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Механізми гормон-медіаторного імпринтингу нейроендокринної регуляції репродукції та стрес-реактивності: Автореф. дис… д-ра біол. наук / Н.Д. Носенко

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3

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

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

IINAIEI Iaae?y Aeieo??aia

OAeE
612.822.43/.45.015.38:

612.6:616003.96

IAOAI?CIE AI?III-IAAe?AOI?IIAI ?II?EIOEIAO IAE?IAIAeIE?EIII?
?AAOEssOe?? ?AI?IAeOEOe??

OA NO?AN-?AAEOEAIINO?

14.01.14 aiaeie?eiieia?y

Aaoi?aoa?ao aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

aeieoi?a a?ieia?/ieo iaoe

Ee?a 1999

Aeena?oaoe??th ? ?oeiien

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

Iaoeiaee eiinoeueoaio

Aeieoi? iaaee/ieo iaoe, i?ioani?, /eai-ei?aniiiaeaio IAI oa AII
Oe?a?ie, caneoaeaiee ae?y/ iaoee ? oaoi?ee Oe?a?ie ?aci?eia Ieaenaiae?
A?eai?iae/, ?inoeooo aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A. I.
Eii?na?aiea AII Oe?a?ie, caa?aeoaa/ a?aeae?eo aiaeie?eiieia??
?ai?iaeoeoe?? oa aaeaioaoe??

Io?oe?ei? iiiiaioe:

aeieoi? iaaee/ieo iaoe I?eioa Ieaen?e Noaiaiiae/, ?inoeooo
aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A. I. Eii?na?aiea AII Oe?a?ie,
caa?aeoaa/ eaai?aoi??? ai?iiiaeueii? ?aaoeyoe?? iai?io ?a/iaei

aeieoi? a?ieia?/ieo iaoe, i?ioani? Eeiainueeee TH??e Iao?iae/, ?inoeooo
o?c?ieia?? ?i. I. I. Aiaiiieueoey IAI Oe?a?ie, caa?aeoaa/ a?aeae?eo
o?c?ieia?? noiaao?a iiceo

aeieoi? iaaee/ieo iaoe Oii?inueea C?ia?aea Ai?en?aia, ?inoeooo
iaae?ao???, aeooa?noaa oa a?iaeieia?? AII Oe?a?ie, caa?aeoth/a
eaai?aoi???th aiaeie?eiieia??

I?ia?aeia onoaiiaa

Oe?a?inueeee iaoeiai-aeine?aeiee ?inoeooo oa?iaeioa?ai?? aiaeie?eiieo
caoai?thaaiue IIC Oe?a?ie, i. Oa?e?a, eaai?aoi??y ?ai?iaeoeoeaii?
aiaeie?eiieia??

Caoeno a?aeaoaeaoueny 7 a?oaeiy 1999 ?. i 13 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 o a?ae?ioaoe? ?inoeoooo
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)

Aaoi?aoa?ao ?ic?neaiee “4” eenoiiaaea 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. Aiiiae?? ?iaeea?aeoaeueiiai ?icaeoeo i?aai?cio ?
eiai ie?aieo o?c?ieia?/ieo nenoai, o oiio /ene? ?ai?iaeoeoeaii? oa
aaeaioaoe?eii?, iiaeooue aeieeaoe aiane?aeie iaoiaaiiiai aieeao
ai?iiiaeueiiai oa iae?io?aini?ooa?iiai aeenaaeaino, no?ano oa ?ioeo
/eiiee?a, ui aieeaathoue ia i?aai?ci, yeee ?icaeaa?oueny, oiaoi
aioo??oiueioo?iaii aai iaaaeiac? i?ney ia?iaeaeaiiy [G. Dorner et al.,
1976, 1983; A.I. Aaae/aa, 1981; B. Jarzab et al., 1990; A.A. Iaoiaiei e
ae?., 1990: A.A. Oaeyieia e ae?., 1995; A.A. ?acieeia, 1994, 1998]. Oea
i?ecaiaeeoue aei oi?ioaaiiy iaoieia?/ii? ?aaeoeaiino?, ii?ooaiiy
noaoaai? iiaaae?iee oa aacie?aeaey. Iaeiae ?ii?eioeiaia? iaoai?cie
?iaeea?aeoaeueiiai ?icaeoeo i?aai?cio oa ?o ?ieue o aeieeiaii?
ooieoe?iiaeueieo ?iceaae?a ?ai?iaeoeoeaii? ooieoe?? oa aaeaioaoe?eieo
i?ioean?a aiane?aeie no?ano oa ai?iiiaeueiiai aeenaaeaino a
ia?eiaoaeueiee ia??iae iioiaaiaco, caeeoathoueny iac’yniaaieie.

Inoaii?ie ?ieaie ana i/aaeaei?oei noa? oa, ui a nenoaio i?aai?caoe?eiiai
aieeao noaoaaeo noa?i?ae?a ia ia?aia? no?oeoo?e, ui ?icaeaathoueny,
caeo/athoueny iae?iiaae?aoi?e. Ia?aeii? ii?ooaiiy iae?i-iaae?aoi?ieo
nenoai aieiaiiai iiceo caeaoi? iiaeeo?eoaaoe aieea noaoaaeo noa?i?ae?a ?
ni?e/eieoe no?ee? ?iceaaee iae?iaiaeie?eiii? ?aaoeyoe?? aiiaaeio?iiii?
nae?aoe?? ? noaoaai? iiaaae?iee [P. Sickmoller et al., 1985; C. Wilson
et al., 1986; A.A. ?acieeia, A.I. Aeai/aiei, 1987; B. Jarzab et al.,
1990;]. ? iia?aeiieaiiy i?i iayai?noue iaae?aoi?ieo c?ooaiue, eio??
aeieeathoue ye a?aeaeaeai? iane?aeee ai?iiiaeueieo aieea?a ia iicie, ui
?icaeaa?oueny [I.Ae. Iinaiei e ae?., 1976; E.E. Na?iaa, A.*. Iae, 1986;
I.I. Aeuaaei, A.A. Iaoiaiei, 1988]. ?acii ?c oei, ?ieue
iae?iiaae?aoi?ieo nenoai o ?aiiueiio ai?iiiaeueiiio ?ii?eioeiao
aeine?aeaeaia ae?ae iaaeinoaoiuei. Na?aae iaae?aoi?ieo c?ooaiue,
i?e/aoieo aei ai?iii-iaae?aoi?iiai ?ii?eioeiao, ia?aaaaeathoue ci?ie
eaoaoieai?ia?a?/ii? nenoaie. I?ioa caeeoa?oueny aeeneon?eiei ieoaiiy i?i
?ieue eaoaoieai?i?a (EA) o iaoai?ci? ?aae?caoe?? ai?iiiaeueieo aieea?a
ia iicie, ui ?icaeaa?oueny. Aei e?ioey ia c’yniaaii, /e
iiina?aaeeiaothoue EA aieea noaoaaeo noa?i?ae?a, /e ae?thoue ?acii ?c
ai?iiiaie. ?nio? i?eiouaiiy, ui noaoaaa aeeoa?aioe?aoe?y
iae?iaiaeie?eiieo iaoai?ci?a ?aaoeyoe?? ?ai?iaeoeoeaii? ooieoe??
ni?yiiaaia ia oi?ioaaiiy iaaiiai naen-niaoeeo?/iiai oeio
iiiiai?ia?a?/ii? ?aaoeyoe?? aieiaiiai iiceo. O oie aea /an, iaoai?cie
oi?ioaaiiy noaoaaeo a?aei?iiinoae o iae?iiaae?aoi?ieo nenoaiao iiceo oa
?o no?oeoo?ia eieae?caoe?y caeeoathoueny iaaea/aieie.

