OA?E?ANUeEEE AeA?AEAAIEE OI?AA?NEOAO

EOCI?OeIA ?AAI A?EOI?IAE*

OAeE. 612.07.2:577.1

IIEAEOEss?I? IAOAI?CIE AAeAIOAOe?? EIEAAAIIAEO NO?OEOO? AeI CI?I
OAIIA?AOO?E

03.00.04 — a?io?i?y

Aaoi?aoa?ao aeena?oaoe??

ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa a?ieia?/ieo iaoe

Oa?e?a – 1998

Aeena?oaoe??th ? ?oeiien.

?iaioa aeeiiaia o IAe* Oa?e?anueeiai aea?aeaaiiai oi?aa?neoaoo,

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

Iaoeiaee ea??aiee:

— aeieoi? a?ieia?/ieo iaoe, noa?oee iaoeiaee ni?a?ia?oiee, Ia?nueeee
?aaai Ao?i?iiae/, i?ioani? eaoaae?e o?c?ieia?? ethaeeie oa oaa?ei
Oa?e?anueeiai aea?aeaaiiai oi?aa?neoaoo.

Io?oe?ei? iiiiaioe:

— aeieoi? a?ieia?/ieo iaoe, i?ioani?, Iao?aiei Ieaenaiae? TH?ue?ae/,
i?ia?aeiee iaoeiaee ni?a?ia?oiee a?aeae?eo e??iiaaeeoeeie ?inoeoooo
i?iaeai e??ia?ieia?? oa e??iiaaeeoeeie IAI Oe?a?ie;

— aeieoi? a?ieia?/ieo iaoe, noa?oee iaoeiaee ni?a?ia?oiee, Oeiioaiei
Ieueaa Iaae?aia, caa. eaai?aoi???th a?io?i?? ? aenia?eiaioaeueiiai
iiaeaethaaiiy Oa?e?anueeiai iaoeiai-aeine?aeiiai ?inoeoooo i?oiiaae?? ?
o?aaiaoieia?? ?i. I.?. N?oaiei IIC Oe?a?ie.

I?ia?aeia i?aai?caoe?y — ?inoeooo a?io?i?? ?i. I.A. Iaeeaae?ia IAI
Oe?a?ie, i. Ee?a.

Caoeno a?aeaoaeaoueny (2( a?oaeiy 1998 p. o 1515 aiaeei? ia can?aeaii?
niaoe?ae?ciaaii? a/aii? ?aaee E 64.051.17 Oa?e?anueeiai
aea?aeaaiiai oi?aa?neoaoo, ca aae?anith: 310077, i. Oa?e?a, ie.
Naiaiaee, 4, aoae. ???-15.

C aeena?oaoe??th iiaeia iciaeiieoeny o Oeaio?aeuei?e iaoeia?e
a?ae?ioaoe? Oa?e?anueeiai aea?aeaaiiai oi?aa?neoaoo: 310077, i. Oa?e?a,
ie. Naiaiaee, 4

Aaoi?aoa?ao ?ic?neaiee (2( _eenoiiaaea 1998 ?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee,

eaiaeeaeao a?ieia?/ieo iaoe
Iaaeaeei A.?.

Caaaeueia oa?aeoa?enoeea ?iaioe

Aeooaeuei?noue ?iaioe. Niieo/ia oeaieia a?ae?a?a? aaaeeeao ?ieue o
i?ioeanao aaeaioaoe?? aaaaoiee?oeiieo oaa?ei aei oaiia?aoo?e
iaaeieeoiueiai na?aaeiaeua. O cia/i?e i??? oe? i?ioeane caaacia/othoueny
no?oeoo?iith noaa?euei?noth eieaaaio – iniiaiiai eiino?oeoe?eiiai
eiiiiiaioa niieo/ii? oeaieie.

A aaaaoueio ?iaioao aoei iieacaii, ui no?oeoo?ia noaa?euei?noue
eieaaaio, yea ioe?ith?oueny ca oaiia?aoo?ith eiai aeaiaoo?aoe??,
aeecueea aei aa?oiuei? iaae? oaiia?aoo?e na?aaeiaeua iaoeaiiy o
iiee?eioa?iieo oaa?ei /e oaiia?aoo?e o?ea – o aiiieioa?iieo
[Ao?aeaeaiaaeca, 1989-1993, Aneiiaa, 1991, 1995, Nhellan , 1993, Rigby,
1990].

A iniia? oe??? caeaaeiino? ciaoiaeeoueny ei?aeyoe?y i?ae oaiia?aoo?ith
aeaiaoo?aoe?? iieaeoe oa ai?noii o ieo ?i?iieeneio, cie?aia a?aeiinii?
e?eueeino? na?aae ieo caeeoe?a ienei?ie?io [Ao?aeaeaiaaeca, 1992, Berg,
1993, Nemethy, 1986, Ramachandran, 1973]. Noia?ia e?euee?noue
?i?iieeneio o eieaaai? oaa?ei ??cieo aeae?a, ui aeeaooue iei?, ?
iane?aeeii o?eaaei? iieaeoey?ii? aaiethoe?? ? cae??ieaia aaiaoe/ii.
Ai?no ienei?ie?io aecia/a?oueny ?ioaineai?noth a?ae?ieneethaaiiy
caeeoe?a i?ie?io, yea cae?enith?oueny oa?iaioii i?ie?ea?ae?ieneeacith a
i?ioean? iinoo?aineyoe?eii? iiaeeo?eaoe?? ia?aeiii? no?oeoo?e eieaaaio
[Eichner, 1988]. Ne?ae i?aee?aneeoe, ui no?oeoo?ia noaa?euei?noue
iaaeiieaeoey?ieo ooai?aiue niieo/ii? oeaieie caeaaeeoue ? a?ae
caaaeueii? eiioeaio?aoe?? eieaaaio, a oaeiae ? a?ae ?ioeo
iinoo?aineyoe?eieo oa iinoneioaoe/ieo eiai iiaeeo?eaoe?e:
ae?eiceethaaiiy , ui aaaea aei ooai?aiiy eieaaai-iie?naoa?eaeieo
eiiieaen?a [Kimura, 1992]; iiia?a/ia ca(ycoaaiiy [Kivirikko, 1990,
Richard-Blum, 1994] oa aca?iiae?y c ?ioeie no?oeoo?ieie eiiiiiaioaie
niieo/ii? oeaieie. On? oe? no?oeoo?i? oa?aeoa?enoeee eieaaaiiaeo
ooai?aiue, ye ? nooi?iue a?ae?ieneethaaiiy caeeoe?a i?ie?io,
aaciina?aaeiuei aaiaoe/ii ia cae?(ieai? ? aecia/athoueny aeoeai?noth
oa?iaio?a, ye? cae?eniththoue i?ioean(ia eieaaaio.

Oaeei /eiii, aaiethoe?eii cae??ieaia ei?aeyoe?y i?ae ai?noii
ienei?ie?io a eieaaai?, eiai no?oeoo?iith noaa?euei?noth ? oaiia?aoo?ith
iaaeieeoiueiai na?aaeiaeua ? eeoa iaeiei c aniaeo?a oaiia?aoo?ii?
aaeaioaoe?? oaa?ei ? a?aeiineoueny aei aeae?a, ye? aaea i?ioyaii
aeiaaiai /ano aeeaooue o iino(eieo oaiia?aoo?ieo oiiaao.

A oie aea /an iiee?eioa?ii? oaa?eie na?aaeiuei? nioae a?ae/oaathoue
naciii? ci?ie oaiia?aoo?e, ui aeinyaathoue aeanyoe?a a?aaeon?a. Oiio
iiaeii? ?nioaaoe ? ?io? — oaeaee? iaoai?cie aaeaioaoe?? eieaaaio aei
cia/ieo ci?i oaiia?aoo?e iaaeieeoiueiai na?aaeiaeua. Iaoai?cie oaeiai
oeio iiaeooue aeei?enoiaoaaoe ? a?aa?iaioi? nnaaoe?, oaiia?aoo?a o?ea
yeeo i?e ia?aoiae? a?ae noaio aaaeuei?ino? aei noaio niey/ee oa iaaiaee
nooo?ai ci?ith?oueny i?ioyaii ei?ioeiai /ano.

Oaeei /eiii, caea?oueny iaaiei, ui oaiia?aoo?a iaaeieeoiueiai
na?aaeiaeua oeyoii ae?? ia aeoeai?noue a?aeiia?aeieo oa?iaio?a neioaco,
i?ioean?iao oa ?iciaaeo eieaaaio iiaea aaciina?aaeiuei aieeaaoe ia
ia?aaoaeiao eiai iaaeiieaeoey?ieo ooai?aiue ?, ye iane?aeie, ci?ithaaoe
?o no?oeoo?io noaa?euei?noue.

