Iaoe?iiaeueia Aeaaeai?y Iaoe Oe?a?ie

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

IAIEAINUeEA Ia??y TH???aia

OAeE 577.113.083:577.218

?AAAIA?AOe?ss IA*?IEE O UO??A :

iieaeoey?ii-a?ieia?/i? i?ioeane,

?o ?aaoeyoe?y oa /aniaa oeaea

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?ieo iaoai?ci?a a?ineioaco a?eea
?inoeoooo iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie (caa. a?aeae?eii
— aeieoi? a?ieia?/ieo iaoe I.I.Ieaoiiia ), a ?inoeooo? aa?iioieia??
AII Oe?a?ie ? a Aeueaa?o-Ethaea?a Oi?aa?neoao? (O?aeao?a, I?ia//eia).

Io?oe?ei? iiiiaioe: aeieoi? a?ieia?/ieo iaoe Nieiiei
Ieaenaiae? Iao?iae/ ?inoeooo iieaeoey?ii? a?ieia?? ? aaiaoeee
IAI Oe?a?ie, Ee?a caa. a?aeae?eii a?io?i?/ii? aaiaoeee;

aeieoi? iaaee/ieo iaoe, i?ioani? Aeei??c Aiaoie?e Eineiiae/
?inoeooo aenia?eiaioaeueii? iaoieia??, iieieia?? ?
?aae?ia?ieia?? ?i. ?.?. Eaaaoeueeiai IAI Oe?a?ie, Ee?a caa.
a?aeae?eii a?ieia?? iooeeiii? ee?oeie;

aeieoi? a?ieia?/ieo iaoe, i?ioani? Noieea ?inoeneaa
Noaiaiiae/ ?inoeooo a?io?i?? ?i. aeaae. I.A. Iaeeaae?ia IAI
Oe?a?ie, Euea?a aee?aeoi? a?aeae?eaiiy ?aaoeyoi?ieo nenoai
ee?oeie.

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

Caoeno aeena?oaoe?? a?aeaoaeaoueny _22 /a?aiy 1999?. i 10 aiaeei? ia
can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae26.237.01 ?inoeoooo
iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie ca aae?anith: 252143, i.
Ee?a-143, aoe.Caaieioiiai 150.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? ?inoeoooo
iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie ca aae?anith: 252143, i.
Ee?a-143, aoe.Caaieioiiai 150.

Aaoi?aoa?ao ?ic?neaiee 22 o?aaiy 1999 ?.

A/aiee nae?aoa? niaoe?ae?ciaaii? a/aii? ?aaee eaiaeeaeao
a?ieia?/ieo iaoe E.E. Eoeao

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue oaie. ?aaaia?aoe?y ia/?iee — neeaaeiee eiiiainaoi?iee
i?ioean a?aeiiaeaiiy iane ? ooieoe?? i?aaio aiane?aeie iaiai?ioiiai
ooeiaeaeaiiy aai iaoai?/iiai aeaeaeaiiy /anoeie eiai ia?aio?ie. Oeae
i?ioean noi?oue a iaeiiio ?yaeo c oaeeie a?ieia?/ieie yaeuaie ye ??no,
?icaeoie, aeeoa?aioe?thaaiiy, aiiioic. ?icoeo?oaaiiy iieaeoey?ieo
iaoai?ci?a, ye? coiiaeththoue oe? i?ioeane, caeeoa?oueny aeooaeueiith
i?iaeaiith no/anii? a?ieia??. Ia?aaeoiiae aeey ae??oaiiy oaeeo caaea/
noai?ai? aeinyaiaiiyie a ??cieo aaeocyo a?ieia??, cie?aia, a oea?ei?
aeni?an?? aaiiio ? i?ioean?a aeeoa?aioe?thaaiiy, aca?iiae?? i?ae
ee?oeiaie ??cii? i?e?iaee oa i?ae nenoaiaie i?aai?cio oiui.

?aaaia?oth/a ia/?iea a?ecoi?a aaaaea?oueny eeane/iith iiaeaeeth aeey
aea/aiiy caaaeueieo caeiiii??iinoae ? oeaieiiniaoeeo?/ieo iniaeeainoae
a?aeiiaethaaeueieo i?ioean?a. Iinooeth?oueny, ui ?aaaia?aoe?eiee i?ioean
a ia/?ioe? ia? oae? neeaaeia?: ?aaeoe?y ia ooeiaeaeaiiy; a?io?i?/ia
aaeaioaoe?y i?aaio aei ooieoe?iiaeueiiai iaaaioaaeaiiy, ui c?inei;
ia?ao?ae ee?oei ?c i?ie?oa?aoeaii iaaeoeaiiai noaio aei ee?oeiiiai
iiae?eo ? ?aaeoe??, ye? coiiaeththoue ci?io noa?iai oaiioeio; aeania
i?ie?oa?aoe?y ee?oei; ?aaeoe??, ye? coiiaeththoue ia?ao?ae a?ae
ee?oeiiiai iiae?eo aei noaio «niieith». A ?ioaa?aeueiiio i?ioean?
?aaaia?aoe?? ia/?iee i?eeiathoue o/anoue an? ee?oeie i?aaio. ?aaoeyoe?y
ee?oeiiniaoeeo?/ieo ooieoe?e oa iiae?eo ee?oei iaeiiai oeio, a oaeiae
aca?iiae?y i?ae ee?oeiaie ??cieo oei?a cae?enith?oueny ca aeiiiiiaith
neaiaeueieo iieaeoe, cie?aia oeo, ui ae?iaeythoueny ee?oeiaie ia/?iee.
I?ioeane, ui a?aeaoaathoueny a ?aaaia?oth/?e ia/?ioe?, eii?aeeiothoueny
oaeiae c iio?aaaie oe?e?niiai i?aai?cio. Ana oea coiiaeth? neeaaeio
???a?o?th ?aaoeyoi?ieo i?ioean?a ia ??cieo ??aiyo i?aai?caoe?? —
noaee?oeiiiio, ee?oeiiiio, i?aaiiiio oa i?aai?ciaiiiio, ? caaacia/o?
/?oeo /aniao oa i?inoi?iao i?aai?caoe?th i?ioeano.

E?ioeaaa iaoa o aea/aii? ?aaaia?aoe?eiiai i?ioeano iieyaa? o
?icoeo?oaaii? eeth/iaeo iaoai?ci?a eiaeii? neeaaeiai? i?ioeano ? cania?a
?o eii?aeeiaoe?? ye i?ae niaith, oae ? c aeiiaaie i?aai?cio, ui
ci?iththoueny. Oe? ciaiiy iiaeii? ?icoe?eoe oyaeaiiy i?i i?eioeeie
ooieoe?iioaaiiy aeeaiai ? iiaeooue aooe aeei?enoai? aeey oi?iaeueiiai
iienaiiy a?aeiiaethaaeueiiai i?ioeano, ui noaei ae nooo?aei aoaiii o
eiai i?ciaii?.

Ca?ac c?a?aiee aaeeeee iaoa??ae i?i a?io?i?/i? i?ioeane, ye?
a?aeaoaathoueny a ia/?ioe? oa a i?aai?ci? i?ae /an a?aeiiaeaiiy ia/?iee,
?niothoue ??ci? a?iioace a?aeiinii i?e?iaee i?ioeano, iaeiae e?ioeaai?
iaoe ua ia aeinyaiooi. A?aenooiy caaaeueia noaia i?ioeano, a ye?e ae
a?aoiaoaaany eiai ?icaeoie a /an? ? a i?inoi?? ??cii??aeiiai ca nai?i
ee?oeiiei neeaaeii i?aaio. sse ? ?ai?oa aeooaeueieie
caeeoathoueny

ieoaiiy i?i i?e?iaeo ?aaoeyoi?ieo oaeoi??a, ui caioneathoue
?aaaia?aoe?th, eii?aeeiothoue ae?yeuei?noue an?o ee?oei i?ioyaii
a?aeiiaethaaeueiiai i?ioeano ? caaa?oothoue eiai. Caeiaooee no/anii?
iieaeoey?ii? a?ieia?? aeathoue iiaeeea?noue caa?oaoeny aei oeeo ieoaiue
? ae??ooaaoe ?o ye ia iiaiio oai?aoe/iiio, oae ? ia iaoiaee/iiio ??ai?.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. ?iaioa
a?aeiia?aea? iniiaiiio ieaio ooiaeaiaioaeueieo aeine?aeaeaiue, ye?
i?iaiaeeeeny o a?aeae?e? iieaeoey?ieo iaoai?ci?a a?ineioaco a?eea ?IA?A
IAI Oe?a?ie (caa. a?aeae?eii aeieoi? a?ieia?/ieo iaoe I.I. Ieaoiiia)
(athaeaeaoi? oaie: ?01.86.0069116 — «Oa?aeoa?enoeea oeeoiieaciaoe/ieo
a?eeiaeo oaeoi??a, ye? aeoeaothoue yaea?iee aaiii ia ?aii?o noaae?yo
?aaaia?aoe?? ia/?iee»; ?01.9100034 — «Aea/aiiy aai?a, aeni?an?y yeeo
caeaaeeoue a?ae ee?oeiiiai oeeeeo aaiaoioeeo?a»; ?0195U005358 —
«Iine?aeiai?noue aeni?an?? i?ioiiieiaai?a ? oeaieiiniaoeeo?/ieo aai?a o
ee?oeiiiio oeeee? aaiaoioeeo?a». *anoeia ?iaioa aeeiioaaeany a ?inoeooo?
aa?iioieia?? AII Oe?a?ie oa a ?inoeooo? ee?oeiii? ? iieaeoey?ii?
a?ieia?? Aeueaa?o-Ethaea?a Oi?aa?neoaoo (O?aeao?a, I?ia//eia) ca
i?iaeoii SFB154 a?ae I?iaoeueeiai iaoeiaiai oiaa?enoaa.

Iaoa ? caaea/? aeine?aeaeaiiy. Iaoa ?iaioe iieyaa? o c’ynoaaii? aeayeeo
iieaeoey?ieo iaoai?ci?a, ye? caaacia/othoue ia?ao?ae
aenieiniaoe?ae?ciaaieo ? iaaeoeaieo o i?ie?oa?aoeaiiio a?aeiioaii?
ee?oei ia/?iee aei i?ie?oa?aoe?? oa eii?aeeiaoe?th ?o
ee?oeiiniaoeeo?/ii? ? i?ie?oa?aoeaii? aeoeaiino? i?ioyaii
a?aeiiaethaaeueiiai i?ioeano, coiiaeaiiai /anoeiaith aaiaoaeoii??th
(*AA). Iniiaio oaaao ieaioaaeiny i?eae?eeoe aea/aiith aeni?an?? aaiiio ?
aeayeeo eiai aai?a, ui eiaeothoue niaoeeo?/i? oa ianiaoeeo?/i? aeey
ia?aio?iieo ? iaia?aio?iieo ee?oei ia/?iee ?aaoeyoi?i? oaeoi?e. Aeey
aeinyaaiiy iinoaaeaii? iaoe ae??ooaaeenue oae? caaea/?: 1.
Aaanoe i???ioe?e aeey noai?aiiy /aniai? oeaee ?aaaia?aoe?? ia/?iee ye
aeeiai?/iiai i?ioeano;
2. Aeine?aeeoe aeaye? iniaeeaino? aeni?an?? aaiiio ?
?aaoeyoi?ieo oaeoi??a i?ioyaii ia?aoiaeo ia/?iee a?ae «niieith» aei
i?ie?oa?aoe??, ine?eueee oeae ia??iae ? e?eoe/iei aeey iiaeaeueoiai
a?aeiiaeaiiy i?aaio; 3. Aeine?aeeoe, ye i?ioyaii a?aeiiaethaaeueiiai
i?ioeano ?aaoeththoueny oa?aeoa?i? aeey aaiaoioeeo?a aeanoeaino?, a naia
caeaoi?noue ciaoeiaeaeoaaoe /oaea??aei? ?a/iaeie ? neioacoaaoe
ee?oeiiniaoeeo?/i? a?eee. Aeey ioe?iee oeeo aeanoeainoae ia?aii
aeni?an?th a?aeiia?aeieo aai?a, ye? eiaeothoue oeeoio?ii ?4502A1, a?eie
ia?oi? oace aeac?ioieneeaoe??, ? ee?oeiiniaoeeo?/iee o?aine?eioe?eiee
oaeoi? — yaea?iee oaeoi? aaiaoioeeo?a 1 SYMBOL 97 \f «Symbol» (ssOA-1
SYMBOL 97 \f «Symbol» ); 4. Aeine?aeeoe ee?oeiiniaoeeo?/io
? i?ie?oa?aoeaio aeoeai?noue iaia?aio?iieo ee?oei ia/?iee ia i?eeeaae?
neioaco, cie?aia, oaeoi?a iae?ico iooeei ? AeIE;

5. I?iaanoe ii??aiyeueia aeine?aeaeaiiy aeni?an?? ee?oeiiniaoeeo?/ieo
(ssOA-1 SYMBOL 97 \f «Symbol» ) ? ee?oeiiianiaoeeo?/ieo (c-Fos, c-Myc)
o?aine?eioe?eieo oaeoi??a a ia/?ioe? o uo??a i?ney *AA ? eaia?ioii??
(EAI) c oei, uia aecia/eoe ?ieue oeeo oaeoi??a o i?ioiaeaeaii?
aaiaoioeeoaie ee?oeiiiai oeeeeo oa o a?aeiia?ae? aaiaoioeeo?a ia o?aaio;

6. C’ynoaaoe ye aieeaathoue iiae?? iinoo?aine?eioe?eiiai ??aiy ia
?aaoeyoe?th aeni?an?? aaiiio oa ie?aieo aai?a o ?aaaia?oth/?e ia/?ioe?,
cie?aia, oeyoii aeine?aeaeaiiy neioaco ? aioo??oiueiee?oeiiiai
?iciiae?eo iiainoai?aii? ?IE, a?aeiiniiai ai?noo ?iaeea?aeoaeueieo
o?aine?eio?a o o?aeoe?yo yaea?ii? ? oeeoiieaciaoe/ii? ?IE, a oaeiae
oeyoii aecia/aiiy aiceiaoe/ii? aeoeaiino? nenoaie
2′,5′-ie?ai(A)neioaoaca-?IEaca L, yea ?aaoeth? ai?no ?IE a ee?oei?;

7. Caaaeath/e ia aaaeeeao ?ieue ieeno acioo ye oi?aa?naeueiiai
a?i?aaoeyoi?a, aecia/eoe eiai i?iaeoeoe?th ??cieie ee?oeiaie
?aaaia?oth/i? ia/?iee ? iiaeeeao ?ieue o a?aeiiaeaii? iane ? ooieoe??
i?aaio.

Iaoeiaa iiaecia iaea?aeaieo ?acoeueoao?a. Aia?oa cai?iiiiiaaii
aeei?enoiaoaaoe iieacieee neioaco AeIE o ee?oeiao ia/?iee ??ciiai oeio
ye i???ioe?e aeey noai?aiiy /aniai? oeaee ?aaaia?aoe?eiiai i?ioeano.

Ia i?aenoaa? aeaieo i?i neioac ?IE ? ?iciiae?e iiainoai?aii? ?IE a
ee?oeiieo no?oeoo?ao aaiaoioeeo?a, a oaeiae aeaieo i?i aeni?an?th aaiiio
oa ?iaeea?aeoaeueieo aai?a aia?oa aeae?eaii aeaa i?eioeeiiai ??cieo
i?ioeane i?ioyaii ia?aoiaeo ia/?iee ?c noaio «niieith» aei noaio
i?ie?oa?aoeaii? aeoeaiino?. Aeey ia?oiai i?ioeano oa?aeoa?i? ieoo?a?
eiiiainaoi?i? ?aaeoe??: aeoeaaoe?y caaaeueiiai neioaco ?IE o
aaiaoioeeoao ? neioaco a yae?ao ? i?oioiiae??yo, i?aeaeuaiiy ai?noo
aeine?aeaeaieo ?iaeea?aeoaeueieo i?IE. Ai?no ee?oeiiniaoeeo?/ieo ?IE
ssOA-1 SYMBOL 97 \f «Symbol» oa ?4502A1 c?inoa? a?eueoith i??ith, i?ae
ai?no ?IE, ye? eiaeothoue ee?oeiiianiaoeeo?/i? o?aine?eioe?ei? oaeoi?e
c-Myc oa c-Fos. Ae?oaee i?ioean oa?aeoa?eco? ci?io oaiioeio ee?oei
ia/?iee ? i?iyaey?oueny oei/aniaei cieaeaiiyi neioaco ?IE, cao?eieith
iiainoai?aii? ?IE a yae?? ? a?aeiia?aeii iaioei iaaeoiaeaeaiiyi ?? a
oeeoiieacio. Aia?oa i?iaeaiiino?iaaii cieaeaiiy ? i?aeaeuaiiy aeoeaiino?
nenoaie 2′,5′-ie?ai(A)neioaoaca-?IEaca L a?aeiia?aeii a yaea?i?e ?
oeeoiieaciaoe/i?e o?aeoe?yo. Aeacaia nenoaia a?aeiia?aea? ca ?icuaieaiiy
aea?aieo iieaeoe ?IE. Aia?oa aeyaeaii i?aeaeuaiiy aeoeaiino? ?ioa?oa?iio
SYMBOL 97 \f «Symbol» / SYMBOL 98 \f «Symbol» , niaoeeo?/iiai
?iaeoeoi?a nenoaie 2′,5′-ie?ai(A)neioaoaca-?IEaca L.

