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Ультраструктурні еквіваленти ендокринної функції серця хребетних в нормі та патології: Автореф. дис… канд. біол. наук / Т.Р. Скибінська, Нац. мед. у

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I?I?NOA?NOAI IOI?IIE CAeI?IA’ss OE?A?IE

IAOe?IIAEUeIEE IAAeE*IEE OI?AA?NEOAO ?i.I.I.Aiaiiieueoey

NEEA?INUeEA OAOssIA ?INOENEAA?AIA

OAeE
611.12-013+611:577.95:616.12

OEUeO?ANO?OEOO?I? AEA?AAEAIOE

AIAeIE?EIII? OOIEOe?? NA?Oess O?AAAOIEO

A II?I? OA IAOIEIA??

14.03.09.- a?noieia?y, oeeoieia?y, aia??ieia?y

Aaoi?aoa?ao aeeca?oaoe??

ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa a?ieia?/ieo iaoe

Ee?a-1999

Aeena?oaoe?y ? ?oeiienii

?iaioa aeeiiaia a eaai?aoi??? aeaeo?iiii? i?e?ineii??
Iaoeiai-aeine?aeiiai eaai?aoi?iiai oeaio?o Iaoe?iiaeueiiai iaaee/iiai
oi?aa?neoaoo ?i.I.I.Aiaiiieueoey

Iaoeiaee ea??aiee: aeieoi? a?ieia?/ieo iaoe, i?ioani? Noa/aiei Ethaeieea
Ieaenaiae??aia, caa?aeoth/a eaai?aoi???th aeaeo?iiii? i?e?ineii??
Iaoeiai-aeine?aeiiai eaai?aoi?iiai oeaio?o Iaoe?iiaeueiiai iaaee/iiai
oi?aa?neoaoo ?i. I.I.Aiaiiieueoey

Io?oe?ei? iiiiaioe: aeieoi? iaaee/ieo iaoe, i?ioani? Eieii?eoeaa
Aiae??e Einoyioeiiae/, caa?aeoaa/ eaoaae?e a?noieia?? Iaaee/iiai
?inoeoooo Oe?a?inueei? anioe?aoe?? ia?iaeii? iaaeeoeeie

aeieoi? a?ieia?/ieo iaoe, i?ioani?
Iaethe A?eoi? ?aaiiae/, i?ioani? eaoaae?e aeieia?? Iaoe?iiaeueiiai
oi?aa?neoaoo “Ee?ai-Iiaeeyinueea aeaaeai?y”

I?ia?aeia onoaiiaa: Oa?iii?euenueea aea?aeaaia iaaee/ia aeaaeai?y

?i. ?.ss. Ai?aa/aanueeiai IIC Oe?a?ie,

eaoaae?a a?noieia??, oeeoieia?? oa
aia??ieia??, i. Oa?iii?eue

Caoeno a?aeaoaeaoueny “ 21 ” o?aaiy 1999 ?. i 14-00 aiaeei? ia
can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae.26.003.04 Iaoe?iiaeueiiai
iaaee/iiai oi?aa?neoaoo ?i.I.I.Aiaiiieueoey, 252057, Oe?a?ia, i.Ee?a-57,
i?.Ia?aiiae, 34, ii?oieia?/iee ei?ion, aoae. N 3.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? Iaoe?iiaeueiiai
iaaee/iiai oi?aa?neoaoo ?i.I.I.Aiaiiieueoey, 252057, i.Ee?a, aoe.
Ciieia?/ia, 1.

Aaoi?aoa?ao ?ic?neaiee “ 20 ” ea?oiy 1999 ?.

A/aiee nae?aoa?

Niaoe?ae?ciaaii? a/aii? ?aaee

aeieoi? iaaee/ieo iaoe, i?ioani?
A.A.*a?eania

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue oaie. Aeine?aeaeaiiy inoaii?o ?ie?a aeathoue i?aenoaae
noaa?aeaeoaaoe, ui na?oea ia? aenieiaoaeoeaio ai?iiiaeueio nenoaio
?aaoeyoe?? ye ooieoe?? na?oeaai-noaeeiii? nenoaie, oae ? i?aeo?eiaiiy
caaaeueiiai aiiainoaco i?aai?cio. ? aaaii? aeieace ooieoe?iioaaiiy a
na?oe? aeanii? aiaeie?eiii?, ia?ae?eiii? oa aooie?eiii? ai?iiiaeueieo
nenoai. Aaaaoi oaaae i?eae?ey?oueny inoaii?i /anii aea/aiith ?iaioe
oeaieiii? ai?iiiaeueii? nenoaie na?oey, i?iaeoeo ae?yeueiino? yei?
ia?aaena?aeiee iao??eo?aoe/iee iaioeae (IIOI), neioaco?oueny a
i?iea?ae? ia?aaena?aeue oa aooie na?oey.