Ooiaeaiaioaeueio iniiao noaoaai? aeeoa?aioe?aoe?? iiceo (NAeI)
noaiiaeoue iaoaaie?/ia o?ainoi?iaoe?y noaoaaeo ai?iii?a o
iae?iaiaeie?eiieo no?oeoo?ao, yea cae?enith?oueny ca aeiiiiiaith
a?iiaoaciiai oa?iaioiiai eiiieaeno. Iaeiae ieoaiiy i?i ?ieue ano?iaaiieo
iaoaaie?o?a o iaoai?ciao ai?iii-iaae?aoi?ii? aca?iiae?? a ia??iae NAeI
caeeoa?oueny iaeaea iaaeine?aeaeaiei. Caeeoa?oueny iac’yniaaiith ?ieue
ano?iaaiieo iaoaaie?o?a, cie?aia eaoaoieano?iaai?a (EA), ? EA ye
?iaeoeoi??a aeic??aaiiy iae?iaiaeie?eiii? nenoaie ?aaoeyoe??
?ai?iaeoeoeaii? ooieoe??.

O aaaaoueio aeiaaeeao ii?ooaiiy NAeI iiaeooue aeieeaoe aiane?aeie no?ano
aai ai?iiiaeueiiai aeenaaeaino a ia??iae i?aiaoaeueiiai iioiaaiaco.
A?aeaeaeaieie aoaeoaie i?aiaoaeueiiai no?ano o ethaeeie oa oaa?ei ?
aeaianeoe?i?caoe?y oa/aai oai?i?caoe?y noaoaai? iiaaae?iee o naioe?a [I.
Ward, 1972; 1984; F. Gotz, G. Dorner, 1980] ? ineaaeaiiy oa?oeeueiino? o
naiie [L. Herrenkohl, 1978]. O ?icaeoeo oeeo oaiiiai?a aaaeeeao ?ieue
a?ae?a?athoue ?iaeoeiaai? no?anii ci?ie EA, noaoaaeo ?
aetheiei?oeei?aeieo ai?iii?a iaoa?? ? ieiaea [I. Ward, J. Weisz, 1980;
W. Rohde et al., 1983, 1984; R. Salisburg et al., 1984]. ?acii ?c oei,
ia aea/aia ieoaiiy i?i aieea i?aiaoaeueiiai no?ano oa ?aiiueiai
ai?iiiaeueiiai aeenaaeaino ia oi?ioaaiiy noaoaaiai aeeii?o?cio
iiiiai?ia?a?/ii? nenoaie a aeene?aoieo no?oeoo?ao iiceo, ye? i?e/aoi?
aei ?aaoeyoe?? ?ai?iaeoeoe?? oa ai?iiiaeueieo i?ioean?a aaeaioaoe??, a
ia??iae ?aiiueiai iinoiaoaeueiiai iioiaaiaco.

Oi?ioaaiiy aaeaioaoe?eieo iaoai?ci?a oaeiae cia/iith i??ith caeaaeeoue
a?ae oeo aieea?a, ui ?o cacia? i?aai?ci o ia??iae aioo??oiueioo?iaiiai
?icaeoeo. ? iecea iia?aeiieaiue i?i ?iaeoeiaai? i?aiaoaeueiei no?anii
ii?ooaiiy ia?aa?ao caaaeueiiai aaeaioaoe?eiiai neiae?iio, yeee
noi?iaiaeaeo?oueny ci?iaie ?aaeoeaiino?
a?iioaeaii-a?iio?ca?ii-iaaeie?eiai? nenoaie (AAIN) [A.A. Iaoiaiei, I.I.
Aeuaaei, 1979; E. Naumenko, L. Maslova, 1985; A.A. Iaoiaiei e ae?.,
1990; M. Weinstock et al., 1992; L. Takahashi et al., 1992]. Aea
caaaeai?e i?iaeai? ia iaaeaii iaeaaeii? oaaae c oi/ee ci?o oi?ioaaiiy
noaoaaeo ? a?eiaeo iniaeeainoae no?an-?aaeoeaiino? AAIN o iauaaee?a.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. ?iaioo aeeiiaii
a ?inoeooo? aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A.I. Eii?na?aiea AII
Oe?a?ie a ?aieao iaoeiai-aeine?aeieo i?ia?ai IIC oa AII Oe?a?ie (??
aea?ae?a?no?aoe?? 834a, 82a, NI.09.02.0002.85, 155, 182, 278), a oaeiae
iaoeiai-aeine?aeii? i?ia?aie, i?aeo?eiaii? AIIC oa a?aioaie
I?aeia?iaeiiai iaoeiaiai oiiaeo Aeae. Ni?ina (a?aioe UAP000, UAP200)

Iaoa ?iaioe aea/aiiy ?ii?eioeiaiaeo iaoai?ci?a iae?iaiaeie?eiii?
?aaoeyoe?? ?ai?iaeoeoeaii? ooieoe?? oa no?an-?aaeoeaiino? a ii?i? oa ??
iiaeeo?eaoe?e aiane?aeie no?aniaeo oa ai?iiiaeueieo aieea?a ia ie?ae ?
i?aai?ci iiaiia?iaeaeaiiai.

Iniiai? caaaeaiiy aeine?aeaeaiiy:

1. Aea/eoe ?ieue ooieoe?iiaeueiiai iaoaaie?cio oanoinoa?iio
(a?iiaoecaoe?? oa a?ae?ieneethaaiiy) o ?icaeoeo aiae?iaaiicaeaaeii?
iaoieia?? ?ai?iaeoeoeaii? nenoaie.

2. I?iaiae?coaaoe ?ieue ooieoe?iiaeueii? aca?iiae?? ano?iaaiieo
iaoaaie?o?a oa eaoaoieai?i?a ye ?iaeoeoi??a aeic??aaiiy
iae?iaiaeie?eiii? nenoaie ?aaoeyoe?? ?ai?iaeoeoe??.