Aia?oa aeyaeaii i?aeaeuaiiy aeni?an?? oaeoi?a iae?ico iooeei i?ioyaii
ia?ao?aeiiai ia??iaeo, ui na?ae/eoue i?i aeoeaaoe?th ee?oeiiniaoeeo?/ieo
ooieoe?e ee?oei Eoioa?a aai «in?eeo» iae?ioaa?a

ia/?iee. I?iaeaiiino?iaaii c?inoath/o ni?eeiyoeea?noue ee?oei ia/?iee
aei oeueiai oaeoi?o aiane?aeie i?aeaeuaiiy aeni?an?? iaio ?aoeaioi??a.
Aia?oa c’yniaaii, ui a ia??iae, eiee a ?aaaia?oth/?e ia/?ioe? iaeii/anii
a?aeaoaathoueny neioac AeIE a aaiaoioeeoao ? oeaieiiniaoeeo?/ia
a?aeiia?aeue ia o?aaio, i?aeaeuaia aeni?an?y o?aine?eioe?eieo oaeoi??a
c-Fos i ssOA-1 SYMBOL 97 \f «Symbol» iia’ycaia c a?aeiia?aeaeth ia
o?aaio. Aia?oa iieacaii, ui a ?aaaia?oth/?e ia/?ioe? ci?ie aeni?an??
oaeoi?o c-Iyc ei?aeththoue c oacaie ee?oeiiiai oeeeeo ? iia’ycai? c
i?ie?oa?aoeaiith a?aeiia?aeaeth c aieo aaiaoioeeo?a.

O oe?e?ni?e oeaiei? ?aaaia?oth/i? ia/?iee oa a iiioeyoe?yo
aaiaoioeeo?a, «in?eeo» iae?ioaa?a ? aiaeioae?aeueieo ee?oei ?c
neioni?ae?a ia/?iee aia?oa aecia/aii neioac ieeno acioo. Aeyaeaii, ui
neioac ieeno acioo iiceoeaii ei?aeth? ?c G1- oa G2-ia??iaeaie ee?oeiiiai
oeeeeo a?aeiia?aeieo ee?oei ? iaaaoeaii — c S-oacith ee?oeiiiai oeeeeo
oeo aea ee?oei.

Ia i?aenoaa? aiae?co e?oa?aoo?ieo aeaieo ? ?acoeueoao?a iaoi? ?iaioe
aia?oa aeneiaeaii aeae?eueea i?eiouaiue a?aeiinii ?ie? niaoeeo?/ii? ?
i?ie?oa?aoeaii? aeoeaiino? ia?aio?iieo ? iaia?aio?iieo ee?oei ia/?iee
i?ioyaii a?aeiiaethaaeueiiai i?ioeano. Neioni?aeaeuei? ee?oeie ia/?iee,
ye? ? ee?oeiaie iacaio?iiiai iioiaeaeaiiy, i?ioyaii ia?ao?aeiiai
ia??iaeo ? ee?oeiiiai oeeeeo aaiaoioeeo?a ?ioaineaii i?iaeoeothoue
neaiaeuei? iieaeoee ? ca ?o aeiiiiiaith ?aaoeththoue ia?ao?ae aei
i?ie?oa?aoe?? ? aeania i?ie?oa?aoe?th aaiaoioeeo?a. C ii/aoeii
ee?oeiiiai oeeeeo neioni?aeaeueieo ee?oei ?o ia?ae?eiia aeoeai?noue
cieaeo?oueny. I?ie?oa?aoeaia aeoeai?noue ee?oei ia/?iee iaeiiai oeio
ii?aeio?oueny c niaoeeo?/iith aeoeai?noth ee?oei ia/?iee ?ioiai oeio.
Aeey eiaeiiai oeio ee?oei ia/?iee ?niothoue ee?oeiiniaoeeo?/i? i?oiaaie,
aeaye? c yeeo iaeii/anii ae?thoue ye ee?oeiiniaoeeo?/i? ?ia?a?oi?e
i?ie?oa?aoe?? aeey ?ioeo ee?oei ?aaaia?oth/i? ia/?iee.

Oai?aoe/ia ? i?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoao?a. Oai?aoe/ia
cia/aiiy iaea?aeaieo ?acoeueoao?a iieyaa? o oiio, ui aiie ?icoe?ththoue
oyao i?i iieaeoey?i? iaoai?cie a?aeiiaethaaeueieo i?ioean?a. Aiie,
cie?aia, na?ae/aoue i?i iaiao?aei?noue aeoeaii? ci?ie «noa?iai» oaiioeio
i?ae /an ia?aoiaeo aenieiaeeoa?aioe?eiaaieo ee?oei aei i?ie?oa?aoe??,
i?i oa?aeoa? aca?iiae?? ee?oei iacaio?iiiai ? ai?oae?aeueiiai
iioiaeaeaiiy, a oaeiae i?i aca?iiae?th ?aaoeyoi?ieo nenoai oaeeo ye ieen
acioo oa 2’5′-ie?ai(A)aaeai?eaoe c oeeoie?iaie. Io?eiai? ?acoeueoaoe
aeathoue oyao i?i i?e?iaeo o?aiene?eioe?eieo oaeoi??a, ye? coiiaeththoue
a?aeiia?aeue ia/?iee ia o?aaio ? i?ioiaeaeaiiy ee?oeiiiai oeeeeo, i?i
iiaea?eio ooieoe?th, i?eiaeii?, aeayeeo ?aaoeyoi??a i?ie?oa?aoe??, ye?
ae?thoue ye ee?oeiiniaoeeo?/i? aeoeaaoi?e ? ee?oeiiniaoeeo?/i?
?ia?a?oi?e i?ie?oa?aoe??.

I?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoao?a iieyaa? o ?o aeei?enoaii?
i?e ieaioaaii? aeine?aeieoeueei? ?iaioe a aaeocyo, iia’ycaieo ?c
aea/aiiyi i?ie?oa?aoe?? oa aeeoa?aioe?thaaiiy ia/?iee. E??i oiai,
io?eiai? ?acoeueoaoe iiaeooue aooe ei?enieie i?e ?ic?iaoe? no?aoaa??
e?eoaaiiy aaaaoueio caoai?thaaiue ia/?iee aai iane?aee?a o??o?a?/iiai
ao?o/aiiy, iai?eeeaae, aeey noeioethaaiiy a?aeiiaethaaeueieo i?ioean?a o
oai?eo ia aaiaoeo aai aeey aaeueioaaiiy ceiye?niiai ?inoo, aeey
eiio?ieth ca ?inoii o?ainieaioiaaii? ia/?iee oiui.

Iniaenoee aeeaae caeiaoaa/a. A?eueo?noue ?acoeueoao?a, ye? aeeeaaeaii
a aeena?oaoe??, iaea?aeai? iiooea/ai naiino?eii. *anoeio
aenia?eiaioaeueieo aeine?aeaeaiue aoei i?iaaaeaii a ni?a?ia?oieoeoa? c
Aa?aneiiaith O.A., A?ee/ E.?., Iia?eiaith E.A., Naciiiaith E.ss.,
Aai?iei A.O., Ii?aea?ioeaaei I.?. ? Ooeueoea-Oiae?ia A. On? aeine?aee
ieaioaaeenue iiooea/ai, i?iaiaeeeenue ? iaaiai?thaaeenue i?ae ??
aaciina?aaei?i ea??aieoeoaii. I?aeaioiaea iaoa??ae?a aei ae?oeo ?
iaienaiiy oaeno?a iaoeiaeo iia?aeiieaiue cae?enithaaeanue aeeeth/ii
iiooea/ai. Iaoa??aee iioae?eiaaii a ni?eueieo ioae?eaoe?yo.

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. ?acoeueoaoe aeine?aeaeaiue aoee
i?aaenoaaeai? ia: IV ananithciiio a?io?i?/iiio c’?cae? (Euea?a, 1979);
IV ananithciiio neiiic?oi? «Iieaeoey?i? iaoai?cie aaiaoe/ieo i?ioean?a »
(Iineaa, 1979); VII ananithciiio neiiic?oi? «No?oeoo?a ? ooieoe??
ee?oeiiiai yae?a» (Oa?e?a, 1980); V ananithciiio neiiic?oi?
«I?aeiieaeoey?i? aca?iiae?? ? eiioi?iaoe?? iieaeoe» (Aeia-Aoa, 1980);
IV neiiic?oi? N?N?-O?I «No?oeoo?a ? o?aine?eioe?y aaiiio» (??aaai,
1981); Oe?a?inueeiio a?io?i?/iiio c»?cae? (Aei?i?iiao?ianuee, 1982); XII
ananithci?e eiioa?aioe?? c aeaeo?iiii? niaeo?ineii?? (Noie, 1982);
?anioae?eainuee?e eiioa?aioe?? «ssaea?i? a?eee ? aeni?an?y aaiiio»
(Eai?a, 1983); VI ananithci?e eiioa?aioe?? «Ieoaiiy ?aaaia?aoe?? ?
ee?oeiiiai ?iciiiaeaiiy» (Iineaa, 1985); V ananithciiio a?io?i?/iiio
c’?cae? (Ee?a, 1986); oeie? «No/ani? i?iaeaie ?aaaia?aoe??» (Eioea?-Iea,
1986); IX ananithciiio neiiic?oi? «No?oeoo?a ? ooieoe?? ee?oeiiiai
yae?a» (*a?iiaieiaea, 1987); V oe?a?inueeiio a?io?i?/iiio c’?cae?
(?aaii-O?aie?anuee, 1987); oeie?-eiioa?aioe?? «No?oeoo?a ? ooieoe?y
a?iiie?ia??a» (Euea?a, 1989); Oaeuee-neiiic?oi? «A?eee a ?aaoeyoe??
aai?a ia/?iee» (Aacaeue, Oaaeoea??y, 1989); Ananithci?e ia?aae?
«Aeooaeuei? ieoaiiy ee?oeiii? a?ieia??» (Eai?ia?aae, 1989); Ananithci?e
oeie?-nai?ia?? «Iieaeoey?ia a?ieia?y ? iaaeeoeeia» (Eai?ia?aae, 1990);
Ananithciiio neiiic?oi? «Iieaeoey?ia ? ee?oeiia iieia?ieia?y» (Euea?a,
1990); VI, VII i VIII i?aeia?iaeieo neiiic?oiao «Ee?oeie neioni?ae?a
ia/?iee» (Aioaa?iai, Aaeuea?y, 1992; E?ioi, ssiii?y, 1994; Ai?aei,
O?aioe?y, 1996); ia?aae? i?iaoeueeiai oiaa?enoaa aeine?aeiee?a ia/?iee
(Aa?e?i, I?ia//eia, 1993); i?aeia?iaeiiio neiiic?oi? «?aaoeyoe?y
aeni?an?? aai?a ia/?iee a ii?i? ? iaoieia?? (Eieae-Ni??ia-Oa?ai?, 1993);

neiiic?oi? aia?eeainueeiai oiaa?enoaa aeine?aeiee?a ia/?iee
«?aaaia?aoe?y ia/?iee» (Aa?e?, NOA, 1993); Oaeuee-neiiic?oi? «Oeeoie?ie
? ia/?iea» (O?aeao?a, I?ia//eia, 1994), Oaeuee-neiiic?oi? ? eiia?an?
«Oai?iae ia/?iee ? ee?i?ea» (Aacaeue, Oaaeoea??y, 1995);
Oaeuee-neiiic?oi? «Neaiae?caoe?y a ia/?ioe?» (O?aeao?a, I?ia//eia,
1997); Oaeuee-neiiic?oi? «Ii?iaeueiee ? ceiye?niee ??no» (Oaeea,
I?ia//eia, 1998).

Ioae?eaoe??. Ca oaiith aeena?oaoe?? iioae?eiaaii 55 ?ia?o, ye?
aeeth/athoue ?icae?e o iiiia?ao??, 26 noaoae, ?c ieo 6 iaeiiin?aieo ? 20
a ni?aaaoi?noa?, 7 — a ca?oa?aeieo aeaeaiyo, ? 28 oac aeiiia?aeae
iaoeiaeo eiioa?aioe?e.

No?oeoo?a ? ianya ?iaioe. Aeena?oaoe?y aeeeaaeaia ia 290 noi??ieao ?
neeaaea?oueny c anooio, iaeyaeo e?oa?aoo?e, ia’?eo?a ? iaoiae?a
aeine?aeaeaiiy, ?acoeueoao?a aeine?aeaeaiiy oa ?o iaaiai?aiiy,
i?aenoie?a ? aeniiae?a. Nienie e?oa?aoo?e aeeth/a? 380 iaeiaioaaiue.
Aeena?oaoe?y i?i?ethno?iaaia 17 oaaeeoeyie ? 57 ?enoieaie.

IA’?EOE OA IAOIAeE AeINE?AeAEAIIss

Aeey ?iaioe aeei?enoiaoaaee naioe?a a?eeo uo??a e?i?? Wistar a?eii 4-5
i?nyoe?a aaaith 200-300 a. ?aaaia?aoe?th ia/?iee iiaeaethaaee ca
aeiiiiiaith iia?aoe?? aeaeaeaiiy e?ai? eaoa?aeueii? oa na?aaeeiii?
/anoie ia/?iee [Higgins, Anderson, 1931], ?aaeoe?th ia/?iee ia o?aaio —
ca aeiiiiiaith eaia?ioii??. Iniiaiei ia’?eoii aeine?aeaeaiiy aoea
ia/?iea ?ioaeoieo ? iia?iaaieo uo??a o ??ci? no?iee i?ney iia?aoe?e.

AeIE-neioaoe/io aeoeai?noue a ee?oeiao ?aaaia?oth/i? ia/?iee aecia/aee
ca aeiiiiiaith 3I-oei?ae?io, yeee aaiaeeee aioo??oiueiaaiii ca 1 aiae
aei aeaeai?oaoe??. Aeae?eaiiy AeIE i?iaiaeeee ca iaoiaeii Oeai?aa ?
Ia?eiaa [1969]. Iieacieee i?ie?oa?aoeaii? aeoeaiino? ee?oei Eoioa?a ?
aiaeioae?aeueieo ee?oei neioni?ae?a aecia/aee ca a?aeiiniei ai?noii
ee?oei c iiaeai?iith e?euee?noth AeIE o iiioeyoe?? a?aeiia?aeieo
?cieueiaaieo ee?oei. Noniaic?th ?cieueiaaieo ee?oei ia?iaeyee ?ic/eiii
?IEace c ianooiiei oa?aoaaiiyi AeIE i?ii?ae?oi?iaeeaeii. A?aeiiniee
ai?no ee?oei c ??ciith e?euee?noth AeIE aecia/aee ia i?eeaae? aeey
ni?ooaaiiy ee?oei c aeei?enoaiiyi i?ia?aie CELLFIT ? iiaeae? SOBR.

