiiy eiai e?eoaaiiy (?o?iia, 1989, Noa?eiaa,
1992). A?aeiia?aeii, /a?ac aeuacacia/ai? oaeoi?e iaeia c oeaio?aeueieo
i?noeue o aeine?aeaeaiiyo iaoai?ci?a ?icaeoeo oeoe?iaiai ae?aaaoo oa
iiaeeeainoae oa?iaeieia?/iiai aieeao ia eiai ia?aa?a iin?ee ieoaiiy
aea/aiiy iaoai?ci?a ?icaeoeo oneeaaeiaiue oeoe?iaiai ae?aaaoo, ye?
aaaaoueia aeine?aeieeaie ?icayaeathoueny ye aaciina?aaei?e i?iya oai?iae
(?o?iia, 1989; Levy, 1994; Stevens et al., 1995). Iniaeeao oaaao na?aae
ieo i?eaa?oathoue ae?aaaoe/i? iae?iiao??, ye aiane?aeie aaaeeino?
ee?i?/iiai ia?aa?ao, oae ? aiane?aeie neeaaeiino? i?aeai?o aaeaeaaoii?
iaoiaaiaoe/ii? oa?ai??. ?iciianthaeaeaiiy oe??? iaoieia?? c?inoa? a?ae
7% na?aae oai?eo c o?eaae?noth oeoe?iaiai ae?aaaoo aei 1 ?ieo aei 50% o
oai?eo c o?eaae?noth oeoe?iaiai ae?aaaoo a?eueoa 25 ?ie?a (Pirart, 1978,
Vinic et al., 1992). A oie aea /an ia an? eaiee iaoiaaiaco ae?aaaoe/ieo
iae?iiao?e ia nueiaiaeiyoi?e aeaiue iiai?noth ?ice?eoi. Ca aeaieie
?o?iiaa oa Ne?iaiinueei? (1998), aeenoaeueia naini?ii-iioi?ia iae?iiao?y
? iaei??th c iaea?eueo ?iciianthaeaeaieo oi?i oe??? iaoieia??. Iniiai?
?ene, ui aeanoea? oe?e oi?i? ae?aaaoe/ii? iae?iiao?? — ii?ooaiiy
oaeaeeino? i?iaaaeaiiy ?iioeuen?a oa caoaeeeaino? ia?aiaeo ee?oei,
?icaeoie aieueiaiai neiae?iio oa a?ia?anoac?e, ia?anoac?e, ae?canoac?e,
ui ci?iththoueny ao?aoith aieueiai? /ooeeaino? — aac noii?ao aeacothoue
ia oa, ui iaei??th c iaoiaaiaoe/ieo eaiie ?o ?icaeoeo ? ii?ooaiiy
eiiiiai aiiainoaco, a oiio /ene? eiiiiai eaeueoe??aiai aiiainoaco,
ine?eueee ?iie Na2+ iaiao?aei? ee?oeiai aeey ?aaoeyoe?? ?o ?inoo,
i?ioean?a aeeoa?aioe?thaaiiy oa cae?eniaiiy nai?o ooieoe?e, i?e/iio
niaeo? ?o ae?? iaaecae/aeii oe?ieee (Eistyuk, 1992, 1998; O?ieuee?n,
1993).

?aeeii? aeoiee noiniaii oa?aeoa?o ci?i eaeueoe??aiai aiiainoaco o
a?aeiia?aeieo ia?aiaeo no?oeoo?ao i?e ae?aaaoe/i?e aeenoaeuei?e
naini?iiiioi?i?e iae?iiao?? ia nueiaiaei? iaia. Oae, a?aeiia?aeii aei
aeaieo Green et al. (1990) i?e

ae?aaaoe/ieo iae?iiao?yo a?aei?/a?oueny ia?aaaaeii cieaeaiiy ??aith Na2+
a ia?aiaeo ee?oeiao, a oie /an ye Hall et al. (1994) a?aei?/aee
i?aeneeaiiy eaeueoe??aeo no?oi?a a naini?ieo iae?iiao i?e oeoe?iaiio
ae?aaao?. Iac`yniaaii, iane?eueee ci?ie aioo??oiueiee?oeiiiai
eaeueoe??aiai aiiainoaco iiaeooue ni?aiaaeaoe c oaeeie i?e ?ioeo
iineeaaeiaiiyo oeoe?iaiai ae?aaaoo, a?ae??ciyoenue a caeaaeiino? a?ae
oeio ae?aaaoo oa ci?ithaaoenue a caeaaeiino? a?ae i?iaeiaaeaiino?
caoai?thaaiiy. Iaeiaio aea/aiei caeeoa?oueny ?aii?e ia??iae ?icaeoeo
ae?aaaoe/ieo naini?ieo iie?iae?iiao?e; /e iiaeooue ia oeeo aoaiao ci?ie
aioo??oiueiee?oeiiiai eaeueoe??aiai aiiainoaco iaoe nooo?aa cia/aiiy —
caeeoa?oueny iac?icoi?eei. Aeine?aeaeaiiy oe??? eaiee iaoiaaiaoe/ieo
iaoai?ci?a ?icaeoeo ae?aaaoe/ii? iae?iiao?? ia? aeiiiiiaoe o ?ica`ycaii?
ieoaiiy i?i iiaeeeaino? oa?iaeieia?/iiai aieeao ia ia?aa?a ae?aaaoe/ieo
iae?iiao?e oa ieoaiiy i?aeai?o iaea?eueo aaeaeaaoiiai ?aaeeio
?inoe?iioa?ai?? aeey iiia?aaeaeaiiy i?ia?anoaaiiy ae?aaaoe/ii?
iae?iiao??.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie oa oaiaie. ?iaioa aeeiioaaeanue a
?aieao ieaiiaeo oai a?aeae?eo ee?i?/ii? ae?aaaoieia?? ?inoeoooo
aiaeie?eiieia?? oa iai?io ?a/iaei ?i. Eii?na?aiea AII Oe?a?ie oa
I?aeia?iaeiiai oeaio?o iieaeoey?ii? o?c?ieia?? ?inoeoooo o?c?ieia?? ?i.
Aiaiiieueoey IAI Oe?a?ie.

Iaoa ?iaioe aea/aiiy noaio aioo??oiueiee?oeiiiai eaeueoe??aiai
aiiainoaco a iae?iiao ??cieo eaiie naini?ieo oeyo?a ia ?aii?o aoaiao
?icaeoeo ae?aaaoe/ii? aeenoaeueii? naini?iiiioi?ii? iie?iae?iiao?? ia
oaa?eiao c ??cieie iiaeaeyie oeoe?iaiai ae?aaaoo (no?aioicioioeeiiaiai
?inoe?icaeaaeiiai oa aaiaoe/ii-aeaoa?i?iiaaiiai ?inoe?iiacaeaaeiiai) oa
c`ynoaaiiy iiaeeeainoae oa?iaeieia?/ii? ei?aeoe?? eiai ii?ooaiue.

Iniiai? caaea/? aeine?aeaeaiiy:

1. Aeine?aeeoe, /e ?icaeaa?oueny naini?iiiioi?ia iie?iae?iiao?y o oaa?ei
c ??cieie oeiaie oeoe?iaiai ae?aaaoo, ? yeui oae, oi ye? aiia ia?
oa?aeoa?enoeee.

2. Anoaiiaeoe, /e aeieeathoue ci?ie aioo??oiueiee?oeiiiai eaeueoe??aiai
aiiainoaco ia ?aii?o aoaiao ?icaeoeo ae?aaaoe/ii? naini?iiiioi?ii?
iie?iae?iiao??, ? yeui oae, oi ye? naini?i? oeyoe ia?aaaaeii
a?aaeathoueny.

3. Aeine?aeeoe, ia yeeo naia eaieao naini?ieo oeyo?a oe? ci?ie
?icaeaathoueny ? /e ? a?aei?ie o ci?iao eaeueoe??aiai aiiainoaco ia
??cieo eaieao ??cieo naini?ieo nenoai.

4. Anoaiiaeoe, ca ?aooiie o?aaeaiiy yeeo naia aioo??oiueiee?oeiieo
eaeueoe?e-?aaoethth/eo no?oeoo? (nenoai iaia?aiiiai o?ainii?oo,
aiaei-ieaciaoe/iiai ?aoeeoeoio, i?oioiiae??e) aeieeathoue ii?ooaiiy
eaeueoe??aiai aiiainoaco.

5. Anoaiiaeoe, /e iiaeooue oae? ii?ooaiiy aooe i?e/eiith nooo?aeo ci?i
iae?iiaeueieo ooieoe?e oa ?icaeoeo ?yaeo ee?i?/ieo neiioii?a i?e
ae?aaaoe/i?e naini?iiiioi?i?e iae?iiao??.

6. Aeine?aeeoe, /e caeaaeeoue oa?aeoa? ci?ie aioo??oiueiee?oeiiiai
eaeueoe??aiai aiiainoaco a?ae oeio oeoe?iaiai ae?aaaoo.

7. Aeine?aeeoe, /e iiaeia aieeaaoe ia ci?ie oeo /e ?ioeo eaiie
aioo?oiueiee?oeiiiai eaeueoe??aiai aiiainoaco i?e iayaiino? ae?aaaoe/ii?
iie?iae?iiao?? a aenia?eiaioaeueieo oiiaao oa?iaeieia?/ieie caniaaie oa
/e aoaea oa?aeoa? ?o aieeao aiaeia?/iee oaeiio o caei?iaeo oaa?ei.

8. Aeine?aeeoe, yeee aieea ia ei?aeoe?th aioo??oiueiee?oeiiiai
eaeueoe??aiai aiiainoaco ia ?aii?o aoaiao ?icaeoeo ae?aaaoe/ii?
naini?iiiioi?ii? iie?iae?iiao?? iathoue eaeueoe??a? aeieaoi?e a?oie
ae?a?ae?ii??eaeei?a i?e ?o o?eaaeiio aaaaeaii? i?aeaeine?aeieie
oaa?eiaie.

9. Aeine?aeeoe, yea cia/aiiy aeey ei?aeoe?? ii?ooaiue eaeueoe??aiai
aiiainoaco iiaea iaoe ?inoe?iioa?ai?y oa ?? ?aaeei.

Iaoeiaa cia/ei?noue ? iiaecia io?eiaieo ?acoeueoao?a. Anoaiiaeaii, ui o
i?aeaeine?aeieo oaa?ei a i?ioean? ?icaeoeo ae?aaaoe/ii? iaoieia??
ca?eueoo?oueny i?ioeaioia ni?aa?aeiioaiiy iia?eueii-i?iaiaey/eo
(iioe?oeaioeaieo) aoa?aioieo ia?aiaeo aieieii a?aeiinii
oaeaeei-i?iaiaey/eo, ui na?ae/eoue i?i nooo?aa o?aaeaiiy ia?oeo aaea ia
?aii?o aoaiao ?icaeoeo ae?aaaoe/ieo iie?iae?iiao?e.

Aia?oa anoaiiaeaii, ui ia ?aii?o aoaiao ?icaeoeo ae?aaaoe/ieo iae?iiao?e
aeieea? ii?ooaiiy aioo??oiueiee?oeiiiai eaeueoe??aiai aiiainoaco ia an?o
eaieao oeyo?a iioe?oeaioeaieo /ooeeaino?, yea iia`ycaia ia?aaaaeii c
iaoaaie?/ieie ci?iaie, oa?aeoa?ieie aeey oeoe?iaiai ae?aaaoo, ? iiaea
aooe iaei??th c i?ia?aeieo i?e/ei oi?ioaaiiy ?yaeo neiioii?a
ae?aaaoe/ii? naini?iiiioi?ii? iie?iae?iiao??.

