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

?inoeooo iieaeoey?ii? a?ieia?? ? aaiaoeee

Aiai?oi Aeieo?i Ieeieaeiae/

OAeE 577.323

577.324

O?CEEI-O?I?*I? IAOAI?CIE

A?IIIEAEOEss?IIAI AI?CIAAAIIss

03.00.03 – iieaeoey?ia a?ieia?y

Aaoi?aoa?ao

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

aeieoi?a a?ieia?/ieo iaoe

Ee?a — 1999 Aeena?oaoe??th ? ?oeiien.

?iaioo aeeiiaii o a?aeae?e? iieaeoey?ii? a?io?ceee ?inoeoooo
iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie, i. Ee?a.

Iaoeiaee eiinoeueoaio: aeieoi? a?ieia?/ieo iaoe, i?ioani?, aeaaeai?e
IAI Oe?a?ie Iaoeoea Aaiiaae?e Oa?eaii?eiae/, ?inoeooo iieaeoey?ii?
a?ieia?? ? aaiaoeee IAI Oe?a?ie, i. Ee?a, aee?aeoi?, caa?aeoaa/
a?aeae?eo no?oeoo?e ? ooieoe?e ioeea?iiaeo eeneio.

Io?oe?ei? iiiiaioe: aeieoi? a?ieia?/ieo iaoe, i?ioani?, aeaaeai?e IAI
Oe?a?ie ?euenueea Aaiia Aaeaioei?aia, ?inoeooo iieaeoey?ii? a?ieia?? ?
aaiaoeee IAI Oe?a?ie, i. Ee?a, caa?aeoaa/ea a?aeae?eo iaoai?ci?a
o?aineyoe?? aaiaoe/ii? ?ioi?iaoe??,

aeieoi? a?ieia?/ieo iaoe, i?ioani? Ceia Aaeaioei Eaii?aeiae/,
Ee?anueeee oi?aa?neoao ?iai? Oa?ana Oaa/aiea, i?ioani? eaoaae?e
a?io?ceee a?ieia?/iiai oaeoeueoaoo,

aeieoi? o?ceei-iaoaiaoe/ieo iaoe, i?ioani? Oa?eyiai Aaea??e
Ieeieaeiae/, ?inoeooo o?ceee IAI Oe?a?ie, i. Ee?a, caa?aeoaa/ a?aeae?eo
o?ceee a?ieia?/ieo nenoai.

I?ia?aeia onoaiiaa: ?inoeooo a?ii?aai?/ii? o?i?? ? iaooio?i?? IAI
Oe?a?ie, i. Ee?a.

Caoeno aeena?oaoe?? a?aeaoaeaoueny 23 eenoiiaaea 1999 ?ieo i 10 aiaeei?

ia can?aeaii? Niaoe?ae?ciaaii? a/aii? ?aaee Ae 26.237.01

a ?inoeooo? iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie

ca aae?anith: 252143, i. Ee?a-143, aoe. Aeaaeai?ea Caaieioiiai, 150.

C aeena?oaoe??th iiaeia iciaeiieoeny a a?ae?ioaoe? ?inoeoooo
iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie (252143, i. Ee?a-143, aoe.
Aeaaeai?ea Caaieioiiai, 150).

Aaoi?aoa?ao ?ic?neaii 21 aeiaoiy 1999 ?ieo.

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

eaiaeeaeao a?ieia?/ieo iaoe,

no. iaoeiaee ni?a?ia?oiee Eoeao E.E.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue oaie. A?iiieaeoey?ia ai?ciaaaiiy ? iaeiei ?c
iaeoa?aeoa?i?oeo ao?eaoo?a aeeaiai – aiii iiina?aaeeiao? ia?aaaaeio
a?eueo?noue a?ieia?/ii aaaeeeaeo i?ioean?a ia iieaeoey?iiio ??ai?
(Aiaaeueaa?aeo, 1976). Oiio iiooe iaea?eueo caaaeueieo, a ?aeaae? –
oi?aa?naeueieo, o?ceei-o?i?/ieo caeiiii??iinoae, eio?? eaaeaoue a eiai
iniia?, ? iaeiei ?c ooiaeaiaioaeueieo iai?yie?a iieaeoey?ii? ?
no?oeoo?ii? a?ieia??. Aeooaeuei?noue oeueiai aeoaea iai?inoiai ? aaeueie
?iceiaiai caaaeaiiy aecia/a?oueny oaeiae ia?niaeoeaaie oe?eani?yiiaaiiai
aieeao ia ia?aa?a ye ia?a-, oae ? iaoaaie?/ieo i?ioean?a o ee?oeiao
i?aai?cio oeyoii neioaco niieoe, aea??eia?noue ca’ycoaaiiy yeeo aeecueea
aei naeaeoeaiino? i?e?iaeieo iieaeoe, oa noai?aiiy ia ?oi?e iniia?
??ciiiai?oieo e?ea?nueeeo ? ae?aaiinoe/ieo iieaeoey?ieo nenoai. Ie??i
oiai, oi?iae?ciaaia ?ioi?iaoe?y uiaei i?e?iaee iaoai?ci?a
a?iiieaeoey?iiai ai?ciaaaiiy ae?ae iaiao?aeia aeey noai?aiiy iaoiaeaie
noi?aiieaeoey?ii? o?ceei-o?i?? iia?oi?o oaoiieia?e, cie?aia
a?iiieaeoey?ii? aeaeo?ii?ee (Lehn, 1995).

Iei? a oea?ei? i?iaeaie a?iiieaeoey?iiai ai?ciaaaiiy i?iaeiaaeo?oueny
?ioaineaia iaeiie/aiiy aenia?eiaioaeueieo aeaieo (a iniiaiiio
?aioaaiino?oeoo?ieo ? ssI?) aeniei? ?icae?eueii? caeaoiino? oa ?oiy
nenoaiaoecaoe?y. I?ioa oe?e?nia ea?oeia o?ceei-o?i?/ii? i?e?iaee
niaoeeo?/iino? aei oeueiai /ano oae ? ia aee?enoae?coaaeany. Anoaiiaeaii
eeoa, ui e?eoe/ieie ia?aiao?aie inoaiiuei? ? ieiua iiaa?oi? iaii?eueiiai
eiioaeoo iieaeoe-ia?oia??a oa ?oiy ao?ii?noue. sseui ia?oee ?c ieo
aecia/a?oueny, a iniiaiiio, neeaie Aai-aea?-Aaaeuena, oi
a?aeiia?aeaeueieie ca ae?oaee ? aiaeiaa? ca’ycee (I-ca’ycee), oae caai?
oi/eia? eiioaeoe. Aaaaea?oueny, ui aianie oi/eiaeo eiioaeo?a o
niaoeeo?/i?noue a?eeiai-ioeea?iiaiai ai?ciaaaiiy ciaioo?oueny a ?yaeo
ioeeaioeaeia iniiaa – oeoe?iaee caeeoie – oinoaoia a?oia. Aeiaaaeaii, ui
a?eie ?ici?cia? ia na?aaei?, a eieaeuei? aa??aoe?? no?oeoo?e AeIE.
Aecia/aeueio ?ieue o ia?aa?ao i?ioean?a a?eeiai-ioeea?iiaiai ai?ciaaaiiy
a?ae?a?a? aaeeeiaiie?ooaeia, iae?i?eia aeeiai?ea iieaeoe-ia?oia??a, a
ia?oo /a?ao AeIE. A?aeiia eeoa iaeia caaaeueiia?ieia?/ia iiaeaeue, yea
eiioeaio?o? a nia? ia?ae?/ai? caeiiii??iino?, — oaiiiaiieia?/ia iiaeaeue
no?oeoo?ii-aeeiai?/ii? a?aeiia?aeiino? i?noeue ai?ciaaaiiy (Koshland,
1958-1971). I?ioa aioo??oiy “iieaeoey?ia eia?ea” i?ioean?a ai?ciaaaiiy
a?iiie?ia?aie iaeei iaeiiai, ye aeioe?eueii aeaooiaaieo nenoai,
i?iaeiaaeo? caeeoaoeny “iica eaae?ii”.

Aiae?c e?oa?aoo?e cana?ae/o?, ui iaeiei ?c “aocueeeo i?noeue” i?iaeaie
a?eeiai-ioeea?iiaiai ai?ciaaaiiy ? iaiaaeai?noue ciaiue i?i o?
no?oeoo?ii-aeeiai?/i? iniaeeaino? AeIE, ye? a?aeiia?aeathoue ca ??
niaoeeo?/io aca?iiae?th c a?eeii. Inue ?o aeaeaei ia iiaiee ia?ae?e:
iecueei/anoioia iae?i?eii-aeeiai?/ia iiaaae?iea, iia’ycaia c
ia?aaeieaaeaiiyi oa, oae caaieie iai?a?ice?eoeie noaiaie; caeaoi?noue
e?i?eii? AeIE aei iecueeiaia?aaoe/ieo eieaeueieo ? ?ioaa?aeueieo
aeaei?a, aaee/eia yeeo ?noioiuei caeaaeeoue a?ae iine?aeiaiino? iniia;
aeaeaeiae?y a AeIE; aa??aoe?? eiioi?iaoe?? oeoe?iai-oinoaoiiai e?noyea
oa a?aenoai? i?ae non?aei?ie eiiieaiaioa?ieie ia?aie iniia AeIE a
caeaaeiino? a?ae iine?aeiaiino? oiui. Oae, ia?aaeienaeueiee c oi/ee ci?o
onoaeaieo iiaeyae?a ia no?oeoo?ii-aeeiai?/io i?aai?caoe?th AeIE
aeniiaie, c?iaeaiee ia iniia? na??? aenia?eiaio?a (Diekmann et al.,
1986-1992), i?i oa, ui aee?eaeai?noue e?i?eii? AeIE iia’ycaia ?c
noae?iaii, iaooiaoo? ia aeoieo, ui no/ani? oyaeaiiy i?i eiai i?e?iaeo
iio?aaothoue yeui ia ia?aaeyaeo, oi i?eiaeii? ye?niiai aaeineiiaeaiiy.
Oa ae naia noino?oueny no?oeoo?ii-aeeiai?/ieo aeanoeainoae eiiiiiaio?a
IE – iniia, ioeeaiceae?a, ?oi?o Oionii-E?ee?anueeeo ia? oiui. sse
aeieeaa? c aiae?co e?oa?aoo?e, aaea iaeiie/eeiny /eiaei a?aoiaio?a ia
i?aeoaa?aeaeaiiy oaei? oi/ee ci?o.

C ?ioiai aieo, ?c i?eioeeiiaeo i?e/ei i?iaeaio a?iiieaeoey?iiai
ai?ciaaaiiy ia iiaeia ae??oeoe, caeeoath/enue a ?? ?aieao. Iaiao?aeii
i?iaiae?coaaoe ?? c a?eueo caaaeueieo iiceoe?e, a naia – ye aeaooiaai?
a?iiie?ia?e, /ei aiie a?ae??ciythoueny a?ae i?aai?/ieo
iie?ia??a-i?aaenoaaiee?a iaaeeaiai na?oo ? a /iio iieyaathoue aeeaeii?
o?ceei-o?i?/i? canaaee (“iieaeoey?ia eia?ea”) ?oiueiai ooieoe?iioaaiiy.
?ioeie neiaaie, i?iaeaia a?iiieaeoey?iiai ai?ciaaaiiy ia iiaea aooe
iiaeieaia ?ai?oa, ai?ae aoaea noai?aia /e, i?eiaeii?, ie?aneaia oe?e?nia
o?ceei-o?i?/ia eiioeaioe?y ooieoe?iioaaiiy a?iiie?ia??a. Ia i?aaaeeeee
aeaeue, caeiaooee o oeueiio iai?yieo a?eueo, i?ae ne?iii?: ia?ii, ii
noo?, eeoa iaeio oaiiiaiieia?/io iiaeaeue “a?eie-iaoeia” (*a?iaaneee,
1987 -1999) ? oaiiiaiieia?/iee i?eioeei iieaeoey?ii? ?aaoeyoe?? (Monod
et al., 1963).

Ioaea, i?iaeaia a?iiieaeoey?iiai ai?ciaaaiiy caeeoa?oueny ae?ae
aeooaeueiith ye c iaeyaeo ia ?? iaia?an?/io oai?aoe/io ? i?aeoe/io
cia/ou?noue, oae ? c o?aooaaiiyi noaio ?? iaaeinoaoiuei? aea/aiino?.
Aeey naiai ae??oaiiy aiia iio?aao? ye?nii iiaeo i?aeoiae?a, ye?
aeoiaeyoue ca onoaeai? ?aiee, ui neeaeeny a oe?e aaeoc? ciaiue.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. Aeena?oaoe?eia
?iaioa oe?eeiaeoi a?aeiia?aea? iniiaiiio ieaiia? ooiaeaiaioaeueieo
aeine?aeaeaiue a?aeae?eo iieaeoey?ii? a?io?ceee ?inoeoooo iieaeoey?ii?
a?ieia?? ? aaiaoeee IAI Oe?a?ie (caa?aeoaa/ a?aeae?eo aei 1995 ?ieo –
eaiaeeaeao a?ieia?/ieo iaoe, no. iaoeiaee ni?a?ia?oiee, aeioeaio
AEaeoianueeee I.A.). ?? aeeiiaii a ?aieao athaeaeaoieo oai “Aea/aiiy
niaoeeo?/iino? a?eeiai-ioeea?iiaeo aca?iiae?e a iiaeaeueieo nenoaiao
??ciiai ??aiy neeaaeiino?” (? aea?ae. ?a?no?aoe?? 01.91.0002584,
1991-1995 ?.?.) ? “Aea/aiiy o?ceei-o?i?/ii? i?e?iaee ia?aa?ao
aeaiaioa?ieo i?ioean?a a?eeiai-ioeea?iiaiai ai?ciaaaiiy a iiaeaeueieo
nenoaiao oeio “iiiiia?-iiiiia?” oa “iiiiia?-iie?ia?” (? aea?ae.
?a?no?aoe?? 0198U005247, 1996-2000 ?.?.).

*anoeiai ?iaioa aeeiioaaeany a ?aieao io?eiaieo ia eiieo?nieo canaaeao
a?o/eciyieo oa i?aeia?iaeieo i?iaeo?a “Aieea iaoeethaaiiy ioeeaioeaeieo
iniia ia ?o ai?ciaaaiiy ai?iieeneioieie ?aaeeeaeaie, ui i?noyoue
ea?aieneeueio a?oio” (i?ia?aia “Oiiae ooiaeaiaioaeueieo aeine?aeaeaiue”
AeEIO Oe?a?ie, oeo? 5/410, 1992-1993 ?.?.), “?ic?iaeoe iaoiaee
iaea?aeaiiy iiaeo aioe?ao?ia??onieo i?aia?ao?a a ?yaeo cai?uaieo ii
oeoe?o neioaoe/ieo ioeeaiceae?a ? ?o aiaeia?a” (i?ia?aia “A?ioaoiieia?y”
AeEIO Oe?a?ie, oeo? 01.11.01/027-92, ? aea?ae. ?a?no?aoe?? 0195U019371,
1992-1995 ?.?.), “Aieea i?ioiioaaiiy oa aeai?ioiioaaiiy ioeeaioeaeieo
iniia oa ai?iieeneio ia i?ioeane oi/eiaiai a?eeiai-ioeea?iiaiai
ai?ciaaaiiy” (i?ia?aia “Oiiae ooiaeaiaioaeueieo aeine?aeaeaiue” AeEIO
Oe?a?ie, oeo? 5.2/97, 1993-1994 ?.?.), “Iiooe ?a/iaei c
?ioiiiiaeoethth/eie oa i?ioeiooeeiieie aeanoeainoyie na?aae ioeeaiceae?a
acai??ei?aeeiiaiai ?yaeo” (i?ia?aia “Caoeno aaiioiiaeo ianaeaiiy
Oe?a?ie” AeEIO Oe?a?ie, oeo? 1.01.01/064-92, ? aea?ae. ?a?no?aoe??
0195U011006, 1994 ?.), “Aea/aiiy no?oeoo?ii-aeeiai?/ieo iniaeeainoae
eiiiiiaio?a a?iiie?ia??a oa ?o aieeao ia niaoeeo?/i?noue oi/eiaeo
a?eeiai-ioeea?iiaeo oa ioeea?iiai-ioeea?iiaeo eiioaeo?a” (i?ia?aia
“Oiiae ooiaeaiaioaeueieo aeine?aeaeaiue” AeEIO Oe?a?ie, oeo? 5.3/184,
1994-1997 ?.?.), “Aea/aiiy no?oeoo?e oa o?ceei-o?i?/ieo aeanoeainoae
iecueeiiieaeoey?ieo aiaeiaaica’ycaieo eiiieaen?a, ui iiaeaeththoue
niaoeeo?/i? a?eeiai-ioeea?iia? eiioaeoe” (a?aio i?aeia?iaeiiai iaoeiaiai
oiiaeo “Oe?a?ia”, oeo? K1F100, 1994-1995 ?.?.), “Aieea i?ioiioaaiiy oa
aeai?ioiioaaiiy ioeeaioeaeieo iniia oa ai?iieeneio ia i?ioeane oi/eiaiai
a?eeiai-ioeea?iiaiai ai?ciaaaiiy, aecia/aeueiiai aeey ?aaoeyoeaieo
iaoai?ci?a ee?oeie” (i?ia?aia “Oiiae ooiaeaiaioaeueieo aeine?aeaeaiue”
AeEIO Oe?a?ie, oeo? 5.2/97, 1995 ?.), “I?ioio?iiia oaooiia??y
ioeeaioeaeieo iniia: iia? i?aeoiaee aei noa?i? i?iaeaie” (i?ia?aia
“Oiiae ooiaeaiaioaeueieo aeine?aeaeaiue” AeEIO Oe?a?ie, oeo? 5.4/77,
1996 – o./an ) oa aeayeeo ?ioeo.

Iaoa ? caaaeaiiy aeine?aeaeaiiy. Iaoa ?iaioe – anoaiiaeoe iaea?eueo
caaaeuei? o?ceei-o?i?/i? caeiiii??iino? ioeea?iiai-ioeea?iiaiai ?
a?eeiai-ioeea?iiaiai ai?ciaaaiiy ia i?e?ino?oeoo?iiio ??ai?, ni?yioaaaoe
iniiai? coneeey ia aeyaeaiiy iiaeo, ooieoe?iiaeueii aaaeeeaeo
no?oeoo?ii-aeeiai?/ieo aeanoeainoae eiiiiiaio?a IE ? iiaeaeueieo
eiiieaen?a oeio “iiiiia?-iiiiia?”, c’ynoaaiiy ?oiuei? eaaioiaio?i?/ii?
i?e?iaee ? noai?aiiy ia oe?e iniia? iae?iiieaeoey?ieo iiaeaeae
ai?ciaaaiiy.

Aeey aeinyaiaiiy oe??? iaoe aoee iinoaaeai? oae? eiiieaeni? caaaeaiiy:

Aea/eoe iniiai? o?ceei-o?i?/i? oi?ie no?oeoo?ii? i?ieeaino?
ioeeaioeaeieo iniia: a ia?oo /a?ao – noa?aio?i?/io iaaei?noe?noue,
no?oeoo?io ?ciia??th oa i?ioio?iiio oaooiia??th, a oaeiae iia? aniaeoe
niiioaiiiai aeacai?ioaaiiy oa iioeiaeaeaiiy eeniaaeie ?aaeeeaeaie ?
aeaoe ?i iaeaaeio eaaioiaio?i?/io ?ioa?i?aoaoe?th.

Aeine?aeeoe i?ioiiiaeiii?i?, aeeth/ath/e NI-eeneioi?noue, oa
i?ioiiiaeoeaioi?i? aeanoeaino? acioenoeo iniia, ui aecia/athoue ?oith
eiiieaenioa??io caeaoi?noue. Anoaiiaeoe, ye ci?ith?oueny no?oeoo?ia
iaaei?noe?noue iniia i?e ?oiueiio i?ioiioaaii? ? aeai?ioiioaaii? oa i?e
aoyaoaaii? o i?aeiieaeoey?i? I-ca’ycee.

Aeine?aeeoe eiioi?iaoe?eio aio/e?noue Oionii-E?ee?anueeeo ia? iniia
AeIE, anoaiiaeoe iniiai? noa?aiaeaeo?iii? /eiieee, ui eaaeaoue a ??
iniia?, oa i?iaiae?coaaoe ?? iiaeeeao a?ieia?/io cia/ou?noue o oea?ei?
a?iiieaeoey?iiai ai?ciaaaiiy.

Aea/eoe aieea noa?aio?i?/ii? iaaei?noeino? iniia AeIE ia iioeei-o?ce/i?
aeanoeaino? ?oi?o e?enoae?a ? ?icaeiooe ia oe?e iniia? iia? oyaeaiiy i?i
noae?ia ioeeaioeaeieo iniia oa ?oi?o Oionii-E?ee?anueeeo ia?.

Aecia/eoe aia?aaoeeo ? eiiia?aoeai? aeanoeaino? i?aeiieaeoey?ieo
I-ca’yce?a o iiaeaeueieo nenoaiao ioeea?iiai-ioeea?iiaiai ai?ciaaaiiy –
e?enoaeao ? ni?ae?enoae?caoao iniia AeIE – oa ?? aianie o noaa?e?caoe?th
inoaii?o.

