I?i?noa?noai ina?oe oe?a?ie

*A?I?AAOeUeEEE AeA?AEAAIEE OI?AA?NEOAO

?i. TH.OAAeUeEIAE*A

E?AAAOeUe Aieiaeeie? Noaiaiiae/

OAeE 581.198

INIAEEAINO? IAOAAIE?CIO CEAE?A I?E Ae?? IECUeEEO OAIIA?AOO? IA ?INEEIE

03.00.04 — a?io?i?y

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

aeieoi?a a?ieia?/ieo iaoe

*a?i?aoe? – 1999

Aeena?oaoe??th ? ?oeiien

?iaioa aeeiiaia a ?inoeooo? o?c?ieia?? ?ineei ? aaiaoeee IAI Oe?a?ie,
i. Ee?a

Iaoeiaee eiinoeueoaio: aeaaeai?e IAI Oe?a?ie, aeieoi? a?ieia?/ieo iaoe,
i?ioani?

Aieiaeeie? Aaneeueiae/ Ii?aoi

?inoeooo o?c?ieia?? ?ineei ? aaiaoeee IAI Oe?a?ie,

aee?aeoi? ?inoeoooo

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

Aaneeue Aaneeueiae/ A?oa?iei

Oa?iii?euenueeee aea?aeaaiee iaaeaaia?/iee

oi?aa?neoao ?i. A.Aiaothea,

i?i?aeoi? c iaoeiai? ?iaioe, caa?aeoaa/ eaoaae?e

aeieoi? a?ieia?/ieo iaoe, i?ioani?

Ieueaa ?ooaai?aia Oa?ae

Euea?anueeee aea?aeaaiee oi?aa?neoao ?i. ?aaia O?aiea,

caa?aeoaa/ eaoaae?e.

aeieoi? a?ieia?/ieo iaoe,

A?eoi??y Eaceie??aia ssai?nueea

?inoeooo o?c?ieia?? ?ineei ? aaiaoeee IAI Oe?a?ie,

caa?aeoaa/ a?aeae?eo

I?ia?aeia i?aai?caoe?y: ?inoeooo ee?oeiii? a?ieia?? oa aaiaoe/ii?
?iaeaia??? IAI Oe?a?ie

Caoeno a?aeaoaeaoueny “20” aeiaoiy 1999 ?. i 14-00 aiae

ia can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee Ae 76.051.05 i?e
*a?i?aaoeueeiio aea?aeaaiiio oi?aa?neoao? ?i. TH. Oaaeueeiae/a ca
aae?anith:

274012, i. *a?i?aoe?, aoe. Eioethaeinueeiai,2.

C aeena?oaoe??th iiaeii iciaeiieoeny a a?a?e?ioaoe? *a?i?aaoeueeiai
aea?aeaaiiai oi?aa?neoaoo ?i. TH. Oaaeueeiae/a

Aaoi?aoa?ao ?ic?neaiee “ 17 ” aa?aniy 1999 ?.

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

Eaiaeeaeao a?ieia?/ieo iaoe aeioeaio
A.I. Eiieeue/oe

2

Caaaeueia oa?aeoa?enoeea ?iaioe

Aeooaeuei?noue oaie. No?oeoo?i? oa ooieoe?iiaeuei? iniaeeaino? aeeaeo
nenoai ??ciiai ??aiy i?aai?caoe?? iai?aaeai? ia caaacia/aiiy ?o
aaeaioaoe?? aei oiia na?aaeiaeua. ?acoeueoaoe aeine?aeaeaiue, aeeiiaieo
a ?yae? eaai?aoi??e, na?ae/aoue, ui a i?ioean? i?enoinoaaiiy aei oieiaeo
a ee?oeiao ?ineei a?aeaoaathoueny ci?ie iaoaaie?cio, ye? caaacia/othoue
?o aeaeeaaiiy ca iani?eyoeeaeo oaiia?aoo?ieo oiia [I?ioeaiei, Eieioa,
1969; Aeanthe, Inoaiethe, 1973; Ooiaiia, 1979; O?oiiaa, 1984; Sutton et
al., 1992; Ii?aoi, Eiaaeiaiei, 1995]. Aaeaioaoe?y ?ineei aei oieiaeo
aeeth/a? eiiieaen a?io?i?/ieo i?ioean?a na?aae yeeo ci?ie neeaaeo
e?i?ae?a, a?ee?a oa aoaeaaiae?a  [Oioeiaa, 1975; Oeoia oa ?i., 1982;
Iiaeoeeay, oa ?i.,1986; Guy,1990; Aieieeia oa ?i., 1998]. Ni?iae,
ni?yiiaai? ia c’ynoaaiiy iieaeoey?ieo iniia aaeaioaoe?? aei oieiaeo,
i?eaaee aei ?aeaioeo?eaoe?? o ?ineei aai?a, aeni?an?y yeeo ?iaeoeo?oueny
ae??th oieiaeo [Thomashow, 1993; Ii?aoi, Eiaaeiaiei, 1995; Monroy,
Dhindsa, 1995; Gana et al.,1997;]. Oa?aeoa?iith iniaeea?noth i?io?eaiiy
aaeaioaoe?eieo ?aaeoe?e ? iiaiooai?aiiy i?IE ? iaeiie/aiiy niaoeeo?/ieo
a?ee?a i?ae aieeaii oaiia?aoo?, ui ni?e/eiythoue ?iaeoeoe?th ?o neioaco
[Jaglo-Ottosen et al., 1998; Sabehat et al., 1998; Voinikov et al.,
1998]. ?yae aaoi??a ?ciethaaee ? ioa?aeoa?ecoaaee aaie, /ooeea? aei
iecueeeo oaiia?aoo? [Takahashi, Shimosaka, 1997; Ouellet et al., 1998],
i?ioa ooieoe?y a?ee?a ?iaeoeiaaieo iecueeeie oaiia?aoo?aie oa ?o ?ieue o
?icaeoeo no?eeino? ee?oei aei oieiaeo caeeoa?oueny aei e?ioey ia
c’yniaaiith.

I?iaeaia no?eeino? ?ineei aei ae?? aeno?aiaeueieo oaiia?aoo? ia? ia eeoa
oai?aoe/ia, a e i?aeoe/ia cia/aiiy. Na?aae ca?iiaeo eoeueoo? Oe?a?ie ye
ca ieiuath iin?ao, oae ? ca aaeiaei cai?ii ca?ia /?eueia i?noea caeia?
eoeo?oaeca. A?iaeaei?noue oe??? eoeueoo?e a cia/i?e i??? aecia/a?oueny
aieeaii ee?iaoe/ieo oiia, a oiio /ene? oaiia?aoo?ieo. Eoeo?oaeca
iaeaaeeoue aei oaieiethaieo ?ineei, i?i?inoaiiy ian?iiy oa ??no yei?
i?eai?/o?oueny i?e iecueeeo oaiia?aoo?ao. Cao?eiea ?icaeoeo eoeo?oaece
iaaani? i?ecaiaeeoue aei iiaeiaaeaiiy aaaaoaoe?eiiai ia??iaeo,
iiiaaeaiiy ?ineei a ia??iae aeic??aaiiy ca?ia a oiiae iiieaeaieo
oaiia?aoo? ainaie ? ye iane?aeie – aei cieaeaiiy o?iaeath ca?ia, aei
oiai ae o?cieia?/ii ia aeic??aoiai ? oaoiieia?/ii iaye?niiai.

sse a?aeiii, i?i?inoaiiy ianiiiy pineei aeeth/a? ?i?oe?aoe?th iecee
iaoaaie?/ieo i?ioean?a i iaeiei ic iaeaieueo aaaeeeaeo napaae ieo ?
iineeaiiy aia?aii?iaeoeoe??, yea noi?iaiaeaeo?oueny i?aeaeuaiiyi
aeoeaiino? aeeoaiiy ?o oeaiei. I?ciaiiy i?e?iaee oeeo i?ioean?a ?
aaaeeeaei, ine?eueee aiapaaoe/ii iiopaae capiaeeo paaeicothoueny
aeeeth/ii ca paooiie iioioiiaepiaeueieo nenoai aeoioeyoei? i
opainoipiaoei? aiapai? a ee?oei? [Naieoaoiaa, 1990]. Inue /iio aea/aiiy
ooieoe?iioaaiiy oapiaioieo eiiieaenia iioioiiaepie ianiiiy oaieiethaieo
pineei, ui i?i?inoa? ipe iecueeeo oaiiapaoopao, aeicaieeoue ia eeoa
noipioaaoe oyaeaiiy ipi paaoeyoeith aiapaicaaacia/aiiy capiaeeo ianiiiy,
a e aiaeiapaoe e?eoa??? ioe?iee ianiiiy ia caeaoi?noue aei i?i?inoaiiy a
oiiaao oieiaeo oa ?ic?iaeoe oaoiieia?/i? i?eeiie, ni?yiiaai? ia
?ioaineo?eaoe?th ?inoiaeo i?ioean?a iaaani? i?e iecueeeo oaiia?aoo?ao
na?aaeiaeua.

Iioeiaeaeaiiy ?ineei a?ae ii?ic?a ? iniiaiith i?e/eiith c??aeaeaiiy, a a
ie?ai? ?iee ? iiaiiai aeia?caiiy iin?a?a ia cia/ieo ieiuao. I?iaeaia
no?eeino? ?ineei ? aeooeueiith aeey iaoi? e?a?ie, ine?eueee a?eueo i?ae
70% n?euenueeiainiiaea?nueeeo iin?a?a caciathoue aieeao aeno?aiaeueieo
oaiia?aoo?. Aeia?caiiy iin?a?a iceieo ca?iiaeo eoeueoo? a ceiiaee
ia??iae i?eaiaeeoue aei ??ceiai ciaioaiiy o?iaeaeiino? oeeo eoeueoo?,
iaiineoue n?euenueeiio ainiiaea?noao cia/iee caeoie. A?i iia??oo?oueny
cieaeaiiyi o?iaeaeiino? oeo ?ineei, ye? aeaeeaathoue i?ney ae?? ii?ic?a,
aea cia/ii cieaeothoue naith i?iaeoeoeai?noue. Ui??/ii ia Oe?a?i? aeia
a?ae ae?? ii?ic?a a?eueoa 20% iin?a?a iceieo ca?iiaeo eoeueoo?, a a
ie?ai? ?iee aeia?caiiy a?aei?/a?oueny ia cia/ieo ieiuao. Oae eeoa a
ceiiaee ia??iae 1993-1994 ??. ia Oe?a?i? a caaeia?eueiiio noai?
ciaoiaeeeenue eeoa 30% iin?a?a iceieo, c??aeaeaieo a?ae aeia?caiiy aoei
38%, ia?an?yii a?eueo i?ae 30% ieiu iceieo ceae?a. Cia/ieo caeoe?a
caciaei n?euenueea ainiiaea?noai aiane?aeie aeia?caiiy iin?a?a iceii?
ioaieoe? ? a 1997 ?ioe?. Iioeiaeaeaiiy ? caaeaaeue ?ineei
niinoa??aa?oueny oaeiae aaniith aiane?aeie ae?? caii?ice?a. Oae o o?aai?
1999 ?ieo aiane?aeie caii?iceo aei –6, -8 °N caaeioee iin?ae a?a/ee,
oiiao?a oa ?yae ?ioeo oaieiethaieo eoeueoo? iaa?oue ia i?aaei? Oe?a?ie.
Aoee iioeiaeaeai? iin?ae iceii? ioaieoe?, a?a?eae?a eoeo?oaece, y/iaith,
oeoe?iaiai ao?yeo, a oaeiae ianaaeaeaiiy ?yaeo ieiaeiaeo aea?aaieo
?ineei.

Iecueeee ??aaiue no?eeino? aei ae?? ii?ic?a o /ooeeaeo ?ineei iiaea
aooe coiiaeaiee a?aenooi?noth niaoeeo?/ieo aei oieiaeo aai?a aai ae
iayai?noth aeaoaeo?a a ?o i?aai?caoe?? oa ooieoe?iioaaii?. Anoaiiaeaii
iayai?noue oaeeo aai?a ye o no?eeeo, oae ? o iaei no?eeeo aei ii?ico
?ineei, aea o inoaii?o aiie oa?aeoa?ecothoueny iecueeei ??aiai aeni?an??
ca ae?? oieiaeo, ui iiaea aooe ni?e/eiaia iacaeaoi?noth ee?oei oeeo
?ineei ni?eeiaoe neaiae i?i ci?io oaiia?aoo?e ciai?oiueiai na?aaeiaeua
oa o?ainoi?ioaaoe eiai o a?aeiia?aei? ia?aaoaeiae a?io?i?/ieo i?ioean?a,
ye? coiiaeththoue aaeaioaoe?th ?ineei aei ae?? iani?eyoeeaeo oaeoi??a
na?aaeiaeua. Iaei aeine?aeaeaieie ia nueiaiaei? ? ia?aeii? i?ioeane
a?io?i?/ii? aaeaioaoe?? ?ineei aei iecueeeo oaiia?aoo?.

Ee?oeie ?aaaothoue ia ci?io oiia, aea ia?aei? iaoai?cie ni?eeiyoy
neaiaeo oa o?ainoi?iaoe?? eiai a eiie?aoio a?io?i?/io ?aaeoe?th, aein?
ia c’yniaaii. Iniaeeaa cia/aiiy a ?aaeoe?? ee?oei ia ae?th ?yaeo
/eiiee?a na?aaeiaeua iaeaaeeoue ia?aeiiei ?aaeoe?yi, ye? a?aeaoaathoueny
a ?ineeiao i?e ci?i? oaiia?aoo?e na?aaeiaeua [ssai?neay, Eaeeiei, 1984;
ssai?neay, 1990]. Aeine?aeaeaiiy inoaii?o ?ieia, i?iaaaeaii ia eeioeiao
oaa?ei, ia?aeiieeai iieacaee, ui aiae?ieic oinoaoeaeeeiiiceoieo iiae
aei?th oinoieiiace C ? neeaaeiaith /anoeiith ia?aaea/i neaiaeo /a?ac
ieaciaoe/io iaia?aio. Iieacaii o/anoue oinoaoeaeeeiiiceoieo o ?aaeoe??
ee?oei ia ae?th naioea oa aiiiiniioe/iiai oieo. O oie aea /an ia
c’yniaaii ia nueiaiaei? iaoai?cie ni?eeiyooy oa o?ainaeoeoe?? neaiae?a a
ee?oeiao ?ineei i?e ci?i? oaiia?aoo?e na?aaeiaeua.

Aaoi?ii noi?ioeueiaaia eiioeaioe?y ?icaeoeo iiae?e a ee?oeiao ?ineei i?e
ae?? iecueeeo oaiia?aoo? ui noi?iaiaeaeothoueny ci?iaie iaoaaie?cio
iie?oinoaoeaeee?iiceoie?a, oi?ioaaiiyi aoi?eiieo iina?aaeiee?a, ye?
caeaoi? aeeeeeaoe ieceo aaeaioaoe?eieo ci?i iai?aaeaiino? oa
?ioaineaiino? iaoaaie?cio ee?oei, aeeth/athoue iiaiooai?aiiy a?ee?a, c
eeano iieaeoey?ieo oaia?ii?a, neioac e?i?ae?a oa aeoeaaoe?th aeanaoo?ac.
Aeacai? ci?ie ni?yiiaai? ia i?aeo?eiaiiy ?yaeo ooieoe?e ee?oeie ? ia?o
ca ana nenoai o?ainoi?iaoe?? oa aeoioeyoe?? aia?a??.

Iaoa ? caaea/? aeine?aeaeaiiy. Aieiaiith iaoith aeine?aeaeaiue aoei
c’ynoaaoe iniaeeaino? iaoaaie?cio ceae?a i?e ae?? iecueeeo oaiia?aoo? ia
?ineeie, anoaiiaeoe i?e?iaeo aoi?eiieo iina?aaeiee?a, ui i?eeiathoue
o/anoue a ?aaeoe?? ?ineei ia ci?io oaiia?aoo?e na?aaeiaeua; c’ynoaaoe
ia?aeii? ?aaeoe?? iaoaaie?cio iie?oinoaoeaeee?iiceoie?a ia ei?ioei/anio
ae?th iecueeeo oaiia?aoo? oa aeyaeoe aoi?eii? iina?aaeieee, ye? caeaoi?
?iaeoeoaaoe i?ioeane aaeaioaoe?? ?ineei aei ae?? iecueeeo oaiia?aoo?
na?aaeiaeua; anoaiiaeoe caeiiii??iino? ?aaeoe?? ia ci?io oaiia?aoo?e
a?eie neioacoth/i? nenoaie ca?iaee?a ?ineei a i?ioean? ?o ?icaeoeo,
aea/eoe iiaiooai?aiiy a ee?oeiao oa ?o i?oioiiae??yo iie?iaioeae?a, yeei
i?eoaiaii? ooieoe?? iieaeoey?ieo oaia?ii?a; aeine?aeeoe ca oiia in vivo
oa in vitro ?aaoeyoe?th o?ainii?oo aeaeo?ii?a o i?oioiiae??yo ?ineei i?e
ae?? oieiaeo.

