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

AA?AOCAIEI IEUeAA AI?EN?AIA

OAeE 577.12:577.112:577.24

AIEEA IAOAI?*IIAI IAAAIOAAEAIIss IA NEIOAC EIEAAAIO, AEANOEIO OA
A(ssCEI-I?OAEI? AEANOEAINO? NIIEO*II? OEAIEIE

03.00.04 – a?io?i?y

Aaoi?aoa?ao

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa a?ieia?/ieo iaoe

Oa?e?a – 1999

Aeena?oaoe??th ? ?oeiien

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

I?i?noa?noaa 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?,
Eae?iai Iaaei

Aaenaio?eiae/, caa. eaoaae?ith a?io?i?? Oa?e?anueeiai aea?aeaaiiai
oi?aa?neoaoo

eaiaeeaeao a?ieia?/ieo iaoe, noa?oee iaoeiaee ni?a?ia?oiee, Eaiioue?aa
O??aea Nieiiii?aia, caa. eaai?aoi???th ee?i?/ii? a?io?i?? Oa?e?anueeiai
iaoeiai-aeine?aeiiai ?inoeoooo i?oiiaae?? ? o?aaiaoieia?? ?i. I.?.
N?oaiea IIC Oe?a?ie

I?ia?aeia onoaiiaa: ?inoeooo a?io?i?? ?i. I.A. Iaeeaae?ia IAI
Oe?a?ie,

a?aeae?e ?aaoeyoe?? iai?io ?a/iaei, i. Ee?a

.

Caoeno a?aeaoaeaoueny «12» aeiaoiy 1999 p. o 1515 aiaeei? ia
can?aeaii? niaoe?ae?ciaaii? a/aii? ?aaee E 64.051.17 i?e Oa?e?anueeiio
aea?aeaaiiio oi?aa?neoao? I?i?noa?noaa ina?oe Oe?a?ie 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?a-nueeiai aea?aeaaiiai oi?aa?neoaoo: 310077, i. Oa?e?a,
ie. Naiaiaee, 4.

Aaoi?aoa?ao ?ic?neaiee “3” aa?aniy 1999 ?.

A/aiee nae?aoa?

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

eaiaeeaeao a?ieia?/ieo iaoe

noa?oee iaoeiaee ni?a?ia?oiee
Iaaeaeei A.?.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue ?iaioe. Iaoai?/i? aeanoeaino? niieo/ii? oeaieie eaaeaoue
a iniia? iaei??? c ?? aaaeeeaeo ooieoe?e — iii?ii?. A on?o aac aeiyoeo
??ciiaeaeao niieo/ii? oeaieie na?aae iniiaieo eiino?oeoe?eieo
aeaiaio?a, ui aecia/athoue oe? aeanoeaino?, ? eieaaai, a a aeayeeo c ieo
(oe??a, noaeeie oa ?io.) — oaeiae aeanoei [Ieeeoei, Ia?neee, Ooaaneay,
1977; Na?ia, Oaooa?, 1985; Turner, Oliver, 1986; Glagov, 1994].
Eiie?aoi? iaoai?/i? aeanoeaino? eiaeiiai c oeeo ??ciiaeae?a niieo/ii?
oeaieie (i?oei?noue, aei?noe?noue, ?icoyaei?noue) a cia/i?e i???
caeaaeaoue a?ae eiioeaio?aoe?? iaio a?ee?a, oa?aeoa?o ?iciiae?eo ?o
iaaeiieaeoey?ieo ooai?aiue — aieieii oa aoaeiae inoaii?o, ia?o ca ana
e?eueeino? a ieo iiia?a/ieo eiaaeaioieo ca(yce?a [Oi?ioeia, 1980; Paoli,
Morchio, 1983]. A naith /a?ao on? oe? no?oeoo?i? ia?aiao?e
aecia/athoueny oa?aeoa?ii iai?io eieaaaio oa aeanoeio — ?ioaineaiinoyie
?o neioaco ? i?i-oean?iao, yeee a?aeaoaa?oueny ye ia iieaeoey?iiio, oae
? ia iaaeiieaeoey?iiio ??aiyo.

O aaee/aci?e e?eueeino? i?ioean?a aeeoo?ae?yeueiino? ( ??no, caai?iiy
?ai, ia??iaee/i? ci?ie aoaeiae i?aai?a ae?ii/iai ?ai?iaeoeoeaiiai
o?aeoo, ??ci? iaoieia?? niieo/ii? oeaieie) iai?i eieaaaio oa aeanoeio
ci?ith?oueny [Neooeeee, 1969]. No?oeoo?ii-ooieoe?iiaeuei? ia?aaoaeiae
niieo/ii? oeaieie i?aai?a, ui a?aeaoaathoueny i?e oeueiio,
a?aeiia?aeathoue ci?iaiei aeiiaai i?aai?cio aei ieo. Na?aae i?e/ei, ui
aeeeeeathoue oaeo ci?io iai?io, aaaeeeaa ?ieue iaeaaeeoue iaoai?/iiio
iaaaioaaeaiith i?aai?a, yea aaaea aei aeaoi?iaoe?? ee?oei niieo/ii?
oeaieie. O aaeee?e e?eueeino? aeine?aeaeaiue in vivo iieacaii ?nioaaiiy
ca(yceo i?ae iaoai?/iei iai?oaeaiiyi niieo/ii? oeaieie oa
eiioeaio?aoe??th a i?e eieaaaio ? aeanoeainoyie eiai iaaeiieaeoey?ieo
ooai?aiue [Oeaaea?n, Neooeeee, 1975; Ay?, Oeeoia?, 1975]. A inoaii? ?iee
aoei aeyaeaii, ui aeaoi?iaoe?y ?cieueiaaieo o?a?iaeano?a oaeiae aaaea
aei ?iaeoeoe?? neioaco ei-eaaaio. O oeueiio aeiaaeeo iiaeeeaee iaoai?ci
ia?aaea/? neaiaeo aeeth/a? niaoeeo?/i? o?ain-iaia?aii? ?aoeaioi?e —
?ioaa?eie, ye? i?e aeaoi?iaoe?? ee?oeiii? iiaa?oi? ia?aaeathoue neaiae
i?i oea ca aeiiiiiaith ?yaeo aioo??oiueiee?oeiieo iina?aaeiee?a ia
i?iiioi?e a?aeiia?aeieo no?oeoo?ieo aai?a [Hynes,1992; Meredith et. al.,
1995; Sundberg,1996;].