Aeine?aeaeaiiy aioo??oiueiee?oeiiiai neioaco ? ?iciiae?eo iiaiooai?aii?
?IE a aaiaoioeeoao i?iaiaeeee iaoiaeii aeaeo?iiii?e?ineii?/ii?
aaoi?aae?ia?ao??. O ieae/acacia/ai? oa?i?ie uo?ai aaiaeeee
aioo??oiuei/a?aaeiii 3I-i?ioiao eeneioo. Ia?iaeo ia/?iee ? i?aeaioiaeo
c??c?a i?iaiaeeee noaiaea?oiei iaoiaeii o iiaeeo?eaoe?? Aa?aneiiai? ?
A?ee/ [1986]. Aeei?enoaia iaoiaeeea aea?

?ici?? ca?ai n??aea iaioee ca iaeiaioo aeine?aeaeaio i?aaiaeo ee?oeie —
i?oioiiae??th, ui aeicaiey? ioioiaeithaaoe ca?ia n??aea c iiaiooai?aiith
?IE, yea iaeaaeeoue oeei ? ?ioei no?oeoo?ai ee?oeie. E?euee?noue ca?ai
n??aea iaae ??cieie eiiia?oiaioaie (yae?i, yaea?oea, i?oioiiae???,
aiaeiieaciaoe/iee ?aoeeoeoi) aaiaoioeeo?a aecia/aee ca aeiiiiiaith
aeaeo?iiiiai i?e?ineiio JEM-IOOB iaoiaeii i?yiiai aeo/aiiy [Iiaeuiia,
Ei?ooeia, 1981].

E?iaoe/io neeaaei?noue yaea?ii? ?IE aecia/aee a ?aaeoe?? a?a?eaeecaoe??
i?ae cieeath/eie e?eueeinoyie ?aae?iaeoeaii i?/aieo o?aeoe?e ee?oeiii?
AeIE ? iaaeeeoeii yaea?ii? ?IE. O?aeoe?? AeIE, a yeeo iine?aeiaiino?
iiaoi?ththoueny aeecueei 105, 103 ? 1-10 ?ac?a, ?icae?eyee ca
oaeaee?noth ?aanioe?aoe?? ? aeae?eyee ca aeiiiiiaith o?iiaoia?ao?? ia
ieneaiaoeo?. AeIE i?oeee a ?aaeoe?? i?e-o?aineyoe?? c 32?-aeOeOO
ca?aeii [Rigby, 1977]. Aeey aeeeth/aiiy ?aanioe?aoe?? AeIE i?ae /an
a?a?eaeecaoe?? ?aaeoe?th cae?enithaaee o i?enooiino? 80% oi?iai?aeo.
Nooi?iue a?a?eaeecaoe?? aecia/aee ye /anoeo ?aae?iaeoeaii? AeIE, yea
aoiaeeoue aei neeaaeo a?a?eaeieo iieaeoe. A?a?eaee a?aeie?aiethaaee a?ae
iaeiieaioethaiai? ioeea?iiai? eeneioe o?iiaoia?ao??th ia ieneaiaoeo?.

A?aeiiniee ai?no ?iaeea?aeoeueieo ?IE aecia/aee a neeaae? yaea?ii? ?
iie?(A)+ oeeoiieaciaoe/ii? ?IE ca iaoiaeii Northern-a?a?eaeecaoe?? oa a
neeaae? oioaeueii? ?IE ca iaoiaeii cai?ioiuei? o?aine?eioe?? —
eaioethaiai? iie?ia?acii? ?aaeoe??. ssaea?io ? oeeoiieaciaoe/io o?aeoe??
ee?oei ia/?iee ?icae?eyee oeaio?eooaoaaiiyi oeaieiiiai aiiiaaiaoo a
?ic/ei? 0,25 I naoa?ice — 2,5% eeiiiii? eeneioe i?e 600 g. Oioaeueio,
yaea?io ? oeeoiieaciaoe/io ?IE aeae?eyee aoai?ae?i?cio?ioe?aiaoiei
iaoiaeii [Cathala et al., 1983]. Iie?(A)+ oeeoiieaciaoe/io ?IE
io?eioaaee oeyoii o?aeoe?iioaaiiy oeeoiieaciaoe/ii? ?IE ia eieiioe? c
ie?ai(aeO)oeaetheicith. eAeIE-ciiaee i?oeee a ?aaeoe?? i?e-o?aineyoe??
aai a ?aaeoe?? c ?icn?yiith cao?aaeith ca?aeii aei ?ino?oeoe?? ae?iaieea
a?aeiia?aeieo ?aaeoea?a (Amersham, Aiae?y). A?a?eaeecaoe?th i?iaiaeeee
ia o?eueo?ao Hybond N+ noaiaea?oiith oaoi?eith Dot- i
Blot-a?a?eae?caoe?? o iiaeeo?eaoe?? Iaoa? oa ?io. [1991]. Nooi?iue
a?a?eaeecaoe?? eAeIE c ?IE e?euee?nii ioe?ithaaee ca aeiiiiiaith
aeaineoiiao??? ?aae?iaaoia?ao?a. A eiaeiee ?c iiiaio?a aeine?aeaeaiiy
aecia/aany ai?no ?iaeea?aeoaeueieo o?aine?eio?a a ?IE o?aeoe?e o %%
noiniaii aei ai?noo a?aeiia?aeieo o?aine?eio?a a ?IE ?c ?ioaeii?
ia/?iee, a oaeiae ai?no ?iaeea?aeoaeueieo o?aine?eio?a o ia?a?aooieo ia
AeIE c a?aooaaiiyi ci?i, ye? noinothoueny ai?noo a?aeiia?aeii? o?aeoe??
?IE a oeaiei? i?ioyaii iia?aoe?eiiai ia??iaeo [Atryzek, Fausto, 1978].

Cai?ioith o?aine?eioe?th oioaeueii? ?IE ? eaioethaiao iie?ia?acio
?aaeoe?th io?eiaii? eAeIE caeenithaaee ca?aeii c i?ioieieii o??ie
Pharmacia (Oaaoe?y). Aeey i?aeaeuaiiy /ooeeaino? eaioethaiai?
iie?ia?e-

caoe?? 5′-i?aeia?e i?oeee ( SYMBOL 103 \f «Symbol» -32?)aeAOO a
O4-iie?ioeeaioeaee?iaci?e ?aaeoe?? ca?aeii aei i?ioieieo o??ie —
ae?iaieea ?aaeoea?a. Oeeee iie?ia?ecaoe?? neeaaeaany ?c aeaiaoo?aoe?? 1
oa i?e 94 SYMBOL 176 \f «Symbol» N, a?aeaeeao 1 oa i?e niaoeeo?/ie
oaiia?aoo?? aeey eiaeii? ia?e i?aeia??a ? aeania iie?ia?ecaoe?? 1 oa.
i?e 72 SYMBOL 176 \f «Symbol» N. Ii/aoeiaa aeaiaoo?aoe?y ? caaa?oaeueia
iie?ia?ecaoe?y o?eaaee a?aeiia?aeii 5 ? 25 oa. ?aaeoe?eio noi?o
i?aeaeaaaee aeaeo?ioi?aco a 5% iie?ae??eai?aeiiio aae? ca
iaaeaiaoo?oth/eo oiia. ?aae?iaeoeai?noue oa?aeoa?ieo nioa ia
?aae?iaaoia?aoao aecia/aee ca iaoiaeii *a?aieiaa. Aeoeai?noue
2′,5′-ie?ai(A)neioaoace aecia/aee a yaea?i?e ? oeeoiieaciaoe/i?e
o?aeoe?? S15 ca e?euee?noth i?/aiiai AIO, yeee aeeth/a?oueny a
ie?aiaaeai?eaoe a i?enooiino? aiceio [Smekens-Etienne et al.,1983].
ssae?a aeae?eyee a ?ic/ei? 0,32 I naoa?ice a i?enooiino? ?ia?a?oi??a
i?ioa?iac ? i/euaee oeaio?eooaoaaiiyi /a?ac 1,5 I naoa?ico [Bush et
al., 1967]. Aicei ?c yaea?ii? ? oeeoiieaciaoe/ii? o?aeoe?? S15 ao?iii
ca’ycoaaee c iaaeeeoeii eiiie?ia?a iie?? — iie?Oe, yeee o?enoaaee ia
iaia?? Whatman 3II. I?iaeoeoe ?aaeoe?? ie?aiia?ecaoe?? ?icae?eyee ca
aeiiiiiaith oiieioa?iai? o?iiaoia?ao?? ia iie?ao?eai?i?iioeaetheic? ?
e?euee?nii aecia/aee ca ?o ?aae?iaeoeai?noth o noeeioeeyoe?eiiio
e?/eeueieeo.

Ai?no ?IEace L aecia/aee iaoiaeii eiaaeaioiiai ca’ycoaaiiy ??
niaoeeo?/iiai aeoeaaoi?a 2-5A4-3′-(32?)oOe [Bisbal et al., 1989].
?aaeoe?th ca’ycoaaiiy ie?aiaaeai?eaoo cae?enithaaee a aeaio aa??aioao —
c ee?oeiiei aeno?aeoii ia/?iee c ianooiiei aeaeo?ioi?acii ? c
Western-aeioii ee?oeiiiai aeno?aeoo. O eiaeiee c iiiaio?a aeine?aeaeaiiy
e?euee?noue ca’ycaiiai ie?aiaaeai?eaoo aecia/aee aeaineoiiao???th
aeaeo?ioi?aa?ai ? ii a?aeiioaiith aei e?eueeino? ie?aiaaeai?eaoo,
ca’ycaiiai c aiceiii ?ioaeoii? ia/?iee, aeaeaeaii? i?ae /an *AA o o???
ae oaa?eie. Aeoeai?noue ?ioa?oa?iio SYMBOL 97 \f «Symbol» / SYMBOL 98
\f «Symbol» a aeno?aeo? oeaieie ia/?iee aecia/aee ca eiai caeaoi?noth
i?eai?/oaaoe oeeoiiaoiaaiio ae?th a??ono aaceeoey?iiai noiiaoeoo a
eoeueoo?? ee?oei [Ho, Enders, 1959]. ??aaiue neioaco ieeno acioo a
ia/?ioe? aecia/aee iaoiaeii aeaeo?iiiiai ia?aiaai?oiiai ?aciiano [Vanin
et al., 1984]. Iaoiae aaco?oueny ia caeaoiino? aeeaoeeo?iea?aaiaoo
(AeAOE), yeee aaiaeeoueny ca 30 oa aei aeaeai?oaoe??, ca’ycoaaoeny c
aiaeiaaiiei iaaaiiaei cae?cii c ooai?aiiyi «ianoee» aeey ieeno acioo.
I?e aca?iiae?? aiaeiaaiiiai ieeno acioo c «ianoeith» ooai?th?oueny
iiiii?o?iceeueiee eiiieaen cae?ca c AeAOE, yeee ? aea? niaoeeo?/iee
AI?-neaiae. Niaeo?e AI? aecia/aee ia ?aae?iniaeo?iiao?? AI? Varian E9.
NO-neioaoe/io aeoeai?noue aaiaoioeeo?a, ee?oei Eoioa?a ?
aiaeioae?aeueieo ee?oei neioni?ae?a aecia/aee ca e?euee?noth i?o?eo?a,
ye? iaeiie/othoueny i?ioyaii iaei??? aeiae o eoeueoo?aeuei?e ??aeei?
a?aeiia?aeieo ?cieueiaaieo ee?oei [Griess et al., 1982].

Aaiaoioeeoe ?ciethaaee eieaaaiaciei iaoiaeii [Tran-Thi et al., 1985], a
ee?oeie Eoioa?a ? aiaeioae?aeuei? ee?oeie neioni?ae?a —
i?iiacii-eieaaaiaciei iaoiaeii c ?o iiaeaeueoei ?icae?eaiiyi a
aetho??aoi?? [Eyhorn et al., 1988]. ?cieueiaai? ee?oeie ?aeaioeo?eoaaee
ca no?oeoo?ith, caeaoi?noth iiaeeiaoe /anoeiee eaoaeno ( SYMBOL 198 \f
«Symbol» 1,16µ), i?eoaiaiio ee?oeiai Eoioa?a, iayai?noth oa?aeoa?iiai
aeey aiaeioae?aeueieo ee?oei neioni?ae?a iiaa?oiaaiai aioeaaio, ui
aca?iiae?? c aioeo?eaie RECA [Duijvestijn et al., 1992] ? ca
niaoeeo?/iith aeey ee?oei Eoioa?a ?aaeoe??th ia oei?aoeaoaoanoa?aco
[Shlayer et al., 1988]. ?cieueiaai? ee?oeie eoeueoeaoaaee ia ieanoeeiaeo
ieaioaoao ca?aeii [Tran-Thi et al., 1985; Eyhorn et al., 1988].

Aeai? aeine?aeaeaiue ia?iaeaii noaoenoe/ii c aeei?enoaiiyi SYMBOL 116
\f «Symbol» -e?eoa??th Nouethaeaioa, a aeai? aeaeo?iiii?e?ineii?/ii?
aaoi?aae?ia?ao?? — ca iaoiaeii iaeii- ? aeaioaeoi?iiai aeenia?n?eiiiai
aiae?co [Ieioeineee, 1975].

AENIA?EIAIOAEUeIA *ANOEIA

*aniaa oa?aeoa?enoeea ?aaaia?aoe?eiiai i?ioeano.

Iieacieee AeIE-neioaoe/ii? aeoeaiino? ee?oei ?aaaia?oth/i? ia/?iee
aea/aee c iaoith aeei?enoaiiy ?o ye /aniaeo i???ioe??a i?ioeano
a?aeiiaeaiiy i?aaio. A?aeiinia eaae?noue aecia/aiiy oeeo iieaciee?a,
??ciiiai?oi?noue ?nioth/eo iaoiae?a, oi?aa?naeuei?noue i?iyao
i?ie?oa?aoeaii? aeoeaiino? c aieo ee?oei on?o oei?a coiiaeththoue oeae
aea??.

?ioaineai?noue neioaco AeIE a ia/?ioe? ?ioaeoieo uo??a ? iacia/iith (
0,66 o 104 ?ii o oa-1 o ia-1 AeIE ), aea /a?ac 18 aiae i?ney *AA ieoiia
?aae?iaeoeai?noue AeIE c?inoa? a?eueoa, i?ae a 100 ?ac?a ? o?eia?oueny
ia oeueiio ??ai? a?ey 10 aiae, ui a?aeiia?aea? o?eaaeino? S-oace
(?en.1). *a?ac 38 ? 50 aiae i?ney *AA aeyaeaii aeiaeaoeia? i?aeeiie
a?ineioaco AeIE. sseui ca?aeii c aeaieie Bucher et al. [1964], ia?oee
i?aeeii iiai?noth coiiaeaiee neioacii AeIE a aaiaoioeeoao, oi ae?oaee
iia’ycaiee c aeoeai?noth ai?oae?th aeia/ieo i?ioie?a ?
ia?eneioni?aeaeue-ieo c??/anoeo ee?oei [Tanaka et al., 1990;
Michalopoulos, 1997], a oaeiae c aeoeai?noth iaaii? o?aeoe??
aaiaoioeeo?a, ye? iiaoi?ii aaeathoueny aei i?ie?oa?aoe??.

I?ie?oa?aoeaia aeoeai?noue ee?oei Eoioa?a aeyaey?oueny iaio
neio?iiiith, i?ae o aaiaoioeeo?a, ine?eueee ca aeaieie FACS-aiae?co
ee?oeie ciaoiaeyoueny a oacao S — G2 i?ioyaii 37-62 aiae (?en. 2A).

A?aeiia?aeia oaca a aiaeioae?aeueieo ee?oei neioni?ae?a i?eiaaea? ia
i?ii? aeie /ano i?ae 72 ? 120 aiae i?ney *AA (?en.2A).

A?aoiaoth/e cacia/ai? ? ?ai?oa a?aeii? iieacieee i?ie?oa-?aoeaii?
aeoeaiino? ee?oei ?aaaia?oth/i? ia/?iee, i?ii?aeie /ano a?ae ii/aoeo
G1-oace ee?oeiiiai oeeeeo aaiaoioeeo?a (aeecueei 3-4 aiae i?ney *AA) [Lu
et al., 1992] aei, i?eiaeii?, 6-i? aeiae i?ney iia?aoe?? iiaeia
?icaeyaeaoe ye ee?oeii? oeeeee aaiaoioeeo?a, ai?oae?y aeia/ieo i?ioie?a,
ia?eneioni?aeaeueieo c??/anoeo ee?oei, ee?oei Eoioa?a ? aiaeioae?aeueieo
ee?oei neioni?ae?a, ui a?aeaoaathoueny a iacaai?e iine?aeiaiino?.