Ii?oieia?/iei aea?aaeaioii aiaeie?eiii? ai?iiiaeueii? nenoaie na?oey ?
niaoeeo?/i? ia?aaena?aei? a?aioee, aia?oa iienai? a 1956 ?ioe? E?oai.
?icoeo?iaea o?i?/ii? i?e?iaee a?aioe, a naia oa, ui aiie i?noyoue
ia?aaena?aeiee iao??eo?aoe/iee iaioeae [De Bold A.I. et al., 1981],
aaaaoi a /iio aeicaieeea i?iynieoe ?ieue na?oey ye aiaeie?eiiiai i?aaio,
yeee i?e aeiiiiic? oeueiai ai?iiio cae?enith? ?aaoeyoe?th ?yaeo aeeoo?ai
aaaeeeaeo ooieoe?e a i?aai?ci? oeyoii aca?iiae?? c iaeaaaeeea?oeie
?aaoeyoi?ieie nenoaiaie: ?ai?i-aia?ioaicei-?ai?i-aeueaeinoa?iiiaith,
neiiaoiaae?aiaeiaith, eae?e?a?i-e?i?iiaith [Aeaenaaiei E.I., I?aoiae/
A.I, 1987; Iinoiia A.TH., 1987; Iieaaiei A.A., Iaaeth/aiei A.A., 1990;
Lang L.E. et al., 1985; Needleman Ph. et al. , 1985; De Bold A. et al. ,
1991; Tummala P. et al., 1992 Anderson S., Vora J.P., 1995; Hirata Y.
et al.,1995; Komers R., 1996; Zietse R. et al., 1997; Chattington P.D.
et al., 1998].

I?aeoaa?aeaeaiiyi ooieoe?iioaaiiy aiaeie?eiii? oa ia?ae?eiii?
ai?iiiaeueieo nenoai a na?oe? aoei aeyaeaiiy i?iea?ae?aeueii?
?ai?i-aia?ioaiceaii? nenoaie, a oaeiae caeaoiino? aiaeioae?th
ei?iia?ieo noaeei ? aaiii?e?ioee?eoeyoi?iiai ?onea na?oey neioacoaaoe
oe?eee ?yae aaciaeoeaieo ?a/iaei.

I?e ee?i?/ieo aei?iaiaoaaiiyo aoei iieacaii, ui an? o? o?c?ieia?/i?
aoaeoe IIOI, ye? aeyaeai? a aeine?aeao ia oaa?eiao, niinoa??aathoueny ?
o ethaeeie [Eaioai I., AEaia AE., 1986]. Oiio aeai? i?i
oeueo?ano?oeoo?i? i?iyae aiaeie?eiii? ooieoe?? i?iea?aea na?oey a o?ei-
oa iioiaaiac? aeiiiiiaeooue ?ice?eoe ia o?eueee iniaeeaino? ?icaeoeo ?
aoaeiae no?oeoo?, a?aeiia?aeaeueieo ca oeth ooieoe?th, aea e
ni?eyoeiooue e?auiio ?icoi?iith oeo ii?oi-ooieoe?iiaeueieo ci?i, ye?
iiaeooue aooe iane?aeeii iioeiaeaeaiiy na?oeaaeo i?ioeeo?a.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie.

Aeena?oaoe?y aeeiiaia a ?aieao aaeocaai? iaoeiai? i?ia?aie
Iaoeiai-aeine?aeiiai eaai?aoi?iiai oeaio?o Iaoe?iiaeueiiai iaaee/iiai
oi?aa?neoaoo IIC Oe?a?ie ii oai?: “Aea/eoe no?oeoo?i? iniaeeaino?
aiaeie?eiii? ooieoe?? na?oey a oiiaao eaaaiaai? a?ia?oaic??”, iiia?
aea?aeaaii? ?a?no?aoe?? 0194EI20877.

Iaoa ? caaea/? aeine?aeaeaiiy. Iaoa ?iaioe – anoaiiaeoe iniaeeaino?
noaio oa ci?i oeueo?ano?oeoo?ieo aea?aaeaio?a aiaeie?eiii? ooieoe??
na?oey a o?ei-, iioiaaiac? o?aaaoieo oa i?e aeayeeo iaoieia?/ieo noaiao.

Aeey aeinyaiaiiy aeacaii? iaoe iinoaaeai? oae? caaea/?:

1. Aea/eoe noai ia?aaena?aeieo a?aioe (ii?oieia?/iiai aea?aaeaioa
aiaeie?eiii? ooieoe?? na?oey) a o?eiaaiaoe/iiio ?yaeo o?aaaoieo ca
aeaieie oeueo?ano?oeoo?iiai oa noa?aieia?/iiai aiae?co.

2. Anoaiiaeoe caeiiii??iino? oeueo?ano?oeoo?ieo ci?i ia?aaena?aeieo
a?aioe a iioiaaiac? uo?a.

3. Aecia/eoe oeueo?ano?oeoo?i? iniaeeaino? aea?aaeaio?a aiaeie?eiii?
ooieoe?? na?oey a oiiaao na?oeaai-noaeeiii? iaoieia?? a ee?i?oe?
(a?iaeaeaiee aeaoaeo i?aeoeoii/eiai? ia?aai?iaeee, oneeaaeiaiee
a?ia?oaic??th o iaeiio eie? e?iaiia?ao) oa aenia?eiaio? (eaaaiaaa
a?ia?oaic?y, aaaaeaiiy iiia?aaeieea neioaco ieneaeo acioo (NO) –
L-a?a?ieio a oiiaao a?ia?oieanoa?eiai??).

4. Aeaoe ii??aiyeueio oa?aeoa?enoeeo ci?i ii?oieia?/ieo aea?aaeaio?a
aiaeie?eiii? ooieoe?? na?oey a o?ei-, iioiaaiac? o?aaaoieo oa i?e
iaoieia??.

Iaoeiaa iiaecia io?eiaieo ?acoeueoao?a. Aeei?enoaiiy a ?iaio?
?noi?e/iiai oa aenia?eiaioaeueiiai i?aeoiae?a, iiaeeeainoae
o?aini?n?eii? aeaeo?iiii? i?e?ineii?? oa i?eioeei?a ii?oiiao?e/iiai
aiae?co iaoa??aeo aeicaieeei io?eiaoe i?eioeeiiai iia? aeai?, ye?
?noioii aeiiiaiththoue oa ?icaeaathoue oyaeaiiy i?i nae?aoi?io ooieoe?th
na?oey:

Aia?oa anoaiiaeai? ?ici??i? oa e?euee?ni? iniaeeaino? o?ueio oei?a
ia?aaena?aeieo a?aioe, ye? a?aeiia?aeathoue ??ciei noaae?yi nae?aoi?iiai
oeeeeo, o ??cieo i?aaenoaaiee?a o?aaaoieo.