Noai?eoe aenia?eiaioaeueio onoaiiaeo aeey i?aoeec?eii? ?a?no?aoe??
?aiai?anueeeo niaeo??a a?iiie?ia??a oa ?oi?o neeaaeiaeo /anoei. Io?eiaoe
iecueei/anoioi? ((<200 ni-1) ?aiai?anuee? niaeo?e eiiiiiaio?a IE, ui iine?aeiaii oneeaaeiththoueny, iecueeiiieaeoey?ieo eiiieaen?a, ui iiaeaeththoue oi/eia? ioeea?iiai-ioeea?iia? oa a?eeiai-ioeea?iia? eiioaeoe, oa ?oi?o eiiiiiaio?a o oaa?ae?e oac?. Anoaiiaeoe i?e?iaeo iecueei/anoioieo eieeaaiue AeIE, aaaeeeaeo aeey ?? ?ici?ciaaaiiy a?eeii. Aea/eoe iniiai? no?oeoo?ii-aeeiai?/i? oa aia?aaoe/i? aeanoeaino? iiaii? iiiaeeie oi/eiaeo eiioaeo?a oeio “ioeeaioeaeia iniiaa – a?/iee ?aaeeeae ai?iieeneioe”, ui noaa?e?cothoueny aeaiia i?aeiieaeoey?ieie I-ca’yceaie. ?ic?iaeoe iia? iiaeaeuei? oyaeaiiy i?i no?oeoo?ii-aeeiai?/i? iniaeeaino? AeIE, ye? eiio?ieththoue ?ici?ciaaaiiy ?? a?eeii, a naia: niiioaii? iai?a?ice?eo? noaie, aee?eaeai?noue, aeaeaeiae?th, ia?aaeieaaeaiiy, aaiioiene/io ae?th OO-ii?ii?iaiiy, eiiia?aoeai?noue. Iaoeiaa iiaecia iaea?aeaieo ?acoeueoao?a. Aia?oa anoaiiaeaii, ui acioeno? iniiae c ai?iia?oiith ? no?oeoo?ii-iaaei?noeeie, aeeiieueii-iano?eeeie iieaeoeaie; ?ioa?eiiaa?n?y yeeo a?aeaoaa?oueny o?ueiia oiiieia?/ii ? aia?aaoe/ii iaaea?aaeaioieie oeyoaie, noi?iaiaeaeo?oueny ?noioiueith aeaoi?iaoe??th ai?iiiai o?aaiaioa ? caeaaeeoue a?ae ?icoaooaaiiy ai?iia?oie a e?eueoe? oa a?ae ?? ioi/aiiy. Aia?oa anoaiiaeaii eaaioiai-o?i?/i? canaaee oe??? a?ieia?/ii aaaeeeai? aeanoeaino? ? c?iaeaii aeniiaie i?i eaaioiao aaiiao??th ioeeaioeaeieo iniia, ye? iathoue o nai?io neeaae? ai?ii- /e iaoeeueio (Ia) a?oio. Aia?oa iieacaii, ui no?oeoo?ia iaaei?noe?noue ioeeaioeaeieo iniia c ai?iia?oiith caeaaeeoue a?ae ?? oaooiia?iiai oa ?ciia?iiai noaio, a oaeiae a?ae ci?ie ?oiueiai ca?yaeiaiai noaio oa aoyaoaaiiy o i?aeiieaeoey?i? I-ca’ycee. Aia?oa aea/aii no?oeoo?io ?ciia??th eaiii?/ieo ioeeaioeaeieo iniia, anoaiiaeaii, ui eeoa aea? c ieo – Ade ? Thy ? aia?aaoe/ii iaeaea?aei?oeie no?oeoo?ieie ?ciia?aie (aeiaaeueieie i?i?ioiaie) a nai?o n?iaenoaao oa iaa?oioiaaii aeniiaie i?i oa, ui c-iii?ae on?o iniia AeIE naia Ade ? iaee?auei no?oeoo?iei aeaiaioii aeey noai?aiiy aaiaoe/ii? eiaeiai? iao?eoe? a i?ioean? aaiethoe??. C?iaeaii aeniiaie i?i oa, ui oa?aeoa?iith a?aei?iiith ?enith IE a?ae i?aai?/ieo iie?ia??a – i?aaenoaaiee?a iaaeeaiai na?oo ? oi?eaeueia ii?aeiaiiy ieie ??ciiiai?oieo oi?i no?oeoo?ii? i?ieeaino?. Aia?oa o iaeneiaeueii iiaeeeaiio oi?iao?, iaiaaeaiiio eeoa oyaeaiiyie i?i i?e?iaeo o?i?/iiai ca’yceo, aeine?aeaeaii i?ioio?iiio oaooiia??th eaiii?/ieo iniia, ?oi?o aeacai?iiaaieo aiaeia?a, a oaeiae iieaeoe – neeaaeiaeo /anoei oeeo niieoe. Aia?oa anoaiiaeaii, ui i?ioio?iiia oaooiia??y acioenoeo iniia ia? iieaeoey?ii-oea?oa??iiiee oa?aeoa?, i?e/iio a i?e aa?ooue o/anoue on? aac aeiyoeo i?ioiie c ea?aii?ioiiaie aeeth/ii. Aia?oa c’yniaaii, ui aeey io?eiiaeo iniia iniiaiei oaooiia?ii-oea?oa??iiii c-iii?ae oeo, ui oi?iothoueny oeyoii i?a?aoe?? ea?aii?ioii?a, ? ?e?aeia oi?ia, eio?a ooai?th?oueny a ?acoeueoao? ia?aoiaeo ea?aii?ioiia i?e aoii? N8 ia non?aei?e c iei aoii acioo, c’?aeiaiee c aoiiii N8 iiaea?eiei ca’yceii, oa a?ae?a?a? aaaeeeao ?ieue o i?ioeanao aiaeiaai-o?eo??aiai iai?io a?oie N8I. Cai?iiiiiaaii iia? oyaeaiiy i?i noae?ia Oionii-E?ee?anueeeo ia? iniia AeIE: aieiaia ?oiy a?aei?ii?noue a?ae ?nioth/eo iieyaa? o iayaiino? niaoeeo?/ieo eoeii?anueeeo eiioaeo?a a?eueii? aeaeo?iiii? ia?e (AAI) ai?iieo aoii?a acioo c aoiiaie aiaeith non?aei?o (aa?oi?o /e ieaei?o) iniia, ye? iiaeia ?icoe?ithaaoe ye aa?oeeaeuei? I-ca’ycee. O eiioaeno? iieaeoey?iiai oi?aae?iiy cai?iiiiiaaii ?icaeyaeaoe ai?iia?oio iniia IE ye “iieaeoey?iee noaeia”, iae?i?eii-aeeiai?/ia ?ooii?noue yeiai ?aaoeth?oueny aoyaoaaiiyi o i?aeiieaeoey?i? I-ca’ycee ye ai?iieo aoii?a aiaeith, oae ? AAI ai?iiiai aoiia acioo. Aia?oa anoaiiaeaii, ui naia oey aeanoea?noue ai?iia?oie caaacia/o? aenieo eiioi?iaoe?eio aio/e?noue Oionii-E?ee?anueeeo ia? iniia a?aeiinii eoo?a "i?iiaea?-oa?no" ? "ceai" oa ie?aneaii a?ieia?/io cia/ou?noue oeueiai aoaeoo aeey no?oeoo?ii-aeeiai?/ii? i?aai?caoe?? IE, ?oiueiai neioaco, eiaeiii-aioeeiaeiiiaeo aca?iiae?e, a oaeiae a?eeiai-ioeea?iiaiai ai?ciaaaiiy. Aia?oa i?aaenoaaeaii i?e?ino?oeoo?i? iiaeae?, ui a?oioothoueny ia iiaeo oyaeaiiyo i?i no?oeoo?io iaaei?noe?noue ioeeaioeaeieo iniia oa ?oi?e noae?ia ? iathoue iine?aeiaia eaaioiaio?i?/ia iaa?oiooaaiiy: aiie noinothoueny oeo no?oeoo?ii-aeeiai?/ieo iniaeeainoae AeIE, ye? aeaoa?i?iothoue ai?ciaaaiiy ?? a?eeii, – niiioaieo iai?a?ice?eoeo noai?a, aee?eaeaiino?, aeaeaeiae??, ia?aaeieaaeaiiy, aaiioiene/ii? ae?? OO_ii?ii?iaiiy oa eiiia?aoeaiino?. Aia?oa aea/aii iniiai? no?oeoo?ii-aeeiai?/i? oa aia?aaoe/i? aeanoeaino? iiaii? iiiaeeie oi/eiaeo eiioaeo?a oeio “ioeeaioeaeia iniiaa-a?/iee ?aaeeeae ai?iieeneioe”, ui noaa?e?cothoueny aeaiia i?aeiieaeoey?ieie I-ca’yceaie. Aia?oa cao?eniaaii ia?ao?ae iniiae ?c iniiaii? o ??aee?nio oaooiia?io oi?io i?ae aieeaii i?aeiieaeoey?ieo aca?iiae?e oa ia?aianaiiy i?ioiio a?ae ai?iieeneioe ia iniiao. Aia?oa anoaiiaeaii, ui aaea ia iaeieae/iio no?oeoo?iiio ??ai? ai?ciaaaiiy a?aeaoaa?oueny ca “noeaia???i” aca?iii? aaeaioeaiino? iieaeoe-ia?oia??a ca o/ano? nooo?ai aca?iicaeaaeieo aioo??oiuei- oa i?aeiieaeoey?ieo I-ca’yce?a, caeaoieo ia eeoa ?iciaaeaoeny ? a?aeiiaethaaoeny, aea e a?oo?eaoeaii ia?aieeaoeny. Aia?oa c?iaeaii aeniiaie i?i iaiaaeai?noue a?aeiii? oi/ee ci?o, yea ioioiaeith? ?iciianthaeaeai?noue, a ioaea – ? a?ieia?/io cia/ei?noue oi/eiaeo a?eeiai-ioeea?iiaeo eiioaeo?a c ?oiueith aia?aaoeeith. Cai?iiiiiaaii aeiiiaieoe “noaoe/i?” noaie oi/eiai? aca?iiae?? iniia c a?/ieie ?aaeeeaeaie ai?iieeneio aeeiai?/ieie, ui aeeth/athoue ye ia?ao?aei? noaie ?ioa?eiiaa?n?? iniia AeIE, oae ? a?/ieo ?aaeeeae?a aeayeeo ai?iieeneio. Iaoiaeaie iecueei/anoioii? ?aiai?anueei? niaeo?ineii?? oa niaeo?aeueii? eaei?eiao??? io?eiaii iia? aeai? i?i o?ceei-o?i?/i? aeanoeaino? oaa?aeioacieo iiaeaeueieo ia’?eo?a – e?enoae?a ? ni?ae?enoae?cao?a ioeeaioeaeieo iniia, ni?ae?enoae?cao?a oeio “iniiaa – iio?aeia ai?iieeneioe”, a oaeiae eiiiiiaio?a IE, ui iine?aeiaii oneeaaeiththoueny. Oai?aoe/ia ? i?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoao?a. Caaaeueiioai?aoe/ia cia/aiiy iaea?aeaieo ?acoeueoao?a iieyaa? o oiio, ui aiie ye?nii ?icoe?ththoue ?nioth/? oyaeaiiy i?i no?oeoo?ii-aeeiai?/io aaoa?iaaii?noue AeIE ? ?? iniiai? eaaioiaio?i?/i? /eiieee oa i?i ia?aa?a aeaiaioa?ieo aeo?a a?eeiai-ioeea?iiaiai oa ioeea?iiai-ioeea?iiaiai ai?ciaaaiiy, i?ae?eiath/e ?o aei ??aiy i?e?ino?oeoo?ieo iiaeaeae, ui iathoue iine?aeiaia eaaioiaio?i?/ia iaa?oiooaaiiy. Aaaeeeaei c oai?aoe/ii? oi/ee ci?o ? iiaeieaiiy i?e oeueiio oe?ei? iecee ?nioth/eo o oe?e oea?ei? noa?aioei?a: oae aaeaeiny, cie?aia, aeiaanoe, ui eaiii?/i? ioeeaioeaei? iniiae iaeaaeaoue aei eeano no?oeoo?ii-iaaei?noeeo iieaeoe, a noae?ia Oionii-E?ee?anueeeo ia? ia iiaea aooe caaaeaiee eeoa aei iaeieae?ciaaieo aca?iiae?e noaoe/ieo ieineiia?aeaeueieo no?oeoo?. Oae? oyaeaiiy aeyaeeeny aeineoue i?iaeoeoeaieie – c’ynoaaeiny, ui aiie eaaeaoue a iniia? oe?ieiai eiea aoaeo?a caaaeueiia?ieia?/ii? cia/ouino?, a naia – iai?a?ice?eoeo noai?a AeIE, ?? aee?eaeaiino?, aeaeaeiae?? a AeIE, ?? ia?aaeieaaeaiiy, eiiia?aoeaiino? oiui, ui eiio?ieththoue ia?aa?a aeaiaioa?ieo aeo?a a?eeiai-ioeea?iiaiai ai?ciaaaiiy ? ye? iienoaaeeny ?ai?oa oaiiiaiieia?/ii. Cai?iiiiiaai? iiaeae?, a oaeiae aeniiaie uiaei oa?aeoa?ii? o?ceei-o?i?/ii? a?aei?iiino? IE a?ae i?aai?/ieo iie?ia??a – i?aaenoaaiee?a iaaeeaiai na?oo aoaeooue noeioethaaoe iiaeaeueo? oai?aoe/i? iiooee o iai?yieo noai?aiiy oe?e?nii? o?ceei-o?i?/ii? eiioeaioe?? ooieoe?thaaiiy IE oa iaiiaiaiiy iiaei ci?noii o?ceei-o?i?/ieo e?eoa???a aeeaiai. sse?nii iiaa eiioeaioe?y noa?aio?i?/ii? iaaei?noeino? iniia AeIE oa ?oiueiai noae?iao iiaea aooe aeei?enoaia ye aaciaa aeey oai?aoe/iiai iiyniaiiy iaia?an?/ieo aenia?eiaioaeueieo oaeo?a, ye? ia iathoue aei oeueiai /ano caaeia?eueiiai i?e?ino?oeoo?iiai oeoia/aiiy ? noinothoueny, cie?aia, oi?eaeueii? iaoai?/ii? aeanoe/iino? AeIE oa ?? caeaoiino? ia?aaeaaaoe aeaeo?e/i? neaiaee, cie?aia oioi?iaeoeiaai?, ia aaeeeo a?aenoaiue aac caoooaiiy. Aiia oaeiae aoaea ni?eyoe oai?aoe/iiio iaa?oiooaaiith iaiao?aeiino? aaeineiiaeaiiy ?nioth/i? no?oeoo?ii? iiiaieeaoo?e IE, ?icaeoeo iiiyooy i?i no?oeoo?ii-aeeiai?/io a?aeiia?aei?noue Oionii-E?ee?anueeeo ia? iniia AeIE ? ?icoe?aiiy o?aaeeoe?eieo ?aiie eiioi?iaoe?eiiai aiae?co IE. Ie??i oiai, aiia aea? ?iioeuen aeey iiaeaeueoiai oai?aoe/iiai ?icaeoeo eeane/ii? ?aea? i?i ?ieue aoaeo?a aeaeo?iiiiai ni?yaeaiiy o ia?aa?ao a?ieia?/ieo i?ioean?a ia iieaeoey?iiio ??ai? ca o/anoth IE: oeaio?aeueiei iiiaioii ooo iiaea aeyaeoeny iiaeeaeaiiy ia oai?aoe/iiio ??ai? aca?iica’yceo i?ae oaeeie aeanoeainoyie, i?eoaiaiieie ye IE, oae ? ioeeaii?ioa?aeiei eiiieaenai, ye eiiia?aoeai?noue, aaoa?iaaii?noue, iae?i?ei?noue ? ia??aiiaaaei?noue ?c no?oeoo?iith iaaei?noe?noth ?oi?o neeaaeiaeo /anoei. Aeai? uiaei aieeao ca?yaeiaiai noaio oinoao?a ia aa?oeeaeuei? aca?iiae?? a AeIE iiaeooue aeyaeoeny aaeueie ei?enieie i?e ?ic?iaoe? neiaia?aaoe/ieo iae?i?eii-aeeiai?/ieo iiaeaeae a?iiieaeoey?iiai oi?aae?iiy ? ?aaoeyoe??, a oaeiae iiaeaeae oa?iiaeeiai?/ii? noaa?eueiino? AeIE ? ?? eiiiaeoecaoe?? a?eeaie. I?eioeeiiai aaaeeeaee aeniiaie i?i eaaioiao aaiiao??th iniia AeIE aeicaieeoue /?oe?oa ie?aneeoe ?aiee i?eaeaoiino? eeane/ieo iaoai?/ieo i?aeoiae?a o oai?aoe/ii-?ic?aooieiaeo aeine?aeaeaiiyo no?oeoo?ii-aeeiai?/ieo aeanoeainoae ye IE, oae ? ioeeaii?ioa?aeieo eiiieaen?a. Aeniiaie i?i iiaeeeao a?ieia?/io cia/ou?noue "i?eoiaaieo" a?ae aaciina?aaeiueiai aenia?eiaioaeueiiai niinoa?aaeaiiy ??aee?nieo oaooiia??a, aeaiaioa?ieo a?eeiai-ioeea?iiaeo eiioaeo?a a ia?ao?aeieo noaiao ?ioa?eiiaa?n?? ?oi?o neeaaeiaeo /anoei oiui aeooae?co? ?ieue oai?aoe/ieo iaoiae?a aeine?aeaeaiue, cie?aia eaaioiaio?i?/ieo. Iniaeeaee oai?aoe/iee ?ioa?an noaiiaeyoue oaeiae ?acoeueoaoe aea/aiiy ??ciiiai?oieo oi?i no?oeoo?ii? i?ieeaino? acioenoeo iniia. Ia aeeeth/a?oueny, ui aiie a?ae?a?athoue iaaaeyeo ?ieue o ia?aa?ao ia?aiaoaaie?/ieo i?ioean?a, ye? iei? ?icoe?iththoueny ye iieaeoey?ia i?aea?oioy noa??iiy ? ana a?eueoa i?eaa?oathoue oaaao aeine?aeiee?a. Io?eiai? a ?iaio? ?acoeueoaoe iiaeooue aooe aeei?enoai? a iaoeiai-aeine?aei?e i?aeoeoe? aeey ?ic?iaee e?euee?nieo iiaeaeae, ui iienothoue i?ioeane a?iiieaeoey?iiai ai?ciaaaiiy ia iae?iiieaeoey?iiio ??ai?. Anoaiiaeai? no?oeoo?ii-aeeiai?/i? aeanoeaino? oi/eiaeo ioeea?iiai-ioeea?iiaeo oa a?eeiai-ioeea?iiaeo eiioaeo?a iiaeooue aooe ei?enieie aeey ciaioaiiy iaiaeiicia/iino? a ?ioa?i?aoaoe?? ?aioaaiino?oeoo?ieo ? ssI?-aeaieo uiaei i?inoi?iai? i?aai?caoe?? IE oa ioeeaii?ioa?aeieo eiiieaen?a. ?acoeueoaoaie aeena?oaoe?? iiaeia oaeiae nei?enoaoeny a noi?aiieaeoey?i?e o?i?? i?e ae?iaeaii? no?aoaa?? neioaco aenieiiieaeoey?ieo niieoe c i?iaiiciaaieie aeanoeainoyie. ?ic?iaeaiee aaoi?ii i?ea?iaeueiee ?aiai?anueeee niaeo?iiao? iiaea ciaeoe canoinoaaiiy anthaee, aea iathoue ni?aao c ?a?no?aoe??th iaei?ioaineaieo niaeo??a. Ie??i oiai, c?iaeai? a ?iaio? aeniiaee aeathoue iiooiao aeey iinoaiiaee iiaeo, ye ?aaeueieo, oae ? eiii’thoa?ieo aenia?eiaio?a, ni?yiiaaieo ia iiaeeaeaiiy ?ioi?iaoe?? uiaei ia?aa?ao aeaiaioa?ieo i?ioean?a a?iiieaeoey?iiai ai?ciaaaiiy. Iaoa??aee aeena?oaoe?? eyaee a iniiao niaoeeo?no, i?i/eoaiiai aeena?oaioii nooaeaioai niaoe?ae?caoe?? "o?ceea a?ieia?/ieo nenoai" oaeoeueoaoo i?e?iaeie/eo iaoe Iaoe?iiaeueiiai oi?aa?neoaoo "Ee?ai-Iiaeeyinueea aeaaeai?y" o 1998-1999 iaa/aeueiiio ?ioe?. Ia?aaaaeia a?eueo?noue /eneiaeo aeaieo, aeeeaaeaieo a aeena?oaoe??, ia’?aeiaia a eiii’thoa?io aaco aeaieo, aeinooiio aeey niiaeeaa/?a. Iniaenoee aianie caeiaoaa/a. Iniaenoee aianie aeena?oaioa ? aecia/aeueiei ia aneo aoaiao aeine?aeaeaiiy ? iieyaa?, ia?o ca ana, o iinoaiiaoe? i?iaeaie oa iaa?oiooaaii? caaaeaiue, o aeai?? ia’?eo?a aeine?aeaeaiue ? ?ic?iaoe? ?oi?o iaoiaeee, aiae?c?, ?ioa?i?aoaoe?? oa ocaaaeueiaii? aenia?eiaioaeueieo ? ?ic?aooieiaeo aeaieo. Eiio iaeaaeaoue an? oai?aoe/i? ?aea? oa aenia?eiaioaeuei? i?aeoiaee, ?aae?ciaai? a aeena?oaoe??. Iniiai? aenia?eiaioaeuei? aeine?aeaeaiiy ? oai?aoe/i? ?ic?aooiee aeeiiai? aaoi?ii iaeiiin?aii. *anoeio ?o io?eiaii o ni?ai?aoe? c ni?a?ia?oieeaie a?aeae?eo iieaeoey?ii? a?io?ceee ?IA?A IAI Oe?a?ie – I. A. AEaeoianueeei, ss. ?. I?uoe, ?. A. Eiiae?aotheii oa N. I. Nai?eeaiei. Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. Iaoa??aee aeena?oaoe?? aeiiia?aeaeeny ia IX, X, XI, XII ? XIV Oeie?-nai?ia?? “Niaeo?ineii?y iieaeoe ? e?enoae?a” (i. Oa?iii?eue, 1989; i. Noie, 1991; i. Oa?e?a, 1993; i. I?aeei, 1995; i. Iaeana, 1999), ???, IV ? VIII ?a?iiaenueeeo eiioa?aioe?yo c niaeo?ineii?? a?ieia?/ieo iieaeoe (i. Aieiiuey, 1989; i. Ei?e, 1991; i. Aioaae, 1999), VIII Ananithci?e eiioa?aioe?? “Aeei?enoaiiy ia/enethaaeueieo iaoei a niaeo?ineii?? iieaeoe ? o?i?/ieo aeine?aeaeaiiyo” (i. Iiainea??nuee, 1989), XII Ananithci?e ia?aae? "Canoinoaaiiy eieeaaeueieo niaeo??a aeey aeine?aeaeaiiy iai?aai?/ieo ? eii?aeeiaoe?eieo niieoe" (i. I?inuee, 1989), ?V Ananithci?e eiioa?aioe?? c niaeo?ineii?? eiia?iaoe?eiiai ?icn?yiiy na?oea (i. Oaeai?iae, 1989), V? Ananithci?e eii?aeeiaoe?ei?e ia?aae? c niaeo?ineii?? iie?ia??a (i. I?inuee, 1989), Ananithci?e oeie? “Iieaeoey?ii-ee?oeii? iaoai?cie ?ioiii? ?aaoeyoe?? aiiainoaco oa i?iaeaie iaoaiaoe/iiai iiaeaethaaiiy” (i. E?aniiy?nuee, 1990), Iaoeiai-i?aeoe/ieo nai?ia?ao “Iioeea ? niaeo?ineii?y oa ?oi? canoinoaaiiy a ia?iaeiiio ainiiaea?noa? ? aeieia??” (i. Iae?oiiieue, 1990; i. Eai’yiaoeue-Iiae?euenueeee, 1992), ??? I?aeia?iaei?e eiioa?aioe?? c eaca?ii? niaeo?ineii?? ?icn?yiiai na?oea oa ae?aaiinoeoe? a?ieia?/ieo ia’?eo?a (i. Iineaa, 1990), ?? I?aeia?iaeiiio eiia?an? oai?aoee?a o aaeoc? i?aai?/ii? o?i?? (i. Oi?iioi, 1990), Ananithci?e oeie? ?c canoinoaaiiy eiii’thoa??a o o?i?? (i. Caaieai?iae, 1990), Oeieao-nai?ia?ao c o?ceee ioeea?iiaeo eeneio (i. Oa?e?a, 1990; i. Oa?e?a, 1993), VI Iieuenueei-?aaeyinueeiio neiiiceoi? c niaeo?ineii?? aiaeiaaiai ca’yceo (i. Iiciaiue, 1990), XX ?a?iiaenueeiio eiia?an? c iieaeoey?ii? niaeo?ineii?? (i. Caa?aa, 1991), V?? ? VIII Eiioa?aioe?yo c niaeo?ineii?? a?iiie?ia??a (i. Oa?e?a, 1991; i. Oa?e?a, 1993), I?aeia?iaeiiio neiiiceoi? c inoaii?o aeinyaiaiue o aeine?aeaeaii? o?i?? oa iieaeoey?ii? a?ieia?? ?aeo (i. Iae?i, 1991), ?X Ananithci?e eiioa?aioe?? c o?i?/ii? ?ioi?iaoeee (i. *i?iiaieiaea, 1992), ? Oe?a?inueei-iieuenueeiio neiiiceoi? c aiaeiaaiai ca’yceo (i. Iaeana, 1992), Eiioa?aioe?? (ca o/anoth i?aeia?iaeieo a/aieo) c eieeaaeueii? niaeo?ineii??, i?enay/ai?e 80-??//th a?ae aeiy ia?iaeaeaiiy A. ?. Noaiaiiaa (i. I?inuee, 1993), XII I?aeia?iaeiiio nai?ia?? c i?aeiieaeoey?ii? aca?iiae?? oa eiioi?iaoe?e iieaeoe (i. Oa?e?a, 1994), I?aeia?iaei?e iaoeia?e eiioa?aioe?? “O?ceea ? o?i?y i?aai?/ieo ethi?iioi??a – 95” (i. Oa?e?a, 1995), ? ? ?? C’?caeao Oe?a?inueeiai a?io?ce/iiai oiaa?enoaa (i. Ee?a, 1996; i. Oa?e?a, 1998), I?aeia?iaei?e eiioa?aioe?? “Iae?i?ei? yaeua a a?ieia??” (i. Ioueii, 1998), THa?eaei?e eiioa?aioe?? Oaaea?aoe?? ?a?iiaenueeeo a?io?i?/ieo oiaa?enoa (i. Eiiaiaaaai, 1998), Eiioa?aioe?? (ca o/anoth i?aeia?iaeieo o/aniee?a) c o?ceee a?ieia?/ieo nenoai (i. Ee?a, 1998), V?? Eiioa?aioe?? c no/anieo oaiaeaioe?e o ia/enethaaeuei?e o?i?? (i. A?enao?a, 1998), XXVII Eiioa?aioe?? i?aaeaiii-no?aeii? anioe?aoe?? o?i?e?a-oai?aoee?a NOA (i. Oaeaoan, 1998), I?aeia?iaei?e eiioa?aioe?? “Aiaeiaaee ca’ycie” (i. Ee?a, 1998), ?? C’?cae? a?io?cee?a ?in?? (i. Iineaa, 1999), XIII Eiioa?aioe?? “Ai?eciioe aeine?aeaeaiue aiaeiaaiai ca’yceo” (i. Oa??aaeoa Cae?oe, 1999) oa Iaoeiaeo nai?ia?ao ?inoeoooo iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie (i. Ee?a, 1990-1999), ?inoeoooo o?ceee IAI Oe?a?ie (i. Ee?a, 1996), Iaoe?iiaeueiiai oi?aa?neoaoo “Ee?ai-Iiaeeyinueea aeaaeai?y” (i. Ee?a, 1997), IAeOe eaaioiai? iaaeeoeeie "A?aeaoe" IIC Oe?a?ie (i. Ee?a, 1999). Ioae?eaoe??. Ia oaio aeena?oaoe?? iaae?oeiaaii 47 noaoae (c ieo 8 - iaeiiin?aieo) o a?o/eciyieo oa ca?oa?aeieo oaoiaeo aeo?iaeao, ui ?aoa?othoueny, oa 33 oace aeiiia?aeae ia iaoeiaeo eiioa?aioe?yo. Aiie o iiaiiio ia’?i? aena?oeththoue ci?no, ?acoeueoaoe, aeniiaee ? ?aeiiaiaeaoe?? aeena?oaoe??. No?oeoo?a oa ianya ?iaioe. Aeena?oaoe?y neeaaea?oueny ?c anooio, /ioe?ueio i?ea?iaeueieo ?icae?e?a, eiaeai c yeeo cae?i/o?oueny aeniiaeaie, aea aeeeaaeaii ?acoeueoaoe aeine?aeaeaiiy ? i?iaaaeaii ?oi? iaaiai?aiiy, i?aenoieiaeo aeniiae?a ? ia?ae?eo aeei?enoaieo e?oa?aoo?ieo aeaea?ae (300 iaeiaioaaiue). Iiaiee ianya aeena?oaoe?? - 300 noi??iie, aiia i?noeoue 37 oaaeeoeue ? 31 ?enoiie. Iaoiaeieia?y, iaoiaee ? ia’?eoe aeine?aeaeaiue. “A?ae i?inoiai aei neeaaeiiai” – oea iniiaiee iaoiaeieia?/iee i?eioeei, yeee aeei?enoaii o ?iaio?. I?inoeie ? aiaeii/an ?ai?acaioaoeaieie /anoeiaie oe?eiai – AeIE ? ioeeaii?ioa?aeieo eiiieaen?a – neoaoaaee ?cieueiaai? ioeeaioeaei? iniiae, ioeeaiceaee, i?ino? iieaeoee, ui iiaeaethaaee a?/i? ?aaeeeaee ai?iieeneio, Oionii-E?ee?anuee? ia?e ? I-ca’ycai? eiiieaene oeio “ioeeaioeaeia iniiaa – a?/iee ?aaeeeae ai?iieeneioe”; aeei?enoiaoaaeeny oaeiae oaa?aeioaci? iecueeiiieaeoey?i? iiaeae? a?eeiai-ioeea?iiaiai ? ioeea?iiai-ioeea?iiaiai ai?ciaaaiiy – e?enoaee eiiiiiaio?a IE, ui iine?aeiaii oneeaaeithaaeeny, ni?ae?enoae?caoe ioeeaioeaeieo iniia, a oaeiae ni?ae?enoae?caoe oeio “ioeeaioeaeia iniiaa –iio?aeia ai?iieeneioe”. Aeei?enoiaoaaeeny ye aenia?eiaioaeuei? (?aiai?anueea niaeo?ineii?y, niaeo?aeueia eaei?eiao??y, a ie?aieo aeiaaeeao ssI? ? ?*- niaeo?ineii?y), oae ? oai?aoe/ii-?ic?aooieia? iaoiaee (iai?aaii??e/i? eaaioiaio?i?/i? iaoiaee AI1, MNDO/3 ? MNDO/I). Iniiaia iioeaaoe?y aeai?o aenia?eiaioaeueieo iaoiae?a – oea aeniea ?ioi?iaoeai?noue, oai?aoe/ii-?ic?aooieiaeo – aeinooii?noue ?aae?caoe?? i?e i?eeiyoi?e oaeaeeino? io?eiaiiy ?ioi?iaoe?? ? ?? yeino?. A eiii’thoa?iiio aenia?eiaio? aaeeea oaaaa i?eae?eyeany eiai ieaioaaiith, a oaeiae ioe?ioe? iaae?eiino? io?eiaieo ?acoeueoao?a. Ia aoai? ieaioaaiiy ia iniia? aiae?co e?oa?aoo?e noai?thaaeany ye?nia iiaeaeue i/?eoaaiiai a eean? niieoe, ui aea/aeeny, aoaeoo. Aeey ioe?iee aeinoia??iino? eiai /eneiai? aa?n?? aeei?enoiaoaaeeny "oanoia?" iieaeoee, ?aoaeueii aea/ai? aenia?eiaioaeueii aai ae ?ic?aooieiaeie iaoiaeaie aenieiai ??aiy neeaaeiino?. O ia?aaaaei?e a?eueoino? aeiaaee?a eiii’thoa?iee aenia?eiaio aoa ni?yiiaaiee ia ia iie?ioaiiy ?nioth/eo /eneiaeo aeaieo, a ia iiooe iiaeo aoaeo?a ui iathoue iaia?an?/ia a?ieia?/ia cia/aiiy. I?e oeueiio ian oe?eaaeee ia no?eueee nai? /eneia? aeai?, ye ?oi? oaiaeaioe??. INIIAIEE CI?NO No?oeoo?ia i?ieea?noue ioeeaioeaeieo iniia: ?acoeueoaoe eaaioiaio?i?/ieo aeine?aeaeaiue Noa?aio?i?/ia iaaei?noe?noue. Aia?oa anoaiiaeaii, ui acioeno? iniiae, ye? iathoue o nai?io neeaae? ai?iia?oio, iaeaaeaoue aei eeano no?oeoo?ii-iaaei?noeeo iieaeoe. ?ioa?eiiaa?n?y ?oi?o ianeiao?e/ieo aiaio?iia??a a?aeaoaa?oueny o?ueiia aia?aaoe/ii ? oiiieia?/ii iaaea?aaeaioieie oeyoaie: /a?ac ieiueiio ?iaa?n?th ai?iiiai o?aaiaioa ?CNH2 , /a?ac ieaia?iee ia?ao?aeiee noai ? nooo?ai ai?cio?iiia iiaa?oaiiy ai?iia?oie iaaeiei aecioeeee?/iiai ca’yceo CN /a?ac aeaa ??cieo ieineineiao?e/ieo ia?ao?aeieo noaie. Oa?aeoa? ?ioa?eiiaa?n?? accioenoeo iniia c ai?iia?oiith nooo?ai caeaaeeoue a?ae iieiaeaiiy inoaiiuei? a e?eueoe? oa a?ae ?? ioi/aiiy. Aeey i??ei?aeeiiaeo iniia iacaeaaeii a?ae iieiaeaiiy ai?iia?oie ? io?eiiaeo iniia c ai?iia?oiith o aoaeue-yeiio iieiaeaii? i??ei?aeeiiaiai e?eueoey iaea?eueo iecueeiaia?aaoe/iei oeyoii ?ioa?eiiaa?n?? ? ieiueiia ?iaa?n?y. I?ioa o aeiaaeeo, eiee ai?iia?oia ciaoiaeeoueny o 8 iieiaeaii? ?i?aeacieueiiai e?eueoey io?eiiaeo iniia, iaea?eueo iecueeiaia?aaoe/iei oeyoii ?ioa?eiiaa?n?? ? aioo??oi? iiaa?oaiiy ai?iia?oie iaaeiei aecioeeee?/iiai ca’yceo CN /a?ac ieineineiao?e/iee ia?ao?aeiee noai c oeen-i???ioaoe??th ai?iii?ioii?a a?aeiinii non?aeiueiai c ai?iiei o?aaiaioii iiaea?eiiai ca’yceo C=N. Aia?oa anoaiiaeaii, ui oaea iiaaae?iea acioenoeo iniia io?eiiaiai ?yaeo c ai?iia?oiith o 8 iieiaeaii? o?nii iia’ycaia c aiaeia?/iith iiaaae?ieith 2-ai?ii?i?aeacieo: eiai aiaethaaiiy c i??ei?aeeiaie ia ci?ith? oa?aeoa?o ?ioa?eiiaa?n??, cao?thth/e eeoa ?? aia?aaoe/i? ia?aiao?e. Aia?oa aaeaeiny iia’ycaoe oa?aeoa? ?ioa?eiiaa?n?? acioenoeo iniia c ai?iia?oiith ?c oa?aeoa?ii i??ai?aeaeueiino? ai?iiiai o?aaiaioa o i?i?ioi? a?ia?iiaa?oi? iioaioe?aeueii? aia?a?? (AIIA): eiee ai?iii?ioiie ?icoaoiaai? ii iaeei a?e a?ae ieiueie e?eueoey, aia?aaoe/ii iaeaea?aei?oei oeyoii ?ioa?eiiaa?n?? ? ieiueiia ?iaa?n?y ai?iiiai o?aaiaioa (oaea neooaoe?y ia? i?noea, cie?aia, o eaiii?/ieo ioeeaioeaeieo iniiaao - Ade, Gua, Cyt); eiee ae ai?iii?ioiie ?ici?uai? aneiao?e/ii ii iaeaeaa aiee a?ae ieiueie e?eueoey, aia?aaoe/ii iaeaea?aei?oei oeyoii ?ioa?eiiaa?n?? ? aioo??oi? iiaa?oaiiy ai?iia?oie iaaeiei aecioeeee?/iiai ca’yceo CN. ?ioa?eiiaa?n?y acioenoeo iniia c ai?iia?oiith noi?iaiaeaeo?oueny ?noioiueith ci?iith aeeiieueiiai iiiaioa (oea iia’ycaii, a iniiaiiio, ?c ci?iith i?inoi?iai? i???ioaoe?? AAI ai?iiiai aoiia acioo), a oaeiae nooo?aith aeaoi?iaoe??th ai?iiiai o?aaiaioa (oae, cie?aia, a ia?ao?aeieo noaiao aioo??oiueiai iiaa?oaiiy ai?iia?oie cia/ii c?inoa? (aei 55° - 57°) eoo iaoeeo ?? ieiueie H'NH'' aei aecioeeee?/iiai ca’yceo CN). Anoaiiaeaii, ui aa?’?? ieiueiii? ?iaa?n?? eaiii?/ieo ioeeaioeaeieo iniia c ai?iia?oiith niaaea? a ?yaeo Gua > Cyt > Ade, i?e/iio aeey Gua a?i aeecueeee aei kT i?e o?c?ieia?/i?e
oaiia?aoo??. Caiaeii oeae aeniiaie aoei i?aeoaa?aeaeaii
eaaioiaio?i?/ieie ?ic?aooieaie aenieiai ??aiy neeaaeiino? (Leszczynski,
?poner et al., 1996). Aia?oa oeyoii c?noaaeaiiy aaee/eie aa?’??o
ieiueiii? ?iaa?n?? c aenia?eiaioaeueiith /anoioith a?yeiaiai
(?iaa?niiai) eieeaaiiy ai?iiiai o?aaiaioa cao?eniaaii iaaeaa?’??iee
oa?aeoa? ieiueiii? ?iaa?n?? a Ade, Cyt ? Gua o aeaeyae? nooo?ai
aiaa?iii?eiiai eieeaaiiy aaeeei? aiie?ooaee, eio?a nyaa? aeanyoe?a
a?aaeon?a. Aia?oa c?iaeaii i?eioeeiiai aaaeeeaee aeniiaie i?i eaaioiao
aaiiao??th oeeo a?ieia?/ii aaaeeeaeo iieaeoe. Oea icia/a?, ui a?aooaaiiy
ioeueiai? eieeaaeueii? aia?a?? nooo?ai ci?ith? aaiiao??th iieaeoee, yea
a?aeiia?aea? aeiaaeueiiio i?i?ioio. Iaa?oue i?e O > 0 E Ade, Cyt ? Gua ?
aoaeoeaii ieaia?ieie iieaeoeaie (aoaeoeaia neiao??y NS), i?e oeueiio
aaee/eio a?aeoeeaiiy a?ae ieiueiiino? ai?iiiai o?aaiaioa o aeiaaeueiiio
i?i?ioi? AIIA iiaeia ?icaeyaeaoe ye ioe?ieo cieco aiie?ooaee eiai
a?yeiaiai eieeaaiiy. Oaeei /eiii, aaeaeiny iiaeieaoe ?nioth/o aeeeaio
«ieaia?i?noue-iaieaia?i?noue» uiaei aoaeiae ioeeaioeaeieo iniia c
ai?iia?oiith. Iei? aenia?eiaioaeuei? aeai? uiaei no?oeoo?ii?
iaaei?noeino? acioenoeo iniia c ai?iia?oiith a?aenooi?, i?ioa, ia iao
iiaeyae, iaee?auei ?? aeieacii ? iai?eaeaoi?noue aa?iii?eiiai
iaaeeaeaiiy aeey ?ioa?i?aoaoe?? /anoio aeaoi?iaoe?eieo eieeaaiue
ai?iiiai o?aaiaioa.