Iaeiae aei oeueiai /ano ia ?nio? ?acoeueoao?a aeine?aeaeaiue aieeao
iaoai?/iiai iaaai-oaaeaiiy ia i?ioean?ia eieaaaio. Ia aea/aii aianie
i?aeee?oeiiiai iao?eeno a iaoai?ci ia?aaea/? neaiaeo i?i aeaoi?iaoe?th o
ee?oeio oa eiai iniaeeaino?.

Aein? iaia? ?ioi?iaoe?? i?i aieea iaoai?/iiai iaaaioaaeaiiy ia neioac
aeanoeio. Iaa?aeiiee ae?aiacii iaoai?/ieo iai?oa, a yeiio iiaea
a?aeaoaaoeny ?iaeoeoe?y neioaco eieaaaio oa aeanoeio, a oaeiae eiai
a?eia? iniaeeaino?.

Ia aea/aeeny iniaeeaino? i?e/eiii-iane?aeeiaeo ca(yce?a o eaioethc?:
iai?i eieaaaio oa aeanoeio — no?oeoo?a ?o iaaeiieaeoey?ieo ooai?aiue —
aeanoeaino? niieo/ii? oeaieie i?e ae?? iaoai?/iiai iai?oaeaiiy.

Iaaeinoaoiuei aea/aia iiaeeea?noue ni?yiiaaii? ae?? a?ieia?/ii aeoeaieo
?a/iaei ??ciiai iioiaeaeaiiy ia oi?ioaaiiy no?oeoo?ii-ooioe?iiaeueieo
aeanoeainoae niieo/ii? oeaieie oeyoii aieeao ia aeaye? aoaie iai?io
eieaaaio.

O ca(yceo c oeei ?acoeueoaoe, io?eiai? i?e aea/aii? aieeao iaoai?/iiai
iaaaioaaeaiiy ia neioac aeanoeio, eieaaaio oa eeth/ia? noaae?? eiai
i?ioean?iao in vivo oa in vitro a ??cieo i?aaiao, iiaeooue aiineoe
iaaiee aeeaae o no/ani? oyaeaiiy i?i ?aaoeyoe?th i?ioean?a ii?oiaaiaco ?
?icaeoeo niieo/ii? oeaieie, a oaeiae iathoue aaaeeeaa cia/aiiy aeey
aa?ii-oieia??, iaae?ao???, oa ee?i?ee caoai?thaaiue niieo/ii? oeaieie.

Ca(ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. ?iaioa aoea
aeeiiaia o IAe* Oa?e?anueeiai oi?aa?neoaoo a iaaeao oai
“No?oeoo?ii-iaoaaie?/i? iaoai?cie aaeaioaoe?? niieo/ii? oeaieie aei
ae?? aeno?aiaeueieo oaeoi??a”, ? aea?ae?a?no?aoe?? 0197U003060 ?
“Aaeaioaoe?ei? ci?ie iaoaaie?cio oa no?oeoo?e iii?ieo a?ee?a niieo/ii?
oeaieie i?ae ae??th aeno?aiaeueieo oaeoi??a”, ? aea?ae?a?no?aoe??
0198U4005758.

Iaoa oa caaaeaiiy aeine?aeaeaiiy. Anoaiiaeoe, ye aieeaa? iaoai?/ia
iaaaioaaeaiiy ia neioac eieaaaio oa aeanoeio, i?ioeaniia eieaaaio,
no?oeoo?o oa a(ycei-i?oaeii aeanoeaino? niieo/ii? oeaieie ??cieo i?aai?a
uo??a e?i?? Ainoa? in vivo oa in vitro. Aea/eoe, ye aieeaa? no?oeoo?a
i?aeee?oeiiiai iao?eeno ia i?ioean ia?aaea/? neaiaeo i?i iaoai?/ia
iaaaioaaeaiiy ia neioac i i?ioeaniia eieaaaio. Aeyaeoe ?ieue aeayeeo
aioo??oiueiee?oeiieo eaiie eaioethao ia?aaea/? neaiaeo i?i iaoai?/ia
iaaaioaaeaiiy a neioac? oa i?ioean?ia? eieaaaio. Aeey oeueiai aoee
iinoaaeaii oaei caaaeaiiy:

Aea/eoe aieea iaoai?/iiai iaaaioaaeaiiy ia neioac eieaaaio oa aeanoeio,
aiae-?ieneethaaiiy oa iiia?a/ia ca(ycoaaiiy eieaaaio o oe???, ai?o?,
nooiaeeeeyo i na?oeaaiio i(yci in vivo oa o oe??? ? ai?o? in vitro.

Anoaiiaeoe iaae? iaoaii/ieo iaaaioaaeaiue, o yeeo aiaeaoaa?oueny
?iaeoeoe?y neioaco eieaaaio oa aeanoeio o oe??? oa ai?o? uo??a in vitro,
oa ?o a?eia? iniaeeaino?.

Aea/eoe aieea iaoai?/iiai iaaaioaaeaiiy ia a(ycei-i?oaeii aeanoeaino?
niieo/ii? oeaieie oe??e oa ai?oe uo??a in vitro.

Aeineiaeeoe aieea naiiea?aaceaeo, oei?i?iiaceio oa iiiia eaeueoe?th ia
neioac oa i?ioeaniia eieaaaio i a(ycei-i?oaeii aeanoeaino? niieo/ii?
oeaieie.