Aia?oa aea/ai? ci?ie ?ici???a, oi?ie ? ia?a?iciiae?e a?aioe a
iioiaaiac? uo?a, ui aeicaiey? ioa?aeoa?ecoaaoe iai?aaeaiiy i?ieeaino?
aiaeie?eiii? ooieoe?? na?oey ia ??cieo aoaiao ?iaeea?aeoaeueiiai
?icaeoeo.

C’yniaaiee noai oeueo?ano?oeoo?ieo aea?aaeaio?a aiaeie?eiii? ooieoe??
na?oey (ia?aaena?aeieo a?aioe ea?ae?ii?ioeeo?a ? nae?aoi?ieo a?aioe
aiaeioae?ioeeo?a) i?e ee?i?/i?e oa aenia?eiaioaeuei?e a?ia?oaic??.

I?aaenoaaeai? aeieace ca?eueoaiiy ?ici???a ? ia?a?iciiae?e oei?a a?aioe
a oiiaao canoinoaaiiy L-a?a?i?io ye iiia?aaeieea neioaco ieneaeo acioo
(NO) a oiiaao a?ia?oieanoa?eiai??.

Anoaiiaeai? iciaee ?aeai?ooeyoe??, ye? oa?aeoa?ecothoue noai
ia?aaena?aeieo a?aioe a iioiaaiac?, a oaeiae a oiiaao ee?i?/ii? oa
aenia?eiaioaeueii? iaoieia?? na?oey.

I?aeoe/ia cia/aiiy io?eiaieo ?acoeueoao?a. Ia i?aenoaa? eiiieaeniiai
aeaeo?iiiii?e?ineii?/iiai oa ii?oiiao?e/iiai aiae?co o?eiaaiaoe/iiai,
iioiaaiaoe/iiai, aenia?eiaioaeueiiai ? a?iin?eiiai iaoa??aeo
i?aaenoaaeaii ocaaaeueiaiiy i?i oeueo?ano?oeoo?i? aea?aaeaioe
aiaeie?eiii? ooieoe?? na?oey a ii?i? ? iaoieia??; ioa?aeoa?eciaaii yaeua
?aeai?ooeyoe?? ia?aaena?aeieo a?aioe a o?ei-, iioiaaiac? oa i?e
iaoieia??; c’yniaaii ? e?euee?nii aecia/aii caeiiii??iino?
ia?a?iciiae?eo a?aioe a iioiaaiac? oa i?e iaoieia??. Io?eiai?
?acoeueoaoe i?aaenoaaeythoue iiaeeea?noue iiooeo iiaeo cania?a
i?io?eaeoeee ? e?eoaaiiy a?ia?oaic?? a iaeiio eie? e?iaiia?ao.
?acoeueoaoe aeine?aeaeaiiy iiaeooue aooe aeei?enoai? a o/aiaiio i?ioean?
ia eaoaae?ao a?noieia??, oeeoieia?? oa aia??ieia??, ii?iaeueii? ?
iaoieia?/ii? o?c?ieia??, ea?ae?ieia??, aiaeie?eiieia??.

Iniaenoee aianie caeiaoaa/a. Aaoi?ii naiino?eii i?iaaaeaiee aiae?c
e?oa?aoo?ieo aeaea?ae, aeeiiai? aeaeo?iiiii?e?ineii?/i? oa ii?oiiao?e/i?
aeine?aeaeaiiy, i?iaaaeaia noaoenoe/ia ia?iaea, aiae?c ? ocaaaeueiaiiy
io?eiaieo ?acoeueoao?a. Aenia?eiaioe ii iiaeaethaaiith
a?ia?oieanoa?eiai?? o e?ie?a oa aaaaeaiiy ia ?? oii? L-a?a?i?io aaoi?ii
i?iaaaeaii ni?eueii c i?a?ia?oieeaie a?aeae?eo o?c?ieia?? e?iaiia?ao
?inoeoooo o?c?ieia?? ?i. I.I.Aiaiiieueoey IAI Oe?a?ie (ea??aiee /e-ei?.
IAI Oe?a?ie A.O.Naaa/a.)*

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. Iaoa??aee aeena?oaoe?eii? ?iaioe
aeiiia?aeaeenue ? iaaiai?thaaeenue ia III c’?cae? aiaoii?a, a?noieia?a,
aia??ieia?a oa oiiia?aoiaiaoii?a O?N? (*a?i?aoe?, 1990 ?.); iaoeia?e
eiioa?aioe?? “I?eiaia-iea yeaeo?iiiie iee?ineiiee a iaaeeoeeia” (Ee?a,
1992 ?.); i?aeaoc?anueeiio neiiic?oi?, i?enay/aiiio 30-??//th IAeEOe
IIO (Ee?a, 1992 ?.); iaoeia?e eiioa?aioe?? “Aeooaeuei? ieoaiiy
ii?oieia??” (?aaii-O?aie?anuee, 1994 ?.); III eiioa?aioe?? anioe?aoe??
na?oeaai-noaeeiieo o??o?a?a Oe?a?ie (Ee?a, 1995 ?.); iaoeiaiio
neiiic?oi? “Aeooaeueiua aii?inu ii?oieiaee na?aeoea” (Aei?i?iiao?ianuee,
1996 ?.); ae?ci?e nan?? Oe?a?inueei? aeaaeai?? iaoe iaoe?i-iaeueiiai
i?ia?ano (Oa?iii?eue, 1997 ?.); eiioa?aioe?? “Ii?oieia?y – aaoa?eia???
? iaaeeoeei?” (A.Oea?eaa,1998 ?.); 3-?e iaoeiai-i?aeoe/i?e eii-oa?aioe??
“Aeooaeuei? i?iaeaie aenia?eiaioaeueii? iaaeeoeeie” (Ee?a, 1999 ?.).

