Ee?anueeee oi?aa?neoao ?iai? Oa?ana Oaa/aiea

?na?ieia Noai?neaa Aaeaioeiiae/

OAeE 577.346

Aieea iiooaeiino? aeice ia aeo?ae iioeiaeaeaiue

AeIE a ee?oeiao ?ineei

03.00.01 — ?aae?ia?ieia?y

Aaoi?aoa?ao aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa
a?ieia?/ieo iaoe

Ee?a — 1999

Aeena?oaoe??th ? ?oeiien

?iaioa aeeiiaia a ?inoeooo? ee?oeiii? a?ieia?? oa aaiaoe/ii? ?iaeaia???
IAI Oe?a?ie

Iaoeiaee ea??aiee — aeaaeai?e IAI Oe?a?ie, aeieoi? a?ieia?/ieo iaoe,
i?ioani?

A?iaec?inueeee Aeieo?i Ieoaeeiae/

?inoeooo ee?oeiii? a?ieia?? oa aaiaoe/i? ?iaeaia??? IAI Oe?a?ie,

caa?aeoaa/ a?aeae?eo a?io?ceee oa ?aae?ia?ieia??

Io?oe?ei? iiiiaioe: aeieoi? a?ieia?/ieo iaoe, aeioeaio,

Aiae??e/aiei Na?a?e Aaaeeiiae/

?inoeooo ioiea?eiaieia?? ?i. i?ioani?a I.N. Eieii?e/aiea IIC Oe?a?ie,
aieiaiee iaoeiaee ni?a?ia?oiee a?aeae?eo a?io?ceee

eaiaeeaeao a?ieia?/ieo iaoe,

Aacae?iaia Ea?ena Einoyioei?aia

?inoeooo yaea?ieo aeine?aeaeaiue IAI Oe?a?ie, caa?aeoaa/ eaai?aoi???
?aae?ia?ieia??.

I?ia?aeia onoaiiaa — Oe?a?inueeee iaoeiai-aeine?aeiee ?inoeooo
n?euenueeiainiiaea?nueei? ?aae?ieia?? I?i?noa?noaa aa?ii?iieneiaiai
eiiieaeno Oe?a?ie.

Caoeno a?aeaoaeaoueny «17» ethoiai i 14 aiaeei? ia can?aeaii?
niaoe?ae?ciaaii? a/aii? ?aaee Ae 26.001.24 i?e Ee?anueeiio
oi?aa?neoao? Oa?ana Oaa/aiea ca aae?anith: Ee?a, aoe. aeaae. Aeooaeiaa
2, a?ieia?/iee oaeoeueoao, aoae.215. Iiooiaa aae?ana: 01033, Ee?a — 33,
aoe. Aieiaeeie?nueea 64, iaoeiaa /anoeia.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? Ee?anueeiai
oi?aa?neoaoo Oa?ana Oaa/aiea ca aae?anith: Ee?a, aoe. Aieiaeeie?nueea,
58.

Aaoi?aoa?ao ?ic?neaiee «27» a?oaeiy 1999 ?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee
A?aeii I.A.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue i?iaeaie.

I?ney oi?eaeueii? ii nai?e i?e?iae? aaa??? ia *AAN iniaeeai aino?i noaei
ieoaiiy i?i aea/aiiy iniaeeainoae aieeao iaeeo aeic ?ii?coth/i?
?aae?aoe?? ia a?ieia?/i? ia`?eoe. *i?iiaeeuenea eaoano?ioa i?ecaaea aei
nooo?aiai ca?eueoaiiy ?aae?aoe?eiiai oiio, aiane?aeie /iai aaeee? ca
nai?th ieiuath oa?eoi??? iino?eii i?aeiaaeathoue i?ae ae?th ii?ii?iaiiy
iecueei? ?ioaineaiino?. Aiae?c aeaieo e?oa?aoo?e na?ae/eoue i?i nooo?aa
i?enei?aiiy iooaoe?eiiai ? ?aeiia?iaoe?eiiai i?ioean?a, ca?eueoaiiy
aaiaoe/iiai aaioaaeo a iiioeyoe?yo ?ineei ? oaa?ei, ui i?aeiaee aieeao
iecueeiaeiciaiai ii?ii?iaiiy (Oaa/aiei e ae?., 1992; Ao?eaeiaa e ae?.,
1996; Aa?anueeei e ae?., 1998). Iaaeineiiae?noue aeieia?/ieo ii?iaoea?a
? aeineoue ooiaiia oyaeaiiy i?i aieea ia aeinenoaie, ui aaaea aei
aeanoaa?e?caoe?? inoaii?o oa cieaeaiiy a?i??ciiiai?ooy, ?iaeoue
aeooaeueiei aiae?c caeiiii??iinoae oi?ioaaiiy ? ?ic?iaeo e?euee?nieo
iaoiae?a ioe?iee aaiaoe/ieo aoaeo?a, ui ?iaeoeiaai? iecueeeie aeicaie
?ii?coth/iai aei?ii?ithaaiiy.

C`yaey?oueny ana a?eueoa aenia?eiaioaeueieo aeaieo, ui na?ae/aoue i?i
iae?i?eiee oa?aeoa? i?iyao a?ieia?/ieo aoaeo?a a iecueeiaeiciaiio
ae?aiacii? (Oaa/aiei e ae?., 1992; Ao?eaeiaa e ae?., 1996; Aa?anueeei e
ae?., 1998). Ine?eueee caeaaei?noue aeica-aoaeo a ae?aiacii? iaeeo aeic
ia? iae?i?eiee oa?aeoa?, ei?aeoia ioe?iea aaiaoe/iiai ?eceeo a oeueiio
aeiaaeeo iiaea aacoaaoeny o?eueee ia /?oeiio ?icoi?ii?
iieaeoey?ieo-ee?oeiieo iaoai?ci?a oi?ioaaiiy ?aaeoe?? ee?oei ia
iecueeiaeiciaa ii?ii?iaiiy.

Iniaeeai aaaeeeaa i?noea a i?iya? oeeo aoaeo?a a?ae?a?a? oaeoi?
?ioaineaiino? aieeao ia a?iia`?eoe. Aoaeoe iiooaeiino? aeice ii?ii?iaiiy
a ae?aiacii? na?aaei?o ? aaeeeeo aeic aea/ai? aeineoue aeaoaeueii
(Oaooiai 1955; A?iaeceineee 1989). ?nio? aaeeea iecea oaeo?a, yea
na?ae/eoue ia ei?enoue oiai, ui i?iyae ?aae?ia?ieia?/ieo aoaeo?a
ci?iththoueny a?aeiia?aeii ?ioaineaiino? ii?ii?iaiiy. A iaeano? aaeeeeo
oa na?aaei?o aeic ii?ii?iaiiy, ye i?aaeei, aoaeoe e?i?eii caeaaeaoue
a?ae iiooaeiino? ii?ii?iaiiy ? a iaai?e i??? iia`ycai? c ?aia?aoe??th
noaeaoaeueieo iioeiaeaeaiue. I?e aenieeo iiooaeiinoyo ii?ii?iaiiy
neeueii iineaaeth?oueny iiaeeea?noue ?aia?aoe?? oaeiai oeio
iioeiaeaeaiue, a i?e iaeeo — iai?ioe, ? iaaia a??ia?aei?noue ?aia?aoe??
noaeaoaeueieo iioeiaeaeaiue.

sse?nii ?ioa ea?oeia niinoa??aa?oueny aeey aoaeo?a iiooaeiino? aeice a
iaeano? iecueeeo aeic ii?ii?iaiiy. I?e aeine?aeaeaiiyo oeeoiaaiaoe/ieo
aoaeo?a a iaeano? iecueeiaeiciaiai ii?ii?iaiiy aea/ai? caeaaeiino? a?ae
iiooaeiino? aeice, i?e/iio oi?ia caeaaeiino? iiaea ci?ithaaoenue ?acii c
aaee/eiith aeice, ui na?ae/eoue i?i neeaaei?noue oeeo yaeu (Ao?eaeiaa e
ae?., 1996). ?niothoue aenia?eiaioaeuei? oaeoe, ye? aeacothoue ia
cai?ioio caeaaei?noue a?ae iiooaeiino? aeice (Oaa/aiei e ae?., 1992). Ia
aeaeue, aeai? i?i aieea iiooaeiino? aeice a ae?aiacii? iecueeeo aeic
ia/eneaii? ? iinyoue noia?a/eeaee oa?aeoa?. Oiio aea/aiiy
?aae?ia?ieia?/ieo aoaeo?a, iaoiiaeaieo ii?ii?iaiiyi ??cii? ?ioaineaiino?
a iaeano? iaeeo aeic, ia? aaaeeeaa cia/aiiy.

Ca iaoei i?eiouaiiyi i?e/eiith iae?i?eiino? i?iyao a?ieia?/ieo aoaeo?a a
iaeano? iaeeo aeic ii?ii?iaiiy iiaea aenooiaoe ?ic?ea
i?ae aeicaie, ui ni?e/eiythoue aeoeaaoe?th ?iaeooeeaaeueieo nenoai
?aia?aoe??, ? aeicaie, ui ia aeoeaothoue aeiaeaoeia? ?aia?aoeai?
nenoaie, a ?iiino? eiinoeoooeaieo nenoai a?aeiiaeaiiy ia aenoa/a?, uia
iiai?noth ae?i?ioaaoe an? iioeiaeaeaiiy, ni?e/eiai? ii?ii?iaiiyi
iecueei? ?ioaineaiino?. Oaeei /eiii, ia? i?aenoaao i?eiouaiiy, ui aeey
?iaeoeoe?? aeiaeaoeiaeo ?aia?aoeaieo nenoai iio??aia iaaia ii?iaiaa
cia/aiiy iioeiaeaeaiue aaiiio ca iaeeieoeth /ano. sseui ?ioaineai?noue
iaaaioaaeaiiy iaioa ca ii?iaiao aaee/eio, oi iiaeeeaa neooaoe?y, eiee
eiinoeoooeai? nenoaie ?aia?aoe?? aaea ia ni?iiiaei? a?aeiiaeoe an?
iioeiaeaeaiiy, ui aoee ni?e/eiai? ii?ii?iaiiyi, a aeoeaaoe??
aeiaeaoeiaeo nenoai ?aia?aoe?? ua ia a?aeaoaa?oueny.

Iiae?aia aea/aiiy ?aaeoe?e ee?oei ?ineei ia ae?th ?ii?coth/eo
aei?ii?ithaaiue aenooia? ye iaoiae ?ice?eooy caaaeueiia?ieia?/ieo
caeiiii??iinoae. Aaaeeea?noue oa aeooaeuei?noue oaeeo iiooe?a
aacia?a/ia, ine?eueee aeno?aiieyoe?y aoaeo?a c iaeano? na?aaei?o oa
aenieeo aeic a iaeanoue iecueeeo aeic ia cian?i ei?aeoia.