Iaoeiaa iiaecia ?iaioe. Aia?oa iieacaii, ui a i?aaiao, ye? caciathoue
??cieo ca aaee/eiith iaoai?/ieo iaaaioaaeaiue in vivo (oe??a, ai?oa,
na?oeaaee i(yc, nooiaeeeey), ?ioaineai?noue neioaco eieaaaio oa
aeanoeio, ??aaiue a?ae?ieneethaaiiy i?ie?io a eieaaai?, a oaeiae
nooi?iue eiai iiia?a/iiai ca(ycoaaiiy ??ci?. Ii??aiyeueiee aiae?c
io?eiaieo aeaieo iieaco?, ui o eiaeiiio i?aai? ni?aa?aeiioaiiy
?ioaineaiinoae neioaco iaio a?ee?a ? aea/aieo noaae?e i?ioean?iao
eieaaaio aaaea aei oi?ioaaiiy a(ycei-i?oaeieo aeanoeainoae niieo/ii?
oeaieie, iioeiaeueieo a?aeiinii aei aaee/eie oa ?iciiae?eo iaoai?/ieo
iai?oa, ui i?eiaaeathoue ia oeae i?aai. Aeyaeaia iiceoeaia ei?aeyoe?y
i?ae aaee/eiith iaoai?/iiai iaaaioaaeaiiy oa ?ioaineaiinoyie neioaco
aeanoeio, eieaaaio, a?ae?ieneethaaiiy i?ie?io oa iiia?a/iei ca(ycoaaiiyi
eieaaaio a ??cieo i?aaiao.

In vivo oa in vitro aia?oa aeyaeaii, ui iaoai?/ia iai?oaeaiiy aeeeeea?
?iaeoeoe?th neioaco aeanoeio.

In vitro aia?oa iieacaii, ui a?aeiia?aeue ee?oei ai?oe oa oe??e 3- oa
12-i?ny/ieo uo??a ia aeaoi?iaoe?th neeaaea?oueny c ?ioaineo?eaoe??
neioaco eieaaaio ? cieaeaiiy nooiaiy a?ae?ie-neethaaiiy o iueiio
i?ie?io. I?e oeueiio ??aaiue iiia?a/iiai ca(ycoaaiiy eieaaaio o
iaeii??/-ieo uo??a c? c?inoaiiyi iaaaioaaeaiiy nii/aoeo i?aeaeuo?oueny,
a iio?i cieaeo?oueny; o 3-i?ny/ieo – oey caeaaei?noue ia? i?ioeeaaeiee
oa?aeoa?

Aeyaeaii, ui iaoai?/i? iai?oae, i?e yeeo niinoa??aathoueny iaeneiaeuei?
aaee/eie ?ioaineaiinoae neioaco oa eiioeaio?aoe?e eieaaaio a oe??? oa
ai?o? uo??a, o 3-i?ny/ieo oaa?ei aeu? i?ae o 12-i?ny/ieo. Oea iiaea aooe
iia(ycaia c a?eueoith aei?noe?noth i?aeee?oeiiiai iao?eeno niieo/ii?
oeaieie iaeii??/ieo oaa?ei, ca ?aooiie aeuiai nooiaiy iiia?a/iiai
ca(ycoaaiiy a iueiio eieaaaio.

Ia i?eeeaae? oe??e in vitro aeyaeaii, ui a oiiaao ae?? iaoai?/iiai
iaaaioaaeaiiy o i?enooiino? nai?ea?aaceaeo — aeieaoi?o ooai?aiiy
iiia?a/ieo eiaaeaioieo ca(yce?a o eieaaai? – niinoa??aa?oueny cieaeaiiy
eiioeaio?aoe?? eieaaaio a oeaiei?, yea iia(ycaia, iiaeeeai, c
ii?ooaiiyie o eaioethc? ia?aaea/? neaiaeo a?ae i?aeee?oeiiiai iao?eeno
aei yae?a.

Aia?oa iieacaii, ui a aaceaeueoe??aiio ioi/aii? o a?aenooiino? iai?oae
niinoa??-aa?oueny i?aeaeuaiiy, a a ioi/aii? c 2,5 iI Na2+ — cieaeaiiy
?ioaineaiino? neioaco eieaaaio, ui iiaea na?ae/eoe i?i o/anoue
eaeueoe?ecaeaaeieo a?ee?a, cie?aia eiiieaeno Na/eaeueiiaeoe?i, a
ia?aaeaaaii? neaiaeo ia i?iiioi?e no?oeoo?ieo aai?a eieaaaio.

Oai?aoe/ia oa i?aeoe/ia cia/aiiy. I?iaaaeai? aeine?aeaeaiiy
aeicaieythoue aeyaeoe aianie i?aeee?oeiiiai iao?eeno a iaoai?ci
ia?aaea/? neaiaeo i?i ae?th iaoai?/iiai iai?oaeaiiy ia no?oeoo?i? aaie
eieaaaio oa aeanoeio ? aaie, ye? a?aeiia?aeathoue ca neioac aeayeeo
oa?iaio?a i?ioean?iao eieaaaio. Aeyaeai? aoaie iai?io eieaaaio, ye?
aaciina?aaeiuei aieeaathoue ia a(ycei-i?oaei? aeanoeaino? niieo/ii?
oeaieie i?e ae?? ia ia? iaoai?/iiai iaaaioaaeaiiy. Aea/aia ae?y
a?ieia?/ii aeoeaieo ?a/iaei (Na2+, AAOA, oei?i?iiacei, nai?ea?aaceae),
ui aeicaieythoue aeyaeoe ?ieue aioo??oiueiee?oeiieo iina?aaeiee?a a
?aaoeyoe?? ?ioaineaiino? ??cieo aoai?a iai?io eieaaaio oa ni?yiiaaii
?aaoethaaoe ? caaeaaaoe a(ycei-i?oaei? aeanoeaino? eiai iaaeiieaeoey?ieo
ooai?aiue ? niieo/ii? oeaieie a oe?eiio.