Iieacaii, ui ii?ooaiiy aioo??oiueiee?oeiiiai eaeueoe??aiai aiiainoaco a
iioe?oeaioeaieo iae?iiao ia? ni?euei? ?ene ia an?o eaieao
iioe?oeaioeaieo ia?aiaeo oeyo?a, a oiio /ene? ? ia aeueo ?o eaieao, ye?
iieyaathoue o ci?i? e?iaoeee eaeueoe??aeo neaiae?a o a?aeiia?aeue ia
caoaeaeaiiy ?, a?aeiia?aeii, ci?i? i?ioean?a aeae?eaiiy iae?iiaae?aoi??a
oa i?ioean?a neiaioe/ii? ia?aaea/?. A oie aea /an, ii?ooaiiy, ui
aeieeathoue ia eiaei?e eaioe? aoa?aioieo oeyo?a, iathoue nai?
iniaeeaino?.

Anoaiiaeaii, ui a i?ioean? ?icaeoeo ae?aaaoe/ii? iie?iae?iiao??
a?aaeathoueny ye iioe?oeaioeai?, oae ? i?ii??ioeaioeai? ia?aeii?
naini?i? iae?iie, i?e/iio oa?aeoa? ci?ie eaeueoe??aeo neaiae?a a ieo
a?ae??ciy?oueny; a iioe?oeaioeaieo iae?iiao oe? ci?ie a?eueo nooo?a?.
A oie aea /an, ia ?aii?o aoaiao

?icaeoeo ae?aaaoe/ii? naini?ii? iie?iae?iiao?? ia aecia/aii nooo?aeo
ci?i aioo??oiueiee?oeiiiai eaeueoe??aiai aiiainoaco a i?ii??ioeaioeaieo
iae?iiao aeueo iiceiaeo no?oeoo?.

Aia?oa i?iaaaeaii ii??aiyeueiee aiae?c oa?aeoa?o iie?iae?iiaoe/ieo ci?i
o oaa?ei c ?inoe?icaeaaeiei oa ?inoe?iiacaeaaeiei oeiaie oeoe?iaiai
ae?aaaoo ia ?aii?o aoaiao ?icaeoeo ae?aaaoe/ii? naini?iiiioi?ii?
iie?iae?iiao??. Anoaiiaeaii, ui aecia/ai? ii?ooaiiy
aioo??oiueiee?oeiiiai eaeueoe??aiai aiiainoaco o iioe?oeaioeaieo
iae?iiao iathoue iiae?aiee ia?aa?a, oi/a ? ia ni?aiaaeathoue iiai?noth.
O oaa?ei c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii a i?ii??ioeaioeaieo
iae?iiao a?aeiia?aeieo ci?i aioo??oiueiee?oeiiiai eaeueoe??aiai
aiiainoaco ia aeyaeaii.

Aia?oa iieacaii ?ioaa?aoeaiee oa?aeoa? ii?ooaiue ooieoe??
aioo??oiueiee?oeiieo Na2+-aeoioethth/eo no?oeoo?o i?e oeoe?iaiio
ae?aaao?: ii?ooaiiy eaeueoe?e-aeoioethth/i? ooieoe?? i?oioiiae??e a
ia?aeiieo naini?ieo iae?iiao oa eaeueoe?e-aeoioethth/i? ooieoe??
eaeueoe?e-/ooeeaiai aiaeiieacia-oe/iiai ?aoeeoeoia o aoi?eiieo naini?ieo
iae?iiao.

Oa?iaeieia?/iee aiae?c anoaiiaea nooo?ao ci?io aieeao
ae?a?ae?ii??eaeei?a ia eaeueoe??ao neaiae?caoe?th iioe?oeaioeaieo
iae?ii?a ia an?o eaieao naini?ieo oeyo?a o oaa?ei c ae?aaaoe/iith
iie?iae?iiao??th, ui na?ae/eoue i?i iayai?noue e?euee?nieo aai ye?nieo
ci?i o ooieoe?? ??cieo oei?a iioaioe?ae-caeaaeieo eaeueoe??aeo eaiae?a
i?e oeoe?iaiio ae?aaao? oa iiaea aooe iaei??th c i?e/ei oi?ioaaiiy
iaoieia?/ieo ?eoi?a ia ??ai? aeueo iiceiaeo no?oeoo?.

Iieacaii, ui o?eaaeee eo?n e?eoaaiiy i?iiae?i?iii o oaa?ei c
no?aioicioioeei-?iaeoeiaaiei ae?aaaoii aieeaa? ia ci?ie eaeueoe??ai?
neaiae?caoe?? a iioe?oeaioeaieo iae?iiao iaiaeiicia/ii. Aea i?iaiaeeoe
iiaio aiaeia?th i?ae aieeaii i?iiae?i?io ia ci?ie eaeueoe??aiai
aiiainoaco o oaa?ei oa o oai?eo ia oeoe?iaee ae?aaao iaiao?aeii c
iaa?aaei?noth.

Aiae?c aieeao ?inoe?iioa?ai?? ia ii?ooaiiy aioo??oiueiee?oeiiiai
eaeueoe??aiai aiiainoaco ia ?aii?o aoaiao ?icaeoeo ae?aaaoe/ii?
naini?iiiioi?ii? iie?iae?iiao?? iieacaa, ui o oaa?ei c
no?aioicioioeei-?iaeoeiaaiei ae?aaaoii iiaii? eiiiainaoe?? aecia/aieo
ci?i ca aeiiiiiaith iiii?inoe?iioa?ai?? aeinyaoe ia iiaeia, ia?aaaaeii
/a?ac o?aaeaiiy eaeueoe?e-aeoioethth/eo aioo??oiueiee?oeiieo no?oeoo?.
Aeey aeinyaiaiiy iaeneiaeueii iiaeeeai? ei?aeoe?? ci?i
aioo??oiueiee?oeiiiai eaeueoe??aiai aiiainoaco iaiao?aeia no?eea
eiiiainaoe?y oeoe?iaiai ae?aaaoo.

I?aeoe/ia cia/aiiy ?iaioe. Iaea?aeai? ?acoeueoaoe ?ice?eaathoue iiaeeea?
iaoiaaiaoe/i? eaiee ?icaeoeo ae?aaaoe/ii? naini?iiiioi?ii?
iie?iae?iiao?? oa

?yaeo ?? neiioii?a, oaeeo ye aieueiaee neiae?ii, a?ia?aeaac?y, a?ia?- oa
a?iianoac?y, oiaoi neiioii?a, ui ia?aaaaeathoue ia ?aii?o aoaiao
?icaeoeo iie?iae?iiao??. Oea aeicaiey? ooeaoe iia? caniae
oa?iaeieia?/iiai aieeao ia ?icaeoie oeueiai oneeaaeiaiiy oeoe?iaai
ae?aaaoo, a?aoiaoth/e oei oeoe?iaiai ae?aaaoo.

Aeine?aeaeaiiy aieeao ?inoe?iioa?ai?? ia ci?ie aioo??oiueiee?oeiiiai
eaeueoe??aiai aiiainoaco aeaei aeiaeaoeiaa iaa?oiooaaiiy iaiao?aeiino?
aeinyaiaiiy o oai?eo ia oeoe?iaee ae?aaao no?eei? eiiiainaoe??
caoai?thaaiiy aeey noaa?e?caoe?? ia?aa?ao ae?aaaoe/ii? iie?iae?iiao??;
aeey aeinyaiaiiy oaei? eiiiainaoe?? aaaeaii aaeeaaiiy ?aaeeio
?ioaineaii? ?inoe?iioa?ai??. A oie aea /an, iaea?aeai? ?acoeueoaoe
na?ae/aoue, ui aeey eiiiainaoe?? aeieeath/eo a i?ioean? ?icaeoeo
ae?aaaoe/ii? iie?iae?iiao?? ii?ooaiue eaeueoe??aiai aiiainoaco aaeeaaiiy
eeoa ?inoe?iioa?ai?? iaaeinoaoi?.

Aeine?aeaeaiiy oa?ai?? ae?a?ae?ii??eaeeiaie na?ae/aoue i?i
iaiaeiicia/i?noue ?acoeueoao?a ?o aeei?enoaiiy i?e o?aaeaii? ia?aiaeo
ee?oei i?e ??cieo oeiao oeoe?iaiai ae?aaaoo, a oaeiae i?i
iaaeinoaoi?noue aeei?enoaiiy i?iiae?i?io ye iiiioa?ai?? i?e ae?aaaoe/i?e
iie?iae?iiao??.

Io?eiai? ?acoeueoaoe aeacothoue ia iaiao?aei?noue oe?eani?yiiaaiiai
iiooeo eiiieaenieo cania?a aeey aoaeoeaii? noaa?e?caoe?? ia?aa?ao
ae?aaaoe/ii? iie?iae?iiao?? c o?aooaaiiyi iaiao?aeiino? eiiiainaoe??
aecia/aieo ii?ooaiue eaeueoe??aiai aiiainoaco, a oaeiae aeeth/aiiy aei
oa?aiaaoe/iiai eiiieaeno i?aia?ao?a, ui aieeaathoue ia ii?ooaiiy
ooieoe?? aioo??oiueiee?oeiieo eaeueoe?e-aeoioethth/eo nooeoo?.

Iniaenoee aianie caeiaoaa/a. Caeiaoaa/ a?aa aaciina?aaeith o/anoue o
ieaioaaii? oa i?iaaaeaii? an?o aenia?eiaioaeueieo aeine?aeaeaiue,
?acoeueoaoe yeeo aeeeaaeai? a aeena?oaoe??. Iei i?iaaaeaiee iiaiee
aiae?c io?eiaieo ?acoeueoao?a, ?o noaoenoe/ia ia?iaea, ni?anoaaeaiiy c
?nioth/eie aeaieie iaoeiai? e?oa?aoo?e oa ioe?iea i?aeoe/iiai cia/aiiy
i?iaaaeaieo aeine?aeaeaiue.

Caeiaoaa/ai i?iaaaeaii ni?anoaaeaiiy io?eiaieo ?acoeueoao?a c ee?i?/ieie
i?iyaaie ae?aaaoe/ii? aeenoaeueii? naini?iiiioi?ii? iie?iae?iiao??, a
oaeiae c?iaeaii ii??aiyeueiee aiae?c aeaieo aenia?eiaioaeueieo oa
ee?i?/ieo aeine?aeaeaiue ?inoe?iioa?ai?? oa e?eoaaiiy
ae?a?ae?ii??eaeeiaie ia ia?aa?a ae?aaaoe/ii? naini?ii? iie?iae?iiao??.

Ai?iaaoe?y oa ioae?eaoe??. Ca iaoa??aeaie aeena?oaoe?? iioae?eiaaii 24
iaoeia? ?iaioe, c ieo 5 naiino?eieo, a iaoeiaeo aeo?iaeao, iaoeiaeo
ca??eao oa oacao eiioa?aioe?e, aea ?iaioe aoei i?eeiyoi ia i?aaao
noaooae.

Iaoa??aee aeena?oaoe?? aeiiia?aeaeenue oa iaaiai?thaaeenue ia
eiioa?aioe?yo oa neiiic?oiao:

1. I eiia?an Oaaea?aoe?? ?a?iiaenueeeo o?c?ieia?/ieo oiaa?enoa,
Iaano??oo, I?aea?eaiaee, na?iaiue 1995?.

2. IV I?aeia?iaeiee n?iiic?oi c ae?aaaoe/ieo iae?iiao?e, Ainoa?aeai,
I?aea?eaiaee, eeiaiue 1997?.

3. VI I?aeia?iaeia oeiea “Eaeoe?? c ae?aaaoo o oaa?ei”, Eiiaiaaaai,
Aeai?y, eeiaiue 1997?.

4. I?aeia?iaeia oeiea “Eaeueoe??aa neaiae?caoe?y”, Ee?a, /a?aaiue 1997?.

5. ??/ia eiioa?aioe?y A?eoainueeiai o?c?ieia?/iiai oiaa?enoaa,
E?aa?ioeue, Aaeeea A?eoai?y, ea?oaiue 1998?.