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

Aeena?oaoe?eia ?iaioa aeeiiaia o a?aeae?e? a?io?ceee oa ?aae?ia?ieia??
?inoeoooo ee?oeiii? a?ieia?? oa aaiaoe/ii? ?iaeaia??? ca?aeii c ieaiii
iaoeiai-aeine?aeieo ?ia?o ?inoeoooo ca oaiaie: «Ei?aeoe?y eoioeyoeaieo
aoaeo?a o?ii?/iiai ii?ii?iaiiy ?ineei ?ii?coth/ith ?aae?aoe??th» oa
«O?ii?/ia ii?ii?iaiiy ?ineei ye oaeoi? oi?ioaaiiy ianoaa?eueiino?
aaiiio». ?iaioa aeeiioaaeanue ca i?aeo?eiee a?aioaie I?i?noa?noaa
Oe?a?ie a ni?aaao iaoee ? oaoiieia?e (i?iaeo: 5.4/389 «Aeine?aeeoe
iieaeoey?ii-a?ieia?/io i?e?iaeo i?ioean?a, ui ?iaeoeiaai? o?ii?/iei
ii?ii?iaiiyi ?ineei»)

Iaoa ?iaioe ? caaea/? aeine?aeaeaiiy.

Aieiaia iaoa ?iaioe iieyaaea o aea/aii? ci?i a AeIE ee?oei ?ineei i?ae
aieeaii ?ii?coth/iai ii?ii?iaiiy ??cii? ?ioaineaiino? a iecueeeo aeicao.
A?aeiia?aeii aei oeueiai aoee iinoaaeai? ne?aeoth/e caaaeaiiy:

— aeaoe ii??aiyeueio ioe?ieo aeoiaeo iioeiaeaeaiue AeIE o
?ineeiieo ee?oei, ui aoee ii?ii?iaii? a iecueeiaeiciaiio ae?aiacii? c
??ciith iiooaei?noth, aeey ??cieo ?ineeiieo nenoai (ian?iiy, i?ioiieanoe
oa i?i?inoee) oa ??cieo aeae?a ?ineei (othothi oa ninia);

— aea/eoe caeiiii??iino? oi?ioaaiiy iioeiaeaeaiue AeIE o ian?ii?
ninie, ui iaoiiaeai? o?ii?/iei ii?ii?iaiiyi a iaeeo aeicao;

— aeyaeoe ?aaeeie ii?ii?iaiiy, a yeiio niinoa??aa?oueny i?aeaeuaiee
aeo?ae iioeiaeaeaiue AeIE aeey oeeo nenoai, oa iiynieoe oea yaeua.

Iaoeiaa iiaecia iaea?aeaieo ?acoeueoao?a.

Aia?oa iieacaii, ui aeo?ae iioeiaeaeaiue AeIE aeey oaeeo ?ineeiieo
ia’?eo?a, ye ninia oa othothi, i?e ae?? ii?ii?iaiiy ??cii? ?ioaineaiino?
a iecueeeo aeicao ia? neeaaeio caeaaei?noue, ye a?ae cia/aiiy aeice, oae
? a?ae ?? iiooaeiino?. Iieacaii, ui oaeoi? iiooaeiino? aeice ? aaaeeeaei
/eiieeii o oi?ioaaii? a?aeiia?ae? ?ineeiii? ee?oeie ia iecueeiaeiciaa
ii?ii?iaiiy. Io?eiai? ?acoeueoaoe na?ae/aoue i?i iaaeinoaoith
iaa?oioiaai?noue canoinoaaiiy iaioth/i? a oaia??oi?e /an
e?i?eii- aacii?iaiai? eiioeaioe?? aeey ioe?iee ?eceeo iecueeiaeiciaiai
ii?ii?iaiiy. Anoaiiaeaii oaeo ?aae?caoe?? “i?eoiaaieo” iioeiaeaeaiue
AeIE, ui noi?ioaaeenue o ii?ii?iaiiai ian?iiy. Ciaeaeaii ?aaeei
iecueeiaeiciaiai ii?ii?iaiiy, i?e yeiio niinoa??aa?oueny i?aeaeuaiee
aeo?ae iioeiaeaeaiue AeIE ii??aiyii c i/?eoaaiei.

I?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoao?a.

Anoaiiaeai? oaeoe iiaeooue aooe ei?enieie aeey a?aeiiaeaiiy e?niaiai
ainiiaea?noaa a ?aeiiao, ui iino?aaeaeaee aiane?aeie ?aae?iaeoeaiiai
caa?oaeiaiiy. Aoei anoaiiaeaii, ui ian?iiy ninie cae/aeii?, yea io?eiaei
ca aeaa ?iee naiai oi?ioaaiiy aeico o?ii?/iiai ii?ii?iaiiy 3 A?, ia?
?yae ia?aaaa ii e?eoa??yi aeeoo?caeaoiino? ii??aiyii c eiio?ieueiei oa
a?eueo neeuei? iiaeeeaino? a?aeiiaeaiiy AeIE ca oiiae aieeao oanooth/iai
aino?iai aaiia-aei?ii?ithaaiiy.

Iniaenoee aianie caeiaoaa/a.

Iniiai? ?aea?, iaoeia? iieiaeaiiy ? aeniiaee ?ic?iaeai? aeena?oaioii
naiino?eii. Aeena?oaioii aaciina?aaeiuei i?iaiaeeeinue ii?ii?iaiiy
ia`?eo?a aeine?aeaeaiiy. A?i aoa caae?yiee ia an?o aoaiao aeae?eaiiy
AeIE, a oaeiae aiae?co nooiai? ooeiaeaeaiiino? oe??? a?iiae?iiieaeoee.
Aeena?oaioii naiino?eii i?iaaaeaiee aiae?c AeIE iaoiaeii eoaeiiai
aeaeo?ioi?aco. A i?ioean? aeeiiaiiy aeena?oaoe?? aaoi? iniaenoi
a?ae?a?aa oa e?eoe/ii i?iaiae?coaaa aeinooiio e?oa?aoo?o, yea ia?
a?aeiioaiiy aei oaie aeine?aeaeaiiy.

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??.

Iaoa??aee ?iaioe aoee i?aaenoaaeai? ia III c’?cae? ii ?aae?aoe?eiei
aeine?aeaeaiiyi, Iineaa, 1997; International Conference on Plant
Ontogenesis in Natural and Transformed environments, Lviv, Ukraine, July
1-4, 1998; XIth International Congress on Photosyntesis, Budapest,
Hungary, August 17-22, 1998; International Conference “Diagnosis and
Treatment of Radiation injury”, Rotterdam, the Nethelands, 30 August —
3 September, 1998; Iaoeiaa-i?aeoe/ia eiioa?aioe?y iieiaeeo a/aieo
“?ioaa?aoeai?noue ooieoe?e ?ineeiiiai i?aai?cio” Aa?anaiue 3 — 4, Ee?a,
Oe?a?ia, 1998; II International Symposium on Plant Biotechnology,
October 4-8, Kyiv, Ukraine, 1998.

No?oeoo?a aeena?oaoe??.

Aeena?oaoe?y aeeeaaeaia ia 155 noi??ieao, aeeth/a? 26 ?ethno?aoe?e,
i?noeoue anooi, o?e ?icae?ee, aiae?c oa ocaaaeueiaiiy ?acoeueoao?a,
aeniiaee oa nienie aeei?enoaieo e?oa?aoo?ieo aeaea?ae (138
iaeiaioaaiue).

CI?NO ?IAIOE

?icae?e 1 .Iaeyae e?oa?aoo?e.

Iaoa??ae ?icae?eo a?aeia?aaea? no/aniee noai ciaiue i?i a?ieia?/i?
aoaeoe iia`ycai? c ae??th ?ii?coth/i? ?aae?aoe?? iecueeeo ??ai?a oa
??cieo iiooaeiinoae ii?ii?iaiiy. Ioa?aeoa?eciaai? oeie
?aae?aoe?eii-?iaeoeiaaieo iioeiaeaeaiue AeIE. ?icaeyioo? oeie
?aia?aoe?eieo nenoai ee?oeie.

?icae?e 2. Iaoa??aee ? iaoiaee aeine?aeaeaiue.

Ian?iiy ninie (Pinus silvestris) aoei io?eiaia a?ae ni?a?ia?oiee?a
Noa?iiao??anueei? e?nii? aeine?aeii? noaioe?? (Ee?anueea iaeanoue). Aeey
ie?aieo aenia?eiaio?a aeei?enoiaoaaee ian?iiy ninie c ??cieo ae?eyiie
Ciie a?ae/oaeaiiy *AAN. Ca aeaieie OEAe-aeiceiao???, ui aoea i?iaaaeaia
?ia?oieeaie Noa?iiao??anueei? e?niai? aeine?aeii? noaioe??, iiaeeiaia
?ineeiaie aeica ?ii?coth/iai ii?ii?iaiiy ca 2 ?iee oi?ioaaiiy oeueiai
ian?iiy ii ciai?oiueiio aaiia-ii?ii?iaiith neeaaeaea: 0,03 A? (?aeii
naea Aeeoyoee), 3 A? (?aeii naea Eiia/?) oa 6 A? (?aeii naea
*enoiaae?aea). I?i?iuoaaee eiai o oai?ya? ia caieiaeaiiio
o?eueo?oaaeueiiio iaia?? ia i?ioyc? 10 ae?a i?e oaiia?aoo?? 25oN. Ia?aae
oa?iinoaooaaiiyi caii/aia ian?iiy ninie aeo?eioaaee 2 aeiae i?e
oaiia?aoo?? 40N a oai?ya? aeey no?aoeo?eaoe??. Aeey aeine?ae?a
a?aeae?aee i?i?inoee aeiaaeeiith 50-60 ii, c yeeo io?eioaaee oaeiae
i?ioiieanoe, oa ian?iiy.

?ineeie othothio (Nicotiana tabacum) aoee io?eiai? c noa?ee?ciaaiiai
ian?iiy, ui i?i?iuoaaee ia oaa?aeiio aaa?eciaaiiio na?aaeiaeu? MS
(Murasige, Skoog, 1962) a eiio?ieueiaaieo oiiaao: 250C, 25 W/i2,
oioiia??iae neeaaeaa 16 aiaeei ia i?ioyc? 5 iaae?eue. Aeey aenia?eiaio?a
a?aeae?aeeny iieiae? eenoee. A /anoei? aenia?eiaio?a c iieiaeeo eenoe?a
othothio io?eioaaee i?ioiieanoe.

A aenia?eiaioao ii?ii?ithaaee i?i?inoee ninie, iieiae? eenoee othothio,
ian?iiy ninie oa i?ioiieanoe oeeo ?ineei. Aeey aeayeeo aa??aio?a
aeine?ae?a i?ioiieanoe oa ian?iiy ninie ii?ii?ithaaee aaiia-i?iiaiyie ca
aeiiiiiaith eiaaeueoiaiai aaiia-i?eeaaeo «Enneaaeiaaoaeue» (?in?y) c
iiooaei?noth aeice 0,063 A?/n. I?ioiieanoe ninie ii?ii?ithaaeenue a
aeicao 20, 40 oa 50 A?. E??i oiai, i?ioiieanoe othothio oa ninie,
i?i?inoee ninie oa iieiae? eenoee othothio, ian?iiy ninie ii?ii?ithaaee
?aioaai?anueeeie i?iiaiyie ca aeiiiiiaith i?eeaaeo ?OI-17 (?in?y). Oe?
ia`?eoe ii?ii?ithaaee c iiooaeiinoyie ii?ii?iaiiy: 4 nA?/oa, 8 nA?/oa,
25 nA?/oa, 46 nA?/oa, 84 nA?/oa oa 118 nA?/oa a aeicao: 9,6 nA?, 24 nA?
oa 48 nA?.