3. Aeine?aeeoe i?ioean oi?ioaaiiy noaoaaiai aeeii?o?cio iiiiai?ia?a?/ii?
nenoaie iiceo a ii?i? oa ca oiia ii?ooaiiy noaoaai? aeeoa?aioe?aoe??
iiceo aiane?aeie no?ano ? ai?iiiaeueiiai aeenaaeaino a i?a- oa
iinoiaoaeueiiio ia??iae? aeeooy.

4. Aea/eoe a?aeaeaeai? iane?aeee aieeao i?aiaoaeueiiai no?ano ia
iae?i-aiaeie?eiio ?aaoeyoe?th ?ai?iaeoeoeaii? ooieoe?? oa
no?an-?aaeoeaiino? a?iioaeaii-a?iio?ca?ii-iaaeie?eiai? nenoaie.

Iaoeiaa iiaecia ?acoeueoao?a aeine?aeaeaiiy. O aeena?oaoe?ei?e ?iaio?
aia?oa oai?aoe/ii oa aenia?eiaioaeueii iaa?oioiaaii iae?io?i?/io
eiioeaioe?th ai?iii-iaae?aoi?iiai ?ii?eioeiao iae?iaiaeie?eiii?
?aaoeyoe?? ?ai?iaeoeoe?? oa aaeaioaoe??. Aeiaaaeaii, ui i?ioeane
?ii?eioeiao o iiceo a?aeaoaathoueny oeyoii aca?iiae?? eaoaoieai?i?a,
noa?i?aeieo ai?iii?a oa ?o aeoeaieo iaoaaie?o?a (cie?aia,
eaoaoieano?iaai?a), ui ooai?ththoueny aaciina?aaeiuei a oeaieiao
oeaio?aeueii? ia?aiai? nenoaie (i?aiioe/i?e ae?eyioe?, iaae?iaacaeueiiio
a?iioa-eaion?), ca iaoai?ciii ei?iaeoeoe??.

Anoaiiaeaii, ui ii?ooaiiy ?ii?eioeiaiaeo iaoai?ci?a aiane?aeie
ia?eiaoaeueieo no?ani?ieo oa ai?iiiaeueieo aieea?a i?ecaiaeeoue aei
?icaeoeo ooieoe?iiaeueieo ?iceaae?a ?ai?iaeoeoeaii? nenoaie (
aiiaoeyoi?ia iaie?aei?noue) oa no?an-?aaeoeaiino? (neiae?ii
i?aiaoaeueiiai no?ano).

Aeyaeai? iia? iaoiaaiaoe/i? iaoai?cie iae?iaiaeie?eiii? iaoieia??,
coiiaeaii? no?aniaeie e ai?iiiaeueieie /eiieeaie a i?ioean? ?icaeoeo
ieiaea oa i?aai?cio a ia??iae ?aiiueiai iioiaaiaco. Aeiaaaeaii, ui ?aii?
iinoiaoaeuei? ci?ie iiiiai?ia?a?/ii? nenoaie iiceo, ui aeieeathoue
aiane?aeie i?aiaoaeueiiai no?ano oa ia?eiaoaeueiiai ai?iiiaeueiiai
aeenaaeaino, caeo/ai? aei ?ii?eioeiaiaeo iaoai?ci?a ? aeaoa?i?iothoue
a?aeaeaeai? ci?ie iae?iaiaeie?eiii? ?aaoeyoe?? ?ai?iaeoeoe?? oa
no?an-?aaeoeaiino? AAIN. Ii?oieia?/iei i?iyaii ooieoe?iiaeueieo
ii?ooaiue ? cieeiaiiy noaoaaeo a?aei?iiinoae uiaei ia’?io yaea?
iae?ioeeo?a o naen-aeeii?oieo ae?eyieao iiceo, cie?aia a
noi?ao?aciaoe/iiio yae?? a?iioaeaiona. Anoaiiaeaii, ui aetheiei?oeei?aee
? iiaeeeaeie ai?iiiaeueieie /eiieeaie, ye? iiina?aaeeiaothoue iaoiaaiiee
aieea no?ano i?ae /an aaa?oiino? ia ?icaeoie iae?iaiaeie?eiii? nenoaie o
iauaaee?a.

Aeiaaaeaii, ui eaoaoieai?ie a?iioaeaiona ? o??th ni?eueiith eaieith a
iaoiaaiac? ii?ooaiue noaoaai? aeeoa?aioe?aoe?? iiceo, ye? aeieeathoue ye
aiane?aeie no?ani?ieo aieea?a o ia??iae aioo??oiueioo?iaiiai ?icaeoeo
i?aai?cio, oae ? ai?iiiaeueiiai aeenaaeaino o iiaiia?iaeaeaieo oaa?ei.

Aia?oa aea/aii iioiaaiaoe/iee i?io?eue (a?ae noaoaaic??eiai a?eo aei
noa?ino?) ci?i no?an-?aaeoeaiino? AAIN o iauaaee?a iaio noaoae, ye?
caciaee aieeao no?ano o ia??iae aioo??oiueioo?iaiiai ?icaeoeo. Aeyaeaii
o?eaaeee oa?aeoa? oeeo ci?i, ye? caa??aathoueny iaa?oue o noa?eo oaa?ei
(neiae?ii i?aiaoaeueiiai no?ano). Aeiaaaeaii, ui i?aiaoaeueiee no?an
caeaoiee iiaeeo?eoaaoe ia?aa?a no?an-a?e-neiae?iio, oiaoi iaaiith i??ith
ci?ithaaoe i?ia?aio iioiaaiaco.

Anoaiiaeaii, ui aieea i?aiaoaeueiiai no?ano ia aaeaioeai? ?aaeoe?? AAIN
o aei?ineeo ? noa?eo oaa?ei ?aae?co?oueny ca ?aooiie iaoai?ci?a, ye?
caeaaeaoue a?ae noao?. Aeyaeaii, ui i?ae aieeaii i?aiaoaeueiiai no?ano
ai?iiiaeueia ?aaeoe?y ei?e iaaeie?eiaeo caeic o aei?ineeo naiie iii??ii
iineeth?oueny, a o naioe?a nooo?ai cieaeo?oueny. C’yniaaii, ui iniiaiith
iaoiaaiaoe/iith eaieith ineaaeaii? no?an-?aaeoeaiino? AAIN o aei?ineeo
i?aiaoaeueii no?aniaaieo naioe?a uo??a ? cia/ia ciaioaiiy aea?eueiaiiy
ii?aae?aiae?io a a?iioaeaion? o a?aeiia?aeue ia no?ania? noeioee.