I?ioa, ii oeae /an ieoaiiy i?i aieea oaiia?aoo?e iaaeieeoiueiai
na?aaeiaeua ia ?ioaineai?noue oeeo noaae?e iai?io eieaaaio i?aeoe/ii ia
aeine?aeaeoaaeinue. I?ae oei eiai aea/aiiy aeicaieeei a aena?oeoe ia
eeoa iieaeoey?i? iaoai?cie oaeaeei? aaeaioaoe?? eieaaaiiaeo ooai?aiue
iiee?eioa?iieo oa a?aa?iaioieo oaa?ei aei oaiia?aoo?e ciai?oiueiai
na?aaeiaeua, a e aianoe iaaio yni?noue o ieoaiiy i?i aeieeiaiiy oa
aaiethoe?th oeeo iaoai?ci?a o o?aaaoieo, a oiio /ene? aiiieioa?iieo
oaa?ei.

Iaea?aeai? ?acoeueoaoe oaeiae iiaeooue aooe aaaeeeaeie ? aeey ?icaeoeo
oyaeaiue i?i caaaeuei? iaoai?cie oaiia?aoo?ii? aaeaioaoe??.

Ca(ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. ?iaioa aoea
aeeiiaia o IAe* Oa?e?anueeiai aea?aeaaiiai oi?aa?neoaoo a iaaeao oaie
“No?oeoo?ii-iaoaaie?/i? iaoai?cie aaeaioaoe?? niieo/ii? oeaieie aei ae??
aeno?aiaeueieo oaeoi??a’’, ? aea?ae?a?no?aoe?? 0197U003060.

Iaoa oa caaaeaiiy aeine?aeaeaiiy. Anoaiiaeoe, ye aieeaa? oaiia?aoo?a
na?aaeiaeua ia neioac, i?ioean?ia, ?iciaae ? no?oeoo?io noaa?euei?noue
eieaaaio o?aaaoieo oaa?ei ??cieo nenoaiaoe/ieo a?oi in vivo oa in
vitro. C’ynoaaoeaaoe, ye? iieaeoey?i? iaoai?cie eaaeaoue a iniia?
oaeaeei? no?oeoo?ii? aaeaioaoe?? eieaaaio aei oaiia?aoo?e. Ii??aiyoe ?o
iniaeeaino? o iiee?eioa?iieo (?ea), a?aa?iaioieo (eaaeai?a) oa
aiiieioa?iieo (uo??a) oaa?ei c iaoith aeyaeaiiy iniiaieo caeiiii??iinoae
aaiethoe?? oeeo iaoai?ci?a. Aeey aeinyaiaiiy iaoe aoee iinoaaeai? oae?
caaea/?:

Aea/eoe oaiia?aoo?i? caeaaeiino? neioaco, a?ae?ieneethaaiiy caeeoe?a
i?ie?io, ae?eiceethaaiiy, iiia?a/iiai ca(ycoaaiiy oa ?iciaaeo eieaaaio a
i?aaiao ?ea in vivo (aaiaoc?y cae/aeia) oa in vitro (ea?i cae/aeiee),
a?eeo uo??a (e?i?y A?noa?) oa eaaeai?a (aaeeeee i?aeeiaii?n) in vitro,
eiee oaiia?aoo?ia ?aaoeyoe?y, oa?aeoa?ia aeey nnaaoe?a, ciyoa.

Aeine?aeeoe oaiia?aoo?io caeaaei?noue ?ic/eiiino? oa oa?iinoaa?eueiino?
eieaaaio ? aeyaeoe ?o ei?aeyoe?th c ??cieie noaae?yie iai?io eieaaaio.

Aeyaeoe iieaeoey?i? iaoai?cie, ui eaaeaoue a iniia? oaiia?aoo?ieo
caeaaeiinoae iai?io oa no?oeoo?e eieaaaio o?aaaoieo oaa?ei ??cieo
nenoaiaoe/ieo a?oi.

I?iaanoe ii??aiyeueiee aiae?c iaoai?ci?a no?oeoo?ii-iaoaaie?/ii?
aaeaioaoe?? eieaaaio aei oaiia?aoo?e o iiee?eioa?iieo, a?aa?iaioieo oa
aiiieioa?iieo oaa?ei. Ciaeoe caaaeuei? caeiiii??iino? oeeo i?ioean?a oa
?icaeyiooe iiaeeeao noaio ?o aaiethoe??.

Iaoeiaa iiaecia ?iaioe. Aia?oa iieacaii, ui oaiia?aoo?ia caeaaei?noue
?ioaineaiino? neioaco eieaaaio a aeine?aeao in vitro o an?o
aeine?aeaeaieo i?aaiao iaio aea/aieo aeae?a nnaaoe?a ia? aeaiai?aee
oa?aeoa? c iaeneioiaie a iaeanoyo 10-15 0N oa 25-35 0N. ?ioaineai?noue
a?ae?ieneethaaiiy i?ie?io ia? aiaeia?/iee oa?aeoa?, cie?aia nooi(iue
a?ae?ieneethaaiiy a on?o aeiaaeeao aeuee a ae?oa?e iaeano? . Nooi?iue
ae?eiceethaaiiy eieaaaio (ye oa?iaioaoeaiiai, oae ? iaoa?iaioaoeaiiai)
oaeiae ? iaeneiaeueiei a iaeano? 35 0N. Ciaeaeaii cai?ioith
caeaaei?noue i?ae oaiia?aoo?ith na?aaeiaeua ? ?ioaineai?noth
aooieieaaaiie?co, iaeneioi yeiai i?eiaaea? ia 10-15 0N. Aia?oa iieacaii,
ui nooi(iue iiia?a/iiai ca’ycoaaiiy a iaaeiieaeoey?ieo eieaaaiiaeo
ooai?aiiyo i?aeaeuo?oueny c? c?inoaiiyi oaiia?aoo?e na?aaeiaeua.
Anoaiiaeaii, ui oa?iinoaa?euei?noue eieaaaio, yea ioe?ith?oueny ca
oaiia?aoo?ith oa aioaeuei??th eiai aeaiaoo?aoe??, aeua aeey iieaeoe,
neioaciaaieo in vitro i?e 35 0N, i?ae aeey iieaeoe, neioaciaaieo i?e 10
0N, ? ei?aeth? c a?eueo aenieei nooiaiai a?ae?ieneethaaiiy,
ae?eiceethaaiiy oa iiia?a/iiai ca’ycoaaiiy eieaaaio, ui niinoa??aa?oueny
i?e oaiia?aoo?? 35 0N.

Aia?oa iieacaii, ui o iiee?eioa?iieo o?aaaoieo ci?ia oaiia?aoo?e
iaaeieeoiueiai na?aaeiaeua, a o a?aa?iaioieo nnaaoe?a ci?ia oaiia?aoo?e
o?ea aaciina?aaeiuei ?aaoeth? aoaeiao oa no?oeoo?io noaa?euei?noue
eieaaaiiaeo ooai?aiue oeyoii ae?? ia ?ioaineai?noue i?ioean?a neioaco,
i?ioean?iao oa ?iciaaeo eieaaaio. Oae? ae nai? no?oeoo?ii-iaoaaie?/i?
iaoai?cie oaiia?aoo?ii? aaeaioaoe?? eieaaaio ciaeaeai? ? o aiiieioa?iieo
nnaaoe?a a oiiaao in vitro.

Aia?oa iieacaii, ui a oiiaao in vitro i?ioeane no?oeoo?ii-iaoaaie?/ii?
ia?aaoaeiae eieaaaiiaeo ooai?aiue iiae?ai? a niieo/i?e oeaiei? oaa?ei
??cieo nenoaiaoe/ieo a?oi, ui iathoue ( ??ci? oeie oaiia?aoo?ii?
?aaoeyoe??.

Oai?aoe/ia oa i?aeoe/ia cia/aiiy ?iaioe. I?iaaaeai? aeine?aeaeaiiy
aeicaieeee aeyaeoe iieaeoey?i? iaoai?cie oaeaeei? aaeaioaoe??
eieaaaiiaeo ooai?aiue aei oaiia?aoo?e ciai?oiueiai na?aaeiaeua. Aeyaeai?
aoaie iai?io eieaaaio, ye? aaciina?aaeiuei ?aaoeththoueny oaiia?aoo?ith
? aaaeooue aei a?aeiia?aeiino? i?ae no?oeoo?iith noaa?euei?noth
niieo/ii? oeaieie ? oaiia?aoo?ith iaaeieeoiueiai na?aaeiaeua. Oe?
?acoeueoaoe aeicaieythoue aeiiiaieoe no/ani? iiaeyaee ia iieaeoey?i?
iaoai?cie aaeaioaoe?? oaa?ei aei oaiia?aoo?e ? iiaeooue aooe ei?enieie o
?ic?iaoe? oyaeaiue i?i iioiaeaeaiiy oa aaiethoe?th oeeo iaoai?ci?a.