Iaeiei c iieaciee?a i?iiaiaaiai o?aaeaiiy ian?iiy ninie aoei
i?i?inoaiiy ian?iiy, ui aecia/aeinue ca oi?ioeith: Ii=nto100 n0, aea:

Ii — i?ioeaio i?i?inoaiiy ian?iiy;

n0 — caaaeueia e?euee?noue ian?iiy, ui aoei iinoaaeaii ia i?i?inoaiiy;

nt — e?euee?noue ian?iei, ui i?i?inee ia 10 aeaiue i?ney caii/oaaiiy.

Aeae?eaiiy i?ioiieano?a. I?ioiieanoe aeae?eyee c i?i?inoe?a ninie oa
iieiaeeo eenoe?a othothio ca noaiaea?oiith iaoiaeeeith (Ae?aeia? e ae?.,
1991). Oa?iaioiee ?ic/ei i?noea: 1 % Onozuka R 10 (Yakult Biochemicals,
ssiii?y), 0,75 % Macerozyme R 10 (Yacult Biochemicals, ssiii?y) 0,75 %
Cellulase (Ferak, I?ia//eia), 0,75 % Pectinase ( Ferak, I?ia//eia), 0,5
I oeoe?ico, 50 iI NaNl2, (pH 5,5). I?ioiieanoe ?ieoaoaaee a 1 ie
na?aaeiaeua A-5 (Gamborg et. al. 1968). *an ?ieoaoaaiiy i?ioiieano?a
i?ney ii?ii?iaiiy a aeayeeo aa??aioao aeine?ae?a nyaaa 6 aiaeei. Aeey
aeayeeo aenia?eiaio?a aei na?aaeiaeua A-5 c i?ioiieanoaie ninie
aeiaeaaaee ?ia?a?oi? iie?ia?ac a oa s — ao?ae?eie?i (Sigma, NOA) a
eiioeaio?aoe?? 50 ia/ie (With et. al., 1995). I?e aeei?enoaii?
i?ioiieano?a aeey aiae?co AeIE iaoiaeii ioeuen-aeaeo?ioi?aco
i?ioiieanoe ninie (0,2 ie) ?anoniaiaeoaaee a ??aiiio ia`?i? 1 % —
iecueeiieaaei? aaa?ice (Biorad, NOA), caaeainiaaii? ia na?aaeiaeu? W-5
i?e 37iN oa aeaaaee canoeaiooe, i?ney /iai aeiaeaaaee ua 0,2 ie
na?aaeiaeua A-5.

AEeoo?caeaoi?noue i?ioiieano?a aecia/aee ca noaiaea?oiith iaoiaeeeith
(Oo?oaa? 1976).

Aiae?c AeIE i?ioiieano?a i?iaiaeeee iaoiaeii ioeuen-aeaeo?ioi?aco (IAO)
a aaa?icieo aaeyo. I?ney ii?ii?iaiiy i?ioiieano?a a aaa?icieo aeieao ?o
e?coaaee, aa??thth/e i?e oeueiio o?eaae?noue i?ii?aeeo /ano i?ae
ii?ii?iaiiyi oa iiaeaeueoei e?cenii. I?aia?aoe i?ioiieano?a e?coaaee 200
iee ?ic/eio: 2,5 I NaCl, 1 % Oriton X-100 (Ferak, I?ia//eia) (pH 9,0).
Aeo?eioaaee i?ioyaii 1,5 aiaeeie i?e 4iN, i?ney /iai ?ic/ei ceeaaee,
aeiaeaaaee 200 iee na?aaeiaeua ESP (Aeaeaen e ae?., 1990) oa ?ieoaoaaee
16 aiaeei i?e 37iN. Aeey i?aeaeuaiiy /ooeeaino? iaoiaeo IAO, i?aia?aoe
AeIE i?aeaeaaaee ia?iaoe? S1-ioeeaacith 100 IA (Amersham,
Aaeeeia?eoai?y) (Maniatis 1982). Aeey oeueiai aaa?ici? anoaaee
a?aeieaaee i?ioyaii 30 oa, aeiaeath/e 1 ie ?ic/eio OA (Maniatis 1982),
i?ney /iai aei anoaaie aeiaeaaaee OA — aooa?, yeee i?noea 0,5 iI
oai?eiaoeenoeueoii?eooi?eaeo (OINO, Serva, I?ia//eia), ? ?ieoaoaaee a
oa?iinoao? i?e 37iN 40 oa. Ia ianooiiiio aoai? anoaaee a?aeieaaee a 1 ie
OA — aooa?o i?ioyaii 30 oa (Maniatis 1982). I?ney oeueiai c?acee
?ici?uoaaee o 150 iee aooa?? aeey S1- ioeeaace (Maniatis 1982) ?
?ieoaoaaee i?e 37iN i?ioyaii 3 aiaeei. ?aaeoe?eio noi?o ioieiaeaeoaaee
aei 0iN, ceeaaee, aaa?ici? c?acee a?aeieaaee a OA-aooa??. I?ney oeueiai
AeIE c?ace?a ?icae?eyee ca aeiiiiiaith IAO. Ia eiaeio aei??aeeo a aae?
iaiineee i?eaeecii 1-3 iea AeIE. Aeaeo?ioi?aoe/ia ?icae?eaiiy AeIE
cae?enithaaee ca aeiiiiiaith i?eeaaeo «Aeeaioeuen» (?in?y) ca?aeii
iaoiaeeee (Elia et. al., 1993) Oiiae i?iaaaeaiiy: iai?oaa 85 A, 199 iA,
/an ioeueno 45 oa i?ioyaii 14 aiaeei. Iio?i i?ioyaii 7 aiaeei /an
ioeueno neeaaeaa 20 oa. Ia ianooiiiio aoai? i?ioyaii 3 aiaeei /an
ioeueno neeaaeaa 30 n. I?ney i?iaaaeaiiy aeaeo?ioi?aco aaeue oa?aoaaee
i?ioyaii 10 oa ?ic/eiii a?iienoiai aoeae?th (Sigma, NOA) a
eiioeaio?aoe?? 20 iea/ie oa oioia?aooaaee (Maniatis 1982).

AeIE aeae?eyee c i?i?inoe?a ninie oa ian?iiy ninie, a oaeiae c iieiaeeo
eenoe?a othothio, ui aoee caio?ai? o ??aeeee acio a ??ciee /an i?ney
ii?ii?iaiiy. Aeae?eaiiy i?iaiaeeee a i?ia??eao oeio Eppendorf 1,5ie ca
aeiiiiiaith AeNI oa i?ioa?iace E ca?aeii noaiaea?oii? iaoiaeeee
(Ae?aeia? e ae?., 1991).

Aiae?c caaaeueii? AeIE ?ineeiieo oeaiei i?iaiaeeee iaoiaeii eoaeiiai
aeaeo?ioi?aco a aaa?icieo aaeyo. Aeaeo?ioi?ac i?iaiaeeee ca iaoiaeeeith
(Maniatis 1982). Oiiae aeaeo?ioi?aco aoee oae?: 28 A, 30 iA ia i?ioyc?
18 aiaeei i?e oaiia?aoo?? 4 iN. I?ney aeaeo?ioi?aco aaa?iciee aaeue
iaeo?ae?coaaee ia i?ioyc? 45 oa a aooa??: 1I Tris-ICl (pH 7,6), 1,5M
NaCl. I?ney oe??? i?ioeaaeo?e aaeue caio?thaaee a ?ic/ei a?iienoiai
aoeae?th (Sigma, NOA) c eiioeaio?aoe??th 20 iea/ie. I?ney i?iaaaeaiiy
aeaeo?ioi?aco aaeue oa?aoaaee i?ioyaii 10 oa ?ic/eiii a?iienoiai
aoeae?th (Sigma, NOA) a eiioeaio?aoe?? 20 iea/ie oa oioia?aooaaee
(Maniatis 1982).

Aeey e?euee?nii? ioe?iee nooiaiy ooeiaeaeaiino? AeIE aeei?enoiaoaaee
iaoiae eoaeii? aeaiaoo?aoe?? c iiaeaeueoei ?icae?eaiiyi o?aeoe??
iaeiiieoeiai? oa aeaiieoeiai? AeIE ia eieiioe? c a?ae?ieneeaiaoeoii ca
iaoiaeeeith, ui iienaia (?aoaoieeia e ae?., 1996). A iaio o?aeoe?yo
e?euee?noue AeIE aecia/aee niaeo?ioioiiao?e/ii ia niaeo?ioioiiao??
NO-2O (?in?y). *anoea iaeiiieoeiai? AeIE a caaaeueiiio ai?no?
a?iiie?ia?o a?aeia?aaeaea nooi?iue ooeiaeaeaiino? ? aoea ae?aaeaia o
a?aeiinieo iaeeieoeyo: N=M/Mk aea M — a?aeiioaiiy e?eueeino?
iaeiiieoeiai? AeIE aei caaaeueii? e?eueeino? AeIE c?aceo, Mk —
a?aeiioaiiy e?eueeino? iaeiiieoeiai? AeIE a eiio?ie? aei caaaeueii?
e?eueeino? AeIE a eiio?ie?.

Aeey ia?aa??ee aeaieo aeine?aee iiaoi?thaaee o?e ?ace. Noaoenoe/ii
ia?iaeo aenia?eiaioaeueieo aeaieo i?iaiaeeee ca aeiiiiiaith
eiii’thoa?ii? i?ia?aie MicroCal Origin 3.1, i?e oeueiio aeey ioe?iee
aeinoia??iino? aeaieo aeei?enoiaoaaee T — oano (aiae?c aeaio
iiioeyoe?e), oaeiae ae?aoiaoaaee noaiaea?oio iiieeeo.

?icae?e 3. Iniaeeaino? ci?i a AeIE ian?iiy ninie cae/aeii?, ui aoei
ii?ii?iaia c ??ciith iiooaei?noth iecueeiaeiciaei ii?ii?iaiiyi.

?acoeueoaoe aea/aiiy i?i?inoaiiy ian?iiy ninie cae/aeii?, ui io?eiaee ca
2 ?iee naiai oi?ioaaiiy ??cio aeico o?ii?/iiai ii?ii?iaiiy, oa ioe?iea
aeeoo?caeaoiino? i?ioiieano?a, ye? io?eiai? c i?i?inoe?a oeueiai
ian?iiy, oa aoee aeiaeaoeiai ii?ii?iaii? aino?i aaiia-i?iiaiyie a aeicao
20 oa 40 A?, na?ae/aoue i?i oa, ui aeey oeeo iieaciee?a niinoa??aa?oueny
neeaaeia caeaaei?noue a?ae aeice. Anoaiiaeaia a?eueoa ?aae?i/ooeea?noue
i?ioiieano?a oa ian?iiy, ui ca aeaa ?iee naiai oi?ioaaiiy io?eiaei
aeico 0,03 A?, i?ae i?ioiieano?a oa ian?iiy, ui ca oie aea ia??iae
io?eiaei 3 A? oa 6 A?, i?e/iio iieacieee aeeoo?caeaoiino? i?e iiaeeiai?e
aeic? o?ii?/iiai ii?ii?iaiiy a 3 A? iaeaea ia a?ae??ciy?oueny a?ae
eiio?ieth. Iae?i?eia caeaaei?noue aoaeoo a?ae aeice iiaea aooe
iaoiiaeaia oei, ui i?ae /an naiai oi?ioaaiiy a ian?ii? ninie i?a
iaeiie/eoenue iaaiee ??aaiue i?eoiaaieo iioeiaeaeaiue, ye? iiaee
?aae?coaaoeny a iiaeaeueoiio.