?en. 2. A?aeiiniee ai?no ee?oei Eoioa?a (A) ? aiaeioae?aeueieo ee?oei
neioni?ae?a (A) a G0- ? G1- oa a S- ? G2-oacao ee?oeiiiai
oeeeeo.

A?eueo aeieeaaeieie i???ioe?aie ? iieacieee ie?aieo oac ee?oeiiiai
oeeeeo ??cieo ee?oei.

I?ii?aeie /ano a?ae iia?aoe?? aei ii/aoeo ia?oiai ee?oeiiiai oeeeeo
aaiaoioeeo?a aeae?ey?oueny ye ia?ao?aeiee, ? aeey eiai ia??iaeecaoe??
aeei?enoai? /enaeuei? iieacieee aeni?an?? aaiiio ? ie?aieo aai?a.

Aeni?an?y aaiiio ? aeaye? iaoai?cie ?? ?aaoeyoe?? i?ioyaii
ia?ao?aeiiai ia??iaeo ?aaaia?aoe?eiiai i?ioeano

Neioac ? aioo??oiueiee?oeiiee ?iciiae?e iiaiooai?aii? ?IE

O aaiaoioeeoao ia/?iee uo??a i?ney *AA ? EAI iaoiaeii
aeaeo?iiii?e?ineii?/ii? aaoi?aae?ia?ao?? ioe?ithaaee neioac ?
aioo??oiueiee?oeiiee ?iciiae?e iiaiooai?aii? ?IE ca na?aaeiueith
/enaeuei?noth ca?ai n??aea iaae ??cieie eiiia?oiaioaie iaeeii/ii?
ee?oeie ? ca a?aeiiniith e?euee?noth ee?oei, a yeeo ca?ia aeyaeythoueny
iaae a?aeiia?aeieie no?oeoo?aie.

?en. 3. ?icoaooaaiiy ca?ai n??aea iaae ee?oeiieie no?oeoo?aie
aaiaoioeeoa.

ss — yae?i, ssAe — yaea?oea, ssI — yaea?ia ii?a, I — i?oioiiae??y, Y? —
aiaeiieaciaoe/iee ?aoeeoeoi, YO — aoo?iiaoei, AO — aaoa?io?iiaoei.

3I-i?ioiao eeneioo (3I-IE) aaiaeeee ia ei?ioeee (ca 15 oa aei caai?o
c?ace?a) ? a?eueo o?eaaeee (aaaaeaiiy a iiiaio iia?aoe??) /an. O ia?oiio
aeiaaeeo iieacieee oa?aeoa?ecothoue ?ioaineai?noue i?ioean?a, a a
ae?oaiio — e?euee?noue ? ?iciiae?e iiaiooai?aii? ?IE aiane?aeie
i?ioean?a, ui a?aeaoaathoueny c ?IE i?ioyaii aeniiceoe?? c ?cioiiii
(?en.3,4).

?en.4. ?ioaineai?noue neioaco ?IE a ioeeaiieaci? (I), yaea?oe? (ssae),
i?oioiiae??yo (I), ?ioaineai?noue ?? iaaeoiaeaeaiiy aei
oeeoiieacie (E?) oa a?aeiinia e?euee?noue caae?yieo a oeeo i?ioeanao
aaiaoioeeo?a a ia/?ioe? uo??a i?ney *AA.

3I-IE aaiaeeee ca 15oa aei caai?o c?ace?a ia/?iee.

t0 i ti — /an caai?o c?ace?a o ?ioaeoieo ? iia?iaaieo uo??a.

Iieacieee aeinoi-a??iino? neee aieeao ca [Ieioeineee, 1970]:

I: O=7>O0,999=5,4; ssae.: O=10,5>O0,999=7,5; I:
O=11,5>O0,999=5,6; Y?: O=4,5>O0,95=2,7

*a?ac 15 oa i?ney aaaaeaiiy 3I-IE iiaiooai?aia ?IE aeyaey?oueny c
??ciith /anoioith a ie?aieo eiiia?oiaioao aaiaoioeeo?a aeiaaeeiaiai
c??co. An? aaiaoioeeoe ?ioaeoii? ia/?iee i?noyoue ca?ia n??aea iaae
aac’yaea?oeaaith /anoeiith yae?a, yaea?oeai ? iaae aiaeiieaciaoe/iei
?aoeeoeoiii ? o?eueee a?ey 10% ee?oei — iaae i?oioiiae??yie (?en.4).
*a?ac 30 oa i?ney *AA no??iei c?inoa? neioac ?IE a yae?? ?
i?aeaeuo?oueny e?euee?noue iiaiooai?aii? ?IE a aiaeiieaciaoe/iiio
?aoeeoeoi?. Cia/ii c?inoa? neioac ?IE o i?oioiiae??yo ie?aii acyoiai
aaiaoioeeoa ? ia ii?yaeie c?inoa? e?euee?noue aaiaoioeeo?a, a yeeo
aeyaeythoueny ca?ia iaae i?oioiiae??yie. I?oioiiae??? c iiaiooai?aiith
?IE ?icoaoiaai? o aaciina?aaei?e aeecueeino? aei yae?a. A ?ioa?aae?
0,5-3,0 aiae i?ney *AA caa??aa?oueny i?aeaeuaiee ??aaiue neioaco ?IE o
aac’yaea?oeaa?e /anoei? yae?a ? oiia?eueith?oueny a yaea?oe? ?
i?oioiiae??yo, i?e/iio cia/iith i??ith ca ?aooiie ciaioaiiy /anoee
aaiaoioeeo?a, ye? i?eeiathoue a iueiio o/anoue. I?ney ciaioaiiy
e?eueeino? iiaiooai?aii? ?IE a aiaeiieaciaoe/iiio ?aoeeoeoi?
a?aeaoaa?onuey ?? iia?eueia c?inoaiiy.

I?e a?eueo o?eaae?e i?enooiino? ?cioiio a i?aai?ci? oaae aeyaey?oueny
aeaioaciee oa?aeoa? ci?i — ii/aoeiaa i?aeaeuaiiy ai?noo ?IE o cacia/aieo
a?aeae?eao ee?oei, yea ci?ith?oueny oei/aniaith cao?eieith iiaiooai?aii?
?IE a yae?? ? a?aeiia?aeii iaioei ?? iaaeoiaeaeaiiyi aei oeeoiieacie.
Oeae i?ioean ca?aa?oueny c ia?aaoaeiaith aia?aoo o?aineyoe?? — c
nei?iieio/ei ciaioaiiyi /enea ca’ycaieo c aiaeiieaciaoe/iei ?aoeeoeoiii
?eainii. Ianooiia ca?eueoaiiy e?eueeino? ?eainii oa ?o aii?yaeeoaaiiy
aoei cao?eniaaii, ii/eiath/e c 3-i? aiaeeie i?ney *AA [Ieaoiiia e ae?.,
1980]. Oaeei /eiii, ca?aeii c iaoeie aeaieie ia?ao?aeiee ia??iae
aeeth/a? oaco aaeaioeaiiai iineeaiiy aeni?an?? yaea?iiai ?
i?oioiiae??aeueiiai aaiii?a ? oaco cai?ie ?nioth/i? i?ia?aie
ae?yeueiino? ee?oei ia iiao.

E?iaoe/ia neeaaei?noue yaea?ii? ?IE

Aeey caaaeueii? oa?aeoa?enoeee yaea?ii? ?IE aoa aeei?enoaiee iaoiae
a?a?eaeecaoe?? iaaeeeoeo ?IE ?c ne?aeiaeie e?eueeinoyie iniiaieo
o?aeoe?e AeIE, ye? aoei ?icae?eaii ca /anoioith iiaoi?aiiy
iine?aeiaiinoae. Aeyaeeiny, ui yaea?ia ?IE ?c ia/?iee /a?ac 3 aiae i?ney
*AA a?a?eaeeco?oueny c iaioith /anoeith oi?eaeueieo iine?aeiaiinoae AeIE
(/anoioa iiaoi?aiiy 1-10 ?ac?a), i?ae yaea?ia ?IE ?c ia/?iee /a?ac 3
aiae i?ney EAI ( 7,9% AeIE a a?a?eaeao i?ioe 11,4 % ) (?en.5). Ca?aene
?ic?aoiaaii e?iaoe/io neeaaei?noue ?IE, yea ia?aaea?oueny aeiaaeeiith
(aai ianith) an?o ??ciiaeaeiinoae ?IE, acyoeo ii iaei?e. Aiia noaiiaeoue
3,0o108 oa 4,8o108 ioeeaioeae?a aeey yaea?ii? ?IE ?c ia/?iee uo??a
a?aeiia?aeii i?ney *AA ? EAI. Ca oa?aeoa?ii ci?i io?eiai? ?acoeueoaoe

ca?aathoueny c aeaieie a?a?eaeecaoe?? oi?eaeueieo iine?aeiaiinoae AeIE c
caaaeueiith ee?oeiiith, iie?(A)-yaea?iith ? iiai-ooai?aiith ?IE
(o?aeoe?y 63 SYMBOL 176 \f «Symbol» N) ?c ?aaaia?oth/i? ia/?iee uo??a
[Grady et al., 1979; 1981; I?eia e ae?., 1982], a oaeiae ei?aeththoue c
aeuaiaaaaeaieie aeaieie i?i neioac ? aioo??oiueiee?oeiiee ?iciiae?e
iiaiooai?aii? ?IE. Cieaeaiiy e?iaoe/ii? neeaaeiino? caaaeueii? ?IE oa ??
o?aeoe?e, cao?eiea iiaiooai?aii? ?IE a yae?? ? ciaioaia ??
iaaeoiaeaeaiiy aei aia?aoo o?aineyoe??, ui

ia?a6oaeiao?oueny, na?ae/eoue i?i oei/aniaa /anoeiaa iaiaaeaiiy
aeni?an?? aaiiio ? a?aeiia?aeii aeayeeo ooieoe?e aaiaoioeeo?a, ye?,
eiia??ii, ia ? i??i?eoaoieie ia oeueiio aoai? ?aaaia?aoe?eiiai i?ioeano.
Ia iiaeaeueoeo aoaiao eiaeoth/? a?eie o?aine?eioe ia a?ae??ciythoueny ca
e?iaoe/iith neeaaei?noth a?ae o?aine?eio?a ?c ia/?iee uo??a i?ney EAI,
oi/a aeaui ?io? ca ye?niei iaai?ii [Colbert et al., 1977; Krieg et al.,
1979].

Iine?aeiaiino? AeIE ?c na?aaeiueith ? aenieith /anoioith iiaoi?aiiy
o?aine?eaothoueny i?ney *AA a a?eueo?e i???, i?ae i?ney EAI (3,6% i?ioe
2,6% /a?ac 3 aiae i?ney eiaeii? iia?aoe??). Ie??i a?aeiieo
?eainiiaeueieo, o?ainii?oieo ? iecueeiiieaeoey?ieo ?IE o?aine?eioe oeeo
iine?aeiaiinoae AeIE inoaoi/ii ia ?aeaioeo?eiaai?. Aeaye? o?aine?eioe
iine?aeiaiinoae c aenieith /anoioith iiaoi?aiiy aoiaeyoue aei neeaaeo
aenieiiieaeoey?ieo ?IE, ye, iai?eeeaae, o?aine?eioe A2 iine?aeiaiinoae a
?IE?4502A1 ? ?IE iniiaiiai eiiieaeno a?noinoi?niino? [E?aia?ia, 1989].

Aeai?, ye? aoei io?eiaii ca aeiiiiiaith iaoiae?a
aeaeo?iii-i?e?ineii?/ii? aaoi?aae?ia?ao?? ? a?a?eaeecaoe?? AeIE-?IE,
na?ae/aoue i?i oa, ui a ia?ao?aeiee ia??iae ?aaaia?aoe?eiiai i?ioeano
a?aeaoaathoueny ci?ie ia ??cieo ??aiyo aeni?an?? aaiiio. Oiio iinoa?
caeiiii??ia ieoaiiy i?i iaoai?cie ? iane?aeee oeeo ci?i.

Aeni?an?y o?aine?eioe?eieo oaeoi??a c-Fos, c-Myc, ssOA-1 SYMBOL 97 \f
«Symbol» ? aenieiniaoe?ae?ciaaiiai a?eea ?4502A1

Aeey ?ica’ycaiiy ieoaiiy i?i iieaeoey?i? iaoai?cie aeyaeaieo ci?i
aeine?aeaeaiiy cae?enithaaee a aeae?eueeio iai?yieao. Cie?aia, aea/aee
aeni?an?th o?aine?eioe?eieo oaeoi??a c-Fos, c-Myc, ssOA-1 SYMBOL 97 \f
«Symbol» ? aenieiniaoe?ae?ciaaiiai a?eea ?4502A1. Aaie c-fos i c-myc
iaeaaeaoue aei aai?a ieoo?ai? a?aeiia?ae?, a oaeiae aei aai?a, ye? nai?
?aaoeththoue naith aeni?an?th, oiaoi ciaoiaeyoueny ia iaeaeu?i uaae?
???a?o?? ?aaoeyoi?ieo a?ee?a. O?aine?eioe?eiee oaeoi? ssOA-1 SYMBOL 97
\f «Symbol» ?aaoeth? /eneaii? oeaieiiniaoeeo?/i? aaie, cie?aia, aai
?4502A1. Aeni?an?y aaio ?4502A1 oiniaeth? iaeneiaeueii niaoe?ae?ciaaiee
oaiioei ia/?iee. I?e iaeiaioeo iciaeao aeaaeeoa?aioe?thaaiiy oeaieie
aeni?an?y oeueiai aaio i?eieiy?oueny iaeia?oith [Padgham et al., 1993].

?en. 6. A?a?eaeecaoe?eiee neaiae iie?(A)+ oeeoiieaciaoe/ii? ?IE
?4502A1.

A?aeiiniee ai?no an?o aeine?aeaeoaaieo ?iaeea?aeoaeueieo ?IE o neeaae?
iie?(A)+ oeeoiieaciaoe/ii? ?IE ci?ith?oueny i?ae aieeaii *AA ? EAI. Ca
oa?aeoa?ii ci?i iaeii/anii aeyaeythoueny noiae?noue ? ??cieoey i?ae
aeaiia i?ioeanaie. *ano?oa e?euee?nii a?eueo ae?aaeaieie ? ci?ie a
?aaaia?oth/?e ia/?ioe?. I?ioyaii ia?ao?aeiiai ia??iaeo ?aaaia?aoe?eiiai
i?ioeano niinoa??aa?oueny ieoo?aa (0 — 0,5 aiae i?ney *AA) i?aeaeuaiiy
ai?noo an?o aeine?aeaeoaaieo o?aine?eio?a o neeaae? iie?(A)+ oeeoi-
ieaciaoe/ii? ?IE ? ianooiia ?aaeoe?y ( 0,5 — 3 aiae i?ney *AA ),
oa?aeoa?ia aeey o?aine?eio?a eiaeiiai oeio (?en. 6 — 8).

Aoaii?noue a aeni?an?? aeine?aeaeoaaieo aai?a a?aeiia?aea?
ia??iaee/iino?, yeo niinoa??aaee a neioac? ? ?iciiae?e? iiaiooai?aii?
?IE. I?aeaeuaia aeni?an?y ??cieo ca oa?aeoa?ii ae?? o?aine?eioe?eieo
oaeoi??a i?ioyaii ieoo?ai? ?aaeoe?? ?ice?eaa? iaoai?ci, ui i?ecaiaeeoue
aei i?aeaeuaiiy aeni?an?? aaiiio (aeea. aeua), cie?aia naiiai
o?aine?eioe?eiiai oaeoi?o ssOA-1 SYMBOL 97 \f «Symbol» [Flodby et al.,
1993] ? aaio ?4502A1 ye iaeiiai c eiai i?oaiae.

?en. 8. Ci?ie a eioeaio?aoe?? iie? (A)+ oeeoiieacia-oe/ii? ?IE ssOA-1
SYMBOL 97 \f «Symbol» a ia/?i-oe? uo??a i?ney *AA ( SYMBOL 209 \f
«Symbol» ) ? EAI ( SYMBOL 168 \f «Symbol» ) ii a?aeii-oaiith aei ??
eiioeaio?aoe?? a ?ioaeoi?e ia/?ioe?.