Iaea?aeai? ?acoeueoaoe aeicaieythoue aeiiiaieoe no/ani? oyaeaiiy i?i
iieaeoey?i? ia-oai?cie noaiiaeaiiy ooieoe?iiaeueieo aeanoeainoae
niieo/ii? oeaieie o ii?i? oa iaoieia??. Aiie iiaeooue aooe aeei?enoai? o
nii?oeai?e iaaeeoeei?, iaae?ao???, aa?iioieia??, a oaeiae o ee?ieoe?
caoai?thaaiue niieo/ii? oeaieie.

Iniaenoee aianie caeiaoaa/a: i?aea??, ia?iaea oa aiae?c e?oa?aoo?ieo
aeaieo; o/anoue o iinoaiiaoe? 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?? i?aeee?oeiiiai iao?eeno niieo/ii?
oeaieie ??cieo i?aai?a uo??a aei aieeao iaoai?/iiai iaaaioaaeaiiy;
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), II eiia?an? iaoio?c?ieia?a Oe?a?ie (Ee?a, 1996), VII
Oe?a?inueeiio a?io?i?/iiio c(?cae? (Ee?a, 1997), XV c(?cae?
Oe?a?inueeiai o?c?ieia?/iiai oiaa?enoaa (Aeiiaoeuee, 1998), III
I?aeia?iaeiiio neiiic?oi? “Aeieiae/aneea iaoaieciu noa?aiey” (Oa?e?a,
1998), II 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 22
iaeiaio-aaiue. ?iaioo aeeeaaeaii ia 120 noi??ieao ae?oeiaaiiai oaenoo,
aiia i?noeoue 5 oaaeeoeue ? 18 ?enoie?a.

E?oa?aoo?iee iaeyae

O ia?oiio ?icae?e? ?iaioe iaaeaii ianya e?oa?aoo?e ca oaiith
aeena?oaoe??. Iniaeeao oaaao i?eae?eaii iieno neioaco oa i?ioean?iao
eieaaaio ? aeanoeio oa aieeao ia aeaye? noaae?? ?o iai?io a?ieia?/ii
aeoeaieo ?a/iaei ??ciiai iioiaeaeaiiy. Iaaaaeai? aeai? i?i aoaeiao oeeo
a?ee?a oa ?o iaaeiieaeoey?ieo ooai?aiue, a oaeiae i?i ca(ycie ?o
no?oeoo?ieo ia?aiao??a c a(ycei-i?oaeieie aeanoeainoyie niieo/ii?
oeaieie i?aai?a.

?icaeyiooa ?ieue iaoai?/iiai iaaaioaaeaiiy o oi?ioaaii?
no?oeoo?ii-ooieoe?iiaeueieo aeanoeainoae niieo/ii? oeaieie ??cieo
i?aai?a.

Iaaaaeai? no/ani? oyaeaiiy i?i iieaeoey?i? iaoai?cie ia?aaea/? neaiaeo
ia i?iiioi?e no?oeoo?ieo aai?a eieaaaio i?e ae?? iaoai?/iiai
iaaaioaaeaiiy ia niieo/io oeaieio.

Iaoa??aee oa iaoiaee aeine?aeaeaiue

Aeine?aeaeaiiy aoee i?iaaaeai? ia 3- ? 12-i?ny/ieo uo?ao — naieao e?i??
A?noa? in vivo oa in vitro. Ia(?eoii aeine?aeaeaiue aoee eieaaai
oainoiaeo oa ao?eiaeo nooiaeeeue, oe??e nieie, ai?oe, na?oeaaiai i’yco;
aeanoei oe??e nieie oa ai?oe ? noia?i? a?ee? oe??e, ai?oe oa na?oeaaiai
i’yco. Aea/aee iai?i eieaaaio oa aeanoeio (?ioaineai?noue neioaco oa
eiioeaio?aoe?th iaio a?ee?a o oeaieiao, nooi?iue a?ae?ieneethaaiiy
i?ie?io a eieaaai? oa nooi?iue eiai iiia?a/iiai ca’ycoaaiiy), a oaeiae
a’ycei-i?oaei? aeanoeaino? (i?oei?noue, aei?noe?noue, ?icoyaei?noue)
nooiaeeeue, oe??e oa ai?oe.

O aeine?aeao in vivo uo?ai ?iaeee aioo??oiuei/a?aai? ?i(?eoe?? 14N- ?
3I- i?ie?io c ?aooieo 0,5 iAe/a iane o?ea ca 48 aiaeei aei ii/aoeo
aeine?aeo.

O aeine?aeaeaiiyo in vitro c?acee oeaiei ?ieoaoaaee o ?ic/ei?
??iaa?a-E?aana, ui i?noea 3I-i?ie?i aai 14N-aeai?i, i?ioyaii 6 aiaeei
i?e 37(N, ye aac iai?oae, oae ? i?ae ae??th iaoai?/iiai iaaaioaaeaiiy
[Eaaaaeaa, 1978; Martens, 1989]. Oe??o oa ai?oo i?ae /an ?ieoaaoe??
i?aeaeaaaee ?icoyaoaaiith a ae?aiacii? noaoe/ieo iai?oa 0-5o105I/i2
ocaeiaae i?iaeieueiiai iai?yieo c?ace?a.

?ioaineai?noue neioaco eieaaaio oa aeanoeio ioe?ithaaee ca ieoiiith
?aae?iaeoeai?noth o ieo 14N-ienei?ie?io aai 3I-ienei?ie?io [Caia?aaaa,
1977; Eaaaaeaa, 1978; Martens, 1989] ? 14N-aeai?io [Foster, 1982]
a?aeiia?aeii.