Aeey aeinyaiaiiy iinoaaeaii? iaoe iaiao?aeii aoei aea/eoe:

· aieea oieiaeiaiai no?ano ??cii? o?eaaeino? ia iaoaaie?ci
iie?oinoaoeaeee?iiceoie?a ee?oei /ooeeaeo aei ae?? oieiaeo ?ineei,
c’ynoaaoe ?o o/anoue o oi?ioaaii? aoi?eiieo iina?aaeiee?a o?ainaeoeoe??
neaiae?a;

· ?aaeoe?th iiaiooai?aiiy iie?iaioeae?a ca?iaee?a ian?iiy eoeo?oaece ia
ae?th aenieeo oa iecueeeo oaiia?aoo? a i?ioean? ?o ?icaeoeo, ii??aiyoe
aieea aenieeo oa iecueeeo oaiia?aoo? na?aaeiaeua ia iiaiooai?aiiy
iie?iaioeae?a, yeei i?eoaiaii? ooieoe?? iieaeoey?ieo oaia?ii?i?a;

· neeaae oinoie?i?ae?a oa ?o aee?ieo eeneio ??cieo oeaiei a?a?eae?a
eoeo?oaece a i?ioean? ?inoo ?ineei i?e iecuee?e oaiia?aoo??;

· ci?ie oinoie?i?ae?a i?oioiiae??e ceae?a a i?ioean? aaeaioaoe?? ?ineei
aei iecueeeo oaiia?aoo? na?aaeiaeua;

· i?ioean oi?ioaaiiy aeueoa?iaoeaiiai o?ainii?oo aeaeo?ii?a a i?ioean?
?inoo oeaiei ?ineei, anoaiiaeoe aeeaae aeueoa?iaoeaiiai oa oeeoio?iiiiai
aeeoaiiy a caaaeueiee aia?aiiai?i ca?iaee?a eoeo?oaece a i?ioean? ?o
?icaeoeo i?e iecueeeo oaiia?aoo?ao na?aaeiaeua.

Iaoeiaa iiaecia iaea?aeaieo ?acoeueoao?a. Noi?iiaaii eiioeaioe?th
iine?aeiaiino? ?icaeoeo iiae?e a ee?oeiao ?ineei i?e cieaeaii?
oaiia?aoo?e na?aaeiaeua a iniiao yei? iieeaaeaii aia?oa io?eiai? oaeoe
uiaei iaoai?ci?a oi?ioaaiiy aoi?eiieo iina?aaeiee?a a ee?oeiao ceae?a
i?e ae?? oieiaeo oa ?o ?ie? a ?iaeoeoe?? aaeaioaoe?eieo ci?i
iai?aaeaiino? oa ?ioaineaiino? iaoaaie?cio ee?oei, ui aeeth/athoue
iiaiooai?aiiy a?ee?a- i?aaenoaaiee?a iieaeoey?ieo oaia?ii?a, neioac
e?i?ae?a oa aeoeaaoe?th aeanaoo?ac.

Iieacaia ?ieue iieaeoey?ieo oaia?ii?a o i?aeo?eiaii? ooieoe?e ee?oei a
i?ioean? ?inoo ? ?icaeoeo ?ineei i?e iecueeeo oaiia?aoo?ao, caaacia/ai?
no?eeino? ee?oei aei ae?? oieiaeo. ?acoeueoaoe aeine?aeaeaiue
aeeoaeueiiai aia?aiiai?io in vivo oa in vitro aeicaieeee aeyaeoe
iaoai?ci ?aaoeyoe?? iioieo aeaeo?ii?a a i?oioiiae??yo, caaacia/eee
?icaeoie oyaeaiiy i?i oeyoe oi?ioaaiiy aia?aaoe/iiai noaoono ee?oei
ceae?a ca oiia oieiaeiaiai oieo.

Aia?oa aeyaeaii ci?ie ioeo iie?oinoaoeae?e?iiceoie?a i?ae aieeaii
ei?ioei/aniiai ioieiaeaeaiiy ia?inoe?a a?a?eae?a eoeo?oaece oa iceii?
ioaieoe?, iieacaii a?ae?ie?c oinoaoeaeee?iiceoie 4,5-a?oinoaoo aei
?iiceoie 1,4,5-o?eoinoaoo oa aeeaoeeeae?oea?ieo i?e ae?? oieiaeiaiai
oieo. C’yniaaii oa?aeoa? oi?ioaaiiy aeueoa?iaoeaiiai oeyoo o?ainii?oo
aeaeo?ii?a a ee?oeiao ia?enoaie, iieacaii c?inoaiiy eiai o/ano? o
o?ainii?o? aeaeo?ii?a aei eenith o aeic??eeo ee?oeiao ?ineei. Aeyaeaii
aeeiai?eo ci?i aeoeaiino? oeeoio?iiiiai oa aeueoa?iaoeaiiai aeeoaiiy a
i?ioean? ?icaeoeo ca?iaee?a ian?iiy eoeo?oaece i?e iecueeeo oaiia?aoo?ao
na?aaeiaeua. Anoaiiaeaii aeoeaaoe?th aeueoa?iaoeaii? ieneaeace a
i?oioiiae??yo ee?oei no?eeeo aei ae?? oieiaeo e?i?e eoeo?oaece i?ae
aieeaii iecueeeo oaiia?aoo?. Io?eiaii ?acoeueoaoe, ye? na?ae/aoue i?i
aeoeaaoe?th aeueoa?iaoeaiiai aeeoaiiy ca ?aooiie neioaco a?eeo
aeueoa?iaoeaii? ieneaeace. Aeyaeaii a aeaeo?ioi?aoe/ieo niaeo?ao
iiaineioaciaaieo a?ee?a i?oioiiae??e ? oeeoiieacie i?i?inoe?a ioaieoe?
oa eoeo?oaece, ui caciaee aieeao oieiaeo, a?eee, ui a?aeiia?aeathoue
iieaeoey?iei ianai 90 ? 70 eAe c ooieoe?yie iieaeoey?ieo oaia?ii?a, a
oaeiae iie?iaioeaee a ae?eyioe? 17–38 eAe. Aia?oa anoaiiaeaii
iiaiooai?aiiy niaoeeo?/iiai aeey iecueei? oaiia?aoo?e iie?iaioeaeo c
iieaeoey?iith ianith aeecueei 120 eAe a oeaieiao i?i?inoe?a
ii?icino?eeeo ni?o?a iceii? ioaieoe?, neioac yeiai aoa neaai ae?aaeaiee
o ni?o?a c iecueeei ??aiai ii?icino?eeino?. Ca?eueoaiiy ?ioaineaiino?
oinoi?eethaaiiy o o?aeoe?? ye i?oioiiae??aeueiiai, oae ? caaaeueiiai
a?eea i?i?inoe?a eoeo?oaece a?aei?/aii ia?o ca ana ia ae?eyioe? 90 eAe,
ui iiaea a?aeiia?aeaoe ?iaeei? oaia?ii?a cpn90.

I?aeoe/ia cia/aiiy io?eiaieo ?acoeueoao?a. Aeyaeai? iniaeeaino? ?aaeoe??
?ineei ia ci?io oaiia?aoo?e na?aaeiaeua oi?ioaaiiyi aoi?eiieo
iina?aaeiee?a, iiaiooai?aiiyi a?ee?a, i?aaenoaaiee?a iieaeoey?ieo
oaia?ii?a, ?aaoeyoe??th ooieoe?e i?oioiiae??e, ye ? c’ynoaaiiy
a?aeiia?ae? ia aeno?aiaeuei? oaiia?aoo?i? oiiae ca?iaee?a ?ineei
iiaeeaeththoue oyaeaiiy i?i i?e?iaeo ii?icino?eeino?, iae?aneththoue
oeyoe ea?oaaiiy oieiaei- oa ii?icino?ee?noth ca aeiiiiiaith
iieaeoey?ii-aaiaoe/ieo i?aeoiae?a ?c iiceoe?e ?aaoeyoe?? aeoeaiino?
aeueoa?iaoeaiiai aeeoaiiy oa neioaco oieiaei-/ooeeaeo a?ee?a.
Cai?iiiiiaaii e?eoa??? a?aeai?o no?eeeo aei oieiaeo a?a?eae?a
eoeo?oaece.

Io?eiai? ?acoeueoaoe iiaeooue aooe aeei?enoaii? a eo?nao eaeoe?e c
a?io?i??, a?iaia?aaoeee, o?c?ieia?? oa naeaeoe?? ?ineei a AOCao
Oe?a?ie.

Iniaenoee aianie caeiaoaa/a. Caeiaoaa/ iniaenoi ?ic?iaea ?iai/?
a?iioace, nieaioaaa oa i?ia?a aenia?eiaioaeuei? aeine?aeaeaiiy.
Cae?eniea aiae?c, ?ioa?i?aoaoe?th ? ocaaaeueiaiiy iaea?aeaieo
?acoeueoao?a.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. Aeena?oaoe?eia
?iaioa aeeiioaaeanue a ?aieao athaeaeaoieo oai a?aeae?eo o?c?ieia??
no?eeino? ?ineei ?O?A IAI Oe?a?ie “Aea/aiiy ?ie? a?ee?a oaia?ii?a oa
oinoaoeaeee?iiceoieo o no?eeino? ceae?a aei aeno?aiaeueieo oaiia?aoo?;
aeine?aeaeaiiy aeieiai-o?c?ieia?/ieo iniaeeainoae ii?icino?eeino? ??cieo
ca aaiioeiaie ni?o?a iceieo ceae?a” (0198U009039), «?aaoeyoe?y aeeoaiiy
oaieiethaeaeo ?ineei a i?ioean? i?i?inoaiiy i?e iecueeeo oaiia?aoo?ao”
0199U000020, “Aeine?aeeoe neioac a?ee?a oa ?ieue e?i?ae?a ye
ianaiaeaea??a o ca`yceo ?c o?c?ieia??th oieiaei- oa ii?icino?eeino?
ceae?a a i?ioean? iioiaaiaco ?ineei” (0198U007708), oa i?iaeo?a Oiiaeo
ooiaeaiaioaeueieo aeine?aeaeaiue Oe?a?ie: “Aeine?aeaeaiiy ?ie?
a?ee?a-oaia?ii?a i?oioiiae??e ca?iaee?a eoeo?oaece i?e ?o ?icaeoeo a
oiiaao ae?? aenieeo oa iecueeeo oaiia?aoo?»(5.4/336).

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. Iniiai? iaoeia? ?acoeueoaoe aoee
i?aaenoaaeai? ia OOV THa?eaei?e I?aeia?iaei?e ia?aae? ? neiiic?oi? ia
oaio:»Aeine?aeaeaiiy a?iaaiaco, no?oeoo?e oa ooieoe?e oioineioaoe/iiai
aia?aoo o ca’ceo c ia?aoai?aiiyi niiy/ii? aia?a?? (Aieaa??y, Oieaoo?i,
1988), I?aeia?iaei?e eiioa?aioe?? » Aeinyaiaiiy oa ia?niaeoeae ?icaeoeo
e??ia?ieia?? ? e??iiaaeeoeeie» (Oa?e?a, 1988), II c’?cae? Ananithciiai
oiaa?enoaa o?c?ieia?a ?ineei (I?inuee, 1990), Ananithci?e eiioa?aioe??
«Aaiaoe/i? iaoai?cie no?eeino? ?ineei aei iani?eyoeeaeo oaeoi??a
na?aaeiaeua » (??eoonuee, 1991 ?.), II c’?cae? Oe?. Oiaa?enoaa
o?c?ieia?a ?ineei (Ee?a, 1993), III c’?cae? Ana?in?enueeiai oiaa?enoaa
o?c?ieia?a ?ineei (Naieo-Iaoa?ao?a, 1993), I?aeia?iaei?e ia?aae?
«Aeooaeuei? ieoaiiy o?c?ieia?? ?ineei a aniaeo? aeieia?/ieo i?iaeai
Oe?a?ie» (*a?i?aoe?, 1995 ?.), I c’?cae? Oe?a?inueeiai oiaa?enoaa
e??ia?ieia?? ? e??iiaaeeoeeie (Oa?e?a, 1995 ?.), I?aeia?iaeiiio
neiiic?oi? «Aeeoaiiy ?ineei: o?c?ieia?/i? oa aeieia?/i? aniaeoe
(Neeoeaea?, 1995), I?aeia?iaeiiio neiiic?oi? «No?an ? anei?eyoe?y
iai?aai?/iiai acioo» (Iineaa, 1996 ?.), 8-io (Aaeuea?y, 1992?.), 10-io
(?oae?y, 1996 ?.) oa 11-io (Aieaa??y, 1998 ?.) eiia?anao Oaaea?aoe??
?a?iiaenueeeo oiaa?enoa o?c?ieia?a ?ineei, I?aeia?iaei?e eiioa?aioe??
«Aieea no?an?a na?aaeiaeua ia iieaeoey?io a?ieia?th ?ineei” (Iieueua,
1997 ?.), I?aeia?iaeiiio nai?ia?? «Ioi?iia aeiae?eey: no/ani?
aeine?aeaeaiiy a aeieia?? ? i?e?ia?ieia??” (Oaeai?iae, 1997), XVIII
Eiia?an? oiaa?enoaa o?c?ieia?a ?ineei Neaiaeeiaa?? (Oaaoe?y,1997), IV
I?aeia?iaeiiio neiiic?oi? “?aaeoe?y iaoaaie?cio ?ineei ia caa?oaeiaiiy
iia?o?y ? aeiaaeuei? ci?ie” (I?aea?eaiaee, 1997), iaoeiaiio nai?ia??
Oi?aa?neoaoo ooaoo I?aaeaia Aeaeioa (NOA, 1999).

Ioae?eaoe??. Iaoeia? ?acoeueoaoe ii oai? aeena?oaoe?? iioae?eiaaii a 37
ae?oeiaaieo ?iaioao, ui aeeth/a? 19 noaoae, a oiio /ene? 7 — aac
ni?aaaoi??a, iioae?eiaaieo a i?ia?aeieo iaoeiaeo aeaeaiiyo.

Ia’?i oa no?oeoo?a ?iaioe. Aeena?oaoe?y, aeeeaaeaia ia 291 noi??ieao
iaoeiiieniiai oaenoo, neeaaea?oueny ?c anooio, 5 aeaa, aeniiae?a, nieneo
aeei?enoaii? e?oa?aoo?e. ?iaioa i?noeoue 25 iaethie?a ? 20 oaaeeoeue.
A?ae?ia?ao?/iee nienie neeaaea? 665 e?oa?aoo?ieo aeaea?ae, ?c ieo 614 ia
?iicaiieo iiaao.

Iniiai? iieiaeaiiy, ye? aeiinyoueny ia caoeno.

1. Iaoaaie?ci iie?oinoaoeae?e?iiceoie?a ?aaao? ia ae?th iecueeeo
oaiia?aoo? oi?ioaaiiyi aoi?eiieo iina?aaeiee?a o?ainaeoeoe?? neaiae?a o
ee?oeiao ?ineei.

2. Ia ?aii?o aoaiao ?icaeoeo ca?iaeee ?ineei ee?oeie ?aae?cothoue
i?ia?aio aeni?an?? aaiiio ? ia ci?iththoue neioac a?ee?a i?ae aieeaii
aenieeo ? iecueeeo oaiia?aoo?.

3. Iecuee? oaiia?aoo?e ?iaeoeothoue iiaiooai?aiiy iie?iaioeae?a ?c
?iaeei AOO70, AOO90 ? iecueeiiieaeoey?ieo a?ee?a (17-35 eAe) o ee?oeiao
ceae?a ? niaoeeo?/iiai aei ae?? iecueeeo oaiia?aoo? a?eea ?c
iieaeoey?iith ianith 120 eAe eeoa o ee?oeiao iceii? ioaieoe?.

4. I?aeo?eiaiiy aia?aaoe/iiai aiiainoaco ee?oei ?ineei ca oiia iecueeeo
oaiia?aoo?ao a?aeaoaa?oueny oeyoii aeoeaaoe?? o?ainii?oo aeaeo?ii?a
/a?ac aeueoa?iaoeaiee oeyo ?o i?oioiiae??e.

Iaoa??aee oa iaoiaee

Ia’?eoe aeine?aeaeaiiy. O aeine?aeaeaiiyo aoee aeei?enoai? ca?iaeee,
eieaiioee? ? eenoee ao?ieueiaaieo i?i?inoe?a e?i?e ? a?a?eae?a
eoeo?oaece oa iceii? ioaieoe?, ye? ae?iuoaaee a oa?iinoao? i?e 25 °N.
Aaeaioaoe?ei? ci?ie iaoaaie?cio iceii? ioaieoe? aea/aee i?e aeo?eioaaii?
a iecueeioaiia?aoo?i?e eaia?? i?e oaiia?aoo?? +3 °N, eoeo?oaece – i?e
+12- +14iN oo?eaae?noth aei o?ueio oeaei?a. Ei?ioei/anio ae?th oieiaeo
ia oeaieie eoeo?oaece (+10 °N ) oa iceii? ioaieoe? (+3 °N) noai?thaaee o
o?en-INl (10 iI, pH 7,2) aooa?? i?ioyaii 5 — 60 oaeeei.

Iaoiaee aeine?aeaeaiiy neeaaeo e?i?ae?a ?ineei. [32P]-i?oioinoao
aaiaeeee a oeaieie eieaiioee?a ? eenoe?a eoeo?oaece oa iceii? ioaieoe?
oeyoii iieoaaoei? o ?ic/ei? c aeoeaiinoth 6,6 IAe ia 1 a oeaiei i?ioyaii
18 aiaeei i?e +25 °N. ?aae?iaeoeaiee oinoi? aaiaeeeany oeyoii aaeooiii?
iioieueo?aoei? i?ioyaii 5 oaeeei aeey aieueo oaeaeeiai i?iieeiaiiy o
oeaieie [Raison et al., 1980]. Ia?aae ioieiaeaeaiiyi ?ineeie aeo?eioaaee
ia na?oe? i?ioyaii 30 oaeeei i?e +25°N aeey caiiaiaaiiy iiai/iiai aieeao
na?oea. Oienaoeith iaoa?iaeo caeienithaaee ?iaeeei acioii.