A?eueo ae?aaeaiee i?e??no ?IE, ui eiaeothoue oeaieiiniaoeeo?/i? a?eee
ssOA-1 SYMBOL 97 \f «Symbol» ? ?4502A1, i?ae ?IE, ye? eiaeothoue
ee?oeiiianiaoeeo?/i? a?eee c-Fos i c-Myc, i?ioyaii ii/aoeiai? ?aaeoe??
na?ae/eoue i?i ?? ni?yiiaai?noue ia i?aeo?eieo oeaieiiniaoeeo?/ieo
ooieoe?e ia/?iee. Oe?eeii eiia??ii, ui a ?aaaia?oth/?e ia/?ioe? /a?ac 1
aiae i?ney *AA ?aae?co?oueny caeaoi?noue oeeoio?iio ?4502A1 eaoae?coaaoe
ooai?aiiy eaoii?a [Koop, 1996], ye? iiaeooue aeei?enoiaoaaoeny o no??iei
c?inoath/iio aetheiiaiaaiac? [Taub, 1996]. Oei/aniaa aeoeaaoe?y
aeni?an?? aaio c-fos ei?aeth? c oei/aniaith (a?ey 1 aiae i?ney *AA)
aeoeai?noth o?aine?eioe?eiiai oaeoi?o c-Fos/c-Jun [Taub, 1996].

Aeuacacia/ai? ci?ie ai?noo iie?(A)+ oeeoiieaciaoe/ieo ?IE c-fos, c-jun
i ssOA-1 SYMBOL 97 \f «Symbol» a?aeaoaathoueny ia oii? iaci?iii? ?o
eiioeaio?aoe?? a yaea?i?e ?IE. Oea na?ae/eoue i?i nooo?ao ?ieue
iinoo?aine?eioe?eieo iiae?e a aeni?an?? oeeo aai?a ia ii/aoeiaiio aoai?
?aaaia?aoe?eiiai i?ioeano. Iaaiaee, iaeii/ania i?aeaeuaiiy ai?noo
yaea?ii? ? iie?(A)+ oeeoiieaciaoe/ii? ?IE ?4502A1 aeaco? ia aeoeaaoe?th
o?aine?eioe?? oeueiai aaio.

Aeoeai?noue nenoaie 2’5′-ie?ai(A)neioaoaca-?IEaca L

Noiniaii iaoai?cio, ui coiiaeth? i?ioyaii ia?ao?aeiiai ia??iaeo
oei/aniaa /anoeiaa iaiaaeaiiy aeni?an?? aaiiio ia ??ai? ?IE, aoei
aenoiooi i?eiouaiiy, ui aiii a?aeaoaa?oueny caaaeyee aeoeaaoe?? nenoaie

2’5′-ie?ai(A)neioaca-?IEaca L. Ae?enii, ?icae?eueia aeine?aeaeaiiy
2’5′-ie?ai(A)neioaoacii? aeoeaiino? yaea?ii? ? oeeoiieaciaoe/ii?
o?aeoe?e iieacaei, a?aeiia?aeii, ciaioaiiy ? i?aeaeuaiiy aiceiaoe/ii?
aeoeaiino? ? e?eueeino? o?eioeeaioeae?a, ui neioacothoueny (oaae. 1).

Oaaeeoey 1. 2’5′-ie?ai(A)neioaoacia aeoeai?noue (2’5′-IAN) ? iiey?i?
ni?aa?aeiioaiiy aee- ? o?eia??a (2A/3A) a ia/?ioe? ?ioaeoieo uo??a ?
uo??a i?ney *AA.

I?ei?oea: 2’5′-ie?ai(A)neioaoacio aeoeai?noue o?aeoe?e iiaeaii a
i?e?iiaeeieoeyo aiceiaoe/ii? aeoeaiino? ia 1 ia noia?iiai a?eea
o?aeoe??.

Ine?eueee ?aeeiith a?aeiiith i?oaiith aeey ie?ai(A)ioeeaioeae?a ? aicei
?IEaca L [Floyd-Smith et al., 1981], oi oae? ci?ie iiaeii? i?ecaiaeeoe
aei oiia?eueiaiiy ?icuaieaiiy (aai i?ioean?iao) yaea?ii? ?IE ?, iaaiaee,
aei i?enei?aiiai ?icuaieaiiy ?IE a oeeoiieaci? ? aei aeenioe?aoe??
iie?nii aiane?aeie o?aaiaioaoe?? ?o i?IE [Oiion e ae?., 1988] (aeey
ii??aiyiiy: aeuacacia/aia ciaioaiiy e?eueeino? iiaiooai?aii? ?IE a
oeeoiieaci? ? ia?a?iciiae?e ?eainii ia aiaeiieaciaoe/iiio ?aoeeoeoi?).
Ca?eueoaiiy eiioeaio?aoe?? ?IEace L, ui aoei aecia/aii ca e?euee?noth
niaoeeo?/ii ca’ycaiiai ?aae?iaeoeaii i?/aiiai ie?aiaaeai?eaoo,
i?aeoaa?aeaeo? eiia??i?noue ooieoe?iioaaiiy cacia/aiiai iaoai?cio.
A?eueoa oiai, o aiiiaaiao? ia/?iee /a?ac 1-3 aiae i?ney *AA aoei
aeyaeaii niaoeeo?/io aeey ?ioa?oa?iio SYMBOL 97 \f «Symbol» / SYMBOL 98
\f «Symbol» aioea??onio aeoeai?noue. ?ioa?oa?ii SYMBOL 97 \f
«Symbol» / SYMBOL 98 \f «Symbol» a?aeiiee ye ?aeeiee ?iaeoeoi? nenoaie
2’5′-ie?ai(A)neioace-?IEace L [Sen, Lengyel, 1992]. Ioaea, i?eiouaiiy
uiaei iiaeeeai? o/ano? nenoaie

2’5′-ie?ai(A)neioaoace-?IEace L a cai?i? oaiioeio ?ioaeoii? ia/?iee ia
oaiioei ?aaaia?oth/i? ia/?iee ia ??ai? ?IE io?eiaei nooo?aa
i?aeoaa?aeaeaiiy.

Oaeoi? iae?ico iooeei SYMBOL 97 \f «Symbol» (OII- SYMBOL 97 \f
«Symbol» ) ye ?aaoeyoi? ia?ai?ia?aioaaiiy ae?yeueiino? ee?oei ia/?iee
aiane?aeie *AA

Iacaaaeath/e ia oa, ui aaaaoi /iai a?aeiii i?i a?aeiiaethaaeueiee
i?ioean a ia/?ioe?, caeeoa?oueny iac’yniaaiei ieoaiiy i?i i?e?iaeo
ioneiaeo iaoai?ci?a. Aeni?an?y niaoeeo?/ieo aeey aaiaoioeeo?a i?oiaai?a
(oaeoi?a ?inoo aaiaoioeeo?a, O?A; ai?aea?iaeueiiai oaeoi?a ?inoo, AO?;
o?ainoi?ioth/iai oaeoi?a ?inoo, OO? SYMBOL 97 \f «Symbol» oiui) oa ?o
?aoeaioi??a cia/ii i?aeaeuo?oueny i?ney *AA [Fausto et al., 1995;
Michalopoulos, DeFrances, 1997]. Uii?aaaea, oaea i?aeaeuaiiy
niinoa??aa?oueny eeoa ia ii/aoeo ia?oiai ee?oeiiiai oeeeeo aaiaoioeeo?a
(3-4 aiae i?ney *AA), ? oiio oe? i?oiaaie ia iiaeooue aaaaeaoeny
ioneiaeie. O iao?e ?iaio? aia?oa aoei iinoaaeaii ieoaiiy i?i ioneiao
ooieoe?th i?iaeoeo?a ee?oei Eoioa?a. Oaea ?aeay a?oiooaaeany ia
caaaeueiiio aeey aoea??ioe/ieo i?aai?ci?a iieiaeaii? i?i ?aaoethth/o
?ieue ee?oei iacaio?iiiai iioiaeaeaiiy a i?ie?oa?aoe?? ee?oei
ai?oae?aeueiiai iioiaeaeaiiy, ia caeaoiino? aeoeaiaaieo ? i?eai?/aieo
ee?oei Eoioa?a a?aeiia?aeii i?enei?thaaoe aai aaeueioaaoe neioac AeIE a
aaiaoioeeoao ?aaaia?oth/i? ia/?iee [Iayineee e ae?., 1977], ia
i?ii?oiaaiieo aeanoeainoyo ii a?aeiioaiith aei aaiaoioeeo?a iniiaiiai
oeeoie?io ee?oei Eoioa?a — OII- SYMBOL 97 \f «Symbol» [Beyer et al.,
1990]. Aia?oa o ia/?ioe? oaa?ei, yeei ia aaiaeeee ?iaeoeoi?e neioaco
OII- SYMBOL 97 \f «Symbol» , aoei aeyaeaii aeni?an?th oeueiai oeeoie?io
c iaeneiaeueiei ?? i?iyaii /a?ac 1 aiae i?ney *AA. Ii/eiath/e c 3-i?
aiaeeie i?ney iia?aoe?? niinoa??aaee oaeiae i?aeaeuaio aeni?an?th ?IE,
ui eiaeothoue iaeaeaa ?aoeaioi?e aei OII- SYMBOL 97 \f «Symbol» , ?OII1
? ?OII2 (?en.9). Oea na?ae/eoue i?i i?aeaeuaio ni?eeiyoeea?noue ee?oei
ia/?iee aei ae?? OII- SYMBOL 97 \f «Symbol» . Aeey a?eueo oi/ii? ioe?iee
e?eueeino? iieaeoe OII-niaoeeo?/ieo ?IE o ?aaaia?oth/?e ia/?ioe?
?aaeoe?th cai?ioiuei? o?ane?eioe?? i?iaiaeeee o i?enooiino? ciai?oiueiai
noaiaea?oo, oiaoi ?IE, yeo aoei io?eiaii in vitro a ?aaeoe??
o?aine?eioe?? iiaeeo?eiaaii? ae?eyiee aaio OII- SYMBOL 97 \f «Symbol» a
neeaae? ieaci?aee ?SPT18. E?euee?noue OII-niaoeeo?/ieo iieaeoe ?IE a 1
iea ee?oeiii? ?IE noaiiaeea (1,6 — 7,2) o 104 aeey ia/?iee /a?ac 1 aiae
i?ney *AA ? (5,6 — 6,0) o 104 oa (0,6 — 3,6) o 103 a?aeiia?aeii aeey
ia/?iee noeioeueiaaieo e?iiiie?naoa?eaeii ? ?ioaeoieo uo??a. A?eueo
aeaoaeueia aeine?aeaeaiiy aeni?an?? OII- SYMBOL 97 \f «Symbol» oa ??
?ie? a i?ioean? a?aeiiaeaiiy ia/?iee i?aeoaa?aeeei iao ?acoeueoao, a
oaeiae aeiaaei, ui OII- SYMBOL 97 \f «Symbol» i?iyaey? iai?yio
i?oiaaiio ae?th ia aaiaoioeeoe [Yamada et al., 1996]. Oey ae?y
iiina?aaeeiao?oueny e?iacaie, o?aine?eioe?eieie oaeoi?aie
c-Fos/c-Jun, yaea?iei oaeoi?ii SYMBOL 107 \f «Symbol» A, a oaeiae
ia?aaeaaa/ai neaiaeo oa

aeoeaaoi?ii o?aine?eioe?? ? ?ioa?eaee?iii-6 [Rai et al., 1996] ?
aeyaey?oueny ?ai?oa ca ae?th ?ioeo i?oiaai?a, ui aieeaathoue ia
aaiaoioeeoe. E??i i?ii?oiaaiii? ae?? ia aaiaoioeeoe, OII- SYMBOL 97 \f
«Symbol» aaeueio? neioac AeIE a aiaeioae?aeueieo ee?oeiao neioni?ae?a
[Sato et al., 1986], ?iaeoeo? i?noeaao ? nenoaiio a?aeiia?aeue c aieo
ia/?iee ia o?aaio [Aaumann, Gauldie, 1994] oa eiio?ieth? ?io?
a?io?i?/i? i?ioeane.

?en. 9. ?aae?iaaoia?ao aiie?o?eiaaieo ?iaeea?aeoaeueieo eAeIE.

* eAeIE ?OII, 55 eAe, ?c ia/?iee /a?ac 3 aiae i?ney *AA aoei
aiie?o?-eiaaii a 20-oe oeeeeao, a c ia/?iee ?ioeo no?ie?a ?aaaia?aoe??
— o 32-o oeeeeao. o.i.i. — oeny/? ia? iniia

Iieaeoey?ii-a?ieia?/i? i?ioeane oa /aniaa oa?aeoa?enoeea
ia?ao?aeiiai ia??iaeo

Ia i?aenoaa? aeaieo, io?eiaieo a iao?e ?iaio? oa a ?iaioao ?ioeo
aeine?aeiee?a, i?iiiio?oueny ?ic??ciyoe aea? oace o ia?ao?aeiiio
ia??iae? ?aaaia?aoe?eiiai i?ioeano — oaco ia?aeiii? eiiiainaoi?ii?
?aaeoe?? (0-0,5 aiae i?ney *AA) ? oaco ia?ai?ia?aioaaiiy ae?yeueiino?
ee?oei (0,5-3,0 aiae i?ney *AA). I?ioyaii ia?oi? oace aeei?enoiaothoueny
ia?aaaaeii ?nioth/? aei iia?aoe?? ?ano?ne i?aaio, cie?aia, aia?aaoe/i?
oa a?eee, aeoeai?noue yeeo iiaeeo?eo?oueny [Taub, 1996]. Ci?ie
iaia?aiiiai iioaioe?aeo, ?iiieo iioie?a, aioo??oiueiee?oeiiiai ?I ?
ieenethaaeueii-a?aeiiaethaaeueiiai iioaioe?aeo aei cia/ii? i??e
iaoiiaeththoue a?io?i?/i? i?ioeane i?ioyaii ia?oi? oace [Koch et al.,
1990]. I?ae /an ae?oai? oace aeni?an?y aai?a caeaaeeoue a?ae a?eeiaiai
neioaco de novo ? ?aaoeth?oueny oaeeie neaiaeueieie iieaeoeaie, ye
oeeoie?ie, ai?iiie oiui [Koch et al., 1990]. Iaeii/anii c
aeoeaaoe??th aeni?an?? iaeieo aai?a (ssOA-1 SYMBOL 97 \f «Symbol» ,

?4502A1, OII- SYMBOL 97 \f «Symbol» , ?OII1, ?OII2 oiui) a?aeaoaa?oueny
oei/aniaa iaiaaeaiiy aeni?an?? ?ioeo. Inoaii? aaaaea?oueny i?iyaii
aeoeaii? cai?ie ?nioth/iai oaiioeio, a nenoaia
2’5′-ie?ai(A)neioaoaca-?IEaca L — ?? eiia??iei /eiieeii ia ??ai? ?IE.