Nooi?iue a?ae?ieneethaaiiy i?ie?io o eieaaai? aecia/aee ca a?aeiioaiiyi
3I-Ii?i/(3I-I?i+3I-Ii?i) [Berg, 1982] i?ney ?iciiae?eo ?aae?iaeoeaieo
i?ie?io oa ienei?ie?io [Ca-ia?aaaa, 1977].

Eiioeaio?aoe?th eieaaaio oa aeanoeio ioe?ithaaee ca ai?noii ienei?ie?io
[Stegemann, Stalder, 1967] i?ney ?iciiae?eo oeeo a?ee?a [Neooeeee,
1969].

Nooi?iue iiia?a/iiai ca(ycoaaiiy eieaaaio nooiaeeeue oa oe??e
ioe?ithaaee ca eiai ?ic-/eii?noth o 1 I ?ic/ei? NaCl [ Ooaaneay,
Ia?neee, 1982], a eieaaaio ai?oe oa na?oeaaiai i(yco — o 0,5 I ?ic/ei?
NI3NIII, ui i?noea 2,5% iaineio [Ieeeoeia, 1993], aeey oeeo ?ic/ei?a ca
aeiiiiiaith aeaeo?ioi?aco a IAAA aoea iieacaia i?aeoe/ii iiaia
a?aenooi?noue iie?iaioeae-ieo eaioetha?a, ui ca(ycai? eiaaeaioii.

Noia?io eiioeaio?aoe?th ?ioeo a?ee?a aei??thaaee ca oe?iceiii
[A?aneiiae/, A?iaeia, 1987].

Aaee/eie i?oeiino? ia ?ic?ea, iiaeoey THiaa a iaeano? i?oaeii?
aeaoi?iaoe?? oa aaee/eio iai?oaeii? aeaoi?iaoe?? ?ic?aoiaoaaee ca
e?eaeie iai?oaa-aeaoi?iaoe?y aeey c?ace?a oeaiei [Aeaenaiaea?, 1970;
Ia?neee, 1987].

O /anoei? aenia?eiaio?a aeine?aeaeoaaee aieea nai?ea?aaceaeo (20 iI),
?ii?a Na2+ (2,5 iI), AAOA (5iI) oa oei?i?iiaceio (0,1 iI) ia
aeine?aeaeoaai? noaae?? iai?io eieaaaio oe??e oa ai?oe ? ?o iaoai?/i?
aeanoeaino? a oiiaao ?ieoaaoe?? in vitro, ui aoee iaaaaeai? ?ai?oa.

Noaoenoe/io ia?iaeo io?eiaieo ?acoeueoao?a i?iaiaeeee ca iaoiaeii
Nouethaeaioa-O?oa?a.

?acoeueoaoe aeine?aeaeaiue oa ?o iaaiai?aiiy

Iai?i, neeaae oa a(ycei-i?oaei? aeanoeaino? niieo/ii? oeaieie ??cieo
i?aai?a.

Aeey aea/aiiy ca(yceo i?ae iai?iii eieaaaio oa aeanoeio, ai?noii oeeo
a?ee?a, oa?aeoa?enoeeith ?o iaaeiieaeoey?ieo no?oeoo? a oeaieiao oa ??
a(ycei-i?oaeieie aeanoeainoyie, aea/ai? i?aaie aoei ?iciiae?eaii ia o?e
a?oie – aeaa oeia nooiaeeeue, oe??a oa ai?oa, no?iee e?aiai ? i?aaiai
oeoii/e?a oa i?aeoeoieiaa ia?aai?iaeea. Iaaeiieaeoey?i? ooai?aiiy
niieo/ii? oeaieie i?aai?a o eiaei?e c oeeo a?oi caciathoue aiaeia?/ieo
ca oa?aeoa?ii, aea ??cieo ca aaee/eiith ooieoe?iiaeueieo (iaoai?/ieo)
iai?oaeaiue, a ?o a?io?i?/iee neeaae ? aeinoaoiuei noiaeei.

Oae, ao?eia? nooiaeeeey a i?aai?ci? caciathoue cia/ii a?eueoeo
iaaaioaaeaiue, i?ae oainoia?. A?aeiia?aeii c oeei, i?oei?noue ia ?ic?ea
? iiaeoeue THiaa a iaeano? e?i?eii? aeaoi?iaoe??, ui oa?aeoa?eco?
aei?noe?noue, i?eaeecii a aeaa ?ace aeua, a iaeanoue iai?oaeii?
aeaoi?iaoe?? i?eaeecii a aeaa ?ace ieae/a o ao?eiaeo, i?ae o oainoiaeo
nooiaeeeue (oaae. 1). A nooiaeeeeyo oe? aeanoeaino? caaaeii
aecia/athoueny eiioeaio?aoe??th eieaaaio oa no?oeoo?ith eiai
iaaeiiea-eoey?ieo ooai?aiue.

Oaaeeoey 1. Iaoai?/i? aeanoeaino? niieo/ii? oeaieie i?aai?a
3-i?ny/ieo uo??a

Iieaciee_I ? A A I___Oainoia? nooiaeeeey_Ao?eia?
nooiaeeeey_Oe??a_Ai?oa__I?oei?noue ia ?ic?ea, (, I/i2 o
106_3,6(0,51_9,5(0,72_4,1(0,63_2,2(0,38__Iiaeoeue THiaa, A, I/i2 o
108_2,1(0,20_4,7(0,51_1,2(0,19_3,2(0,29__Iai?oaeia aeaoi?iaoe?y, (,
%_6_3,7_14_50__ sse aeaeii, eiioeaio?aoe?y eieaaaio a ao?eiaeo ?
oainoiaeo nooiaeeeeyo i?aeoe/ii iaeiaeiaa. A oie aea /an nooi?iue eiai
iiia?a/iiai ca(ycoaaiiy ? ??aaiue a?ae?ieneethaaiiy a iueiio i?ie?io
aeua a ao?eiaeo nooiaeeeeyo. A?aeiii, ui iaeaea? oe? iiaeeo?eaoe??
ca?eueoothoue no?oeoo?io noaa?euei?noue eieaaaiiaeo ooai?aiue ye ia
iieaeoey?iiio, oae ? ia iaaeiieaeoey?iiio ??aiyo [Berg, Prockop, 1973;
Viidik, 1973].