Ioae?eaoe??. Ca iaoa??aeaie aeena?oaoe?eii? ?iaioe iioae?eiaaii 12
?ia?o, ?c ieo 3 noaoo? a i?io?eueieo aeo?iaeao.

No?oeoo?a oa ia’?i aeena?oaoe??. Aeena?oaoe?y neeaaea?oueny ?c anooio,
iaeyaeo e?oa?aoo?e, iieno iaoa??aeo ? iaoiae?a aeine?aeaeaiiy, ?icae?eo
?acoeueoao?a aeanieo aeine?aeaeaiue, ?icae?eo aiae?co ? ocaaaeueiaiiy
?acoeueoao?a aeine?aeaeaiiy, aeniiae?a oa nieneo aeei?enoaieo aeaea?ae
(225, ?c ieo 191 – ?iicaiieo). ?iaioa aeeeaaeaia ia 152 noi??ieao,
aeeth/a? 5 oaaeeoeue, ?ethno?iaaia 47 ?enoieaie.

INIIAIEE CI?NO ?IAIOE

Iaoa??ae ? iaoiaee aeine?aeaeaiiy.

Iaoa??aeii aeine?aeaeaiiy aoee eoni/ee i?iea?aea ??ciiiai?oieo
i?aaenoaaiee?a o?eiaaiaoe/iiai ?yaeo. Aea/aii i?iea?ae ?ea (Pisces):
ei?ii

______________________________________

* Aaoi? ue?i aaey/ia ni?a?ia?oieeai iacaaiiai a?aeae?eo ca iino?eio
oaaao ? aeiiiiiao o ni?eueieo aeine?aeao.

(Cyprinus carpio) – 5; caiiiaiaeieo (Amphibia): ica?ia aeaaa (Rana
ridibunda) – 5; ?aioee?e (Reptilia): yu??ea (Lacerta agilis L.) – 3,
/a?aiaoa (Testudo g.) – 2; ioao?a (Aves): aeiiaoiy eo?ea (Gallus
domesticus) – 5; nnaaoe?a (Mammalia): uo? e?i?? A?noa? (Rattus w. ) – 5.
Onueiai aeine?aeaeaii 25 i?aaenoaaiee?a o?aaaoieo. Aea/ai? ??ci?
?aa?iie na?oey – ia?aaena?aey ? oeoii/ee.

Aeey c’ynoaaiiy noaiiaeaiiy oeueo?ano?oeoo?ieo aea?aaeaio?a
aiaeie?eiii? ooieoe?? na?oey a iioiaaiac? aeine?aeaeoaaeiny i?aaa
ia?aaena?aey ? i?aaee oeoii/ie a?ae 15-oe eaai?aoi?ieo uo??a o
??ciiiai?oi? ia??iaee ?iaeea?aeoaeueiiai ?icaeoeo: 20-oe aeiaiaee
aia??ii, 1-ii, 7-ie, 30-oe aeiaiaee ? 6-oe i?ny/iee uo?.

Aeey aea/aiiy eiiiainaoi?ieo iiaeeeainoae na?oey, iia’ycaieo ?c eiai
aiaeie?eiiith ooieoe??th, ? aecia/aiiy aeayeeo ?aeai?ooeyoe?eieo iciae
o?eiaaiaco a iaoieia?? aeine?aeaeaii i?iea?ae i?e aeayeeo
na?oeaai-noaeeiieo caoai?thaaiiyo. Aea/ai? a?iioaoe i?aaiai aooea
na?oey oai?eo, ye? no?aaeaeaee a?iaeaeaieie aaaeaie na?oey. Oe? ia’?eoe
io?eioaaee a Ee?anueeiio IAe? na?oeaai-noaeeiii? o??o?a??, aea oai??
i?ioiaeeee iaiao?aeia caaaeueiiee?i?/ia oa niaoe?aeueia ianoaaeaiiy
(AEA, OEA, OCAe, aiaeiaaneoey?ia ciiaeoaaiiy, a?io?i?/ia aeine?aeaeaiiy
aac?a e?ia? e ?i.). Iaoa??ae ocyoee a?ae 9 oai?eo c a?iaeaeaieie
aeaoaeoaie i?aeoeoii/eiai? ia?aai?iaeee, oneeaaeiaieie a?ia?oaic??th o
iaeiio eie? e?iaiia?ao ??ciiiai?oieo a?eiaeo a?oi (a?ae 6 i?nyoe?a aei
1 ?ieo, a?ae 1,5 aei 4 ?ie?a, a?ae 4 ?ie?a ? a?eueoa) ii 3 o eiaei?e
a?oi?. Eiio?ieai neoaeea iaoa??ae a?ae 3-o oai?eo c a?iaeaeaieie aaaeaie
na?oey aac eaaaiaai? a?ia?oaic??.