Ee?oeiiee oeeee aaiaoioeeo?a ? ?aaeoe?y aino?i? oace

Iaei??th ?c neeaaeiaeo ?aaaia?aoe?eiiai i?ioeano ? ?aaeoe?y aino?i?
oace c aieo ia/?iee. Oey aenieiniaoeeo?/ia nenoaiia ?aaeoe?y ia/?iee ia
o?aaio aoaeue-yeei /eiieeii ? aoaeue-yei? eieae?caoe?? iieyaa? a
nae?aoe?? niaoeeo?/ieo a?ee?a, ye? aaeueiothoue ?oei?ai? ? aeoeaothoue
caaithaaeuei? i?ioeane a i?noe? ooeiaeaeaiiy. Ia /ania?e oeae?
?aaaia?aoe?eiiai i?ioeano ?aaeoe?y aino?i? oace ca?aa?oueny c ia?oei
ee?oeiiei oeeeeii aaiaoioeeo?a, a iaeneiaeueiee ?? i?iya — c
iaeneiaeueiith AeIE-neioaoe/iith aeoeai?noth [Sobczak et al., 1989].
Naii ii nia? oea noi?uaiiy aeaco? ia iaaecae/aeio ni?iiiaei?noue
aaiaoioeeo?a ii?aeioaaoe oeaieiiniaoeeo?/i? oa i?ie?oa?aoeai? ooieoe?? ?
a oie aea /an aeeeeea? ieoaiiy, ye oe? ooieoe?? ?aaoeththoueny. Aeey
a?aeiia?ae? ia oea ieoaiiy i?iaaaeaii ii??aiyeueia aeine?aeaeaiiy
aeni?an?? aai?a c-fos, c-myc, ssOA-1 SYMBOL 97 \f «Symbol» ? ?4502A1 o
cacia/aiiio i?ii?aeeo /ano i?ney *AA ? EAI. A inoaiiueiio aeiaaeeo
a?aeiia?aeue ia/?iee ia o?aaio, ni?e/eiaio ?icoeiii /a?aaii? ii?iaeieie,
iinoa? ye naiino?eia ?aaeoe?y. Aeyaeaii, ui i?ney *AA ? i?ney EAI a
ia/?ioe? c?inoa? ai?no aeine?aeaeaieo o?aine?eio?a, ye? ciaoiaeyoueny a
neeaae? iie?(A)+ oeeoiieaciaoe/ii? ?IE (?en. 7). Iaeiae, ia a?aei?io
a?ae iiae?e ia ii/aoeo ?aaaia?aoe?eiiai i?ioeano, i?e??no e?eueeino?
oeaieiiniaoeeo?/ieo o?aine?eio?a ssOA-1 SYMBOL 97 \f «Symbol» ? ?4502A1
iaeaea aei??aith? i?e?inoo i?ney EAI, oi/a i?ney *AA ooieoe?iio? o?eueee
1/3 ia/?iee, a iio?aaa a a?eeao aino?i? oace i?ney *AA aeaea?oueny ia
iaioith, a a?eueoith. Noiae? ci?ie niinoa??aathoueny c o?aine?eioaie
aaio c-fos. E?euee?nii ? ca /anii i?iyao aiie nooo?ai ia
a?ae??ciythoueny i?ney iaio iia?aoe?e. Ia oe?e i?aenoaa? aeni?an?y
i?ioiiieiaaio c-fos a ?ioa?aae? 15-30 aiae i?ney *AA aaaaea?oueny
iaiao?aeiith a ia?oo /a?ao aeey a?aeiia?ae? ia/?iee ia o?aaio. Oeae
aeniiaie a?aeiia?aea? oiio, ui eiai aoei c?iaeaii ia i?aenoaa?
aeine?aeaeaiue in vitro [Karin et al., 1997], ? ni?inoiao?
?iciianthaeaeaio aeoieo i?i aeeeth/io ?ieue i?ioiiieiaaio c-fos a
ee?oeiiiio oeeee?. I?ioeeaaeiee aei aeuaiaaaaeaiiai oa?aeoa? ci?i
niinoa??aa?oueny c iie?(A)+ oeeoiieaciaoe/ieie o?aine?eioaie aaio c-myc.
I?e??no ?o ai?noo iaeaea a o?e ?ace ia?aaeuo? oie, ui aeyaey?oueny i?ney
EAI, ? ai?no o?aine?eio?a ci?ith?oueny neio?iiii c oacaie ee?oeiiiai
oeeeeo. Oaeee ?acoeueoao in vivo io?eiaii aia?oa. A?i i?aeoaa?aeaeo?
aeai?, ui ?o aoei io?eiaii in vitro, ? ia?aaeaa/a? ?aaoeyoi?io ?ieue
oaeoi?o c-Myc o i?ioiaeaeaii? ee?oeiiiai oeeeeo [Seth et al., 1993].

Ieen acioo (NO) — eiia??iee iaei?ino?oee aioo??oiuei- ee?oeiiee ?
i?aeee?oeiiee ?aaoeyoi? ?aaaia?aoe?eiiai i?ioeano

Aaeeeo oaaao i?e aeine?aeaeaii? oaeoi??a, ye? ?aaoeththoue
a?aeiiaethaaeueiee i?ioean a ia/?ioe?, i?eaa?oa? ieen acioo. Oea
iaei?ino?oa ei?ioeiaeeao/a e, i/aaeaeii, aaiethoe?eii aeaaiy niieoea,
yea aca?iiae?? c? nai?ie i?oaiyie aac o/ano? aicei?a. Aeaye? ?aaeoe??
ieeno acioo iiaeooue aooe cai?ioieie, ui ? nooo?aith aeiiaith aeey
ooieoe?iioaaiiy ?aaoeyoi?iiai oaeoi?a. I?oaiyie aeey ieeno acioo
aenooiathoue o?iee a?ee?a, aoiie/?iie cae?ca oa ?ioeo ia?ao?aeieo
iaoae?a, iieaeoee eenith oa noia?ieneae, oe?iceiiaee ?aaeeeae, oiui
[Marletta et al., 1988]. Ieen acioo iiaea aooe ye aioo??oiueiee?oeiiei
?aaoeyoi?ii, oae ? oaeoi?ii i?aeee?oeiiiai ni?eeoaaiiy.

Ca aeiiiiiaith ae?th/i? in vivo «ianoee» aeey NO aoei aia?oa aecia/aii
e?euee?noue ieeno acioo, ui ooai?th?oueny i?ioyaii 30 oa a ??ci?
ia??iaee ?aaaia?aoe?eiiai i?ioeano. C’yniaaii, ui neioac ieeno acioo
ci?ith?oueny i?ney *AA. Iaeneiaeueia c?inoaiiy niinoa??aa?oueny /a?ac 1
? 6 aiae, a oaeiae a ?ioa?aae? 25 — 36 aiae i?ney *AA (?en. 10). Ae?oaee
? o?ao?e ia??iaee c?inoaiiy ca?aathoueny c G1- i G2-ia??iaeaie ia?oiai
ee?oeiiiai oeeeeo aaiaoioeeo?a. ?o ?icae?ey? ia??iae i?i?iaeueii?
i?iaeoeoe?? NO, ui i?eiaaea? ia S-oaco aaiaoioeeo?a.

*an i?ney *AA (aiae)

?en. 10. Neioac ieeno acioo ? AeIE a ?aaaia?oth/?e ia/?ioe? uo??a.

Aeey c’ynoaaiiy oiai, ye? ee?oeie ?aaaia?oth/i? ia/?iee i?iaeoeothoue
ieen acioo aoei i?iaaaeaii aeine?aeaeaiiy c ?cieueiaaieie a ??ci? /ane
i?ney *AA aaiaoioeeoaie, ee?oeiaie Eoioa?a ? aiaeioae?aeueieie ee?oeiaie
neioni?ae?a. Aeyaeeiny, ui e?euee?noue neioaciaaiiai ieeno acioo o
caaaeaieo ee?oeiao ciaioo?oueny a ii?yaeeo ?o iaaaaeaiiy, i?e/iio a on?o
ee?oeiao neioac a?aeaoaa?oueny i?ioyaii G1- ? G2-oac ? niaaea? i?ioyaii
S-oace ee?oeiiiai oeeeeo (?en.11). Iiee ui iaa?aeiii, ye? c iiaeeeaeo
i?oaiae ieeno acioo caae?yi? a ?aaoeyoe?? ee?oeiiiai oeeeeo. Aeineoue
ae?ia?aeii, ui NO ca?aeii c? nai?th aeanoea?noth aca?iiae?yoe c
eeth/iaei aiceiii neioaco AeIE — ?eaiioeeaioeae?aaeoeoacith [Moncada et
al., 1991], i?eai?/o? ?? aeoeai?noue i?ioyaii G1- ? G2-oac ? «aeicaiey?»
a?aeaoaaoeny neioaco AeIE ca oiia, eiee neioac naiiai NO a?aenooi?e.
Ci?ie aeoeaiino? aai eiioeaio?aoe?? aaaaoueio niieoe a ?aaaia?oth/?e
ia/?ioe? iiaeia iiynieoe ia??iaee/iith ae??th NO. Cie?aia, ciaioaiiy
eiioeaio?aoe?? eiiieaenii ca’ycaiiai iaaaiiaiai cae?ca a yae?ao ee?oei
?aaaia?oth/i? ia/?iee (19,0 SYMBOL 177 \f «Symbol» 2 ia% i?ioe 27
SYMBOL 177 \f «Symbol» 2 ia% a ?ioaeoiiio i?aai?) iiaea aooe iane?aeeii
niaoeeo?/ii? ae?? NO ? nooo?aei aeey ee?oeiiiai oeeeeo.

*an i?ney *AA (aiae)

?en. 11. Neioac ieeno acioo ?cieueiaaieie ee?oeiaie aiaeioae?th
neioni?ae?a, ee?oeiaie Eoioa?a ? aaiaoioeeoaie.

Eii?aeeiaoe?y ooieoe?e ee?oei ia/?iee ca aeiiiiiaith neaiaeueieo
iieaeoe

Iaie aoa i?iaaaeaiee aiae?c aeanieo ? e?oa?aoo?ieo aeaieo c iaoith
anoaiiaeoe, ye ci?ith?oueny iaa?? neaiaeueieo iieaeoe i?ioyaii
a?aeiiaethaaeueiiai i?ioeano ? ye oe? ci?ie iia’ycai? c
iieaeoey?ii-a?ieia?/ieie i?ioeanaie, ui a?aeaoaathoueny a ?aaaia?oth/?e
ia/?ioe?. Iaea?eueoa ??ciiiai?oi?noue neaiaeueieo iieaeoe aeyaey?oueny
ia ?aii?o aoaiao i?ioeano — aei ii/aoeo ? i?ioyaii ia?oiai ee?oeiiiai
oeeeeo aaiaoioeeo?a (oaae. 2).

Oaaeeoey 2. Aeni?an?y neaiaeueieo iieaeoe a /ania?e oeae?
?aaaia?aoe?eiiai i?ioeano.

Ia?ao?aeiee I?ie?oa?aoeaia aeoeai?noue ee?oei ?aaaia?oth/i? ia/?iee

ia??iae Aaiaoioeeoe Ee?oeie Aiaeioa?aeuei? Eoioa?a
ee?oeie

0 — 3* 3 — 12 15 — 24 24 — 30 30 — 72 62 — 120

Oen-A2 — — — — —

IA-F2 SYMBOL 97 \f «Symbol» IA-F2 SYMBOL 97 \f «Symbol» ** — —
— —

IA-A2 IA-A2 IA-A2** IA-A2** IA-A2 —

OII- SYMBOL 97 \f «Symbol» ** OII- SYMBOL 97 \f «Symbol» — —
— —

?OI- SYMBOL 97 \f «Symbol» / SYMBOL 98 \f «Symbol» ** ?OI- SYMBOL 97 \f
«Symbol» / SYMBOL 98 \f «Symbol» ** — — — —

O?A O?A O?A** O?A O?A —

?E-1 SYMBOL 97 \f «Symbol» ** ?E-1 SYMBOL 97 \f «Symbol» ?E-1 SYMBOL
97 \f «Symbol» — — —

?E-1 SYMBOL 98 \f «Symbol» ** ?E-1 SYMBOL 98 \f «Symbol» ?E-1 SYMBOL
98 \f «Symbol» — — —

— ?E-6** ?E-6** — — —

OO?- SYMBOL 98 \f «Symbol» 1 OO?- SYMBOL 98 \f «Symbol» 1 OO?- SYMBOL 98
\f «Symbol» 1 OO?- SYMBOL 98 \f «Symbol» 1 OO?- SYMBOL 98 \f «Symbol»
1** —

O?O O?O O?O O?O O?O O?O**

NOaai NOaai — NOaai NOEeE NOAEeN

AO? AO?** — — — —

OO?- SYMBOL 97 \f «Symbol» 1 OO?- SYMBOL 97 \f «Symbol» 1 OO?- SYMBOL 97
\f «Symbol» 1** OO?- SYMBOL 97 \f «Symbol» 1 — —

I?ei?oea: * *an i?ney *AA (aiae). ** Iaeneiaeueia aeni?an?y neaiaeueieo
iieaeoe. Nei?i/aiiy: aai — aaiaoioeeoe, EeE — ee?oeie Eoioa?a, AEeN —
aiaeioae?aeuei? ee?oeie neioni?ae?a, Oen — o?iiaienai, IA —
i?inoaaeaiaeei. Eo?neaii aeae?eai? aeai?, io?eiai? a iao?e ?iaio?.

?en. 12. Aeeoa?aioe?eia ?aaoeyoe?y i?ie?oa?aoeaii? aeoeaiino?
aaiaoioeeo?a.

?en. 13. Aeeoa?aioe?eia ?aaoeyoe?y i?ie?oa?aoeaii? aeoeaiino? ee?oei
Eoioa?a.

*enaeuei? neaiaeuei? iieaeoee i?iaeoeothoueny iaia?aio?iieie ee?oeiaie
? ca?aeii c? nai?ie aeanoeainoyie ?aaoeththoue ??ciiiai?oi? a?io?i?/i?
i?ioeane. ?aii?e ae?eiaaiie?c, caoaeaiiy noaeei, aeoeaaoe?y e?ia’yieo
aeyoie ei?aeththoue c aeoeai?noth o?iiaienaio A2; ianooiia ?icoe?aiiy
noaeei, i?aeaeuaiiy ??aiy oeeee?/iiai AIO, i?iaeoeoe?y e ae?y
ai?aea?iaeueiiai oaeoi?a ?inoo — c ae??th i?inoaaeaiae?io A2; ieoo?aa
?aaeoe?y ia o?aaio — c ae??th OII- SYMBOL 97 \f «Symbol» oa
?ioa?eaee?io 1 (?E-1), oiui. Iniiaieie i?oiaaiaie aeey aaiaoioeeo?a ?
OII- SYMBOL 97 \f «Symbol» , oaeoi? ?inoo aaiaoioeeo?a oa
ai?aea?iaeueiee oaeoi? ?inoo, eii?oiaaiii — i?inoaaeaiaeei A2
[Michalopoulos, DeFrances, 1997]. I?ae /an i?ie?oa?aoe?? aaiaoioeeo?a
aeaye? neaiaeuei? iieaeoee (OII- SYMBOL 97 \f «Symbol» , i?inoaaeaiaeei
A2, ?E-1) i?eai?/othoue i?ie?oa?aoe?th iaia?aio?iieo ee?oei [Hortelano
et al., 1995; Holloway et al., 1997] (?en.12). Aiaeia?/ia neooaoe?y
niinoa??aa?oueny c ?ioeie ee?oeiaie ia/?iee. I?ie?oa?aoe?y ee?oei
Eoioa?a oa aiaeioae?aeueieo ee?oei neioni?ae?a ni?aiaaea? c oei/aniaith
aeoeai?noth ?o niaoeeo?/ieo i?ii?oiaai?a, o?ainoi?ioth/iai oaeoi?a ?inoo
SYMBOL 98 \f «Symbol» [Holloway et al., 1997] ? oaeoi?a ?inoo
aiaeioae?th noaeei, a?aeiia?aeii [Yamane et al., 1994], ye?
i?iaeoeothoueny ia?eneioni?aeaeueieie c??/anoeie ee?oeiaie ?
aaiaoioeeoaie (?en. 13, 14). Iaeii/anii c aeoeaaoe??th neioaco AeIE a
ee?oeiao Eoioa?a o?ainoi?ioth/ee oaeoi? ?inoo SYMBOL 98 \f «Symbol»
i?eai?/o? neioac AeIE a aaiaoioeeoao oa aiaeioae?aeueieo ee?oeiao
neioni?ae?a [Holloway et al., 1997; Fausto et al., 1995]. E??i oiai, a?i
aeoeao? neioac NO a ee?oeiao Eoioa?a ?, oaeei /eiii, ni?ey? cieaeaiith
?o ee?oeiiniaoeeo?/ii? aeoeaiino? [Stadler et al., 1995] (?en. 13).