Oaaeeoey 2. No?oeoo?ii-iaoaaie?/i? oa?aeoa?enoeee niieo/ii? oeaieie
i?aai?a

3-i?ny/ieo uo??a

I?aai_Ieoiia ?aae?iaeoeai?noue 3I-Ii?i a eieaaai? (?ii/oa/ia
eieaaaio)_Eiioeaio-?aoe?y eieaaaio (% ia nooo iano)_?ic/ei-i?noue
eieaaaio, %_Nooi?iue a?ae?iene-ethaaiiy i?ie?io

a eieaaai?_Eiioeaio-

?aoe?y aeanoeio (% ia nooo iano)__Oainoia? nooiaeeeey
_180(11,3_95,4(1,41*_50,0(0,81_0,45(0,012*_-__Ao?eia? nooiaeeeey
_85(7,5_92,0(2,19*_39,1(0,85_0,51(0,017*_-__Oe??a
_1800(125_59,0(2,72_1,6(0,25_0,50(0,011*_5,1(0,85__Ai?oa

_5000(341_15,1(1,42*_4,0(0,68_0,46(0,015*_15,6(3,17__E?aee oeoii/ie
_21000(1027_15,9(1,39*_17,0(4,29_0,45(0,014*_-__I?aaee
oeoii/ie
_13000(853_27,2(2,14_32,1(5,91*_0,49(0,012*_-__I?aeoeoieiaa ia?aai?iaeea
_18000(941_21,3(1,71_29,0(5,10*_0,47(0,011*_-__I?ei?oea: ooo oa a on?o
ianooiieo oaaeeoeyo ciaeii “*” iicia/athoueny ??cieoe? i?ae
?acoeueoaoaie, ui iaa??ia?aei? (?>0,05)

Oa?aeoa?ii, ui ?ioaineai?noue iai?io eieaaaio a oainoiaeo nooiaeeeeyo
aeua, i?ae a ao?eiaeo, i?i ui na?ae/eoue iaioa ieoiia ?aae?iaeoeai?noue
a ieo 3I-ienei?ie?io a eieaaai? inoaii?o (oaae. 2). Oea oaeiae aeaco? ia
a?eueoo c??e?noue ? no?oeoo?ii-iaoaaie?/io noaa?euei?noue ao?eiaeo
nooiaeeeue o ii??aiyii? c oainoiaeie.

I?e iiae?aiiio oa?aeoa?? ?iciiae?eo iaoai?/ieo iai?oaeaiue a oe??? oa
no?ioe? ai?oe, aaee/eie oeeo iai?oa o oe??? aeua. O oie aea /an no?iea
ai?oe cacia? cia/ii a?eueoeo iai?ioieo a?aeiinieo aeaoi?iaoe?e. Oe?
ooieoe?iiaeuei? iniaeeaino? iaio i?aai?a caaacia-/othoueny ??cieoeyie ?o
iaoai?/ieo aeanoeainoae.

Oae, i?oei?noue oe??e i?aeoe/ii a aeaa ?ace aeua, a ?? ?icoyaei?noue a
iaeano? iai?oaeii? aeaoi?iaoe?? a o?e ?ace ieae/a, i?ae no?iee ai?oe.
I?e oeueiio inoaiiy a iaeano? i?oaeii? aeaoi?iaoe?? ia? a?eueoo
aei?noe?noue, ui caaacia/o? iai?ioi?noue a oeeeeao ??
?icoyaoaaiiy-coyaoaaiiy (oaae. 1). O naith /a?ao ??cieoe? iaoai?/ieo
aeanoeainoae oe??e oa ai?oe aec-ia/athoueny ?o no?oeoo?ii-iaoaaie?/ieie
oa?aeoa?enoeeaie. Naia oae a?eueoa i?oei?noue ? iai-oa ?icoyaei?noue
oe??e o ii??aiyii? c ai?oith caaacia/othoueny cia/ii a?eueoith
eiioeaio?aoe??th a i?e eieaaaio ? a?eueo aenieei nooiaiai eiai
iiia?a/iiai ca(ycoaaiiy (oaae. 2). Ia a?aei?io a?ae nooiaeeeue
a(ycei-i?oaei? aeanoeaino? oe??e ? ai?oe aaaaoi a /iio caeaaeaoue a?ae
ai?noo aeanoeio. Eiai eiioeaio?aoe?y a ai?o? i?eaeecii a o?e ?ace aeua,
i?ae o oe??? (oaae. 2), ui aecia/a? cia/ii a?eueoo cae?ai?noue ?? no?iee
aei aenieeo aaee/ei iai?ioieo aeaoi?iaoe?e.

Oae naia, ye ? o aeiaaeeo nooiaeeeue a?eueoa no?oeoo?ia noaa?euei?noue
iaaeiieaeoey?ieo eieaaaiiaeo ooai?aiue oe??e iia(ycaia c ieae/ei ??aiai
iai?io oeueiai a?eeo a i?e o ii-??aiyii? c ai?oith (oaae. 2).