Aenia?eiaioaeueio a?oio oaa?ei neeaaeaee niaaee c eaaaiaaith
a?ia?oaic??th, iiaeaethaaiiy yei? cae?enithaaee oeyoii i?e?iaiaie?caoe??
noaeei eaaaiue aaoa?iaaiieie a?eo?ioeeoaie (5-8 ie/ea 50% noniaic??
a?aeieoeo eo?y/eo a?eo?ioeeo?a ia o?c?ieia?/iiio ?ic/ei?). Aea/ai? i?aaa
ia?aaena?aey ? i?aaee oeoii/ie na?oey 5-oe niaae /a?ac 6 i?nyoe?a i?ney
ii/aoeo aenia?eiaioo oa 3-o eiio?ieueieo oaa?ei.

Aenia?eiaioaeueia a?ia?oieanoa?eiai?y iiaeaethaaeany ia 3-o e?ieeeao
ii?iaee oeioeeea oeyoii aaaaeaiiy ia i?ioyc? 4 i?nyoe?a /a?ac ciiae
uiaeiy 0,2 a ?ic/eiaiiai a niiyoieeia?e ie?? oieanoa?eio ia 1 ea iane
oaa?eie. ?ioo a?oio neeaaeaee 3 oaa?eie, ye? e??i aoa?iaaiii? ae??oe
iaea?aeoaaee eiaeiee aeaiue i?ioyaii 4 i?nyoe?a aioo??oiueiaaiii 25
ia/ea L-a?a?i?io (o??ia “Sigma”, NOA). E??i oiai aeine?aeaeoaaeany
a?oia a?aa??eiiai eiio?ieth (3 oaa?eie). Aiae?coaaany i?iea?ae i?aaiai
ia?aaena?aey.

Iaoa??ae o?enoaaee a 1% ?ic/ei? /ioe?ueioieeno ini?th ca Eieo?eueaeii
i?ioyaii aeaio aiaeei ia oieiae?. Ia’?eoe ciaaiaeithaaee a aoaiie?
c?inoath/i? eiioeaio?aoe??, a aoeaoii? ? caeeaaee a noi?o aiiio c
a?aeae?oii ca caaaeueiii?eeiyoith noaiith [Ea?oio A.ss., 1984].
Oeueo?aoiie? c??ce iaea?aeoaaee ia oeueo?aoiiao LKB ? Reichert,
eiio?anooaaee ?o 2% ?ic/eiii o?ai?eaoeaoaoo ia 70% aoaiie? i?ioyaii
oa oeeo?aoii naeioeth ? iio?i aea/aee a aeaeo?iiiiio i?e?ineii?
AIA-100A?, AI-125E. C iaoith e?euee?nii? ioe?iee ci?i ia?aaena?aeieo
a?aioe na?oey o o?ei-, iioiaaiac? ? i?e aeayeeo iaoieia?/ieo noaiao
i?iea?aea, canoiniaaiee iaoiae ii?oiiao?e/iiai aiae?co, a iniiao yeiai
iieeaaeai? i?eioeeie noa?aiiao??? [Aaeaaeue, 1970; Aaoaiaeeeia, 1979,
1980, 1984]. Ii?oiiao?e/iee aiae?c i?iaiaeeany ca aeiiiiiaith
niaoe?aeueii ?ic?iaeaii? eiii’thoa?ii? i?ia?aie “I?aaiaea” ia
iai?aaaoiiaoe/iiio i?eno?i? ia?iaee a?ao?/ieo cia?aaeaiue. Aecia/ai?
oae? iieacieee ye ia?eiao?, ieiua, oaeoi? oi?ie niaoeeo?/ieo
ia?aaena?aeieo a?aioe I, II ? III oei?a, a oaeiae ?oi? a?aenioeiaa
ni?aa?aeiioaiiy. Ii eiaeiiio ?c ia?ae?/aieo iieaciee?a aecia/aeenue
na?aaei? cia/aiiy aaee/eie, na?aaei? eaaae?aoe/ia a?aeoeeaiiy,
iioeaea na?aaeiuei?, eiao?oe??io aa??aoe?? oa eiao?oe??io oi/iino?
aecia/aiiy na?aaeiuei? ie?aii aeey eiaeii? aeine?aeaeoaaii? oaa?eie.
E?euee?noue a?aioe i?ae?aoiaoaaeanue ia 10 noaiaea?oieo o?aaiaioao
ea?ae?ii?ioeeo?a aeey eiaeiiai ia’?eoo. Noaoenoe/ia ia?iaea io?eiaieo
?acoeueoao?a cae?enithaaeanue a?aeiia?aeii ?aeiiaiaeaoe?e, ye?
i?noyoueny a iin?aieeao c iaoaiaoe/ii? noaoenoeee [O?aao A.TH., 1964;
Eaeei A.A., 1973]. Ii??aiyiiy ?acoeueoao?a cae?enithaaeinue ca
aeiiiiiaith ia?aiao?e/iiai e?eoa??th Nouethaeaioa. Ii – a?iioaca i?i
a?aenooi?noue eiia??ieo ?ica?aeiinoae i?ae ii??aiyeueieie na?aaei?ie
a?aeoeeyeany i?e ? 0,05.

?acoeueoaoe aeine?aeaeaiiy oa ?o iaaiai?aiiy

Ii??aiyeueiee aiae?c oeueo?ano?oeoo?e i?iea?aea na?oey a o?eiaaiac?