Aeey ye?niiai aiae?co oa?aeoa?o ci?i a AeIE i?ioiieano?a ninie c ??cieo
ae?eyiie 30-ei ciie *AAN aeei?enoiaoaaee iaoiae ioeuen-aeaeo?ioi?aco
AeIE a aaa?icieo aaeyo. Aeey ia?aa??ee aoaeoeaiino? ?iaioe nenoai
?aia?aoe?? AeIE, a oaeiae aea/aiiy e?iaoeee aeaa?aaeaoe?? AeIE,
i?ioiieanoe c ??ciith eoioeyoeaiith aeicith ii?ii?iaiiy aeiaeaoeiai
ii?ii?ithaaee aino?i aaiia-i?iiaiyie a aeicao 20 ? 50 A?. I?e aino?iio
ii?ii?iaii? i?ioiieano?a a aeic? 20 A?, aeey an?o aa??aio?a, aea e?cen
a?aeaoaaany aaciina?aaeiuei i?ney ii?ii?iaiiy, aeanoeaa iayai?noue o?aeo
o?aaiaioiaaii? AeIE, ui aeaco? ia i?io?eaiiy i?ioean?a aeaa?aaeaoe??
AeIE. A c?aceao, ye? aoee ?ieoaiaai? i?ioyaii 4 aiaeei i?ney ii?ii?iaiiy
a aeic? 20 A?, o?aee o?aaiaioiaaii? AeIE aoee iaeaea iaiii?oi?, aea oeae
oaiiiai noino?oueny eeoa oeo i?ioiieano?a, ye? caciaee aieeao o?ii?/iiai
ii?ii?iaiiy. sseui ian?iiy ia caciaaaei oaeiai aieeao (eiio?ieue),
oa?aeoa? o?aeo o?aaiaioiaaii? AeIE i?ney 4 aiaeei ?ieoaaoe?? ia
a?ae??ciyany ca oa?aeoa?ii a c?aceao aac aeiaeaoeiai? ?ieoaaoe??. AeIE
c?ace?a, ui io?eiaee aeico o?ii?/iiai ii?ii?iaiiy 0,03 A?, i?e
aeiaeaoeiaiio ii?ii?iaii? a aeic? 50 A? AeIE aeoaea neeueii aeaa?aaeo?.
O aeiaaeeo eiio?ieueieo c?ace?a oa c?ace?a, ui io?eiaee aeico o?ii?/iiai
ii?ii?iaiiy 6 A?, iaaiee ??aaiue aeaa?aaeaoe?? niinoa??aa?oueny ia 4
aiaeei? i?ney aeiaeaoeiaiai ii?ii?iaiiy. AeIE c i?ioiieano?a, ui
io?eiaei ca 2 ?iee aeico 3 A?, i?aeoe/ii ia ia? oaiaeaioe?? aei
aeaa?aaeaoe??. Oaeei /eiii iiaeia i?eionoeoe, ui ian?iiy, yea ca /an
naiai oi?ioaaiiy io?eiaei aeico 3 A?, ia? a?eueo aenie? ?aia?aoe?ei?
caeaoiino?, ii??aiyii c ?ioeie c?aceaie.

I?e aiae?c? AeIE o?iiaoia?ao?/ii aoee aeyaeai? «i?eoiaai?» iioeiaeaeaiiy
AeIE a i?ioiieanoao, ui aeine?aeaeoaaeenue aac aeiaeaoeiaiai
ii?ii?iaiiy. Aiae?c aeiciai? caeaaeiino? a?aeiinii? e?eueeino?
iaeiiieoeiai? AeIE a i?ioiieanoao i?iaeaiiino?oaaa (?en. 1), ui i?e
aeic? aeiaeaoeiaiai aino?iai ii?ii?iaiiy a 20 A? e?euee?noue
iaeiiieoeiai? AeIE ciaioo?oueny ii??aiyii c oeei iieacieeii i?e aeic? a
40 A?. I?e aeiaeaoeiaiio aino?iio ii?ii?iaii? a aeic? 20 A? i?ioiieano?a
c ian?iiy, ui io?eiaee aeice o?ii?/iiai ii?ii?iaiiy 3 A? oa 6 A?,
iieaciee e?eueeino? iaeiiieoeiai? AeIE iaeaea ia a?ae??ciyany a?ae
eiio?ieth aaea /a?ac 30 oaeeei i?ney aino?iai ii?ii?iaiiy. Oe?eaai oa,
ui i?e ?ieoaaoe?? ee?oei i?ioyaii 4 aiaeei ??aaiue iaeiiieoeiai? AeIE
?a?no?oaaany ieae/ei, i?ae cia/aiiy oeueiai iieacieea aeey iaii?ii?iaieo
aeiaeaoeiai c?ace?a. Aiaeia?/i? caeaaeiino? niinoa??aathoueny ? i?e
ii?ii?iaii? oeeo i?ioiieano?a a aeic? 40 A?. Oaeei /eiii, i?ioean
ae?i?iaoe?? a?eueo ?ioaineaii a?aeaoaa?oueny o i?ioiieano?a, ian?iiy
yeeo io?eiaei eoioeyoeaio aeico 3 A?.

I?e o?iiaoia?ao?/iiio aiae?c? AeIE ian?iiy ninie cae/aeii? ii?ii?iaiiai
?aioaai?anueeeie i?iiaiyie a aeicao 48 nA? oa 24 nA? c ??cieie
iiooaeiinoyie aeice aeyaeaii, ui a?aeiinia e?euee?noue iaeiiieoeiai?
AeIE i?e ii?ii?iaii? cia/ii aeua ca eiio?ieueiee ??aaiue (?en. 2). Aea
oeae iieaciee ia ? iino?eiith aaee/eiith aeey iaaii? aeice, ?
oa?aeoa?eco?oueny a?aenooi?noth i?yii? caeaaeiino? a?ae iiooaeiino?
aeice. Aeiyoeii i?e oeueiio aenooia? ??aaiue iaeiiieoeiai? AeIE aeey
c?ace?a, ui aoee ii?ii?iai? a aeic? 48 nA? ? iiooaeiino? 84 nA?/oa, o
oeueiio aeiaaeeo oey oa?aeoa?enoeea i?aeoe/ii ia a?ae??ciyeanue a?ae
eiio?ieueiiai ??aith.

I?e ii?ii?iaii? ian?iiy ninie a aeic? 24 nA? iieaciee e?eueeino?
iaeiiieoeiai? AeIE a i?i?inoeao oeueiai ian?iiy aoa aeuee ca
eiio?ieueiee aeey iaeeo iiooaeiinoae aeic: 4 nA?/oa, 8 nA?/oa, 25
nA?/oa, a aeey ?ioeo iiooaeiinoae aeic ??aaiue iaeiiieoeiai? AeIE aoa
iacia/ii ieae/ee ca eiio?ieueiee (?en. 3). ??aaiue iaeiiieoeiai? AeIE a
i?i?inoeao c ian?iiy, ui aoei ii?ii?iaia a aeic? 48 nA?, eieeaa?oueny
oaeeaiiae?aii ? ia? o?e iaeneioie aeey iiooaeiinoae ii?ii?iaiiy 8
nA?/oa, 46 nA?/oa oa 118 nA?/oa oa aeaa i?i?ioie aeey iiooaeiinoae 25
nA?/oa oa 84 nA?/oa. Aeey ?aaeei?a ii?ii?iaiiy c iiooaeiinoyie 25
nA?/oa oa 84 nA?/oa a?aeiinia e?euee?noue iaeiiieoeiai? AeIE i?aeoe/ii
ia a?ae??ciy?oueny a?ae eiio?ieth aeea.?en. 3.

Oaeei /eiii, a i?ioiieanoao ninie, ui io?eiaee aeice o?ii?/iiai
ii?ii?iaiiy 3 A? oa 6 A? i?e aeiaeaoeiaiio ii?ii?iaii? a aeicao 20 A? oa
40 A?, iiaeooue ?i?oe?thaaoeny iiooaei? ?iaeooeeaaeuei? nenoaie
a?aeiiaeaiiy, ui a?ae?aia?iaothoue iane?aeee aino?iai ii?ii?iaiiy. Oe?
nenoaie ia ?iaeoeothoueny, yeui aeica o?ii?/iiai ii?ii?iaiiy neeaaeaea
0,03 A?. I?iaaaeai? aeine?aee iieacothoue, ui aeiaaio?eaaea ii?ii?iaiiy
iecueei? ?ioaineaiino? ian?iiy, ui oi?io?oueny, a iaaieo aeiciaeo
ae?aiaciiao, iiaea ca?eueoeoe caeaoi?noue a?aeiiaethaaoe AeIE a ee?oeiao
aei?ineiai i?aai?cio i?e ae?? aino?iai ii?ii?iaiiy a aeineoue aenieeo
aeicao oa iiaeeo?eoaaoe aeaye? iieacieee aeeoo?caeaoiino? ian?iiy. C
aeuacacia/aiiai iaoa??aeo aeaeii, ui iaa?oue a?aeiinii ei?ioei/ania
ii?ii?iaiiy ian?iiy iecueeiaeiciaei ii?ii?iaiiyi a aeayeeo aeiaaeeao
iiaea i?ecaiaeeoe aei ca?eueoaiiy ??aith iioeiaeaeaiue AeIE a
iiaeaeueoeo iieie?iiyo ee?oei, ui iaoiiaeaia ?aae?caoe??th /anoeie
«i?eoiaaieo» iioeiaeaeaiue AeIE.

?icae?e 4. Aieea iecueeiaeiciaiai ?aioaai?anueeiai ii?ii?iaiiy ??cii?
?ioaineaiino? ia aeo?ae iioeiaeaeaiue AeIE o ?ineeiieo i?ioiieano?a.

I?e ii?ii?iaii? i?ioiieano?a ninie c iiooaei?noth aeice 4 nA?/oa a aeic?
9,6 nA? ??aaiue iaeiiieoeiai? AeIE aoa ieae/ee ca eiio?ieueiee aanue /an
i?ney ii?ii?iaiiy (?en. 4). sseui aeica ii?ii?iaiiy neeaaeaea 24 nA?, oi
a ia?o? oaeeeie i?neyii?ii?iaiiy ??aaiue iaeiiieoeiai? AeIE aoa aeuee ca
eiio?ieue, a i?e iiaeaeueo?e ?ieoaaoe?? i?ioiieano?a i?ney ii?ii?iaiiy
a?i cieaeoaaany ? eieeaaany a iaaeao eiio?ieueiiai ??aith. I?e
ii?ii?iaii? a aeic? 48 nA?, a ia?o? oaeeeie i?ney ii?ii?iaiiy
niinoa??aaany i?aeaeuaiee aeo?ae iioeiaeaeaiue AeIE, aea c /anii ??aaiue
ooeiaeaeaiino? AeIE cieaeoaaany, ? ia 6 aiaeei? i?ney ii?ii?iaiiy a?i
aoa iaioee ca eiio?ieueiee. Iiaeia i?eionoeoe, ui o?eueee i?e
ii?ii?iaii? a aeic? 48 nA? ii/eiathoue ae?yoe ie?ai? eaiee
?iaeooeeaaeueieo ?aia?aoeaieo i?ioean?a, ai ??aaiue iaeiiieoeiai? AeIE
nooo?ai ieae/ee eiio?ieueiiai ia 6 aiaeei? ?ieoaaoe?? i?ney ii?ii?iaiiy.