Na?oeaaee i(yc /eieoue ia o?eueee ianeaiee (ca ?aooiie eieaaaiiaeo), aea
e aeoeaiee (ca ?aooiie i(yciaeo aieieii) ii?? i?eeeaaeaiiio aei iueiai
iai?oaeaiith. Oiio ?nio? ca(ycie i?ae ai?noii eieaaaio, i(yciaeo a?ee?a
? iaoai?/iei iaaaioaaeaiiyi, yea ?icaeaa?oueny a ??cieo a?aeae?eao
na?oey. Oae, ai?no eieaaaio ? i?i?iaeueiei a no?ioe? e?aiai,
iaeneiaeueiei a no?ioe? i?aaiai oeoii/e?a ? ia? i?ii?aeia cia/aiiy a
i?aeoeoieia?e ia?aai?iaeoe? (oaae. 2). Oea ei?aeth? c oei, ui no?iea
e?aiai oeoii/ea, yea cacia? iaeneiaeueiiai e?ia(yiiai oeneo, ai?uo?
iaea?eueoo e?euee?noue i?io?a?ee, a a no?ioe? i?aaiai oeoii/ea, yeee ia
cacia? oaeiai oeneo, ?o cia/ii iaioa [Neooeeee, 1969].

Oae naia, ye ? a ?ioeo aeine?aeaeaieo i?aaiao, ?ioaineai?noue neioaco
eieaaaio a a?aeae?eao na?oey iaa?iaiii?iii?oe?iiaeueia nooiaith
noaa?eueiino? eiai iaaeiieaeoey?ieo ooai?aiue (oaae. 2).

Oaeei /eiii, caaaeuei? caeiiii??iino? ca(yceo i?ae ?ioaineai?noth iai?io
eieaaaio oa aeanoeio ? no?oeoo?iith ?ieeth, yeo aiie aeeiiothoue o
aea/aieo ??ciiaeaeao niieo/ii? oea-ieie, na?ae/aoue, ui oe?
no?oeoo?ii-iaoaaie?/i? iniaeeaino? o?nii ei?aeththoue c iaoai?/ieie
iai?oaeaiiyie, yeeo caciathoue oe? ??ciiaeaee.

Aieea iaoai?/iiai iaaaioaaeaiiy ia iai?i, neeaae oa a(ycei-i?oaei?
aeanoeaino? oe??e oa ai?oe in vitro. ?acoeueoaoe aeine?aeaeaiue
a(ycei-i?oaeieo aeanoeainoae iaio i?aai?a na?ae/aoue, ui i?ae ae??th
aea/aieo iaaaioaaeaiue aiie noathoue a?eueo aei?noeeie ? iaio
?icoyaeieie (oaae.3). Ui ae noino?oueny i?oeiino?, oi a iaeano? iai?oa,
aea a?aeaoaa?oueny i?aeaeuaiiy aei?noeino?, aiia aeinoia??ii ia
ci?ith?oueny, a i?e cieaeaii? aei?noeino? — ciaioo?oueny.

Oaaeeoey 3. Iaoai?/i? aeanoeaino? oei?e oa ai?oe uo?ia iiney iieoaaoeii

i?e ?icieo iaaaioaaeaiiyo.

Iieaciee_I ? a a i___O e i ? a_A i ? o a___( = 0_( = 2,4o105 I/i2_( =
4,8o105 I/i2_( = 0_( = 2,2o105 I/i2_( = 4,4o105 I/i2__I?oei?noue ia
?ic?ea, (, I/i2 o
106_3,3+0,44*_3,4+0,35*_2,6+0,32_3,3+0,48_2,2+0,21*_1,8+0,11*__ Iiaeoeue
THiaa, A, I/i2 o 107_7,5
+0,91_12,0+1,38_9,7+1,08_0,7+0,09_2,0+0,41*_1,8+0,36*__Iai?oaeia
aeaoi?iaoe?y, (, %_16_7_10_65_31_60__

I?e?iaeii, ui oaea ci?ia a(ycei-i?oaeieo aeanoeainoae niieo/ii? oeaieie
iaio i?aai?a iiaeiia caeaaeaoe a?ae aieeao iaoai?/iiai iaaaioaaeaiiy ia
iai?ii? i?ioeane a i?e.

Ia ?en. 1 iaaaaeai? caeaaeiino? eiioeaio?aoe?e eieaaaio, aeanoeio oa
oe?iceio a?ae iai?oaeaiiy a oe??? oa ai?o? i?ney ?o ?ieoaaoe??. A iaio
i?aaiao aeey eieaaaio oa aeanoeio oe? caeaaeiino? yaeythoue niaith e?ea?
c iaeneioiii, yeee i?eiaaea? i?aeoe/ii ia oo ae naio iaeanoue iai?oa –
(2,1-2,4) o 105 I/i2

_

?en. 1. Aieea iaoai?/iiai iaaaioaaeaiiy ia eiioeaio?aoe?th eieaaaio (1),
aeanoeio (2) ? oe?iceio (3) a oe??? oa ai?o? uo??a i?ney ?ieoaaoe?? in
vitro (% ia nooo iano).

_

?en. 2. Aieea iaoai?/iiai iaaaioaaeaiiy ia ieoiio ?aae?iaeoeai?noue
3I-ienei?ie?io a caaaeueiiio eieaaai? (1), 14N-aeai?io a caaaeueiiio
aeanoei? (2), 3I-ienei?ie?io a na?aeineioaciaaiiio eieaaai? (3) oe??e oa
ai?oe uo??a i?ney ?ieoaaoe??.

C ?en. 2 aeaeii, ui caaaeuei? ?ioaineaiino? neioaco iaio a?ee?a, yea
aecia/a?oueny ca ieoiiith ?aae?iaeoeai?noth 3I-ienei?ie?io aeey
noia?iiai eieaaaio oa 14N-aeai?io aeey noia?iiai aeanoeio, oaeiae
yaeythoue niaith e?ea? c iaeneioiii i?e (0,4-0,8) o105 I/i2. Oea aeaco?,
ui caeaaei?noue eiioeaio?aoe?? oeeo a?ee?a a oe??? oa ai?o? a?ae iai?oae
? ?acoeueoaoii ci?ie ?ioaineaiino? ?o neioaco. Aaaeeeai, ui a iaio
i?aaiao iaeano?, aea niinoa??aathoueny iaeneioie aeeth/aiiy
?aae?iaeoeaieo ai?iieeneio aei eieaaaio oa aeanoeio oa iaeneioie
eiioeaio?aoe?e oeeo a?ee?a, ca?aathoueny. Oea iiaea na?ae/eoe i?i oa, ui
a ??cieo i?aaiao a?aeiia?aeue niieo/ii? oeaieie ia aeaoi?iaoe?th ye?nii
iiae?aia ? niaoeeo?/ia aeey ee?oei, ui neioacothoue eieaaai oa aeanoei.