Aeaeieaiiy caaaeueieo eiiiaeia i?iaiaeeee ca iaoiaeii Oie/a (1957).
Aeno?aeoeith i?iaiaeeee noiiooth oei?ioi?i-iaoaiie (2:1 ca ia’?iii ) c
?ic?aooieo 21 ie noiioi ia 1 a oeaiei. Oei?ioi?iio o?aeoeith eiiiaeia
oia?thaaee o no?oii acioo oa ?ic/eiyee o noiioi oei?ioi?i-iaoaiie (2:1).

Aeno?aeoe?y iie?oinoaoeaeee?iiceoie?a. Aeey aeae?eaiiy
iieioinoaoeaeeeiiiceoieia iaie aoea iiaeeo?eiaaia iaoiaeeea *i oa *ai
[Crookston et al., 1974]. Inaae oeaiei, yeee caeeoeany ia oieueo?i iiney
aeno?aeoei? caaaeueieo eiiiaeia, ia?iaeyee 0,05 I AAeOA i?ioyaii 20 oa.
oa ia?aiineee inaae c o?eueo?o o ?ic/ei oei?ioi?i-iaoaiie-eHCl
(200:100:1) ? i?iaiaeeee aeno?aeoeith i?e 37 °N i?ioyaii 1 aiaeeie c
ia?iiaee/iei iii?ooaaiiyi. I/enoeo aiae aiaei?ic/eiieo eiiiiiaioia
caeienithaaee oeyoii aeiaeaaaiiy 0,2 ia’?io 1i HCl c iiaeaeueoei
oeaio?eooaoaaiiyi 10 oaeeei i?e 3000 g. Iiaa?oiaaee oa? i?iieaaee o?e/?
?ic/eiii iaoaiie-1iHCl (1:1), eiaeiiai ?aco oeaio?eooaoth/e c?acee 10
oaeeei i?e 3000 g. Oei?ioi?iiee aeno?aeo oia?thaaee o no?oii acioo oa
?ic/eiyee o noiioi oei?ioi?i-iaoaiie (2:1).

Oiieioa?iaa o?iiaoia?ao?y e?i?ae?a. ?iciiaeie eiiiaeia i?iaiaeeee ia
neeieaaaeaaeo ieanoeiao «Sorbfil» IAOO NOO-1A (AI «Ni?aiieiia?»)
iaoiaeii aeaii??ii? o?iiaoia?ao??: 1 iai?yiie oei?ioi?i-iaoaiie-7i
aiiae (60:30:4), 2 iai?yiie oei?ioi?i-iaoaiie-ioeoiaa eeneioa-aiaea
(170:25:25:6) , oinoieiiiaee- o nenoaiao ?ic/eiieeia: 1) aoeaoii-ioeoiaa
eeneioa-aiaea (100:2:1); 2) oei?ioi?i-iaoaiie-aiaea (65:25:4) [Roughan,
Slack, 1982].

Aiaeic iieioinoaoeaeeeiiiceoieia i?iaiaeeee ia iii?aaiiaaieo 1%
ienaeaoii eaeith (75 n) neeieaaaeaaeo ieanoeieao «Sorbfil» IAOO NOO-1A .
Aeey ?icaeieaiiy aoea ia?aia nenoaia ?ic/eiieeia oei?ioi?i-iaoaiie-4i
aiiae (9:7:2) [Cao?aeia e ae?., 1990].

Aaoi?aae?ia?ao?y oa noeeioeeyoe?eiee ?aooiie. Iiney ?icaeieaiiy o
iaaaaeaieo aeua nenoaiao ?ic/eiieeia ieanoeiee c [32?]-ii/aieie
oinoieiiiaeaie oa oinoaoeaeeeiiiceoieueiith o?aeoei?th aiaeicoaaee
iaoiaeii aaoi?aaeiia?aoi? ia ?aioaaiianueeee ieiaoei. Eieueeinio
aeoeaiinoue c?ace?a aecia/aee iaoiaeii noeeioeeyoe?eiiai ?aooieo ia
??aeeiiiio noeeioeeyoi?i Rac-Beta (Oiieyiaeiy) c noeeioeeyoeieiith
?iaeeiith AEN-7.

E?euee?nia aecia/aiiy caaaeueieo oinoie?i?ae?a. Eieueeinoue caaaeueieo
oinoieiiiaeia oa ?o ie?aieo eeania aecia/aee ii iai?aaii/iiio oinoi?o ca
iaoiaeii Ai?? i Eiia [Bowyer,King,1977].

Aaci??aeeiia o?iiaoia?ao?y aee?ieo eeneio. Iaoiae iniiaaiee ia
ia?aaaaeaiii aee?ieo eeneio o eaaei iaoeeiai ao??e c iiaeaeueoei
aecia/aiiyi ?o e?eueeino? aaci??aeeiiith o?iiaoia?ao??th [Neiye e
ae?.,1976]. E?euee?noue aee?ieo eeneio aecia/aee iaoiaeii aioo??oiuei?
ii?iae?caoe?? [Aieueaa?o, Aeaaea?aaoc,1990]. sse?nio ?aeaioeo?eaoe?th
o?iiaoia?ai i?iaiaeeee ca noaiaea?oiei ?ic/eiaie iaoeeiaeo ao???a
aee?ieo eeneio oa ca /anii oo?eiaiiy [Iae-Iae?, Aiiaeei,1970].

E?euee?nia aecia/aiiy ?iaeea?aeoaeueieo iiiceoie-oinoaoia.
O?aeoe?iioaaiiy oinoiiiiceoieia cae?enithaaee iaoiaeii ??aeeiii?
?iiiiai?iii? o?iiaoia?ao?? [Bartlet 1982]. Ci?ie ai?noo ?iiceoieoinoao?a
aecia/aeeny ca ??aiai iai?aaii/iiai oinoi?o o a?aeiia?aei?e i?ia?
aethaioo i?ney ?iiiiai?iii? o?iiaoia?ao??.

Aeine?aeaeaiiy iiaiooai?aiiy iie?iaioeae?a. Aeine?aei? ? eiio?ieuei?
i?i?inoee iiae??aithaaee ia oiaoi/ee aeiaaeeiith 4–5 ii oa ?ieoaoaaee o
?ic/ei? 35S-iao?ii?io (ieoiia aeoeai?noue 80–100 ieE?/ie) aai noi?o?
14C-ai?iieeneio (ieoiia aeoeai?noue 20–60 ieE?/ie) i?ioyaii 3–5 aiae.
i?e 24°C. O ?ioeo na??yo aenia?eiaio?a aeeth/aiiy i?oee i?iaiaeeee i?e
oaiia?aoo?? +3°C i?ioyaii inoaiiuei? aeiae ae?? oieiaeo. I?/aiiy
32P-i?oioinoaoii (ieoiia ?aae?iaeoeai?noue aeecueei 200 ieE?/ie)
i?iaiaeeee a oiiaao ae?? oaiia?aoo?iiai /eiieea. Ia?aae iiaeaeueoeie
iia?aoe?yie ?ineeiiee iaoa??ae a?aeieaaee a?ae ?aae?iaeoeaii? i?oee.
*anoeia iaoa??aeo aoea aeei?enoaia aeey iaea?aeaiiy noia?ii? o?aeoe??
iie?iaioeae?a i?i?inoe?a, c ?aooe io?eioaaee o?aeoe?th Io. Caaaeueiee
a?eie aeno?aaoaaee iiaea?eiei aooa?ii Eaiie?, io?eiai? c?acee (i?ney
1-oaeeeiiiai i?ia??aaiiy ia aiaeyi?e aai? i?e 100 oN) ?icae?eyee
iaoiaeii aeene aeaeo?ioi?aco o iie?ae?eeai?aeiiio aae? c ania?a?iiaei
aooa?ii o i?enooiino? SDS. Aaoia?aoe io?eioaaee ia ?aioaai?anuee?e
ie?aoe? oeio ?I-A c aenooaieo aaeae i?ney oa?aoaaiiy ?o eiei?aeiei
?ic/eiii eoian? R250 (Reanal, Oai?ueia). sse iieaeoey?i? ia?ea?e
aeei?enoiaoaaee iaa?? «Rainbow» (Amersham, Aaeeeia?eoai?y) ?c 7 a?ee?a c
iieaeoey?ieie ianaie a?ae 14,3 aei 200 eAe.

Aeeoaeueiee aia?aiiai?i ?ineei. Aeey aeine?aeaeaiiy aaciiai?io oeaiei
eoeo?oaece aeei?enoiaoaaee iiey?ia?ao II-105 (Oai?ueia) oa cae?eoee
ieaoeiiaee aeaeo?iae oeio Eea?ea. Aei??thaaiiy i?iaiaeeee o na?aaeiaeu?,
yea i?noeoue NaNl2 (0,2%) oa o?en-INl aooa? (10iI). Oaiia?aoo?a eii??ee
i?aeo?eioaaeany ca aeiiiiiaith oeueo?aoa?iinoaoo, c oieiaeeeueiei
aa?aaaoii, ui caaacia/oaaa oaiia?aoo?o a?ae 0 aei 30iN. ?ineeie ia?aae
aei??aie ?ic??caee ca aeiiiiiaith eaca ia ae?eyiee caaoe?oee 2 ii oa
aeiaaeeiith 3-4 ii. Ia?aae aei??thaaiiyi aaciiai?io oeaieie aeo?eioaaee
o ?ic/ei? ?ieoaaoe??, aai ae a aa???iaaieo ?ic/eiao a?aeiia?aeieo
?ia?a?oi??a i?ioyaii 20 oaeeei. Aeey aeieoaaiiy o?ainii?oo aeaeo?ii?a
/a?ac oeeoio?iiiee eaioetha aeei?enoiaoaaee aceae iao??th (5 iI),
aeueoa?iaoeaio ieneaeaco aeieoaaee ?ic/eiii aaica?ae?ienaiiai? eeneioe
(10 iI). Aeeaae ieneaeac iai?oioiiae??aeueiiai iioiaeaeaiiy a caaaeueia
iiaeeiaiiy eenith aecia/aee ye caeeoeiaa aeeoaiiy i?ney noi?niiai
aaaaeaiiy a oeaieie ei?ai?a eoeo?oaece aceaeo iao??th oa
aaica?ae?ienaiiai? eeneioe [Azcon-Bieto et. al, 1983]. Aeoeai?noue
oeeoio?iiiiai oeyoo i?oioiiae??e ee?oei ei?aiy aecia/aee, ye ??cieoeth
i?ae ?ioaineaiinoyie iiaeeiaiiy eenith oeaieiaie a i?enooiino?
aaica?ae?ienaiiai? eeneioe oa caeeoeiaiai aeeoaiiy. Iioaioe?aeueio
aeoeai?noue aeueoa?iaoeaiiai oeyoo ?icaeyaeaee ye ?ioaineai?noue
aeeoaiiy a i?enooiino? aceaeo iao??th i?ney a?ae?aooaaiiy caeeoeiaiai
aeeoaiiy. Caeiyo?noue aeueoa?iaoeaiiai oeyoo ?ic?aoiaoaaee ca
a?aeiioaiiyi nooiai? ?ia?aoaaiiy aeeoaiiy aaica?ae?ienaiiaith
eeneioith aei nooiai? no?eeino? aeeoaiiy a i?enooiino? aceaeo iao??th
(Lambers et.al.,1983). I?oioiiae??? io?eioaaee ca aeiiiiiaith iaoiaeo
aeeoa?aioe?aeueiiai oeaio?eooaoaaiiy ca iienaiith iaoiaeeeith (E?aaaoe,
Aieieeia, 1984).

?AAEOe?ss IAOAAIE?CIO IIE?OINOAOEAeEE?IICEOIE?A IA Ae?TH OIEIAeIAIAI
NO?ANO

Aeey aeine?aeaeaiiy ae?? oieiaeiaiai oieo ia iaoaaie?ci
iie?oinoaoeaeee?iiceoie?a aoea ia?aia oaiia?aoo?a +10°N ia o?e
i?aenoaa?, ui oey oaiia?aoo?a ia aeeeeea? iioeiaeaeaiiy oeaiei ?ineei
eoeo?oaece ? ? ciiith oaiia?aoo?iiai ae?aiaciio, a yeiio aeaia eoeueoo?a
caeaoia i?aeaeuoaaoe no?ee?noue aei ae?? iecueeeo oaiia?aoo?. Naia i?e
oe?e oaiia?aoo?? a ee?oeiao ii/eiathoueny i?ioeane, ye?
oa?aeoa?ecothoueny oa?i?iii «aaeaioaoe?y aei oieiaeo». A i?ioean? ?inoo
i?e oeeo oaiia?aoo?ao a ee?oeiao ?ineei a?aeaoaathoueny aaeaioeai? ci?ie
iaoaaie?cio.

Aoei aeine?aeaeaii aieea oieiaeiaiai oieo (5 — 20 oa.) ia iaoaaie?ci
e?i?ae?a eieaiioee?a 4-aeiaiaeo ao?ieueiaaieo ia?inoe?a eoeo?oaece.
Oeaieie eieaiioee?a ? iaeii??aeieie, caaa?oeee ?inoia? i?ioeane ? a
i?e?iaeieo oiiaao caciathoue aieeao oieiaeiaiai no?ano o ia?oo /a?ao.

Aeine?aeaeaiiy io?eiaieo o?iiaoia?ai aeyaeei aenieee ??aaiue aeeth/aiiy
oinoi?o o oinoaoeaeeeaoaiieai?i (OAA) oa oinoaoeaeeeoie?i (OO). Aeaui
ieae/ee ??aaiue aeeth/aiiy oinoi?o aoei a?aei?/aii o o?aeoe??
oinoaoeaeeeae?oea?ieo (OA) ii??aiyii c OAA oa OO. Aeineoue iii?oiee
??aaiue ?aae?iaeoeaiino? ca?a?no?iaaii o o?aeoe?yo oinoaoeaeeena?eio
(ON), oinoaoeaeee?iiceoieo (O?) oa oinoaoeaeii? eeneioe (OE). Na?aae
o?aeoe?? iie?oinoaoeaeee?iiceoie?a aeyaeaii aeeth/aiiy i?oee aei
oinoaoeaeee?iiceoieoinoaoo (O?O) oa oinoaoeaeee?iiceoiea?oinoaoo (O?O2).

??cea ioieiaeaeaiiy eieaiioee?a eoeo?oaece i?ioyaii 5 — 20 oaeeei i?e
+10°N ia aeeeeeaei aeinoia??ii? ci?ie aeoeaiinoae o o?aeoe?yo caaaeueieo
oinoie?i?ae?a (?en. 1), a oie /an ye na?aae iie?oinoaoeaeee?iiceoie?a
a?aei?/aii ??cea cieaeaiiy aeoeaiino? O?O oa O?O2 /a?ac 10 oaeeei ae??
iecueei? oaiia?aoo?e (?en. 2), yea noi?iaiaeaeoaaeinue c?inoaiiyi ??aiy
?iiceoie 1,4,5-o?eoinoaoo (?ic(1,4,5)O3) (?en. 3). Iiaeaeueoa
aeo?eioaaiiy oeaiei ?ineei i?e oaiia?aoo?? +10°N i?ioyaii 20 oaeeei
iaoiiaeei c?inoaiiy e?eueeino? aeeth/aiiy i?oee o O?O oa O?O2, ui aoei
i?aeoaa?aeaeaii aeaieie noeeioeeyoe?eiiai ?aooieo (?en. 2). O
iiaeaeueoeo aeine?aeaeaiiyo ae?? iecueeioaiia?aoo?iiai no?ano ia
iaoaaie?ci oinoie?i?ae?a oa iie?oinoaoeaeee?iiceoie?a o eenoeao oa
eieaiioeeyo 4-aeiaiaeo ao?ieueiaaieo ia?inoe?a eoeo?oaece oeaieie ?ineei
ioieiaeaeoaaee i?ioyaii 10 oaeeei i?e oaiia?aoo?? +10°N. Aiae?c
io?eiaieo o?iiaoia?ai iieacaa, ui ? a eenoeao ? a eieaiioeeyo
aeine?aeaeaieo oeaiei a?aeaoaaeiny ??cea cieaeaiiy e?eueeino? O?O oa
O?O2 i?e ae?? oieiaeo +10°N i?ioyaii 10 oaeeei.

E?euee?noue caaaeueieo oinoie?i?ae?a caeeoaeanue o iaaeao eiio?ieueieo
cia/aiue .