Io?eiai? ?acoeueoaoe aeicaieythoue aeiiiaieoe aeaye? oyaeaiiy i?i
iieaeoey?i? iaoai?cie aaeaioaoe?? a?ee?a aei oaiia?aoo?e na?aaeiaeua.
Aiie oaeiae iiaeooue iaoe cia/aiiy i?e aee?iaoe?? oaa?ei oa o aeieia??.

Iniaenoee aianie caeiaoaa/a: i?aea??, ia?iaea oa aiae?c e?oa?aoo?ieo
aeaieo; iinoaiiaea iaoeiai? caaea/?; aea?? ? ?aae?caoe?y aiae?oe/ieo
iaoiae?a ?? ae??oaiiy; o/anoue o ?ic?iaoe? noaie iiaeeeaeo iaoai?ci?a
no?oeoo?ii? aaeaioaoe?? eieaaaiiaiai iao?eeno niieo/ii? oeaieie oaa?ei c
??cieie oeiaie oa?ii?aaoeyoe?? aei ci?ie oaiia?aoo?e iaaeieeoiueiai
na?aaeiaeua; aiae?c oa ?ioa?i?aoaoe?y iaea?aeaieo ?acoeueoao?a.

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. ?acoeueoaoe ? iniiai? iieiaeaiiy
?iaioe aeiiia?aeaeenue ia: XIV c(?cae? Oe?a?inueeiai o?c?ieia?/iiai
oiaa?enoaa (Ee?a, 1994), neiiic?oi? “Aeieiae/aneea iaoaieciu noa?aiey”
(Oa?e?a, 1994), ?? eiia?an? iaoio?c?ieia?a Oe?a?ie (Ee?a, 1996), ??
Oe?a?inueeiio a?io?i?/iiio c(?cae? (Ee?a, 1997), OV c(?cae?
Oe?a?inueeiai o?c?ieia?/iiai oiaa?enoaa (Aeiiaoeuee, 1998), ???
I?aeia?iaeiio neiiic?oi? (Aeieiae/aneea iaoaieciu noa?aiey( (Oa?e?a,
1998), ?? c(?cae Oe?a?inueeiai a?io?ce/iiai oiaa?enoaa (Oa?e?a, 1998).

Ioae?eaoe??. Ca ?acoeueoaoaie aeena?oaoe?? iioae?eiaaii 11 ?ia?o.

No?oeoo?a oa ianya aeena?oaoe??. Aeena?oaoe?eia ?iaioa neeaaea?oueny c
anooio, ?icae(e?a: e(oa?aoo?iee iaeyae, iaoa??aee oa iaoiaee
aeine?aeaeaiue, ?acoeueoaoe aeanieo aeine?aeaeaiue oa ?o iaaiai?aiiy,
i?aenoieiaeo aeniiae?a oa ia?ae?eo aeei?enoaii? e?oa?aoo?e ?c 220
iaeiaioaaiue. ?iaioo aeeeaaeaii ia 137 noi??ieao ae?oeiaaiiai oaenoo,
aiia i?noeoue 5 oaaeeoeue ? 18 ?enoie?a.

?icae?e 1. E?oa?aoo?iee iaeyae

O ia?oiio i?ae?icae?e? e?oa?aoo?iiai iaeyaeo iaaaaeaii oai?aoe/i? oa
aenia?eiaioaeuei? aeai? i?i no?oeoo?o eieaaaiiaeo ooai?aiue niieo/ii?
oeaieie. ?icaeyiooi ?acoeueoaoe aea/aiiy neioaco, iinoo?aineyoe?eieo oa
iinoneioaoe/ieo iiaeeo?eaoe?e oeeo niieoe ia ??cieo ??aiyo ?o
i?aai?caoe??. Iieacaii. yeo ?ieue a?ae?a?a? eieaaai o noai?aii?
niieo/ii-oeaieiiiai iao?eeno. O ae?oaiio i?ae?icae?e? iaaaaeai? aeai?
i?i aeaye? iaoai?cie oaiia?aoo?ii? aaeaioaoe?? oaa?ei. ?icaeyaeathoueny
iaoai?cie aaeaioaoe?? a?ee?a aei ??cieo oaiia?aoo?ieo oiia. Iaaaaeai?
aeai? i?i ei?aeyoe?th i?ae no?oeoo?ith eieaaaiiaeo ooai?aiue oa
oa?iiaeeiai?/ieie ia?aiao?aie eiai aeaiaoo?aoe?? aeey oaa?ei, ui
aeeaooue i?e iino?eieo oaiia?aoo?ieo oiiaao.

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

Aenia?eiaioe aoee i?iaaaeai? ia o?aaaoieo oaa?eiao, ui a?aeiinyoueny aei
o?ueio nenoaiaoe/ieo a?oi c ??ciei oeiii oa?ii?aaoeyoe??.
I?aaenoaaieeaie iiee?eioa?iieo aoee ?eae aeaio aeae?a: a aeine?aeao in
vivo – aaiaoc?y cae/aeia (Gambusia affinis), in vitro – ea?i cae/aeiee
(Cyprinus carpio). I?aaenoaaieeaie a?aa?iaioieo nnaaoe?a a aeine?aeao in
vitro aoee eaaeaie aeaeo aaeeeee i?aeeiaii?n (Rhinolophus
ferrum-equinum), a aiiieioa?iieo –uo?e-naiee e?i?? A?noa? (Rattus
norveg?nus).

A aenia?eiaioao in vivo c iiee?eioa?iieie oaa?eiaie aaiaoc(e iiae?eyee
ia aea? a?oie oa ai?uoaaee a aeaa??oie c oaiia?aoo?ith aiaee 14 oa 24
0N. I?ney 35-aeiaiai? aee?iaoe?? a aeaa??oie aiineee 14N-i?ie?i. Ia 45-o
aeiao ?ea oia?oaeyee ? a aenia?eiaio a?aee oe??o, o?aaao ? oainoiaee
ieaaaoeue. O aeine?aeao in vitro aeine?aeaeoaaee oe??o oa ieaaaeueiee
i?oo? ea?ia. Eiai aiiiaaiao ai?uoaaee o ?ic/ei ??iaa?a-E?aana, ui i?noea
3I-i?ie?i. ? ?ieoaoaaee i?e oaiia?aoo?ao 15 oa 25 0N [Eaaaaeaa, 1978,
Martens, 1989].

Aeine?aee c nnaaoeyie i?iaiaeeee in vitro aeey ciyooy oa?aeoa?ii? aeey
ieo oaiia?aoo?ii? ?aaoeyoe??. O uo??a aea/aee oe??o nieie oa na?oea, a o
eaaeai?a — oe??yio ia?aoeieo e?eea. Ia(?eoii aeine?aeaeaiiy o ??cieo
aenia?eiaioao aoa eieaaai oeio ? oa noia?iee eieaaai i?aai?a.
?ieoaoaaiiy aiiiaaiao?a i?iaiaeeee oae naia. ye ? a aenia?eiaioao c
?eaaie in vitro, i?e oaiia?aoo?ao ?ieoaaoe?? 10, 15, 20, 25, 30,
35 oa 40 0N o uo??a ? 5, 10, 15, 20, 25, 30, 35 0N o eaaeai?a.

I?ney ?ieoaaoe?? eieaaai aeeo/aee c i?aai?a [Neooeeee, 1969, Ieeeoeia,
1992], a?ae/euaee oeaio?eooaoaaiiyi oa ae?ae?cii ? aeei?enoiaoaaee o
iiaeaeueoeo aeine?aeaeaiiyo.

Ai?no eieaaaio ioe?ithaaee ca ienei?ie?iii [Stegemann, Stalder, 1967].

I?i ?ioaineai?noue neioaco eieaaaio noaeeee ia i?aenoaa? ieoiii?
?aae?iaeoeaiino? a iueiio 14N-ienei?ie?io (a aeine?aeao in vivo) oa
3I-ienei?ie?io (a aeine?aeao in vitro) [Caia?aaaa, 1977, Eaaaaeaa, 1978,
Martens, 1989].

Nooi?iue a?ae?ieneethaaiiy i?ie?io a eieaaai? aecia/aee ca [Berg, 1982].
?iciiae?e ?aae?iaeoeaiiai i?ie?io oa ienei?ie?io i?iaiaeeee ca
Caia?a?aith [Caia?aaaa, 1977].