Io?eiai? ?acoeueoaoe na?ae/aoue, ui o i?iea?ae? an?o aea/aieo iaie
i?aaenoaaiee?a o?eiaaiaoe/iiai ?yaeo o?aaaoieo i?enooi? nei?i/oaaeuei?
i?ioeeoe, ui i?noyoue niaoeeo?/i? nae?aoi?i? a?aioee, a?aeii? a
e?oa?aoo?? ye ia?aaena?aei? [B.Kisch, 1956]. O ?ea, aio?a?e, ?aioee?e
oa ioao?a oae? ea?ae?ii?ioeeoe aeyaeythoueny ? a ia?aaena?aeyo, ? a
oeoii/eao, aea ia?aaaaeia ?o a?eueo?noue cina?aaeaeaia a ia?aaena?aeyo.
O nnaaoe?a, aeeth/ath/e ethaeeio, a?aioee a ii?i? aeyaeythoueny o?eueee
a ia?aaena?aeyo ? aooeao na?oey.

E?euee?noue a?aioe a ee?oeiao aa??th? – aeaye? c ieo iathoue cia/io
e?euee?noue a?aioe, oiae? ye ?io? i?noyoue iiiaeeiie? a?aioee. A?aioee
?ici?uai? ii on?e oeeoiieaci? ee?oeie: i?ae i?io?a?eeaie, a
i?aena?eieaiaeuei?e cii?, aea a?eueoa ?o /anoeia ciaoiaeeoueny a cii?
eiiieaeno Aieueaeae?.

Ca oeueo?ano?oeoo?iith i?aai?caoe??th na?aae iiioeyoe?? a?aioe iiaeia
aeae?eeoe o?e oeie. Ia?oee oei oa?aeoa?eco?oueny aenieith aeaeo?iiiith
u?euei?noth ? aiiiaaieciaaiei iao?eenii, yeee ioi/aiee iaia?aiith
? aeaeo?iiiii?ici?ei iaaeieiiaia?aiiei ia?aeeii. A?aioee ae?oaiai
oeio i?noyoue iaio aeaeo?iiiiu?eueiee iao?een, iathoue iaia?aio,
i?ioa iaaeieiiaia?aiiue ia?aeie i?aeoe/ii a?aenooi?e. A?aioee o?aoueiai
oeio i?noyoue iaea?eueo aeaeo?iiiii?ici?ee iao?een, ciai?oiy iaia?aia
a ieo a?aenooiy. Oe? oeie a?aioe a?aeia?aaeathoue iine?aeiai? noaae??
nae?aoe?? ai?iiio (IIOI). Ia?oee oei ie a?aeiineii aei iieiaeeo a?aioe,
ui noi?ioaaeeny. Ae?oaee – oea c??e? no?oeoo?e, o?ao?e oei – a?aioee, ui
aeeooiaeothoue, oiaoi oae?, ui aeaiaeyoueny c ee?oeie oeyoii aeeooc??.

E?euee?nia ni?aa?aeiioaiiy o?ueio oei?a a?aioe o ??ciiiai?oieo
i?aaenoaaiee?a o?aaaoieo ia? nai? aeaeia? iniaeeaino? (oaae.1) ?,
eiia??ii, iiaea a?aeia?aaeaoe ni?aa?aeiioaiiy i?ioean?a neioaco,
iaeiie/aiiy ? aeaaaeaiiy iaioeaeo c ee?oeie.

Oaaeeoey 1

Noa?aieia?/i? iieacieee ia?aaena?aeieo a?aioe ea?ae?ii?ioeeo?a

o ??ciiiai?oieo i?aaenoaaiee?a o?aaaoieo

Aeae

oaa?eie Oeie a?aioe Ia?aaena?aey Oeoii/ieNa?aaeiy ieiua,

10-2 iei2 Oaeoi? oi?ie E-noue a?aioeNa?aaeiy ieiua,

10-2 iei2Oaeoi? oi?ieE-noue a?aioe % Aani-ethoia%Aani-ethoia Ea?i I
2,14? 0,25 0,85? 0,02172,53? 0,240,81? 0,0119 II 1,89?
0,180,79?0,0147462,93?0,350,87?0,015234III1,94?0,290,83?0,01362,24?0,500
,75?0,0129AEaaa
I2,31?0,090,86?0,01302,48?0,200,91?0,0110II1,99?0,100,82?0,01303472,26?0
,300,91?0,014397III1,78?0,070,79?-,01401,69?0,110,79?0,0247ssu??eaI1,93?
0,200,75?0,03551,66?0,370,81?0,0448II2,11?0,210,89?0,0122513,94?0,070,91
?0,021918III2,34?0,440,79?0,05231,58?0,250,89?0,0433O?eoiiI1,78?0,210,85
?0,02491,47?0,090,92?0,0158II1,63?0,220,85?0,0333631,46?0,160,92?0,01423
3III3,16?0,650,91?0,0218—*a?aiaoaI1,40?0,130,74?0,05452,66?0,430,78?0,
0957II1,48?0,210,73?0,0629382,84?0,580,93?0,011414III1.50?0,240,81?0,032
63,38?0,240,93?0,0229Eo?eaI2,89?0,160,89?0,02262,04?0,200,85?0,0424II2,3
2?0,320,89?0,0127412,59?0,310,88?0,013528III2.15?0,130,91?0,01471,76?0,2
40,90?-,0141Uo?I1,89?0,160,89?0,0227—II2,72?0,360,89?0,012941—III1.9
5?0,170,91?0,0144—NiaaeaI2,09?0,150,90?0,0218—II2,59?0,260,91?0,0135
61—III1.78
?0,110,89?0,0146—E?ie?eI3,27?0,200,89?0,0221—II2,98?0,210,89?0,01387
8—III2,76?0,200,91?0,0141—

sse aeaeii ?c a?noia?aie (?en.), o ?ea, aeaa, ioao?a ? nnaaoe?a ia?oee
oei a?aioe neeaaea? i?eaeecii iaeio o?aoeio an??? iiioeyoe??,
oiae? ye a?aenioeiaa

EA?I

AEAAA

ssU??EA

O?EOII

*A?AIAOA

EO?EA

NIAAEA

UO?