I?e ii?ii?iaii? i?ioiieano?a ninie c iiooaei?noth aeice 8 nA?/oa, aeey
aeice ii?ii?iaiiy 9,6 nA? a?aei?/aiee ??aaiue iaeiiieoeiai? AeIE, ui
ieae/ee ca eiio?ieue. I?e ii?ii?iaii? a aeic? 24 nA?, iieaciee
e?eueeino? iaeiiieoeiai? AeIE ia 6 aiaeei? i?ney ii?ii?iaiiy noa?
ieae/ei ca eiio?ieueiee. Iaiaaii, i?e oae?e iiooaeiino? ii?ii?iaiiy
a?aeaoaa?oueny aeoeaaoe?y ie?aieo eaiie aaeaioeaii? a?aeiia?ae? aaea i?e
aeic? ii?ii?iaiiy 24 nA?. O aeiaaeeo ii?ii?iaiiy i?ioiieano?a c
?ioaineai?noth 8 nA?/oa a aeic? 48 nA? ??aaiue iaeiiieoeiai? AeIE ia 6
aiaeei? ?ieoaaoe?? i?ney ii?ii?iaiiy iaaeeaeaany aei eiio?ieueiiai, ui,
iaiaaii, iaoiiaeaii iane/ai?noth ?iiino?, ye aaea aeoeaiaaieo, oae ?
eiinoeoooeaieo ?aia?oth/eo nenoai.

sseui i?ioiieanoe ninie ii?ii?ithaaee c iiooaei?noth aeice 25 nA?/oa a
aeic? 9,6 nA?, oi niinoa??aaany iiieaeaiee ??aaiue iaeiiieoeiai? AeIE,
ii??aiyii c eiio?ieai. A??ia?aeii, ui oaeee iiieaeaiee ??aaiue
iaoiiaeaiee oei, ui eiinoeoooeaia nenoaia ?aia?aoe?? ae?i?io? an?
iioeiaeaeaiiy aaiiio. sseui aeica ii?ii?iaiiy neeaaeaea 24 nA?,
niinoa??aaeiny i?aeaeuaiiy aeoiaeo iioeiaeaeaiue AeIE, oiae? ye i?e
aeic? ii?ii?iaiiy 48 nA? a?aeaoaaeiny cieaeaiiy iieacieea e?eueeino?
iaeiiieoeiai? AeIE ia 1 oa 6 aiaeei? i?ney ii?ii?iaiiy, ui iiaea aooe
iaoiiaeaii caioneii iiaiiai eaneaaeo i?ioean?a aaeaioeaii? a?aeiia?ae?.
A oaeiio aeiaaeeo ii?ii?iaiiy c oaeith ?ioaineai?noth a aeic? 24 nA? ?
oei ?ioa?aaeii, aea ?iaeoeoe?? aaeaioeaii? a?aeiia?ae? a iiaiiio ianyc?
ia a?aeaoaa?oueny, a ?ii?noue ?nioth/eo ?aia?aoeaieo nenoai aaea
ae/a?iaia, oiio aeo?ae iioeiaeaeaiue AeIE ii/eia? ia?aaeuoaaoe
eiio?ieue.

Aoa i?iaaaeaiee aeine?ae, aea ee?oeie ii?ii?ithaaee c aea?aaeaioiei
o?aeoe?iioaaiiyi a aeic? 48 nA? (24 nA?+24 nA?) oa iiooaeiinoyie 25
nA?/oa oa 8 nA?/oa (?en 5). Aeaeii, ui i?e aeiaeaoeiaiio ii?ii?iaii? c
iiooaei?noth aeice 8 nA?/oa /a?ac 1 aiaeeio a aea?aaeaioi?e aeic?,
cia/ii cieaeo?oueny iieaciee e?eueeino? iaeiiieoeiai? AeIE, ? noa?
ieae/ei ca eiio?ieue. Aeiaeaoeiaa ii?ii?iaiiy ia 4 aiaeei? i?ney ia?oiai
oaeiae cieaeo? oeae iieaciee, aea a?eueo iia?eueii. Iiaeeeai, ui i?e
ae?oaiio ii?ii?iaii? c ?ioaineai?noth 8 nA?/oa /a?ac aiaeeio i?ney
ia?oiai iiaea a?aeaoaaoenue aeoeaaoe?y aeiaeaoeiaiai neioaco oa?iaio?a
?aia?aoe??, ui i?enooi? a ee?oei? ? a ii?iaeueieo oiiaao. O aeiaaeeo,
eiee ae?oaa ii?ii?iaiiy a?aeaoaaeiny /a?ac aiaeeio i?ney ia?oiai c
?ioaineai?noth 25 nA?/oa, niinoa??aa?oueny neooaoe?y, eiee iieaciee
iaeiiieoeiai? AeIE i?aeaeuo?oueny. Oaeo iiaaae?ieo e?eai? iiaeia
iiynieoe c iiceoe?e oiai, ui ii?ii?iaiiy a oaeee ?aae?i/ooeeaee iiiaio
aaaea aei oiai, ui /anoeia iioeiaeaeaiue, ni?e/eiaieo ii?ii?iaiiyi,
iiaea i?ecaiaeeoe aei iineaaeaiiy ei?aeoi?nueei? ooieoe?? aeayeeo
AeIE-iie?ia?ac (Iineaeueiaa, Eethoia 1990; Nie?ei 1990). C ?ioiai aieo,
eiee iiaoi?ia ii?ii?iaiiy i?iaiaeeee /a?ac 4 aiaeeie i?ney ia?oiai, oi a
oe?e neooaoe?? iaaooue a?aeaoaa?oueny iiaia ae?i?iaoe?y an?o
iioeiaeaeaiue, ai ia iiiaio 4-aiaeeiii? ?ieoaaoe?? i?e iaeii?aciaiio
ii?ii?iaii? a aeic? 24 nA? ??aaiue iaeiiieoeiai? AeIE iaaeeaeaany aei
eiio?ieueiiai.

Eiee ii?ii?iaiiy i?ioiieano?a ninie i?iaiaeeee a aeic? 24 nA? c
iiooaeiinoyie 8 nA?/oa ? 25 nA?/oa oa a i?enooiino? ao?ae?eie?io —
?ia?a?oi?o iie?ia?ac ? oa ? (?en. 6), oa?aeoa? caeaaeiinoae aeoiaeo
iaeiiieoeiai? AeIE a?ae iiooaeiino? aeice ii?ii?iaiiy ci?ith?oueny. I?e
ii?ii?iaii? c iiooaei?noth aeice 8 nA?/oa iieaciee e?eueeino?
iaeiiieoeiai? AeIE ca?eueooaaany. Oaea ca?eueoaiiy iaoiiaeaia oei, ui
ao?ae?eie?i aeieo? ?iaioo AeIE-iie?ia?ac ? oa ?, oiio iiaeeeaino?
?aia?aoeaii? nenoaie ciaioothoueny ? a?aeaoaa?oueny i?aeaeuaiiy ??aith
iaeiiieoeiai? AeIE. Aeaeii, ui i?e ii?ii?iaii? i?ioiieano?a c
iiooaei?noth 25 nA?/oa a i?enooiino? ao?ae?eie?io, caiaeii aeo?ae
iioeiaeaeaiue AeIE ciaioo?oueny. sse aoei cacia/aii ?ai?oa, oeae aoaeo
iiaea aooe iaoiiaeaiee oei, ui ii?ii?iaiiy c oaeith ?ioaineai?noth a
aeic? 24 nA? caeaoia iineaaeyoe ei?aeoi?nuee? ooieoe?? AeIE-iie?ia?ac
i?e ?ioeec?eiiio i?ioean?, aiane?aeie /iai ca?eueoo?oueny e?euee?noue
iiieeie i?e ?aia?aoe??, ? aeo?ae iioeiaeaeaiue AeIE ca?eueoo?oueny
(Nie?ei 1990).

I?e ii?ii?iaii? i?ioiieano?a ninie c iiooaei?noth aeice 46 nA?/oa a
aeic? 48 nA?, aeaeii nooo?aa cieaeaiiy ??aith iaeiiieoeiai? AeIE ia 6
aiaeei? ?ieoaaoe??. Oaeee i?iya aoaeoo iiaeia iiynieoe oei, ui i?e
ii?ii?iaii? a oaeiio ?aaeei?, e??i eiinoeoooeaieo nenoai ?aia?aoe??,
aeiaeaoeiai aeoeao?oueny neioac ?aia?aoe?eieo oa?iaio?a, ui ?niothoue a
ee?oei? a ii?iaeueieo oiiaao. Iaiaaii, i?e iiooaeiino? aeice ii?ii?iaiiy
46 nA?/oa a?aeaoaa?oueny ci?uaiiy a aeiciaee ae?aiacii 9,6 nA?
aeoeaaoe?? neioaco ?aia?oth/eo oa?iaio?a, ui ooieoe?iiothoue o ee?oeiao
? i?e ii?iaeueieo oiiaao. O aeiaaeeo, eiee aeica ii?ii?iaiiy neeaaeaea
24 nA?, a iiooaei?noue ii?ii?iaiiy 46 nA?/oa, niinoa??aa?oueny
neooaoe?y, i?e ye?e ?nioth/? a ee?oei? ?aia?aoeai?
nenoaie i?aoeththoue ia iaae? ia?aiaaaioaaeaiiy, aea aeoeaaoe??
anueiai eaneaaeo caoenieo iaoai?ci?a aaeaioeaii? a?aeiia?ae? ia
a?aeaoaa?oueny.

sseui iiooaei?noue aeice neeaaeaea 84 nA?/oa. a aeica 24 nA?, ??aaiue
iaeiiieoeiai? AeIE noaaaa iaioa eiio?ieueiiai. Oaeaeoa ca ana, ui i?e
ii?ii?iaii? a aeicao 24 oa 48 nA? a?aeaoaa?oueny aeoeaaoe?y nenoai
aaeaioeaii? a?aeiia?ae? a iiaiiio ianyc?. O aeiaaeeo, eiee i?ioiieanoe
ninie cae/aeii? ii?ii?ithaaee a oaeiio ?aaeei?, aea a aeic? 9,6 nA?,
niinoa??aa?oueny iiaia ae?i?iaoe?y an?o iioeiaeaeaiue AeIE nenoaiaie
eiinoeoooeaii? ?aia?aoe??, e??i oiai aeiaeaoeiai neioacothoueny
?aia?aoeai? oa?iaioe, ui ooieoe?iiothoue a ee?oei? ? a ii?iaeueieo
oiiaao.