C?iaeaii ocaaaeueiaiiy uiaei ai?cio?ii?? iiaa?oaiiy ai?iia?oie –
iaea?eueoa ?? aaee/eia niinoa??aa?oueny o oeo aeiaaeeao, eiee ai?iia?oia
ioi/aia c iaeiiai aieo aiaeioeeee?/iei aoiiii acioo, a c ?ioiai –
aecioeeee?/iei ca’yceii NH /e CH, ye oea ia? i?noea a Cyt (5 eeae/iieue)
? Gua (4,4 eeae/iieue ca aeaieie AI1). Ca oeeo oiia ai?iia?oi? cia/ii
aea?aei?oa iiaa?oaoeny ai?iii?ioiiaie o a?e non?aeiueiai iiaea?eiiai
ca’yceo C=N, ai ia?ao?aeiee noai iiaa?oaiiy o oeueiio aeiaaeeo
noaa?e?co?oueny eoeii?anueeith aca?iiae??th ai?iieo aoii?a aiaeith c
aiaeioeeee?/iei aoiiii acioo ? AAI ai?iiiai aoiia acioo c aoiiii aiaeith
non?aeiuei? a?oie NH /e CH. Oeth aca?iiae?th iiaeia ?icaeyaeaoe ye
aioo??oiueiiieaeoey?i? I-ca’ycee, ui aca?iiiineeththoueny. Iiaa?oaiiy
ai?iia?oie o cai?ioi?e a?e cia/ii oneeaaeiaia aeanoaa?e?caoe??th
ia?ao?aeiiai noaio eoeii?anueeei a?aeooiaooaaiiyi AAI ai?iiiai ?
non?aeiueiai aiaeioeeee?/iiai aoiia acioo, a oaeiae ai?iieo ? aoiia
aiaeith a?oie NH /e CH. Oaeei /eiii, aia?oa anoaiiaeaii, ui ai?iia?oia a
eaiii?/ieo ioeeaioeaeieo iniiaao ia iaa?oa?oueny ?cio?iiii ia 360° aac
aeaoi?iaoe?e, ye oea o?aaeeoe?eii aaaaeaeiny ?ai?oa, a ia?aaaaeii
iiaa?oa?oueny a iaiaaeaiiio naeoi?? (180°) aeaia?aiieo eoo?a, ?noioiuei
i?e oeueiio aeaoi?ioth/enue. C?iaeaii aeniiaie, ui iniiaieie
noa?aio?i?/ieie /eiieeaie no?oeoo?ii? iaaei?noeino? acioenoeo iniia c
ai?iia?oiith ? ??-ni?yaeaiiy AAI ai?iiiai aoiia acioo c ?-aeaeo?iiiith
nenoaiith e?eueoey, Oa?i?- a?aeooiaooaaiiy i?ae ai?iiei ca’yceii NI ?
non?aei?i ca’yceii NI /e CI, eoeii?anueea aca?iiae?y AAI ai?iiiai aoiia
acioo ye c AAI non?aei?o aoii?a, oae ? c aoiiii aiaeith non?aeiueiai
ca’yceo NI /e CI. Aia?oa anoaiiaeaii aca?iicaeaaei?noue no?oeoo?ii?
iaaei?noeino? acioenoeo iniia c ai?iia?oiith a?ae oaooiia??? ?
aaiiao?e/ii? ?ciia???.

Iayai?noue iecueeeo (( 200 ni-1) /anoio a eieeaaeueieo niaeo?ao
acioenoeo iniia, ui a?aeiia?aeathoue iaieaia?iei aeaoi?iaoe?yi e?eueoey,
aeaco? ia eiai eaac?aei?noe?noue: ?ic?aooiie ca oi?ioeith aa?iii?eiiai
inoeeeyoi?a c aeei?enoaiiyi e?oa?aoo?ieo aeaieo aeey neeiaeo iino?eieo
iaieaia?ieo eieeaaiue e?eueoey (10 — 20 eeae/iieue) aea? aeey ioeueiai?
aiie?ooaee aaee/eio, ui ia?aaeuo? 10°. Oey aeaiaioa?ia ioe?iea
oe?eeiaeoi ocaiaeaeo?oueny c aeine?aeaeaiiyie eiioi?iaoe?eii? aio/eino?
aaoa?ioeeee?a ioeeaioeaeieo iniia eaaioiaio?i?/ieie iaoiaeaie aenieiai
??aiy neeaaeiino? (Oeoe?i, 1999).

Aia?oa iieacaii, ui i?e ia?aoiae? a?ae iniiae aei ioeeaiceaeo
niinoa??aa?oueny neaaea oaiaeaioe?y aei i?aeaeuaiiy aa?’??o ieiueiii?
?iaa?n?? ai?iiiai o?aaiaioa ? a?aeiia?aeii cieaeaiiy aa?’???a iiai?ioii?
?ooiiino? ai?iia?oie. Ia a?aei?io a?ae iniia oa ?oi?o iaoeeueiaaieo ii
ae?eiceaeiiio acioo aiaeia?a, aea ieiueiia ?iaa?n?y ai?iiiai o?aaiaioa
a?aeaoaa?oueny o aeca?eaeueii-neiao?e/iiio aae?aaaoe/iiio iioaioe?ae?, a
ioeeaiceae? oeae iioaioe?ae noa? aneiao?e/iei, i?e/iio aaee/eia
aneiao??? iii?oii iaioa ca aa?’?? ?iaa?n??. Oea icia/a?, ui aeey
ioeeaioeaeieo iniia c ai?iia?oiith o neeaae? ioeeaiceae?a iiyaey?oueny,
iaoae ? o aeeiai?/iiio nain?, aeae?eaiee iai?yiie «aa?o-iec».

Aeei?enoiaoth/e iaoiae MNDO/I, aia?oa aaeaeiny iieacaoe, ui eaiii?/i?
ioeeaioeaei? iniiae ioiieai? i?inoi?iaith n?oeith aca?iicaeaaeieo
aioo??oiueiiieaeoey?ieo I-ca’yce?a oeio NI…I ? NI…N c aia?a??th 0,7-3
eeae/iieue (yeo o?aaa ?icaeyaeaoe ye ioe?ieo caa?oo). Oae? oyaeaiiy
i?aee??ieththoueny aenia?eiaioaeueieie aeaieie: oae, cie?aia, iaeneaaoee
(0,7 eeae/iieue) aioo??oiueiiieaeoey?iee I-ca’ycie N6I’’…N7 a Ade
cao?eniaaii ?aioaaiino?oeoo?ieie iaoiaeii o e?enoae?/iiio noai?
(Jeffrey, Saenger, 1994), a iaeneeuei?oee (3 eeae/iieue) N4I’…N3 a Cyt
ca?a?no?iaaii iaie iaoiaeii ssI?-niaeo?ineii?? o aacaiaeiiio
aeeiaoeenoeueoieneae?. Aeey aioo??oiueiiieaeoey?ieo I-ca’ycea? ca
o/anoth ai?iia?oie oa?aeoa?ii oa, ui i?e ?? ai?cio?iiiiio iiaa?oaii?
aiie ?noioiuei ci?iththoue ia eeoa naith aia?aaoeeo, ia?aieeath/enue c
iaeieo aoiiieo a?oi ia ?io?, a e e?euee?noue: ?oiy aia?a?y, eiiia?aoeai?
aeanoeaino?, oei ? iaa?oue e?euee?noue ia ? noaoe?iia?ieie
oa?aeoa?enoeeaie.

I?ioio?iiia oaooiia??y. Aia?oa a i?aeoeoe? eaaioiaio?i?/iiai
aeine?aeaeaiiy yaeua i?ioio?iiii? oaooiia??? acioenoeo iniia «c
i?eoe?eii» ia ?? a?ieia?/io cia/ei?noue oey caaea/a ?ica’ycaia o
iaeneiaeueii iiaeeeaiio oi?iao?, iaiaaeaiiio eeoa oyaeaiiyie i?i
i?e?iaeo o?i?/iiai ca’yceo, aeey aeineoue aaeeei? e?eueeino? iieaeoe, ui
iine?aeiaii oneeaaeiththoueny: eaiii?/ieo iniia AeIE, ?oi?o
aeacai?iiaaieo aiaeia?a oa neeaaeiaeo /anoei — io?eio, i??ei?aeeio ?
?i?aeacieo (Im). Oaeee i?aeo?ae, aeicaieea cao?enoaaoe ieceo iiaeo
o?ceei-o?i/ieo caeiiii??iinoae i?ioio?iiii? oaooiia??? acioenoeo iniia.