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0000000000?en. ?iciiae?e I, II ? III oei?a (a %) niaoeeo?/ieo
ia?aaena?aeieo a?aioe a ea?ae?ii?ioeeoao ia?aaena?aeue ? oeoii/e?a
na?oey o ??cieo i?aaenoaaiee?a o?aaaoieo

ni?aa?aeiioaiiy ae?oaiai ? o?aoueiai oei?a o ??cieo i?aaenoaaiee?a
o?aaaoieo aeaui ??ciyoueny. Aeii?ioaaiiy ae?oaiai oeio a?aioe a
ea?ae?ii?ioeeoao ?eae na?ae/eoue i?i ia?aaaaeaiiy i?ioean?a ?o
iaeiie/aiiy o oeueiai i?aaenoaaieea. O ioao?a ? nnaaoe?a i?ioeane
iaeiie/aiiy ? aeaaaeaiiy a?aioe i?eaeecii caaeainiaai?. O i?aaenoaaiee?a
?aioee?e – yu??ee, o?eoiia ? /a?aiaoe a ea?ae?ii?ioeeoao
ia?aaaaeathoue a?aioee ? oeio. E?euee?noue aea a?aioe o ee?oei?
iacia/ia, i?e/iio, /anoi a?aioee ? oeio niinoa??aathoueny a
i?aena?eieaiaeuei?e cii?. Oiio iiaeia i?eionoeoe, ui a ?aioee?e ii?yae
?c aeeooc??th IIOI aeaaaeaiiy oeueiai iaioeaeo a?aeaoaa?oueny ? oeyoii
aecioeeoico. A?aeiinii aaniethoii? e?eueeino? a?aioe iaiao?aeii
a?aei?oeoe, ui oi/a e?euee?noue ?o aa??th?, aea nooo?ai oeae iieaciee
ia a?ae??ciy?oueny a ??cieo i?aaenoaaiee?a o?aaaoieo. Aeaui ie?aii
noi?oue eean caiiiaiaeieo, aea e?euee?noue a?aioe a ea?ae?ii?ioeeoao
a??ia?aeii a?eueoa (iaeaea a 6-7 ?ac?a), i?ae a ?ioeo i?aaenoaaiee?a
o?aaaoieo.

O aiaeioae?? e?iaiiinieo i?e?inoaeei a on?o aea/aieo i?aaenoaaiee?a
o?aaaoieo niinoa??aathoueny a?aioee oeio Aaeaaey-Iaeeaaea ? a?aioee
ie?oaei? oi?ie ii nai?e no?oeoo?? iiae?ai? aei ia?aaena?aeieo a?aioe.

I?eioeeiiao iiae?ai?noue no?oeoo?ii? i?aai?caoe?? a?aioe o aeaeaei
noiy/eo o o?eiaaiaoe/iiio ?yaeo o?aaaoieo iiaeia iiynieoe yaeuaie
ia?aeae?cio, ui iia’ycai? c caaaeueiei iioiaeaeaiiyi ? ooieoe?iiaeueiei
i?ecia/aiiyi. A?aei?iiino? a o?eaaeino? oa ni?aa?aeiioaii? ??cieo oac
nae?aoi?iiai oeeeeo ? i?iyaii aeeaa?aaioiiai ?icaeoeo. I/aaeaeii,
aeaaa, yea i?enoiniaaia ? aei aiaeiiai, ? aei iacaiiiai ?nioaaiiy,
iio?aao? i?aeaeuaii? aiaeie?eiii? ooieoe?? na?oey.

Ii??aiyeuei? ii?oieia?/i? aeai? ia?aeiiothoue oaeiae e o oiio, ui
o?eiaaiaoe/ii iaoiiaeaia ?aeiino?oeoe?y oeaio?aeueiiai i?aaio
e?iaiia?ao, iia’ycaia c ia?aoiaeii a?ae aeaieaia?iiai aei o?ueio- ?
aeae? – aei /ioe?ueioeaia?iiai na?oey, ui noi?iaiaeaeo?oueny ci?iith
aaiiaeeiai?ee, i?ecaiaeeoue aei c?inoaiiy aiaeie?eiii? ooieoe?? a
ia?aaena?aeyo, aea eiioeaio?o?oueny i?iaeoeoe?y niaoeeo?/ieo
ia?aaena?aeieo a?aioe. ?icaeoie /ioe?ueioeaia?iiai na?oey o o?aaaoieo
i?eaiaeeoue aei i?aeaeuaiiy oeneo a i?aaiio ia?aaena?ae? a ii??aiyii? c
aaiiciei na?oeai ?ea, ui o naith /a?ao iicia/a?oueny ia i?ioeanao
neioaco ? aeaaaeaiiy niaoeeo?/ieo ia?aaena?aeieo a?aioe, ye? i?noyoue
IIOI. Iiaeeeai, ui naia ci?ie aaiiaeeiai?ee iaoiiaeththoue oa, ui
ea?ae?ii?ioeeoe oeoii/e?a ia?anoathoue neioacoaaoe ia?aaena?aei?
a?aioee.

Aa?oi i?aee?aneeoe, ui ci?ia na?aaeiaeua ?nioaaiiy, iaa?oue oaea
?aaeeeaeueia ye aeo?ae ?c aiaee ia nooo, ?noioii ia aieeaa? ia
no?oeoo?io i?aai?caoe?th ia?aaena?aeieo a?aioe ea?ae?ii?ioeeo?a
o?aaaoieo, a ci?ith? ?o e?euee?nia i?aaenoaaieoeoai ? ia?a?iciiae?e ii
oeiao, oiaoi, i?ioeane nae?aoe?? IIOI.