Aeyaeaii, ui i?ae /an noa??iiy o i?aiaoaeueii no?aniaaieo naioe?a uo??a
caa??aa?oueny caeaoi?noue eaoaoieai?i?a a?iioaeaiona ?aaaoaaoe ia
no?ania? noeioee ia oe? cia/ii? aeoea?caoe?? ooieoe?? ei?e iaaeie?eiaeo
caeic. Aeiaaaeaii, ui i?ae aieeaii i?aiaoaeueiiai no?ano o noa?eo naiie
cieaeothoueny aaeaioeai? iiaeeeaino? eaoaoieai?iiai? nenoaie
a?iioaeaiona, i?ioa caa??aa?oueny aae?aiiei?oeeaeueia ?aaeoe?y ia
no?ania? noeioee, ui na?ae/eoue i?i a?eueo i?ci? caanaiiy aaeaioeaieo
iaoai?ci?a.

I?aeoe/ia cia/aiiy ?iaioe. Oai?aoe/i? iieiaeaiiy, ye? aeieeaathoue ?c
?acoeueoao?a i?iaaaeaii? ?iaioe, ni?eythoue ?icoe?aiith oa ?icaeoeo
i?iaeaie “O?c?ieia?/i? iaoai?cie ae?? ai?iii?a” ?, cie?aia, oaei? ??
/anoeie, ye “Ai?iiie oa iicie”. Iaea?aeai? ?acoeueoaoe ?ice?eaathoue
aaaeeeaa cia/aiiy ?ii?eioeiai-aeo iaoai?ci?a ?icaeoeo iae?iaiaeie?eiii?
iaoieia?? ?ai?iaeoeoe?? oa no?an-?aaeoeaiino?, yea aeieea? aiane?aeie
ia?eiaoaeueii? iiaeeo?eaoe?? i?ia?aie ?iaeea?aeoaeueiiai ?icaeoeo
i?aai?cio, ? aeathoue i?aenoaao cai?iiiioaaoe eiioeaiooaeueio iiaeaeue
ai?iii-iaae?aoi?iiai ?ii?eioeiao iae?iaiaeie?eiii? nenoaie. C iiceoe?e
oe??? eiioeaioe?? io?eiai? aeai? aeicaieythoue ii-iiaiio inieneeoe
caaaeuei? caeiiii??iino? iaoiaaiaco no?an-?aaeoeaiino? AAIN,
aiaeie?eiiiai aacie?aeaey, a oaeiae ii?ooaiue noaoaai? iiaaae?iee.

?acoeueoaoe aeine?aeaeaiue aeaiiino?othoue iioaioe?eio iaaaciaeo
aeiiieiaiaiai no?ano aeey ?ai?iaeoeoeaiiai caei?ia’y oa aaeaioaoe?eiiai
iioaioe?aeo o iauaaee?a ? iiaeooue neoaeeoe oai?aoe/iei iaa?oiooaaiiyi
aeey oe?eani?yiiaaiiai iiooeo cania?a ? iaoiae?a i?io?eaeoeee oa
ei?aeoe?? ii?ooaiue iae?iaiaeie?eiii? ?aaoeyoe?? ?ai?iaeoeoeaii? oa
aaeaioaoe?eii? nenoai. Io?eiai? aeai? uiaei a?aeaeaeaieo iane?aee?a
aieeao no?ano, aetheiei?oeei?aeieo oa noaoaaeo ai?iii?a o ia??iaee
aioo??oiueioo?iaiiai oa ?aiiueiai iinoiaoaeueiiai ?icaeoeo i?aai?cio ia
iae?iaiaeie?eiio nenoaio iiaeooue aooe ei?eni? oe?ieiio eieo
ee?i?oeeno?a: aeooa?ai-a?iaeieiaai, iaae?ao?ai, naeniiaoieiaai oa
aa?iioieiaai. Iniaenoee aianie caeiaoaa/a. Caeiaoaa/ a?aa
aaciina?aaeith o/anoue o ?ic?iaoe? i?ia?aie aeine?aeaeaiue oa aeeiiaii?
an?o aenia?eiaioaeueieo aeine?aeaeaiue, ?acoeueoaoe yeeo aeeeaaeai? a
aeena?oaoe??. Ii?oieia?/ia aeine?aeaeaiiy eaoaoieai?ia?a?/ieo oa?i?iaeae
na?aaeeiiiai ocaeooy i?iaaaeaii ca o/anoth e.a.i. E.O. Ao?/einueei?,
noa?oiai iaoeiaiai ni?a?ia?oieea ?inoeoooo o?c?ieia?? ?i. I.I.
Aiaiiieueoey IAI Oe?a?ie; ?aooa ii?oieia?/ieo aeine?aeaeaiue – ca
o/anoth ae.a.i. O.?. Aiaaeaiiai?, e.a.i. A.A. Eice?eoeueeiai oa
iaoeiaiai ni?a?ia?oieea E.?. Iieyeiai? ia aac? eaai?aoi??? ii?oieia??
aiaeie?eiii? nenoaie oa a?aeae?eo aiaeie?eiieia?? ?ai?iaeoeoe?? oa
aaeaioaoe?? ?inoeoooo aiaeie?eiieia?? oa iai?io ?a/iaei ?i. A.I.
Eii?na?aiea AII Oe?a?ie.

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. Iaoa??aee aeena?oaoe??
aeiiia?aeaeenue oa iaaiai?thaaeenue ia: I?aeia?iaeiiio neiiic?oi?,
i?enay/aiiio i?iaeaiai iae?iiioiaaiaco (I?aaa, 1985); ??? Ananithci?e
eiioa?aioe?? c i?iaeai iae?i-aiaeie?eiieia?? (Oa?e?a, 1988);
I?aeia?iaei?e eiioa?aioe?? c i?iaeai iae?i-aiaeie?eiieia?? (Othaeiaai,
1989); ??? I?aeia?iaei?e eiioa?aioe?? c i?iaeai naenieia?? (Aa?e?i,
1990); 5-e eiioa?aioe?? ?a?iiaenueeiai oiaa?enoaa iae?i-aiaeie?eiieia?a
(Aoaeaiaoo, 1991); Ananithciiio neiiic?oi? “Iae?ia?ieia?/i? aniaeoe
no/anii? aiaeie?eiieia??” (Iineaa, 1991); ?O I?aeia?iaeiiio eiia?an? c
ai?iiiaeueieo noa?i?ae?a (Aeaean, 1994); 3-io I?aeia?iaeiiio eiia?an? c
iae?i-aiaeie?eiieia?? (Aoaeaiaoo, 1994); O?O c’?cae? Oe?a?inueeiai
o?c?ieia?/iiai oiaa?enoaa ?i. ?.I. Iaaeiaa (Ee?a, 1994); ?O
Ana?in?enuee?e eiioa?aioe?? “Iae?i-aiaeie?eiieia?y-95” (Naieo-Iaoa?ao?a,
1995); ?? Iaoe?iiaeueiiio eiia?an? iaoi-o?c?ieia?a Oe?a?ie (Ee?a, 1996);
I?aeia?iaeiiio neiiic?oi? “Iae?iaiaeie?eii? aniaeoe ?ai?iaeoeoe??”
(Aa?oaaa, 1996); C’?cae? Oe?a?inueeiai o?c?ieia?/iiai oiaa?enoaa
(Aeiiaoeuee, 1998). Aeena?oaoe?y i?ieoea ai?iaaoe?th ia ni?eueiiio
can?aeaii? ai?iaaoe?eii? eii?n?? c aenia?eiaioaeueii? aiaeie?eiieia?? oa
i?io?eueieo a?aeae?e?a ? eaai?aoi??e ?inoeoooo aiaeie?eiieia?? oa iai?io
?a/iaei ?i. A.I. Eii?na?aiea AII Oe?a?ie 4 /a?aiy 1999 ?.