OII- SYMBOL 97 \f «Symbol»

OO?- SYMBOL 98 \f «Symbol» AEeN ?OAN

IA-A2

?en. 14. Oaeoi?e, ui ?aaoeththoue i?ie?oa?aoe?th aiaeioae?aeueieo ee?oei
neioni?ae?a. ?OAN — ?inoiaee oaeoi? aiaeioae?th
noaeei

Oaeei /eiii, i?ioyaii ?aaaia?aoe?eiiai i?ioeano ee?oeii-niaoeeo?/ia
aeoeai?noue iaeieo ee?oei iaiao?aeia aeey caaacia/aiiy i?ie?oa?aoeaii?
aeoeaiino? ?ioeo. A iaaeao ee?oei iaeiiai oeio i?ie?oa?aoeaia
aeoeai?noue noi?iaiaeaeo?oueny cieaeaiiyi ?o ee?oeiiniaoeeo?/ii?
aeoeaiino?. Ii?yae c oi?aa?naeueieie aeey an?o ee?oei ?aaoeyoi?aie
i?ie?oa?aoe?? (oeeee?ie, ieen acioo oiui) a ia/?ioe? ooieoe?iiothoue
ee?oeiiniaoeeo?/i? aeoeaaoi?e oa ?ia?a?oi?e. ?o aca?ii?aaoeyoe?y a
cia/i?e i??? coiiaeth? ia?aa?a ?aaaia?aoe?eiiai i?ioeano a ia/?ioe?
uo??a.

Cae/aeii, oey i?aoey ia aea? a?aeiia?ae? ia an? ieoaiiy, ui noiyoue
ia?aae aeine?aeieeii a?aeiiaethaaeueieo i?ioean?a o
aenieiaeeoa?aioe?ei-aaieo oeaieiao, iaeiae, aea? iia? oyaeaiiy i?i
/aniao oeaeo ?aaaia?aoe?? oa i?i iieaeoey?ii-a?ieia?/i? i?ioeane ia
eiaeiiio ?c ?? aoai?a, aeicaiey? cai?iiiioaaoe eiioeaioe?th i?i
aca?iiae?th ee?oei ??cieo oei?a ? i?eioeeie ci?ie i?ia?ai ae?yeueiino?
ee?oei ?, ia?aoo?, aeaco? ia iaiao?aei?noue i?iaaaeaiiy iiaeaeueoeo
aeine?aeaeaiue.

AENIIAEE

1. Aeine?aeaeaiiy ia?aiao??a ee?oeiiiai oeeeeo, aeni?an?? aaiiio oa
aeoeaiino? aeayeeo ?aaoeyoi?ieo oaeoi??a a ia/?ioe? uo??a i?ney *AA
aeicaieeee cai?iiiioaaoe: /aniao oeaeo i?ioeano; a?iioaco, ui iiynith?
iaoai?ci ia?aoiaeo aenieiaeeoa?aioe?eiaaieo ? iaaeoeaieo a
i?ie?oa?aoeaiiio a?aeiioaii? ee?oei ia/?iee aei i?ie?oa?aoe??;
iieiaeaiiy i?i i?ia?aeio ?ieue iaia?aio?iieo ee?oei o ia?aoiae?
aaiaoioeeo?a aei i?ie?oa?aoe??; eiioeaioe?th i?i ni?aa?aeiioaiiy
i?ie?oa?aoeaii? ? ee?oeiiniaoeeo?/ii? aeoeaiino? ee?oei ia/?iee i?ioyaii
a?aeiiaethaaeueiiai i?ioeano.

2. Anoaiiaeaii, ui a ?aaaia?oth/?e ia/?ioe? uo??a aaiaoioeeoe, ee?oeie
Eoioa?a oa aiaeioae?aeuei? ee?oeie neioni?ae?a anooiathoue o ee?oeiiee
oeeee a ii?yaeeo ?o iaaaaeaiiy. Cai?iiiiiaaii aeei?enoiaoaaoe ia?aiao?e
ee?oeiieo oeeee?a ee?oei ??cieo oei?a ye /ania? i???ioe?e
?aaaia?aoe?eiiai i?ioeano, a ia??iae i?ae iia?aoe??th ? ii/aoeii
ee?oeiiiai oeeeeo aaiaoioeeo?a ?icaeyaeaoe ye ia?ao?aeiee a?ae
i?ie?oa?aoeaii iaaeoeaiiai aei i?ie?oa?aoeaii aeoeaiiai noaio oeaieie.

3. Cai?iiiiiaaii ?ic??ciyoe aeaa aoaie o ia?ao?aeiiio ia??iae?
?aaaia?aoe?eiiai i?ioeano: aoai ia?aeiii? eiiiainaoi?ii? ?aaeoe??
(0-0,5aiae i?ney *AA) oa aoai ia?ai?ia?aioaaiiy ae?yeueiino? ee?oei
(0,5-3 aiae i?ney *AA). Aeey ia?oiai oa?aeoa?i? i?aeaeuaiiy neioaco
oioaeueii? ?IE ? ?? o?aeoe?e, i?enei?aiiy ia?aoiaeo iiaiooai?aii? ?IE c
yae?a a oeeoiieacio, i?aeaeuaiiy aeni?an?? oeaieiiniaoeeo?/ieo
(ssOA-1 SYMBOL 97 \f «Symbol» ,

?4502A1) ? ianiaoeeo?/ieo a?ee?a (c-myc, c-fos). Aeey ae?oaiai aoaio
oa?aeoa?i? cieaeaiiy neioaco ? iaaeoiaeaeaiiy iiaiooai?aii? ?IE a
oeeoiieacio, ciaioaiiy ??ciiiai?oiino? yaea?ii? ?IE ? cao?eiea ?? a
yae??.

4. Aia?oa aeyaeaii aeoeaaoe?th nenoaie 2’5′-ie?ai(A)neioaoaca-?IEaca L
oa ?? niaoeeo?/iiai ?iaeoeoi?a ?ioa?oa?iio SYMBOL 97 \f «Symbol» /
SYMBOL 98 \f «Symbol» i?ioyaii ia?ao?aeiiai ia??iaeo ?aaaia?aoe?eiiai
i?ioeano. Cai?iiiiiaaii a?iioaco, ca yeith aeoeaia aea??eiaa aeaeaeaiiy
«noa?eo» ?IE, cie?aia ca aeiiiiiaith aeuacacia/aii? nenoaie,
aaaaea?oueny iaiao?aeiei aoaiii aeey ia?aoiaeo aeeiaeeoa?aioe?eiaaii?
oeaieie a?ae noaio niieith aei i?ie?oa?aoe??.

5. Aia?oa o ?aaaia?oth/?e ia/?ioe? aecia/aii aeni?an?th aai?a, ye?
eiaeothoue oaeoi? iae?ico iooeei ? eiai aeaa ?aoeaioi?e, oa aeyaeaii
cia/ia i?aeaeuaiiy ?o aeni?an?? i?ioyaii ia?ao?aeiiai ia??iaeo
?aaaia?aoe??;

6. Iaa?oioiaaii iieiaeaiiy i?i i?ia?aeio ?ieue iaia?aio?iieo ee?oei
ia/?iee a caioneo ?aaaia?aoe?eiiai i?ioeano, ui caaacia/o?oueny
aeoeaaoe??th ?o ee?oeiiniaoeeo?/ieo ooieoe?e i?ioyaii ia?ao?aeiiai
ia??iaeo ? ia?oiai ee?oeiiiai oeeeeo aaiaoioeeo?a.

7. Aia?oa aecia/aiee neioac ieeno acioo ? eiai ?ioaineai?noue o
?aaaia?oth/?e ia/?ioe? uo??a. E?euee?noue NO, yea i?iaeoeothoue
aaiaoioeeoe, ee?oeie Eoioa?a oa aiaeioae?aeuei? ee?oeie neioni?ae?a,
ciaioo?oueny a ii?yaeeo iaaaaeaiiy oeeo ee?oei. Neioac NO a?aeaoaa?oueny
i?ioyaii G1- ? G2-oac ? niaaea? a S-oac? ee?oeiiiai oeeeeo.

8. Aia?oa ia iiaeae? ?aaaia?oth/i? ia/?iee in vivo i?iaeaiiino?iaaii
aca?iica’ycie i?ae aei?an??th aaio c-myc ? oacaie ee?oeiiiai oeeeeo
aaiaoioeeo?a. ??aaiue aeni?an?? aaio c-myc i?aeaeuaiee i?ioyaii G1- ?
G2-oac ? cieaeaiee a S-oac? ee?oeiiiai oeeeeo .

9. Aia?oa c’yniaaii, ui aeni?an?y aai?a n-fos, ssOA-1 SYMBOL 97 \f
«Symbol» ? ?4502A1 a i?ii?aeeo /ano, eiee iaeii/anii a?aeaoaathoueny
i?ie?oa?aoe?y aaiaoioeeo?a ? a?aeiia?aeue ia/?iee ia o?aaio, iia’ycaia c
a?aeiia?aeaeth ia/?iee ia o?aaio.

10. Aeoiaey/e c aeaieo, io?eiaieo o oe?e ?iaio?, cai?iiiiiaaii
eiioeaioe?th i?i neia?a?ci ee?oeiiniaoeeo?/ii? aeoeaiino? iaeieo ee?oei
ia/?iee ? i?ie?oa?aoeaii? aeoeaiino? ?ioeo. Iieacaia iiaea?eia ooieoe?y
neaiaeueieo iieaeoe, ye? ?aaoeththoue i?ie?oa?aoe?th ee?oei a
?aaaia?oth/?e ia/?ioe?, — aeaye? c ieo (i?inoaaeaiae?i A2, OII- SYMBOL
97 \f «Symbol» , OO?- SYMBOL 98 \f «Symbol» ) iiaeooue iaeii/anii
aeeiioaaoe ?ieue ee?oeiiniaoeeo?/ieo aeoeaaoi??a ? ee?oeiiniaoeeo?/ieo
?ia?a?oi??a i?ie?oa?aoe??.

NIENIE I?AOeUe IIOAE?EIAAIEO CA OAIITH AeENA?OAOe??

Iiiia?ao?y

1. ?aaoeyoeey aeineioaca aaeea o yoea?eio/ Aeueneay A.A.,
Noa?iaeoa I.O., Iioaiia A.I., Eiaaeaiei I.E., Ia/a?aiei A.E., Iaieaineay
I.TH., Eaaiia E.A./ Iiae ?aae. A.A. Aeueneie.-E.: Iaoeiaa Aeoiea, 1990.
— 278 n.

Noaoo? o oaoiaeo iaoeiaeo aeaeaiiyo

2. Iaieaineay I.TH. Aic?anoiua iniaaiiinoe neioaca AeIE a
?aaaia?e?othuae ia/aie aaeuo e?un.// Oeeoieiaey. — 1976. — O.18, ? 7. —
N. 857 — 861.

3. Iaieaineay I.TH. Aaeee a ?aaoeyoeee yeni?annee aaiia ia/aie. //
Aeiiieeia?u e eeaoea. — 1990. — O.6, ? 5. — N. 106 — 108.

4. Obolenskaya M.Yu. The potential targets

of Kupffer Cells activity during liver regeneration.// Aeiiieeia?u e
eeaoea. — 1997. — O. 13, ? 2. — N. 168-172.

5. Obolenskaya M.Yu. Cytokines and liver regeneration.// EOS — J.
Immunology and Immunopharmacology. — 1997. — V.17, ? 2. — ?. 51 -58.

6. Iaieaineay I.TH. Neaiaeueiua iieaeoeu e ?aaaia?aoeey ia/aie. //
Aeiiieeia?u e eeaoea. — 1998. — O. 14, ? 3. — N. 210 — 222.

7. Iaieainueea I.TH., Aa?aneiiaa A.A. ?ieue eiiieaenii ca»ycaiiai cae?ca
a iiae?e? ee?oeie.// Oe?. a?io?i. aeo?iae. — 1972. — O. 44,? 5. — N.
577-579.

8. Aa?aneiiaa O.A., Iaieaineay I.TH., Ieaoiiia I.I.
Aaoi?aaeeia?aoe/aneia eco/aiea neioaca e auoiaea a oeeoiieacio ?IE ia
?aiieo noaaeeyo ?aaaia?aoeee ia/aie.// Oeeoieiaey e aaiaoeea. — 1980. —
O.14, ? 3. — N. 77 — 82.

9. Iaieaineay I.TH., I?eia A.E., Ieaoiiia I.I. Aea?eaeecaoeey
ioeeaeiiauo eeneio yoea?eio a i?enoonoaee aunieeo eiioeaio?aoeee
oi?iaieaea.// Studia biophysica. — 1984 — V. 98, ? 2. — ?. 103 — 110.

10. Iaieaineay I.TH., I?eia A.E., Eoeeeianeay E. A., Ieaoiiia I.I.
ssaea?iua ?IE ia ?aiiai yoaia ?aaaia?aoeee ia/aie.// Aeiiieeia?u e
eeaoea. -1987. — O. 3, ? 1. — N. 27 -35.

11. Iaieaineay I.TH,. Aa?aneiiaa O.A., Aeee/ E.E., Ieaoiiia I.I.
Aioo?eeeaoi/iia ?ani?aaeaeaiea iiaiia?aciaaiiie ?IE ia ?aiiai yoaia
?aaaia?aoeee ia/aie.// Oeeoieiaey e aaiaoeea. — 1987. — O. 21,? 5. —
N.376 — 382.

12. Iaieaineay I.TH., I?eia A.E., Aa?aneiiaa O.A., Ieaoiiia I.I.
*anoe/iia ia?aie/aiea yeni?annee aaiiia — eiiiiiaio ia?ano?ieee

aai ?aaiou a ?aaaia?e?othuae ia/aie ieaeiieoathueo.// Aeiiieeia?u e
eeaoea. — 1989. — O. 5, ? 2. — N. 79 — 88.

13. I?eia A.E., Eei/ei O.A., Iaieaineay I.TH., Eaaiia I.E., ?uneia
A.I., Ieaoiiia I.I. Neio?iiiua eciaiaiey eiioeaio?aoeee c-fos ?IE e
A2i?IEo a i?a?aieeeaoeaiii ia?eiaea ?aaaia?aoeee ia/aie.// Aeiiieeia?u e
eeaoea. — 1989. — O. 5, ? 6. — N. 83 -86.

14. Iaieaineay I.TH., I?eia A.E., Eei/ei O.A., Iaoa? E.A., Oieieaiei
E.I., Ieaoiiia I.I. Oeeeeicaaeneiay yeni?anney aaia, eiaee?othuaai
A2i?IEo.// Iieaeoey?iay aeieiaey, ae?onieiaey e iee?iaeieiaey. — 1991.-
? 6. — N. 31 — 33.

15. Obolenskaya M.Yu., Bernauer H., Tran — Thi T.-A., Decker K. Levels
of RNA for

TNF and TNF receptors during the prereplicative period of liver
regeneration.// Aeiiieeia?u e eeaoea. — 1994. — O.10, ? 4. — N.
152 — 156.

16. Obolenskaya M.Yu., Vanin A.F., Mordvintcev P.I., Mulsch A., Decker
K. EPR

evidence of nitric oxide production by the regenerating rat liver.//
Biochem. Biophys. Res. Comm. — 1994. — V. 202, ? 1. — ?. 571 — 576.

17. Obolenskaya M., Schulze-Specking A., Plaumann B., Frenzer K.,
Freudenberg N. and K.

Decker. Nitric oxide production by cells isolated from regenerating rat
liver. // Biochem.

Biophys. Res. Comm. — 1994. — V. 204, ? 3. — ?. 1305 -1311.

18. Iaoa? E.A., O?eioiia A.A., Eaeoieoeeay A.A., I?eia A.E., Iaieaineay
I.TH., Ieaoiiia I. I. Yeni?anney i?ioiiiieiaaiia c-myc e c-fos ai a?aiy
ia?aiai eeaoi/iiai oeeeea a ?aaaia?e?othuae ia/aie e?un.// Aeiiieeia?u e
eeaoea. — 1994. — O. 10, ? 2. — N. 57 -60.

19. Decker K., Obolenskaya M. Cytokines, nitric oxide synthesis and
liver regeneration.//

J. Gastroenterology and Hepatology. — 1995. — ? 10. — S12-S17.

20. Iaieaineay I.TH., Aaiei A.O., Ii?aeaeioeaa I.E., Aeaea? E. Ieenue
acioa a ?aaaia?e?othuae ia/aie e?un. // Aeiiieeia?u e eeaoea. — 1995. —
O. 11, ? 2. — N. 76 — 82.

21. Iaieaineay I.TH., Eyeueaa I., Naciiiaa E.ss. 2″5″ieeai(A)
neioaoaciay aeoeaiinoue ia ?aiieo yoaiao ?aaaia?aoeeiiiiai i?ioeanna a
ia/aie e?un. // Aeiiieeia?u e eeaoea. — 1997. — O.13, ? 5. — N. 391 —
396.