??aaiue caaaeueiiai ae?eiceethaaiiy eieaaaio aecia/aee ca
n??/aii-oaiieueiei iaoiaeii. Aeey aecia/aiiy oa?iaioaoeaiiai
ae?eiceethaaiiy aeei?enoiaoaaee iaoiae eiieo?aioiiai ?ia?a??oaaiiy
o?ainoa?ac [Dubous, 1986, Smith, 1989].

?ioaineai?noue aooieieaaaiie?co aecia/aee ia i?aenoaa? a?aeiioaiiy
a?eueiiai ienei?ie?io, ui aeeoia c? c?ace?a o ?ieoaaoe?eia na?aaeiaeua a
i?ioean? ?ieoaaoe??, aei caaaeueii? e?eueeino? ienei?ie?io, yeee
ciaoiaeeany o c?aceao aei ii/aoeo ?ieoaaoe?? [Ia?nueeee, 1997].

Iiia?a/ia ca(ycoaaiiy eieaaaio ioe?ithaaee a aeine?aeao in vitro ca
eiai ?ic/eii?noth a aeno?aeoe?eiiio ?ic/ei? [Ooaaneay, Ia?neee, 1982]. A
aeine?aeao c ?eaaie in vivo eiai ioe?ithaaee oaeiae ca a?aeiioaiiyi
aeaiaoo?iaaiiai eieaaaio aei caaaeueii? e?eueeino? eieaaaio o c?aceao
aei ?o iaa??ao [Ieeeoeia, 1992].

Oa?iinoaa?euei?noue eieaaaio, neioaciaaiiai a aeine?aeao in vitro i?e
??cieo oaiia-?aoo?ao, ioe?ithaaee ca oa?iiaeeiai?/ieie ia?aiao?aie eiai
aeaiaoo?aoe?? [Ia?neee, Aei-iaeaae/, 1990, Eociaoeia, Ia?neee,
Nieiaueaaa, 1998].

?iaioa aeeiiaia c aeei?enoaiiyi niaoe?aeueiiai i?ia?aiiiai
caaacia/aiiy: Microsoft Word for Windows 6.0, Borland Quadro Pro 5.0,
CorelDRAW 5.0, Global DeLorme Explorer.

?icae?e 3. ?acoeueoaoe aeine?aeaeaiue oa ?o iaaiai?aiiy.

Aenia?eiaioe ia ?eaao in vivo .

Ia ?en. 1 A iaaaaeaii caeaaei?noue ?ioaineaiino? neioaco eieaaaio oe??e,
oainoiaiai ieaaoey oa o?aaoa a?ae oaiia?aoo?e iaaeieeoiueiai
na?aaeiaeua. Aeaeii, ui iaea?eueoa ?ioaineai?noue neioaco eieaaaio
niinoa??aa?oueny i?e oaiia?aoo?? 24 0N. I?e oe?e aea oaiia?aoo?? aeuee
? nooi?iue a?ae?ieneethaaiiy caeeoe?a i?ie?io a eieaaai? oe??e (?en. 1
A). Oa?aeoa?ii, ui oeae ia?aiao? eaaeeoue o iaaeao 27-38 %, ui
a?aeiia?aea? na?aaei?i cia/aiiyi nooiaiy a?ae?ieneethaaiiy caeeoe?a
i?ie?io a eieaaai? ?ea na?aaei?o oe?io.

?en. 1. Oaiia?aoo?ia caeaaei?noue ?ioaineaiino? neioaco eieaaaio oe??e
(1A), o?aaoo (2A), oainoiaiai ieaaoey (3A) aaiaoc?e; ?ioaineaiino?
a?ae?ieneethaaiiy eieaaaio oe??e(1A), o?aaoo (2A) oa oainoiaiai ieaaoey
(3A) aaiaoc?e.

Eiioeaio?aoe?y ienei?ie?io a oe??? ?ea oaeiae aeua i?e oaiia?aoo?? aiaee
24 0N (?en. 2 A). I?e oe?e oaiia?aoo?? a?aeaoaa?oueny ?
?ioaineo?eaoe?y i?ioean?a iiia?a/iiai ca(ycoaai-

?en. 2. Oaiia?aoo?ia caeaaei?noue eiioeaio?aoe?? eieaaaio a oe???
aaiaoc?e (A); ?ic/eiiino? eieaaaio oe??e aaiaoc?e a 1I NaCl (A); aeoiaeo
eieaaaio c? oe??e aaiaoc?e i?e ?? iaa??a? o aiae? (A).

iy eieaaaio a oe???. I?i oea na?ae/eoue cieaeaiiy aeoiaeo c? oe??e ?ea
aeaiaoo?iaaiiai eieaaaio i?e iaa??aaii? oeaieie o aiae? i?e 60 0N, a
oaeiae iaeo?aeueii?ic/eiiiai eieaaaio i?e eiai aeno?aeoe?? 1M NaCl (?en.
2 A oa 2 A).

Iaaaaeai? aeai? na?ae/aoue i?i oa, ui i?aeaeuaiiy oaiia?aoo?e
iaaeieeoiueiai na?aaeiaeua aeeeeea? o ?ea no?oeoo?io ia?aaoaeiao
eieaaaiiaeo ooai?aiue. Aiia iieyaa? o c?inoaii? noaa?eueiino? eieaaaio
ye ia iieaeoey?iiio, oae ? ia iaaeiieaeoey?iiio ??aiyo ?
a?aecia/a?oueny ocaiaeaeaieie ci?iaie ?ioaineaiino? iine?aeiaieo noaae?e
eiai iai?io.

Aenia?eiaioe ia ?eaao in vitro.

O oaae.1 iaaaaeai? aeai? i?i ?ioaineai?noue ??cieo noaae?e iai?io
eieaaaio ieaaaeueiiai i?oo?a ?ea o aeine?aeao in vitro. Iaea?eueoa
?ioaineai?noue neioaco niinoa??aa?oueny i?e 25 0N. Nooi?iue
a?ae?ieneethaaiiy i?ie?io o eieaaai? i?e oe?e oaiia?aoo?? oaeiae aeuee.
Oe? aeai? na?ae/aoue i?i c?inoaiiy no?oeoo?ii? noaa?eueiino?
?iaeea?aeoaeueieo iieaeoe eieaaaio, neioaciaaieo i?e aeu?e oaiia?aoo??
iaaeieeoiueiai na?aaeiaeua. ??aaiue ?iciaaeo eieaaaio i?e oe?e
oaiia?aoo?? iaeiaioee. O ii?aeiaii? c ?ioaineo?eaoe??th neioaco i?e 25
0N oea aaaea aei i?aeaeuaiiy eiioeaio?aoe?? eieaaaio a oeaiei? i?e oe?e
oaiia?aoo?? ?ieoaaoe??. Nooi?iue iiia?a/iiai ca(ycoaaiiy a eieaaai?
oaeiae iaea?eueoee i?e 25 0N, i?i ui na?ae/eoue cieaeaiiy aeoiaeo
?ic/eiiiai eieaaaio a i?ioean? aeno?aeoe?? 1I NaCl. C?inoaiiy
eiioeaio?aoe?? eieaaaio a oeaiei?, a oaeiae i?aeaeuaiiy nooiaiy eiai
iiia?a/iiai cay(coaaiiy i?e aeu?e oaiia?aoo?? na?aaeiaeua aaaea aei
ca?eueoaiiy no?oeoo?ii? noaa?eueiino? ? iaaeiieaeoey?ieo eieaaaiiaeo
ooai?aiue.

Oaaeeoey 1

Aieea oaiia?aoo?e ?ieoaaoe?eiiai na?aaeiaeua ia ??ci? noaae?? iai?io
eieaaaio a ieaaaeueiiio i?oo?? ea?ia cae/aeiiai.

Oaiia?a-

oo?a ?ieoaaoe??,

O, 0C

Ieoiia ?aae?iaeoeai?noue

3I-Ii?i,

?ii/ oa/iea Ii?i Nooi?iue a?ae?i-

eneethaaiiy i?ie?io,

3I-Ii?i_____

3I-Ii?i+3I-I?i Nooi?iue

ae?eice-ethaaiiy ei-eaaaio, %

??aaiue

aooieiea-aaiie?co, % ?ic/ei-i?noue eieaaaio a 1I NaCl, %

15 195 + 9,8 0,37 + 0,026 3,2 + 0,25 2,9 + 0,12 2,3 + 0,08

25 810 + 37 0,46 + 0,037 5,7 + 0,38 1,3+0,09 1,4+ 0.04

Oaeei /eiii, ie aa/eii, ui a aeine?aeao in vitro i?aeaeuaiiy oaiia?aoo?e
iaaeieeoiueiai na?aaeiaeua aeeeeea? oaeaeeo ia?aaoaeiao iai?io eieaaaio,
yea aaaea aei c?inoaiiy no?oeoo?ii? noaa?eueiino? ia on?o ??aiyo eiai
i?aai?caoe??. Iaaii, ui oaea ia?aaoaeiaa ? iaeiei c iaoai?ci?a oaeaeei?
aaeaioaoe?? iiee?eioa?iieo i?aai?ci?a aei ??ceeo ci?i oaiia?aoo?e
iaaeieeoiueiai na?aaeiaeua.