Ioae?eaoe??. Ca iaoa??aeaie aeena?oaoe?? iioae?eiaaii 46 iaoeiaeo ?ia?o,
c yeeo 22 noaoo? o oaoiaeo iaoeiaeo aeaeaiiyo (6 iaeiiin?aieo ? 16 o
ni?aaaoi?noa?) ? 24 ioae?eaoe?? ca iaoa??aeaie i?aeia?iaeieo ?
a?o/eciyieo iaoeiaeo c’?cae?a, neiiic?oi?a, eiioa?aioe?e.

Ia’?i ? no?oeoo?a aeena?oaoe??. Aeena?oaoe?th aeeeaaeaii ia 293
noi??ieao ae?oeiaaiiai oaenoo. ?iaioa neeaaea?oueny ?c anooio, iaeyaeo
e?oa?aoo?e, iaoiaee/iiai ?icae?eo, i’yoe ?icae?e?a aeanieo
aeine?aeaeaiue, iaaiai?aiiy ?acoeueoao?a, aeniiae?a oa nieneo
e?oa?aoo?e, yeee i?noeoue 570 aeaea?ae. ?iaioo ?ethno?iaaii 35
oaaeeoeyie oa 13 ?enoieaie.

CI?NO ?IAIOE

Iaoa??aee, iaoiaee oa iiaeae? aeine?aeaeaiue

Oaa?eie. Aeine?aeaeaiiy i?iaaaeai? ia 3000 naioeyo ? naieao uo??a e?i??
A?noa? ??ciiai a?eo c aeaoiaaiei aeiai ia?iaeaeaiiy: iiaiia?iaeaeaieo ?
?aiiueiai iinoiaoaeueiiai a?eo (1- 7, 8, 10 ? 12 -aeaiieo),
noaoaaic??eeo (3-6-i?ny/ieo) ? noa?eo (20-22-i?ny/ieo). Aeaiue
ia?iaeaeaiiy iauaaee?a aaaaeaee ca ia?oo aeiao aeeooy. Oaa?ei
oo?eioaaee i?e e?iiaoi?e oaiia?aoo?? oa i?e?iaeiueiio oioiia??iae?. An?
oaa?eie ciaoiaeeeeny ia cae/aeiiio ?aoe?ii? a?aa??th. Aaaaeaiiy
ai?iiiaeueieo oa ?ioeo i?aia?ao?a, no?anoaaiiy, a oaeiae caa?e oaa?ei
cae?enithaaee a iaeei ? oie aea /an aeiae i?ae 10 oa 11 aiaeeiaie ?aieo
aei i?eeiaiiy ?aee. Aenia?eiaioe noaaeee ia?aaaaeii o ceiiai-aaniyiee
ia??iae (ethoee-ea?oaiue). Eiaeia aeine?aeia a?oia oaa?ei aea/aeany
iaeii/anii c a?aeiia?aeiith eiio?ieueiith a?oiith. Oaa?ei caaeaaee
oaeaeeith aeaeai?oaoe??th ca?aeii c oiiaaie aenia?eiaio?a.

Ai?iiiaeuei? i?aia?aoe. Oanoinoa?ii-i?ii?iiao (5% ?ic/ei aeey
aioo??oiueii’yciaeo ?i’?eoe?e a aiioeao), ?ic/eiaiee o e?noi/eia?e aai
neeaia?e ie??, aaiaeeee naieai uo??a i?ae oe??o iaeii?aciai a aeic? 50
iea ia oaa?eio ia ia?oo aai i’yoo aeiao aeeooy, aai a aeic? 250 iea ia
o?ao?e aeaiue aeeooy a ia’?i? 40 iee. Caa?e oaa?ei i?iaiaeeee ia 7, 8,
10 ? 12-oo aeiao aeeooy, a oaeiae o 3-4-i?ny/iiio a?oe?.

2-A?ae?ieneano?ii (2-IIA1), 2-a?ae?ieneano?aae?ie-17 (2-IIA2-17)
(“Steraloids”, NOA), 4-a?ae?ieneano?aae?ie-17 (4-IIA2-17), a oaeiae
4-a?ae?iene-ano?aae?ie-17 (4-IIA2-17) (inoaii? aeaa i?aia?aoe etha’ycii
iaaeai? i?io. R. Knuppen, I?ia//eia) ?ic/eiyee o i?ii?eaiae?eie? c 0,01%
anei?a?iiai? eeneioe ? aaiaeeee naieai uo??a i?ae oe??o iaeii?aciai aai
aaaaoi?aciai a aeic? 5, 10 aai 25 iea ia oaa?eio a ia’?i? 40-100 iee ia
i?ioyc? ?aiiueiai iinoiaoaeueiiai ia??iaeo ?icaeoeo. A ie?aieo
aeine?aeao 5-aeaiiei naieai uo??a i?iaiaeeee i?yi? ?io?aoea?aa?aeuei?
?i’aeoe?? 2-IIA1 ? 2-IIA2-17 a aeic? 5 iea ia oaa?eio a ia’?i? 10 iee
ca iaoiaeii N. Hasegawa et al. (1991). Eiio?ieuei? oaa?eie aiaeia?/iei
/eiii (nenoaiii aai ?io?aoea?aa?aeueii) io?eioaaee aea?aaeaioio
e?euee?noue ?ic/eiieea aeine?aeaeoaaieo i?aia?ao?a. A caeaaeiino? a?ae
iio?aa aenia?eiaioo oaa?ei aeaeai?ooaaee /a?ac 30 oa, 1, 2, 3, 4 oa
24 aiae, a oaeiae /a?ac 2, 5 oa 7 ae?a i?ney inoaiiuei? ?i’?eoe??
a?aeiia?aeiiai i?aia?aoo. ?aooo oaa?ei aeaeai?ooaaee o 3-4-i?ny/iiio
a?oe?.