6. I c`?cae Oe?a?inueeiai oiaa?enoaa iae?iiaoe, Ee?a, aeiaoaiue 1998?.

7. Can?aeaiiy a?aeae?eo ae?aaaoieia?? ?inoeoooo aiaeie?eiieia?? oa
iai?io ?a/iaei oa Nai?ia? iieaeoey?ii? o?c?ieia?? ?inoeoooo o?c?ieia??,
a?oaeaiue 1998 — n?/aiue 1999? .

8. Ai?iaaoe?eia eii?n?y niaoe?ae?ciaaii? a/aii? ?aaee ?inoeoooo
aiaeie?eiieia?? oa iai?io ?a/iaei ?i. Eii?na?aiea AII Oe?a?ie, a?oaeaiue
1998?.

Ianya oa no?oeoo?a aeena?oaoe??. Aeena?oaoe?y aeeeaaeaia ia 264
noi??ieao iaoeiiieniiai oaenoo, neeaaea?oueny c anooio, iaeyaeo
e?oa?aoo?e, ?icae?e?a “Iaoa??aee oa iaoiaee aeine?aeaeaiiy”, “Aeani?
?acoeueoaoe” (7 i?ae?icae?e?a), “Iaaiai?aiiy ?acoeueoao?a”, aeniiae?a,
iieacieea e?oa?aoo?e (392 aeaea?ae). ?iaioa ?ethno?iaaia 18 iaethieaie,
14 ae?aa?aiaie, 13 oaaeeoeyie.

IA’?EOE OA IAOIAeE AeINE?AeAEAIUe

Iiaeae? oeoe?iaiai ae?aaaoo c?o/i? oei, ui aiie, oi/a ? ia iiai?noth
a?aeiia?aeathoue oaeei o ethaeae, caaaeii caa??aathoue iniiai?
iaoiaaiaoe/i? iaoai?cie ?icaeoeo caoai?thaaiiy oa eiai iineeaaeiaiue,
aeicaieythoue aea/eoe aeaoaeueii ?yae iaoai?ci?a, ye? ia iiaeia
aeine?aeeoe o ee?i?/ieo oiiaao (iai?eeeaae, ?yae iaoaaie?/ieo aniaeo?a,
aioo??oiueiee?oeii? ci?ie), a?eueo aeaoaeueii iaa?oiooaaoe aeei?enoaiiy
e?eoaaeueieo i?aia?ao?a, a oaeiae aa?eo?eoaaoe caaaeuei? caeeth/aiiy oa
oai???, ui io?eiothoueny i?e aeine?aeieoeueeeo ?iaioao o ee?i?oe?. Oiio
ie ae??oeee caa?iooenue aei i?iaaaeaiiy ?yaeo aeine?aeaeaiue ia
aenia?eiaioaeueieo iiaeaeyo oeoe?iaiai ae?aaaoo. Na?aae iineeaaeiaiue,
ui iiaeooue ?icaeiooenue o oaeeo oaa?ei, aeyaeai? ?aoeiiiao?y,
iao?iiao?y, aeaiaioe aia?iiao??, a oaeiae ae?aaaoe/i? iae?iiao??.

Oa?aeoa?enoeea oaa?ei, ui aeei?enoiaoaaeenue o aeine?aeao. Aeoiaey/e
c aeaieo e?oa?aoo?e, iaie i?e i?iaaaeaii? aeine?aeaeaiue aoei
aeei?enoaii aeae?eueea a?oi oaa?ei: ieo? e?i?? AC57Black6 oa uo?e e?i??
W?star, ia yeeo iiaeaethaaany no?aioicioioeeiiaee (NOC) ae?aaao
(iiaeaeue ?COeAe), ? ieo? e?i?? db/db c aaiaoe/ii-aeaoa?i?iiaaiei
ae?aaaoii (iiaeaeue ?IOeAe). Aeei?enoiaoaaeenue eiio?ieuei? a?oie oaa?ei
oa oaa?eie /a?ac 3 aei? oa 1-2 oeaei? i?ney aaaaeaiiy NOC (aeey
ia?aa??ee aaciina?aaeiueiai aieeao no?aioicioioeeio oa aieeao ci?i
??aith ae?eai?? ia ia?aia? ee?oeie a i?ioean? ?icaeoeo oeoe?iaiai
ae?aaaoo) oa oaa?eie c ?icaeiooei oeoe?iaei ae?aaaoii. Iayai?noue
oeoe?iaiai ae?aaaoo aecia/aeanue ??aiai ae?eai?? (aeua 14 iiieue/e aeey
NOC-?iaeoeiaaiiai oa aeua 10 iiieue/e aeey aaiaoe/ii-aeaoa?i?iiaaiiai
ae?aaaoo).

O ieoae e?i?? AC57Black6 ae?aaao iiaeoethaaany oeyoii i`yoe
iine?aeiaieo aioo??oiueia?thoeiieo ?i`?eoe?e NOC (noia?ia aeica
40ia/ea). NOC aaiaeeany oaa?eiai o a?oe? 7-8 oeaei?a; a aenia?eiaio
oaa?eie a?aeenue o a?oe? 12 oeaei?a. Uo?ai e?i?? Wistar NOC aaiaeeany
iaeii?aciai, c ?ic?aooieo 60 ia/ea. Aiae?c aeic i?aia?aoo oa ?aaeei eiai
aaaaeaiiy i?iaaaeaii ca aeaieie e?oa?aoo?e (Aa?aiia,1983, Caiaaeithe oa
ni?aaao., 1983, Aaeaaiee?i, 1984, Shafrir, 1990).

Ieo? e?i?? db/db c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii a?aeenue o
aeine?ae o oiio ae a?oe?, ui ? ieo? e?i?? AC57Black6.

Ieoai e?i?? AC57Black6 c NOC-?iaeoeiaaiei ae?aaaoii i?iaiaeeeanue
?inoe?iioa?ai?y ia i?ioyc? oeaeiy; e?eoaaiiy i?iiae?i?iii — ia i?ioyc? 3
oeaei?a.

I?ae /an i?iaaaeaiiy aeine?aeaeaiue oaa?eie oo?eioaaeenue ia
noaiaea?oi?e ae??o?, a iaeiaeiaeo oiiaao; aeey ni?ia a?aeenue ia?aaaaeii
naioe? (ye a?eueo /ooeea? aei no?aioicioioeeio) iaeiaeiaeo a?eiaeo a?oi.

Aei??thaaiiy ae?eai?? i?iaiaeeeenue aeae?eueea ?ac?a, a?aeiia?aeii o
eiio?ieueieo oaa?ei, /a?ac iaaiee /an i?ney aaaaeaiiy no?aioicioioeeio
oa aaciina?aaeiuei ia?aae acyooyi o ni?iao aeey anoaiiaeaiiy iayaiino?
?icaeiooiai oeoe?iaiai ae?aaaoo. O eiaeio na??th ni?ia oaa?eie
i?aeae?aeenue c aeecueeeie iieacieeaie ae?eai??. E?ia aeey aei??thaaiiy
caae?aeanue c oainoiai? aaie ye o ieoae, oae ? o uo??a. Aei??thaaiiy
i?iaiaeeeenue ca aeiiiiiaith aetheiiao?o Reflolux S.F. (c ia?aeaeueiei
eiio?ieai ii eieuei?ia?e oeae?).

Aecia/aiiy oaeaeeino? i?iaaaeaiiy ia?aiaiai ?iioeueno ii aoa?aioiei
ia?aiaei aieieiai oa aecia/aiiy iioaioe?aeo aei?caeueii? iiaa?oi?
nieiiiai iiceo (IAeI). Oaa?eie ia?eioecoaaeenue iaiaooaeii 60 ia/ea
aioo??oiueia?thoeiii). A?aeaaaeaiiy iioaioe?ae?a aei?caeueii? iiaa?oi?
(IAeI) nieiiiai iiceo i?iaiaeeeinue ia ??ai? L4-L5 ca aeiiiiiaith
a?aea?aeiiai aeaeo?iaeo. ?aoa?aioiee aeaeo?iae aeiethaaany o i`yce
nieie. Aeey iineeaiiy

iioaioe?ae?a aeei?enoiaoaaany i?aeneethaa/ ci?iiiai no?oio c aenieei
eiao?oe??ioii i?aeneeaiiy. Neaei?/iee ia?a ?ici?uoaaany ia
iiae?acithth/? aeaeo?iaee c i?aeaeaeo?iaeioth a?aenoaiith 1,5 ii.

Aecia/aiiy ??aith aioo??oiueiee?oeiiiai eaeueoe?th [Ca2+]i oa
eaeueoe??aeo neaiae?a a ia?aeiieo naini?ieo iae?iiao oa i?oioiiae??yo.
Aeey aecia/aiiy [Ca2+]i a iae?iiao aei?caeueieo aaiae??a nieiiiai iiceo
(AeANI) aeei?enoiaoaaeenue eiio?ieuei? ieo? e?i?? AC57Black6, ieo? oe???
ae eei?? /a?ac 3 aei? oa 1-2 oeaeaeiy i?ney aaaaeaiiy NOC oa c
?icaeiooei NOC-?iaeoeiaaiei ae?aaaoii, a oaeiae ieo? e?i?? db/db. AeANI
aeae?eyeenue c nieiiiiceiaiai eaiaeo (iiia?aeiai-e?eaeiai? iaeano?) o
noa?eeueieo oiiaao i?ae i?e?ineiiii. Aeae?eai? aaiae?? i?ioiaeeee
noaiaea?oio aiceiaoe/io ia?iaeo o ioieiaeaeaiiio ?ic/ei? O??iaea, ui
i?noea 1 ia/ie i?iiace (oei XIV, “Sigma”, NOA). I?ney cae?i/aiiy ia?iaee
ee?oeie aeae?eyeenue iaoai?/ii oa ia?aiineeenue ia iie?eaieo oeaeueoeyo
a aenia?eiaioaeueio eaia?o c aiiioiaaiei ?iaa?oiaaiei oethi?enoeaioiei
i?e?ineiiii. Aeey i?iaaaeaiiy aenia?eiaioo aaeeaaeanue nenoaia oaeaeei?
aie?eaoe?? ?ic/ei?a i?ae eiio?ieai AII (Usachev et al., 1993).

Aioo??oiueiee?oeiiee a?eueiee [Ca2+]i aecia/aany noaiaea?oiith
iaoiaeeeith ca aeiiiiiaith eaeueoe?e-/ooeeaiai oethi?enoeaioiiai
aa?aieea indo-1AM (o ?ic/ei oaeiae aeiaeaaaany aeaoa?aaio ieth?ii?e
F-127). Oethi?enoeaioe?y ?ic/eio aeeeeeaeanue oeueo?a-o?ieaoiaei
aei?ii?ithaaiiyi ia aeiaaeei? oaee? 360 ii, iiaea?eia ai?n?y
aei??thaaeanue ia aeiaaeeiao oaeeue 395-415 oa 490-510 ii; [Ca2+]i
aecia/aany ii ci?i? ni?aa?aeiioaiiy oethi?enoeaioieo neaiae?a a
iaaaaeaieo ae?aiaciiao c aeei?enoaiiyi oi?ioee Grynkiewich et al.
(1985). Ia?iaea neaiae?a i?iaiaeeeanue aeiiiiiaith i?ia?aiii-aia?aoiiai
eiiieaeno Win Tyda (Battelle, Germany).