A aeine?aeaeaiiyo ?aaeoe?? iaoaaie?cio iie?oinoaoeaeee?iiceoie?a ia
ae?th oieiaeiaiai no?ano aoei aeei?enoaii oeaieie ia?inoe?a iceii?
ioaieoe?. O yeino? no?aniaiai /eiieea o aeaiiio aeiaaeeo aoea aea?aia
oaiia?aoo?a +5°N (oaiia?aoo?a, yea ia aeeeeea? iioeiaeaeaiue, aea
?iaeoeo? aaeaioeai? ia?aaoaeiae o ee?oeiao ?ineei ioaieoe?). Io?eiai?
?acoeueoaoe na?ae/aoue, ui aaea /a?ac 5 oaeeei ae??
iecueeioaiia?aoo?iiai no?ano niinoa??aa?oueny ciaioaiiy e?eueeino? O?O
oa O?O2 o oeaieiao o?ueio aeiaiaeo ao?ieueiaaieo eieaiioee?a ia?inoe?a
ioaieoe? (?en. 4). Aeanyoe oaeeeiia aeniiceoe?y i?ecaiaeeea aei
iiaeaeueoiai ?iciaaeo ie?aieo eiiiiiaio?a iie?oinoaoeaeee?iiceoie?a, ia
iecueeei aoa ??aaiue oeeo niieoe ? /a?ac 20 oaeeei ae?? oieiaeo.
E?euee?noue caaaeueieo oinoie?i?ae?a oeaiei ia?inoe?a iceii? ioaieoe?
aeinoia??ii ia ci?ithaaeanue ia i?ioyc? anueiai aenia?eiaioo.

Io?eiai? ?acoeueoaoe na?ae/aoue i?i /ooeea?noue iaoaaie?cio O?O oa O?O2
aei ei?ioei/aniiai ioieiaeaeaiiy oeaiei ?ineei ia a?aei?io a?ae
iaoaaie?cio iniiaieo oinoie?i?ae?a. A?aeiii, ui O?O2 o ee?oeiao ?ineei
a?ae?ie?co?oueny i?ae ae??th oinoie?iace N (oinoi?iiceoeaeace) aei
?ic(1,4,5)O3 oa AeAA. Io?eiai? ?acoeueoaoe aeacothoue ia oa, ui i?e ae??
oieiaeiaiai no?ano a?aeaoaa?oueny aeoeaaoe?y O?O2-niaoeeo?/ii?
oinoie?iace N.

Ia i?eeeaae? ae?? ?yaeo /eiiee?a (ina?oeaiiy, ci?ia iniioe/iiai oeneo,
ae?y iaoiaaiieo ae?n?oi??a oa noi?ani??a) c’yniaaii, ui ?ic(1,4,5)O3
aieeaa? ia ia?a?iciiae?e ai?noo ?ii?a Na2+ o eiiia?oiaioao ee?oeie,
cie?aia, iaoiiaeth? aeo?ae Na2+ c aaeoie?. A?aeiii, ui Na2+ ? aoi?eiiei
ianaiaeaea?ii, yeee aaciina?aaeiuei caeo/a?oueny aei aaaaoueio
neaiaeueieo i?ioean?a [Bush, 1995]. Iieacaii, ui i?aeaeuaiiy ??aiy
eaeueoe?th o a?aeiia?aeue ia ae?th oieiaeo a?aeaoaa?oueny ca ?aooiie ye
ciai?oiuei ee?oeiiiai Na2+, oae ? ?ic(1,4,5)O3-?iaeoeiaaiiai ca?eueiaiiy
eaeueoe?th ?c aaeoie? [Knight et al., 1996]. Iaaaaeai? aeai? ?acii ?c
?acoeueoaoaie i?iaaaeaieo aeine?aeaeaiue na?ae/aoue i?i iiaeeeao o/anoue
oinoaoeaeee?iiceoieueii? nenoaie o ni?eeiyoo? iecueeioaiia?aoo?iiai
aieeao oa caioneo ?aaeoe?e ee?oeiiiai a?aeaoeo ia ae?th oieiaeo.
I?aeaeuaiiy aeoeaiinoae O?O ? O?O2 a i?ioean? iiaeaeueoiai aeo?eioaaiiy
oeaiei eoeo?oaece ia oieiae? iiaea na?ae/eoe i?i aeoeaaoe?th O?4- ?
O?O5-e?iac, ye? oi?iothoue O?O oa O?O2 ? eieae?ciaai? o oeeoineaeaoi?e
o?aeoe?? ieaciaoe/ii? iaia?aie [Tan, Boss, 1992].

?IEUe E?I?Ae?A A AAeAIOAOe?? EE?OEI ?INEEI AeI IANI?EssOEEAEO OIIA
NA?AAeIAEUA.

Ci?ie neeaaeo oinoie?i?ae?a ee?oei ?ineei i?e cieaeaii? oaiia?aoo?e
na?aaeiaeua

Aaeaioaoe?y ia?inoe?a eoeo?oaece aei ae?? iecueei? oaiia?aoo?e
noi?iaiaeaeoaaeanue ca?eueoaiiyi caaaeueii? e?eueeino? oinoie?i?ae?a o
an?o aeine?aeaeaieo i?aaiao ?ineei. O eenoeao noia oinoie?i?ae?a c?inea
o 2,5 ?ace, oiae? ye o eieaiioeeyo oa iacieioeeyo — o 1,5 oa 1,2 ?ace,
a?aeiia?aeii. ?icaeoie oieiaeiaeo iioeiaeaeaiue niinoa??aaany
caeaa?eueoiai ia iacieioeeyo ia?inoe?a oa aeayeith i??ith — ia
eieaiioeeyo. Eenoee ye caaa?oiaaieo, oae ? iacaaa?oiaaieo ?ineei ca
aeaieo oiia ioieiaeaeaiiy aoee no?ee?oeie aei ae?? oieiaeo. Oaeei /eiii,
?ioaineai?noue iaeiie/aiiy oinoie?i?ae?a a ie?aieo i?aaiao ?ineei
a?aeiia?aeaea nooiaith ?o no?eeino? aei ae?? iecueei? oaiia?aoo?e, ui
na?ae/eoue i?i o/anoue oeeo niieoe o ?icaeoeo no?eeino? aei
ioieiaeaeaiiy.

Aeeiai?ea neeaaeo e?i?ae?a a i?ioean? aaeaioaoe?? ia?inoe?a ceae?a aei
iecueei? oaiia?aoo?e na?aaeiaeua. Iniiaieie eiiiiiaioaie oinoie?i?ae?a
i?oioiiae??aeueieo iaia?ai iceii? ioaieoe? ? OO,OAA, OA ? ea?ae?ie?i?i
(EE). ?aeaioeo?eiaaii oaeiae O?, ? OE (oaae.1). Aaeaioaoe?y ?ineei aei
iecueeeo oaiia?aoo? noi?iaiaeaeo?oueny iaeiie/aiiyi OO. OAA ? OA o
i?oioiiae??yo iaaii?a iceii? ioaieoe?, oae a i?ioean? ?inoo ia i?ioyc?
ia?oeo 7 ae?a i?e 5 °C e?euee?noue OO ca?eueoo?oueny ia 28%, OAA ia 22%
? OA ia 17% ii??aiyii ?c eiio?ieai.

Oaaeeoey 1

Aeeiai?ea iaeiie/aiiy oinoie?i?ae?a i?oioiiae??e ia?inoe?a iceii?
ioaieoe? ni?oo Ie?ii?anueea 808 a i?ioean? aaeaioaoe?? ?ineei aei
iecueeeo oaiia?aoo? na?aaeiaeua (oinoie?i?aee a iI ia ia a?eeo )

Oaiia?aoo?a ae?iuoaaiiy ?ineeiOinoie?i?aeeEiio?ieue,25 °C7 aei?a i?e

5 °C7 aei?a i?e 5 °C oa 7 aei?a i?e

2 °C7 aei?a i?e 5 °C,

7 aei?a i?e 2 °C oa 5 aei?a i?e-2 °COinoaoeaeee-oie?i81,0 ± 2,0103,5 ±
2,5112,4 ± 2,2110,5 ± 2,8Oinoaoeaeee-aoaiieai?i65,5 ± 2,580,2 ± 0,888,0
±1,088,4 ± 2,0Oinoaoeaeee-ae?oea?ei21,0 ± 0,524,5 ± 0,525,6 ±1,026,0 ±
0,7Ea?ae?ie?i?i

10,2 ± 0,711,2 ± 0,412,0 ± 0,811,5 ± 0,5Oinoaoeaeee-?iiceoie7,2 ± 0,38,6
± 0,58,5 ± 0,28,8 ± 0,3Oinoaoeaeia eeneioa4,5 ± 0,24,2 ± 0,54,4 ± 0,34,7
± 0,5Iiaeaeueoee ??no ?ineei i?e 2 °C iaoiiaeth? c?inoaiiy aeacaieo
e?i?aeieo eiiiiiaio?a ii??aiyii c eiio?ieai ia 39, 34 ? 22%
a?aeiia?aeii. Ia a?aei?/aii aeinoia??ieo ci?i ai?noo EE ? OE ia i?ioyc?
anueiai ia??iaeo ?inoo ?ineei i?e iecueeeo oaiia?aoo?ao (oaae.1).
I?iaaaeai? iaie aeine?aeaeaiiy na?ae/aoue i?i c?inoaiiy ai?noo
oinoie?i?ae?a o i?oioiiae??yo iceii? ioaieoe? Ie?ii?anueea 808 a
i?ioean? aaeaioaoe?? aei iecueeeo oaiia?aoo?. Oe? ci?ie caaacia/othoue
a?eueo aenieee ??aaiue aia?aii?iaeoeoe?? a i?aaiaeao ii?icino?eeiai
ni?oo i?e iecueeeo oaiia?aoo?ao, i?aeo?eioth/e iaiao?aeiee ??aaiue
aeo?ao ia i?aeo?eiaiiy no?oeoo? ee?oei ca iani?eyoeeaeo oaiia?aoo?ieo
oiia.

?ieue aee?iieeneioiiai neeaaeo iaia?ai ?ineei a i?aeo?eiaii? ooieoe?e
ee?oei i?e aeno?aiaeueieo oaiia?aoo?ao na?aaeiaeua

Aea/aiiy a?aeiiniiai neeaaeo aee?ieo eeneio oinoie?i?aeii? o?aeoe??
eenoe?a, eieaiioee?a oa iacieioee?a 4 aeiaiaeo ao?ieueiaaieo ia?inoe?a
eoeo?oaece iieacaei, ui iane/ai? aee?i? eeneioe, ia?aaaaeii,
i?aaenoaaeai? iaeuei?oeiiaith (N16:0) oa noaa?eiiaith (N18:0), a
iaiane/ai? — iea?iiaith (N18:1), e?iieaaith (N18:2) oa e?iieaiiaith
(N18:3) eeneioaie. Na?aae iane/aieo aee?ieo eeneio o i?aaiao ?ineei, ye?
ie aea/aee ia?aaaaea? iaeuei?oeiiaa (N16:0), oiae? ye na?aae iaiane/aieo
— e?iieaaa (N18:2). I?e ae?iuoaaii? ?ineei ca oiia iecueei? oaiia?aoo?e
(+10°N) niinoa??aaeeny ci?ie a?aeiiniiai ai?noo aee?ieo eeneio o an?o
aeine?aeaeaieo i?aaiao ?ineei. Aiae?c ie?aieo a?oi aee?ieo eeneio aeyaea
ciaioaiiy e?eueeino? iane/aieo iaeuei?oeiiai? (N16:0) oa noaa?eiiai?
(N18:0) eeneio o an?o i?aaiao ?ineei, yea a?aeiia?aeaei o?eaaeino? ae??
iecueeioaiia?aoo?iiai oaeoi?a. Ne?ae a?aecia/eoe, ui o aeiaaeeo
iacieioee?a oe? ci?ie a?aeaoaathoueny eeoa i?e 10 aeiaia?e aeniiceoe?? a
oiiaao caaa?oiaoth/i? oaiia?aoo?e, oiae? ye i?e 4 aeiaia?e ?o ??aaiue
caeeoa?oueny o iaaeao eiio?ieth. O eenoeao oa eieaiioeeyo e?euee?noue
iane/aieo iaeuei?oeiiai? oa noaa?eiiai? eeneio nooo?ai cieaeoaaeanue
aaea i?e 4 aeiaao caaa?ooaaiiy, oa caa??aaeanue ia oiio ae ??ai? ? i?e
10 aeiaao ae?? oieiaeo. I?e aeniiceoe?? i?ioyaii 21 aeiae o eenoeao oa
eieaiioeeyo ciia niinoa??aaeiny ianooiia nooo?aa cieaeaiiy a?aeiiniiai
ai?noo oeeo aee?ieo eeneio.

Aiae?c io?eiaieo ?acoeueoao?a iieacaa, ui a i?ioean?
iecueeioaiia?aoo?iiai caaa?ooaaiiy a?aeaoaa?oueny ca?eueoaiiy e?eueeino?
iaiane/aieo aee?ieo eeneio oinoie?i?ae?a o an?o i?aaiao ao?ieueiaaieo
ia?inoe?a eoeo?oaece, ye? ie aea/aee. Io?eiai? aenia?eiaioaeuei? aeai?
na?ae/aoue i?i oa, ui oea ? iane?aeeii c?inoaiiy a?aeiiniiai ai?noo
e?iieaaiai (N18:2) oa e?iieaiiaiai (N18:3) aee?iieeneioieo eaioetha?a o
neeaae? oinoie?i?ae?a.

Neioac a?ee?a a ee?oeiao ceae?a oa ?o i?oioiiae??yo ca oiia iecueeeo
oaiia?aoo? na?aaeiaeua.

Aeine?aeaeaiiy aieeao ??cieo oiia ae?? oieiaeo ia neeaae oa neioac
a?ee?a i?oioiiae??e ceae?a. A i?ioean? aeine?aeaeaiiy aieeao iecueeeo
oaiia?aoo? ia iiaiooai?aiiy iie?iaioeae?a a ee?oeiao eoeo?oaece
anoaiiaeaii, ui ci?ia neioaco caaaeueiiai a?eea ? a?eueo /ooeeaith aei
ae?? oaiia?aoo?iiai /eiieea ? a?aeaoaa?oueny ?ai?oa, i?ae ia?aaoaeiae,
ui niinoa??aathoueny o niaeo?? iiaineioaciaaieo i?oioiiae??aeueieo
a?ee?a.

Oi?ioi aeine?aeaeaieie ? ci?ie neioaco i?aaenoaaiee?a ?iaeei AOO, ui
aeieeathoue a oiiaao aieeao aenieeo oaiia?aoo? [Vierling, 1991]. Oea
aeicaiey? aeei?enoiaoaaoe aa??aio oaieiaiai oieo a yeino? ae?oaiai
eiio?ieth i?e aeine?aeaeaii? aieeao oieiaeo ia iiaiooai?aiiy
iioioiiae?iaeueieo aieeia. Ae?y cieaeaii? oaiia?aoo?e ia i?i?inoee
eoeo?oaece a?eo 4–5 ae?a i?ecaiaeeoue aei ci?i niaeo?a iiaineioaciaaieo
a?ee?a (?en. 5) Ia?o ca ana, oea noino?oueny oaia?iiieo ae?eyiie, – 70 ?
90 eAe. E??i oiai, iiaeia aeae?eeoe ua aeaye? oa?aeoa?i? iie?iaioeaee,
ieoiiee neioac yeeo nooo?ai iineeth?oueny aiane?aeie oieiaeiaiai aieeao,
oea a?eee c iieaeoey?ieie ianaie aeecueei 23, 32 oa 58 eAe. Iiaeaeueoa
eia?ea aeine?aeaeaiue i?eaaea aei iaiao?aeiino? ?icae?eaiiy aoaeoo ae??
oieiaeiaiai /eiieea aeey ??cieo oei?a oeaiei. O ca’yceo c oeei
ii??aithaaee aieea oieiaeo ia ai?eaeueio oa aacaeueio /anoeio ?ineei
eoeo?oaece. Io?eiaii ?acoeueoaoe, ye? na?ae/aoue, ui ia?enoaiaoe/ia
oeaieia ia ?icaeaa? a?aecia/aio ?aaeoe?th (iineeaiiy ?ioaineaiinoae
neioaco iie?iaioeae?a o oaia?iiieo ae?eyieao 70 ? 90 eAe), ia oaieiaee
oie aeeth/ii. Oa?aeoa?ia a?aeiia?aeue ? iaa?ae’?iiith ?enith aacaeueieo
oeaiei, ee?oeie yeeo i?eieieee iiae?e . Oaiia?aoo?a +10 °C aeeeeea?
ci?ie neioaco a?eea, ui aeaoaeoothoueny aaea i?ney 10 aiae., iaeiae
iioeiaeueia o?eaae?noue aeey i?iyao ?aaeoe?? aei ae?? oieiaeo ia ??ai?
iiaiooai?aiiy Io a?ee?a neeaaea? ia iaioa 20 aiae (?en. 5).

Aieea iecueeeo oaiia?aoo? ia neioac a?ee?a ao?ieueiaaieo i?i?inoe?a
iceii? ioaieoe? A aeine?aeaeaiiyo aoee aeei?enoai? i?i?inoee ni?oe
iceii? ioaieoe?, ui a?ae??ciythoueny ca ii?icino?ee?noth: Ethoanoeain 7
(E7), Ie?ii?anueea 808 (I808) – ii?icino?ee?, Ie?ii?anueea 40 (I40) –
na?aaeiy ii?icino?ee?noue, Ie?ii?anueea 29 (I29) – ii?icino?ee?noue
ieae/a ca na?aaeith. I?e ae?? aeniei? oaiia?aoo?e (+41°C) o niaeo?ao
iiaineioaciaaieo caaaeueieo oa i?oioiiae??aeueieo a?ee?a
niinoa??aathoueny oa?aeoa?i? nioae AOO, ui a?aeiia?aeathoue
iieaeoey?iei ianai (II) 73, 82 ? 98 eAe, a oaeiae a ae?eyioe? 16–34 eAe.
Aeo?eioaaiiy ?ineei ioaieoe? ca oiia iecueeeo aeiaeaoi?o oaiia?aoo?
i?ecaiaeeoue aei i?aeaeuaiiy ?ioaineaiino? iiaiooai?aiiy iie?iaioeae?a o
ae?eyieao 70 oa 90 eAe (?en. 6, o?aee 4 ? 9). A?aecia/ai? ci?ie noiae?
ia o?, ui niinoa??aa?oueny aeey aa??aioa oaieiaiai oieo, aea aeyaeyoueny
o iaio?e i???.