I?e ii?ii?iaii? i?ioiieano?a ninie c iiooaei?noth aeice 118 nA?/oa oa
aeic? 24 nA? ne?ae caa?iooe oaaao ia oa, ui ?a?no?o?oueny iacia/ia
i?aeaeuaiiy iieacieea e?eueeino? iaeiiieoeiai? AeIE, ? aaee/eia oeueiai
iieacieea i?aeoe/ii ia ci?ith?oueny o /an?. Oaeee aoaeo iaoiiaeaiee oei,
ui a?aeaoaa?oueny ia?aaaioaaeaiiy ?iaioe eiinoeoooeaieo ? aaea /anoeiai
?iaeoeiaaieo nenoai ?aia?aoe??.

sse oea ia aeeaii, aea AeIE i?ioiieano?a othothio ia? a?eueoo
/ooeea?noue aei ae?? iecueeeo ??ai?a ?aae?aoe??, i?ae o i?ioiieano?a
ninie. O aeiaaeeo ii?ii?iaiiy i?ioiieano?a othothio c iiooaei?noth aeice
4 nA?/oa (?en. 7) iaeieae/?e ??aaiue iaeiiieoeiai? AeIE niinoa??aa?oueny
i?e ii?ii?iaii? a aeic? 24 nA?. Oeae iieaciee i?aeoe/ii ia ci?ith? nai?o
oa?aeoa?enoee o /an?. Aeoaea noiaea neooaoe?y niinoa??aa?oueny ? i?e
ii?ii?iaii? i?ioiieano?a othothio c iiooaei?noth 8 nA?/oa a o?e aea
nai?e aeic?. I?e aeic? ii?ii?iaiiy 9,6 nA? oa iiooaeiino? 4 nA?/oa
iieaciee iaeiiieoeiai? AeIE cieaeo?oueny iaeaea aei eiio?ieueiiai ca
aeineoue ei?ioeee i?ii?aeie /ano, a o aeiaaeeo c ii?ii?iaiiyi i?e
iiooaeiino? 8 nA?/oa oeae iieaciee ?a?no?oaaany iaa?oue ieae/a
eiio?ieueiiai ??aith. Iiaeia i?eionoeoe, ui i?e ii?ii?iaii? a aeic? 9,6
nA? c iiooaei?noth 4 nA?/oa oa 8 nA?/oa eiinoeoooeaia ?aia?aoeaia
nenoaia ee?oeie ni?iiiaeia i?aeoe/ii iiai?noth a?aeiiaeoe an?
iioeiaeaeaiiy AeIE. I?i?iaeueiiai cia/aiiy iieaciee iaeiiieoeiai? AeIE
i?ioiieanoe c othothio iaaoaa? i?e ii?ii?iaii? a aeic? 24 nA?. I?e
ca?eueoai? aeice aei 48 nA? aeo?ae iioeiaeaeaiue AeIE ii/eia? ciiao
ca?eueooaaoenue, ui iaoiiaeaia iinooiiaei iane/aiiyi ?iiino? ?aia?oth/eo
nenoai ee?oeie sse?nii ?ioa neooaoe?y niinoa??aa?oueny i?e ii?ii?iaii?
i?ioiieano?a othothio c iiooaei?noth 25 nA?/oa, i?i?iaeueia e?euee?noue
iaeiiieoeiai? AeIE niinoa??aa?oueny, eiee aeica ii?ii?iaiiy neeaaea? 24
nA?. E??i oiai i?aeaeuaiiy aeoiaeo iaeiiieoeiai? AeIE ?a?no?o?oueny ?
i?e ii?ii?iaii? a aeic? 9,6 nA?. Iiae?aiee oa?aeoa? aeoiaeo
iaeiiieoeiai? AeIE niinoa??aa?oueny ? i?e ii?ii?iaii? i?ioiieano?a
othothio c iiooaei?noth aeice 46 nA?/oa. I?e iiaeaeueoiio ca?eueoaii?
iiooaeiino? ii?ii?iaiiy aei 84 oa 118 nA?/oa a?aei?/aii aeayea ciaioaiiy
??aith iaeiiieoeiai? AeIE o aeiaaeeo io?eiaii? aeice 9,6 nA?. Iaiaaii,
oaea cieaeaiiy ??aiy iaeiiieoeiai? AeIE iaoiiaeaia aeoeaaoe??th
aeiaeaoeiaiai neioaco oa?iaio?a ?aia?aoe??, ui ooieoe?iiothoue a
ee?oei? ? ca ii?iaeueieo oiia. Nenoaie aaeaioeaii? a?aeiia?ae?
ii/eiathoue i?aoethaaoe a iiaiiio ianyc? eeoa i?e aeic? ii?ii?iaiiy 48
nA?. Ne?ae cacia/eoe, ui i?e ii?ii?iaii? i?ioiieano?a othothio a aeic?
48 nA? oa iiooaeiino? 118 nA?/oa nenoaia aaeaioeaii? a?aeiia?ae?
i?aoeth? ia iaae? iane/aiiy, oiio ui ??aaiue iaeiiieoeiai? AeIE
iaaeeaea?oueny aei eiio?ieueieo iaae.

O?aaa caoaaaeeoe, ui aaeaioeaia a?aeiia?aeue — neeaaeiee ? aaaaoiaoaiiee
i?ioean, ? iiaeeeaa neooaoe?y, eiee i?e iaaieo aeiciaeo iaaaioaaeaiiyo
aaea ii/eiathoue i?aoethaaoe aeaye? ii/aoeia? eaiee oeueiai i?ioeano,
aea oeueiai iaaaioaaeaiiy ia aeinoaoiuei aeey caioneo iiaiiai eaneaaeo
?aaeoe?e aaeaioeaii? a?aeiia?ae?. Aeoeaaoe?? ii/aoeiaeo eaiie
aaeaioeaii? a?aeiia?ae?, cie?aia aeoeaaoe?? neioaco oa?iaio?a
?aia?aoe??, ui ooieoe?iiothoue a ee?oei? ? a ii?iaeueieo oiiaao, iiaea
ia aenoa/aoe uia iiai?noth ae?i?ioaaoe an? iioeiaeaeaiiy AeIE, a caioneo
iiaiiai eaneaaeo ?aaeoe?e aaeaioeaii? a?aeiia?ae? ia a?aeaoaa?oueny. A
oae?e neooaoe?? ne?ae i/?eoaaoe i?aeaeuaiiy aeoiaeo iioeiaeaeaiue AeIE o
ii?ii?iaieo ee?oei. Oaeei /eiii, i?e ii?ii?iaii? i?ioiieano?a ninie oa
othothio c ??cieie iiooaeiinoyie a iecueeiaeiciaiio ae?aiacii?, aeica
ii?ii?iaiiy 24 nA? iiaea aooe oith e?eoe/iith aaee/eiith, i?e ye?e iiaea
a?aeaoaaoeny i?aeaeuaiiy, ii??aiyii c eiio?ieai, aeoiaeo iioeiaeaeaiue
AeIE, ui iaoiiaeaii oaeoii iani?iiiaeiino? ae?i?iaoe?? an?o
iioeiaeaeaiue AeIE, ui ooai?eeenue aiane?aeie ii?ii?iaiiy.

?icae?e 5. Oa?aeoa? ooeiaeaeaiino? AeIE ee?oei i?i?inoe?a ninie, ui aoee
ii?ii?iai? ?aioaai?anueeeie i?iiaiyie ??cii? ?ioaineaiino? a
iecueeiaeiciaiio ae?aiacii?.

?acoeueoaoe eoaeiiai aeaeo?ioi?aco c?ace?a AeIE i?i?inoe?a ninie, ui
aoee ii?ii?iai? i?e ??ci?e ?ioaineaiino? oa a?ae?aco i?ney oe???
i?ioeaaeo?e e?ciaai?, aeaiiino?othoue, ui i?e aeic? ii?ii?iaiiy 24 nA?
iaea?eueoee ??aaiue iioeiaeaeaiue AeIE ii a?aeiioaiith aei eiio?ieth
niinoa??aa?oueny aeey c?ace?a c iiooaei?noth aeice 8 nA?/oa. sseui
aeica ii?ii?iaiiy neeaaeaea 48 nA?, i?e ?ioaineaiino? aeice 8 nA?/oa oa
118 nA?/oa niinoa??aa?oueny iaea?eueoee ??aaiue ooeiaeaeaiino? AeIE.
O?iiaoia?ao?/iee aiae?c AeIE i?iaeaiiino?oaaa, ui i?e iiooaeiino? aeice
ii?ii?iaiiy 4 nA?/oa oa aeic? 24 nA? ?a?no?o?oueny iaeaeuee iieaciee
iaeiiieoeiai? AeIE a?ae?aco i?ney ii?ii?iaiiy, aea c /anii cia/aiiy
oeueiai iieacieeo ciaioo?oueny (?en. 8). Iiae?aia ea?oeia
niinoa??aa?oueny ? i?e ii?ii?iaii? i?i?inoe?a a aeic? 24 nA? c
iiooaei?noth 8 nA?/oa.

O oeueiio aeiaaeeo aeoiaeeoue, ui iaeaeuiio ??aith iaeiiieoeiai? AeIE
a?ae?aco i?ney ii?ii?iaiiy a?aeiia?aea? iaeiaioa cia/aiiy oeueiai
iieacieea ia 6 aiaeei? i?ney ii?ii?iaiiy, ui aeaco? ia iia?eueia
i?io?eaiiy i?ioean?a a?aeiiaeaiiy AeIE. I?e ii?ii?iaii? i?i?inoe?a a
aeic? 9,6 nA? c oeie naieie iiooaeiinoyie niinoa??aa?oueny iacia/ia
i?aeaeuaiiy ??aith iaeiiieoeiai? AeIE, ui i?aeoe/ii ia ci?ith?oueny
aanue oanoiaaiee ia??iae /ano. Oe?eeii eiia??ii, ui i?e oaeeo oiiaao
ii?ii?iaiiy a?aeiiaeaiiy iioeiaeaeaiue a?aeaoaa?oueny ca ?aooiie
eiinoeoooeaieo ?aia?aoe?eieo nenoai ee?oei. O aeiaaeeo aeice ii?ii?iaiiy
48 nA? oa iiooaeiino? aeice 4 nA?/oa o i?i?inoe?a ninie ?a?no?o?oueny
ciiao i?aeaeuaiiy iieacieea iaeiiieoeiai? AeIE, ui aeaco? ia neaae?
caeaoiino? a?aeiiaeaiiy. I?e ii?ii?iaii? i?i?inoe?a c iiooaei?noth 8
nA?/oa, c /anii ?a?no?o?oueny iinooiiaa ciaioaiiy iieacieea
ooeiaeaeaiino? AeIE. I?e/iio, ooo oaeiae niinoa??aa?oueny, oi/a ? a
iaio?e nooiai?, oaiaeaioe?y a?aeiia?aeiino? — /ei aeua ooeiaeaeai?noue
AeIE a?ae?aco i?ney ii?ii?iaiiy, oei ieae/ee aoaea oeae iieaciee caiaeii
i?ney ii?ii?iaiiy.