?io?ai?oioiiae??aeueiee ??aaiue Na2+ aeine?aeaeoaaee ca e?euee?noth
eiai aea?eueiaiiy c i?aaiae o oeeoicieue. Aeey oeueiai o
ciai?oiueiee?oeiiee ?ic/ei O??iaea aeiaeaaaany i?oioiiae??aeueiee
i?ioiiioi? ea?aii?eoe?ai?ae m-oei?oai?ea?ae?acii (NNN?).

Neeaae ?ic/eio O??iaea (iI): NaCl-140, KCl-4, CaCl2-2, MgCl2 -2,
EGTA-0,5, HEPES/NaOH-10, pH-7,3. Aeey aeaiiey?ecaoe?? ee?oeiii? iaia?aie
aeei?enoiaoaaeenue ?ic/eie O??iaea, a yeeo aoei i?iaaaeaia aea?iiey?ia
cai?uaiiy 50 iI iao??th eae??i.

Aecia/aiiy [Ca2+]i oa eaeueoe??aeo neaiae?a o iae?iiao nieiiiai iiceo
(aoi?eii? naini?i? iae?iie), iae?iiao iici/eo, oaeaiono oa ei?e aaeeeeo
i?aeoeue. Aeey aecia/aiiy [Ca2+]i aeei?enoiaoaaeenue eiio?ieuei? uo?e
e?i?? Wistar, a oaeiae oaa?eie oe??? ae e?i?? /a?ac 1-2 oeaei? i?ney
aaaaeaiiy NOC oa c ?icaeiooei

NOC-?iaeoeiaaiei oeoe?iaei ae?aaaoii. Aeine?aeaeoaaeenue ia ?cieueiaai?
iae?iie, a c??ce a?aeiia?aeieo aeueo no?oeoo? iiceo. Oaa?eie
i?eneieyeenue iaiaooaeii, 60 ia/ea.

Aeey aeaioiaeaiiy neaen?a (c??c?a) nieiiiai iiceo ae??caeanue eiai
/anoeia ia ??ai? L5-L7. O oieiaeiiio ?ic/ei? E?aana ia oinoaoiiio aooa??
i?ae i?e?ineiiii i?iaiaeeeinue i?aia?oaaiiy oa i/euaiiy iio??aiiai
a?aeae?eo. ?ic/ei aacia?a?aii ieneaaioaaany ea?aiaaiii (95% I2 + 5%NI2).
C??ce ia??caeenue ia a?a?ioii? (oiaueia c??co 300 iei). C??ce iici/eo,
oaeaiono oa ei?e aaeeeeo i?aeoeue aeaioiaeyeenue ca aiaeia?/iith
iaoiaeeeith c aeei?enoaiiyi ?ic/eio ??iaa?a.

I?ney ?ieoaaoe?? oa caaa?aeaiiy aeaioiaeai? c??ce ?acii c i?aaeiaoiei
oeaeueoeai ia?aiineeenue o aenia?eiaioaeueio eaia?o, ciiioiaaio ia
i?aenoaaoe? oethi?enoeaioiiai i?e?ineiio. Ie?ai? ee?oeie o c??cao
a?coae?coaaeenue ca aeiiiiiaith ae?noaioe?eiiai aiaei-?ia?n?eiiai
ia`?eoeao. Aecia/aiiy [Ca2+]i i?iaiaeeeinue ca aeiiiiiaith
oethi?enoeaioiiai aa?aieea oo?a-2AM. Aeey caoaeaeaiiy aa?aieea ee?oeie
ina?oethaaeenue ia oaeeyo 360 ± 5 ii oa 390 ± 5 ii. Aeae? na?oei
caoaeaeaiiy cae?aeinue ia aeiaaeei? oaee? 530 ± 10 ii. ??ai? [Ca2+]i
ia/enethaaeenue oaeiae ca oi?ioeith Grynkiewich et al. (1985).

Neeaae ?ic/eio ??iaa?a ia oinoaoiiio aooa?? (a iI): NaCl-125,0
KCl-5,0 CaCl2-2,5, MgSO4-1,5, NaHCO3-25,0, KH2PO4-1,5, aetheica-10,0, ?I
7,4.

Aecia/aiiy [Ca2+]i a aiaeiieaciaoe/iiio ?aoeeoeoi?. Aeey aecia/aiiy
[Ca2+]? aeei?enoiaoaaeenue a?oie oaa?ei, aiaeia?/i? oaeei i?e
aeaioiaeaii? iiceiaeo c??c?a. Aeaioiaeaiiy c??c?a oa ?o caaa?aeaiiy
oethi?enoeaioiei aa?aieeii i?iaiaeeeinue ca aiaeia?/iith iaoiaeeeith.
Aeey aei??thaaiiy aea?eueiaiiy [Ca2+]i c aioo??oiueiee?oeiieo aeaii
c??ce ia?aiineeenue o ?ic/ei aac CaCl2, a yeee aeiaeaaaeinue 30 iI
eioa?io; ai?no MgCl2 i?aeaeuoaaany aei 3iI c ianooiiith aeiaea/ath aei
?ic/eio 1iI EGTA.

Aecia/aiiy aeoeaiino? Na2+ AOOace o aaiae?yo aei?caeueieo ei??ioe?a,
aiiiaaiaoao nieiiiai oa aieiaiiai iiceo. Aeey aecia/aiiy Na2+-AOOacii?
aeoeaiino? aeei?enoiaoaaeenue aiaeia?/i? a?oie ieoae. Oaa?eie
i?eneieyeenue ao??ii. Aea?aiee nieiiee oa aieiaiee iicie aiiiaai?coaaee
ia oieiaeo o oeeyiiio aiiiaai?caoi?? (?ic/ei aeey aiiiaai?caoe?? i?noea
250 iI naoa?ice, 20 iI o??n-Nl, i?e ?I 7,5). Ia?aae aecia/aiiyi
aeoeaiino? aiceio aei aiiiaaiaoo aeiaeaaaee ?ic/ei ae?a?oieneio
(e?ioeaaa eiioeaio?aoe?y 0,2%). Caaaeueio AOOacio aeoeai?noue aecia/aee
o ?ic/ei? ianooiiiai neeaaeo: o??n-Cl (pH 7,5)-50iI, MgCl2-4iI, CaCl2
-1iI, AOO-4iI, ioaa?i-0,15iI. Aeoeai?noue Mg2+-AOOace aecia/aee o oeo
aea oiiaao, aea a i?enooiino? AGOA. Ai?no iai?aai?/iiai oinoi?o
aecia/aee ca Fiske & Subbarow (1955), ai?no a?eeo — ca Lowry et al.
(1951). Aeoeai?noue ae?aaeaee o i?e?iiieyo aea?eueiaiiai
iai?aai?/iiai oinoi?o ia 1 ia a?eeo a aiaeeio. Aeoeai?noue Na2+-AOOace
ae?aoiaoaaee ii ??cieoe? i?ae caaaeueiith AOOaciith aeoeai?noth oa
aeoeai?noth Mg2+-AOOace.

?ACOEUeOAOE AeINE?AeAEAIUe OA ?O IAAIAI?AIIss

Oaeaee?noue i?iaaaeaiiy aoa?aioieo ia?aiaeo ?iioeuen?a o
i?aeaeine?aeieo oaa?ei. Iaei??th c iniiaieo iciae ?icaeoeo aeenoaeueieo
ae?aaaoe/ieo naini?iiiioi?ieo iie?iae?iiao?e ? cieaeaiiy oaeaeeino?
i?iaaaeaiiy ia?aiaiai ?iioeueno ii aeenoaeueieo aoa?aioieo ia?aiaeo
aieieiao. Iayai?noue oeueiai oaiiiaio i?aeoaa?aeaeo?oueny ye a ee?i?oe?
(aea a?i ? ae?aaiinoe/iith iciaeith ?icaeoeo iie?iae?iiao?e), oae ? a
oiiaao aenia?eiaioo ca aeiiiiiaith aeaeo?ii?ia?ao?/ieo iaoiaeee oa
o?aineooaiiiai iiae?aciaiiy ia?aiaeo aieieii. A iaoeo aeine?aeaeaiiyo
aeey aecia/aiiy oaeaeeino? i?iaaaeaiiy ia?aiaiai ?iioeueno ii ia?aiaei
aoa?aioiei aieieiai aeei?enoiaoaaeanue noaiaea?oia iaoiaeeea
a?aeaaaeaiiy iioaioe?aeo aei?caeueii? iiaa?oi? nieiiiai iiceo (IAeI),
aeeeeeaiiai iiae?aciaiiyi n?aeie/iiai ia?ao. I?e iiiiiiey?i?e
?a?no?aoe?? eiiiiiaioiee neeaae IAeI ia? o?eooaciee oa?aeoa?: oaeaeeee
eiiiiiaio, ui oa?aeoa?eco? iaeneiaeueio oaeaee?noue i?ioiaeaeaiiy
aoa?aioii? oaee? (ia?aiaiai caoaeaeaiiy) i?ae a?aea?aeiei aeaeo?iaeii,
oiaoi ii/aoie oa o?eaae?noue ?? ia?aa?ao; iia?eueiee iaaaoeaiee
eiiiiiaio (N1 oa N2 oaee?), yeee a?aeia?aaea? noia?io iinoneiaioe/io
aeoeai?noue iae?ii?a a?eueo iiaa?oiaaeo oa??a aei?caeueiiai ?iao
nieiiiai iiceo (CINI) aoa?aioaie c ??ciith oaeaee?noth i?iaaaeaiiy ?,
ia?aoo?, caeeth/iee aeaeo?iiiceoeaiee eiiiiiaio (? oaeey), ui aeieea?
i?e ia?ai?uaii? caoaeaeaiiy o a?eueo aeeaie? oa?e nieiiiai iiceo
(iae.1).

Ia iae.2 i?aaenoaaeai? cia/aiiy ??aiae ae?eai?? (A), oaeaeeino?
i?iaaaeaiiy aoa?aioii? oaee? (A) oa o?eaaeino? iiceoeaii? oace oe???
oaee? (A) o ieoae eiio?ieueii? a?oie (E), ieoae c NOC-?iaeoeiaaiei
ae?aaaoii (NOC) oa ieoae c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii.

Io?eiai? ?acoeueoaoe na?ae/aoue i?i oa, ui ca a?aenooi?no? ci?i o
iaeneiaeuei?e oaeaeeino? i?iaaaeaiiy ia?aiaiai caoaeaeaiiy o oaa?ei c
oeoe?iaei ae?aaaoii ia ?aii?o aoaiao ?icaeoeo caoai?thaaiiy, o ieo
aeyaeythoueny ci?ie, oa?aeoa?i? aeey ii/aoeiaeo aoai?a ?icaeoeo
ae?aaaoe/ieo naini?iiiioi?ieo iie?iae?iiao?e. Ca?eueoaiiy o?eaaeino?
oaeaeeiai iiceoeaiiai eiiiiiaioo

Iae.1. Iioaioe?aee aei?caeueii? iiaa?oi? nieiiiai iiceo, aeeeeeai?
iiae?aciaiiyi n?aeie/iiai ia?aa o ieoae eiio?ieueii? a?oie (A), c
NOC-?iaeoeiaaiei (A) oa aaiaoe/ii-aeaoa?i?iiaaiei (A) ae?aaaoii. Aeey
eiaeii? a?oie iaaaaeai? i?eeeaaee iioaioe?ae?a (1,2), ca?a?no?iaaieo o
??cieo oaa?ei. Aa?oi? caiene – ie?ai? ?a?no?aoe??, ieaei? – noiiaai?
caiene

Iae.2. Cia/aiiy ??aiy ae?eai?? (A), oaeaeeino? i?iaaaeaiiy aoa?aioii?
oaee? (A) oa o?eaaeino? ?? iiceoeaii? oace o ieoee eiio?ieueii? a?oie
(E), c NOC-?iaeoeiaaiei (NOC) oa aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii
(db).

na?ae/eoue i?i ci?io i?ioeaioiiai ni?aa?aeiioaiiy ia?aiaeo aieieii c
aenieith oa iecueeith oaeaee?noth i?iaaaeaiiy ia?aiaeo ?iioeuen?a o a?e
ca?eueoaiiy a?aeiinii? e?eueeino? inoaii?o, ui ni?aiaaea? c aeaieie
Terenghi et al., (1994).