Aeey i?i?inoe?a ioaieoe? aeyaeaii niaoeeo?/io ae?eyieo i?iyao aoaeoo
oieiaeo o aenieiiieaeoey?i?e /anoei? niaeo?aeueiiai neeaaeo
iiaineioaciaaieo iie?iaioeae?a – aeecueei 120 eAe. C?inoaiiy
?ioaineaiino? neioaco aeacaieo a?ee?a a?aecia/a?oueny i?ney o?eaaeino?
ae?? oieiaeo ia iaioa 10–12 aiae. oa ca oiia ?iioeueniiai
?aae?iaeoeaiiai i?/aiiy i?e iecuee?e oaiia?aoo??. Ci?ie o
aenieiiieaeoey?i?e ae?eyioe? iaeaea ia a?aei?/aii o c?aceao,
?aae?iaeoeaia i?/aiiy yeeo a?aeaoaaeinue i?e ii?iaeuei?e (+24°C)
oaiia?aoo?? i?ney oieiaeiaiai aieeao (?en. 6, o?aee 5 ? 10), ui
iiyniththoueny, iiaeeeai, oaeaeeei i?eieiaiiyi neioaco iie?iaioeae?a
aeacaieo iieaeoey?ieo ian i?ney ia?aianaiiy i?i?inoe?a o ii?iaeuei?
oaiia?aoo?i? oiiae.

Iiaeia aeae?eeoe ua aeaye? oa?aeoa?i? iie?iaioeaee, ieoiiee neioac yeeo
nooo?ai iineeth?oueny aiane?aeie oieiaeiaiai aieeao – oea iie?iaioeaee c
II aeecueei 16, 22, 24, 35, 38 ? 54eAe . Aeey c?ace?a a?eea Io iniiai?
ci?ie ?ioaineaiino? nioa iiaineioaciaaieo iie?iaioeae?a i?e ae??
iecueei? oaiia?aoo?e aeaoaeoothoueny o ae?eyieao 70 eAe, 38 eAe, 35 eAe
oa 16eAe. Iiae?aia i?ieea?noue o aeacaieo ae?eyieao ? ni?eueiith aeey
c?ace?a i?oioiiae??aeueiiai oa caaaeueiiai a?eea. Aeine?aeaeaiith
iniaeeainoae i?ieeaino? neioaco iie?iaioeae?a o oaia?iiieo ae?eyieao
aoei i?eae?eaii iniaeeao oaaao (oaae. 2).

Oaaeeoey 2

Ieoiia ?ioaineai?noue neioaco a?ee?a i?i?inoe?a ioaieoe? a oiiaao
oieiaeo o oaia?iiieo ae?eyieao (ca aeeth/aiiyi ?aae?iaeoeaii? i?oee), %
aei eiio?ieth

Oiiae oa ?ioaineai?noue neioaco a?ee?aNi?oO?eaae?noue
caaaeueieoI?oioiiae??aeueieoaieeao71 82 eAe71 eAe 82 eAeOieiae I29+2°C
10 ae?a122 ± 18115 ± 16111 ± 18110 ± 8I40+3°C 7 ae?a––127 ± 19122 ±
15E7+3°C 7 ae?a––138 ± 21128 ± 17I808+2°C 10 ae?a183 ± 27179 ±38132 ±
14119 ± 12I?ei?oea. “–” – aeaieo iaia?.

O i?i?inoeao ni?o?a, a?eueo /ooeeaeo aei ae?? iecueeeo oaiia?aoo?,
?iaeoeoe?y neioaco a?ee?a o ae?eyieao AOO70 ? AOO90 ae?aaeaia neaae?oa,
i?ae o a?eueo no?eeeo ni?o?a.

O i?iaaaeaieo aeine?aeaeaiiyo aoei aeyaeaii, ui oaieiaee oie aeeeeea?
iineeaiee neioac AOO70 c II 73 eAe , oiae? ye i?e oieiaeia?e aaeaioaoe??
ca?eueoo?oueny ??aaiue iiaiooai?aiiy AOO c II 71 eAe. A??ia?aeii, oea
iia’ycaii c ??cieie ooieoe?yie oeeo AOO: iiaeia aeoiaoe, ui ia?oee c ieo
ia? ni?aao c aeaiaoo?iaaieie aiane?aeie no?an?a a?eeaie, ?ioee
iaiao?aeiee aeey i?ioean?a o?ainii?ooaaiiy aai neeaaeaiiy
iiaineioaciaaieo a?ee?a i?e cieaeai?e oaiia?aoo??.

Iie?iaioeae c II 35 eAe (iineeaiee neioac yeiai niinoa??aa?oueny) iiaea
yaeyoe niaith iiiiia? aeueoa?iaoeaii? ieneaeace, aaeaea aeoeaaoe?y
oeueiai oeyoo aeicaiey? caoenoeoe ee?oeio a?ae a?eueii ?aaeeeaeueiiai
iioeiaeaeaiiy a oiiaao aa?ioe/ieo no?an?a [Purvis, Shewfelt, 1993].
I?iya i?ieeaino? neioaco o ieaei?e /anoei? niaeo?o (16–18 eAe)
iia’ycaiee, iaea??ia?aei?oa, ?c ?ioaineo?eaoe??th iiaiooai?aiiy
iecueeiiieaeoey?ieo AOO [Sabehat et al., 1998].

Neioac a?ee?a i?oioiiae??e ca?iaee?a eoeo?oaece i?e ?o ?icaeoeo a oiiaao
ae?? aenieeo oa iecueeeo oaiia?aoo?. ?yae aai?a oaieiaiai oieo
aeni?anothoueny ia i?ioyc? ?aii?o aoai?a ?icaeoeo aia??iiaeueieo oeaieo
aaea??io, aea niaoeeo?/ia ?ieue AOO a aia??iaaiac? c’yniaaia neaai.
Aeine?aeaeaiiy ae?? oaeeo oeiiai no?aniaeo oaeoi??a, ye i?aeaeuaia /e
iecueea oaiia?aoo?a, ia ?aii?o aoaiao ?icaeoeo ?ineei i?aaenoaaey?
cia/iee ?ioa?an ye c oi/ee ci?o ooiaeaiaioaeueii? iaoee, oae ? c
i?aeoe/ieo i??eoaaiue. Aaaeeeaa ?ieue nenoaie aia?aicaaacia/aiiy o
?aae?caoe?? no?an-?aaeoe?? ee?oei aei iani?eyoeeaeo oiia coiiaeth?
i?aeaeuaio oaaao aei a?ee?a i?oioiiae??iio [Martinus et al., 1996].

Aeiaaio?eaaeee aieea oieiaeo ia ca?iaeee a?a?eaeo eoeo?oaece Eieaeoeaiee
100 NA aeine?aeaeoaaee a na??? aenia?eiaio?a, o yeeo ae?iuoaaiiy
aeine?aeieo ?ineei a?aeaoaaeinue i?e cieaeai?e (+14 °N) oaiia?aoo??. A
?io?e na??? aeine?ae?a i?ney i?i?iuoaaiiy ian?iiy o ii?iaeueieo
oaiia?aoo?ieo oiiaao (+24 °N) /anoeio ca?iaee?a aeo?eioaaee i?ioyaii 24
aiae. i?e +10 °N (oieiae) aai 2 aiae. i?e +42 °N (oaieiaee oie).
?cieueiaai? ?c ian?iiy ca?iaeee ?ieoaoaaee o ?ic/ei? 35S-iaoeii?io.
Aiae?c ?iciiae?eo a?eeiaeo nioa ia ?aae?iaaoia?aoao aeyaea, ui iniiai?
iii?i? nioae, a naia, c iieaeoey?ieie ianaie 10, 60, 70 ? 90 eAe, ye?
a?aeiia?aeathoue a?eeai ?c ?iaeeie oaia?ii?a, iayai? i?aeoe/ii o an?o
aeine?aeaeaieo aa??aioao (?en. 7). Aeeeth/aiiy neeaaethoue noo?
ca?iaeee, aea ia niinoa??aa?oueny acaaae? i?yeiai iii?oiiai ??aiy
neioaco a?eea de novo. O o?aeao, ui a?aeiia?aeathoue 6 aiae. ?icaeoeo
ca?iaee?a i?e oaiia?aoo?? +24 °N oa 12 aiae. – i?e oaiia?aoo?? +14 °N,
?ioaineai?noue aeeth/aiiy ?aae?iaeoeaii? i?oee i?eaeecii iaeiaeiaa ?
ciaoiaeeoueny ua ia aeineoue iecueeiio ??ai?. Oea na?ae/eoue i?i oa, ui
o aeacaieo oiiaao neioac a?eea Io o?eueee ii/eia?oueny. Iiae?aiee
ia?aeae?ci i?ae aa??aioaie ??cieo oaiia?aoo?ieo oiia ?icaeoeo
niinoa??aa?oueny i?e aiae?c? ?ioaineaiinoae nioa, ui a?aeiia?aeathoue
iniiaiei a?eeai-oaia?iiai: ae?eyiea 70 eAe- ii/aoie neioaco i?ae 12 ? 24
aiae. i?e oaiia?aoo?? +24 °N oa i?eaeecii 48 aiae. – i?e +14 °N;
ae?eyiea 60 eAe – iiyaa, a iio?i iinooiiaa cieeiaiiy nioae c
iieaeoey?iith ianith i?eaeecii 65 eAe ? aeae? iiyaa nioae c ieae/ith
iieaeoey?iith ianith (i?eaeecii 62,5 eAe) ye o aeine?aeieo, oae ? a
eiio?ieueieo c?aceao i?aeoe/ii c oei naiei (o 2–2,5 ?ace) cnoaii o /an?.
Aiaeia?/iee oa?aeoa? i?ieeaino? niinoa??aa?oueny ? a ae?eyioe? 90 eAe.
?acoeueoaoe aeine?aeaeaiiy iiaiooai?aiiy iie?iaioeae?a a ca?iaeeao
eoeo?oaece aeathoue i?aenoaao aeey i?eiouaiiy i?i oa, ui
aeine?aeaeoaaiee iecueeioaiia?aoo?iee aieea (+14 °N) i?e ?icaeoeo
ca?iaee?a aeaiiai a?a?eaeo eoeo?oaece ia ni?e/eiy? niaoeeo?/ii?
aeoeaaoe?? neioaco oieiaeo a?ee?a, na?aae yeeo cae/aeii niinoa??aa?oueny
iineeaiiy iiaiooai?aiiy a?ee?a-oaia?ii?a, yeei a?aeiia?aeathoue /ioe?e
ae?eyiee c iieaeoey?ieie ianaie aeecueei 70, 60, 10 ? 90 eAe. O aeai?e
na??? aenia?eiaio?a a?aeaoaa?oueny c?inoaiiy ?o neioaco, aea, eiia??ii,
ia aiane?aeie aieeao iecueei? oaiia?aoo?e, a a ?acoeueoao? ?aae?caoe??
i?ia?aie ?icaeoeo ca?iaee?a. Aaaeeeai a?aei?oeoe, ui neioac a ee?oeiao
ca?iaee?a eoeo?oaece iie?iaioeae?a, ui a?aeiinyoueny aei oeiiai
«oieiaeo» ae?eyiie a?eeiaiai niaeo?o, niinoa??aa?oueny ia iaaieo
noaae?yo ?icaeoeo. A?aeiii, ui, iai?eeeaae, i?aaenoaaieee ?iaeeie AOO 90
caeaoi? ci?ithaaoe o i?inoi?? ? /an? i?i oeane aia??iaaiaco ??cieo
nenoai oaa?eiiiai iioiaeaeaiiy [Csermely et al., 1998]. Aiae?c
?acoeueoao?a, io?eiaieo o ianooii?e na??? aenia?eiaio?a, aeicaiey?
c?iaeoe oae? aeniiaee. Iie?iaioeae c iieaeoey?iith ianith 70 eAe
neioaco?oueny aaea ia naieo ?aii?o aeine?aeaeoaaieo noaae?yo ?icaeoeo
ca?iaee?a – ii/eiath/e c 12 aiae. i?ney ii/aoeo caii/oaaiiy (?en. 8). Ia
oe?e noaae?? aieea oaiea /e oieiaeo i?aeoe/ii ia ni?e/eiy? ci?i
?ioaineaiino? eiai neioaco. Iiae?aia niinoa??aa?oueny oaeiae ? i?ney 24
aiae. o?eaaeino?

?icaeoeo, iaeiae aeey aa??aioo oaieiaiai oieo ? iii?oiei oaeiae neioac ?
ieaeiuei? neeaaeiai? ?c iaioith iieaeoey?iith ianith (i?eaeecii 68 eAe),
ui, nei??o ca ana, iiynith?oueny cae/aeiei i?enei?aiiyi ia?aoai?aiue i?e
i?aeaeuaii? oaiia?aoo?e aai a?eueo neeueiith iai?oaeai?noth no?aniaiai
oaeoi?a ii??aiyii aei iecueei? oaiia?aoo?e.

Aeae?, i?e ia?aoiae? aei 48 aiae. o?eaaeino? ?icaeoeo oa?aeoa?
i?ieeaino? o oe?e ae?eyioe? aeey an?o o?ueio aa??aio?a – eiio?ieue,
oaieiaee oie oa oieiae – i?aeoe/ii iaeiaeiaee, ca aeeeth/aiiyi
eiiiiiaioe a?eeiaiai niaeo?o c ?iaeeie 70 eAe: niinoa??aa?oueny neioac o
aa??aio? oaieiaiai oieo, a?aecia/a?oueny aeey aa??aioo oieiaeiaiai
aieeao ? i?aeoe/ii a?aenooi?e o eiio?ie?. Aeey ?ioeo iii?ieo ae?eyiie
niaeo?o a?ee?a ca?iaee?a eoeo?oaece – 90, 60 ? 10 eAe niinoa??aa?oueny
iiae?aia oaiaeaioe?y, ye ? o AOO70. Iieacaii, ui, iai?eeeaae, aia??iie
?ea iaaoaathoue caeaoiino? aei aeoioeyoe?? mRHK AOO 47, AOO 70 ? AOO 90
i?e ae?? oaieiaiai oieo eeoa i?ney i?ioiaeaeaiiy noaae?? aeanooee [Lele
et.al., 1997]. I?e aeinyaiaii? noaae?? aeineoue noi?iiaaieo i?i?inoe?a
eoeo?oaece(48 aiae.) nooi?iue ?acenoaioe?? cieaeo?oueny ? aieea
no?aniaeo oaiia?aoo? i?ecaiaeeoue aei niaoeeo?/ieo ci?i ??aiy neioaco
iie?iaioeae?a c iieaeoey?ieie ianaie 70–72 eAe. A oiiaao ae?? oieiaeo
?aaeoe?y i?i?inoe?a i?ney 2 ae?a ?icaeoeo iiae?aia aei oaei?, ui
niinoa??aa?oueny aeey aa??aioo oaieiaiai oieo . Io?eiai? ?acoeueoaoe
na?ae/aoue, ia ?aii?o noaae?yo ?icaeoeo ca?iaee?a eoeo?oaece neioac
iie?iaioeae?a o oeiiaeo oaia?iiieo ae?eyieao a?eeiaiai niaeo?o
aecia/a?oueny naiith noaae??th ?icaeoeo ca?iaee?a ? iaei caeaaeeoue a?ae
ciai?oi?o oaiia?aoo?ieo oiia.

?ioaineai?noue aeeoaiiy oeaiei ?ineei o i?ioean? ?o ?inoo.