Noaoo? a iaoeiaeo ca??ieeao

22. Iaieaineay I.TH., Ieaoiiia I.I., A?aaei A. Iaeioi?ua iaoiaee/aneea
iniaaiiinoe iae?iiieaeoey?iie aea?eaeecaoeee ?IE e AeIE.// Iaoiaeu
iieaeoey?iie aeieiaee. — E.: Iaoeiaa Aeoiea, 1979. — N. 45 — 55.

23. Iaieaineay I.TH., I?eia A.E., Eoeeeianeay E.A., Nioaiue Y.I.,
Ei?ithoeia O.A., Ieaoiiia I.I. Aea?eaeecaoeey ioeeaeiiauo eeneio
yoea?eioia a ?anoai?ao oi?iaieaea n aunieie eiioeaio?aoeeae.//
Iieaeoey?iay aeieiaey. — E.: Iaoe. Aeoiea, 1982. — O. 31. — N. 27 — 32.

24.Iaieaineay I.TH. Yeni?anney aaiiia a ?aaaia?e?othuae ia/aie
ieaeiieoathueo.// Nia?aiaiiua i?iaeaiu ?aaaia?aoeee. Oa?iaeiei-ae/aneay
?aaoeyoeey ?aaaia?aoeee. — Eioea?-Iea. — 1987. — N. 139 -152.

Iaoa??aee iaoeiaeo eiioa?aioe?e ? neiiic?oi?a

25. Obolenskaya M., H. Bernauer, K. Decker. Do Kupffer cells take part
in triggering of

liver regeneration? // Proc. of the Sixth Intern. Symposium on Cells of
the Hepatic Sinusoid. Antwerpen (Belgium) 1992. — Cells of the hepatic
sinusoid (eds. D.L.Knook, E.Wisse). — AkLeiden, Netherlands, 1993. — V.
4. — P. 76 — 78

26. Obolenskaya M.Yu., Decker K. Liver regeneration: interplay between
two systems of

intercellular communication, cytokines and nitric oxide.// Proc. of the
Seventh Intern. Symposium on Cells of the Hepatic Sinusoid. Kyito
(Japan) 1994.- Cells of the hepatic sinusoid

(eds. D.L.Knook, E.Wisse, E.Wake). — AkLeiden, Netherlands, 1995. — V.5.
— ?. 339 — 342.

27. Obolenskaya M., L. Pater, L. Sazonova, A. Elskaya and K. Decker.
Kupffer Cells and Liver

Regeneration.// Proc. of the Eighth Intern. Symposium on Cells of the
Hepatic Sinusoid. Bordeaux (France) 1996. — Cells of the Hepatic
Sinusoid (eds. E.Wisse, D.L.Knook, C.

Balabaud).- AkLeiden, Netherlands, 1997. — V.6. — P. 340 — 343.

28. Obolenskaya M., H. Bernauer, K. Decker. Do Kupffer cells take part
in triggering of

liver regeneration?// Abstracts of the Sixth International Symposium on
Cells of the hepatic sinusoid. — Antwerpen, Belgium, 23-30 August,
1992. — P. 134.

29. Obolenskaya M.Yu., Pater L.V., Decker K. Expression of
transcriptional factors c-Fos, c-Myc and HNF-1 in the liver after
partial hepatectomy and laparatomy.// Abstracts of the International
Falk Workshop «Normal and malignant liver cell growth». — Halle
(Germany). — 1998. — P. 24.

AIIOAOe??

Iaieainueea I.TH. ?aaaia?aoe?y ia/?iee o uo??a: iieaeoey?ii-a?ieia?/i?
i?ioeane, ?o ?aaoeyoe?y oa /aniaa oeaea. — ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy aeieoi?a a?ieia?/ieo iaoe
ca niaoe?aeuei?noth 03.00.03 — iieaeoey?ia a?ieia?y. — ?inoeooo
iieaeoey?ii? a?ieia?? ? aaiaoeee IAI Oe?a?ie, Ee?a, 1999.

Aeine?aeaeaii neioac AeIE, aioo??oiueiee?oeiiee ?iciiae?e ? e?euee?noue
iiaiooai?aii? ?IE, e?iaoe/io neeaaei?noue yaea?ii? ?IE, aeni?an?th
ie?aieo aai?a (c-fos, c-myc, P4502E1, ssOA-1 SYMBOL 97 \f «Symbol» ,
oaeoi?o iae?ico iooeei ? eiai ?aoeaioi??a), aeoeai?noue nenoaie
2’5′-ie?ai(A)neioaoace-?IEace L ? neioac ieeno acioo a ?aaaia?oth/?e
ia/?ioe? uo??a. Cai?iiiiiaaii /aniao oeaeo i?ioeano. Noi?ioeueiaaii
iieiaeaiiy i?i /anoeiaa iaiaaeaiiy aeni?an?? aaiiio a ia?ao?aeiee
ia??iae ?aaaia?aoe??, i?i neia?a?ci i?ae i?ie?oa?aoeaiith ?
ee?oeiiiniaoeeo?/iith aeoeai?noth ee?oei ia/?iee ??ciiai oeio ? i?i
aca?iia iaiaaeaiiy ee?oeiiniaoeeo?/ii? ? i?ie?oa?aoeaii? aeoeaiino? a
ee?oeiao ia/?iee iaeiiai oeio.

Eeth/ia? neiaa: ?aaaia?aoe?y ia/?iee; iaia?aio?ii? ee?oeie; oaeoi?
iae?ico iooeei; 2’5′-ie?ai(A)neioaoaca-?IEaca L; ieen acioo; n-fos;
c-myc; ssOA-1 SYMBOL 97 \f «Symbol» ; ?4502A1.

Obolenskaya M.Yu. Rat liver regeneration: iolecular processes, their
regulation and the time scale. — Manuscript.

Thesis for a doctor’s degree by speciality 03.00.03 — molecular
biology. — The Institute of Molecular Biology and Genetics of National
Academy of Sciences of Ukraine, Kyiv, 1999.

This thesis is focused on DNA synthesis, amount and intracellular
distribution of newly formed RNA, kinetic complexity of nuclear RNA,
gene expression (c-fos, c-myc, P4502E1, HNF-1 SYMBOL 97 \f «Symbol» ,
tumor necrosis factor and its rceptors), activity of
2’5′-oligo(A)synthetase-RNAse L system and nitric oxide production in
regenerating rat liver. The time-scale of the process is proposed. The
following statements are formulated: about the partial restriction of
genome expression during the transition period, about synergism of
proliferative and cell-specific activities between the cells of
different type and about mutual restriction of proliferative and
cell-specific activities in the cells of the same type.

Key words: liver regeneration; nonparenchymal cells; tumor necrosis
factor; 2’5′-oligo(A)synthetase-RNAse L; nitric oxide; c-fos; c-myc;
HNF-1 SYMBOL 97 \f «Symbol» ; P4502E1.

Iaieaineay I.TH. ?aaaia?aoeey ia/aie o e?un: iieaeoey?ii-aeieiae/aneea
i?ioeannu, eo ?aaoeyoeey e a?aiaiiay oeaea. — ?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. Aeenna?oaoeey
iinayuaia enneaaeiaaieth iieaeoey?iuo iaoaieciia, eioi?ua iaania/eaatho
ia?aoiae aunieiniaoeeaeece?iaaiiuo e iaaeoeaiuo a i?ieeoa?aoeaiii
ioiioaiee eeaoie ia/aie e i?ieeoa?aoeee e eii?aeeiaoeeth eo
eeaoi/iiniaoeeoe/aneie e i?ieeoa?aoeaie aeoeaiinoe a ?aaaia?e?othuae
ia/aie e?un. ?aaaia?aoeeth ia/aie e?un iiaeaee?iaaee iooai oaeaeaiey 2/3
i?aaia. Yoa iia?aoeey aucuaaao neiaeioth ioaaoioth ?aaeoeeth, eioi?ay
aeeth/aao a ea/anoaa ninoaaiuo yeaiaioia ?aaeoeeth ino?ie oacu ni
noi?iiu ia/aie, eiiiainaoi?iua ?aaeoeee iiaeaea?aeaiey i?ei?eoaoiuo
ooieoeee i?aaia, nianoaaiii i?ieeoa?aoeeth eeaoie e niaio oaiioeia
“iieiyueony” eeaoie ia oaiioei aeaeyueony eeaoie. ?aaeoeey ino?ie oacu
ni noi?iiu ia/aie, eioi?ay i?iyaeyoany a ioaao ia iia?aaeaeaiea
i?aaiecia ethaie yoeieiaee e ethaie eieaeecaoeee e iai?aaeaia ia
ia?aie/aiea aeano?oeoeaiuo e aeoeaaoeeth ?aia?aoeaiuo i?ioeannia,
iiaeaee?iaaeanue n iiiiuueth eaia?ioiiee (EAI). Ia?aeeaeueiia
enneaaeiaaiea iaeieo e oao aea iieacaoaeae ai a?aiy ?aaeoeee ino?ie oacu
ni noi?iiu ia/aie e a i?ioeanna ?aaaia?aoeee aeaei aiciiaeiinoue ioianoe
iaeioi?ua ec iaia?oaeaiiuo eciaiaiee e ii?aaeaeaiiui iaoioeceieiae/aneei
i?ioeannai. Iniiaiia aieiaiea a aeenna?oaoeee auei oaeaeaii
enneaaeiaaieth yeni?annee aaiiia e ioaeaeueiuo aaiia, eioi?ua eiaee?otho
eeaoi/iiniaoeeoe/aneea e ianiaoeeoe/aneea aeey ia?aioeiiuo e
iaia?aioeiiuo eeaoie ia/aie ?aaoeyoi?iua oaeoi?u. Ii?aaeaeaiu eiee/anoai
e aioo?eeeaoi/iia ?ani?aaeaeaiea iiaiia?aciaaiiie ?IE, eeiaoe/aneay
neiaeiinoue yaea?iie ?IE, yeni?anney eeaoi/iiniaoeeoe/aneeo (P4502E1,
ssOA-1 SYMBOL 97 \f «Symbol» , oaeoi? iae?ica iiooieae) e
ianiaoeeoe/aneeo (c-fos, c-myc, ?aoeaioi?u oaeoi?a iae?ica iiooieae)
aaeeia, aeoeaiinoue nenoaiu 2’5′-ieeai(A)neioaoacu — ?IEacu L e neioac
ieene acioa a ia/aie e?un iinea *AA e EAI. Aeey nicaeaiey /aniaie oeaeu
?aaaia?aoeeiiiiai i?ioeanna enneaaeiaaeenue iieacaoaee i?ieeoa?aoeaiie
aeoeaiinoe eeaoie ia/aie. Onoaiiaeaii, /oi a ?aaaia?e?othuae ia/aie e?un
aaiaoioeeou, eeaoee Eoioa?a e yiaeioaeeaeueiua eeaoee neionieaeia
anooiatho a eeaoi/iue oeeee a i?eaaaeaiiie iineaaeiaaoaeueiinoe.
I?aaeeiaeaii eniieueciaaoue ia?aiao?u eeaoi/iuo oeeeeia eeaoie ia/aie
?aciuo oeiia a ea/anoaa a?aiaiiuo i?eaioe?ia ?aaaia?aoeeiiiiai
i?ioeanna, a ia?eiae iaaeaeo iia?aoeeae e ia/aeii eeaoi/iiai oeeeea
aaiaoioeeoia ?anniao?eaaoue eae ia?aoiaeiue io i?ieeoa?aoeaii
iaaeoeaiiai ninoiyiey oeaie e i?ieeoa?aoeaii aeoeaiiio. I?aaeeiaeaii
?acee/aoue aeaa yoaia a oa/aiea ia?aoiaeiiai ia?eiaea ?aaaia?aoeeiiiiai
i?ioeanna: yoai ia/aeueiie eiiiainaoi?iie ?aaeoeee (0-0,5 / iinea *AY) e
yoai ia?ai?ia?aiie?iaaiey aeayoaeueiinoe eeaoie (0,5-3 / iinea *AY).
Ia?aue oa?aeoa?ecoaony iiauoaieai neioaca oioaeueiie ?IE e aa o?aeoeee,
onei?aieai ia?aoiaea iiaiia?aciaaiiie ?IE ec yae?a a oeeoiieacio,
iiauoaieai yeni?annee eeaoi/iiniaoeeoe/aneeo (ssOA-1, ?4502A1) e
eeaoi/iiianiaoeeoe/aneeo (c-myc, c-fos) aaeeia. Aeey aoi?iai yoaia
oa?aeoa?iu nieaeaiea neioaca e iinooieaiey iiaiia?aciaaiiie ?IE a
oeeoiieacio, oiaiueoaiea ?aciiia?acey yaea?iie ?IE e caaea?aeea aa a
yae?a. Aia?aua iaia?oaeaia aeoeaaoeey nenoaiu 2″5″ieeai(A)neioacau —
?IEacu L e aa niaoeeoe/aneiai eiaeoeoi?a eioa?oa?iia SYMBOL 97 \f
«Symbol» / SYMBOL 98 \f «Symbol» a oa/aiea ia?aoiaeiiai ia?eiaea
?aaaia?aoeeiiiiai i?ioeanna. I?aaeeiaeaia aeiioaca, niaeanii eioi?ie
aeoeaiia ecae?aoaeueiia oaeaeaiea «noa?uo» ?IE, a /anoiinoe, n iiiiuueth
auoaoeacaiiie nenoaiu, ?anniao?eaaaony eae iaiaoiaeeiue yoai i?e
ia?aoiaea aunieiaeeooa?aoee?iaaiiie oeaie io iieiy e i?iee-oa?aoeee.
Aia?aua a ?aaaia?e?othuae ia/aie iaia?oaeaia yeni?anney aaiia,
eiaee?othueo oaeoi? iae?ica iiooieae SYMBOL 97 \f «Symbol» e aai aeaa
?aoeaioi?a, e ii?aaeaeaii cia/eoaeueiia iiauoaiea eo yeni?annee a
oa/aiea ia?aoiaeiiai ia?eiaea ?aaaia?aoeee. Iainiiaaii iieiaeaiea i
aaaeouae ?iee iaia?aioeiiuo eeaoie ia/aie a caionea ?aaaia?aoeeiiiiai
i?ioeanna, /oi iaania/eaaaony aeoeaaoeeae eo eeaoi/iiniaoeeoe/aneeo
ooieoeee a oa/aiea ia?aoiaeiiai ia?eiaea e ia?aiai eeaoi/iiai oeeeea
aaiaoioeeoia. Aia?aua ioa?aeoa?eciaai neioac ieene acioa a
?aaaia?e?othuae ia/aie e?un. Eiee/anoai NO, eioi?ia i?iaeooee?otho
aaiaoioeeou, eeaoee Eoioa?a e yiaeioaeeaeueiua eeaoee neionieaeia,
oiaiueoaaony a i?eaaaeaiiii ?yaeo eeaoie e ei??aee?oao n oacaie eo
eeaoi/iuo oeeeeia — iiauoaii a G1- e G2- e nieaeaii a S-oaca. Aia?aua ia
iiaeaee ?aaaia?e?othuae ia/aie in vivo i?iaeaiiino?e?iaaia acaeiinaycue
iaaeaeo yeni?anneae aaia c-myc e oacaie eeaoi/iiai oeeeea aaiaoioeeoia —
iiauoaiea a oa/aiea G1- e G2- e nieaeaiea a S-oaca, e iaaeaeo
yeni?anneae aaiia n-fos, ssOA-1 SYMBOL 97 \f «Symbol» e ?4502A1 e
ioaaoii ia/aie ia o?aaio. Ia iniiaaiee aiaeeca iieo/aiiuo e eiathueony
aeaiiuo i?aaeeiaeaia eiioeaioeey i neia?aecia eeaoi/iiniaoeeoe/aneie
aeoeaiinoe iaeieo eeaoie ?aaaia?e?othuae ia/aie e i?ieeoa?aoeaiie
aeoeaiinoe ae?oaeo e i acaeiiia?aie/aiee yoeo aeoeaiinoae a eeaoeao
iaeiiai oeia.

Eeth/aaua neiaa: ?aaaia?aoeey ia/aie; iaia?aioeiiua eeaoee; oaeoi?
iae?ica iiooieae; 2’5′-ieeai(A)neioaoaca-?IEaca L; ieenue acioa; c-fos;
c-myc; ssOA-1 SYMBOL 97 \f «Symbol» ; P4502E1.

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