I?e aeine?aeaeaii? noia?ii? aeoeaiino? iinoneiaioe/ieo aeaiaio?a o
aei?caeueiiio ?ic? nieiiiai iiceo aea/aeenue ci?ie iia?eueieo
eiiiiiaio?a IAeI. sse o oaa?ei c NOC-?iaeoeiaaiei ae?aaaoii, oae ? o
oaa?ei c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii a?aei?/aeinue i?aeaeuaiiy
aiie?ooaee N-oaee?, ui na?ae/eoue i?i i?aeneeaiiy noia?ii?
iinoneiaioe/ii? aeoeaiino? iae?ii?a aei?caeueiiai ?iao nieiiiai iiceo o
oeeo ieoae ii a?aeiioaiith aei eiio?ieueieo

oaa?ei (iae.3). O oaa?ei e?i?? db/db oaea i?aeaeuaiiy aoei iaio cia/iei,
i?ae o oaa?ei c NOC-?iaeoeiaaiei ae?aaaoii (?<0,05). Iae.3. Ci?ie aiie?ooaee iia?eueiiai iaaaoeaiiai eiiiiiaioo iioaioe?aeo aei?caeueii? iiaa?oi? nieiiiai iiceo o ieoae c NOC-?iaeoeiaaiei (A) oa aaiaoe/ii-aeaoa?i?iiaaiei (A) ae?aaaoii ii a?aeiioaiith aei eiai aiie?ooaee o eiio?ieueieo oaa?ei (A) Oiio a iiaeaeueoiio iaie aoei i?iaaaeaii aeine?aeaeaiiy aioo??oiuei-ee?oeiiiai eaeueoe??aiai aiiainoaco ia an?o eaieao naini?ieo ia?aiaeo oeyo?a. I?e oeueiio aeine?aeaeoaaany ia eeoa aacaeueiee aioo??oiueiee?oeiiee ??aaiue [Ca2+]i aea ? eaeueoe??a? neaiaee (“eaeueoe??a? o?aice?ioe”), ye? oa?aeoa?ecothoue i?ioeane iaaeoiaeaeaiiy (i?eiaa aiie?ooaea) oa aeaaaeaiiy (ia??iae oa?i?iaoe??) Na2+ c ee?oei a i?ioean? ?o caoaeaeaiiy.. Ii?ooaiiy ooieoe?? ia?aeiieo naini?ieo iae?ii?a aei?caeueieo aaiae??a nieiiiai iiceo. Iae?iie a AeANI iiaeia ?icae?eeoe ia aea? iniiai? a?oie ca ?ici??ii ?o niie - aaeee? oa iae?, ui a?aeiia?aeii a?aeaeathoue aoa?aioi? aieieia aaeeeiai oa iaeiai ae?aiao?o (c aenieith oa iecueeith oaeaee?noth i?iaaaeaiiy ia?aiaeo ?iioeuen?a). Iae?iie aaeeeiai ?ici??o iathoue aeniie oeio Aab, i?iaiaeyoue ia?aaaaeii i?ii??ioeaioeai? oa oaeoeeuei? aenoa?ioeaioeai? aoa?aioi? neaiaee, a oie /an ye iae?iie iaeiai ae?aiao?o c aeniiaie oeio Ad oa N (iai??e?i?ciaaieie) ia?aaeathoue ia?aaaaeii iioe?oeaioeai? oa oaiia?aoo?i? neaiaee. E??i oiai, oe? aea? a?oie iae?ii?a o ieoae a?ae??ciythoueny ca nai?ie iaoai?ciaie ?aaoethaaiiy aioo??oiueiee?oeiiiai Na2+. O aaeeeeo iae?iiao ?noioia cia/aiiy ia? iaeiie/aiiy oa ianooiia ca?eueiaiiy Na2+ nenoaiith aiaeiieaciaoe/iiai ?aoeeoeoio (eaeueoe?e-?iaeoeiaaia ca?eueiaiiy ), a oie /an ye o iaeeo iae?iiao oeae iaoai?ci ia ooieoe?iio? (Shmigol et al., 1994). Aoee aeei?enoai? eiio?ieuei? ieo? AN57Black6, ieo? oe??? ae e?i?? /a?ac 3 aei? i?ney aaaaeaiiy NOC, ieo? c ?icaeiooei NOC-?iaeoeiaaiei ae?aaaoii oa ieo? c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii. Na?aaei? ??ai? aacaeueiiai [Ca2+]i aeey aeuacacia/aieo a?oi oaa?ei a?aeiia?aeii neeaaeaee: 114,00 ± 4,49 iI, 117,05 ± 6,77 iI, 118,83 ± 8,64 iI oa 121,12 ± 4,85 iI aeey iioe?oeaioeaieo iae?ii?a oa 82,97 ± 6,36 iI, 85,38 ± 7,43 iI, 65,25 ± 4,86 iI oa 82,67 ± 3,90 iI aeey i?ii??ioeaioeaieo iae?ii?a. sse aeaeii c iaaaaeaieo aeua aeaieo, o eiio?ieueieo oaa?ei a?aei?/aeanue aeinoia??ia ??cieoey i?ae ??aiai aaciaiai [Na2+]i a iioe?oeaioeaieo oa i?ii??ioeaioeaieo iae?iiao (?<0,05). Oey ??cieoey caa??aaeanue ? o ieoae iaio e?i?e c oeoe?iaei ae?aaaoii. A oie aea /an nooo?ai? ??cieoe? a aaciaiio ??ai? [Ca2+]i. o iioe?oeaioeaieo iae?iiao ieoae c oeoe?iaei ae?aaaoii ii a?aeiioaiith aei eiio?ieueieo oaa?ei aecia/aii ia aoei. O?eueee o i?ii??ioeaioeaieo iae?iiao ieoae c ?icaeiooei NOC- ?iaeoeiaaiei ae?aaaoii a?aei?/aeanue oaiaeaioe?y aei cieaeaiiy aaciaiai ??aiy [Ca2+]i (0,1>?>0,05).

I?e aeine?aeaeaii? eaeueoe??aeo neaiae?a o iae?iiao AeANI, ui
aeeeeeaeenue aeaiiey?ecaoe??th iaia?aie aino?ith aie?eaoe??th ?ic/eio c
i?aeaeuaiei ai?noii ENl, aea/aeenue ?o i?eiaa aiie?ooaea oa
oa?aeoa?enoeee ia??iaeo oa?i?iaoe?? i?ae aieeaii aeaiiey?ecaoe??
ee?oeiii? iaia?aie, a naia /an iai?aniaaeo eaeueoe??aiai neaiaeo oa
??aaiue caeeoeiaiai eaeueoe?th. ??aaiue caeeoeiaiai [Ca2+]i aecia/aany
a?aeiia?aeii aei oi?ioee: R=(D[Ca2+]60/D[Ca2+]peak)o100%, aea D[Ca2+]60
-oea ??aaiue Na2+ a ee?oei? /a?ac 60 nae i?ney i?eieiaiiy
aeaiiey?ecaoe?? i?ion ??aaiue Na2+ a niiei?, a D[Ca2+]peak — oea
??cieoey i?ae aaniethoiei cia/aiiyi i?eiai? aiie?ooaee oa ??aiai Na2+ a
niiei? (Shmigol et al., 1995). Oaeee can?a aei??o aeicaieya ii??aithaaoe
ci?ie ??aith caeeoeiaiai Na2+ o ee?oeiao c ??ciith aiie?ooaeith
ii/aoeiai eaeueoe??aiai neaiaeo.

Na?aaei? cia/aiiy i?eiaeo aiie?ooae o iioe?oeaioeaieo iae?iiao oaa?ei
c oeoe?iaei ae?aaaoii aeinoia??ii ia a?ae??ciyeenue a?ae oaeeo o
eiio?ieueieo oaa?ei ? noaiiaeee 470 ± 97 iI, 499 ± 88 iI oa 546 ± 89 iI
aeey eiio?ieueieo oaa?ei, ieoae c ?icaeiooei NOC-ae?aaaoii oa ieoae c
aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii a?aeiia?aeii.

I?e aeine?aeaeaii? oa?i?iaoe?? eaeueoe??aeo o?aice?io?a aoei
anoaiiaeaii, ui o ieoae ye c NOC-?iaeoeiaaiei, oae ? c
aaiaoe/ii-aeaoa?i?iiaaiei oeoe?iaei ae?aaaoii a iioe?oeaioeaieo iae?iiao
a?aei?/a?oueny nooo?aa ca?eueoaiiy o?eaaeino? eaeueoe??aeo neaiae?a,
ia?aaaaeii ca ?aooiie caeeoeiaiai i?aeaeuaiiy ??aiy [Na2+]i ia i?ioyc?
aeanyoe?a naeoiae i?ney i?eieiaiiy aeaiiey?ecaoe??.

Oaae.1

I?eia? cia/aiiy aiie?ooae oa ??ai? caeeoeiaiai [Ca2+]i o iioe?oeaioeaieo
iae?iiao eiio?ieueieo oaa?ei oa oaa?ei c oeoe?iaei ae?aaaoii

————————————————————————
——————————-

cia/aiiy aiie?ooae caeeoeiaee [Ca2+]?