Oi?ioaaiiy aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a a i?oioiiae??yo
ei?ai?a eoeo?oaece. Aea/aee ?ioaineai?noue iiaeeiaiiy eenith ei?aiyie
a?a?eaeo eoeo?oaece Eieaeoeaiee 100 NA ciie ia?enoaie (ae?eyiea 0-2 ii
a?ae ii/aoeo ei?aiy), ciie ?icoyaoaaiiy (ae?eyiea 2-4 ii) oa ciie
aeic??eeo ee?oei (10-20 ii). A i?ioean? ?inoo ei?ai?a eoeo?oaece
?ioaineai?noue aeeoaiiy ?o ee?oei, a ia?a?aooieo ia nooo ?a/iaeio,
cieaeo?oueny a?ae 450,9 ieI I2 (a noo.?a/ia.)-1aiae-1 a ee?oeiao ciie
ia?enoaie aei 249,2 ieI I2 (a noo.?a/ia.)-1aiae-1 a aeic??eeo ee?oeiao
ei?ai?a. Iieaciee ?ioaineaiino? aeeoaiiy ee?oei ciie ?icoyaiaiiy caeia?
i?ii?aeia iieiaeaiiy i?ae ee?oeiaie ia?enoaie oa aeic??eeie ee?oeiaie
ei?ai?a (oaae. 3, eiio?ieue). Aaaaeaiiy a oeaieie ia?enoaie ?ia?a?oi?a
oeeoio?iiiiai oeyoo o?ainii?oo aeaeo?ii?a (aceaeo iao??th) cieaeoaaei ia
56% ?ioaineai?noue aeeoaiiy ee?oei a ia??iae ?o iiae?eo. ?acii c oei,
ee?oeie ia?enoaie neaai ?aaaoaaee ia ae?th ?ia?a?oi?a aeueoa?iao?aiiai
oeyoo o?ainii?oo aeaeo?ii?a — aaica?ae?ienaiiai? eeneioe. Oeaieie ciie
ii/aoeo ?icoyaoaaiiy ee?oei ei?ai?a eoeo?oaece aoee a?eueo no?ee? aei
ae?? aceaeo iao??th ? aeinoia??ii ia a?ae??ciyeeny ca /ooeea?noth aei
ae?? aaica?ae?ienaiiai? eeneioe, ii??aiyii c oeaieiaie ia?enoaie (oaae.
3, eiio?ieue). Aeic??e? ee?oeie ei?ai?a eoeo?oaece (ciia 10-20 ii a?ae
ii/aoeo ei?aiy) aoee iaeaea aaea?/? /ooeea?oeie aei ae?? ?ia?a?oi?a
aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a ii??aiyii aei ee?oei
ia?enoaie. Oae nooi?iue ?ia?aoaaiiy ?ioaineaiino? iiaeeiaiiy eenith a
i?enooiino? aaica?ae?ienaiiai? eeneioe neeaea 16,4% a?ae
eiio?ieth.Noi?nia aaaaeaiiy a oeaieie ei?ai?a a?a?eae?a eoeo?oaece
aceaeo iao??th oa aaica?ae?ienaiiai? eeneioe i?eai?/oaaei aeeoaiiy aei
90%, ui na?ae/eoue i?i aeii?ioth/o ?ieue i?oioiiae??aeueieo oa?iaio?a o
iiaeeiaii? eenith. sse na?ae/aoue ?acoeueoaoe i?iaaaeaieo
aeine?aeaeaiue, a i?ioean? ?inoo ee?oei ei?ai?a niinoa??aa?oueny
cieaeaiiy ?ioaineaiino? iiaeeiaiiy eenith (oaae.3), ui ocaiaeaeo?oueny
?c ?acoeueoaoaie aeine?aeaeaiiy aeeoaiiy i?ioiieano?a y/iaith, io?eiao
?c naaiaio?a, ?icoaoiaaieo aeua oa ieae/a eioa?eaey?ii? ia?enoaie
i?i?inoe?a y/iaith [Owen et al.,1986]. Ci?ie a ?ioaineaiino? iiaeeiaiiy
eenith ee?oeiaie ia?enoaie ei?ai?a eoeo?oaece i?e aeieoaaii? o?ainii?oo
aeaeo?ii?a ii oeeoio?iiiiio eaioethao i?oioiiae??e ? iiae?ai? aei
nooiaith no?eeino? ee?oei ?ioa?eaey?ii? ia?enoaie eenoe?a y/iaith aei
ae?? aiaeia?/iiai ?ia?a?oi?a [Owen et al.,1986].

Io?eiai? ?acoeueoaoe aeine?aeaeaiue aia?aiiai?io ceae?a na?ae/aoue i?i
oa, ui oi?ioaaiiy aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a a
i?oioiiae??yo a?aeaoaa?oueny i?ae /an iiae?eo ee?oei eoeo?oaece.
I?aaiaee ee?oei ia?enoaie caeaoi? o?ainii?ooaaoe aeaeo?iie ii
aeueoa?iaoeaiiio oeyoo, aea oeae oeyo a ee?oeiao ia?enoaie ooieoe?iio?
neaai. Caeiyo?noue oe?ai?ae?acenoaioiiai oeyoo o?ainii?oo aeaeo?ii?a
aeaui ca?eueoo?oueny a cii? ii/aoeo ?icoyae?iiy ee?oei ei?i?a eoeo?oaece
. Iaea?eueoo aeoeai?noue aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a
i?iyaeythoue aeic??e? ei?ai? eoeo?oaece. Oaeei /eiii iieacaii, ui
aeueoa?iaoeaiee oeyo o?ainii?oo aeaeo?ii?a a i?oioiiae??yo oi?io?oueny,
ye ? oeeoio?iiiee, ia ?aii?o aoaiao ?inoo ee?oei, aea iaea?eueo aeniea
aeoeai?noue oe?ai?ae?acenoaioiiai aeeoaiiy niinoa??aa?oueny o aeic??eeo
ee?oeiao ceae?a.

Aieea oieiaeiaiai oieo ia aeoeai?noue oeeoio?iiiiai oa aeueoa?iaoeaiiai
oeyo?a o?ainii?oo aeaeo?ii?a a ee?oeiao ia?enoaie, aeic??eeo ee?oei
ei?ai?a oa eenoe?a eoeo?oaece. Aea/aiiy aieeao oieiaeiaiai oieo ia
?ioaineai?noue iiaeeiaiiy eenith ee?eeiaie ceae?a aoei ?icii/aoi c
aecia/aiiy /ooeeaino? aeeoaeueiiai aia?aiiai?io aei iiieaeaiiy
oaiia?aoo?e na?aaeiaeua ee?oei ?ineei, ui ciaoiaeyoueny ia ??cieo aoaiao
?inoo. sse ia’?eo aeine?aeaeaiue aoee aeei?enoai? ei?ai? a?a?eae?a
eoeo?oaece Eieaeoeaiee 100 NA. Iiey?ia?ao?/iee aiae?c iiaeeiaiiy eenith
ei?aiyie ??cieo ae?eyiie ei?ai?a a?a?eae?a eoeo?oaece aeyaea ??cio
?aaeoe?th ia oieiaeiaee no?an ee?oei ei?ai?a eoeo?oaece a i?ioean? ?o
?inoo. Oae, ia aeyaeaii aeinoia??ieo ci?i ?ioaineaiino? aeeoaiiy ee?oei
ia?enoaie i?ae aieeaii ae?? oieiaeo (oaae.3), a oie /an ye ee?oeie ciie
ii/aoeo ?icoyae?iiiy (2-4 ii a?ae e?i/eea ei?aiy) ?aaaothoue iacia/iei
i?aeaeuaiiyi ?ioaineaiino? iiaeeiaiiy eenith i?ney ae?? oieiaeiaiai
no?ano. Iaea?eueo ae?aaeaia ?aaeoe?y ia oieiaeiaee oie o ee?oei ei?ai?a
a?a?eae?a eoeo?oaece, ui caaa?oeee ??no (ae?eyiea 10-20 ii), aea
a?aei?/aii cia/ia i?aeaeuaiiy aeeoaeueii? aeoeaiino? (oaae.3).
Ioieiaeaeaiiy ?ineei i?aeoe/ii ia ci?ith? /ooeea?noue an?o
aeine?aeaeoaaieo cii ei?ai?a aei ?ia?a?oi?a oeeoio?iiiiai oeyoo
o?ainii?oo aeaeo?ii?a a i?oioiiae??yo — aceaeo iao??th. ?acii c oei, o
an?o aeine?aeaeoaaieo cii ?inoo ei?ai?a oieiaeiaee no?an ci?ith?
?aaeoe?th aeeoaiiy ia aaaaeaiiy a na?aaeiaeua ?ieoaaoe?? ?ia?a?oi?a
aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a a i?oioiiae??yo
aaica?ae?ienaiiai? eeneioe. Niinoa??aa?oueny ca?eueoaiiy nooiaith
?ia?aoaaiiy aeeoaiiy aaica?ae?ienaiiaith eeneioith 9,6 aei 11,5% a
ee?oeiao ia?enoaie; c 11,5 aei 17,0% a cii? ii/aoeo ?icoyaoaaiiy ee?oei;
c 16,4 aei 24,3% a cii? aeic??eeo ee?oei ei?ai?a a?a?eae?a eoeo?oaece,
a?aeiia?aeii aei oa i?ney oieiaeiaiai oieo.

Oaaeeoey 3

Aieea iecueeeo oaiia?aoo? ia aeoeai?noue oeeoio?iiiiai ?
aeueoa?iaoeaiiai oeyo?a o?ainii?oo aeaeo?ii?a i?oioiiae??e ee?oei
ei??iiy a?a?eaeo eoeo?oaece Eieaeoeaiee 100 NA i?e 25°C oieM O2(a
noo.?a/ia.)-1 aiae-1

Eiio?ieue, 25 NOieiae, 10 N, 24 aiaeAe?eyiea ei??iiyAe?eyiea ei??iiy0-2
ii2-4 ii10-20 ii0-2 ii2-4 ii10-20 ii?ioaineai?noue aeeoaiiy

450,9±6,3

338,8±8,9

249,2±2,8

451,6±9,1

363±10,1

343,5±4,9Aeoeai?noue oeeoio?iiiiai aeeoaiiy

359 ± 4

265 ± 8

177 ± 3

359 ± 6

266 ± 8

231 ± 12Iioaioe?eia aeoeai?noue aeueoa?iaoeaiiai aeeoaiiy

150 ± 2

144 ± 16

100 ± 6

157 ± 6

148 ± 5

138 ± 7?ioaineai?noue aeeoaiiy a i?enooiino?NaN3 oa AAE

48,4 ± 4,6

34,9 ± 0,5

31,0 ±2,9

40,8 ± 2,7

35,0 ±4,2

29,3 ± 2,3

Io?eiai? ?acoeueoaoe na?ae/aoue i?i aeoeaaoe?th aeueoa?iaoeaiiai oeyoo
i?e iecueeeo oaiia?aoo?ao a ee?oeiao, ui caaa?oeee ??no. Ca?eueoaiiy
aeoeaiino? aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a i?e iiieaeaii?
oaiia?aoo?e na?aaeiaeua iiaea aooe iaoiiaeaii iiaiooai?aiiyi a?eea
[Vanlerberghe and McIntosh, 1992], aiane?aeie aeoeaaoe?? aeni?an??
aai?a, ye oea aoei iieacaii ia oae aeaaii aeey ?eno (Ito et al., 1997).

?aaoeyoe?y aeoeaiino? oeeoio?iiiiai oa aeueoa?iaoeaiiai aeeoaiiy a
i?ioean? ?icaeoeo ca?iaee?a eoeo?oaece i?e i?i?inoaii? ian?iiy. Ci?ie a
?ioaineaiino? iiaeeiaiiy eenith oeaieiaie ?ineei a?aeia?aaeothoue ci?ie
a aeoeaiino? iaoaaie?cio ee?oei, yeee o?nii iia’ycaiee c oacith ?o
?icaeoeo. I?i?inoaiiy ian?iiy eoeo?oaece noi?iaiaeaeo?oueny ??ceei
i?aeaeuaiiyi ?ioaineaiino? aeeoaiiy (oaae.4). No?eea aei ae?? aceaeo
aeeoaiiy oeaiei aia??ii?a niinoa??aa?oueny ia an?o noaae?yo i?i?inoaiiy
ian?iiy. ?acii c oei /ooeea?noue oeaiei ?ineei aei ae?? SHAM ??cei
c?inoa? ia i?ioyc? 24 – 120 aiaeei i?i?inoaiiy ian?iiy eoeo?oaece i?e
iecueeeo oaiia?aoo?ao.

Oaaeeoey 4

Ci?ie ?ioaineaiino? iiaeeiaiiy eenith oa aeoeaiino? oeeoio?iiiiai ?
aeueoa?iaoeaiiai oeyo?a o?ainii?oo aeaeo?ii?a ca?iaee?a ian?iiy a?a?eaeo
eoeo?oaece Eieeaeoeaiee 100 NA i?e 12°C o ieM O2(a noo. ?a/ia.)-1
oaee.-1

O?eaae?noue i?i?inoaiiy ian?iiy i?e 12 °C (aiae.)?ioaineai?noue aeeoaiiy
ca?iaee?aAeoeai?noue oeeoio?iiiiai ueyooAeoeai?noue aeueoa?iaoeaiiai
oeyooCaeeoeiaa aeeoaiiy

24

380,2 ± 12

290,2 ± 15

53,2 ± 2,8

20,0 ± 3

72

564,2 ± 72

358,4 ± 32

160,8 ±18

45,0 ± 5

120

1114,5± 82

684,3 ± 72

287,5 ± 25

112 ± 8

192

670,4 ± 58

502,6 ± 35

116,0 ± 11

52,0 ± 6

Ca oeeo oiia niinoa??aa?oueny ca?eueoaiiy nooiaiy caeiyoino?
aeueoa?iaoeaiiai oeyoo a i?oioiiae??yo oeaiei aia??ii?a eoeo?oaece
(oaae. 4). A i?iaaaeaieo aeine?aeaeaiiyo a?aei?/aii iaeneioi aeoeaiino?
aeueoa?iaoeaii? ieneaeace o ia??iae 72 — 120 aiaeei i?i?inoaiiy i?e
iecueeeo oaiia?aoo?ao. Caeeoeiaa aeeoaiiy neeaaeaei a?ey 10% a?ae
caaaeueai aeeoaiiy ? niinoa??aaeinue ia an?o noaae?yo ?icaeoeo
aia??ii?a. ?acoeueoaoe i?iaaaeaieo aeine?aeaeaiue na?ae/aoue i?i
?iaeoeoe?th aeueoa?iaoeaiiai oeyoo aeeoaiiy o oeaieiao ca?iaee?a
eoeo?oaece a i?ioean? i?i?inoaiiy ian?iiy ca oiia ae?? iecueeeo
oaiia?aoo?. A i?ioean? aeine?aeaeaiiy iieiaeeo n?i’yaeieae ? oeaiei in?
ni? iieacaia caeaaei?noue i?ae i?aeaeuaiiyi aeoeaiino? aeueoa?iaoeaiiai
oeyoo i?oioiiae??e oa ca?eueoaiiyi ai?noo a?eea aeueoa?iaoeaii?
ieneaeace a oeeo oeaieiao (Obenland et al., 1990).

Aieea iecueeeo oaiia?aoo? ia ?aaoeyoe?th o?ainii?oo aeaeo?ii?a o
i?oioiiae??yo ?ineei. Aeyaeaii aeoeaaoe?th in vivo aeueoa?iaoeaii?
ieneaeace oeaiei ?ineei i?e iecueeeo oaiia?aoo?ao na?aaeiaeua. C iaoith
c’ynoaaiiy aieeao iecueeeo oaiia?aoo? ia ooieoe?? i?oioiiae??e ?ineei
iaie aoee i?iaaaeai? aeine?aeaeaiiy in vitro aieeao oieiaeo ia ia?aiao?e
i?oioiiae??aeueii? aeoeaiino?. sse na?ae/aoue i?iaaaeai? aeine?aeaeaiiy,
ia?aianaiiy ia?inoe?a ?ineei iceii? ioaieoe? ?c na?aaeiaeua c
oaiia?aoo?ith 25 N ia iecueeo oaiia?aoo?o (3 N) aeeeeeathoue ci?ie
ia?aiao??a i?oioiiae??aeueii? aeoeaiino?. *a?ac 24 aiaee ae?? oieiaeo ia
?ineeie a?aei?/aii c?inoaiiy ?ioaineaiino? iiaeeiaiiy eenith i?aia?aoaie
i?oioiiae??e a o?aoueiio (ia 12%) oa /aoaa?oiio (ia 34,7%) iaoaaie?/iiio
noaiao i?aaiae (?en. 9).

C?inoaiiy ?ioaneaiino? aeeoaiiy ?cieueiaaieo i?oioiiae??e
noi?iaiaeaeoaaeinue cieaeaiiyi aeeoaeueiiai eiio?ieth oa a?aeiioaiiy
AAeO:I. Ca aeacaiee i?ii?aeie /ano a?aeaoeiny c?inoaiiy ?ioaineaiino?
iiaeeiaiiy eenith a i?enooiino? aceaeo iao??th (aei 67,1%). A?aei?/aii
oaeiae i?aeaeuaiiy nooiaith ?ia?aoaaiiy iiaeeiaiiy eenith i?oioiiae??yie
i?ae aieeaii AAE (aei 38%). Iiaeaeueoee aieea iecueeeo oaiia?aoo? ia
ia?inoee ioaieoe? ia i?ioyc? 48 aiaeei ia aeeeeea? aeinoia??ieo ci?i
aeoeaiino? i?aaiae o o?aoueiio iaoaaie?/iiio noai?, a oie /an ye o
/aoaa?oiio iaoaaie?/iiio noai? a?aeaoaa?oueny aeinoia??ia cieaeaiiy
?aoaineaiino? iiaeeiaiiy eenith ii??aiyii ?c ??aiai, cao?eniaaiei /a?ac
24 aiaeeie ae?? oieiaeo ia ?ineeie (?en.9). A?aei?/aii oaeiae
i?aeaeuaiiy /ooeeaino? aeeoaiiy i?aaiae aei ae?? aceaeo iao??th, ?acii c
oei niinoa??aaaeiny iinooiiaa cieaeaiiy /ooeeaino? ieenethaaeueii?
aeoeaiino? i?oioiiae??e aei ae?? ?ia?a?oi?a aeueoa?iaoeaii? ieneaeace –
AAE.