Oae, aia?oa anoaiiaeaii, ui oea yaeua ia? iieaeoey?ii-oea?oa??iiiee
oa?aeoa? – a iueiio aa?ooue o/anoue on? aac aeiyoeo i?ioiie, aeeth/ath/e
ea?aii?ioiie. Iieacaii, ui io?eiia? iniiae oa Im iathoue cia/ii a?eueoo
noeeuei?noue aei oea?oa??iiii? oaooiia???, ai?ae i??ei?aeeiia?. Aeyaeaii
ni?eueio no?oeoo?io ?eno iieaeoey?ii-oea?oa??iiieo oaooiia??a iniia, ui
iathoue o nai?io neeaae? aoiie eenith, – ia?aaaaeio oeen-i???ioaoe?th
a?ae?ieneeueieo a?oi a?aeiinii non?aeiueiai aiaeioeeee?/iiai aoiia
acioo. Aeiaaaeaii, ui ieaia?ia aoaeiaa iniiaiiai oaooiia?a ia ?
iane??ciith aeanoea?noth an??? iiiaeeie oaooiia??a.

Aia?oa c’yniaaii, ui aeey eiaeii? acioenoi? iniiae io?eiiaiai ?yaeo oa
Im iniiaiei oaooiia?ii-oea?oa??iiii c-iii?ae oeo, ui oi?iothoueny oeyoii
i?a?aoe?? ea?aii?ioii?a, ? ?e?aeia oi?ia, ui ooai?th?oueny a ?acoeueoao?
ia?aoiaeo i?ioiia i?e aoii? aoaeaoeth N8 (N2 a Im) ia non?aei?e aoii
acioo (N7 — o io?eiao, N2 — a Im). A?ieia?/ia cia/ou?noue ??, cie?aia,
iieyaa? a oiio, ui aiia ? ia?ao?aeiei noaiii ?aaeoe?? I > T(D) — iai?io
a?oie N8I io?eiiaeo iniia ? a?oie N2I Im o aiae? i?e ni?eyoeeaeo pH:
iieaeoey?ii-e?iaoe/iee iaoai?ci ?? oi?ioaaiiy — oea anoaoaoia
i?ioiioaaiiy (aoiia N7 io?ei?a ? aoiia N3 Im) – aeai?ioiioaaiiy (ca’yceo
N8I io?ei?a ? N2I Im) ca o/anoth i?ioii?a c aiaeiiai na?aaeiaeua.
Anoaiiaeaii, ui o aeiaaeeo io?eiiaeo iniia, ye? iathoue o nai?io neeaae?
ai?iia?oio, iaa?oue ia iecueeiiieaeoey?iiio ??ai? i?ioean iai?io ?
eiioi?iaoe?eii-/ooeeaei, ine?eueee ia?ao?ae c iniiaii? oaooiia?ii? oi?ie
a ?e?aeio noi?iaiaeaeo?oueny ?noioiei cao?aiiyi no?oeoo?ii-aeeiai?/ieo
aeanoeainoae ai?iiiai o?aaiaioa. Oea icia/a?, ui oaeaee?noue iai?io a
a?oi? N8I Ade ? Gua aecia/a?oueny ia eeoa eiioi?iaoe??th IE, aea e ??
cao?aiiyi i?e ia?aoiae? a ?e?aeio oi?io. C?iaeaii i?eiouaiiy i?i o/anoue
?e?aeieo oaooiia??a io?eiiaeo iniia a i?ioeanao ?oiueiai iioeiaeaeaiiy
eeniaaeie ?aaeeeaeaie.

Iaa?oioiaaii oi/eo ci?o i?i cia/ii a?eueoo a?ieia?/io cia/ou?noue
i?ioio?iiii? oaooiia??? acioenoeo iniia, ai?ae oea i?eeiyoi iei?. Oea
noino?oueny, ianaiia?aae, aenieiaia?aaoe/ieo oaooiia??a, cie?aia
oea?oa??iiieo, ye? aaciina?aaeiuei ia o?enothoueny o?ceei-o?i?/ieie
iaoiaeaie, a ia?oo /a?ao — niaeo?ineii?/ieie, a oiiaao oa?iiaeeiai?/ii?
??aiiaaae ? aiiiaaiiino? ioi/aiiy (aaeooi, iecueeioaiia?aoo?ia ?ia?oia
iao?eoey, ?ic/ei oiui), — oae caaieo «i?eoiaaieo no?oeoo?».
Noaa?aeaeo?oueny, ui a oiiaao nooo?ai? aaoa?iaaiiino? eieaeueiiai
ioi/aiiy, ye oea ia? i?noea o aenieiiieaeoey?ieo a?eeiai-ioeea?iiaeo
eiiieaenao, a?eueo?noue, yeui ia an? oaooiia?e o??? /e ?ioi?
ioeeaioeaeii? iniiae, iiaeooue aooe cao?eniaai? I-ca’yceaie o
a?aeiia?aeieo i?noeyo ai?ciaaaiiy. I?eioeeiiaeo o?ceei-o?i?/ieo
ia?aoeiae aeey oeueiai ia ?nio?, ine?eueee oa?aeoa?iee aeey ieo
ae?aiacii aia?aaoe/ii? noaa?eueiino? (( 30 eeae/iieue) c iaaeeeoeii
iiaea aooe ia?ae?eoee aoyaoaaiiyi ??aee?nieo oaooiia??a o I-ca’ycee ?c
e?eueeiia a?/ieie ?aaeeeaeaie ai?iieeneio (iniaeeai ca?yaeaeaieie), ui
ooai?ththoue i?noea ai?ciaaaiiy. Oa?aeoa?iei ooo ? i?eeeaae a?ineioaco
io?eiiaeo ioeeaioeae?a — iaiao?aeiith eiai ia?aaeoiiaith, ye a?aeiii, ?
ia?aaaaeaiiy iniiae ?c iniiaii? o ??aee?nio oaooiia?io oi?io N7I c
a?aee?eoei aeey i?e?aeiaiiy oeoe?iaiai caeeoeo aoiiii acioo N9
?i?aeacieueiiai oeeeeo, yea ?aae?co?oueny oa?iaioii. Ie??i oiai, a
e?enoae? isoCyt ni?a?niothoue aea? oaooiia?i? oi?ie — iniiaia, c
eieae?caoe??th ?i?iii?ioiia i?e aoii? N3, ? ??aee?nia, c eieae?caoe??th
?i?iii?ioiia i?e aoii? N1, aia?a?? yeeo ?noioiuei a?ae??ciythoueny – ia
12,2 eeae/iieue (ca aeaieie AI1). Oea iieaciaee i?eeeaae oiai, ye
iaa?oue o aiiiaaiieo oiiaao ca ?aooiie caeaaeiino? eiiieaenioa??ii?
caeaoiino? iniiae a?ae ?? oaooiia?iiai noaio i?aeiieaeoey?i? I-ca’ycee
noaa?e?cothoue ??aee?nio oaooiia?io oi?io c aeineoue aenieith aia?a??th.
Ioaea, iai?aaii??ii noaaeoe o i?yio caeaaei?noue, ye oea o?aaeeoe?eii
?iaeoueny, a?ieia?/io cia/ou?noue o??? /e ?ioi? oaooiia?ii? oi?ie a?ae
?? aia?aaoe/ii? noaa?eueiino?.

Aia?oa iinooeueiaaii iiaeeea?noue aca?iiia?aoai?aiiy aeayeeo oaooiia??a
acioenoeo iniia ca ?aooiie ia?aianaiiy i?ioiio acaeiaae
aioo??oiueiiieaeoey?iiai I-ca’yceo. I?ci?o? ?ic?aooiee aenieiai ??aiy
neeaaeiino? eii?aeeiaoe ?aaeoe?? aioo??oiueiiieaeoey?iiai ia?aianaiiy
i?ioiio a Gua (a?ae aoiia N1 ia aoii eenith), Cyt (c ai?iia?oie ia aoii
N3) ? aeayeeo ?ioeo iniiaao AeIE (Leszczynski, Gorb, 1998-1999)
iiai?noth oea i?aeoaa?aeeee. A?eueoa oiai, oe? ?ic?aooiee ? /e ia
iaee?auei aeiaeaoeiaei na?ae/aiiyi iayaiino? a ioeeaioeaeieo iniiaao
aioo??oiueiiieaeoey?ieo I-ca’yce?a.

Caeaoi?noue i?e?aeioaaoe aai a?aeuaiethaaoe i?ioii. I?ea?iaeuei?noue
cai?iiiiiaaiiai i?aeoiaeo aeey ae??oaiiy oe??? caaea/? iieyaa? o
aeei?enoaii? oe?ieiai (oaeiai ae, ye ? o aeiaaeeo aea/aiiy i?ioio?iiii?
oaooiia???) iaai?o ia’?eo?a aeine?aeaeaiiy. Oea aeicaieeei anoaiiaeoe
ni?euei? ? a?aei?ii? ?ene ye i?ioiiiaeiii?ieo-i?ioiiiaeoeaioi?ieo
aeanoeainoae acioenoeo iniia, oae ? ?oiuei? caeaoiino? aeaoi?ioaaoeny
i?e ci?i? ca?yaeiaiai noaio, a oaeiae i?ine?aeeoaaoe, ye aiie
ci?iththoueny c oneeaaeiaiiyi no?oeoo?e iieaeoee.

Oae, aeaoaeueiee aiae?c aaee/eie oa oa?aeoa?o no?oeoo?ieo cao?aiue, ui
iathoue i?noea i?e i?ioiioaaii? oa aeai?ioiioaaii? iniiae, cana?ae/o?,
ui iaeneiaeuei? ci?ie aaee/eie aaiiao?e/ieo ia?aiao??a (aaeaioieo
ca’yce?a, aaeaioieo oa aeaia?aiieo eoo?a) niinoa??aathoueny iiaeeco
i?noeue i?e?aeiaiiy /e a?aeuaieaiiy i?ioiia, ciaiooth/enue ia ia?eoa???,
i?e oeueiio cao?aiiy aaiiao?e/ieo ia?aiao??a noi?iaiaeaeo?oueny ?noioiei
ia?a?iciiae?eii ca?yae?a ia aoiiao. Acioeno? iniiae, ye? ia iathoue o
nai?io neeaae? ai?iia?oie, ia ci?iththoue ieiueiiiai oa?aeoa?o nai??
aoaeiae i?e i?ioiioaaii? /e aeai?ioiioaaii? iacaeaaeii a?ae eiai i?noey.
Aia?oa anoaiiaeaii, ui acioeno? iniiae c ai?iia?oiith iiaiaeyoue naaa
?iaeoa: iae?noioi?oeo cao?aiue, ye? nyaathoue aeanyoe?a a?aaeon?a i?e
ci?i? ?oiueiai ca?yaeiaiai noaio oeyoii i?ioiioaaiiy /e aeai?ioiioaaiiy,
caciathoue «iaei’yeo?» aaiiao?e/i? ia?aiao?e – aeaia?aii? eooe, ui
oa?aeoa?ecothoue i??ai?aei?noue ai?iiiai o?aaiaioa (CNI2. I?ioiioaaiiy
iniia c ai?iia?oiith, iacaeaaeii a?ae eiai i?noey (aeiyoie neeaaea? eeoa
ai?iiee aoii acioo), i?ecaiaeeoue aei iiaiiai ?oiueiai nieiuaiiy – ?oiy
?ioa?eiiaa?n?y a?aeaoaa?oueny eeoa oeyoii ai?cio?iiiiai iiaa?oaiiy
ai?iia?oie iaaeiei aecioeeee?/iiai ca’yceo CN, i?e/iio aaee/eie aa?’???a
iiaa?oaiiy c?inoathoue o ii??aiyii? c aeaeo?iiaeo?aeueiith iniiaith.
Aeai?ioiioaaiiy iniia c ai?iia?oiith iaaiaee – cia/ii iineeth?
iaieiueii?noue ?oiueiai ai?iiiai o?aaiaioa ? aeaui cii?uo? e?eueoea. I?e
oeueiio aaee/eia aoaeoo oa eiai oa?aeoa? caeaaeeoue a?ae i?noey
aeai?ioiioaaiiy iniiae. ?oiy ?ioa?eiiaa?n?y a?aeaoaa?oueny, ye ? o
aeaeo?iiaeo?aeueieo aiaeia?a, o?ueiia oiiieia?/ii ? aia?aaoe/ii
iaaea?aaeaioieie oeyoaie, aea ia a?aei?io a?ae inoaii?o, c?inoa? aa?’??
ieiueiii? ?iaa?n?? ? a?aeiia?aeii ciaioothoueny aa?’??e aioo??oiueiai
iiaa?oaiiy ai?iia?oie. O ai?ii?a Ade, iai?eeeaae, aa?’?? ieiueiii?
?iaa?n?? c?inoa? a?eueoa, i?ae ia ii?yaeie o ii??aiyii? c
aeaeo?iiaeo?aeueiith iniiaith, ? iiaeia aiaaiaii aiai?eoe i?i ?oith
iaieaia?io eiioi?iaoe?th. Inoaii? noino?oueny aneo aac aeiyoeo ai?ii?a
iniia c ai?iia?oiith. Oa?aeoa? ia?aaoaeiae AIIA ioeeaioeaeieo iniia c
ai?iia?oiith i?e ci?i? ?oiueiai ca?yaeiaiai noaio ia?aeiieeai aeaco? ia
oa, ui iaeiei ?c aieiaieo noa?aiaeaeo?iiieo /eiiee?a, ui aecia/athoue
?oith no?oeoo?io iaaei?noe?noue, ? ??-ni?yaeaiiy AAI ai?iiiai acioo c
?-aeaeo?iiiith nenoaiith e?eueoey. I?i oea oaeiae aiai?eoue ca?eueoaiiy
aeiaaeeie aecioeeee?/iiai ca’yceo CN i?e i?ioiioaaii? ai?iiiai aoiia
acioo (ia ~ 0,09 – 0,1 A ca aeaieie AI1) aiane?aeie oiaiiaeeeaeaiiy
??-ni?yaeaiiy. Aia?oa cao?eniaaii aaaeeeaee c a?ieia?/ii? oi/ee ci?o
aoaeo – iiaeeea?noue aeenoaioe?eiiai oi?aae?iiy no?oeoo?iith
iaaei?noe?noth iniiae c ai?iia?oiith /a?ac ci?io ca?yaeiaiai noaio
ioeeaiceaeo oeyoii aeai?ioiioaaiiy eiai oeoe?iaiai caeeoea ii i?noeyo
I3’ ? I5’.

Aia?oa iiaoaeiaaii ?yaee eeneioieo (aeeth/ath/e NI-eeneioi?noue) ?
eoaeieo aeanoeainoae aeey aneo aeine?aeaeaieo iieaeoe. Anoaiiaeaii, ui
aiie ia ei?aeththoue c oaeeie aeaeo?iiieie oa?aeoa?enoeeaie
a?aeiia?aeieo aoiiieo a?oi, ye ca?yaee ia aoiiao /e ?oi? aonoeie. Oea
iiynith?oueny, cie?aia, oei, ui AAI aoiia-aeoeaioi?a i?ioiio a oeeo
iieaeoeao i?eioeeiiai ia iiaeooue aooe ioioiaeiai? c iaei??th
iaca’ycoth/ith iieaeoey?iith i?a?oaeeth, no?iai ia iueiio eieae?ciaaiith
(E?aoaiieeia e ae?., 1978). Iieacaii, ui c-iii?ae aneo aeine?aeaeaieo
eaiii?/ieo iniia AeIE iaeiioeiaeuei?oa ni?a?nioaaiiy eeneioii-eoaeieo
aeanoeainoae ia? i?noea a Gua. Ia a?aei?io a?ae Gua, Cyt ?
iaei?ioio?euei?oith iieaeoeith c iaea??oeie eeneioieie aeanoeainoyie; c
?ioiai aieo – iieaeoea Thy ? iaeeene?oith iniiaith. Aia?oa iiyniaii, ui
cao?eniaaia aenia?eiaioaeueii (Greco et al., 1990) iaa?iaiiy ?yaeo
i?ioio?eueiino? (Gua>Cyt) oe?eeiaeoi iia’ycaia c i?ioio?iiiith
oaooiia???th oeeo iniia. Iaa?oue o aea?/? iaoeeueiaaiiio aoai?i?
m21,9Gua eiai i?ioio?euei?noue (228,8 eeae/iieue ca aeaieie AI1)
iii?oii iaioa, i?ae i?ioio?euei?noue m1Cyt (230,3 eeae / iieue).

Ia i?eeeaae? aeaio aia?aaoe/ii iaeaea?aei?oeo oaooiia??a N7I ? N9I Xan ?
Hyp aia?oa iieacaii, ui eeneioii-eoaei? aeanoeaino? iniia ?noioiuei
caeaaeaoue a?ae ?oiueiai oaooiia?iiai noaio. Ia i?eeeaae? io?eio oa eiai
neeaaeiaeo /anoei – i??ei?aeeio ? Im i?ine?aeeiaaii, ye i?ioiioaaiiy oa
aeai?ioiioaaiiy iniiae aieeaa? ia ?? eeneioii-eoaei? aeanoeaino?.
Anoaiiaeaii, ui a i?ioiiiaaieo ? aeai?ioiiiaaieo niieoeao
i??i?eoaoi?noue i?noeue i?ioiioaaiiy ? aeai?ioiioaaiiy ?noioiuei
ci?ith?oueny o ii??aiyii? c aeaeo?iiaeo?aeueieie aiaeiaaie.

Ieeueio oaaao i?eae?eaii NI-eeneioiei aeanoeainoyi acioenoeo iniia. Oea
iia’ycaii c oei, ui I-ca’ycee ca o/anoth a?oi NI, ye aeiii?a i?ioiio,
iii?e naith a?aeiinio neaae?noue o ii??aiyii? c eeane/ieie I-ca’yceaie,
a?ae?a?athoue iaia?an?/io ?ieue o no?oeoo?i?e ? iieaeoey?i?e a?ieia??
(Zhurkin, 1998). Aia?oa anoaiiaeaii, ui acioeno? iniiae ye NI-eeneioe
?noioiuei a?ae??ciythoueny a?ae «ii?iaeueieo» i?aai?/ieo NI-eeneio,
i?ioiiiaeiii?ia caeaoi?noue eio?eo aecia/a?oueny aaee/eiith ca?yaeo ia
aoii? aiaeith a?oie NI (Iiai?aeue e ae?., 1984). Ia cao?eniaaii
i?eeiyoiiai aeey i?aeoe/iiai canoinoaaiiy ??aiy ei?aeyoe?? i?ae
aia?a??th aeai?ioiioaaiiy a?oi NI acioenoeo iniia, c iaeiiai aieo, ?
oaeeie ia?aiao?aie, ye ca?yae ia ?oiueiio aoii? aiaeith, aeiaaeeia
ca’yceo NI ? /anoioa eiai aaeaioiiai eieeaaiiy, c ?ioiai.

No?oeoo?ia ?ciia??y. Aia?oa aeine?aeaeaii no?oeoo?io ?ciia??th acioenoeo
iniia oa ?? iniiai? o?ceei-o?i?/i? caeiiii??iino?. Oae, aiae?c
aia?aaoe/ieo oa?aeoa?enoee iiaiiai n?iaenoaa no?oeoo?ieo ?ciia??a
ai?iiio?eio, ai?iiieneio?eio, ai?iiienei??ei?aeeio oa N-iaoeeo?aoeeeo
aeicaieea aia?oa aeyaeoe aeiyoeiao aeanoea?noue aeaio eaiii?/ieo
ioeeaioeaeieo iniia — Ade ? Thy : a nai?o n?iaenoaao aiie ? aia?aaoe/ii
iaeaea?aei?oeie no?oeoo?ieie ?ciia?aie. Ia a?aei?io a?ae Ade oa
eiiieaiaioa?iiai eiio Thy aea? ?io? eiiieaiaioa?i? iniiae AeIE – Gua oa
Cyt ia ? aia?aaoe/ii iaeaea?aei?oeie iieaeoeaie a nai?o n?iaenoaao
no?oeoo?ieo ?ciia??a. Na?aae n?iaenoaa no?oeoo?ieo ?ciia??a ai?iiio?eio
aia?aaoe/ii iaeaea?aei?oei ? i8Ade – i?iaeoeo ieeneaiiy Ade – oaieioa
eiai ooai?aiiy ia 11,7 eeae/iieue ieae/a, i?ae aiaeia?/ia aaee/eia aeey
Gua; na?aae iiaii? iiiaeeie ?ciia??a ai?iiienei??ei?aeeio aia?aaoe/ii
iaeaea?aei?oith no?oeoo?ith ? 5-aminoi4Pur – a?i ia? oaieioo ooai?aiiy
ia 7,2 eeae/iieue ieae/o, i?ae Cyt. Ie??i oiai, Ade ia? ua iaeio
aeiyoeiao ?eno: eeoa aeey oe??? iniiae aia?aaoe/ia a?aenoaiue i?ae iei ?
non?aei?i o n?iaenoa? no?oeoo?iei ?ciia?ii, ui ia? ?io? eiaeia?
aeanoeaino?, iaea?eueoa (5,3 eeae/iieue). I?e oeueiio 2- aminoPur, yeee,
ye a?aeiii (Sowers et al., 1987), ? aeineoue neeueiei iooaaaiii, a
n?iaenoa? aaiiao?e/ieo ?ciia??a ai?iiio?eio ia? iaeaeuo aia?a?th –
oaieioa eiai ooai?aiiy ia?aaeuo? aiaeia?/io aaee/eio aeey Ade ia 7,8
eeae/iieue (ca aeaieie AI1).

Io?eiai? ?acoeueoaoe aeicaieythoue, ia iao iiaeyae, ?icaeyaeaoe
eaiii?/i? iniiae AeIE a eiioaeno? iieaeoey?ii? aaiethoe?? ? iia’ycaoe
ia?oi/a?aia?noue ooai?aiiy Ade – ?aeeii? iniiae, yea ia ia? o nai?io
neeaae? eenith, – o aa?iaaiiiio neioac? c eiiiiiaio?a ia?aeiii?
aoiinoa?e ? iiaeaeueoa «aeaeeaaiiy» a i?e (Iaaeianeay, 1973) ia eeoa c
iaea?eueoith aia?a??th aeaeo?iiiiai ni?yaeaiiy (Pullman, 1962), a e c
iaeiaioith oaieioith eiai ooai?aiiy o nai?io n?iaenoa? no?oeoo?ieo
?ciia??a. I?i iniaeeaa i?noea Ade na?aae eaiii?/ieo ioeeaioeaeieo iniia
na?ae/aoue oaeiae ? iaeiaioa eiai noeeuei?noue c-iii?ae iniia AeIE aei
oaeeo aaiioiene/ieo no?oeoo?ieo ia?aoai?aiue, ye i?ioio?iiia oaooiia??y
i??ei?aeeiiaiai e?eueoey, niiioaiia aeacai?ioaaiiy ? ieeneaiiy o aiae?
oiui. An? oe? aeai? iiaeia ?icoe?ithaaoe ye a?aoiaioe ia ei?enoue
a?aeiii? a?iioace (Rich, 1964) uiaei oiai, ui ia ?aii?o aoaiao
aaiethoe?? no?oeoo?a ia?aeiieo iie?ioeeaioeae?a aoea cia/ii
iiiioiii?oith ? aiie i?noeee eeoa aea? eiiieaiaioa?i? iniiae – Ade ?
Thy. Ie??i oiai, ?o iiaeia ?icaeyaeaoe ye iaeia ?c iiaeeeaeo iiyniaiue,
/iio Ade ia? o aaiii? iaeaeuo eiaeiao oe?ii?noue (Iacei, 1980), /iio
naia AOO aeei?enoiao?oueny o aeea?e i?e?iae? ye oi?aa?naeueia
«aia?aaoe/ia aaethoa» oiui.