o?eaae.aeaiie.
iI DA60nae, %

eiio?. oaa?eie (0,5 nae) 311,98±36,08
1,16±0,22

(5,0 nae) 468,83±63,18
6,83±0,85

?>0,05
?<0,05 NOC-eiio?ieue (0,5 nae) 320,04±80,04 1,12±0,39 (5,0nae) 469,78±114,69 6,34±0,88 ?>0,5
? <0,05 NOC-ae?aaao 1 oae. (0,5nae) 362,71±74,96 3,72±3,57 (5,0nae) 497,51±50,56 16,63±3,84 ??>0,5
?<0,001 NOC-ae?aaao 3 oi. (0,5nae) 269,10±32,19 2,19±0,39 (5,0nae) 451,78±52,09 17,13±2,94 ?<0.05 ?<0,001 db/db oaa?eie (0,5nae) 442,41±48,11 2,61±0,66 (5,0nae) 546,89±19,93 17,00±1,33 ?>0,5
?<0,001 ------------------------------------------------------------------------ -------------------------------- Iae.4. I?eeeaaee ?a?no?aoe?? aeeeeeaieo 5-oe naeoiaeiith aeaiiey?ecaoe??th iaia?aie eaeueoe??aeo neaiae?a o aaeeeiio (A) oa iaeiio (A) ia?aeiieo naini?ieo iae?iiao eiio?ieueii? oaa?eie (e?ai?o/) oa oaa?eie c NOC-?iaeoeiaaiei ae?aaaoii (i?aai?o/) Cia/aiiy aiie?ooae oa caeeoeiaiai eaeueoe?th i?e ??ci?e o?eaaeino? aeaiiey?ecaoe?? iaaaaeaii o oaae. 1. I?aeaeuaiiy caeeoeiaiai [Ca2+ ]i noa? iaea?eueo cia/iei i?e o?eaaeino? aeaiiey?ecaoe?? 5,0 nae (?<0,001) ? caa??aa?oueny iaa?oue /a?ac 1 oaeeeio i?ney cae?i/aiiy aeaiiey?ecaoe?? (iae.4). ?ieue i?oioiiae??e a ii?ooaii? eaeueoe??ai? neaiae?caoe?? a ia?aeiieo naini?ieo iae?iiao i?e ae?aaao?. Iiaeiaaeai?noue eaeueoe??aeo neaiae?a oa ?io? ?o ci?ie iiaeooue aooe i?e/eiith iioaioe?aoe?? neiaioe/ii? ia?aaea/? oaeeo neaiae?a ia ianooii? iae?iie iioe?oeaioeaii? nenoaie ? aeieeiaiiy ee?i?/ieo neiioii?a iae?iiao?e. Iaoai?ciii oaeiai yaeua ia? aooe ii?ooaiiy ee?oeiieo iaoai?ci?a, a?aeiia?aeaeueieo ca aeaeaeaiiy Na2+ c oeeoicieth i?ney ??ceiai i?aeaeuaiiy ?o ??aiy i?ae/an caoaeaeaiiy ee?oeie. Iaeiei c eiiiiiaio?a oaeeo ii?ooaiue iiaeooue aooe ci?ie a i?oioiiae??yo. Inoaii?i /anii anoaiiaeaii, ui i?oioiiae??? iiaeooue aoaeoeaii iiaeeiaoe oe? eiie c oe?oicieth caaaeyee aeoeaiino? oi?ii?oa?ii? nenoaie ?o aioo??oiuei? iaia?aie. Oey nenoaia aeei?enoiao? i?ioiiiee a?aae??io, ui ?nio? ia aioo??oi?e iaia?ai?, aeey aoaeoeaiiai o?ainii?oo Na2+ a i?oioiiae???. Ii?yae c oeei, i?oioiiae??aeueia iaia?aia ia? oaeiae nenoaie Ca2+/Na+ oa Ca2+/H+ iai?iiee?a, ui i?ney cae?i/aiiy caoaeaeaiiy iiaeooue aea?eueiyoe Na2+ iacaae o oeeoicieue. Aeey aeeeth/aiiy i?oioiiae??aeueiiai iaeiie/aiiy Na2+ aeei?enoiaoaaeanue ae?y i?ioiioi?o NNN?, ui aeeeeea? aeeneiaoe?th i?ioiiiiai a?aae??ioo ia aioo??oi?e iaia?ai?. Oaea ae?y naia ii nia? iaeaea ia aeeeeeaea i?aeaeuaiiy aaciaiai ??aith a?eueiiai Na2+ a oeeoicie?. Ae?y NNN? i?eaiaeeea aei ae?acieo aeaiyeeo ci?i a oa?aeoa?enoeeao eaeueoe??aeo o?aice?io?a, aeeeeeaieo aoiaeii o ee?oeio Na2+ /a?ac aeoeaiaai? iaia?aiiith aeaiiey?ecaoe??th eaeueoe??a? eiii? eaiaee: aiia ca?eueooaaea i?e oeeo o?aice?io?a (ca ?aooiie i?aeaeuaiiy aiie?ooaee oa cia/iiai eiai iiaeiaaeaiiy), a oie aea /an nei?i/oth/e caeeoeiaa i?aeaeuaiiy oeeoicieueiiai ??aith Na2+. Oe? ci?ie aeacothoue ia i?eai?/aiiy NNN? aeoioeyoe?? Na2+ i?oioiiae??yie c oeeoicieth i?ae /an i?ea o?aice?ioa oa ianooiiiai iia?eueiiai aea?eueiaiiy ?o iacaae o oeeoicieue i?ney i?eieiaiiy caoaeaeaiiy. Aeniiaie i?i oa, ui iia?eueia aea?eueiaiiy Na2+ iacaae o oeeoicieue i?ney iaaaioaaeaiiy oeeie eiiaie i?oioiiae??e iacaae o oeeoicieue cae?enith?oueny naia nenoaiaie Ca2+/Na+ oa Na2+/H+ iai?io, i?aeoaa?aeaeo?oueny eiai i?eai?/aiiyi niaoeeo?/iei ?ia?a?oi?ii oaeiai iai?io oao?aoai?e-oinoii??i (O??+). I?oioiiae??aeueiee i?ioiiioi? NNN? o iae?iiao oaa?ei c NOC-?iaeoeiaaiei ae?aaaoii, ye ? o iae?ii?a eiio?ieueieo oaa?ei, aeeeeeaa ca?eueoaiiy aiie?ooaee oa iiaeiaaeaiiy /ano i?aniaaeo aeeeeeaieo iaia?aiiith aeaiiey?ecaoe??th eaeueoe??aeo o?aice?io?a, aea iaio cia/ia, i?ae o eiio?ie?. O oaa?ei c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii i?aeaeuaiiy aiie?ooaee o?aice?io?a acaaae? aoei iaiii?oiei. O?eaae?noue i?eo (ui aei??thaaeanue /anii eiai iai?aniaaeo) a iaeeo iae?iiao oaa?ei c NOC-ae?aaaoii i?ae NNN? oaeiae ca?eueooaaany iaio nooo?ai, i?ae a aiaeia?/ieo iae?iiao caei?iaeo oaa?ei. Ua a?eueo ?noioii ci?ithaaany caeeoeiaee ??aaiue oeeoicieueiiai Na2+. Oa?aeoa?ia aeey ae?aaaoe/iiai noaio o?eaaea i?aeaeuaiiy oeueiai ??aiy a cia/i?e i??? onoaaeinue ia oii? ae?? i?ioiiioi?o. A?aeiia?aei? noaoenoe/i? aeai? i?i ci?ie eaeueoe??aeo neaiae?a a aaeeeeo oa iaeeo iae?iiao noiiaai? ia Oaae.2. Oaae.2 Ci?ie oa?aeoa?enoee eaeueoe??aeo neaiae?a naini?ieo iae?ii?a i?ae aieeaii i?eeeaaeaiiy i?oioiiae??aeueiiai i?ioiiioi?o NNN? ------------------------------------------------------------------------ ----- Aaeee? iae?iie Iae? iae?iie aei NNN? i?ney NNN? aei NNN? i?ney NNN? eiio?ieue aiie.(ieI) 711,8±72,4 2108,0±173,1 556,0±57,9 869,0±124,0 i?aniaae (nae) 3,8±0,6 18,6±3,1 7,8±2,9 28,5±3,4 ?<0.001 ?<0.001 caeeoe.i?aea.(%) 8,4±1,2 6,4±3,9 8,1±1,1 5,8±1 ?>0,5
??>0,5