Ai?iaeiaae aeiaaio?eaaeiai aeo?eioaaiiy ?ineei i?e iecueeeo oaiia?aoo?ao
a i?aaiaeao a?aei?/aii c?inoaiiy aia?aaoe/ii? aoaeoeaiino? aeeoaiiy oa
iinooiiaa i?aeaeuaiiy aeeoaeueiiai eiio?ieth i?oioiiae??e. I?e a?eueo
aeiaaio?eaae?e ae?? oieiaeo (a?ae 3 aei 6 ae?a) a?aeaoaa?oueny
iiaeaeueoa cieaeaiiy ?ioaineaiino? iiaeeiaiiy eenith i?aaiaeaie o
o?aoueiio oa /aoaa?oiio iaoaaie?/ieo noaiao ii??aiyii ?c ??aiai
cao?eniaaiei /a?ac 24 aiaeeie ae?? iecueeeo oaiia?aoo? ia ?ineeie (?en.
9). I?oioiiae??? i?iyaeyee iaioo /ooeea?noue aei ae?? AAE ?, iaaiaee o
a?eueo?e i??? i?eai?/oaaee ?ioaineai?noue aeeoeiiy i?e aaaaeaii? a
noniaic?th ?cieueiaaieo i?aaiae aceaeo iao??th. Aeacai? ci?ie
noi?iaiaeaeoaaeeny i?aeaeuaiiyi aia?aaoe/ii? aoaeoeaiino? aeeoaiiy.

Aeine?aeaeaiiy c ?cieueiaaieie i?oioiiae??yie aeyaeee aeoeaaoe?th
aeueoa?iaoeaii? ieneaeace i?e ioieiaeaeai? ?ineei i?ioyaii 24 aiae. I?e
a?eueo aeiaaio?eaae?e ae?? iecueeeo oaiia?aoo? niinoa??aa?oueny
cieaeaiiy aeoeaiino? aeueoa?iaoeaiiai oeyoo o?anii?oo aeaeo?ii?a.

Oaeei /eiii, aeine?aeaeaiiy aieeao iecueeeo oaiia?aoo? ia aeeoaeueiee
aia?aiiai?io ceae?a, i?iaaaeai? ia oeaieiiio oa noaee?oeiiio ??aiyo
i?aai?caoe?? na?aeaoue i?i aeoeaaoe?th aeueoa?iaoeaii? ieneaeace i?e
ae?? oieiaeo. Ca?eueoaiiy ?ioaineaiino? iiaeeiaiiy eenith /a?ac
aeueoa?iaoeaiee oeyo iiaea aooe iaoiiaeaia ?iaeoeoe??th neioaco
aeueoa?iaoeaii? ieneaeace [Ito et al., 1997], aai ae ?? aeoeaaoe??th
i?aai?/ieie eeneioaie [Wagner et.al.,1995; Day, Wiskich, 1995] ?/aai
aiane?aeie oeieueii? iiaeeo?eaoe?? oa?iaioo [Umbach, Siedow, 1993].

Cia/aiiy aeueoa?iaoeaiiai aeeoaiiy iiaea iieyaaoe a oiio, ui: 1)
iineeaia oaieii?iaeoeoe?y a i?ioean? o?ainii?oo aeaeo?ii?a ii
aeueoa?iaoeaiiio oeyoo caeaoia aaciina?aaeiuei caoeuaoe ee?oeie i?ioyaii
iaaiiai /ano [Moynihan et.al., 1995]; 2) iio?e aeaeo?ii?a /a?ac
aeueoa?iaoeaiee oeyo cieaeo? caeaoi?noue aei aaia?aoe?? aenieiai ??aiy
noia?ieneae?a oa H2O2 (Purvis and Shewfelt , 1993), yeee c?inoa? i?e
aeieoaaii? o?ainii?oo aeaeo?ii?a ii oeeoio?iiiiio eaioethao o oeaiei,
ui caciaee aieeao oieiaeo (Anderson et al., 1995); 3)aeoeaaoe?y oeueiai
oeyoo ni?ey? ooieoe?iioaaiith oeeeeo o?eea?aiiiaeo eeneio ca oiia
aenieiai ??aiy a?aeiiaeaiiino? eiiiiiaio?a oeeoio?iiiiai oeyoo [Wagner,
Krab, 1995; Day, Wiskich, 1995].

Aeniiaee

1. Iie?oinoaoeaeee?iiceoiee aa?ooue o/anoue o o?ainaeoeoe?? neaiae?a,
iaoiiaeaieo ??ceei iiieaeaiiyi oaiia?aoo?e na?aaeiaeua ee?oei ?ineei, ui
i?aeoaa?aeaeo?oueny ?iaeoeoe??th a?ae?ie?co oinoaoeaeee?iiceoie
4,5-a?oinoaoo oa oi?ioaaiiyi aoi?eiiiai iina?aaeieea o?ainaeoeoe??
neaiae?a: ?iiceoie 1,4,5-o?eoinoaoo o ee?oeiao ?ineei, i?ae aieeaii
ei?ioei/aniiai ioieiaeaeaiiy ia?inoe?a eoeo?oaece oa iceii? ioaieoe?.

2. Ia ?aii?o aoaiao ?icaeoeo ca?iaee?a a?aeaoaa?oueny neioac a?ee?a,
iia’ycaieo ?c ?aae?caoe??th i?ia?aie ?icaeoeo, ye ca iioeiaeueieo oiia,
oae ? i?e ae?? aeno?aiaeueieo oaiia?aoo?; neioac “no?aniaeo a?ee?a” a
ee?oeiao ?ineei a?aeaoaa?oueny eeoa i?ney i?ioiaeaeaiiy iaaieo noaae?e
?o ?icaeoeo.

3. No?ee?noue ?ineei aei ae?? iecueeeo oaiia?aoo? o?nii iia’ycaia ?c
iiaiooai?aiiyi a?ee?a, i?aaenoaaiee?a iieaeoey?ieo oaia?ii?a, i?i ui
na?ae/eoue iiyaa a niaeo?ao iiaineioaciaaieo a?ee?a i?oioiiae??e ?
oeeoiieacie i?i?inoe?a ioaieoe? oa eoeo?oaece, ui caciaee aieeao oieiaeo
a?ee?a, c iieaeoey?ieie ianaie 90 ? 70 eAe c ooieoe?yie iieaeoey?ieo
oaia?ii?a, a oaeiae iie?iaioeae?a a ae?eyioe? 17–38 eAe.

4. Ae?y iecueei? oaiia?aoo?e ia i?i?inoee eoeo?oaece aeeeeea?
ca?eueoaiiy ?ioaineaiino? oinoi?eethaaiiy o o?aeoe?? a?eea ye
i?oioiiae??aeueiiai, oae ? caaaeueiiai o ae?eyioe? 90 eAe, ui iiaea
a?aeiia?aeaoe ?iaeei? oaia?ii?a cpn90.

5. Ii?icino?ee? ni?oe iceii? ioaieoe? ?aaaothoue ia cieaeaiiy
oaiia?aoo?e iiaiooai?aiiyi niaoeeo?/iiai aei ae?? iecueei? oaiia?aoo?e
iie?iaioeaeo c iieaeoey?iith ianith aeecueei 120 eAe , neioac yeiai
neaai ae?aaeaiee o ni?o?a c iecueeei ??aiai ii?icino?eeino?; aeacaiee
a?eie oa?aeoa?eco?oueny aenieith ?ioaineai?noth oinoi?eethaaiiy o
niaeo?ao i?oioiiae??aeueiiai ? caaaeueiiai a?eea.

6. ?aaeoe?y iaoaaie?cio ee?oei ?ineei ia ae?th oieiaeo o?nii iia’ycaia
c? ci?iaie ooieoe?e i?oioiiae??e, ia?o ca ana ?c aeoeaaoe??th o?ainii?oo
aeaeo?ii?a ii aeueoa?iaoeaiiio oeyoo.

7. Oi?ioaaiiy aeueoa?iaoeaiiai oeyoo o?ainii?oo aeaeo?ii?a a
i?oioiiae??yo, ye ? oeeoio?iiiiai, a?aeaoaa?oueny a ee?oeiao ia?enoaie,
iaeneiaeueia aeoeai?noue aeueoa?iaoeaiiai oeyoo a?aei?/aia o ee?oei, ui
caaa?oeee ??no.

8. ?icaeoie ca?iaee?a ian?iiy eoeo?oaece i?e iecueeeo oaiia?aoo?ao
na?aaeiaeua o?nii ca’ycaiee ?c aeoeaaoe??th aeueoa?iaoeaiiai aeeoaiiy.

9. Aenieee ??aaiue oieiaeino?eeino? o?nii iia’ycaiee ?c aeoeaaoe??th
aeueoa?iaoeaii? ieneaeace i?ae aieeaii iecueeeo oaiia?aoo? a
i?oioiae??yo ee?oei no?eeeo aei ae?? oieiaeo e?i?e eoeo?oaece.

10. Ca?eueoaiiy ai?noo iniiaieo oinoie?i?ae?a, a oiio /ene?
oinoaoeaeeeoie?io ? oinoaoeaeeeaoaiieai?io a i?oioiiae??yo i?i?inoe?a
iceii? ioaieoe? a i?ioean? aaeaioaoe?? aei iecueeeo oaiia?aoo?
iai?aaeaia ia noaa?e?caoe?th aia?aaoe/iiai noaoono ee?oei ?ineei ca
iani?eyoeeaeo oiia na?aaeiaeua.

11. ?ioaineai?noue i?aeaeuaiiy ai?noo oinoie?i?ae?a a i?ioean?
aaeaioaoe?? eoeo?oaece a?aeiia?aea? ??aith oieiaeino?eeino? ie?aieo
i?aai?a. Iaea?eueo ?ioaineai? ci?ie a?aeaoaathoueny o a?eueo no?eeeo aei
oieiaeo eenoeao ?ineei.

12. Aaeaioaoe?y eoeoe?oaece aei iecueeeo oaiia?aoo? na?aaeiaeua
noi?iaiaeaeo?oueny c?inoaiiyi ??aiy iaiane/aiino? oinoie?i?ae?a.

Iniiai? ioae?eaoe?? ca oaiith aeena?oaoe??

1. E?aaaoe A.N. ?acaeoea i?aaenoaaeaiee ia aaeaioaoeee ?anoaiee e ieceei
oaiia?aoo?ai // Oeceieiaey e aeioeiey eoeueo.?anoaiee.- 1996.- O.28,
N3.- N.167-182.

2. E?aaaoeue A.N. ?ieue e?i?ae?a a aaeaioaoe?? ee?oei ?ineei aei
iani?eyoeeaeo oiia na?aaeiaeua // Aeooaeuei? i?iaeaie o?c?ieia??
aiaeiiai ?aaeeio ?ineei oa iinooino?eeino? ?ineei: Ca.iaoe. i?aoeue
?inoeoooo o?c?ieia?? ?ineei ? aaiaoeee IAI Oe?a?ie.- Ee?a, 1997.- N. 56-
59.

3. Kravets V. The alternative respiration of maize embryo tissues during
germination of seeds at low temperatures // Responses of plant
metabolism to air pollution and global change / Edited by L.J. De Kok
and I.Stulen.- Backhuys Publishers, Leiden, 1998.- P.355-358.

4. E?aaaoe A.N. Iieaeoey?iay aeieiaey aeueoa?iaoeaiie ieneaeacu
?anoaiee// Aeiiieeia?u e eeaoea.-1999.- O.15, N 2.- N.133-137.

5. E?aaaoe A.N. Aeeyiea ieceeo oaiia?aoo? ia iai?aaeaiiinoue iioiea
yeaeo?iiia a ieoioiiae?eyo ?anoaiee // E?eiaeieiaey.- 1999.- ? 2.- N.
34-37.

6. E?aaaoeue A.N.Aieea iecueeeo oaiia?aoo? ia o?ainii?o aeaeo?ii?a
aeeoaeueiiai eaioethaa i?oioiiae??e ?ineei // Oe?a?inueeee a?io?i?/iee
aeo?iae.- 1999.- N5.- C. 23-28.

7. Kravets V.S., Morgun V.V. Effect of low temperatures on the
alternative respiration of maize leaf tissues // Environment protection:
Modern studies in ecology and microbiology.- Uzhgorod, 1997.- P.
291-297.

8. E?aaaoe A.N., Aieieeia A.E. Aeaenoaea ieceeo oaiia?aoo? ia
aeoeaiinoue ?anoeoaeueiuo ieoioiiae?ee (Iaoiaee/aneea aniaeou) //
Onoie/eainoue ?anoaiee e aeaenoaeth io?eoeaoaeueiuo oaiia?aoo?:
Ca.iao/i. ?aaio ?inoeoooa oeceieiaee ?anoaiee e aaiaoeee IAI Oe?aeiu.-
Eeaa: Iaoe.aeoiea.,1984.- N.79-90.

9. E?aaaoeue A.N., Iio??ia E.I. ?aaeoe?y iaoaaie?cio
iie?oinoaoeaeee?iiceoie?a ia ae?th oieiaeiaiai oieo // Aeiiia?ae? AI
Oe?a?ie.- 1998.- N 11.- N. 166-169.

10. E?aaaoeue A.N., Iaei? I.N., Iieiaeie?i ?.A. Neioac a?ee?a
i?oioiiae??e ca?iaee?a eoeo?oaece i?e ?o ?icaeoeo a oiiaao ae?? aenieeo
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N.154-157.

11. Iaei? I.N.,Iieiaeie?i ?.A., E?aaaoeue A.N. Aieea iecueeeo oaiia?aoo?
ia caoiieaiiy oa neioac a?ee?a ao?ieueiaaieo i?i?inoe?a iceii? ioaieoe?.
// Oeceie. e aeioei. eoeueo. ?anoaiee.- 1999.- O.31, N 2.- N.130-134.

12. Iio??ia E.I., E?aaaoeue A.N. AEe?iieeneioiee neeaae oinoie?i?ae?a
eoeo?oaece a i?ioean? ?? aaeaioaoe?? aei oieiaeo ia ?aii?o aoaiao
iioiaaiaco // Oeceie. e aeioei. eoeueo. ?anoaiee. – 1998. – O.30, N 4. —
N. 288-293.

13. Iio??ia E.I., E?aaaoeue A.N. Ci?ie oinoie?i?aeiiai neeaaeo
i?i?inoe?a eoeo?oaece a i?ioean? caaa?ooaaiiy aei iecueeeo oaiia?aoo? //
Oeceie. e aeioei. eoeueo. ?anoaiee. – 1998.- O.30, N5.- C.368-373.

14. Eiia?ieoeeee I.A., Eoeueyiia E.N., Necia N.N., E?aaaoe A.N.,
Oaoa?eiia A.A., Eaaaaeaa N.E. Aeuoaiea, ?ino e ii?icinoieeinoue ii/ae
/a?aoie a oneiaeyo thaa Oe?aeiu // Oeceie. e aeioei. eoeueo. ?anoaiee.-
1981.- O.13, N5.- N.501-506.

15. Iaei? I.N., Iieiaeie?i ?.A. E?aaaoeue A.N. Neioac caaaeueieo oa
i?oioiiae??aeueieo a?ee?a iceii? ioaieoe? i?e ae?? iecueeeo oaiia?aoo?
oa ca oiia a?iien?? // Aeiiia?ae? IAI Oe?a?ie.- 1999.- N8.- N.152-156.

16. Iieiaeie?i ?.A., Iaei? I.N., E?aaaoeue A.N. Neioac
i?oioiiae??aeueieo a?ee?a eoeo?oaece ca oiia iecueeioaiia?aoo?iiai
aieeao// Oeceie. e aeioei. eoeueo. ?anoaiee.- 1999.- O.31, N4.-
N.250-254.

17. E?aaaoeue A.N. Oa?aeoa?enoeea oa ooieoe?? a?ee?a, ?iaeoeiaaieo
ae??th iecueeeo oaiia?aoo? // Oeceie. e aeioei. eoeueo. ?anoaiee.-
1999.- O.31, N5.- N.323-332.

18. E?aaaoe A.N., Eieioa I.E., Naiaithe A.A., Aa?eeaiei ?.N. Aeeyiea
aanoeeciaie eeneiou e aeeiaoeenoeueoieneaea ia ooieoeee ieoioiiae?ee e
ii?icionoie/eainoue iceiie ioaieoeu // Aiiaeiiaiai e onoie/eainoue
?anoaiee.- Ecae-ai EacAO, 1993.- N.110-116.

19. Aiaoo I.A., E?aaaoeue A.N. Ci?ie oinoi?iiceoeae?a iceii? ioaieoe? o
a?aeiia?aeue ia oaiia?aoo?iee no?an // Ca??iee iaoeiaeo i?aoeue
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20. Kravets V. Relationship between inhibition of respiration and frost
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for Plant Physiology.- Uppsala, Sweden, 12-17 June 1997.- P.125.

21. Kravets V. The effect of cold stress on the respiration of cereals
in vitro and in situ // International Workshop. Plant respiration:
physiological and ecological aspects.- Syktyvkar, 1995.- P.80-82.

22. E?aaaoe A.N. Ooieoeeiie?iaaiea aeueoa?iaoeaiie ieneaeacu
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23. Kravets V. Effect of low temperature stress on plants: physiological
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34(suppl.).- P. 76-77.