Aenia?eiaioe ia a?aa?iaioieo oa aiiieioa?iieo oaa?eiao in vitro.

Ia ?enoieao 3-7 oa a oaae. 2 ? 3 iaaaaeai? aeai? i?i aieea oaiia?aoo?e
ia ??ci? noaae?? iai?io eieaaaio aeayeeo i?aai?a nnaaoe?a o aeine?aeao
in vitro. Ine?eueee eieaaai oeio ? neeaaea? iniiaio iano (a?eueo 90%)
eieaaaio oe??e noaoaaic??eeo oaa?ei, eiai aeanoeaino? o cia/i?e i???
aecia/athoue oa?iinoaa?euei?noue iaaeiieaeoey?ieo eieaaaiiaeo
ooai?aiue. C ?en. 3 oa

?en. 3. Oaiia?aoo?ia caeaaei?noue ?ioaineaiino? neioaco eieaaaio a oe???
uo??a (1) oa a

oe??yi?e ia?aoeioe? e?eea eaaeai?a (2).

aeaieo oaae. 2 aeaeii, ui caeaaeiino? ieoiii? ?aae?iaeoeaiino?
ienei?ie?io a eieaaai? oe??e oaa?ei iaio aeae?a ? a eieaaai? na?oey
uo??a a?ae oaiia?aoo?e iathoue aeaa iaeneioie.

A an?o aeiaaeeao ia?oee iaeneioi niinoa??aa?oueny i?e 15 0N, ae?oaee aea
– i?e 25 0N aeey oe??e eaaeai?a oa 35 0N – aeey oe??e oa na?oey uo??a.
Caeaaeiino? ?ioaineaiino? a?ae?ieneethaaiiy caeeoe?a i?ie?io aeey
eieaaaio oe??e uo??a oa oe??yii? ia?aoeiee e?eea eaaeai?a a?ae
oaiia?aoo?e iaaaaeai? ia ?en. 4. Aiie oaae iathoue aeaa iaeneioie, i?e
oaiia?aoo?ao 15 oa 35 0N.

Oaiia?aoo?ia caeaaei?noue a?ae?ieneethaaiiy caeeoe?a i?ie?io o
eieaaai? na?oey uo??a ia? eeoa iaeei iaeneioi i?e 35 0N (oaae. 2).

?en. 4. Oaiia?aoo?ia caeaaei?noue ?ioaineaiino? a?ae?ieneethaaiiy
eieaaaio a oe??? uo??a (1) oa a oe??yi?e ia?aoeioe? e?eea eaaeai?a (2).

Oaaeeoey 2

Aieea oaiia?aoo?e ?ieoaaoe?eiiai na?aaeiaeua ia ??ci? noaae?? iai?io
eieaaaio oa ?iciaaeo caaaeueiiai a?eea na?oey 3-i?ny/i?o uo??a-naiie
e?i?? A?noa?.

Oaiia?a-oo?a ?ieoaaoe??, O, 0N Ieoiia ?aae?iaeoeai?noue

3I-Ii?i, ?ii/oa/iea Ii?i Nooi?iue a?ae?ieneethaaiiy i?ie?io,

3I-Ii?i_____

3I-Ii?i+3I-I?i ??aaiue aooiei-eaaaii-e?co, % ??aaiue ?iciaaeo
caaaeueiiai

a?eea, %

10 310+14 0,28+0,012 0,6+0,04 4,5+0,22

15 845+37 0,31+0,015 4,8+0,37 8,6+0,36

20 182+16 0,33+0,014 0,7+0,06 5,7+0,28

25 183+10 0,30+0,025 0,6+0,05 3,5+0,17

30 304+23 0,35+0,018 0,8+0,03 3,3+0,19

35 507+41 0,46+0,011 0,98+0,04 2,8+0,15

40 160+6,5 0,27+0,013 0,7+0,02 3,1+0,17

Ia ?en. 5 iaaaaeai? aeai? i?i caaaeueiee nooi?iue ae?eiceethaaiiy
eieaaai-iie?naoa?eaeieo eiiieaen?a a oe??? uo??a. Oea ae?eiceethaaiiy
aeeth/a? aei naaa ye oa?iaioaoeaia, oae ? iaoa?iaioaoeaia i?e?aeiaiiy
aaenic aei eieaaaio. O a?aeiia?aeiino? c iaaaaeaieie aeaieie
niinoa??aa?oueny eeoa iaeei i?e ye caaaeueiiai, oae ? oa?iaioaoeaiiai
ae?eiceethaaiiy – i?e 35 0N.

?en. 5. Oaiia?aoo?ia caeaaei?noue nooiaiy caaaeueiiai ae?eiceethaaiiy
eieaaaio (1) oa nooiaiy oa?iaioaoeaiiai ae?eiceethaaiiy eieaaaio (2).

Oaiia?aoo?i? caeaaeiino? e?eueeino? ?ic/eiiiai eieaaaio, ui
aeno?aao?oueny c? oe??e oaa?ei iaio aeae?a, iaaaaeai? ia ?en. 6. sse
aa/eii, i?ae ?ic/eii?noth eieaaaio ? oaiia?aoo?ith ?ieoaaoe?? ? cai?ioiy
e?iaeia caeaaei?noue, i?e oeueiio iaeneioi ?ic/eiiino? niinoa??aa?oueny
i?e 5 0N, a i?i?ioi – i?e 40 0N. Oea na?ae/eoue i?i ?ioaineo?eaoe?th
iiia?a/iiai

?en. 6. Oaiia?aoo?ia caeaaei?noue ?ic/eiiino? na?aeineioaciaaiiai
eieaaaio oe??e uo??a, ui aeeoia o ?ic/ei o i?ioean? aeno?aeoe?? 1M NaCl
(1), oa na?aeineioaciaaiiai eieaaaio oe??yii? ia?aoeiee e?eea eaaeai?a,
ui aeeoia o ?ic/ei a i?ioean? aeno?aeoe?? 0,5I NI3NIII (2).

eiaaeaioiiai ca(ycoaaiiy iieaeoe eieaaaio i?e i?aeaeuaii? oaiia?aoo?e
na?aaeiaeua, ui aaaea aei c?inoaiiy noaa?eueiino? eiai iaaeiieaeoey?ieo
ooai?aiue.

Ia ?en. 7 oa a oaae.2 iaaaaeai? aeai? i?i oaiia?aoo?io caeaaei?noue
aooieieaaaiie?co o oe??? oaa?ei iaio aeae?a ? a na?oe? uo??a. On? oe?
aeai? ye?nii iiae?ai?. Iaeneioi ?ioaineaiino? ?iciaaeo eieaaaio o oe???
oa na?oe? uo??a a?aecia/a?oueny i?e 15 0N, a o oe??? eaaeai?a – i?e
10 0N. Oaeei /eiii, ie aa/eii, ui oaiia?aoo?a na?aaeiaeua
aaciina?aaeiuei aieeaa? ia aeoeai?noue oa?iaio?a neioaco oa i?ioean?iao
eieaaaio. Oea aaaea aei ia?aaoaeiae iai?io, ni?yiiaaiiai ia ci?io
no?oeoo?ii? noaa?eueiino? eieaaaiiaeo ooai?aiue ? i?eaaaeaiiy ?o o
a?aeiia?aei?noue c oaiia?aoo?ith na?aaeiaeua. Aeey aeyaeaiiy oeueiai
aoaeoo aoea aei??yia oa?iinoaa?euei?noue eieaaaio, neioaciaaiiai i?e
??cieo oaiia?aoo?ao.

?en. 7. Oaiia?aoo?ia caeaaei?noue aooieieaaaiie?co a oe??? uo??a (1) oa
a oe??yi?e ia?aoeioe? e?eea eaaeai?a (2).

Ia ?en.8 cia?aaeai? e?ea? oaieiiiaeeiaiiy o?a?ee, ooai?aieo c eieaaaio,
neioaciaaiiai i?e 15 oa 35 0N, a a oaae. 3 iaaaaeai? ?ac?aoiaai? ca
oeeie e?eaeie oa?iiaeeiai?/i? ia?aiao?e eiai aeaiaoo?aoe??. ?acoeueoaoe
?ic?aooie?a na?ae/aoue, ui oa?iinoaa?euei?noue eieaaaio, neioaciaaiiai
i?e 35 0N, cia/ii aeua i?ae oa?iinoaa?euei?noue eieaaaio, neioaciaaiiai
i?e 15 0N. Oea aeaco? ia ca(ycie i?ae oaiia?aoo?ieie caeaaeiinoyie
aea/aieo aoai?a iai?io eieaaaio ? no?oeoo?ii? noaa?eueiino? oeueiai
a?eea, neioaciaaiiai i?e ??cieo oaiia?aoo?ao.