Io?eiai? ?acoeueoaoe uiaei no?oeoo?ii? ?ciia??? ioeeaioeaeieo iniia
i?ieeaathoue oaeiae na?oei ia oa, /iio naia caaaa/ai? ia ia?e Gua:Cyt
ae?eyiee AeIE ? iioaioe?eiei aeaea?aeii oi/eiaeo iooaoe?e (Bernardi,
1995). Oe?eeii eiia??ii, ui iaei??th c i?e/ei oeueiai ? aeaa?aaeaoe?y
Gua ? Cyt a iecueeiaia?aaoe/i? no?oeoo?i? ?ciia?e, ?iaeoeiaaia
??ciiiai?oieie o?ceei-o?i?/ieie /eiieeaie aeciaaiiiai iioiaeaeaiiy,
cie?aia OO-ii?ii?iaiiyi.

Niiioaiia aeacai?ioaaiiy. A?aoiaoth/e, ui ?iaeoeiaaia oaieii ca
iayaiino? aiaee aeacai?ioaaiiy iniia AeIE a?ae?a?a? iaaaeyeo ?ieue o
niiioaiiiio iooaaaiac?, ni?e/eiyth/e o?aiceoe?? (Aeaieeia oa ?i., 1971),
aoei aeine?aeaeaii aia?aaoe/i? (aioaeuei?th) oa no?oeoo?i? (ci?ie
noa?aio?i?/ii? iaaei?noeino? oa oaooiia?iiai noaio) eiai iniaeeaino?
aeey eaiii?/ieo ioeeaioeaeieo iniia oa iecee ?oi?o aiaeia?a – i?ii?ieo
iniia AeIE, i?iaeoeo?a aee?eoaaiiy oa iioeiaeaeaiiy eeniaaeie
?aaeeeaeaie, no?oeoo?ieo ?ciia??a, iieaeoe, ui iathoue oa?aiaaoe/ia
cia/aiiy, iooaaai?a, iiaeaeueieo niieoe (anueiai 38 no?oeoo?), ui
iathoue ai?iia?oio, o a?eueiiio noai?.

Aia?oa io?eiaii oae? ?acoeueoaoe.

Ca noeeuei?noth aei niiioaiiiai aeacai?ioaaiiy eaiii?/i? iniiae AeIE
ooai?ththoue ?yae Gua > Cyt > Ade. Eeoa aeey Ade oa ia?aaaaeii?
a?eueoino? eiai iio?aeieo oeae i?ioean ? aiaeioa?i?/iei; aeey Gua, Cyt
oa aneo aac aeiyoeo ?oi?o iio?aeieo a?i ia? aecioa?i?/iee oa?aeoa?.
Na?aae no?oeoo?ieo ?ciia??a ai?iiio?eio iaeiaioo noeeuei?noue aei
niiioaiiiai aeacai?ioaaiiy ia? eeoa Ade. Iaoeeueiaai? ii ae?eiceaeiiio
ca’yceo eaiii?/i? iniiae AeIE ca nai?th caeaoi?noth niiioaiii
aeacai?ioaaoeny ooai?ththoue ?yae m1Cyt > Cyt, m9Gua < Gua ? m9Ade >
Ade. Iaoeiiaa iiaaae?iea Gua ? m9Gua iiynith?oueny o?enaoe??th
iaoeethaaiiyi aenieiaia?aaoe/ii? oaooiia?ii? oi?ie i?iaeoeoo
aeacai?ioaaiiy inoaiiueiai a oi?i? m9Xan. I?iaeoeoe aeacai?ioaaiiy aneo
aeine?aeaeaieo niieoe, ui iaeaaeaoue aei eeano no?oeoo?ii-iaaei?noeeo
iieaeoe, ? ieaia?ieie eaac?aei?noeeie iieaeoeaie. O ia?aaaaei?e
a?eueoino? aeiaaee?a aeacai?ioaaiiy ia ni?e/eiy? oaooiia?ieo ia?aoiae?a.

Niiioaiia iioeiaeaeaiiy eeniaaeie ?aaeeeaeaie. A?aoiaoth/e a?ieia?/io
cia/ou?noue oeueiai i?ioeano aoei aea/aii eiai aia?aaoe/iee
(aioaeuei?th) oa no?oeoo?iee (cai?io noa?aio?i?/ii? iaaei?noeino? ?
oaooiia?iiai noaio) aniaeoe. Caaea/a c iiae?aiith iinoaiiaeith (iniaeeai
a ia/enethaaeueii-oai?aoe/i?e ieiuei?) ia iiiaaeaea a iiea ci?o
aeine?aeiee?a. I?eeiath/e aei oaaae, ui iaea?eueoo a?ieia?/io
cia/ou?noue iathoue i?iaeoeoe iioeiaeaeaiiy eeniaaeie ?aaeeeaeaie naia
io?eiiaeo iniia (Iieoaa, A?oneia e ae?., 1993), ia’?eoaie aeine?aeaeaiiy
aea?aii Ade ? Gua, ?oi? iaoeeueiaai? ii aeaa’yoiio iieiaeaiith aiaeiae,
ui ? iaei?ino?oeie iiaeaeyie ioeeaiceae?a, i?iaeoeoe aeacai?ioaaiiy oeeo
niieoe, a oaeiae aeaye? “iiaeaeuei?” iieaeoee (Im, Pur, m1Im oa m9Pur).

Aia?oa io?eiaii oae? ?acoeueoaoe. Anoaiiaeaii, ui aeey aneo aac aeiyoeo
aeine?aeaeaieo iieaeoe iae?iia??i?oei i?noeai i?e?aeiaiiy eenith a
i?ioean? ?oiueiai ieeneaiiy ? aoii N8 ?i?aeacieueiiai e?eueoey (N2 – o
aeiaaeeo Im ? m1Im).

Ca noeeuei?noth aei ieeneaiiy aeine?aeaeai? niieoee ooai?ththoue ?yaee
Gua>Ade, m9Gua>m9Ade, i?e/iio m9Ade>Ade, m9Gua>Gua. Inoaii? aeaa
ni?aa?aeiioaiiy iiaeooue na?ae/eoe i?i oa, ui i?e ia?aoiae? a?ae iniiae
aei ioeeaiceaeo noeeuei?noue aei ooeiaeaeaiue eeniaaeie ?aaeeeaeaie
i?aeaeuo?oueny. I?e oeueiio iieaeoee-i?iaeoeoe aeacai?ioaaiiy o
ii??aiyii? c aeo?aeieie iniiaaie c ai?iia?oiith oa ?oi?ie iaoeeueiaaieie
a aeaa’yoiio iieiaeaii? iio?aeieie i?iyaeythoue iaioo caeaoi?noue aei
oeueiai i?ioeano. Ii??aiyiiy a?aeiia?aeieo aioaeuei?e aeey Im (m1Im) oa
io?eiiaeo iniia (iaoeeueiaaieo a aeaa’yoiio iieiaeaii? aiaeia?a) aeaco?
ia cia/io ?oith caeaaei?noue a?ae eiiaeainaoe?? ?i?aeacieueiiai e?eueoey
?c i??ei?aeeiiaei. O aneo aeiaaeeao, ca aeiyoeii Xan, oey eiiaeainaoe?y
i?ecaiaeeoue aei nooo?aiai cieaeaiiy aioaeuei?? ?aaeoe?? ieeneaiiy.
Ieeneaiiy io?eiiaeo iniia nooo?ai ci?ith? ?oith eaac?aei?noe?noue. Oea
iia’ycaii c oei, ui iieaeoea i2Im o ii??aiyii? c Im ? cia/ii i’yeoith
no?oeoo?ith ii a?aeiioaiith aei iaieaia?ieo aeaoi?iaoe?e. Ci?i
oaooiia?iiai noaio i?e ooeiaeaeaii? acioenoeo iniia io?eiiaiai ?yaeo
eeniaaeie ?aaeeeaeaie ia cao?eniaaii.

Aeeeaaeai? aeua ?acoeueoaoe aeicaieythoue c?iaeoe aeoaea aaaeeeaee, ia
iiaeyae aaoi?a, aeniiaie: aieiaia a?aei?ii?noue ioeeaioeaeieo iniia, a
a?aeoae – ? IE a?ae i?aai?/ieo iie?ia??a — i?aaenoaaiee?a iaaeeai?
i?e?iaee e?e?oueny, i/aaeaeii, o oiio, ui aiie ii?aeiothoue o nia?
oi?eaeueii oe?ieee niaeo? o?ceei-o?i?/ieo oi?i no?oeoo?ii? i?ieeaino?,
eiaeia c yeeo i?eoaiaiia ie?aiei eeanai i?aai?/ieo iieaeoe —
i?aaenoaaiee?a iaaeeaiai na?oo. Aiaeii/an oey oa?aeoa?ia aeanoea?noue IE
oa?oue o nia?, ai/aaeaeue, ? iaaaoeaiee nain c oi/ee ci?o o?eaaeino?
aacia?aa?eiiai ?oiueiai ooieoe?thaaiiy. Oe?eeii iiaeeeai, ui naia aiia ?
aeaea?aeii oae caaieo aeiaeaoeiaeo a?io?i?/ieo aeoeaiinoae (Bartosz,
1981), ye? ia «aienothoueny» a eeane/i? noaie iaoaaie?cio ?
ni?e/eiythoue ia?aa?a ia?aiaoaaie?/ieo i?ioean?a, ui i?ecaiaeyoue
a?aoo?-?aoo aei noa??iiy (Aieoaaa, 1996). Iiaeia oaeiae i?eionoeoe, ui
?icia?ooy o?ceei-o?i?/ieo oi?i no?oeoo?ii? i?ieeaino? IE, i?i ye? eoeiny
aeua, ? eiie?aoiei i?e?ino?oeoo?iei iaiiaiaiiyi ?oiuei? «e?iaoe/ii?
aeiaa?oaiino?», yea ?icoe?ith?oueny ye can?a ? ye ?acoeueoao
aaiethoe?eiiai i?ioeano (Oiieue, 1979).

Niaeo?ineii?/ia aeine?aeaeaiiy oaa?aeioacieo eiiieaen?a, ui
iiaeaeththoue oi/eia? a?eeiai-ioeea?iia? ? ioeea?iiai-ioeea?iia?
eiioaeoe

Ia aac? na??eiiai i?eeaaeo AeON-24 noai?aii aeniei/ooeeaee eaca?iee
o?ueioi?iiaiaaee ?aiai?anueeee niaeo?iiao? aeey ?a?no?aoe??
aenieiye?nieo niaeo??a a?iiie?ia??a ? ?oi?o eiiiiiaio?a o oe?ieiio
ae?aiacii? /anoio, ui ia? iie?ooieoe?iiaeuei? iiaeeeaino?, aenie?
iao?ieia?/i? oa?aeoa?enoeee ? ??aaiue aaoiiaoecaoe??. O neeaae? oe???
onoaiiaee aeei?enoaii ieceo aeanieo ?ic?iaie, c-iii?ae ieo: i?ea?iaeuei?
eiino?oeoe?? iaai?oieo aeaoieonoth/eo ianaaeie aeey i?aeaeuaiiy
ii?iaiai? /ooeeaino? OAI; i?inoo oeeo?iao noaio onoiaiiy aieeao
ianoaa?eueiino? eaca?a ia ye?noue io?eioaaieo niaeo??a, a oaeiae
oeeo?iaee i?eno??e, ui aeicaiey? ii?ioaaoe oioi/ooeeaino? iniiaiiai ?
iii?iiai eaiae?a niaeo?iiao?a ia iioe/iee neaiae a?ae ei?eaoth/iai
aeaea?aea; aenieiaoaeoeaiee iaoiae cieaeaiiy ?icn?yiiai iiiio?iiaoi?ii
na?oea iiaeeco e?i?? caoaeaeaiiy, ui ?icoe?th? ieaeith a?aieoeth
niaeo?aeueiiai ae?aiaciio aei 10 ni-1 i?e aeine?aeaeaiiyo
neeueii?icn?thth/eo c?ace?a. ?ic?iaeaii oaeiae iaoiaeeeo caoaeaeaiiy
?aiai?anueeeo niaeo??a eiiiiiaio?a a?iiie?ia??a oa ?oi?o eiiieaen?a a
iie?e?enoae?/iiio noai?. Aei ia? aoiaeeoue i?aeaioiaea c?ace?a aeey
aeine?aeaeaiiy oa aeei?enoaiiy i?ea?iaeueieo ethaao – oea aeicaiey?
nooo?ai iie?ioeoe ni?aa?aeiioaiiy neaiae/ooi ia ao?aei?e u?eei?
iiiio?iiaoi?a ca i?i?iaeueii? e?eueeino? ?a/iaeie ? iaaaeeei?
iiooaeiino? eaca?a.

Aenia?eiaioaeueii iaa?oioiaaii canoinoaaiiy iecueei/anoioii?
?aiai?anueei? niaeo?ineii?? ye aoaeoeaiiai iaoiaeo aea/aiiy
no?oeoo?ii-aeeiai?/ieo aeanoeainoae oaa?aeioacieo ioeea?iiai-ioeea?iiaeo
? a?eeiai-ioeea?iiaeo eiiieaen?a.