__________________________________________________________

NOC-ae?aaao

aiie.(ieI) 818,4±58,9 1875,0±360,4 791,1±79,3 938,9±80,7

i?aniaae (nae) 3,4±1,3 12,3±2,3 6,7±1,1
14,8±4,0

?<0.001 ?<0.05 caeeoe.i?aea.(%) 11,8±1,9 7,0±1,8 20,9±3,7 9,2±1,8 ?<0.05 ?<0.001 __________________________________________________________ db/db aiie.(ieI) 994,1±140,5 897,9±126,3 827,5±16,8 744,8±68,7 i?aniaae (nae) 2,8±0,7 10,7±1,9 6,8±2,5 16,3±3,3 ?<0.001 ?<0.001 caeeoe.i?aea.(%) 8,8±2,1 3,4±0,2 10,2±2.2 3,9±1,9 ?<0.001 ?<0.001 Ia iae.5 iaaaaeai? i?eeeaaee ci?i eaeueoe??aeo neaiae?a o aaeeeiio (A) oa iaeiio (A) naini?ieo iae?iiao eiio?ieueii? (e?ai?o/) oa NOC-ae?aaaoe/ii? (i?aai?o/) oaa?eie ia oii? i?eeeaaeaiiy i?oioiiae??aeueiiai i?ioiiioi?o NNN?. Iae.5. I?eeeaaee ci?i eaeueoe??aeo neaiae?a aaeeeiai (A) oa iaeiai (A) naini?ieo iae?ii?a eiio?ieueii? oaa?eie oa oaa?eie c NOC-?iaeoeiaaiei ae?aaaoii ia oii? ae?? i?oioiiae??aeueiiai i?ioiiioi?o NNN?. 5-oe naeoiaei? aeaiiey?ecaoe?? iaia?aie 50 iI KCl a?aei?/ai? no??eeaie Oaea?aeai? ?acoeueoaoe iiaeooue aeacoaaoe ia oa, ui caoiieaiiy eii?a Na2+ i?oioiiae??yie i?ae /an NOC-?iaeoeiaaiiai ae?aaaoo ana ua ia? i?noea, aea noa? iaio aoaeoeaiei. Ca?eueoaia caeeoeiaa i?aeaeuaiiy ??aith Na2+ a oeeoicie?, iniaeeai oa?aeoa?ia aeey iaeeo naini?ieo iae?ii?a o ae?aaaoe/ieo oaa?ei, ia oii? ae?? NNN? iiaa?oaeinue aei ??aiy, oa?aeoa?iiai aeey iae?ii?a eiio?ieueieo oaa?ei. Oea aea? i?aenoaae aaaaeaoe, ui a iineeaii? oaeiai i?aeaeuaiiy i?e ae?aaao? iaaio ?ieue a?a? naia cao?eiaia aea?eueiaiiy Na2+ c i?oioiiae??e. I?e aaiaoe/ii-aeaoa?i?iiaaiiio ae?aaao? iae?iiaeuei? iaoai?cie, a?aeiia?aeaeuei? ca oi?ioaaiiy eaeueoe??aeo neaiae?a i?ae /an caoaeaeaiiy ee?oeie, a oiio /ene? Na2+-aeoioethth/a ooieoe?y i?oioiiae??e, oaeiae ii?ooothoueny, aea ii ?ioiio, i?ae i?e ?COeAe, aeeeeeaiiio aaaaeaiiyi no?aioicioioeeio. A?aenooi?noue i?aeaeuaiiy aiie?ooaee eaeueoe??aiai o?aice?ioo i?ae aieeaii i?eeeaaeaiiy i?oioiiae??aeueiiai i?ioiiioi?o NNN? na?ae/eoue i?i oa, ui ??cieoey uaeaeoa anueiai iieyaa? a ci?iao i?ioeano iaeiie/aiiy Na2+ i?oioiiae??yie. Aeey i?yiiai aeine?aeaeaiiy ci?i aeeeaeo Na2+, iiia?aaeiuei iaeiie/aiiai o i?oioiiae??yo, i?oioiiae??aeueiee i?ioiiioi? NNN? i?eeeaaeaany aaciina?aaeiuei i?ney aeeeeeo eaeueoe??aiai o?aice?ioo oa a?aeiia?aeiiai iiaeeiaiiy eii?a i?oioiiae??yie. Oaea i?eeeaaeaiiy aeeeeaaei iiyao ianiaaiiai aeeeaeo Na2+ a oeeoicieue, i?e yeiai a iae?iiao oaa?ei c NOC-?iaeoeiaaiei ae?aaaoii ciaiooaaany o aeae?eueea ?ac?a (c 1423,3 ± 115,1 iI aei 58,0 ± 13,5 iI o aaeeeeo iae?iiao oa c 1053,5 ± 105,4 iI aei 69,7 ± 11,4 iI o iaeeo, ?<0,001). A?aeiia?aeiee i?eeeaae iaaaeaiee ia iae.6. Iae.6. ?a?no?aoe?? aeeeeeaieo aeaiiey?ecaoe??th eaeueoe??aeo neaiae?a o aaeeeiio (A) oa iaeiio (A) naini?ieo iae?iiao eiio?ieueii? ieo? (aa?oi?e ?yae) oa ieo? c NOC-?iaeoeiaaiei ae?aaaoii (ieaei?e ?yae). Ii??aithaaeenue ?aaeoe?? aei i?eeeaaeaiiy i?ioiiioi?o NNN? oa i?e eiai i?eeeaaeaii? i?ney aeinyaiaiiy eaeueoe??aeie neaiaeaie i?eiaiai cia/aiiy Iaea?aeai? ?acoeueoaoe i?aeoaa?aeaeothoue c?iaeaiee aeua aeniiaie i?i oa, ui aeieeiaiiy ae?aaaoe/iiai noaio iia??oo? iaeiie/aiiy eii?a Na2+ o i?oioiiae??yo ? iniaeeai eiai aeeeae iacaae o oeeoicieue nenoaiith eiiieo iai?iiee?a. Cae/aeii, ?io? eiiiiiaioe iaoai?cio i?aeo?eiaiiy eaeueoe??aiai aiiainoaco a ia?aiaeo ee?oeiao oaeiae iiaeooue ii?oooaaoenue i?e ae?aaaoe/i?e iaoieia??. Ooieioe?iioaaiiy nenoai aeoeaiiai o?ainiaia?aiiiai o?ainii?oo Na2+. Nooo?ao ?ieue o ?aaoeyoe?? aioo??oiueiee?oeiiiai aiiainoaco Na2+, a oiio /ene? o a?aeiiaeaii? aaciaiai ??aith oeeo eii?a i?ney caoaeaeaiiy, i?eeia? aeoeaiee o?ainii?o ?o /a?ac ee?oeii? iaia?aie ca aeiiiiiaith eaeueoe??aeo AOOac ?-oeio. A ?yae? aeine?aeaeaiue iieacaii ii?ooaiiy aeoeaiino? Na+/K+-AOOace i?e ??cieo iineeaaeiaiiyo oeoe?iaiai ae?aaaoo, a oiio /ene? ? i?e ae?aaaoe/i?e iie?iae?iiao?? (Kaur & Lackman, 1994; Ver et al., 1995). A iaoeo aeine?aeaeaiiyo ??ai? ae?eai?? o eiio?ieueii? a?oie ieoae e?i?? AC57Black6 noaiiaeee 4.04 ± 0.17 iiieue/e, a o ieoae c NOO-?iaeoeiaaiei oa aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii aiie a?aeiia?aeii c?inoaee aei 21.15 ± 0.50 oa 11.45 ± 0.50 iiieue/e. Aeoeai?noue Na2+-AOOace a aiiiaaiao? nieiiiai iiceo o eiio?ieueii? a?oie ieoae noaiiaeea 5,75 ± 0,70 ieI ?/ia a?eea/aiae, a o ieoae c NOC-?iaeoeiaaiei ae?aaaoii neeaaeaea 6,07 ± 0,9 ieI ?/ia a?eea/aiae oa o ieoae c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii 9.40 ± 1.51 ieI ?/ia a?eea /aiae (?<0,05). Aeoeai?noue Na2+-AOOace a aiiiaaiao? aieiaiiai iiceo eiio?ieueii? a?oie ieoae noaiiaeea 11,96 ± 2,37 ieI ?/ia a?eea/aiae oa 9,3 ± 1,4 ieI ?/ia a?eea/aiae i?e NOC-?iaeoeiaaiiio oeoe?iaiio ae?aaao?; aeinoia??ii? ??cieoe? aeyaeaii ia aoei. O ieoae c aaiaoe/ii-aeaoa?i?iiaaiei ae?aaaoii ??aaiue aeoeaiino? o aieiaiiio iiceo oaeiae aoa i?aeaeuaiei (14.83 ± 1.28 ieI ?/ia a?eea/aiae). A oie aea /an ne?ae a?aecia/eoe oe?ieo aa??aaaeuei?noue io?eiaieo aeaieo. O oaa?ei e?i?? db/db aaeeei? aaae (a?ey 100 a) c ?icaeiaiei iaee??iiyi oa aaeeeith o?eaae?noth ae?aaaoo ie io?eiaee iecuee? iieacieee AOO-ace a i?aia?aoao nieiiiai oa aieiaiiai iiceo ii a?aeiioaiith aei oaa?ei a?eii 10-12 oeaei?a. Oaeei /eiii, i?e ?icaeoeo iaio oei?a oeoe?iaiai ae?aaaoo o i?aeaeine?aeieo oaa?iei ia a?aecia/a?oueny aeinoia??ii? ci?ie noia?ii? Na2+-AOOacii? aeoeaiino? a oeaiei? ye nieiiiai, oae ? aieiaiiai iiceo. Iiaeeeai, ui aecia/aiiy noia?ii? AOOacii? aeoeaiino?, ui aeeth/a? aeoeai?noue ye ieaciaeaiaeueieo, oae ? aiaei?aoeeoey?ieo AOOac, a aeaiiiio ?ac? ? iaaeinoaoiuei /ooeeaeo iaoiaeii, ? a?aenooi?noue ci?i a aeoeaiino? NE?EA iiaea ianeoaaoe iaaaeee? ci?ie aeoeaiino? IMEA, ? iaaiaee. Eaeueoe??aa neaiae?caoe?y a iae?iiao aei?caeueiiai ?iao nieiiiai iiceo i?e NOC-?iaeoeiaaiiio ae?aaao?. Aecia/ai? aeua aeeaie? ci?ie o eaeueoe??a?e neaiae?caoe?? a ia?aeiieo naini?ieo iae?iiao, iniaeeai cia/i? o iae?iiao iaeiai ae?aiao?o, ui a?aeiia?aeathoue ca ia?aaea/o iioe?oeaioeaieo neaiae?a, iiaeooue i?eaiaeeoe aei cia/ieo ci?i o ia?aaea/? ia?aiai? ?iioeuenaoe?? aei aeueo iiceiaeo no?oeoo?, o ia?oo /a?ao ia iae?iie aei?caeueiiai ?iao nieiiiai iiceo. Iae?iie aei?caeueiiai ?iao, aieiaiei /eiii iae?iie aeaeao?iicii? noanoaioe??, ? aoi?eiieie naini?ieie iae?iiaie, ui aeathoue ii/aoie aeno?aeiei iioe?oeaioeiei oeyoai ? ia?aaeathoue nai? neaiaee o a?aeiia?aei? no?oeoo?e aieiaiiai iiceo. sse ? o aeiaaeeo ia?aeiieo naini?ieo iae?ii?a, ie ia aecia/eee ?noioieo ci?i aaciaiai ??aith (??aith niieith) [Ca2+]i i?e ii??aiyii? iaio a?oi oaa?ei. Oeae ??aaiue a aeai?e na??? noaiiaea 68 ± 15 iI aeey iae?ii?a eiio?ieueieo oaa?ei oa 82 ± 17 iI aeey iae?ii?a oaa?ei c NOC-?iaeoeiaaiei ae?aaaoii. ?noioieo ci?i i?eiaeo cia/aiue aiie?ooaee o?aice?io?a oa oaeaeeino? ?o ia?inoaiiy i?ae iae?iiaie iaio a?oi oaa?ei oaeiae ia a?aei?/aii. Na?aaei? cia/aiiy aiie?ooaee [Ca2+]i o?aice?io?a noaiiaeei 719 ± 132 iI aeey eiio?ieueieo oaa?ei oa 694 ± 155 iI aeey ae?aaaoe/ieo oaa?ei. Iino?eia /ano ia?inoaiiy oeeo o?aice?io?a (yea aecia/aeanue ca aeiiiiiaith ai?ieneiaoe?? iiiiaeniiiaioe?aeueiith ooieoe??th) noaiiaeea 3,0 ± 0,8 nae ye aeey eiio?ieueieo iae?ii?a, oae ? aeey iae?ii?a ae?aaaoe/ieo oaa?ei. Iniiaiith a?aei?iith aeeeeeaieo iaia?aiiith aeaiiey?ecaoe??th eaeueoe??aeo o?aice?io?a a iae?iiao ae?aaaoe/ieo oaa?ei a ii??aiyii? c iae?iiaie eiio?ieueieo aoei cia/ia oiia?eueiaiiy aeaaaeaiiy Na2+ c oeeoiieacie i?ney cae?i/aiiy noeioeyoe??. A ee?oeiao eiio?ieueieo oaa?ei ??aaiue [Ca2+]i i?ney i?eieiaiiy aeaiiey?ecaoe?? (20 nae aie?eaoe?? ciai?oiueiai ?ic/eio c 50 iI ENl) iiiiaeniiiaioe?aeueii iiaa?oaany aei ??aith niieith c iino?eiith /ano 8,0 ± 0,5 nae. Ia a?aei?io a?ae oeueiai, a iae?iiao ae?aaaoe/ieo oaa?ei a?aeiiaeaiiy a?aeaoaaeinue a?eueo neeaaeii; o an?o aeiaaeeao iiaeia aoei aeae?eeoe ae?oao, aeoaea iia?eueio aeniiiaioe?aeueio eiiiiiaioo (iae. 7). Iae.7. I?eeeaaee eaeueoe??aeo o?aice?io?a, aeeeeeaieo aeaiiey?ecaoe??th iaia?aie oa i?eeeaaeaiiyi eioa?io a iae?iiao aei?caeueiiai ?iao nieiiiai iiceo eiio?ieueiiai uo?a (A) oa uo?a c NOC-?iaeoeiaaiei ae?aaaoii Iino?ei? /ano oeeo aeaio oac i?ioeano a?aeiiaeaiiy aaciaiai ??aith [Ca2+]i a iae?iiao ae?aaaoe/ieo oaa?ei noaiiaeee a?aeiia?aeii 7,2 ± 0,5 nae oa 23,0 ± 0,6 nae. ??aaiue aioo??oiueiee?oeiiiai Na2+, ui aecia/aany /a?ac 90 nae i?ney i?eieiaiiy noeioeyoe??, aoa cia/ii aeuei, i?ae eiai aaciaee ??aaiue o eiio?ieueieo oaa?ei (?<0,001). Ae?aa?aia ci?i a?aeiiniiai ??aith caeeoeiaiai eaeueoe?th o iae?iiao eiio?ieueieo oaa?ei oa oaa?ei /a?ac 1 oa 7 oeaei?a i?ney aaaaeaiiy NOC, o ii??iyii? aei ??aith ae?eai??, iaaiaeeoueny ia iae.8 (e?ai?o/). C iaaaaeaii? ae?aa?aie aeaeii, ui ci?ie ??aith caeeoeiaiai Na2+ aoee iia'ycai? c nooiaiai ?icaeoeo oeoe?iaiai ae?aaaoo (??aiai a?ia?ae?eai??). Eaeueoe??a? neaiaee, aeeeeeai? aeeeaeii Na2+ c aiaeiieaciaoe/iiai ?aoeeoeoio i?e ae?? eioa?io. Iaei??th c i?e/ei a?eueo nooo?aiai ii?ooaiiy eaeueoe??ai? neaiae?caoe?? a iae?iiao aei?caeueiiai ?iao iiaea aooe ii?ooaiiy aea?eueiaiiy Na2+ c Na2+-/ooeeaeo aeaii aiaeiieaciaoe/iiai ?aoeeoeoio. Aenia?eiai-oaeueiei iaoiaeii aeoeaaoe?? nenoaie oaeiai aea?eueiaiiy Na2+ ? ae?y eioa?io aaciina?aaeiuei ia ?aoeaioi?e iaia?aie aiaeiieaciaoe/iiai ?aoeeoeoio. I?eeeaaeaiiy 30 iI eioa?io aei iae?ii?a aei?caeueiiai ?iao a c??cao nieiiiai iiceo ye eiio?ieueieo, oae ? ae?aaaoe/ieo oaa?ei aeeeeeaei o?aiceoi?ia i?aeaeuaiiy a ieo oeeoicieueiiai ??aith Na2+. Aaee/eia oaeiai i?aeaeuaiiy ia ci?ithaaeanue i?e 5-oe oaeeeii?e noia?ooc?? c??c?a aaceaeueoe??aei ?ic/eiii, ui i?aeoaa?aeaeo? iaaeoiaeaeaiiy eii?a naia c A?. Na?aaei? cia/aiiy aiie?ooae oeeo ?aaeoe?e noaiiaeee 268 ± 29 iI a eiio?ieueieo ee?oeiao, 149 ± 27 oa 31 ± 9 iI a iae?iiao /a?ac 1 oa 7 oeaei?a i?ney aaaaeaii? NOC (iae.8, i?aai?o/). Iae.8. ??aaiue caeeoeiaiai Na2+ /a?ac 90 nae i?ney i?eieiaiiy aeaiiey?ecaoe?? (e?ai?o/) oa aeeeae Na2+ c aiaeiieaciaoe/iiai ?aoeeoeoio (i?aai?o/) a iae?iiao aei?caeueiiai ?iao eiio?ieueieo uo??a oa uo??a /a?ac 1 oa 7 oeaei?a i?ney aaaaeaiiy NOC. I?ae ae?aa?aiith aeacai? a?aeiia?aei? ??ai? ae?eai??. Aieea aeieaoi?a eaeueoe??aeo eaiae?a i?oaae?i?io ia eaeueoe??a? neaiaee a iae?iiao aei?caeueiiai ?iao. Aeey aecia/aiiy ?ie? iaaieo oei?a eaeueoe??aeo eaiae?a o aaia?aoe?? eaeueoe??aeo neaiae?a a iae?iiao aei?caeueiiai ?iao iaie aoa aeine?aeaeaiee aieea ia iienai? aeua eaeueoe??a? o?aice?ioe aeieaoi?a eaeueoe??aeo eaiae?a L-oeio i?oaae?i?io a eiioeaio?aoe?? 50-100 ieI. Aieea oeueiai aeieaoi?a ia?aa??yany ia iae?iiao eiio?ieueieo oaa?ei ? oaa?ei c NOC-?iaeoeiaaiei ae?aaaoii. Aiie?ooaea aeeeeeaieo aeaiiey?ecaoe??th o?aice?io?a a iae?iiao eiio?ieueieo oaa?ei cieaeoaaeanue i?ae aieeaii i?oaae?i?io a?ae 719 ± 132 iI aei 556 ± 24 iI A oie aea /an o oaa?ei c NOC-ae?aaaoii i?eai?/oth/a ae?y i?oaae?i?io cia/ii iineethaaeanue - a?ae 694 ± 155 iI aei 278 ± 28 iI, ?<0,001. Oaeei /eiii, a aaia?aoe?? eaeueoe??aeo neaiae?a a aeine?aeaeaieo iae?iiao ?noioio ?ieue a?athoue naia i?oaae?i?i-/ooeea? eaeueoe??a? eaiaee; i?e ?icaeoeo NOC-?iaeoeiaaiiai ae?aaaoo i?eai?/oth/ee aoaeo i?oaae?i?io ?noioii iineeth?oueny, ui iiaea aeacoaaoe aai ia i?aeaeuaiiy aeni?an?? a?aeiia?aeieo eaiae?a, aai ia i?aeaeuaiiy aoaeoeaiino? ae?? ia ieo aeacaiiai aeieaoi?a. Io?eiai? aeai? na?ae/aoue i?i oa, ui ci?ie eaeueoe??aiai aiiainoaco ia ??ai? aoi?eiieo naini?ieo iae?ii?a nieiiiai iiceo ? aeoaea cia/ieie. Ci?iththoueny aieiaiei /eiii iaoai?cie, ui a?aeiiaeththoue aaciaee ??aaiue Na2+ i?ney eiai ??ceiai i?aeaeuaiiy o ca`yceo ?c caoaeaeaiiyi ee?oeie ? aaia?aoe?? a i?e eaeueoe??aeo neaiae?a. A?aeiia?aeii oa?aeoa?enoeee ia?aaea/? ia?aiaiai caoaeaeaiiy i?e i?e?iaei?e aeey ia?aiai? nenoaie ?eoi?/i?e ?iioeuenaoe?? aoaeooue oaeiae ?noioii ii?oooaaoenue. Eaeueoe??aa neaiae?caoe?y a iae?iiao oaeaiono, iici/eo oa aaeeeeo i?aeoeue aieiaiiai iiceo i?e NOC-?iaeoeiaaiiio ae?aaao?. A ca`yceo c iaaaaeaieie a iiia?aaei?o ?icae?eao ?acoeueoaoaie aoei aaaeeeaei c`ynoaaoe, /e aeieeathoue ?iaeoeiaai? ae?aaaoii ci?ie o eaeueoe??a?e neaiae?caoe?? oaeiae o iae?iiao aeueo iiceiaeo no?oeoo?, iniaeeai o iae?iiao oaeaiono oa naini?ii? iaeano? ei?e aaeeeeo i?aeoeue, ? oaeei /eiii anoaiiaeoe, iane?eueee oae? ci?ie ?ai?acaioothoue caaaeuei? ?ene ae?aaaoe/ii? iaoieia??. ?acoeueoaoe aeacothoue ia aeayeo oaiaeaioe?th aei ca?eueoaiiy aiie?ooaee eaeueoe??aeo neaiae?a o ei?oeeaeueieo iae?ii?a ae?aaaoe/ieo oaa?ei; aea aiia ia aoea noaoenoe/ii cia/eiith. *an iai?aniaaeo aeeeeeaieo iaia?aiiith aeaiiey?ecaoe??th eaeueoe??aeo neaiae?a o eiio?ieueieo oaa?ei aoa ?aeaioe/iei a iae?iiao iaio no?oeoo?; a?i ?noioii ia ci?ithaaany i?e ?icaeiooiio NOC-?iaeoeiaaiiio ae?aaao?. Iaaiaee, caeeoeiaa i?aeaeuaiiy [Ca2+]i aoei a?eueo cia/iei o oaa?ei c oeoe?iaei ae?aaaoii (Iae. 9, Oaae.3). Oe? ci?ie aoee, iaeiae, cia/ii iaio aecia/aieie, i?ae o ia?eoa?e/ieo iae?iiao. Iae.9. I?eeeaaee eaeueoe??aeo o?aice?io?a, ca?a?no?iaaieo a?ae iae?ii?a ei?e aaeeeeo i?aeoeue iiceo o a?aeiia?aeue ia aeaiiey?ecaoe?th iaia?aie eiio?ieueii? oaa?eie (e?ai?o/) oa oaa?eie c NOC-?iaeoeiaaiei ae?aaaoii (i?aai?o/) Oaae.3 Oa?aeoa?enoeee eaeueoe??aeo o?aice?io?a a iae?iiao ei?e oa oaeaiono ----------------------------------------------------------------------- ---------------------------------------------- eiio?ieue n NOC-eiio?. n NOC-ae?aaao n aiie?ooaee (ii) ei?a 280±55 12 275±61 10 317±98 18 oaeaion 278±72 10 284±48 7 296±63 10 ?>0,5 ?>0,5