A?ae?iei?oecii aoeaoao (“Aaaeaii ?eooa?”, Oai?ueia) o aeic? 50 ia/ea
iane o?ea aaiaeeee naieai uo??a i?ae oe??o uiaeaiii ia i?ioyc?
inoaiiueiai oeaeiy aaa?oiino?, oiaoi c 15-i? ii 21-oo aeiao. Iauaaee?a
uo??a iaio noaoae aeaeai?ooaaee ia 10-oo aeiao i?ney ia?iaeaeaiiy aai o
3-4-i?ny/iiio a?oe?.

Oa?iaeieia?/i? i?aia?aoe. ?ia?a?oi?e a?iiaoace oanoinoa?iio
aiae?ino-4-?i-3,6,17-o??ii (AO) o aeic? 0,5 ia ia oaa?eio oa
aiae?ino-1,4,6-o???i-3-17-ae?ii (AOAe) (“Steraloids”, NOA) o aeic? 1 ia
aai 0,1 ia aaiaeeee naieai uo??a iiiaeeioe? aai ?acii c OI ia i?ioyc?
ia?oeo ae?a i?ney ia?iaeaeaiiy. Caa?e oaa?ei i?iaiaeeee ia 7, 8, 10 oa
12-o aeiao iinoiaoaeueiiai ?icaeoeo, a oaeiae o 3-i?ny/iiio a?oe?.

?ia?a?oi? eaoaoie-I-iaoeeo?ainoa?ace o?iiieii (“Sigma“, NOA) ?ic/eiyee a
caaooa?aiiio o?c?ieia?/iiio ?ic/ei? (?I 6,5) ? aaiaeeee naieai uo??a o
aeicao 100, 300 aai 600 iea ia oaa?eio o ??ciee oa?i?i ?aiiueiai
iinoiaoaeueiiai ia??iaeo aeeooy iiiaeeioe? aai ?acii ?c OI aai
2-IIA2-17. Oaa?ei aeaeai?ooaaee /a?ac 3, 6 oa 24 aiae i?ney inoaiiuei?
?i’?eoe?? i?aia?aoo, a oaeiae ia 10-o aeiao iinoiaoaeueiiai ?icaeoeo. A
ie?aieo aeine?aeao oaa?ei aeaeai?ooaaee o 3-i?ny/iiio a?oe?.

Iiia?aaeiee eaoaoieai?i?a L-AeIOA ( L–(3,4-aeea?ae?ieneoai?e)-aeai?i;
“Koch-Light“, Aiae?y) ?ic/eiyee o i?aeeeneaiiio 1i nieyiith eeneioith
o?c?ieia?/iiio ?ic/ei? ? aaiaeeee naieai uo??a o aeic? 1 ia ia
oaa?eio c 4-i? ii 10-o aeiae aai o aeic? 0,2 ia ia 5-o oa 7-o aeiao
aeeooy. Eiio?ieuei? oaa?eie iaea?aeoaaee ?ic/eiiee i?aia?aoo. Caa?e
oaa?ei i?iaiaeeee o 3-i?ny/iiio a?oe?.

6-A?ae?ieneaeioai?i a?ae?ioei?eae (6-II-AeA) (“Serva”, I?ia//eia)
?ic/eiyee o o?c?ieia?/iiio ?ic/ei? c 0,1% anei?a?iiai? eeneioe ?
aaiaeeee naieai uo??a o aeic? 1 ia ia oaa?eio (100 ia/ea iane o?ea) ia
i?ioyc? ia?oeo aeaio ae?a i?ney ia?iaeaeaiiy. Oaa?ei aeaeai?ooaaee o
10-aeaiiiio oa 3-i?ny/iiio a?oe?.

On? oa?iaeieia?/i? i?aia?aoe aaiaeeee iiaiia?iaeaeaiei oaa?eiai i?ae
oe??o a ia’?i? 40 iee.

I?aiaoaeueiee no?an. A iniiao aenia?eiaioaeueii? iiaeae? i?aiaoaeueiiai
no?ano iieeaaeaii oaeo, yea aoea cai?iiiiiaaia I. Ward, J. Weisz
(1984). Naiie o noaae?? ano?ono i?aenaaeaeoaaee aei noaoaaic??eeo
naioe?a (ii aea? naiee ia naioey). Aeniiaie i?i ianoaiiy aaa?oiino?
?iaeee ia i?aenoaa? aeyaeaiiy nia?iaoici?ae?a o aaa?iaeueieo iaceao
ianooiiiai ?aieo. *anoeia naiie c 15-i? ii 21-oo aeiao aaa?oiino?
i?aeaeaaaeanue uiaeaiiiio no?anoaaiith oeyoii ?iiia?e?caoe?? ia niei?
i?ioyaii 1 aiae. Iano?aniaai? aaa?oi? naiee neeaaeaee eiio?ieueio
a?oio.

A?io?i?/i? iaoiaee. Ai?no ii?aae?aiae?io (IA) oa aeioai?io (AeA)
aecia/aee a no?oeoo?ao aieiaiiai iiceo (oe?eiio a?iioaeaion?,
i?aiioe/i?e ae?eyioe? oa iaae?iaacaeueiiio a?iioaeaion?) oeoi?eiao?e/iei
iaoiaeii [D. Jacobowitz, J. Richardson, 1978]. I?aiioe/io ae?eyieo oa
iaae?iaacaeueiee a?iioaeaion iiaiia?iaeaeaieo uo??a aeae?eyee i?ae
noa?aii?e?ineiiii ca iaoiaeii V. Luine et al. (1975) ia oieiaeo i?e + 4
0 N. C?acee aeine?aeaeoaaieo ae?eyiie iiceo ia’?aeioaaee a?ae 3-4
uo?aiyo a?aeiia?aeii? noao?. Ca noaiaea?oe aeei?enoiaoaaee
L-ii?aae?aiae?i a?oa?o?ao oa nieyiieeneee 3-a?ae?ieneoe?ai?i
(“Koch-Light”, Aiae?y). Oethi?anoeaioe?th aei??thaaee ia
niaeo?ioethi?eiao?? “O?oa/? MPF-2A” (ssiii?y).

Ia?a IA ? AeA o i?aiioe/i?e ae?eyioe? oa iaae?iaacaeueiiio a?iioaeaion?
iiaiia?iaeaeaieo uo??a aeine?aeaeoaaee ca aeiiiiiaith ?ia?a?oi?a
oe?iceia?ae?ieneeace -iaoee-ia?a-oe?iceio (IIO; “Sigma”, NOA), yeee
aaiaeeee iaeii?aciai i?ae oe??o o aeic? 250 iea a?eueii? iniiae ia 1 ea
iane o?ea ca iaeio aai aea? aiaeeie aei aeaeai?oaoe?? oaa?ei. Eiinoaioo
oaeaeeino? ia?ao, oaeaee?noue ia?ao oa /an ia?ao iiiiai?i?a
?ic?aoiaoaaee ca iaoiaeii B. Brodie et al. (1966) ia i?aenoaa?
aecia/aiiy ai?noo iiiiai?i?a aei oa i?ney aaaaeaiiy IIO.