?en. 8. Oaiia?aoo?ia caeaaei?noue oaieiiiaeeiaiiy o?a?ee, ooai?aieo c
eieaaaio, ui aoa neioaciaaiee i?e 15 0N (1) oa i?e 35 0N (2).

Oaaeeoey 3

Nooi?iue a?ae?ieneethaaiiy eieaaaio, neioaciaaiiai i?e ??cieo
oaiia?aoo?ao,

oa oa?iiaeeiai?/i? ia?aiao?e ieaaeaiiy ooai?aieo c iueiai o?a?ee.

Oaiia?a- oo?a ?ieoaaoe??, O, 0N Nooi?iue a?ae?ieneethaai-iy eieaaaio, %
Oaiia?aoo?a aeaiaoo?aoe??, E Aioaeuei?y aeaiaoo?aoe??, (eAeae/iieue) o
102 Aio?ii?y aeaiaoo?aoe??,

Aeae/iieue

15 41,5 320,5 0,89 + 0,12 0,28 + 0,02

35 52,3 321,5 1,45 + 0,23 0,45 + 0,02

Aiae?c oaiia?aoo?ieo caeaaeiinoae aeine?aeaeaieo noaae?e iai?io
aeicaiey? aeyaeoe iaoai?cie, ui aaaeooue aei i?aeaeuaiiy
oa?iinoaa?eueiino? eieaaaio, neioaciaaiiai i?e 35 0N. ?ioaineai?noue
neioaco eieaaaio i?e oe?e oaiia?aoo?? o 4 ?ace aeua, i?ae o ?ioa?aae?
oaiia?aoo? 20-30 0N, a ?ioaineai?noue ?iciaaeo iaeiaioa. Aeuee i?e oe?e
oaiia?aoo?? ? nooi?iue a?ae?ieneethaaiiy caeeoe?a i?ie?io o eieaaai?, a
ye iane?aeie iaea?eueo aeniea no?oeoo?ia noaa?euei?noue
?iaeea?aeoaeueieo iieaeoe. I?e 35 0N a?eueo aenieee ? ??aaiue
iinoneioaoe/ieo iiaeeo?eaoe?e eieaaaio. Oaeei /eiii, i?e oaiia?aoo??, ui
aeecueea aei oaiia?aoo?e o?ea aiiieioa?iieo nnaaoe?a, aeyaey?oueny
ocaiaeaeaiiy ?ioaineo?eaoe?? ? oiia?eueiaiiy noaae?e iai?io eieaaaio,
ni?yiiaaieo ia ooai?aiiy aaeaeaaoii? oe?e oaiia?aoo?? no?oeoo?ii?
noaa?eueiino? eieaaaio.

I?e cieaeaii? oaiia?aoo?e aeyaey?oueny, ui i?e 15 0N iaeii/anii ? ??cei
c?inoathoue neioac, ?iciaae oa ?ic/eii?noue eieaaaio, o oie /an, ye
nooi?iue eiai a?ae?ieneethaaiiy cieaeo?oueny. Aeey iiyniaiiy aeyaeaiiai
aoaeoo ne?ae ?icaeyiooe iniaeeaino? aenia?eiaioo. O aeeo/aieo c?aceao,
ui ciaoiaeyoueny o ?ieoaaoe?eiiio ?ic/ei?, o?c?ieia?/ia oaiia?aoo?ia
?aaoeyoe?y a?aenooiy. I?e iecuee?e oaiia?aoo?? ?ieoaaoe?? no?oeoo?ia
noaa?euei?noue eieaaaiiaeo ooai?aiue o ieo aeua, i?ae iaiao?aeia. Oiio o
c?aceao iiaeiia a?aeaooeny ia?aaoaeiaa oeeo ooai?aiue, ni?yiiaaia ia
i?eaaaeaiiy o a?aeiia?aei?noue ?o no?oeoo?ii? noaa?eueiino? ciai?oi?e
oaiia?aoo??. sse aa/eii c io?eiaieo aeaieo, iaeaea oea ? a?aeaoaa?oueny
o c?aceao oeaieie. ??cea ?ioaineo?eaoe?y ?iciaaeo aaaea aei aeeo/aiiy
iieaeoe eieaaaio, no?oeoo?ia noaa?euei?noue yeeo ia a?aeiia?aea?
oaiia?aoo??. Ia ?o i?noea i?eoiaeyoue iia?, na?aeineioaciaai? oa iaio
noaa?euei? iieaeoee, ye? iathoue ieae/ee ??aaiue a?ae?ieneethaaiiy,
ae?eiceethaaiiy oa iiia?a/iiai ca(coaaiiy. Oaeei /eiii, o c?aceao oe??e
a?aeaoaa?oueny ci?ia iai?io eieaaaio, ui aaaea aei aaeaioaoe?? oeaieie
aei a?eueo iecueei? oaiia?aoo?e na?aaeiaeua, yea ia ? oa?aeoa?ia aeey
aea/a?iiai aeaeo nnaaoe?a.

Oeae i?ioean a?aeaoaa?oueny eeoa oiio, ui o eieaaai-neioacoth/eo
ee?oeiao nnaaoe?a ? oa?iaioao?ai? nenoaie, iioeioi ae?? yeeo
ciaoiaeeoueny i?e iecuee?e oaiia?aoo??. Oa?aeoa?ii, ui i?e cieaeaii?
oaiia?aoo?e ?ieoaaoe?? aaea aei 10 0N, a aeine?aeao in vitro c? uo?aie,
?ioaineai?noue ye neioaco, oae ? ?iciaaeo ??cei cieaeo?oueny. I?e ye?nii
iiae?aiiio oa?aeoa?? oaiia?aoo?ieo caeaaeiinoae aea/aieo noaae?e iai?io
eieaaaio uo??a oa eaaeai?a i?ae ieie a?aei?/a?oueny ? aeayea ??cieoey.
Ia?o ca ana i?eaa?oa? aei naaa oaaao oa, ui ye aaniethoi? aaee/eie, oae
? iaeneioie e?eaeo, ye? a?aeia?aaeothoue ?ioaineai?noue neioaco oa
?iciaaeo eieaaaio eaaeai?a, ia ca?aathoueny c oaeeie o uo??a. Oae, a
iaeano? aenieeo oaiia?aoo? iaeneioi ?ioaineaiino? neioaco eieaaaio o
eaaeai?a a?aei?/a?oueny ia i?e 35, a i?e 25 0N (?en. 3). Iaeneioi
?iciaaeo eieaaaio o eaaeai?a oaeiae cnoiooi o iaeanoue a?eueo iecueeeo
oaiia?aoo? (?en. 7).

Oey ?icieoey iia(ycaia, iiaeeeai, c ?icieie oiiaaie inioaaiiy oeeo
aeaeia oa c ?icieie oeiaie iaoaiiciia ?o oa?ii?aaoeyoei?. Oaeei /eiii,
aeyaeaii caeaaeii aiae oaiia?aoo?e ia?aaoaeiae eieaaaiiaeo ooai?aiue o
eaaeaiia iiaeooue aooe i?enoinoaaiiyi aei aeaio ?icieo noaiia –
aeoeaiiai oa ianeaiiai.

Iaea?aeaii aeaii iaaeathoue iiaeeeainoue c?iaeoe aeniiaie, ui o i?ioeani
aaiethoei? o?aaaoieo oaa?ei ae?iaeany ?aeeiee oiiaa?naeueiee iaoaiici
oaeaeei? no?oeoo?ii-iaoaaiei/ii? aaeaioaoei? eieaaaiiaiai iao?eeno aei
ciii oaiia?aoo?e iaaeieeoiueiai na?aaeiaeua. A i?ioeani iiaeaeueoi?
aaiethoei? nnaaoeia oeae iaoaiici i?iaeiaaeo? aeei?enoiaoaaoeny oieueee
aiaa?iaioieie oaa?eiaie. O aiiieioa?iieo aii caeeoeany, iiaeeeai,
oieueee o aeaeyaei ?aeieoa.