————————————————————————
———

caeeoeiaee Na2+ (%)

ei?a 18,35±3,20 12 16,43±2,40 10 20,62±2,10

?>0,5
?<0,05 oaeaion 17,35±5,50 10 19,62±3,70 7 22,80±3,20 ?>0,5
?<0,05 Aoei oaeiae i?iaaaeaia aeine?aeaeaiiy ae?? ??cieo aeieaoi??a Na2+ eaiae?a ia iae?iie eiio?ieueieo oa ae?aaaoe/ieo oaa?ei (oaae.4). O ei?oeeaeueieo iae?iiao niinoa??aaeanue noaoenoe/ii aeinoia??ia ci?ia a aoaeoeaiino? i?eai?/oth/i? ae?? Ni2+ oa i?iiae?i?io: aeeeeeaiee i?iiae?i?iii aoaeo aoa a?eueo ?ioaineaiei o ii??aiyii? c ae??th Ni2+, aoaeoeai?noue yei? cieaeoaaeanue. Oae? ii?ooaiiy o ae?aaaoe/ieo oaa?ei iiaeooue aooe iia`cai? c? ci?iaie a aeni?an?? eaiae?a L-oeio oa a?aeiia?aeii a ni?aa?aeiioaii? ??cieo oei?a iioaioe?ae-ea?iaaieo Na2+ eaiae?a. Oaae.4 I?eai?/aiiy aiie?ooae eaeueoe??aeo o?aice?io?a (a %%) a iae?iiao ei?e aaeeeeo i?aeoeue oa oaeaiono, i?ae aieeaii Ni2+ oa i?iiae?i?io ------------------------------------------------------------------------ ---------------------------------------------- eiio?ieue n NOC-1 oeae. n NOC-7 oeaei. n iae?iie ei?e 50 mM Ni2+ 26±13 10 22±8 11 13±7 13 ?<0,01 100 mM i?iiae?i?i 36±16 10 32±10 15 54±15 17 ?<0,05 ------------------------------------------------------------------------ ---------------- iae?iie oaeaiono 50 mM Ni2+ 21±10 7 27±10 10 30±7 9 100 mM i?iiae?i?i 8±8 6 10±8 12 6±4 13 ?>0,5

Eaeueoe??a? neaiaee, aeeeeeai? a ei?eiaeo oa oaeai?/ieo iae?iiao
aeoeaaoe??th io??iiaeo ?aoeaioi??a. Aeey oaeai?/ieo oa ei?oeeaeueieo
iae?ii?a oa?aeoa?iith ?enith ? ?noioiee i?iya io??ia?a?/ieo
iae?iiaae?aoi?ieo iaoai?ci?a, ui cae?eniththoue naith ae?th /a?ac
aeoeaaoe?th ca aeiiiiiaith AOO ye eiiio?iiieo, oae ? iaoaaio?iiieo
?aoeaioi??a (Lalo et al., 1998). Ie i?ine?aeeoaaee oaeiae iiaeeeao ci?io
oeeo iaoai?ci?a i?e NOC-?iaeoeiaaiiio ae?aaao? (oaae.5).

O oaa?ei c oeoe?iaei ae?aaaoii na?aae iae?ii?a ei?e eaeueoe??a?
o?aiece?ioe aeieeaee i?e ae?? AOO ia?aaaaeii eeoa i?e iayaiino? Na2+ o
iicaee?oeiiieio ?ic/ei?. A aaceaeueoe??aiio iicaee?oeiiiio ?ic/ei? aiie
aoee ca?a?no?iaai? eeoa o 3o c 7ie iae?ii?a, oiaoi eiia??iei iaoai?ciii
aaia?aoe?? oaeeo o?aiceaio?a aoea aeoeaaoe?y eiiio?iiieo ?aoeaioi??a, a
oie /an ye ooieoe?y iaoaaio?iiieo ?aoeaioi??a, eiia??ii,
i?eai?/oaaeanue.

Oaae.5

Ci?ie eaeueoe??aeo o?aice?io?a, aeeeeeaieo o oaeai?/ieo oa ei?eiaeo
iae?iiao aie?eaoe??th 0,5 iI AOO.

————————————————————————
—-

eiio?. oaa?eie
NOC-ae?aaao

oaeai?/i? iae?iie

i?eiaa aiie?ooaea (iI) 118 ± 35 iI 100 ± 55 iI

e?euee?noue aeoeaieo ee. 11 c 16oe (69%) 3 c 11oe
(27%)

?<0,01 ------------------------------------------------------------------------ ---- ei?eia? iae?iie i?eia? aiie?ooaea (iI) 134 ± 31 iI 122 ± 20 iI e?euee?noue aeoeaieo ee. 13 c 18oe (72%) 7 c 12oe (58%) 0,1

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