Io?eiaii iaea?eueo iiai? iecueei/anoioi? (10 ni-1 < i < 200 ni-1) ?aiai?anuee? niaeo?e (aeey aeayeeo ia’?eo?a oae? aeai? io?eiaii aia?oa) eiiiiiaio?a ioeea?iiaeo eeneio, ui iine?aeiaii oneeaaeiththoueny (eaiii?/ieo ioeeaioeaeieo iniia ? ?o iaoeeueiaaieo a i?noe? ae?eiceaeiiai ca’yceo iio?aeieo, ioeeaiceae?a ? ioeeaioeae?a), a oaeiae aeayeeo oaa?aeioacieo eiiieaen?a (Cyt:F-Gly, Cyt:Benz-Gly), ui iiaeaeththoue "oi/eia?" a?eeiai-ioeea?iia? eiioaeoe, a iie?e?enoae?/iiio noai?. Ia iniia? ye?nii? ?ioa?i?aoaoe?? oeeo niaeo??a iaea?aeaii aeai? i?i aeaye? no?oeoo?ii-aeeiai?/i? oa o?ceei-o?i?/i? iniaeeaino? aeine?aeaeaieo ia’?eo?a. Oae, iaoiaeii iecueei/anoioii? ?aiai?anueei? niaeo?ineii?? anoaiiaeaii, cie?aia, ui i?e e?iiaoi?e oaiia?aoo?? o?eueee o aeiaaeeo Cyt a?ae?aoaoe?y noi?iaiaeaeo?oueny ?noioiueith ia?aaoaeiaith aeaiaei??ii? n?oee eiiia?aoeaieo I-ca’yce?a o o?eaei??io c o/anoth iieaeoe e?enoae?caoe?eii? aiaee ye ??aiii?aaieo ia?oia??a. A ?ioeo aeiaaeeao ( Ura, Thy, Ade) aeiaeaoeia? I-ca’ycee c o/anoth e?enoae?caoe?eii? aiaee ia ia?aaoaeiaothoue ?noioiuei i?e e?iiaoi?e oaiia?aoo?? aeo?aeio n?oeo eiiia?aoeaieo I-ca’yce?a – ?o iiaeia ?icaeyaeaoe ye neaaea cao?aiiy, ui i?aeoe/ii ia a?aeaeaa?oueny ia oiiiii?e aeeiai?oe? e?enoae?a. I?i oea na?ae/eoue iaeaea ?aeaioe/i?noue a?aoeiaeo ?aiai?anueeeo niaeo??a aacaiaeieo e?enoae?a ? ?oi?o e?enoaeia?ae?ao?a. Eeoa ia /anoioao ~ 10 ni-1 e?enoaeia?ae?aoe aeaiiino?othoue i?aeneeaia aeeoocia ?icn?yiiy ia eaac?eieaeueieo iiaeao e?enoae?caoe?eii? aiaee. A?aenooi?noue caaeia?eueii? ei?aeyoe?? i?ae aenia?eiaioaeueieie /anoioaie a?aoeiaeo eieeaaiue, ui niinoa??aathoueny o ?aiai?anueeeo niaeo?ao e?enoae?a nooiai Cyt ? eiai iiiia?ae?aoa, ? ?ic?aooieaie aiaeia?/ieo /anoio, aeeiiaieie o iaaeeaeaii? aei?noei? iieaeoee (Weber, Kirin, 1992), aeaco? ia iai?eeiyoi?noue inoaiiueiai – no?oeoo?ia iaaei?noe?noue iniia c ai?iia?oiith neeueii aieeaa? ia iecueei/anoioio aeeiai?eo e?enoae?/ii? a?aoee /a?ac aca?iiae?th aioo??oiueiiieaeoey?ieo ? ciai?oi?o eieeaaeueieo nooiai?a a?eueiino?. I/aaeaeii, oaea ae naia neooaoe?y ia? i?noea ? a AeIE – ?? iecueei/anoioi? aioo??oiueiiieaeoey?i? eieeaaiiy neeueii aca?iiae?thoue i?ae niaith oa c iicaeiaaei?ie aeonoe/ieie oiiiiaie. A e?enoaeao Thy, m1Thy, m1Ura, m1Cyt ? m9Ade ca?a?no?iaaii ooiaeueia ?icuaieaiiy aioo??oiueiiieaeoey?iiai oi?n?eiiai eieeaaiiy Ia-a?oie, yea i?iyaey?oueny o ?aiai?anueeeo niaeo?ao ye neaaea aeoaeaoia nioaa ? ia? aaee/eio 172,2-154,5=17,7 , 164,7-137,8=26,9 (aeey Ia-a?oie i?e aoii? N1), 176,7-150,2=26,5, 153,8-139,8=14,0 ? 155,8-130,5=25,3 ni-1 a?aeiia?aeii. Ia iniia? aenia?eiaioaeueieo /anoio ooiaethaaiiy ? aaiiao?e/ii? no?oeoo?e iniia aecia/aii aia?aaoe/i? ? e?iaiaoe/i? oa?aeoa?enoeee Ia-a?oie – aa?’?? iaa?oaiiy (1,05;0,68 (aeey Ia-a?oie i?e aoii? N1); 0,69; 1,25 ? 0,74 eeae/iieue), na?aaeith aiie?ooaeo oi?n?eieo e?a?aoe?e i?e o?c?ieia?/i?e oaiia?aoo?? (12,6; 13,8 (aeey Ia-a?oie i?e aoii? N1); 12,6; 14,1 ? 14,4°) oa oi?n?eio neeiao iino?eio (6,28; 5,85 (aeey Ia-a?oie i?e aoii? N1) 6,83; 5,51 ? 5,24 eeae/iieue)(aeey Thy, m1Thy, m1Ura, m1Cyt ? m9Ade a?aeiia?aeii). Anoaiiaeaii, ui Ia-a?oia ? cia/ii i’yeoei (eaa?euei?oei) no?oeoo?iei o?aaiaioii, i?ae aaoa?ioeeeee iniia – ?? oi?n?eia neeiaa iino?eia iaioa, i?ae neeiaa iino?eia ia caeiaiiy aaoa?ioeeeeo i?eaeecii o 2 (aeey i??ei?aeei?a) – 3 (aeey io?ei?a) ?ace. Oe? aeai? aeicaieythoue a?aeoai?eoe aeeiai?/io iiaaae?ieo Ia-a?oie o aeine?aeaeaieo niieoeao i?e o?c?ieia?/i?e oaiia?aoo??: ?? oi?n?eia eieeaaiiy (e?a?aoe?y) a?aeiinii aecioeeee?/iiai ca’yceo NN /e NN, ui a?aeaoaa?oueny a ieie? oei/aniaiai iieiaeaiiy ??aiiaaae c /anoioith aeecueei 150 ni-1 ? aiie?ooaeith ~ 13,5°, ia??iaee/ii ia?a?eaa?oueny (iiaeoeth?oueny) iecueei/anoioiei (~ 22 ni-1) ooiaeueiei i?ie?o/oaaiiyi ia eoo 120° o ianooiia oei/aniaa iieiaeaiiy ??aiiaaae. Oaeei /eiii, ioeeaioeaei? iniiae c Ia-a?oiith ? oeiiaeie iieaeoeaie, ui iathoue eaaioiao aaiiao??th. *anoioa oi?n?eiiai eieeaaiiy Ia-a?oie Thy aeecueea aei /anoioe aeniei?ioaineaiiai aeia?ioiiai eieeaaiiy, yea niinoa??aa?oueny a ?aiai?anueeeo niaeo?ao AeIE ia /anoio? ~ 20 ni-1 (Urabe et al., 1984), ui aeaco? ia ?aciiainio aca?iiae?th oeeo iecueeiaia?aaoe/ieo eieeaaeueieo ?oo?a ? oei naiei ia aaaeeeaee aeeaae Thy a aeeiai?eo AeIE. Ocaaaeueiaiiy io?eiaieo aeaieo uiaei caeiiii??iinoae iiaaae?iee iecueei/anoioieo ?aiai?anueeeo niaeo??a iiaeaeueieo ia’?eo?a AeIE, ui iine?aeiaii oneeaaeiththoueny, ii/eiath/e iniiaaie ? cae?i/oth/e ioeeaioeaeaie, ? oi?n?eieo neeiaeo iino?eieo aeey Ia-a?oi iniia i?e ae?eiceaeiiio aoii? acioo, a oaeiae i?inoeo iiaeaeueieo ?ic?aooie?a ? e?oa?aoo?ieo aeaieo uiaei i?e?iaee iae?ioaineai?oeo a?aoeiaeo eieeaaiue oa?oaaoeo e?enoae?a, ui neeaaeathoueny ?c aaoa?ioeeee?/ieo iieaeoe (Oaaaiia e ae?., 1989), na?ae/aoue i?i oa, ui caaaeoaaia aeua iiaea i?e ~ 20 ni-1 ? ia ui ?ioa, ye aioo??oiueiiieaeoey?ia iioe/ia eieeaaiiy – eieaeoeaia e?a?aoe?y nia?aieo iniia AeIE iaaeiei ae?eiceaeieo ca’yce?a, ui ?iciianthaeaeo?oueny acaeiaae iae?iiieaeoee. Oei naiei oe?eeiaeoi i?aeoaa?aeaeothoueny iiaeaeuei? oyaeaiiy i?i iioiaeaeaiiy iecueei/anoioii? ?ooiiino? AeIE, aia?oa cai?iiiiiaai? Iae??aei ua a 1965 ?ioe?. Iaoiaeii “niaeo?aeueii? eaei?eiao???” (Eiaainai, 1981) aia?oa aecia/aii aia?aaoeeo i?aeiieaeoey?ieo I-ca’yce?a o e?enoaeao ? ni?ae?enoae?caoao ioeeaioeaeieo iniia oa ?oi?o iaoeeueiaaieo ia ae?eiceaeiiio aoii? acioo iio?aeieo – iiaeaeueieo ia’?eoao ioeea?iiai-ioeea?iiaiai ai?ciaaaiiy (anueiai aeine?aeaeaii 12 niieoe). Anoaiiaeaii, ui aiie iaeaaeaoue aei ?ic?yaeo na?aaei?o (~6 eeae/iieue), i?e oeueiio aioaeuei?y i?aeiieaeoey?ieo I-ca’yce?a a aeine?aeaeaieo e?enoaeao ? ni?ae?enoae?caoao ioeeaioeaeieo iniia ei?aeth? c ?oi?ie ?aioaaiino?oeoo?ieie aeiaaeeiaie. I?e ia?aoiae? c aaciaiai noaio a e?enoae?/iee aai a iecueeioaiia?aoo?io ?ia?oio iao?eoeth aeiaaeeie i?aeiieaeoey?ieo I-ca’yce?a nei?i/othoueny, ui i?ecaiaeeoue aei i?aeaeuaiiy ?oiuei? aia?aaoeee. Oae, iai?eeeaae, a oeeee?/i?e I-ca’ycai?e ia?? m1Ura:m1Ura aioaeuei?y i?aeiieaeoey?ieo I-ca’yce?a i?e ia?aoiae? c a?eueiiai noaio a e?enoae?/iee c?inoa? ia ~25%, i?e/iio, a iecueeioaiia?aoo?i?e ?ia?oi?e iao?eoe? ?oiy aioaeuei?y ia iaioa, i?ae o e?enoae?. Aia?oa anoaiiaeaii, ui i?aeiieaeoey?i? I-ca’ycee a e?enoaeao ? ni?ae?enoae?caoao ioeeaioeaeieo iniia iathoue iaaaeeoeaiee oa?aeoa?: a oaa?aeioacieo eiiieaenao ioeeaioeaeieo iniia, ui i?noyoue ai?iia?oie, noia?ia aioaeuei?y aiaeiaaeo ca’yce?a ana?aaeei? ia?e iaioa aiaeia?/ii? aaee/eie aeey aaeooio (a?aeaoaa?oueny aca?iia iineaaeaiiy I-ca’yce?a), a I-ca’ycee c o/anoth ?i?iia?oi i?oei?o?, i?ae ca’ycee c o/anoth ai?iia?oi o oiio ae naiiio eiiieaen?. A e?enoaeao, aea no?oeoo?iei iioeaii ? oeeee?/i? ia?e, ye? ia ooai?ththoue i?aeia?ieo I-ca’yce?a, noia?ia aioaeuei?y I-ca’yce?a a ia?? ia?aaeuo? aiaeia?/io aaee/eio aeey aaeooio – a?aeaoaa?oueny, cie?aia, ?oi? aca?iii?aeneeaiiy. Aia?oa iieacaii, ui a aeine?aeaeaieo e?enoaeao ? ni?ae?enoae?caoao ioeeaioeaeieo iniia aianie i?aeiieaeoey?ieo I-ca’yce?a a aia?a?th e?enoae?/ii? a?aoee ia ? aecia/aeueiei – ia?aaaaeathoue ianiaoeeo?/i? neee iaeoaaiiy. I?e oeueiio aieea i?aeoa?iaeo aca?iiae?e ia aia?aaoeeo aioo??oiueioa?iaeo I-ca’yce?a iaaaeeeee. Oea aea? aeiaeaoeiao ?ioi?iaoe?th i?i ni?aa?aeiioaiiy aiane?a ai?eciioaeueieo (I-ca’ycee) ? aa?oeeaeueieo (noae?ia) aca?iiae?e a noaa?euei?noue AeIE. Anoaiiaeaii, ui o aieiaeo ie?aeao AeIE aioaeuei?y I-ca’yce?a c o/anoth ai?ii- ? ?i?iia?oi iaeiaeiaa ? iaioa aiaeia?/ieo aaee/ei o ni?ae?enoae?cao? eth9Gua:m1Cyt, ui i?noeoue Oionii-E?ee?anuee? ia?e iniia. Aeaoaeueiee aiae?c iecee iai?aeeia?ieo (c oi/ee ci?o onoaeaieo iiaeyae?a ia ?oith aoaeiao) o?ceei-o?i?/ieo aeanoeainoae e?enoae?a ioeeaioeaeieo iniia c ai?iia?oiith, a naia – iaaii?yaeeiaaiino? nooi? ?ao?oee, yea nooo?ai cieaeo?oueny i?e caieiaeoaaii? oa aeaeoa?oaaii? (Aoiaithe, 1980) oa iayaiino? aeaoaeo?a-ianoie iin??a ca?yaeo (Inoaiaiei oa ?i., 1990), i?iiaea?iiiae?aii? eiioi?iaoe?? I-ca’ycaieo iieaeoey?ieo oa??a (Craven et al., 1980), oaiia?aoo?ieo aiiiae?e ae?aeaeo?e/ii? i?iieeiino? ? oaiaaina eooa ao?ao oa iia’ycaieo c ieie no?oeoo?ieo ia?aoiae?a (Aiaao?ia oa ?i., 1980), ii?ooaiue i?aaee a?aeai?o a iiey?eciaaieo eieeaaeueieo niaeo?ao (Weber et al., 1992) ? ?icuaieaiiy iecueei/anoioieo (?<200 ni-1) nioa eieeaaiue i?e e??iaaiieo oaiia?aoo?ao (Beetz et al.,1980), eiioaeoii? eiioi?ia??? (Ci?eee e ae?., 1979) oiui aeicaieea ae?eoe aeniiaeo i?i oa, ui ni?eueiith i?e?ino?oeoo?iith i?e/eiith oeeo aoaeo?a ? no?oeoo?ia iaaei?noe?noue naieo iniia oa iniaeeaino? noae?iaiaeo aca?iiae?e i?ae ieie, ye? aeoiaeyoue ca ?aiee o?aaeeoe?eieo oyaeaiue ? ia caiaeyoueny aei iani?yiiaaieo, ianiaoeeo?/ieo aca?iiae?e i?ae ia?aeaeueii /e eaac?ia?aeaeueii (Sponer at al., 1998) i???ioiaaieie noaoe/ieie aaoa?ioeeee?/ieie iieaeoeaie. Aeiaaaeaii, ui i?e eiiaeainaoe?? ioeeaioeaei? iniiae c ai?iia?oiith ooai?ththoue ssi-Oaeea??anuee? e?enoaee, oa?aeoa?iith iniaeea?noth eio?eo ? iayai?noue, ii?yae c o?aaeeoe?eiei noae?iaii, niaoeeo?/ieo eoeii?anueeeo eiioaeo?a AAI ai?iieo aoii?a acioo c aoiiaie aiaeith iieaeoe ?c non?aei?o (aa?oi?o /e ieaei?o) oa??a, ye? iiaeia ?icoe?ithaaoe ye i?aeoa?ia? (aa?oeeaeuei?) I-ca’ycee. Oae? i?aeoa?ia? aca?iiae?? iineeththoue i??ai?aeaeuei?noue ai?iieo o?aaiaio?a (aa?’?? ieiueiii? ?iaa?n?? c?inoa? i?e ia?aoiae? ?c aa?oiueiai noaio o e?enoae?/iee a?eueoa, i?ae ia ii?yaeie – iaea?eueo iieaciaeie o oeueiio a?aeiioaii? i?eeeaaeaie ? e?enoaee m9Ade (Craven et al., 1980) ? AI? (Kraut et al., 1963)), a caeaaei?noue nooiaith ei?aeueiaaiino? oeeo no?oeoo?ii-aeeiai?/ieo cao?aiue aaeeei? aiie?ooaee, yea eaaeeoue a iniia? eiiia?aoeaiiai aoaeoo ssia-Oaeea?a, a?ae oaiia?aoo?e, ni?e/eiy? no?oeoo?i? ia?aoiaee oeio "eaae-aaceaae". Anoaiiaeaii, ui oe? oyaeaiiy ia iaiaaeothoueny eeoa e?enoaeaie ioeeaioeaeieo iniia ? iiaeooue aooe ?iciianthaeaeai? ia AeIE. I?i oea na?ae/eoue, cie?aia, ?icoiia ioe?iea aaee/eie aa?’??o ieiueiii? ?iaa?n?? ai?iia?oie Ade AeIE (~1,7 eeae/iieue), io?eiaia ca oiiae, ui oaiia?aoo?a, i?e ye?e niinoa??aa?oueny ia?ao?ae i?ioiiii? nenoaie AeIE o noai? ie?aee oeio "eaae-aaceaae" (Ruprecht et al., 1994) ca?aa?oueny c oaiia?aoo?ith eiiia?aoeaiiai ssi-Oaeea??anueeiai ia?aoiaeo, ui a?aeiia?aea? "ieaaeaiith" aa?oeeaeueieo I-ca’yce?a ia caaaa/aieo ia?aie Ade:Thy ?? ae?eyieao. Aa?o/e aei oaaae aeaeaei ia aeiaaeeiao aiaeia?th i?ae AeIE ? neeii (Aieueaeaineee, 1987),a oaeiae oie oaeo, ui a AeIE, ye ? o nee?, aeaiyii? eiio?ao?aoe?? iathoue iaeii/anii ye iaoai?/i?, oae ? aeaeo?e/i? aeanoeaino?, iiaeia i/?eoaaoe iayai?noue o AeIE oaeeo naieo iae?i?eii-aeeiai?/ieo aoaeo?a, ui e o nee? (Niieyeia, Oaeiiae/, 1982; Eeeiaa?, 1987). C-iii?ae ieo – aaia?aoe?y eiaa?aioieo aeaeo?e/ieo ? aeonoe/ieo neaiae?a-a?aeaoe?a, iae?i?eia aeonoe/ia i?ici??noue, ?noioiy caeaaei?noue oeueo?acaoeiaiai iiaeeiaiiy a?ae iiooaeiino? aeaeo?iiaai?oiiai iiey ? ?aciiainia iiaaae?iea o caeaaeiino? a?ae /anoioe inoaiiueiai, iayai?noue aeaeo?iiaai?oiiai aei?ii?ithaaiiy i?e no?oeoo?ieo ia?aoiaeao oa nooo?aa caeaaei?noue iiaeeco ieo ae?aeaeo?e/ii? i?iieeiino? a?ae ciai?oiueiai aeaeo?e/iiai iiey oiui – yaeu, ui iiaeooue aaciina?aaeiuei noinoaaoeny ooieoe?thaaiiy AeIE. I?e?ino?oeoo?i? iiaeae? a?iiieaeoey?iiai ai?ciaaaiiy Ia iniia? aeine?aeaeaiiy iai?aaii??e/iei eaaioiaio?i?/iei iaoiaeii MNDO/I AIIA Oionii-E?ee?anueeeo ia? Ade:Thy ? Gua:Cyt o a?eueiiio noai? cai?iiiiiaaii no?oeoo?ii-aeeiai?/io iiaeaeue niiioaiieo iai?a?ice?eoeo noai?a aeaieaioethaiai? AeIE. A ?aieao oe??? iiaeae? aeuacaaaeai? noaie AeIE ioioiaeiththoueny c ia?ao?aeieie noaiaie aioo??oiueiai iiaa?oaiiy ai?iia?oie Ade, Gua ? Cyt o Oionii-E?ee?anueeeo ia?ao Ade:Thy ? Gua:Cyt a oaeiae c eiio?ao?aoe?eiith iano?ee?noth ia?e Ade:Thy. O ia?oiio aeiaaeeo oa?aeoa?iith no?oeoo?iith iniaeea?noth oeeo noai?a ? iayai?noue ia?e a?oo?eaoeaieo I-ca’yce?a ca o/ano? aea?aaeaioieo ai?iieo aoii?a aiaeith ? aoiia eenith eiiieaiaioa?ii? iniiae, ui caaacia/othoueny cia/iith aeaoi?iaoe??th ai?iia?oie, a naia – c?inoaiiyi eooa iaoeeo ?? ieiueie aei aecioeeee?/iiai ca’yceo NN aei ~ 52° - 55° o ia?ao?aeiiio noai? ?? aioo??oiueiai iiaa?oaiiy, a oaeiae i???ioaoe?y AAI ai?iieo aoii?a acioo, ui iathoue cia/ii aeuo i?ioio?euei?noue o ii??aiyii? c iniiaiei eiioi?iaoe?eiei noaiii ia?e, o a?aeiia?aeio ai?icaieo AeIE. Iacaaaeath/e ia oa, ui oe? noaie noaa?e?cothoueny a?eueoith e?euee?noth (ia iaeei) aioo??oiueiiieaeoey?ieo I-ca’yce?a, i?ae iniiai? eiioi?iaoe?ei? noaie Oionii-E?ee?anueeeo ia?, ?oiy noia?ia aia?aaoeea aeyaey?oueny cia/ii iaioith, i?ae aia?a?y eeane/ieo Oionii-E?ee?anueeeo I-ca’yce?a, oiaoi a ia?ao?aeieo noaiao aioo??oiueiai iiaa?oaiiy ai?iia?oie aca?iiae?y i?ae eiiieaiaioa?ieie iniiaaie ?noioiuei iineaaeth?oueny. O ae?oaiio aeiaaeeo ia?a Ade:Thy iiaiaeeoue naaa ye "aieaeaeea" – ia?aoiaeeoue ?c iniiaiiai, eeane/iiai noaio o aenieiaia?aaoe/iee iaoanoaa?eueiee, ui noaa?e?co?oueny aeaiia aioo??oiueiia?ieie I-ca’yceaie N3I(Thy)…N1(Ade) ? N2I(Ade)…I2(Thy) ?c cia/ii iineaaeaiei I-ca’yceii ca o/anoth ai?iia?oie Ade ? aoiia eenith I4 Thy. Aiae?c o/ano? niiioaiieo iai?a?ice?eoeo noai?a AeIE o i?ioeanao a?eeiai-ioeea?iiaiai ai?ciaaaiiy, ui i?ecaiaeyoue aei ?icie?oaiiy AeIE a?eeii, aeicaiey? c?iaeoe aeniiaie i?i i?ia?aeio ?ieue o oeueiio i?ioean? ae?eyiie AeIE, caaaa/aieo ia ia?e Ade:Thy. Naia ia oaeeo ae?eyieao noai?ththoueny iaeni?eyoeea?o? oiiae aeey ?icie?oaiiy AeIE a?eeii, ui niaoeeo?/ii aca?iiae?? c ?? aaeeeith ai?icaieith. ?ic?iaeaii oaeiae no?oeoo?ii-aeeiai?/io iiaeaeue i?inoi?iai? aee?eaeaiino? e?i?eii? AeIE. A iniiao ?? iieeaaeaii iia? o?ceei-o?i?/i? oyaeaiiy i?i noae?ia Oionii-E?ee?anueeeo ia? ioeeaioeaeieo iniia oa i?i ai?iia?oio inoaii?o ye iae?i?eii-aeeiai?/iee "iieaeoey?iee noaeia", ?ooii?noue yeiai eiio?ieth?oueny ye eeane/ieie ai?eciioaeueieie I-ca’ycaie ca o/anoth ai?iieo aoii?a aiaeith, oae ? aa?oeeaeueieie I-ca’ycaie, aea ?ieue aeoeaioi?a i?ioiia a?ae?a?a? AAI ai?iieo aoii?a acioo. Eieaeueia aee?eaeai?noue e?i?eii? AeIE, yea nooo?ai caeaaeeoue a?ae iine?aeiaiino?, aeieea? aiane?aeie iaeiieaia?iino? Oionii-E?ee?anueeeo ia? ioeeaioeaeieo iniia oa eeeiiiiae?aii? i???ioaoe?? non?aei?o ia?. Oi?eaeueiei c oe??? oi/ee ci?o i?eeeaaeii ? iie?ioeeaioeae poly dA · poly dT – "i?oeiee ai??oie" no?oeoo?ii? a?ieia?? – c aiiiaeueii aenieith i?inoi?iaith aee?eaeai?noth. Aiae?c ?? eaaioiai-o?i?/iiai iioiaeaeaiiy iaoiaeii MNDO/I a ?aieao cai?iiiiiaaii? iiaeae? na?ae/eoue i?i oa, ui iniiaiei noa?aiaeaeo?iiiei /eiieeii, yeee caaacia/o? eeeiiiiae?aio i???ioaoe?th non?aei?o ia? Ade:Thy, ? aa?oeeaeuei? I-ca’ycee …N6I…N6I…N6I… i?ae ai?iieie ca’yceaie aaeai?i?a, ui aa?ooue o/anoue o ai?eciioaeueieo Oionii-E?ee?anueeeo I-ca’yceao. Oe? aa?oeeaeuei? i?aeieaia?i? "ne??iee", ye? iathoue eiiia?aoeaiee oa?aeoa? oa nooo?ai aeeiai?/ia iioiaeaeaiiy ? iineeththoueny ?c c?inoaiiyi ?oiuei? e?eueeino?, aneiao?e/ii i??ai?aeae?cothoue ai?iia?oio Ade, ca?eueooth/e oei naiei ?? aoaeoeaiee ?ici?? acaeiaae aeiaai? in? iie?ioeeaioeaeo ? ye iane?aeie – a?aenoaiue i?ae non?aei?ie ai?iia?oiaie: aeieea? eeeiiiiae?aia i???ioaoe?y non?aei?o ia? Ade:Thy ?c aiiiaeueii aocueeith iaeith ai?icaieith ? aiiiaeueii aenieei cia/aiiyi eooa i?iiaea?-oa?no eiaeii? ia?e, yea aeiaeaoeiai noaa?e?co?oueny i?aeia?ieie I-ca’ycaie N2I(Aden+1)…O2(Thyn), ye? a?oo?eothoue c aioo??oiueiioeeaioeaeieie I-ca’yceaie O2(Thyn)… I1’N1’(Thyn). Oae? oyaeaiiy aeicaieythoue ia eeoa aeoeoia/eoe ia i?e?ino?oeoo?iiio ??ai? c ?aeeieo o?ceei-o?i?/ieo iiceoe?e oe?eo ieceo aoaeo?a, iia’ycaieo ?c caeaaei?noth i?inoi?iai? aee?eaeaiino? e?i?eii? AeIE oa ei?aeyoe?eii? iiaaae?iee ?? ai?iii?ioii?a a?ae ?? iine?aeiaiino?, a e iiaeieaoe aeineoue no?ee? noa?aioeie, ui iiaooothoue o?eaaeee /an o no?oeoo?i?e ? iieaeoey?i?e a?ieia??. Ia?oee ?c ieo noino?oueny i?e?iaee i?inoi?iai? aee?eaeaiino? AeIE o aenieiniaoeeo?