AENIIAEE

O iiee?eioa?iieo o?aaaoieo – aaiaoc?? oa ea?ia – ?ioaineai?noue neioaco
caaaeueiiai eieaaaio a oe???, ieaaaeueiiio i?oo??, oainoiaiio ieaaoe? ?
o?aao? a iaeano? 24-25 0N aeua, i?ae a iaeano? 14-15 0N. O a?aa?iaioieo
o?aaaoieo – eaaeai?a – oaiia?aoo?ia caeaaei?noue ?ioaineaiino? neioaco
eieaaaio o oe??yi?e ia?aoeioe? e?eea ia? aeaa iaeneioia – a iaeano?
15 0 oa 25 0N. A aenia?eiaioao in vitro a oe??? oa na?oeaaiio i’yc?
aiiieioa?iieo nnaaoe?a – uo??a – oey caeaaei?noue oaeiae ia? aeaa
iaeneioie – a iaeanoyo 15 0 oa 35 0N.

Nooi?iue a?ae?ieneethaaiiy caeeoe?a i?ie?io a iieaeoeao eieaaaio oaeiae
ei?aeth? c oaiia?aoo?ith na?aaeiaeua. Iaeneioi niinoa??aa?oueny a
iaeano? 24-25 0N o ?ea oa i?e 35 0N o nnaaoe?a.

??aaiue ae?eiceethaaiiy eieaaai-iie?naoa?eaeieo eiiieaen?a ?
iaeneiaeueiei a ieaaaeueiiio i?oo?? oa oe??? ea?ia i?e 25 0N, a o oe???
uo??a i?e 35 0N.

C? c?inoaiiyi oaiia?aoo?e na?aaeiaeua o iiee?eioa?iieo oa a?aa?iaioieo
o?aaaoieo oa oaiia?aoo?e ?ieoaaoe?? a aenia?eiaioao in vitro o
aiiieioa?iieo nooi?iue iiia?a/iiai ca’ycoaaiiy eieaaaio a on?o aea/aieo
i?aaiao c?inoa?.

O aea/aieo i?aaiao on?o aeine?aeaeaieo aeae?a o?aaaoieo ??aaiue
aooieieaaaiie?co iia’ycaiee c oaiia?aoo?ith iai?ioiueii?iii?oe?eiith
caeaaei?noth ? ? iaeneiaeueiei i?e oaiia?aoo?? 10-15 0N.

Eieaaai, neioaciaaiee in vitro i?e 35 0N, a?eueo oaiinoaa?eueiee — ia?
aeuo oaiia?aoo?o oa aioaeuei?th aeaiaoo?aoe?? — i?ae neioaciaaiee i?e 15
0N.

Anoaiiaeaii, ui ci?ia oaiia?aoo?e na?aaeiaeua iiaea aaciina?aaeiuei
?aaoethaaoe ?ioaineai?noue iai?io eieaaaio, aieeaath/e ia eeth/ia?
noaae?? eiai neioaco, i?ioean?iao oa ?iciaaeo. ?niothoue aeaa oeie oaei?
?aaoeyoe??. I?e ia?oiio ?ioaineai?noue eiie?aoii? noaae?? iai?io
aecia/a?oueny oaiia?aoo?iith caeaaei?noth aeoeaiino? oiai ae naiiai
oa?iaioa (ye o aeiaaeeo ae?eiceethaaiiy). I?e ae?oaiio – oaiia?aoo?a
aeoea?co? iaeio c ?cioi?i oa?iaioa, iaea?eueo aeoeaio a aecia/aiiio
oaiia?aoo?iiio ?ioa?aae?, ye oea iiaeeeai ia? i?noea i?e
a?ae?ieneethaaii? i?ie?io.

Ai?no ienei?ie?io a iieaeoeao eieaaaio ia ? aeaeiaith iciaeith
i?aai?cio. Eeoa iaeneiaeueiee nooi?iue a?ae?ieneethaaiiy i?ie?io, yeee
caeaaeeoue a?ae ia?aeiii? no?oeoo?e iieaeoee, iiaea aeei?enoiaoaaoeny ye
nenoaiaoe/iee iieaciee, aea ia a?eueo aeuiio oaeniiii?/iiio ??ai?, i?ae
aeae.

Ia?aaoaeiaa eieaaaiiaeo ooai?aiue, yea aeeeeeaia ci?iaie oaiia?aoo?e
na?aaeiaeua, ? iaeiei c iaoai?ci?a oaeaeei? aaeaioaoe?? niieo/ii?
oeaieie aei oeeo ci?i. Oe? iaoai?cie aeei?enoiaothoueny iiee?eioa?iieie
oa a?aa?iaioieie o?aaaoieie, ye? ?niothoue a oiiaao ??ceeo naciiieo ci?i
oaiia?aoo?e aeiae?eey. O aiiieioa?iieo nnaaoe?a oe? iaoai?cie
caeeoeeenue eeoa o aeaeyae? ?ae?eoo.

Nienie iioae?eiaaieo i?aoeue ca oaiith aeena?oaoe??:

Ia?neee A.Y., Eociaoeia E.A., Aa?aocaiei I.A. Oaiia?aoo?iay caaeneiinoue
iaeioi?uo noaaeee iaiaia eieeaaaia a eiaea e?un in vitro // AeAI
Oe?aeiu. – 1997. — ? 1. – N. 170-173.

Garbuzenko O.B., Kuznetsov I.V., Naglov A.V., Nikitina N.A., Persky
E.E., Utevskaya L.A. Structural and metabolic mechanisms of collagen
adaptation to endogenous and exogenous factors effects // School of
fundamental medicine journal. – 1997. – V.3, ? 1 – N. 27-30.

Eociaoeia E.A., Aa?aocaiei I.A., Eocueien A.Y., Ieeeoeia I.A., Ia?neee
A.Y., Ooaaneay E.A. Iniaaiiinoe iaiaia eieeaaaia eiaee iicaiii/iuo
aeeaioiuo i?e oaiia?aoo?iie aaeaioaoeee // I?iaeaiu e?eiaeieiaee. –
1997. — ? 3. – N. 27-31.

Eociaoeia E.A., Nieiaueaaa A.N., Ia?neee A.Y., Cei/aiei A.A. Auno?ay
no?oeoo?ii-iaoaaiee/aneay aaeaioaoeey eieeaaaia e oaiia?aoo?a //
I?iaeaiu e?eiaeieiaee. – 1998. — ? 1. – N. 67-68.

Ia?neee A.Y., Eociaoeia E.A., Iaaeia A.A., Ieeeoeia I.A., Ieia I.A.,
Ooaaneay E.A. Iniaaiiinoe iaoaieciia, iaania/eaathueo iaaeaaeiinoue
eieeaaaiiauo no?oeoo? a iioiaaiaca // Neiiiceoi “Aeieiae/aneea iaoaieciu
noa?aiey”. — Oac. aeiee., Oa?ueeia. – 1994. – N. 62.

Ia?nueeee ?.A., Aa?aocaiei I.A., Eoci?oeia ?.A., Iaaeia I.A., I?e?o?ia
I.A., Ooaanueea E.A. ?ieue ie?aieo aoai?a neioaco a “oaeaee?e”
aaeaioaoe?? eieaaaiiaeo no?oeoo? o?aaaoieo aei ??cieo oaiia?aoo? // O?V
c(?cae Oe?a(inueeiai o?c?ie. oia. — Oac. aeii., Ee?a. – 1994. — N.59.

Aa?aocaiei I.A., Eoci?oeia ?.A., Iaaeia I.A., I?e?o?ia I.A. A?io?i?/i?
iaoai?cie ?acenoaioiino? eieaaaiiaeo no?oeoo? niieo/ii? oeaieie aei ae??
iioeiaeaeoth/eo oaeoi??a // ?? eiia?an iaoio?c?ieia?a Oe?a?ie. — Oac.
aeii., Ee?a. – 1996. – N.46.

Eoci?oeia ?.A., Aaneeue?aa E.A., Eocuei?n A.A., I?e?o?ia I.A., Ia?nueeee
?.A. No?oeoo?ii-iaoaaie?/i? iaoai?cie aaeaioaoe?? niieo/ii? oeaieie
nnaaoe?a aei oaiia?aoo?e // V?? Oe?a?inueeee a?io?i?/iee c(?cae. — Oac.
aeii., Ee?a. – 1997. –O.??. – N. 148.

Eoci?oeia ?.A., Iaaeia I.A., Ooaanueea E.A. Iaoai?cie oaeaeei?
no?oeoo?ii-iaoaaie?/ii? aaeaioaoe?? eieaaaio oe??e aei oaiia?aoo?e //
Iao. OV c(?caeo Oe?a?inueeiai o?c?ie. oia. — O?c?ieia?/iee aeo?iae. —
1998. — O.44, ? 3. — N.295.