I?e ii?ii?iaii? c iiooaei?noth aeice 25 nA?/oa oa aeic? ii?ii?iaiiy 9,6
nA? iaeaeuee ??aaiue iaeiiieoeiai? AeIE ?a?no?o?oueny a?ae?aco i?ney
ii?ii?iaiiy, a iaeieae/ee ia 6 aiaeei? i?ney ii?ii?iaiiy. Iaiaaii oaea
ca?eueoaiiy iiooaeiino? aeice ii?ii?iaiiy i?ecaiaeeoue aei ci?uaiiy
caioneo i?ioean?a a?aeiiaeaiiy a iaeanoue iaioeo aeic. I?e ii?ii?iaii? a
aeicao 24 nA? oa 48 nA? aeaeii, ui iieaciee e?eueeino? iaeiiieoeiai?
AeIE ia 6 aiaeei? i?ney ii?ii?iaiiy aeuee, i?ae ca?a?no?iaaiee a?ae?aco
i?ney ii?ii?iaiiy. ??aaiue iaeiiieoeiai? AeIE ia 6 aiaeei? i?ney
ii?ii?iaiiy i?aeoe/ii ia a?ae??ciy?oueny aeey oeeo aeic. Iiaeeeai oea ?
? oie naiee aeiaaeie, eiee ??aaiue iioeiaeaeaiue AeIE aeuee
eiio?ieueiiai ? ia caeaaeeoue a?ae aaee/eie aeice ii?ii?iaiiy (iaeanoue
«ieaoi»), ui iaoiiaeaii ?iaioith nenoai SOS-a?aeiia?ae? (Aa?anueeei
1998).

sseui i?i?inoee ninie ii?ii?ithaaee i?e iiooaei?noth aeice 46 nA?/oa,
??aaiue iaeiiieoeiai? AeIE i?e aeic? ii?ii?iaiiy 9,6 nA? ca?eueooaaany c
/anii. Iiaeeeai, ui i?e oae?e ?ioaineaiino? ii?ii?iaiiy oaeiae
niinoa??aa?oueny aeoeaaoe?y nenoai SOS-a?aeiia?ae?. Neooaoe?y, yea
niinoa??aa?oueny i?e ii?ii?iaii? a a?eueo aenieeo aeicao, iaiaaii aaea
iaoiiaeaia oei, ui a oeueiio aeiaaeeo a?aeaoaa?oueny ia?ao?ae ee?oei a
«aaa??eiee» ?aaeei ?iaioe (Aa?anueeei 1998). A??ia?aeii, ui aoaeoe, ye?
?a?no?o?oueny i?e ii?ii?iaii? i?i?inoe?a ninie c iiooaeiinoyie aeice 84
nA?/oa oa 118 nA?/oa, iaoiiaeai? iiae?aieie i?ioeanaie, ui ? i?e
ii?ii?iaii? c iiooaei?noth aeice 46 nA?/oa. Eiia??ii, ui i?e iineeaii?
?ioaineaiino? ii?ii?iaiiy a?aeaoaathoueny ci?ie ia o?eueee ia ??ai?
ee?oei, aea ? ia ??ai? oe?eiai i?aai?cio. C oaei? iiceoe?? eieeaaiiy
iieacieea iaeiiieoeiai? AeIE, ui ?a?no?o?oueny, iaoiiaeaia ia o?eueee
?aia?aoe?eieie i?ioeanaie, aea ? noaiii AeIE a ee?oeiiiio yae??, ui
iia`ycaiee c noaae??th ee?oeiiiai oeeeeo oa o?c?ieia?/iith ooieoe??th
ee?oeie.

C aeuacacia/aiiai iaoa??aeo aeaeii, ui ii e?eoa??th aeoiaeo
iaeiiieoeiai? AeIE a?ae iiooaeiino? aeice/aeice ii?ii?iaiiy ee?oeie
i?i?inoe?a ninie ? a?eueo /ooeeaeie aei aieeao iecueeiaeiciaiai
ii?ii?iaiiy, i?ae i?ioiieanoe oe??? ?ineeie. A iaaaaeaieo aeiciaeo
ae?aiaciiao oaeoi? iiooaeiino? aeice ii?ii?iaiiy ? aaaeeeaei /eiieeii,
ui iiaea ci?ithaaoe oa?aeoa? i?ioean?a a?aeiiaeaiiy.

AENIIAEE

1. O?ii?/ia ?ii?coth/a ii?ii?iaiiy ian?iiy ninie a aeicao 0,03 A?, 3 A?
oa 6 A? ca ia??iae eiai oi?ioaaiiy ?iaeoeo? «i?eoiaai?» iioeiaeaeaiiy
AeIE. Iioeiaeaeaiiy, ui noi?ioaaeenue aiane?aeie o?ii?/iiai ii?ii?iaiiy
a aeicao 3 oa 6 A?, aenooiathoue aeoeaaoi?aie nenoai a?aeiiaeaiiy.

2. I?e ii?ii?iaii? nooiai ian?iiy ninie a aeicao 24 nA? oa 48 nA?
niinoa??aa?oueny c?inoaiiy ??aith ooeiaeaeaiiino? AeIE. E?euee?noue
iioeiaeaeaiue AeIE aecia/a?oueny cia/aiiyi iiooaeiino? aeice
ii?ii?iaiiy. Aeey i?i?inoe?a, ui io?eiai? c ii?ii?iaiiai ian?iiy, ia
a?aei?/aii i?yii? caeaaeiino? ??aiy ooeiaeaeaiino? AeIE c oeei
iieacieeii a ian?ii?.

3. I?e ii?ii?iaii? i?i?inoe?a ninie oa?aeoa?ia a?eueoa /ooeea?noue AeIE
aei aieeao iecueeiaeiciaiai ii?ii?iaiiy, i?ae aeey i?ioiieano?a oe???
?ineeie.

4. I?e ii?ii?iaii? i?ioiieano?a ninie a aeic? 24 nA? c iiooaei?noth
aeice a 25 nA?/oa oa 118 nA?/oa niinoa??aa?oueny c?inoaiiy ??aith
ooeiaeaeaiino? AeIE. Oe? ianoaaeie na?ae/aoue i?i oa, ui i?e oaeeo
?aaeeiao ii?ii?iaiiy nenoaie ?aia?aoe??, ui a aeaiiee iiiaio
ooieoe?iiothoue a ee?oei?, ia ni?iiiaei? ae?i?ioaaoe an? iioeiaeaeaiiy
AeIE.

5. Ii?ii?iaiiy i?ioiieano?a ninie a aeic? 24 nA? oa a i?enooiino?
?ia?a?oi?a oa?iaio?a ?aia?aoe?? — ao?ae?eie?io iiaea i?ecaanoe ye aei
ca?eueoaiiy, oae ? aei ciaioaiiy ??aiy ooeiaeaeaiiino? AeIE, ui
aecia/a?oueny iieacieeii iiooaeiino? aeice ii?ii?iaiiy.

6. I?e ii?ii?iaii? i?ioiieano?a othothio a?aei?/a?oueny oioiaeia
caeaaei?noue i?ae ??aiai iioeiaeaeaiiy AeIE, ?aaeeiii ii?ii?iaiiy oa
oaeoi?ii /ano i?ney ii?ii?iaiiy. AeIE i?ioiieano?a othothio ? a?eueo
/ooeeaith aei aieeao iecueeiaeiciaiai ii?ii?iaiiy ??cii? ?ioaineaiino?,
i?ae AeIE i?ioiieano?a ninie.

7. I?e iecueeiaeiciaiio ii?ii?iaii? c ??ciith iiooaei?noth aeice ee?oei
ninie oa othothio, ?niothoue oae? iiooaeiino? aeice ? aeice ii?ii?iaiiy,
aieea yeeo ia i?ecaiaeeoue aei aeoeaaoe?? aeiaeaoeiaeo eaioetha?a
nenoaie a?aeiiaeaiiy, a iino?eii ?nioth/? a ee?oei? a?aeiiaethaaeuei?
nenoaie aaea ia a ciic? ae?i?ioaaoe an? ?aae?aoe?eii-?iaeoeiaai?
iioeiaeaeaiiy AeIE.

IA?AE?E ?IA?O, IIOAE?EIAAIEO CA OAIITH AeENA?OAOe??

1. Nieieia I.A., ?naaieia N.A., Ni?i/einueeee A.A. O?ii?/ia ii?ii?iaiiy
a iaeeo aeicao caeaoia iiaeeo?eoaaoe iieacieee aeeoo?caeaoiino? ian?iiy
ninie cae/aeii? ( Pinus silvestris) // Oeeoie. e aaiaoee.- 1998.- O.
32., ?4.- N. 65 — 71.

2. Enaaieia N.A., Nieieia N.A., Ni?i/eineee A.A. Aeieiae/aneea yooaeou
eiiece?othuaai eceo/aiey a iaeuo aeicao. Aiaeec nia?aiaiiuo iiaeoiaeia e
eiioeaioeee // I?iae. *i?iiaeeuenei? Ciie a?ae/oaeaiiy — 1998.- ?5.- N.
102 — 112.

3. Isaenkov S.V., Sokolov N.V., Sorochinsky B.V. Genom changes in pine
seeds after irradiation // Photosyn. : Mechanisms and Effects. — 1998. —
Vol. V. — P. 4101 — 4104.

4. Enaaieia N.A., Ni?i/eineee A.A., Nieieia I.A., A?iaeceineee Ae.I.
?aaeeaoeeiiii-eiaeooee?iaaiiua eciaiaiey a aaiiia naiyi niniu,
noi?ie?iaaaoeony a oneiaeyo o?iie/aneiai iaeo/aiey a Ciia io/oaeaeaiey
*AYN // Oe?. aeioei. aeo?i. — 1999. — O. 71., ? 4. — N. 103 — 106.

5. Enaaieia N.A., Nieieia N.A., Ni?i/eineee A.A. ?aaeno?aoeey ne?uouo
iia?aaeaeaiee aaiiia iaeo/aiiuo naiyi niniu iaoiaeii
ioeuen-yeaeo?ioi?aca // 3 Nuacae ii ?aaeeaoeeiiiui enneaaeiaaieyi
(Iineaa, 14 — 17 ieoya?y 1997). — Oacenu aeieeaaeia. — Ioueii, 1997. —
Oii 1. — N. 169

6. ?na?ieia N.A., Nieieia N.A., Ni?i/einueeee A.A. O?ii?/ia ii?ii?iaiiy
ian?iiy ninie cae/aeii? i?ae /an eiai oi?ioaaiiy aieeaa? ia aaiii oa
iiaeeo?eo? iieacieee aeeoo?caeaoiino? ian?iiy // I?aeia?iaeia
eiioa?aioe?y «Iioiaaiac ?ineei a i?e?iaeiiio oa o?ainoi?iiaaiiio
na?aaeiaeu?» (Euea?a, 1-4 eeiiy 1998). — Iaoa??aee i?aeia?iaeii?
eiioa?aioe?? — Euea?a, 1998. — N. 202 — 203.

7. Isaenkov S.V., Sokolov N.V., Sorochinsky B.V. Pine seeds genome
changes after irradiation// XI International Congress on Photosintesis
(Budapest, 17 — 22 August 1980): Abstr. — Budapest, 1998. — P. 207.