Ai?no na?ioii?io (NO) oa 5-a?ae?iene?iaeieioeoiai? eeneioe (5-A?IE) o
aeene?aoieo no?oeoo?ao iiceo aecia/aee iaoiaeii aenieiaoaeoeaii?
??aeeiii? o?iiaoia?ao?? i?ae aenieei oeneii ?c oethi?eiao?e/iei
aeaoaeoi?ii [A. Krstulovic, C. Matzura, 1972]. Iaoaaie?/iee ia?a
na?ioii?io ?ic?aoiaoaaee ii ni?aa?aeiioaiith eiioeaio?aoe?e eiai
iniiaiiai iaoaaie?oo 5-A?IE oa NO ( 5-A?IE/NO).

Aeoeai?noue eaoaoie-I-iaoeeo?ainoa?ace (EIIO) o a?iioaeaion? aecia/aee
o i?e?iniiaeuei?e o?aeoe?? ca iaoiaeii R. McCaman (1965) o iiaeeo?eaoe??
W. Ladosky, H. Schneider (1981). Ca noano?ao aeei?enoiaoaaee
3,4-aeea?ae?ieneaaicieio eeneioo (iaea?aeaia c
3,4-aeea?ae?ieneaaicieiiai aeueaeaa?aeo, E. Merck, a eaai?aoi???
i?aneioaco). Aeiiaoi?ii iaoeeueieo a?oi aoa
S-aaeaiicee-iaoee-[14C]-iaoeii?i (NEN Research Products, NOA). I?ney
cae?i/aiiy ?ieoaaoe?? i?e 27 0 N iaoeeueiaai? i?iaeoeoe aeno?aaoaaee
aoeeaoeaoaoii ? i?iaiaeeee ?aae?iiao??th aeno?aeo?a ia aaoa-niaeo?iiao??
“Beckman” (NOA). Ai?no a?eeo aecia/aee iaoiaeii O. Lowry et al.
(1951).

??aaiue ei?oeeinoa?iio (11-ieneei?oeeinoa?i?ae?a) o ieaci? e?ia?
aecia/aee oethi?eiao?e/iei iaoiaeii [P. De Moor et al., 1960], ai?no
oanoinoa?iio (O) ? ano?aae?ieo o ieaci? e?ia? ?ioiioa?iaioiei iaoiaeii
?c aeei?enoaiiyi noaiaea?oieo iaai??a Testosterone Kit ? Estradiol Kit
o??ie “Bio-Rad “(?oae?y).

No?an-?aaeoeai?noue AAIN o aei?ineeo (3-5-i?ny/ieo) oa noa?eo
(20-22-i?ny/ieo) ?ioaeoieo ? i?aiaoaeueii no?aniaaieo naioe?a ? naiie
uo??a aeine?aeaeoaaee ca ?aaeoe??th eaoaoieai?i?a a?iioaeaiona oa ei?e
iaaeie?eiaeo caeic ia aeiciaaiee ?iiia?e?caoe?eiee no?an. Uo??a eiaeii?
a?eiai? a?oie a?aeiia?aeii? noao? (?ioaeoi? oa i?aiaoaeueii no?aniaai?)
aoei i?aeaeaii ?iiia?e?caoe?? ia niei? i?ioyaii 30 aai 60 oa. Oaa?ei
caaeaaee oaeaeeith aeaeai?oaoe??th a?ae?aco i?ney cae?i/aiiy no?ano aai
/a?ac 30 oa i?ney cae?i/aiiy iaeiiaiaeeiiiai ?iiia?e?caoe?eiiai no?ano.
Eiio?ieuei? a?oie neeaaeaeeny c ?ioaeoieo oa i?aiaoaeueii no?aniaaieo
naioe?a ? naiie uo??a a?aeiia?aeiiai a?eo, ye? ia caciaee aino?iai
no?ano.

Aeine?aeaeaiiy aaeaioaoe?eieo iiaeeeainoae i?aai?cio i?iaiaeeee ca
oanoii aoiie [E.I. O?aooaiaa?a e ae?., 1987] o naioe?a ? naiie uo??a
a?eii 3-5, 11 oa 21 i?n. ?icaeoie o?ce/ii? aoiie i?aai?cio ioe?ithaaee
ca /anii (nae) oo?eiaiiy oaa?ei ia aa?oeeaeuei?e aea?aeei? ae?aiao?ii 2
ni oa aeiaaeeiith 50 ni.

Ooieoe?iiaeueiee noai ?ai?iaeoeoeaii? nenoaie o naiie uo??a ioe?ithaaee
ca oa?i?iii noaoaaiai aeic??aaiiy ?c ?a?no?aoe??th aeiae a?aee?eooy
i?oae oa no?oeoo?ith ? o?eaae?noth ano?aeueieo oeeee?a oeyoii uiaeaiiiai
i?e?ineii?/iiai ianoaaeaiiy i?oa’yieo iace?a i?ioyaii aeaio i?nyoe?a c
aeiae a?aee?eooy i?oae, a oaeiae i?ioyaii ia?oeo aeaio oeaei?a eiaeiiai
ae?oaiai i?nyoey o a?aeaeaeai? ia??iaee aeeooy (3, 4, 6, 8, 10, 12, 14,
16, 18, 20 ? 22- i?n).

Ii?oieia?/i? aeine?aeaeaiiy aaeaiia?iio?ca, ei?e iaaeie?eiaeo caeic, a
oaeiae eaoaoieai?ia?a?/ieo oa?i?iaeae na?aaeeiiiai ocaeooy i?iaiaeeee ca
aeiiiiiaith aeaeo?iiii? oa na?oiai? i?e?ineii??. Aeey oa?aeoa?enoeee
naen-aeeii?oieo yaea? iiceo (noi?ao?aciaoe/ieo yaea? oa a?eoaoiiai yae?a
a?iioaeaiona) aeei?enoiaoaaee ii?oiiao?e/iee aiae?c ia’?io yaea?
iae?ioeeo?a.

Noaoenoe/io ia?iaeo ?acoeueoao?a aeine?aeaeaiue i?iaiaeeee ?c
aeei?enoaiiyi t-e?eoa??th Nothaeaioa, iaia?aiao?e/iiai e?eoa??th U
A?eeieniia – Iaiia – O?oi?, a oaeiae iaoiaeo aeueoa?iaoeaiiai
aa??thaaiiy. ?ic?aooiie na?aaei?o iieaciee?a cae?enithaaee ca i?ia?aiith
aeey AII ia i?e?ieaeueeoeyoi?? “Yeaeo?iieea-IE-61“.