Oeueo?ano?oeoo?i? aea?aaeaioe aiaeie?eiii? ooieoe?? na?oey a iioiaaiac?

A i?aiaoaeueiiio ia??iae? iioiaaiaco uo??a ea?ae?ii?ioeeoe, ui i?noyoue
ia?aaena?aei? a?aioee, i?enooi? o aooeao, ia?aaena?aeyo ? oeoii/eao. O
iinoiaoaeueiiio iioiaaiac? aiie aeyaeythoueny o?eueee o aooeao ?
ia?aaena?aeyo. sse ? o o?eiaaiac?, o ??ciiiai?oi? ia??iaee iioiaaiaco
na?aae ia?aaena?aeieo a?aioe iiaeia aeae?eeoe o?e oeie.

O 20-aeiaiaiai aia??iia ia?aaaaeia a?eueo?noue ea?ae?ii?ioeeo?a ? a
ia?aaena?aeyo, ? oeoii/eao i?noeoue niaoeeo?/i? ia?aaena?aei? a?aioee
an?o o?ueio oei?a. Ea?ae?ii?ioeeoe iathoue aaeeeo e?euee?noue a?aioe,
na?aae yeeo a?eueoa i?ae 50% neeaaeathoue a?aioee II oeio (oaae.2).
Oea na?ae/eoue i?i ia?aaaao i?ioean?a iaeiie/aiiy i?iai?iiio a ee?oeiao.

Oaaeeoey 2

Noa?aieia?/i? iieacieee ia?aaena?aeieo a?aioe ea?ae?ii?ioeeo?a

uo??a a iioiaaiac?

Oa?i?i niinoa?aaeaiueOeie a?aioeIa?aaena?aey Oeoii/ie Na?aaeiy ieiua,

10-2 iei2Oaeoi? oi?ie E-noue a?aioeNa?aaeiy ieiuaaeue,

10-2 iei2Oaeoi? oi?ie E-noue a?aioe%Aani-ethoia%Aani-ethoia 20-
aeiaiaee I2,39?0,200,87?0,0113 1,99? 0,35 0,83? 0,02 4aia??ii
II2,52?0,110,85?0,01521762,89?0,170,82?0,0164
118uo?aIII1,90?0,09*0,81?0,01352,03?0,150,82?0,02321-ii aeiaiaee
I2,43?0,270,85?0,048–uo?II4,05?0,43*,x0,91?0,0139119—III3,56?0,24*,x0
,91?0,0153—7-ie
aeiaiaeeI2,69?0,180,92?0,0145—uo?II4,07?0,56*0,91?0,022020
—III2,69?0,42*,x0,93?0,0135—1-ii i?ny/iee
I——uo?II5,23?0,050,91?0,0125201—III3,90?0,190,87?0,0175—6-oe
i?ny/iee
I1,89?0,160,89?0,0227—uo?II2,72?0,36*,x0,89?0,012941—III1,95?0,17×0,
91?0,0144—

O 1-ii aeiaiaiai uo?a ee?oeie c? niaoeeo?/ieie a?aioeaie aeyaeythoueny
aaea o?eueee a ia?aaena?aeyo. A ea?ae?ii?ioeeoao ciaioo?oueny
caaaeueia /enei

a?aioe ? nooo?ai i?aeaeuo?oueny e?euee?noue a?aioe III oeio, oiaoi
a?aioe, ui aeeooiaeothoue. Ca?a?no?iaaia iaie ??cea aea?eueiaiiy
a?aioe ?c ee?oei a?aeaoaa?oueny a ia??iae, eiee i?aeaeuo?oueny oene o
aaeeeiio eie? e?iaiia?ao, a ii?? oieo e?ia? oa oene o iaeiio eie?
nooo?ai cieaeo?oueny [Aieeiaa I.A., Iaea?neee I.E., 1976]. Ca?aeii
aeaieo e?oa?aoo?e a?aeiii, ui eiioeaio?aoe?y IIOI a ia?oocao? ia? i?yio
caeaaei?noue a?ae oeneo o i?aaiio ia?aaena?ae? [Ruskoaho H. et al.,
1986; Burnett J. et al., 1989], a i?aeaeuaiiy ??aiy ai?iiio, ui
oee?eoeth? a e?ia?, noi?iaiaeaeo?oueny ciaioaiiyi /enea a?aioe ? iaaiaee
[Choi K. C. et al., 1994; Baralis G. et al., 1997; Wu S. Q. et al.,
1998]. Oiio iiaeia i?eionoeoe, ui nooo?aa ciaioaiiy i?ney ia?iaeaeaiiy
e?eueeino? ia?aaena?aeieo a?aioe a ea?ae?ii?ioeeoao ia?aaena?aeue ?
iiaia ?o cieeiaiiy ?c ea?ae?ii?ioeeo?a oeoii/e?a iia’ycaia ia c? ci?iaie
oeneo o iaeiio eie? e?iaiia?ao, a nei??oa iiaea aooe iane?aeeii ?ioeo
oaeoi??a, oaeeo ye i?aeaeuaiiy oeneo o aaeeeiio eie? e?iaiia?ao aai
no?ano, yeiai cacia? i?aai?ci i?ae /an ia?iaeaeaiiy.

*- eiiia??ii ii a?aeiioaiith aei iiia?aaeiueiai oeio a?aioe (?

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