24. Kravets V.S., Semenyuk V.E. Response of plant mitochondria in situ
to extreme temperatures// International conference “Achievements and
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25. E?aaaoe A.N., *oaae I.TH., Iaeaia?/oe N.I. Oeeaieae?acenoaioiia
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Iaaeaeoia?.niaau. e neiiiceoia ia oaio: “Enneaaeiaaiea aeiaaiaca,
no?oeoo?u e ooieoeee oioineioaoe/aneiai aiia?aoa a nayce n
i?aia?aciaaieai nieia/iie yia?aee” (10-15 ieoya?y
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26. E?aaaoe A.N., Aieoaie/ay O.A., A?iieaiei E.I. Oeeaieae?acenoaioiia
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?anoaiee (24-29 naioya?y 1990 a. Ieine): Oacenu aeieeaaeia.- Ieine,
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27. E?aaaoe A.N., Ieaeiee E.I., ss?ioaiei I.A. Aeeyiea oieiaeiaiai oiea
ia oeeaieae?acenoaioiia aeuoaiea oieiaeionoie/eauo e /oanoaeoaeueiuo e
oieiaeo eeiee eoeo?ocu // Ananithciay eiioa?aioeey «Aaiaoe/aneea
iaoaieciu onoie/eainoe ?anoaiee e iaaeaaii?eyoiui oaeoi?ai n?aaeu (8-12
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28. Kravets V.S., Boreyko V., Yaroshenko N., Oleynik I. The effects of
sodium aride and benzylhydroxamic acid on the respiration of growing
tissues of roots of mayze (Zea mays L) // Physiologia Plantarum.-
1992.-Vol.85(3) Part 2:A53.

29. E?aaaoeue A.N, Ai?aeei A.N., ss?ioaiei I.A. Aieea iecueeeo
oaiia?aoo? ia aeoeai?noue aeueoa?iaoeaiiai oeyoo i?oioiiae??e e?i?e ?
a?a?eae?a eoeo?oaece // Oace ?? c’?caeo Oiaa?enoaa o?c?ieia?a
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30. Bucolova T.P., Volovik N.V., Kravets V.S. A possible role of
phosphatidylinositols in transduction of temperature signal in winter //
Biol. plant.- 1994.- 36.- P.45.

31. E?aaaoeue A.N., Iieiaeieei E.A., Iaei? I.N. N?aaieoaeueiia
enneaaeiaaiea neioaca ieoioiiae?eaeueiuo aaeeia a i?i?inoeao ioaieoeu
i?e aeaenoaee aunieeo e ieceeo oaiia?aoo? // Aeooaeuei? ieoaiiy
o?c?ieia?? ?ineei a aniaeo? aeieia?/ieo i?iaeai Oe?a?ie.- Ee?a, 1995.-
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32. E?aaaoe A.N., Iieiaeieei E.A., Iaeii? I.N. I aiciiaeiie ?iee
iieaeoey?iuo oaia?iiia a ?aaeoeee i?i?inoeia eoeo?ocu ia aeaenoaea
oieiaea // I c’?cae Oe?a?inueeiai oiaa?enoaa e??ia?ieia?? ?
e??iiaaeaoeeie: oace aeiiia?aeae.- Oa?e?a, 1995.- N.116-118.

33. Kravets V.S Polovinkin I.G., Major P.S. A possible of chaperones in
the cold stress response of maize seedlings mitochondria // J.Plant
Physiology and Biochemistry, Special iss.- 1996.- P.227.

34. Kravets V.S., Bucolova T.P., Volovik N.V. Phosphatidylinositols
changes in winter wheat in response to cold shock // Fifth International
Plants Cold Hardiness Seminar.- Oregon, USA, 5-8 Aug. 1996.- 1996.- P.
42.

35. Kravets V.S., Polovinkin I.G., Major P.S. A possible of chaperones
in the cold stress response of maize seedlings mitochondria // J.Plant
Physiology and Biochemistry, Special iss.- 1996.- P.227.

36. Krfvets V.S., Major P.S., Polovinkin I.G. Protein synthesis in maize
seedlings mitochondria under low temperature conditions // International
symposium on stress and inorganic nitrogen assimilation & the 2nd fohs
biostress symposium. — Moscow, 1996.- P.59.

37. Yankes E.P., Kravets V.S.- Effect of low-temperature stress on
phosphatidylinositol metabolism of chilling sensitive plants. //
International symposium on stress and inorganic nitrogen assimilation &
the 2nd fohs biostress symposium.- Moscow, 1996.- P.122.

E?aaaoeue A.N. “ Iniaeeaino? iaoaaie?cio ceae?a i?e ae?? iecueeeo
oaiia?aoo? ia ?ineeie” — ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaith aeieoi?a a?ieia?/ieo iaoe
ca niaoe?aeuei?noth 03.00.04- “A?io?i?y”. *a?i?aaoeueeee aea?aeaaiee
oi?aa?neoao ?i.TH. Oaaeueeiae/a, *a?i?aoe?, 1999.

Aeena?oaoe?th i?enay/aii aeine?aeaeaiith ei?ioeioa?i?iiaiai oa o?eaaeiai
aieeao iecueei? oaiia?aoo?e ia neeaae iie?oinoaoeaeee?iiceoie?a oeaiei
i?i?inoe?a eoeo?oaece oa iceii? ioaieoe?, neioac a?ee?a oa aeoeai?noue
oeeoio?iiiiai oa aeueoa?iaoeaiiai oeyo?a o?ainii?oo aeaeo?ii?a a
i?oioiiae??yo ceae?a.

Io?eiai? ?acoeueoaoe na?ae/aoue, ui ei?ioei/aniaee aieea iecueeeo
oaiia?aoo? cieaeo? a oeaieiao i?i?inoe?a ??aaiue ?aae?iaeoeaiino? o
o?aeoe?yo ?iiceoieoinoie?i?ae?a O?O oa O?O2 , a oie /an ye ??aaiue
?aae?iaeoeaiino? o o?aeoe?? ?iiceoieoinoao?a c?inoa?.

Anoaiiaeaii, ui aieea ye iecueeeo oae ? aenieeo oaiia?aoo? ia i?i?inoee
ceae?a iaoiiaeth? i?aeaeuaiiy ??aiy neioaco a?oie a?ee?a a ae?aiacii?
(17-35 oa 58-120 eAe), aeeth/ath/e AOO70 ? AOO90. C’yniaaii, ui
?iaeoeiaaiee ci?iith oaiia?aoo?e neioac a?ee?a i?oioiiae??e a?eueo /?oei
ae?aaeaiee a ee?oeiao, ye? caaa?oeee ??no, ii??aiyii c ai?eaeueiith
ia?enoaiith.

Iieacaii, ui aaeaioaoe?y eoeo?oaece aei iecueeeo oaiia?aoo? na?aaeiaeua
noi?iaiaeaeo?oueny iaeiie/aiiyi a ?? oeaieiao a?oie oinoie?i?ae?a oa
i?aeaeuaiiyi ??aiy iaiane/aiino? ?o aee?ieo eeneio.

Cieaeaiiy oaiia?aoo?e ca?eueoo? caeiyo?noue aeueoa?iaoeaiiai oeyoo
o?ainii?oo aeaeo?ii?a i?oioiiae??e eoeo?oaece.

Iaaiai?th?oueny ?ieue oinoaoeaeee?iiceoieueii? nenoaie a o?ainaeoeoe??
neaiae?a a ?aaeoe?? ee?oei ?ineei oa cia/aiiy iieaeoey?ieo oaia?ii?a a
ee?oeiao ?ineei i?e ae?? iecueeeo oaiia?aoo?.

Eeth/ia? neiaa: iecuee? oaiia?aoo?e, o?ainaeoeoe?y neaiae?a,
iie?oinoaoeae?e?iiceoiee, aaeaioaoe?y aei oieiaeo, oaia?iie,
i?oioiiae???, aeeoaiiy.

E?aaaoe A.N. “Iniaaiiinoe iaoaaieecia ceaeia i?e aeaenoaee ieceeo
oaiia?aoo? ia ?anoaiey ”. -?oeiienue

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie aeieoi?a aeieiae/aneeo iaoe ii
niaoeeaeueiinoe- 03.00.04- “Aeioeiey”. *a?iiaeoeeee ainoaea?noaaiiue
oieaa?neoao ei.TH. Oaaeueeiae/a, *a?iiaoeu, 1999.

Aeenna?oaoeey iinayuaia enneaaeiaaieth e?aoeia?aiaiiiai e
i?iaeieaeeoaeueiiai aicaeaenoaey ieceie oaiia?aoo?u ia ninoaa
iieeoinoaoeaeeeeiiceoieia oeaiae i?i?inoeia eoeo?ocu e iceiie ioaieoeu,
neioac aaeeia e aeoeaiinoue oeeoio?iiiiai e aeueoa?iaoeaiiai iooae
o?ainii?oa yeaeo?iiia a ieoioiiae?eyo ceaeia, a oaeaea aeeyiea ieceeo
oaiia?aoo? ia iiaeiuaiea eenei?iaea oeaiyie ca?iaeuoae ei?iae eoeo?ocu,
iaoiaeyueony ia ?acee/iuo noaaeeyo ?acaeoey.

Aaoi?ii noi?ioee?iaaia eiioeaioeey iineaaeiaaoaeueiinoe ?acaeoey
niauoee a eeaoeao ?anoaiee i?e iiieaeaiee oaiia?aoo?u n?aaeu,
iniiaaiiay ia aia?aua iieo/aiiuo oaeoao oi?ie?iaaiey aoi?e/iuo
iin?aaeieeia a eeaoeao ceaeia i?e aeaenoaee oieiaea e eo ?iee a
eiaeoeoeee aaeaioaoeeiiiuo eciaiaiee iai?aaeaiiinoe e eioaineaiinoe
iaoaaieecia eeaoie, eioi?ua aeeth/atho iiaiia?aciaaiea aaeeia ec eeanna
iieaeoey?iuo oaia?iiia, neioac eeieaeia e aeoeaaoeeth aeanaoo?ac.

Aia?aua auyaeaiu eciaiaiey ioea iieeoinoaoeaeeeeiiceoieia iiae aeeyieai
e?aoeia?aiaiiiai ioeaaeaeaiey i?i?inoeia aea?eaeia eoeo?ocu e iceiie
ioaieoeu, iieacai aeae?ieec oinoaoeaeeeeiiceoie 4,5-aeoinoaoa ia
eiiceoie 1,4,5-o?eoinoaoa e aeeaoeeeaeeoea?ie i?e aeaenoaee oieiaeiaiai
oiea.

Onoaiiaeai oa?aeoa? oi?ie?iaaiey aeueoa?iaoeaiiai iooe o?ainii?oa
yeaeo?iiia a eeaoeao ia?enoaiu, iieacaii oaaee/aiea aai o/anoey a
o?ainii?oa yeaeo?iiia e eenei?iaeo o caaa?oeaoeo ?ino eeaoie ?anoaiee.

I?ineaaeaii aeeiaieeo eciaiaiee aeoeaiinoe oeeoio?iiiiai e
aeueoa?iaoeaiiai aeuoaiey a i?ioeanna ?acaeoey ca?iaeuoae a naiaiao
eoeo?ocu i?e ieceeo oaiia?aoo?ao n?aaeu. Auyaeaii aeoeaaoeeth
aeueoa?iaoeaiie ieneaeacu a ieoioiiae?eyo eeaoie onoie/eauo e aeaenoaeth
oieiaea eeiee eoeo?ocu iiae aeeyieai ieceeo oaiia?aoo?. Iieo/aiu
?acoeueoaou, eioi?ua naeaeaoaeuenoaotho i oii, /oi aeoeaaoeey
aeueoa?iaoeaiiai aeuoaiey inouanoaeyaony ca n/ao neioaca aaeea
aeueoa?iaoeaiie ieneaeacu.

Iieacaii, /oi aeaenoaea ieceeo oaiia?aoo? iaoneaaeeaaao oaaee/aiea
aeueoa?iaoeaiiai e oeeoio?iiiiai aeuoaiey oeaiae ei?iae o/anoea 10-20
ii. Ionoonoaea eciaiaiee a aeoeaiinoe enneaaeoaiuo oa?aeoa?enoee
aeuoaiey eeaoie ia?enoaiu iiae aeeyieai ieceeo oaiia?aoo?
naeaeaoaeuenoaoao i oii, /oi a ia?eiae eioaineaiiai aeaeaiey eeaoie, eo
iaoaaieeci, a oii /enea aeuoaoaeueiue, iai?aaeai ia ?aaeecaoeeth
i?ioeannia ?inoa eeaoie.

Yoeiee?iaaiiua 2,5 nooi/iua i?i?inoee iceiie ioaieoeu ioeaaeaeaee i?e 3N
a oa/aiea 24-144 /ania. Iinea 24 /ania aeaenoaey ieceeo oaiia?aoo? ia
?anoaiey, eioaeoiua ieoioiiae?ee iieacuaaee aunieoth noaiaiue aeoeaaoeee
aeueoa?iaoeaiie ieneaeacu, eioi?oth ii?aaeaeyee eae onoie/eaia e
aeaenoaeth aceaea, /oanoaeoaeueiia e aaicaeae?ienaiaoo aeuoaiea i?aaiaee
n noeoeeiaoii a ea/anoaa noano?aoa.

?acoeueoaou enneaaeiaaiee aeuoaoaeueiiai yia?aiiaiaia in vivo e in vitro
iicaieeee auyaeoue iaoaieci ?aaoeyoeee iioiea yeaeo?iiia a
ieoioiiae?eyo, iaania/eee ?acaeoea i?aaenoaaeaiee i iooyo oi?ie?iaaiey
yia?aaoe/aneiai noaoona ceaeia iiae aeaenoaeai oieiaeiaiai oiea.
Ianoaeaeaaony ?ieue aeueoa?iaoeaiie ieneaeacu a nieaeaiee o?iaiy
oi?ie?iaaiey ?aaeoeaiuo oi?i eenei?iaea a oeaiyo ?anoaiee, eioi?ua auee
iiaeaa?aioou aeaenoaeth no?anna.

Onoaiiaeaii, /oi ?anoaiey ?aaae?otho ia iinoaiaiiia nieaeaiea oaiia?aoo?
eiaeoeoeeae eee iiauoaieai o?iaiy neioaca no?anniauo aaeeia, a oii
/enea niaoeeoe/aneeo ia aeaenoaea oieiaea. Ianoaeaeaaony ?ieue
ieceiiieaeoey?iuo AOO, AOO70, AOO90 a i?ioeannao aaeaioaoeee ?anoaiee e
aeaenoaeth oieiaea. Iieacaia ?ieue iieaeoey?iuo oaia?iiia a
iiaeaea?aeaiee ooieoeee eeaoie a i?ioeanna ?inoa e ?acaeoey ?anoaiee i?e
ieceeo oaiia?aoo?ao, iaania/aiee onoie/eainoe eeaoie e aeaenoaeth
oieiaea.

Aaoi? aeaeaao caeeth/aiea i oii, /oi iieaeoey?iua oaia?iiu ea?atho
eeth/aaoth ?ieue a onoie/eainoe eeaoie ?anoaiee e aeaenoaeth oieiaea e
caii?aaeeaathueo oaiia?aoo?.

Aia?aua onoaiiaeaii iiaiia?aciaaiea niaoeeoe/aneiai e aeaenoaeth ieceie
oaiia?aoo?u iieeiaioeaea n iieaeoey?iui aanii ieiei 120 eAe a oeaiyo
i?i?inoeia ii?icinoieeeo ni?oia iceiuo ioaieoe, neioac, eioi?iai aue
neaai au?aaeai o ni?oia n ieceei o?iaiai ii?icionoie/eainoe.

Oaaee/aiea eioaineaiinoe oinoi?eee?iaaiey ai o?aeoeee eae
ieoioiiae?eaeueiiai, oae e iauaai aaeea i?i?inoeia eoeo?ocu auyaeaii,
i?aaeaea anaai, a o/anoea 90 eAe, /oi iiaeao niioaaonoaiaaoue naiaenoao
oaia?iiia cpn90.

Eeth/aaua neiaa: ieceea oaiia?aoo?u, o?ainaeoeoeey neaiaeia,
iieeoinoaoeaeeeeiiceoieu, aaeaioaoeey e oieiaeo, oaia?iiu, ieoioiiae?ee,
aeuoaiea.

Kravets V.S. “Features of metabolism of cereal plants under low
temperature conditions”. Dissertation thesis as manuscript is presented
for academic degree the Doctor of sciences (Biology) on specialty
03.00.04- “Biochemistry”, Yu.Fedkovych Chernivtsy State University,
Chernivtsy, 1999.

Dissertation thesis is devoted to the investigations the effects of
short-term and prolonged period of low temperature on
polyphosphatidylinositol phospholipids composition of maize and winter
wheat seedlings tissues, synthesis of polypeptides and activity of
cytochrome and alternative pathway of electron transport of cereals
mitochondria.

The results obtained show that under low temperature stress the level
of radioactivity in inositolphospholipids PIP and PIP2 sharp decreased,
while the level of radioactivity in inositol phosphates increased during
10 min. incubation of seedlings. The results show that
polyphosphoinositides take part in early   responsible reactions of
plant cells to cold stress.

The study has revealed that exposure of seedlings both to cold and to
heat caused the increasing of level synthesis of a common group of
proteins (17-35 and 58-120 kD) including heat shock protein 70 and
90kD.   Presented results of in vivo radiolabeling experiments indicate
that the temperature-induced changes in mitochondrial newly
synthesized proteins are more expressed in the whole stems than in
the apical meristem.

It has been found that low temperature acclimation of maize was
accompanied by the accumulation in the tissues of the main groups of
phospholipids and increasing the unsaturation of their fatty acid
chains. Lower temperatures increased the degree of engagement of the
alternative pathway of electron transport in maize mitochondria.

The role phosphoinositide system in the transduction of temperature
signals into cellular reactions of plant cells and signification of
molecular chaperones in the plant cells under low temperature stress is
discussed.

Key words: low temperature, signal transduction,
polyphosphatidylinositols, cold acclimation, chaperones, mitochondria,
respiration.

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