?ACOEUeOAOE AeINE?AeAEAIUe OA ?O IAAIAI?AIIss

?ieue ooieoe?iiaeueiiai iaoaaie?cio noaoaaeo ai?iii?a o ?icaeoeo
aiae?iaaiicaeaaeii? iaoieia?? ?ai?iaeoeoeaii? nenoaie

Ii?ooaiiy iae?iaiaeie?eiii? ?aaoeyoe?? ?ai?iaeoeoe?? aeine?aeaeoaaee ia
eeane/i?e iiaeae? iaiiaoaeueii aiae?iaai?ciaaieo naiie uo??a, yeo
a?aeoai?thaaee ca aeiiiiiaith ?i’?eoe?e aeciaaiiiai oanoinoa?iio
i?ii?iiaoo o e?eoe/iee ia??iae NAeI. A?aeiii, ui aieiaiei i?iyaii oeeo
ii?ooaiue ? ?icaeoie aiiaoeyoi?iiai neiae?iio, yeee oa?aeoa?eco?oueny
a?aenooi?noth oeeee?/ii? nae?aoe?? aiiaaeio?iiieo ai?iii?a. Iaeiae
iaoai?cie, ye? ciaoiaeyoueny o i?aea?oio? ia?aaoaeiae a?iioaeai?/ieo
oeaio??a ca /ieia?/ei oeiii i?ae /an NAeI, caeeoathoueny iac’yniaaieie.
I?e ?ica’ycaii? oe??? i?iaeaie iniiaia oaaaa aoea i?eae?eaia aiae?co
ooieoe?iiaeueii? aca?iiae?? i?ae noaoaaeie noa?i?aeaie oa eaoaoieai?iaie
o a?iioaeaion? iaiiaoaeueii aiae?iaai?ciaaieo naiie uo??a.

Aieiaiith ia?aaeoiiaith aeey oeueiai iai?yieo aeine?aeaeaiue aoei
aeyaeaiiy i?e/eiiiai ca’yceo i?ae i?aeaeuaiiyi ??aiy IA o a?iioaeaion? o
e?eoe/iee ia??iae NAeI oa ?icaeoeii aiiaoeyoi?ii? iaie?aeiino? o
iaiiaoaeueii aiae?iaai?ciaaieo naiie uo??a [A. Reznikov, N. Nosenko,
1983]. Oyaeaiiy i?i ?ieue EA o aiae?iaaiicaeaaei?e NAeI ae?noaei na?e
iiaeaeueoee ?icaeoie i?e aea/aii? noaoaaeo a?aei?iiinoae EA a?iioaeaiona
o ia??iae ?aiiueiai iinoiaoaeueiiai iioiaaiaco. Io?eiai? aeai?
na?ae/aoue i?i aeuee ??aaiue IA o a?iioaeaion? naioe?a, i?ae o naiie, ia
10-oo aeiao i?ney ia?iaeaeaiiy (eiio?ieuei? naioe?: 3,39 0,26 iiieue/a
oeaieie; eiio?ieuei? naiee: 2,56 0,16 iiieue/a oeaieie, P? ??H??????????? ???????yyy???????????? „i??? ????'yy?????????? ????yyy???? ???????uoy??????1/4???I???"Arial Cyr?I????-?????uoy?????? 2-II-A1 4-II-A2-17 2-II-A2 4-II-A2-17 (30 oa) (1 aiae) (4 aiae) (24 aiae) Eiio?ieue I?aiaoaeueiee no?an wmetafile8? ??c??????????? ???????yyy???????????? „i??? ????'yy?????????? ????yyy???? ???????uoy??????1/4???I???"Arial Cyr?I????-?????uoy?????? wmetafile8? ??oe??????????? ???????yyy???????????? _I??? ????'yy?????????? ????yyy???? ???????uoy??????1/4???I???"Arial Cyr?I????-?????uoy?????? 5 11 21 A?e oaa?ei (i?n) Naiee Naioe? Naioe? Naiee wmetafile8? ???? ????????? ???????yyy???????????? ‰+??? ????'yy?????????? ????yyy???? ???????uoy??????1/4???I???"Arial Cyr?I????-?????uoy?????? Eiio?ieue I?aiaoaeueiee Eiio?ieue I?aiaoaeueiee no?an no?an Aacaeueiee ??aaiue 1aiae ?iiia?e?caoe?y Oaaeeoey 1 Ai?no oa ia?a eaoaoieai?i?a ? na?ioii?io a i?aiioe/i?e ae?eyioe? oa iaae?iaacaeueiiio a?iioaeaion? uo??a 10-aeaiiiai a?eo, ye? caciaee aieeao a?ae?iei?oeciio a i?aiaoaeueiee ia??iae aeeooy (Im, n=6-11) IieacieeEiio?ieueA?ae?iei?oeciinaieenaioe?naieenaioe?I?aiioe/ia ae?eyieaIA, iiieue/a oeaieie4,22 0,48 2,74 0,38 (a) 2,64 0,37 (a) 2,16 0,31AeA, iiieue/a oeaieie3,45 0,68 4,12 0,36 2,54 0,23 3,24 0,22Oaeaee?noue ia?ao IA, iiieue/a/aiae.0,99 0,21 1,89 0,45 0,45 0,20 0,33 0,09 (b)Oaeaee?noue ia?ao AeA, iiieue/a/aiae0,33 0,11 0,73 0,19 0,69 0,12 (a) 1,13 0,27NO, ia/ia oeaieie0,43 0,03 0,35 0,02 0,43 0,04 0,52 0,03 (b)5-A?IE, ia/ia oeaieie0,73 0,14 0,87 0,08 1,71 0,09 (a) 1,42 0,07 (a, b, c)5-A?IE/NO1,80 0,36 2,49 0.23 4,22 0,58 (a) 2,76 0,15 (a, c)Iaae?iaacaeueiee a?iioaeaionIA, iiieue/a oeaieie3,76 0,33 3,35 0,23 4,32 0,24 3,86 0,19AeA, iiieue/a oeaieie3,59 0,24 3,21 0,22 2,63 0,67 3,51 0,19Oaeaee?noue ia?ao IA, iiieue/a/ aiae.0,53 0,09 0,78 0,13 0,72 0,12 0,62 0,17Oaeaee?noue ia?ao AeA, iiieue/a/ aiae0,42 0,11 0,69 0,12 0,61 0,19 0,62 0,29NO, ia/ia oeaieie0,44 0,05 0,64 0,15 0,36 0,02 0,33 0,045-A?IE, ia/ia oeaieie0,74 0,08 1,03 0,14 1,51 0,14 (a, b) 1,11 0,06 (a, c)5-A?IE/NO1,85 0,31 2,23 0,57 4,19 0,39 (a, b) 3,57 0,59 (a)I?ei?oea: ??cieoey aeinoia??ia P

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