/ieo a?eeiai-ioeea?iiaeo eiiieaenao. ?ai?oa aeey iiyniaiiy oeueiai oaiiiaio aeei?enoiaoaaee aeeeth/ii aeaeo?inoaoe/i? oyaeaiiy a ?aieao iiaeae? AeIE ye i?oaeiiai no?eaeiy, ye? caiaeeee eieaeuei? aeaeie AeIE aei i?inoi?iai? iaiaeii??aeiino? eiiiainaoe?? ca?yae?a oinoaoieo a?oi, ui oyaia ca niaith /a?ac a?aae??io eoeii?anueeeo nee eieaeueio aeaoi?iaoe?th AeIE, ye i?oaeiiai noa?aeiy aac cao?aiiy eiai aioo??oiuei? no?oeoo?e. Oaia? noaei c?icoi?ei, ui oeae i?ioean a?aeaoaa?oueny ca ni?eyiiy ai?iia?oi iniia AeIE ye "iieaeoey?ieo noaeia?a", ine?eueee ci?ia ca?yaeo ia oinoao? aeenoaioe?eii (/a?ac nenoaio o?i?/ieo ca’yce?a) ? iaeaieiai?oa – niaoeeo?/ii aieeaa? ia ?oith iae?i?eii-aeeiai?/io ?ooii?noue, yea a naith /a?ao ni?e/eiy? eieaeueiee aeaei AeIE. Ae?oaee noa?aioei noino?oueny iayaiino? o iie?ioeeaioeae? poly dA · poly dT iaeaiii?/ieo I-ca’yce?a a?aeiia?aeaeueieo ca eiai "aiiiaeueio" i?inoi?iao a?o?oaeoii?eo, – c’ynoaaeiny, ui oae caai? i?aeeaioethaia? iioee? I-ca’ycee (Klug et al., 1987) ia iathoue aei oeueiai aoaeoo aaciina?aaeiueiai a?aeiioaiiy, i?e/iio naia ?oi? ?nioaaiiy ? aaeueie i?iaeaiaoe/iei ? ioe?ith?oueny iei? ia ??ai? e?eueeio a?aenioe?a (Levitt et al., 1996). C aeuaaeeeaaeaieie iiaeaeyie o?nii iia’ycaia cai?iiiiiaaia aaoi?ii no?oeoo?ii-aeeiai?/ia iiaeaeue ia?aaeieaaeaiiy AeIE. Ca?aeii c iath i?ioean ia?aaeieaaeaiiy AeIE ?ioa?i?aoo?oueny ye oa?iio?iiiee iaciaaiacen no?oeoo?ii iaaei?noeiai eaac?iaeiiaei??iiai e?enoaeo: oa?iio?iii? iacioace aeieeathoue aiane?aeie iiaoaiii? ao?aoe iei aeeiai?/ii? no?eeino?, coiiaeaii? ai?cio?iiiei ia?inoaiiyi aiaa?iii?cio eiioi?iaoe?eieo nooiai?a a?eueiino?, iia’ycaieo cie?aia c ?ioa?eiiaa?n??th ai?iiiai o?aaiaioa oa aio/e?noth e?eaoeue ioeeaioeaeieo iniia c i?aeaeuaiiyi oaiia?aoo?e. Oaeee iae?i?eii-aeeiai?/ee noeaia??e iinoaae?eiiai ?icaeoeo no?oeoo?ieo ia?aoai?aiue, ui ia?aaeothoue ieaaeaiith AeIE, ocaiaeaeo?oueny c aeaieie ssI? (Lane et al., 1989) ? aeicaiey? eeaneo?eoaaoe ?o ca aeeiai?/ieie iciaeaie, nie?ath/enue ia oai?aio a???aea. Iaea?eueoa a?ieia?/ia cia/ou?noue oe??? iiaeae? iieyaa? a oiio, ui aiia ia?aaeaa/a? iiaeeea?noue aeenoaioe?eii? aeoeaaoe?? AeIE iicaeiaaei?ie iae?i?eieie eiioi?iiiaie, iai?eeeaae o i?ioeanao ?aie?eaoe??, ca a?eueo neeaaeiei ? ooieoe?iiaeueii i?eaaaeea?oei iaoai?ciii, ai?ae i?inoa a?ia?ia noaia "ana – aai i?/iai", ye oea o?aaeeoe?eii aaaaeaeiny ?ai?oa. Cai?iiiiiaaii i?e?ino?oeoo?io iiaeaeue aeaeaeiae?? a AeIE – iaeiiai ?c ooiaeaiaioaeueieo ? aiaeii/an /e ia iaeiaio c?icoi?eeo oaiiiai?a iieaeoey?ii? a?ieia??. Nooue eiai iieyaa? a oiio, ui ye iieaco? aenia?eiaio (Pohl et al., 1972; Kamzolova et al., 1973), aca?iiae?y a?eeiai? iieaeoee P1 ?c naeoii S1 AeIE iiaea aieeaaoe ? iaa?oue ci?ithaaoe oa?aeoa? aca?iiae?? a?eeiai? iieaeoee P2 ?c naeoii S2 AeIE – i?e oeueiio a?aenoaiue i?ae naeoaie S1 ? S2 iiaea nyaaoe oeny/ ia? iniia AeIE. Aieiaiee iaaeie?e ?nioth/eo iae?i?eii-aeeiai?/ieo i?aeoiae?a aei iiyniaiiy o?ceei-o?i?/iiai iaoai?cio aeaeaeiae?? a AeIE (Yakushevich, 1998) iieyaa? a oiio, ui aiie a?oioothoueny ia aenieiaia?aaoe/ieo caoaeaeaiiyo, oa?aeoa?ia aia?a?y eio?eo a?eueoa, i?ae ia ii?yaeie ia?aaeuo? eO i?e o?c?ieia?/i?e oaiia?aoo??. Ia? i?noea iaa?aeiia?aei?noue ?ioi?iaoe?eiiai oa?aeoa?o iae?i?eieo, ia??aiiaaaeieo eiioi?iii?a, ui ?iciianthaeaeothoueny acaeiaae AeIE ? caaacia/othoue aeaeaeiae?th, ?oi?e aia?aaoeoe?. Ie??i oiai, a?aeii? iiaeae? ia ? aacaeiaaiieie ? uiaei iaoai?ci?a ?i?oe?aoe?? oa ?aoeaioe?? eiioi?iii?a – iin??a aeaeaeiae??, ye? iaeiicia/ii ia iia’ycothoueny c ?oiueith o?ceei-o?i?/iith i?e?iaeith. Ianaie?iaoeue, aiie i?aeoe/ii ia aaeaioiaai? aei oiia aioo??oiueiee?oeiii? eiiia?oiaioae?caoe??, ui ? iaeiei ?c ao?eaoo?a aeeaiai. Cai?iiiiiaaiee iae?i?eii-aeeiai?/iee iaoai?ci aeaeaeiae?? ? iecueeiaia?aaoe/iei ? ia? iine?aeiaia eaaioiaio?i?/ia iaa?oiooaaiiy, ine?eueee aaco?oueny ia aeia?a aea/aieo iaie yaeuao noa?aio?i?/ii? iaaei?noeino? eaiii?/ieo ioeeaioeaeieo iniia ? caeaaeiino? ?? a?ae ca?yaeiaiai noaio inoaii?o, aeeiai?/ii? noa?ai?ciia??? eiiieaiaioa?ieo ia? iniia AeIE oa ia iiaeo oyaeaiiyo i?i noae?ia inoaii?o. A?i aea? a?aeiia?aeue ye ia o?ceei-o?i?/io i?e?iaeo neaiaeo aeaeaeiae??, oae ? ia iaoai?cie eiai ?i?oe?aoe?? ? ?aoeaioe??. Ia?oa aaa/a?oueny o aaeeeiaiie?ooaeieo aeeiieueii-aeoeaieo a?yeiaeo eieeaaiiyo ai?iia?oi nia?aieo iniia AeIE, ye? nooo?ai aca?iiae?thoue c iecueei/anoioieie eieaeoeaieie e?a?aoe?yie iniia iaaeiei ae?eiceaeieo ca’yce?a ? aeey yeeo aeeiiothoueny eeane/i? oiiae (Aeaaeaeia, 1988; ss?aiei, 1996) aaoieieae?caoe?? o noaoe?iia?i? nie?oiie. I?e oeueiio iaoai?ci caioneo ?ioi?iaoe?eii? iae?i?eii? oaee? acaeiaae AeIE caiaeeoueny aei eieaeueiiai cao?aiiy ai?iia?oi AeIE ca ?aooiie aoyaoaaiiy a?aeiia?aeieo oinoaoieo a?oi AeIE o I-ca’ycee c eeneeie ai?iieeneioieie caeeoeaie a?eea, iiaeeeai iaa?oue c a??ooaeueiei ia?aianaiiyi i?ioiia ia oinoaoe, – ui c iaiao?aei?noth ni?e/eiy? eieaeueio ci?io ca?yaeiaiai noaio nenoaie c oneia iane?aeeaie, ui c oeueiai aeieeaathoue. I?eioeeiiaei iiiaioii cai?iiiiiaaii? iiaeae? ? oa, ui ?iciianthaeaeaiiy iae?i?eiiai eiioi?iiia acaeiaae AeIE noi?iaiaeaeo?oueny eieaeueiei aee?eaeaiiyi inoaiiuei?. Naia caaaeyee oaeiio aeeiai?/iiio aee?eaeaiith ? ?aae?co?oueny ea?iaaiee caeaeaeo niaoeeo?/iee eieaeueiee eiioaeo AeIE c a?eeii P2, yeee noa? iaiiaeeeaei i?e a?aenooiino? neaiaeo /a?ac /enoi noa?e/i? ia?aiiie. I?e ?aae?caoe?? oeueiai aae?aniiai eiioaeoo a ae?th anooia? eeane/iee iaoai?ci no?oeoo?ii-aeeiai?/ii? a?aeiia?aeiino? i?noeue ai?ciaaaiiy. Ia aeeeth/aii, ui oey no?oeoo?ii-aeeiai?/ia a?aeiia?aei?noue, yea caaacia/o? aaciiieeeia?noue i?ioeano ai?ciaaaiiy, ?aae?co?oueny oiae?, eiee ? ii a?eeia?e iieaeoe? neio?iiii ?iciianthaeaeo?oueny, oae ae iiaeoe “eiiieaiaioa?iee” nie?oii ? ooieoe?iiaeueii cia/ouee eiioaeo ?aae?co?oueny eeoa i?e ?oi?e cono??/? ? aca?iiae??. Aeoaea aaaeeeai, ui nie?oiie a AeIE, ye ? nie?oiie a a?eeao (ss?aiei, 1996) ? aeeiieueii-aeoeaieie caoaeaeaiiyie. Oea aeicaiey? aoaeoeaii aca?iiocaiaeeoe i?ioean ?oiueiai aenieinaeaeoeaiiai, aaeaioeaiiai caeeaeaiiy iacono??/ iaeei iaeiiio ca ?aooiie, cie?aia, ?aciiainieo aeaeo?iiaai?oieo aca?iiae?e. Cai?iiiiiaaia iiaeaeue aeaeaeiae?? a AeIE aaeaioiaaia aei oiia aioo??oiueiee?oeiii? eiiia?oiaioae?caoe??, ine?eueee ia aeiaaa? aeey nai?? ?aae?caoe?? iaaeeeoeiaiai a?eueiiai ia’?io. Ooo, ye ? i?e ia?aa?aia? o?i?/ieo ?aaeoe?e o eiiaeainiaai?e oac? ca oiia aeao?oeeoo a?eueiiai ia’?io, i?ia?aeio ?ieue i/aaeaeii a?ae?a?athoue eieaeuei? oeoeooaoe?? aio?ii??, ye? aeicaieythoue AeIE ? a?eeai anooiaoe o aca?iiae?th i?ae niaith o ooieoe?iiaeueii-cia/ou?e eiioi?iaoe?? (ia?ao?aeiiio noai?). I?ioa ia a?aei?io a?ae eiiaeainiaaiiai na?aaeiaeua, aea ia?aa?a iiae?aieo i?ioean?a eiio?ieth?oueny ??aiiaaaeiith aonoeiith oaeeo oeoeooaoe?e (Ao?ooaei, 1978), o ee?oei? aiie, i/aaeaeii, ?aaoeththoueny /anoioith ?oiueiai ieaiiaiai oi?ioaaiiy, ine?eueee ia a?aei?io a?ae o?ce/iiai na?aaeiaeua, o aeea?e ee?oei? ia?aaaaeia a?eueo?noue, yeui ia an? i?ioeane ? aca?iiiaeaoa?i?iiaaieie, ?iaeoeiaaieie, a ia niiioaiieie, ? aei oiai ae a?aeaoaathoueny a oiiaao nooo?ai? oa?iiaeeiai?/ii? ia??aiiaaaeiino? oa i?inoi?iai? aaoa?iaaiiino?. Aeoaea aaaeeeai c a?ieia?/ii? oi/ee ci?o, ui cai?iiiiiaaia iiaeaeue aeiionea? oeeee?/iee ?aaeei naiai ooieoe?iioaaiiy. Iaeiei c iaea?eueo iaaaoeaieo o?ceei-o?i?/ieo /eiiee?a, ui ia?aoeiaeaeathoue aeieeiaiith ? aoaeoeaiiio ?iciianthaeaeaiith acaeiaae AeIE iae?i?eieo ia??aiiaaaeieo eiioi?iii?a, a?aeiia?aeaeueieo ca aoaeo aeaeaeiae??, ? iayai?noue aiaeiiai ioi/aiiy, yea ? ?oi?i iiooaeiei aeaiioa?ii (Lindsay, 1986). I/aaeaeii, naia c oe??? i?e/eie ia aaea?oueny aenia?eiaioaeueii ?o niinoa??aaoe a AeIE iica ee?oeiith. A ?aaeueieo oiiaao ooieoe?iioaaiiy AeIE o ee?oei? oeae oaeoi?, i/aaeaeii, a?aenooi?e (Verkman, 1993). ?icaeiooi iia? oyaeaiiy i?i iieaeoey?i? iaoai?cie aaiioiene/ii? ae?? OO-ii?ii?iaiiy ia IE. Iaeei ?c ieo a?oioo?oueny ia aia?oa cao?eniaaiiio aoaeo? ?aaeeeaeueii? ci?ie oiiieia?? AIIA ioeeaioeaeieo iniia c ai?iia?oiith i?e ia?aoiae? ?c iniiaiiai aeaeo?iiiiai noaio S0 o ia?oee caoaeaeaiee S1 : a?aeaoaa?oueny o?ainoi?iaoe?y aeaiyiiiai aeca?eaeueii-neiao?e/iiai aae?aaaoe/iiai iioaioe?aeo ieiueiii? ?iaa?n?? ai?iiiai o?aaiaioo ?NNI2 o iaeiiyiiee (eiai i?i?ioi a?aeiia?aea? ieaia?i?e no?oeoo??), oa?aeoa? iiaa?oaeueii? ai?cio?ii?? ai?iia?oie i?e oeueiio ia ci?ith?oueny (aeaui c?inoathoue eeoa aa?’??e iiaa?oaiiy). Oaea iiaaae?iea iniia iia’ycaia ?c cia/iei iineeaiiyi ??-ni?yaeaiiy AAI ai?iiiai aoiia acioo c ?-aeaeo?iiiith nenoaiith e?eueoey ? cia/ii caaaeueiiao? eiioi?iaoe?eio aio/e?noue Oionii-E?ee?anueeeo ia? AeIE a?aeiinii eoo?a "i?iiaea?-oa?no" ? "ceai". Oaeei /eiii, OO-ii?ii?iaiiy caeaoia cao?thaaoe eieaeueio aee?eaeai?noue AeIE, ine?eueee inoaiiy aecia/a?oueny noa?aio?i?/iith iaaei?noe?noth iniia c ai?iia?oiith oa aa?oeeaeueieie I-ca’yceaie i?ae ieie, ii?oooth/e oei naiei ia?aa?a neio?ii?ciaaieo o i?inoi?? ? /an? aenieiniaoeeo?/ieo i?ioean?a a?eeiai-ioeea?iiaiai ai?ciaaaiiy. Ie??i oiai, i?e ?aeaenaoe?? ii?ii?iaii? AeIE a i?e iiaeooue ?i?oe?thaaoeny iicaeiaaei? iae?i?ei? eiioi?iiie – iin?? aeaeaeiae??. ?iciianthaeaeoth/enue acaeiaae AeIE, aiie caii/aoeiaothoue ianaieoe?iiiaai? ?? eieaeuei? aeaeie, ui oaae i?ecaiaeeoue aei ii?ooaiiy i?e?iaeiueiai ia?aa?ao ooieoe?iiaeueii aaaeeeaeo a?eeiai-ioeea?iiaeo aca?iiae?e. Noeaia??e ?icaeoeo iiae?e iiae?aiiai oeio ia iieaeoey?iiio ??ai? iniaeeai iaaacia/iee c oi/ee ci?o aoaeoeaiiai aeieoaaiiy ?oiueiai iiaeaeueoiai ?icaeoeo, a cia/eoue – ? iane?aee?a, oa?iaioieie nenoaiaie, ine?eueee OO-na?oei ia ni?e/eiy? o?i?/ii? iiaeeo?eaoe?? AeIE, cao?thth/e eeoa ia ei?ioeee /an ?? eiioi?iaoe?th. I?e?iaeiuei, ui oaeiai oeio a??ooaeuei? "iioeiaeaeaiiy" ia iiaeooue aooe ca?a?no?iaai? a?io?i?/ieie iaoiaeaie, ine?eueee i?ney ii?ii?iaiiy AeIE aeineoue oaeaeei iiaa?oa?oueny o iaoeaiee noai. Ia?aoo? aaiioiene/ia ae?y OO-ii?ii?iaiiy iiaea aooe iia’ycaia c a?aeiiei aoaeoii oioi?iaeoeiaaii? ?ciia?ecaoe?? (Aeuey/eia, Eaaei, 1987). Oeae iaoai?ci iiaea ?icaeaaoeny ia ae?eyieao AeIE, caaaa/aieo ia ia?e Gua:Cyt, ine?eueee i? Gua, i? Cyt ia ? aia?aaoe/ii iaeaea?aei?oeie no?oeo?ieie ?ciia?aie, ii?oooth/e oei naiei eiaeia? aeanoeaino? AeIE. Aea/aii o?ceei-o?i?/i? canaaee eiiia?aoeaiino? – iaei??? c iaeoa?aeoa?i?oeo iciae a?ieia?/ieo nenoai ia iieaeoey?iiio ??ai? (Karreman, 1980). Nie?ath/enue ia ?acoeueoaoe aeanieo aeine?aeaeaiue noa?aio?i?/ii? iaaei?noeino? ioeeaioeaeieo iniia ? aeayeeo ioeeaiceae?a oa ?oiuei? eaaioiaio?i?/ii? i?e?iaee, a oaeiae ia eiioeaioe?th a?a?iiieo aca?iiae?e (Aa?noea?, 1987), aeiaaaeaii, ui aeaiaioa?ieie no?oeoo?ieie eaieaie – iin?yie eiiia?aoeaieo aeanoeainoae IE ? ioeeaiceaee – ssi-Oaeea??anuee? iieaeoey?i? no?oeoo?e, a iecueeiaia?aaoe/ia eiiia?aoeaia iiaaae?iea IE coiiaeaia eieaeoeaieie a?a?iiieie aca?iiae?yie. Ia i?aenoaa? oeeo ?acoeueoao?a c?iaeaii i?eioeeiiai aaaeeeaa i?eiouaiiy i?i oa, ui naia iecueeiaia?aaoe/i? (oiaoi oae?, aia?a?y eio?eo ni?ai??ia c na?aaeiueith oaieiaith aia?a??th i?e o?c?ieia?/i?e oaiia?aoo??) aeaeo?iiii-eiioi?iaoe?ei? caoaeaeaiiy IE, aeioe?eueii ?i?oe?eiaai? o oie /e ?ioee niin?a (?aciiainiee /e ia?aciainiee) oeyoii a?a?iiiiai oi?aae?iiy – cao?aiiyi iaaae?aaaoe/iiai ci?ooaaiiy aeaeo?iiieo noai?a, a?ae?a?athoue /e ia iaeaaaeeea?oo ?ieue o ia?aa?ao oe?ieiai eiea a?ieia?/ii aaaeeeaeo i?ioean?a, i?aai?/ii iiina?aaeeiaaieo aenieiniaoeeo?/ieie aeoaie a?iiieaeoey?iiai ?ici?ciaaaiiy. Eaaioiaio?i?/ia aeine?aeaeaiiy iiaii? iiiaeeie oi/eiaeo eiioaeo?a oeio "ioeeaioeaeia iniiaa – a?/iee ?aaeeeae ai?iieeneioe", ui noaa?e?cothoueny aeaiia i?aeiieaeoey?ieie I-ca’yceaie Aia?oa a ?aieao i?iaeaie a?eeiai-ioeea?iiaiai ai?ciaaaiiy o iaeneiaeueii iiaeeeaiio oi?iao?, yeee iaiaaeo?oueny eeoa no/anieie oyaeaiiyie i?i i?e?iaeo I-ca’yceo, ?c canoinoaaiiyi iai?aaii??e/iiai eaaioiaio?i?/iiai iaoiaeo MNDO/I ?icaeyiooi iiaio iiiaeeio oi/eiaeo eiioaeo?a oeio "ioeeaioeaeia iniiaa – a?/iee ?aaeeeae ai?iieeneioe", ui noaa?e?cothoueny aeaiia i?aeiieaeoey?ieie I-ca’yceaie, ? aeine?aeaeaii iniiai? ?? o?ceei-o?i?/i? oa?aeoa?enoeee – aaiiao?e/io, aeaeo?iiio ? aia?aaoe/io no?oeoo?o, ui aaciina?aaeiuei ni?aa?aeiinyoueny c ia?aa?aii aeaiaioa?ieo aeo?a a?eeiai-ioeea?iiaiai ai?ciaaaiiy ia i?e?ino?oeoo?iiio ??ai?. An? aac aeiyoeo a?/i? ?aaeeeaee ai?iieeneio, caeaoi? iaeii/anii aoyaoaaoeny o aeaa i?aeiieaeoey?i? I-ca’ycee ye aeiii? ? aeoeaioi?, iiaeaethaaeeny oaeeie niieoeaie: na?eio, o?aii?io ? oe?iceio – iaoeeiaei nie?oii, e?ceio – iaoeeai?iii, aeooai?io ? ania?aa?io – oi?iai?aeii, a?aei?io – ai?aeeiii, aeooai?iiai? ? ania?aa?iiai? eeneioe – io?aoeiith eeneioith (o aeiaaeeo e?ceio ? a?aei?io – ye aeaeo?iiaeo?aeueiith oae ? i?ioiiiaaiith oi?iaie; o aeiaaeeo aeooai?iiai? ? ania?aa?iiai? eeneioe – ye aeaeo?iiaeo?aeueiith oae ? aeai?ioiiiaaiith oi?iaie). Ie??i oiai, aeey io?aoeii? eeneioe i?eeiaee aei oaaae iaeaea? eiioi?iaoe?? a?ae?ieneeo – iniiaio (oeen) ? aenieiaia?aaoe/io (o?ain); aeey ioeeaioeaeieo iniia ?icaeyaeaeeny ye iniiai? ?oi? oaooiia?i? noaie, oae ? aeaye? aenieiaia?aaoe/i? (??aee?ni?). Aia?oa anoaiiaeaii oae? caeiiii??iino?. O ia?aaaaei?e a?eueoino? aeiaaee?a ?aae?co?oueny aoaeoeaii-ieaia?i? eiiieaene – iaea?eueoa a?aeoeeaiiy a?ae ieaia?iino? niinoa??aa?oueny o oiio aeiaaeeo, eiee ea?aieneeao-ai?ii aca?iiae?? c NI– /e NI– a?oiith, ui ciaoiaeeoueny a ioi/aii? aeaio aeaeo?iiaaaoeaieo aoii?a (N2I Ade, N3I Thy oiui). Aiie noaa?e?cothoueny i?inoi?iaith n?oeith aca?iicaeaaeieo aioo??oiuei- oa i?aeiieaeoey?ieo I-ca’yce?a c aoaeoii eiiia?aoeaiino?, ui ia ia?aaeuo? 10%. Ca?yaeaeai? oi?ie ai?iieeneioieo caeeoe?a ? a?eueo naeaeoeaieie i?noeyie ?ici?ciaaaiiy, i?ae aeaeo?iiaeo?aeuei?, ine?eueee aiie caaacia/othoue cia/ii oe?oee aia?aaoe/iee ae?aiacii aca?iiae?e a caeaaeiino? a?ae iniiae oa i?noey ca’ycoaaiiy, a oaeiae oa?aeoa? ? aaee/eio a?aeiia?aeieo no?oeoo?ieo cao?aiue iniia. Cao?eniaaii iaiiiaeeiie? aeiaaeee ia?aianaiiy i?ioiio acaeiaae i?aeiieaeoey?iiai I-ca’yceo, cie?aia a?ae iniiae ia ea?aieneeao-ai?ii ? a?ae i?ioiiiaaieo ai?iieeneioieo a?/ieo ?aaeeeae?a ia iniiao, – oea iiaeia ?icoe?ithaaoe ye ci?io oaooiia?iiai noaio eiiieaeno. Iieacaii, ui aca?iiae?y a?/ieo ?aaeeeae?a ai?iieeneio c ioeeaioeaeieie iniiaaie iiaea ?iaeoeoaaoe ia?ao?ae inoaii?o o ??aee?nio oaooiia?io oi?io – iaea?eueoa a?ieia?/ia cia/ou?noue oeueiai aoaeoo i/?eo?oueny a i?ioeanao neioaco IE. Anoaiiaeaii aeiyoeiao aeanoea?noue ea?aieneeao-ai?iio: oea ?aeeiee a?/iee ?aaeeeae ai?iieeneioe, ui caeaoiee ni?e/eiyoe ?oeiaoe?th iaio Oionii-E?ee?anueeeo ia? iniia AeIE oeyoii eiieo?aioe?? ca I-ca’ycoaaiiy. Ia aeeeth/aii, ui oea iaeei ?c iiaeeeaeo i?e?ino?oeoo?ieo iaoai?ci?a ?icie?oaiiy AeIE a?eeaie. Aia?oa cai?iiiiiaaii c?iaeoe aeiiiaiaiiy iiiaeeie oi/eiaeo I-eiioaeo?a, i?i ye? eoea iiaa aeua, ?oi?ie "a??ooaeueieie" aiaeiaaie, oiaoi eiiieaenaie oaeiai ae oeio, aea a ia?ao?aeieo noaiao ?ioa?eiiaa?n?? iaei??? aai iaio iieaeoe-ia?oia??a. Oeae e?ie ? ei?eniei aeey ?icoi?iiy oiai, ui i?ioeane a?iiieaeoey?iiai ai?ciaaaiiy iiaeooue a?aeaoaaoeny ia eeoa a iniiaiiio, a e a ia?ao?aeieo eiioi?iaoe?eieo noaiao. C?iaeaii iniaeeaee iaaiein ia ?ie? no?oeoo?ii? iaaei?noeino? ioeeaioeaeieo iniia o caaacia/aii? iia?aoeaiino? ? aaciiieeeiaino? ia?aa?ao aeaiaioa?ieo aeo?a a?eeiai-ioeea?iiaiai ai?ciaaaiiy: naia oey aeanoea?noue aa?aioo? naeaeoeaia caeeaeaiiy iieaeoe-ia?oia??a o oe?ieiio ae?aiacii? noa?oiaeo eoo?a aoaee, oiaoi ?iaeoue ?o aea?aeieie c aio?ii?eii? oi/ee ci?o. C?noaaeaiiy aia?aaoeee aeine?aeaeaieo eiiieaen?a c ?oiueith ?iciianthaeaeai?noth o ?aaeueieo aenieiiieaeoey?ieo a?eeiai-ioeea?iiaeo eiiieaenao i?eaiaeeoue aei aaaeeeaiai aeniiaeo i?i iai?aaii??i?noue ioioiaeiaiiy a?ieia?