Ieeeoeia I.A., Eociaoeia E.A., Iaaeia A.A., Ooaaneay E.A. Aic?anoiua
iniaaiiinoe auno?ie aaeaioaoeee eieeaaaia ?ua e oaiia?aoo?a // ???
Iaaeaeoia?iaeiue neiiiceoi (Aeieiae/aneea iaoaieciu noa?aiey(. — Oac.
aeiee., Oa?ueeia — 1998. — N.26.

Eoci?oeia ?.A., Aaneeue?aa E.A., Eocuei(n A.A., Ooaanueea E.A.,
Ia?nueeee ?.A. Iieaeoey?iee iaoai??ci oaeaeei? ci?ie oa?iinoaa?eueiino?
eieaaaio i?ae ae??th ciai?oiuei? oaiia?aoo?e // ?? c(?cae Oe?a?inueeiai
a?io?c. oia. — Oac. aeii, Oa?e?a. — 1998. — N.48.

Eoci?oeia ?.A. Iieaeoey?i? iaoai?cie aaeaioaoe?? eieaaaiiaeo no?oeoo?
aei ci?i oaiia?aoo?e. — ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe
c niaoe?aeuei?noth 03.00.04 — a?io?i?y — Oa?e?anueeee aea?aeaaiee
oi?aa?neoao, Oa?e?a, 1998.

I?iaaaeaii ii??aiyeueia aeine?aeaeaiiy iniiaieo aoai?a iai?io eieaaaio,
ui caaacia/othoue eiai noaa?euei?noue ia iieaeoey?iiio oa
iaaeiieaeoey?iiio ??aiyo o i?ioean? oaiia?aoo?ii? aaeaioaoe?? o
iiee?eioa?iieo, a?aa?iaioieo oa aiiieioa?iieo oaa?ei. Iieacaii, ui a
on?o aeiaaeeao iniiai? aoaie iai?io eieaaaio ci?iththoueny iaeiaeiai, ui
caaacia/o? iiceoeaio ei?aeyoe?th i?ae oa?iinoaa?euei?noth eieaaaio oa
oaiia?aoo?ith iaaeieeoiueiai na?aaeiaeua. Oaea ia?aaoaeiaa iai?io
iia(ycaia c? ci?iith neioaco, i?ioean?iao oa ?iciaaeo eieaaaio. A
aenia?eiaioao in vitro i?iaeaiiino?iaaii oaeiae, ui i?e cieaeaii?
oaiia?aoo?e ?ieoaaoe?eiiai na?aaeiaeua ?ioaineo?eo?oueny ?iciaae a?eueo
oa?iinoaa?eueiiai oa neioac iaio oa?iinoaa?eueiiai eieaaaio.

Iieacaii, ui oae? iaoai?cie oaeaeei? aaeaioaoe?? eieaaaiiaeo ooai?aiue
aei oaiia?aoo?e na?aaeiaeua iiaeooue aaciina?aaeiuei aeei?enoiaoaaoeny
iiee?eioa?iieie o?aaaoieie, ui ?niothoue a oiiaao ??ceeo naciiieo ci?i
oaiia?aoo?e. Noiae? iaoai?cie aeei?enoiaothoueny ? a?aa?iaioieie
nnaaoeyie, oaiia?aoo?a o?ea yeeo oaeiae iiaea oaeaeei ci?ithaaoeny
i?ioyaii ei?ioeiai /ano i?e ia?aoiae? a?ae noaio aaaeuei?ino? aei noaio
niey/ee.

C?iaeaii aeniiaie, ui o aiiieioa?iieo nnaaoe?a o oiae? aaiethoe?? oaeiae
caeeoeany ?ae?eoiaee iaoai?ci oaiia?aoo?ii? aaeaioaoe?? eieaaaiiaiai
ea?eano niieo/ii? oeaieie.

Eeth/ia? neiaa: aaeaioaoe?y, eieaaai, oaiia?aoo?a, iai?i,
aooieieaaaiie?c, oa?iiaeeiai?/i? ia?aiao?e, aeaiaoo?aoe?y, no?oeoo?ia
noaa?euei?noue.

Eociaoeia E.A. Iieaeoey?iua iaoaieciu aaeaioaoeee eieeaaaiiauo no?oeoo?
e eciaiaieyi oaiia?aoo?u. — ?oeiienue.

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie eaiaeeaeaoa aeieiae/aneeo iaoe
ii niaoeeaeueiinoe 03.00.04. — aeioeiey. -Oa?ueeianeee ainoaea?noaaiiue
oieaa?neoao, Oa?ueeia, 1998.

I?iaaaeaii n?aaieoaeueiia enneaaeiaaiea iniiaiuo yoaiia iaiaia
eieeaaaia, iaania/eaathueo aai noaaeeueiinoue ia iieaeoey?iii e ia
iaaeiieaeoey?iii o?iaiyo a i?ioeanna oaiia?aoo?iie aaeaioaoeee
iieeeeioa?iiuo, aeaa?iaioiuo e aiiieioa?iiuo aeeaioiuo. Iieacaii, /oi ai
anao neo/ayo iniiaiua yoaiu iaiaia eciaiythony noiaeiui ia?acii,
iaania/eaay iieiaeeoaeueioth ei??aeyoeeth iaaeaeo oa?iinoaaeeueiinoueth
eieeaaaia e oaiia?aoo?ie ie?oaeathuae n?aaeu. Oaeay ia?ano?ieea iaiaia
naycaia, a iniiaiii, n eciaiaieai neioaca, i?ioeanneiaa e ?aniaaea
eieeaaaia. A yenia?eiaioao in vitro i?iaeaiiino?e?iaaii oaeaea, /oi i?e
nieaeaiee oaiia?aoo?u eieoaaoeee eioaineoeoee?oaony ?aniaae aieaa
oa?iinoaaeeueiiai e neioac iaiaa oa?iinoaaeeueiiai eieeaaaia.

Iieacaii, /oi oaeea iaoaieciu auno?ie aaeaioaoeee eieeaaaiiauo no?oeoo?
e ?aceei eciaiaieyi oaiia?aoo?u n?aaeu iiaoo iaiin?aaenoaaiii
eniieueciaaoueny iieeeeioa?iiuie iicaiii/iuie, iaeoathueie a oneiaeyo
?aceeo naciiiuo eieaaaiee oaiia?aoo?u ie?oaeathuae n?aaeu. Noiaeiua
iaoaieciu eniieuecothony, ii-aeaeeiiio, e aeaa?iaioiuie ieaeiieoathueie,
oaiia?aoo?a oaea eioi?uo oaeaea iiaeao ?acei eciaiyoueny i?e ia?aoiaea
io ninoiyiey aiae?noaiaaiey e ninoiyieth niy/ee.

Naeaeai auaiae, /oi o aiiieioa?iiuo ieaeiieoathueo a oiaea yaiethoeee
oaeaea nio?aieeny ?aeeeoiaue iaoaieci oaiia?aoo?iie aaeaioaoeee
eieeaaaiiaiai ea?eana.

Eeth/aaua neiaa: aaeaioaoeey, eieeaaai, oaiia?aoo?a, iaiai,
aooieieeaaaiieec, o

a?iiaeeiaie/aneea ia?aiao?u, aeaiaoo?aoeey, no?oeoo?iay noaaeeueiinoue.

Kuznetsov I.V. Molecular mechanisms of collagen structures adaptation to
the temperature influence. — Manuscript.

Thesis for degree of candidate of biological sciences on the speciality
03.00.04 — biochemistry. Kharkov State University, Kharkov, 1998.

The comparative investigation of the collagen metabolism main stages
were done. These stages provide the stability on collagen molecular and
supramolecular levels at temperature adaptation of cold-blooded,
hibernating and warm-blooded animals.

It was shown that in these three cases all metabolism stages change in
the same way which provides positive correlation between collagen
thermostability and the environment temperature. Such rebuildings of
metabolism connect mainly with synthesis, processing and degradation
changes.

In experiments in vitro there is the intensification of more
thermostable collagen degradation and synthesis of less thermostable
collagen with the decreasing of incubation temperature.

These mechanisms of collagen structures quick adaptation to the sharp
environment temperature changes can be used by cold-blooded animals,
which live in such conditions. The same mechanisms have hibernating
animals, because their body temperature changes at the transition
awaking to sleeping state.

We have made the conclusion that warm-blooded animals also have these
mechanisms of collagen quick adaptation to the temperature influence and
they exist in relict form.

Key words: adaptation, collagen, temperature, metabolism,
autocollagenolysis, thermodynamic parameters, denaturation, structure
stability.

I?aeienaii aei ae?oeo 20.10.98

Ia’?i 1 ae?.a Iae.-ae?oe. a. – 0,75

Oi?iao iaia?o 60o84 1/16

_____ Iaeeaae 100 i?.___________________________________________________
Cai. 22/142___

AI “E?i?-??ci”, i. Eiinoeoooe??, 6

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