8. ?naaieia N.A. Aieea ao?ae?eie?io oa iiooaeiino? aeice ia oa?aeoa?
aeoiaeo iioeiaeaeaiue AeIE a i?ioiieanoao ninie, ii?ii?iaieo a aeic?
0,25 A? //Iaoeiai-i?aeoe/ia eiioa?aioe?y iieiaeeo a/aieo
“?ioaa?aoeai?noue ooieoe?e ?ineeiiiai i?aai?cio” (Ee?a, 3-4 aa?anaiue
1998). — Oace aeiiia?aeae — Ee?a, 1998. — N. 39 — 41.

9. Isaenkov S.V., Sorochinsky B.V. the role of chronic irradiation
intensity in the induction of DNA singlebreakes // International
Conference » Diagnosis and Treatment of Radiation injury ( de Doelen
Rotterdam, 30 August — 3 September 1998): Abstr. — Rotterdam, 1998 — P.
46.

10. Sorochinsky B.V., Zelenaya L.B., Isaenkov S.V. Microevolution
processes among coniferous plants from the Chernobyl Exclusion zone //
II International Symposium on Plant Biotechnology (Kiyv, 4-8 October
1998): Abstr. — Kyiv, 1998. — P. 125.

SUMMARY

Isaenkov S.V. The influence of dose intensity irradiation on DNA-damages
issuer in plant cells. — Manuscript.

Thesis for competition on scientific degree of the candidate of
biological sciences on the speciality 03.00.01 — radiobiology. Kiyv
University by Taras Shevchenko. Kyiv, 1999.

The thesis includes data about issuer of DNA-damages in plant cells
irradiated with different dose intensities at low level dose. The dose
intensity influence at low dose range on the character of DNA-damages
issuer has been investigated. It was demonstrated that the dose
intensity is sufficient for forming the answer of Nicotiana tabacum and
Pinus silvestris cells by DNA damages index on influence of low dose
irradiation. The dependence of DNA-damages issuer on dose intensity
irradiation has complicated character and does not correlate with the
value of dose intensity and absorbed dose. It was detected the increased
DNA-damages issuer for some condition of irradiation. The fact of hidden
damages realisation in pine seeds after low dose irradiation was
registrated. Higher radiosensivity of pine shoots in comparison with
pine protoplasts on low dose irradiation was registrated.

Key words: dose intensity, low dose irradiation, DNA-damages,
DNA-repair, protoplasts, shoots, seeds.

AIIOAOe?ss

?na?ieia N.A. Aieea iiooaeiino? aeice ia aeo?ae iioeiaeaeaiue AeIE a
?ineeiieo ee?oeiao. ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe
ca niaoe?aeuei?noth 03.00.01. — ?aae?ia?ieia?y, Ee?anueeee oi?aa?neoao
?iai? Oa?ana Oaa/aiea. Ee?a 1999.

Caoeua?oueny aeena?oaoe?eia ?iaioa, yea i?noeoue aeaii? i?i aeo?ae
iioeiaeaeaiue AeIE a ?ineeiieo ee?oeiao, ui aoee ii?ii?iai? c ??ciith
iiooaei?noth aeice ?ii?coth/ei iecueeiaeiciaei ii?ii?iaiiyi.
Aeine?aeaeaii aieea oaeoi?o iiooaeiino? aeice a iecueeiaeiciaiio
ae?aiacii? ia oa?aeoa? aeoiaeo iioeiaeaeaiue AeIE. Aoei iieacaii,
iiooaei?noue aeice ? aaaeeeaei /eiieeii a oi?ioaaii? ?aaeoe??
a?aeiia?ae? ee?oei othothio (Nicotiana tabacum) oa ninie cae/aeii?
(Pinus silvestris), ii e?eoa??th aeoiaeo iioeiaeaeaiue AeIE, ia
iecueeiaeiciaa ii?ii?iaiiy. Caeaaeiino? aeoiaeo iioeiaeaeaiue AeIE a?ae
iiooaeiino? aeice ii?ii?iaiiy iathoue neeaaeiee oa?aeoa? ? ia
ei?aeththoue c aaee/eiith iiooaeiino? oa aeice ii?ii?iaiiy. I?e aeayeeo
?aaeeiao ii?ii?iaiiy aeyaeaiee i?aeaeuaiee aeo?ae iioeiaeaeaiue AeIE a
ii??aiyii? c eiio?ieai. Ca?a?no?iaaiee oaeo ?aae?caoe?? i?eoiaaieo
iioeiaeaeaiue a ian?ii? ninie, ui aoei ii?ii?iaia a iecueeiaeiciaiio
ae?aiacii?. Iieacaia a?eueoa ?aae?i/ooeea?noue i?i?inoe?a ninie, i?ae
i?ioiieano?a oe??? ?ineeie ia aieea iecueeiaeiciaiai ii?ii?iaiiy.

Eeth/ia? neiaa: iiooaei?noue aeice, iecueeiaeiciaa ii?ii?iaiiy,
o?ii?/ia ii?ii?iaiiy, iioeiaeaeaiiy AeIE, ?aia?aoe?y AeIE, i?ioiieanoe,
i?i?inoee, ian?iiy.

AIIIOAOeEss

Enaaieia N.A. Aeeyiea iiuiinoe aeicu ia auoiae iia?aaeaeaiee AeIE a
?anoeoaeueiie eeaoea. — ?oeiienue

Aeenna?oaoeey ia nieneaiea iao/iie noaiaie eaiaeeaeaoa aeieiae/aneeo
iaoe ii niaoeeaeueiinoe 03.00.01 — ?aaeeiaeieiaey, Eeaaneee oieaa?neoao
eiaie. Oa?ana Oaa/aiea. Eeaa, 1999.

Caueuaaony aeenna?oaoeeiiiay ?aaioa, eioi?ay niaea?aeeo aeaiiua i
auoiaea iia?aaeaeaiee AeIE a ?anoeoaeueiuo eeaoeao, eioi?ua auee
iaeo/aiu n ?aciie iiuiinoueth aeicu a ieceiaeiciaii aeeaiaciia.
Iiyaeyaony ana aieueoa yenia?aiaioaeueiuo aeaiiuo, naeaeaoaeuenoaothueo
i iaeeiaeiii oa?aeoa?a i?iyaeaiey aeieiae/aneeo yooaeoia a ieceiaeiciaii
aeeaiaciia. Nouanoaoao, ii e?aeiae ia?a, aeaa i?e/eiu, eioi?ua
iaoneiaeeaatho iniaaiiinoe aeieiae/aneiai aeaenoaey ieceiaeiciaiai
eiiece?othuaai iaeo/aiey. Ia?aay naycaia n aiciiaeiinoueth ea/anoaaiiuo
eciaiaiee a ?ani?aaeaeaiee ii eeaoea niauoee iiaeiuaiey yia?aee. Aoi?ay
naycaia n iieaeoey?iuie iaoaieciaie, eioi?ua oi?ie?otho ?aaeoeee eeaoee
ia iaeo/aiea. Iiaeii i?aaeiieiaeeoue, /oi a oi?ie?iaaiee ?aaeoeee eeaoee
ia neaaia aicaeaenoaea aiaoiaai oaeoi?a aeaaiia cia/aiea eiatho
o?eaaa?iua iieaeoey?iua iaoaieciu, eioi?ua iaoneaaeeaatho ia?aeeth/aiea
eeaoee a iiaue ?aaeei ooieoeeiie?iaaiey.

Enneaaeiaaii aeeyiea oaeoi?a iiuiinoe aeicu iaeo/aiey a aeeaiaciia
ieceeo aeic ia oa?aeoa? auoiaea iia?aaeaeaiee AeIE. Iieacaii, /oi
iiuiinoue aeicu yaeyaony aaaeiui oaeoi?ii a oi?ie?iaaiee ioaaoiie
?aaeoeee eeaoie oaaaea (Nicotiana tabaccum) e niniu iaueiiaaiiie (Pinus
silvestris), ii e?eoa?eth auoiaea iia?aaeaeaiee AeIE, ia ieceiaeiciaia
iaeo/aiea. Caaeneiinoe auoiaea iia?aaeaeaiee AeIE io iiuiinoe aeicu
iaeo/aiey eiatho neiaeiue oa?aeoa? e ia ei?aee?otho n aaee/eiie iiuiinoe
e aeicu iaeo/aiey. I?e iaeioi?uo ?aaeeiao iaeo/aiey ioia/ai iiauoaiiue
auoiae iia?aaeaeaiee AeIE a i?ioiieanoao, eeaoeao naiyi e i?i?inoeao ii
n?aaiaieth n eiio?ieueiui o?iaiai. Ca?aaeno?e?iaai oaeo ?aaeecaoeee
ne?uouo iia?aaeaeaiee AeIE o naiyi niniu, iaeo/aiiuo eiiece?othuei aaiia
— e ?aioaaiianeei eceo/aieai a ieceiaeiciaii aeeaiaciia. Ioia/aia ?ieue
iaeioi?uo AeIE-iieeia?ac a i?ioeannao yeeieiaoeee iia?aaeaeaiee AeIE a
i?ioiieanoao niniu.

Onoaiiaeaii, /oi o?iie/aneia eiiece?othuaa iaeo/aiea ieceie
eioaineaiinoe iiaeao iiaeeoeoee?iaaoue iaeioi?ua iieacaoaee
aeecianiiniaiinoe naiyi niniu, a a iaeioi?uo neo/ayo oaaee/eaaoue
ainnoaiiaeoaeueiua niiniaiinoe i?e aeiiieieoaeueiii ino?ii aaiia
-iaeo/aiee. Iieacaii, /oi i?i?inoee niniu ioee/athony aieueoae
?aaee/oanoaeoaeueiinoueth, /ai eciee?iaaiiua i?ioiieanou yoiai ?anoaiey
i?e iaeo/aiee a ieceeo aeicao.

Iieo/aiiua ?acoeueoaou naeaeaoaeuenoaotho a iieueco iaaeinoaoi/iie
iainiiaaiiinoe i?eiaiaiey iauai?eiyoie ia aeaiiue iiiaio
eeiaeii-aacii?iaiaie eiioeaioeee. Iiauoaiiue auoiae iia?aaeaeaiee AeIE a
oanoe?oaiii aeiciaii aeeaiaciia iauyniyaony ia iniiaa i?aaeeiaeaiiie
aeiioacu, eioi?ay ninoieo a oii, /oi nouanoaoao ii?aaeaeaiiue aeeaiacii
aeic, a eioi?ii eiinoeoooeaiua ?aia?aoeaiua nenoaiu oaea ia a ninoiyiee
yeeieie?iaaoue ana iia?aaeaeaiey AeIE, eiaeooee?iaaiiua iaeo/aieai, a
aeoeaaoeee eiaeooeeaaeueiie ?aia?aoeaiie eiiiiiaiou ia i?ienoiaeeo
aneaaenoaea iaaeinoaoi/iie aaee/eiu eiaeoeoeee iia?aaeaeaiee AeIE
eiiece?othuei iaeo/aieai ca a?aiaiiie eioa?aae.

Eeth/aaua neiaa: iiuiinoue aeicu, ieceiaeiciaia iaeo/aiea,
o?iie/aneia iaeo/aiea, iia?aaeaeaiey AeIE, ?aia?aoeey AeIE, i?ioiieanou,
i?i?inoee, naiaia.

Похожие записи