/ii? cia/ouino? oiai /e ?ioiai oi/eiaiai eiioaeoo c eiai aia?aaoeeith, yea oi?ioethaaeiny ?ai?oa ye i?aaeei – "aeaa I-ca’ycee – oea e?aua, i?ae iaeei" (Rich, 1992). AENIIAEE Anoaiiaeaii iaea?eueo caaaeuei? o?ceei-o?i?/i? caeiiii??iino? a?iiieaeoey?iiai ai?ciaaaiiy. Aia?oa iieacaii, ui aaea ia iaeieae/iio no?oeoo?iiio ??ai? ioeea?iiai-ioeea?iiaa ? a?eeiai-ioeea?iiaa ai?ciaaaiiy a?aeaoaa?oueny ca noeaia???i aca?iii? no?oeoo?ii-aeeiai?/ii? aaeaioeaiino? iieaeoe-ia?oia??a ca o/anoth aca?iicaeaaeieo aioo??oiuei- oa i?aeiieaeoey?ieo I-ca’yce?a, caeaoieo ia eeoa ooai?thaaoeny ? ?ic?eaaoeny, a e a?oo?eaoeaii ia?aieeaoeny, ? caaacia/o?oueny, a iniiaiiio, noa?aio?i?/iith iaaei?noe?noth ioeeaioeaeieo iniia, yeo ne?ae ?icoe?ithaaoe ye caii?oeo aeniei? oi/iino? oa iia?aoeaiino? ai?ciaaaiiy a oiiaao aeao?oeeoo a?eueiiai ia’?io. Aia?oa ?ic?iaeaii ieceo aca?iiiia’ycaieo ni?eueiei i?e?ino?oeoo?iei iioiaeaeaiiyi ? o/anoth o aenieinaeaeoeaieo i?ioeanao a?eeiai-ioeea?iiaiai ai?ciaaaiiy no?oeoo?ii-aeeiai?/ieo iiaeaeae niiioaiieo iai?a?ice?eoeo noai?a AeIE oa ?? ia?aaeieaaeaiiy, aeaeaeiae?? a AeIE oa ?? i?inoi?iai? aee?eaeaiino?, aaiioiene/iiai aieeao ia ia? OO-ii?ii?iaiiy ? eiiia?aoeaiino?, ye? iathoue iine?aeiaia eaaioiaio?i?/ia oeoia/aiiy. An? aiie a?oioothoueny ia aia?oa io?eiaieo aeaieo uiaei noa?aio?i?/ii? iaaei?noeino? ioeeaioeaeieo iniia oa ?oiueiai noae?iao. Aia?oa iieacaii, ui ioeeaioeaei? iniiae oi?eaeueii ii?aeiothoue o nia? oi?eaeueio ieceo aca?iiiia’ycaieo o?ceei-o?i?/ieo oi?i no?oeoo?ii? i?ieeaino?, oaeeo ye noa?aio?i?/ia iaaei?noe?noue, i?ioio?iiia oaooiia??y, aaiiao?e/ia ?ciia??y, caeaoi?noue i?e?aeioaaoe aai a?aeuaiethaaoe i?ioii, niiioaiii aeacai?ioaaoeny ? ieenethaaoeny i?e eiioaeo? c iieaeoeaie aiaee oiui. Naia a oeueiio aaa/a?oueny oa?aeoa?ia a?aei?ii?noue IE a?ae i?aai?/ieo iie?ia??a – i?aaenoaaiee?a iaaeeai? i?e?iaee. Oeae aeniiaie ?acii ?c ?ic?iaeaieie iiaeaeyie a?iiieaeoey?iiai ai?ciaaaiiy aaoi? ?icoe?ith? ye ia?oee e?ie o iai?yieo noai?aiiy oe?e?nii? o?ceei-o?i?/ii? eiioeaioe?? ooieoe?iioaaiiy IE. ?ic?iaeaii ye?nii iiaa oyaeaiiy i?i no?oeoo?io i?ieea?noue ioeeaioeaeieo iniia. Oae cie?aia, aia?oa anoaiiaeaii, ui: - iniiae c ai?iia?oiith ? noa?aio?i?/ii-iaaei?noeeie iieaeoeaie, aeeiieueii-aeoeaia ?ioa?eiiaa?n?y eio?eo a?aeaoaa?oueny o?ueiia oiiieia?/ii ? aia?aaoe/ii iaaea?aaeaioieie oeyoaie, noi?iaiaeaeo?oueny nooo?aith aeaoi?iaoe??th ai?iiiai o?aaiaioa ? ?noioiuei caeaaeeoue a?ae ?icoaooaaiiy ai?iia?oie a e?eueoe? oa ?? ioi/aiiy, a oaeiae oaooiia?iiai oa ?ciia?iiai noaio iniiae. Aia?oa iieacaii, ui o aeiaaeeo eaiii?/ieo iniia IE ?oiy iecueeiaia?aaoe/ia ?ioa?eiiaa?n?y ? ieiueiiith ?iaa?n??th ai?iiiai o?aaiaioa, ui yaey? niaith iaaeaa?’??ia a?yeiaa eieeaaiiy aaeeei? aiie?ooaee, a aenieiaia?aaoe/ia ?aae?co?oueny ye nooo?ai ai?cio?iiia i?aeaa?’??ia iiaa?oaiiy ai?iia?oie iaaeiei aecioeeee?/iiai ca’yceo CN. Aia?oa c’yniaaii eaaioiaio?i?/i? i?aeaaeeie oeeo aaaeeeaeo c a?ieia?/ii? oi/ee ci?o aeanoeainoae; - i?ioio?iiia oaooiia??y acioenoeo iniia ia? iieaeoey?ii-oea?oa??iiiee oa?aeoa?, i?e/iio a i?e aa?ooue o/anoue on? aac aeiyoeo i?ioiie c ea?aii?ioiiaie aeeth/ii. Aeey aoaeue-yei? io?eiiai? iniiae iniiaiei oaooiia?ii-oea?oa??iiii c-iii?ae oeo, ui oi?iothoueny oeyoii i?a?aoe?? ea?aii?ioii?a, ? ?e?aeia oi?ia, yea ooai?th?oueny a ?acoeueoao? ia?aoiaeo ea?aii?ioiia i?e aoii? N8 ia non?aei?e aoii acioo, c’?aeiaiee c aoiiii N8 iiaea?eiei ca’yceii; - eeoa aea? eaiii?/i? iniiae AeIE – Ade ? Thy ? aia?aaoe/ii iaeaea?aei?oeie no?oeoo?ieie ?ciia?aie a nai?o n?iaenoaao. Aia?oa i?iaeaiiino?iaaii, ui a?ieia?/ia cia/ou?noue i?ioio?iiii? oaooiia??? ioeeaioeaeieo iniia ia ae/a?io?oueny ?? o?aaeeoe?eieie ?aieaie: iaeei ?c iaeaaaeeea?oeo, aea ia ?aeeiee a?ieia?/iee i?ioean, yeee eiio?ieth?oueny oaooiia???th, ? neioac io?eiiaeo ioeeaioeae?a. E?eoa???i a?ieia?/ii? cia/ouino? ??aee?nieo oaooiia??a cai?iiiiiaaii aaaaeaoe ni?aa?aeiioaiiy ?oiuei? aia?aaoe/ii? noaa?eueiino? ia c na?aaeiueith oaieiaith aia?a??th, ye oea o?aaeeoe?eii aaaaeaeiny ?ai?oa , a c aa?’??aie a?aeiia?aeieo a?io?i?/ieo ?aaeoe?e, ui iiaeooue ia?aaeuoaaoe inoaiith a aeanyoee ?ac?a i?e o?c?ieia?/i?e oaiia?aoo??. Aia?oa i?iaiae?ciaaii caeiiii??iino? no?oeoo?ii? ?ciia??? ioeeaioeaeieo iniia i?ae eooii ci?o iieaeoey?ii? aaiethoe?? IE. ?noioiuei ?icoe?aii o?ceei-o?i?/i? oyaeaiiy i?i noae?ia ioeeaioeaeieo iniia c ai?iia?oiith oa ?oi?o aiaeiaai-ca’ycaieo ia?: c’yniaaii , ui aiie ia iiaeooue aooe caaaeai? eeoa aei ianiaoeeo?/ieo aca?iiae?e noaoe/ieo, ieineineiao?e/ieo no?oeoo?, ye oea o?aaeeoe?eii aaaaeaeiny ?ai?oa, - nooo?aee aianie o noae?ia aeathoue niaoeeo?/i? eoeii?anuee? eiioaeoe AAI ai?iieo aoii?a acioo c aoiiaie aiaeith non?aei?o (aa?oi?o /e ieaei?o) iniia, ye? iiaeia ?icoe?ithaaoe ye aa?oeeaeuei? I-ca’ycee. Aia?oa o eiioaeno? iieaeoey?iiai oi?aae?iiy cai?iiiiiaaii ?icaeyaeaoe ai?iia?oio iniia IE ye “iieaeoey?iee noaeia”, iae?i?eia ?ooii?noue eio?iai iiaea ?aaoethaaoeny, cie?aia i?aeiieaeoey?ieie I-ca’yceaie, ye eieaeueii, oae ? aeenoaioe?eii. Aia?oa iieacaii, ui Oionii-E?ee?anuee? ia?e AeIE ? iaaecae/aeii iia?eueieie no?oeoo?aie a?aeiinii eoo?a “i?iiaea?-oa?no ? “ceai” ? iiaeooue eaaei i?eeiaoe iaieaia?io eiioi?iaoe?th i?ae ae??th ciai?oiueiai iecueeiaia?aaoe/iiai cao?aiiy, iai?eeeaae aa?oeeaeueieo I-ca’yce?a. Aia?oa i?aaenoaaeaii aaiiao?e/io, aeaeo?iiio ? aia?aaoe/io no?oeoo?o iiaii? iiiaeeie oi/eiaeo eiioaeo?a oeio “ioeeaioeaeia iniiaa – a?/iee ?aaeeeae ai?iieeneioe”, ui noaa?e?cothoueny aeaiia i?aeiieaeoey?ieie I-ca’yceaie. Cai?iiiiiaaii aeiiiaieoe ?? eiiieaenaie ca o/anoth ioeeaioeaeieo iniia ? aeayeeo ai?iieeneioieo caeeoe?a o ia?ao?aeieo noaiao ?oiuei? ?ioa?eiiaa?n??. Anoaiiaeaii, ui a?ieia?/ia cia/ou?noue oi/eiaeo eiioaeo?a, a naia – ?oiy ?iciianthaeaeai?noue o aenieiiieaeoey?ieo a?eeiai-ioeea?iiaeo eiiieaenao, ia iiaea aooe ioioiaeiaia c ?oiueith aia?aaoe/iith noaa?euei?noth, ye oea aaaaeaeiny ?ai?oa. Aia?oa i?iaeaiiino?iaaii aenieo aoaeoeai?noue iecueei/anoioii? ?aiai?anueei? niaeo?ineii?? aeey aeine?aeaeaiiy no?oeoo?ii-aeeiai?/ieo aeanoeainoae iiaeaeueieo oaa?aeioacieo ioeea?iiai-ioeea?iiaeo ? a?eeiai-ioeea?iiaeo eiiieaen?a. Aia?oa aecia/aii /anoioo oi?n?eiiai ooiaethaaiiy Ia-a?oie Thy, eiai aia?aaoe/iee aa?’?? oa na?aaeiueieaaae?aoe/io aiie?ooaeo oi?n?eieo e?a?aoe?e. I?aeoaa?aeaeaii aaeaeaaoi?noue ? aeaoae?ciaaii eeane/io iiaeaeue Iae??aa iecueei/anoioii? ?ooiiino? AeIE: iecueei/anoioia aeia?ioia iiaea, ui niinoa??aa?oueny a AeIE ? neioaoe/ieo aeaieaioethaiaeo iie?ioeeaioeaeao a ?aiai?anueeeo niaeo?ao, ?ioa?i?aoiaaia ye iicaeiaaeiy eieaeoeaia e?a?aoe?y nia?aieo iniia a?aeiinii ae?eiceaeieo ca’yce?a, ui aoaeoeaii aca?iiae?? ye c iicaeiaaei?ie aeonoe/ieie oiiiiaie, oae ? c eieaeoeaieie ?iaa?nieie eieeaaiiyie ai?iia?oi, ? a?aeaoaa?oueny i?ae “aeiiiaiaiaio” oi?n?eiiai ooiaethaaiiy Ia-a?oi Thy. A?ieia?/ia cia/ou?noue oe??? iiaee aaa/a?oueny o ?? o/ano? o i?ioeanao ?aie?eaoe?? AeIE, a oaeiae o ia?aaea/? neaiae?a aeaeaeiae?? acaeiaae AeIE. Aia?oa aecia/aii aia?aaoeeo ? eiiia?aoeai? aeanoeaino? i?aeiieaeoey?ieo I-ca’yce?a o e?enoaeao ? ni?ae?enoae?caoao ioeeaioeaeieo iniia – iiaeaeueieo ia’?eoao ioeea?iiai-ioeea?iiaiai ai?ciaaaiiy – oa ?? aianie o noaa?e?caoe?th inoaii?o. IA?AE?E INIIAIEO IAOEIAEO I?AOeUe, IIOAE?EIAAIEO IA OAIO AeENA?OAOe?? Aiai?oi Ae.I., Ieuoe ss.?., AEaeoianeee I.A. Iecei/anoioiua oiiiiiua niaeo?u eiiaeiaoeeiiiiai ?annayiey naaoa eiiiiiaioia ioeeaeiiauo eeneio: io?eiiaua iniiaaiey // Aeiiieeia?u e eeaoea.-1990.-6,N3.-N.24-31. Aiai?oi Ae.I., Ieuoe ss.?., AEaeoianeee I.A. Iecei/anoioiua oiiiiiua niaeo?u eiiaeiaoeeiiiiai ?annayiey naaoa eiiiiiaioia ioeeaeiiauo eeneio: ie?eieaeeiiaua iniiaaiey // Aeiiieeia?u e eeaoea.-1990.-6,N3.- N.31-39. Aiai?oi Ae.I., Ieuoe ss.?., AEaeoianeee I.A. E aii?ino i i?e?iaea iecei/anoioiuo niaeo?ia eiiaeiaoeeiiiiai ?annayiey naaoa aeiiieeia?ia // Aeiiieeia?u e eeaoea.-1990.-6,N4.-N.79-83. Aiai?oi Ae.I., Eiiae?aothe E.A., Ieuoe ss.?., AEaeoianeee I.A. Eneeth/aiea aeeyiey ianoaaeeueiinoe iiuiinoe eaca?a ia oi/iinoue ?aaeno?aoeee niaeo?ia eiiaeiaoeeiiiiai ?annayiey naaoa aeiiieeia?ia // Aeiiieeia?u e eeaoea.-1990.-6,N4.-N.105-107. Aiai?oi Ae.I., Ieuoe ss.?., AEaeoianeee I.A. 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IE, oi?eaeueii ii?aeiothoue o nia? oe?eo ieceo aca?iiiia’ycaieo oi?i no?oeoo?ii? i?ieeaino? – i?ioio?iiio oaooiia??th, no?oeoo?io iaaei?noe?noue, aaiiao?e/io ?ciia??th, caeaoi?noue i?e?aeioaaoe aai a?aeuaiethaaoe i?ioii, niiioaiii aeacai?ioaaoeny ? ieenethaaoeny oiui. ?ic?iaeaii iiaeae? niiioaiieo iai?a?ice?eoeo noai?a AeIE oa ?? ia?aaeieaaeaiiy, aeaeaeiae?? a AeIE oa ?? i?inoi?iai? aee?eaeaiino?, aaiioiene/iiai aieeao ia ia? OO- ii?ii?iaiiy, eiiia?aoeaiino?; anoaiiaeaii iniiai? i?e?ino?oeoo?i? /eiieee no?oeoo?ii-aeeiai?/ii? aaoa?iaaiiino? AeIE. I?aaenoaaeaii iecueei/anoioi? ?aiai?anuee? niaeo?e eiiiiiaio?a IE, ui iine?aeiaii oneeaaeiththoueny, oa ni?ae?enoae?cao?a oeio “iniiaa – iio?aeia ai?iieeneioe”. I?aeoaa?aeaeaii eeane/io iiaeaeue Iae??aa iecueei/anoioii? ?ooiiino? AeIE. Aeine?aeaeaii aia?aaoeeo ? eiiia?aoeai? aeanoeaino? I-ca’yce?a o e?enoaeao ? ni?ae?enoae?caoao iniia AeIE. Eeth/ia? neiaa: a?eeiai-ioeea?iiaa ? ioeea?iiai-ioeea?iiaa ai?ciaaaiiy; AeIE; I-ca’ycie; eiii’thoa?ia iiaeaethaaiiy; ?aiai?anueea niaeo?ineii?y; no?oeoo?ii-aeeiai?/i? aeanoeaino?; ai?iia?oia. Aiai?oi Ae.I. Oeceei-oeie/aneea iaoaieciu aeiiieaeoey?iiai ociaaaiey. – ?oeiienue. Aeenna?oaoeey ia nieneaiea o/aiie noaiaie aeieoi?a aeieiae/aneeo iaoe ii niaoeeaeueiinoe 03.00.03-iieaeoey?iay aeieiaey. - Einoeooo iieaeoey?iie aeieiaee e aaiaoeee IAI Oe?aeiu, Eeaa, 1999. Onoaiiaeaiu iaeaieaa iauea oeceei-oeie/aneea iaoaieciu aaeeiai-ioeeaeiiaiai e ioeeaeiiai-ioeeaeiiaiai ociaaaiey. ?anoe?aiu i?aaenoaaeaiey i no?oeoo?ii-aeeiaie/aneeo naienoaao ioeeaioeaeiuo iniiaaiee e eo noaeeiaa. Iieacaii, /oi iniiaaiey IE, eae e naie IE, oieeaeueii iauaaeeiytho a naaa oeaeue ?yae acaeiinaycaiiuo oi?i no?oeoo?iie eciai/eainoe - i?ioio?iiioth oaooiia?eth, no?oeoo?ioth iaaeanoeinoue, aaiiao?e/aneoth eciia?eth, niiniaiinoue i?eniaaeeiyoue eee iouaieyoue i?ioii, niiioaiii aeacaieie?iaaoueny e ieeneyoueny a aiaea e o. i. ?ac?aaioaiu iiaeaee niiioaiiuo iieo?ane?uouo ninoiyiee AeIE e aa i?aaeieaaeaiey, aeaeaeiaeaenoaey a AeIE e aa i?ino?ainoaaiiie ene?eaeaiiinoe, aaiioiene/aneiai aeeyiey ia iaa OO- iaeo/aiey, eiiia?aoeaiinoe; onoaiiaeaiu iniiaiua iee?ino?oeoo?iua oaeoi?u no?oeoo?ii-aeeiaie/aneie aaoa?iaaiiinoe AeIE. I?aaenoaaeaiu iecei/anoioiua ?aiaiianeea niaeo?u iineaaeiaaoaeueii oneiaeiythueony eiiiiiaioia IE e nie?enoaeeecaoia oeia “iniiaaiea - i?iecaiaeiay aieiieeneiou”. Iiaeoaa?aeaeaia eeanne/aneay iiaeaeue Iaeaaaa iecei/anoioiie iiaeaeaeiinoe AeIE. Enneaaeiaaia yia?aaoeea e eiiia?aoeaiua naienoaa I-naycae a e?enoaeeao e nie?enoaeeecaoao iniia AeIE. Eeth/aaua neiaa: aaeeiai-ioeeaeiiaia e ioeeaeiiai-ioeeaeiiaia ociaaaiea; AeIE; I-naycue; eiiiuethoa?iia iiaeaee?iaaiea; ?aiaiianeay niaeo?ineiiey; no?oeoo?ii-aeeiaie/aneea naienoaa; aieiia?oiia. Hovorun D.M. Physico-chemical mechanisms of biomolecular recognition. - Manuscript. Thesis for the Doctor of sciences' degree by speciality 03.00.03-molecular biology. - Institute of Molecular Biology and Genetics of Ukrainian National Academy of Sciences, Kyiv, 1999. The dissertation is devoted to the study of the most general physico-chemical mechanisms of a protein-nucleic acid and nucleic acid-nucleic acid recognition. It was established that the NA bases with aminogroup are molecules with non-rigid structure, dipole-active interconversion of which occurs via three energetically and topologically non-equivalent ways - the "over-barrier" large amplitude inverse vibration of amino fragment and two essentially anisotropic "under-barrier" rotations of aminogroup. It was shown, that stacking of the NA bases and their Watson-Crick pairs does not boil down to non-specific interactions of static planar-parallel structures: contribution of specific Coulomb contacts of amino nitrogen atoms' lone-pairs with hydrogen atoms of adjacent (upper or lower) bases - so called vertical H-bonds - are substantial. Implying molecular regulation it was suggested to consider the NA bases' aminogroup as "a molecular joint", non-linear mobility of which could be controlled by H-bonds both directly and at a distance. The Watson-Crick base pairs were established to be extremely soft as to "propeller-twist" and "sharp bend" angles and capable of adoption non-planar conformation under influence of small external perturbations, as it takes place in DNA. There are developed microstructural models of spontaneous semiopen states of DNA and its premelting, long-range action in DNA and its spatial curvature, genotoxic effects of UV irradiation. Microstructural sources of a DNA structural-dynamic heterogeneity which dominates in protein-nucleic acid recognition were considered. The conclusion was made that leading stereochemical principle of DNA spatial structure is not isomorphism of Watson-Crick pairs but their structural-dynamical conformity. It was shown that the NA bases like NA themselves accumulate uniquely whole gamma of mutually conjugated forms of structural changeability - prototropic tautomerism, structural non-rigidity, geometric isomerism, capability to accept or to give a proton, their spontaneous deamination or oxidation under the contact with water and so on. It was demonstrated that prototropic tautomerism is far not confined to point mutagenesis. Biological significance of high energy tautomers, inaccessible to direct physico-chemical detection, so called "latent structures", is discussed. It is noted, that their biological importance is not directly connected with their energetic stability. A complete multitude of point H-contacts of the "a nucleotide base - an amino acid residue" type, stabilised by two H-bonds, were investigated. It was shown, that already at this level the recognition occurs according to scenario of mutual structural-dynamical adaptivity with participation of essentially interdependent intra- and intermolecular H-bonds capable not only of forming and breaking but also of switching over in a way of bifurcations. A structural non-rigidity of the bases is believed to be a security of unerring and expeditious recognition in conditions of the free volume deficit which is characteristic part for a living cell. The low-frequency Raman spectra of successively complicated NA components, as well as cocrystallizates of the "a base - an amino acid derivative" type are presented. Dynamic and energetic features of torsional tunnel junction of methyl group of Thy and the bases substituted at glycosidic nitrogen atoms are determined. An adequacy of the classical Maleev's model is confirmed: low-frequency qualitative mode observed in DNA Raman spectra is interpreted as collective extensional optical oscillations, exactly - librations of paired bases around glycosidic bond, which effectively interact with both acoustic extensial phonons and collective inverse vibrations of amino groups, are accompanied by collective torsional tunnel junction of the Thy methyl groups. A biological applications of this mode are discussed. An energetics and cooperative properties of H-bonds in crystals and cocrystals of nucleotide bases – model of nucleic acid-nucleic acid recognition – are investigated. Key words: protein - nucleic acid and nucleic acid-nucleic acid recognition; DNA; H-bond; computer simulation; Raman spectroscopy; structural-dynamical properties; aminogroup. PAGE 1

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