Ne?ae aeacaoe, ui caaaeueia ?ioaineai?noue neioaco – oea ?ioaa?aeueiee
iieaciee, yeee caeaaeeoue ye a?ae an??? e?eueeino? neioacoth/eo ee?oei,
oae ? a?ae ?ioaineaiino? neioaco a eiaei?e c ieo. Aeai? ?en. 2
aeacothoue, ui a iaeano? (0,9-5) o 105 I/i2 ia oii? cieaeaiiy ieoiii?
?a-ae?iaeoeaiino? 3I-ienei?ie?io a caaaeueiiio eieaaai? oe??e oa ai?oe,
ieoiia ?aae?iaeoeai?noue na?aeineioaciaaiiai eieaaaio i?iaeiaaeo? ?inoe.
Oaeee aoaeo iiaea na?ae/eoe i?i i?aeaeuaiiy ?ioaineaiino? neioaco
eieaaaio a ie?aieo ee?oeiao i?ioyaii anueiai aeine?aeaeaiiai ae?aiaciio
iai?oa. Cieaeaiiy ae caaaeueii? ?ioaineaiino? neioaco iaio a?ee?a iiaea
aooe iia(ycaia c ii?ooaiiyi aai oe?e?niino? /anoeie neioacoth/eo ee?oei,
aai no?oeoo?e i?aeee?oeiiiai iao?eeno i?ae aieeaii aaeeeeo iai?oa, aai
iaiia aoaeoaie ?acii. A ca(yceo c oeei ia?aaea/a neaiaeo i?i iaoai?/ia
iaaaioaaeaiiy ia ee?oeiio iiaa?oith ci?ith?oueny.

_

?en. 3. Aieea iaoai?/iiai iaaaioaaeaiiy ia ?ic/eii?noue eieaaaio oe??e
(1) oa ai?oe (2) uo??a i?ney ?ieoaaoe??.

C ?en. 3 aeaeii, ui, noaey/e c aeeiai?ee caeaaeiino? ?ic/eiiino?
eieaaaio, nooi?iue iiia?a/iiai ca(ycoaaiiy eiai iaaeiieaeoey?ieo
no?oeoo? nii/aoeo cieaeo?oueny ((0-2,4) o 105 I/i2), a iio?i ciiao
i?aeaeuo?oueny, oi/a e ia aeinyaa? ia?aeiiiai ??aiy. Iiaeia i?eionoeoe,
ui oea cieaeaiiy iiia?a/iiai ca(ycoaaiiy iiaea aooe iane?aeeii ci?ie
oa?aeoa?o ia?aaea/? neaiaeo a oiiaao aaeeeeo aeaoi?iaoe?e oeaieie.

Aeyaeaia ciaioaiiy nooiaiy iiia?a/iiai ca(ycoaaiiy eieaaaio ? oaeiae
iaei??th c i?e-/ei ciaioaiiy i?oeiino? niieo/ii? oeaieie (oaae.3), ui
niinoa??aa?oueny a aeine?aeao in vitro.

??aaiue a?ae?ieneethaaiiy i?ie?io a eieaaai? oe??e oa ai?oe i?e ae??
iaaenieeo ((0-1)(105 I/i2) iaaaioaaeaiue noaiiaeoue aeecueei 0,5, a i?e
iiaeaeueoiio ?ino? iaaaioaaeaiiy cieaeo?oueny, aeinyaath/e o oe???
aaee/eie 0,38+0,012, a a ai?o? – 0,35+0,011.

Oaeei /eiii, o 3-i?ny/ieo uo??a in vitro a?aeiia?aeue ee?oei niieo/ii?
oeaieie ia aeaoi?iaoe?th, ie??i ?iaeoeoe?? neioaco eieaaaio oa aeanoeio,
aaaea, iiaeeeai, aei ?ia?a??oaaiiy aeayeeo noaae?e i?ioean?iao
eieaaaio.

Caeaaeiino? eiioeaio?aoe?? oe?iceio a?ae iaaaioaaeaiiy a oe??? oa ai?o?
iaaaaeai? ia ?en.1. O oe??? i?ae aieeaii aeaoi?iaoe?? ai?no oe?iceio
ci?ith?oueny oae naii, ye ? ai?no eieaaaio oa aeanoeio, a a ai?o?
aeeiai?ea eiai eiioeaio?aoe?? ia? i?ioeeaaeiee oa?aeoa?. Ine?eueee
oe?i-cei ? iieacieeii a?ee?a, ye? a ia?oo /a?ao aoiaeyoue aei neeaaeo
ae?eii?ioa?i?a oa i?ioai-ae?eai?a ? yaeythoue niaith no?oeoo?i?
eiiiiiaioe i?aeee?oeiiiai iao?eeno [Robert, Moczar, 1985], iaea?aeai?
aeai? na?ae/aoue, ui ia?aaoaeiae eiai no?oeoo?e, a yeeo aa?ooue o/anoue
i?ioaiae?eaie oa ae?eii?ioa?ie, iathoue a oe??? oa ai?o? i?ioeeaaeiee
oa?aeoa?. Oea, i/aae-aeii, iia(ycaia c iniaeeainoyie ooieoe?iiaeueieo
aeanoeainoae oeeo i?aai?a.

Oaeei /eiii, i?iaaaeaia aeine?aeaeaiiy aeyaey?, ui iaa?oue in vitro a
oeaieiao caa??-aathoueny iaoai?cie, ye? cae?eniththoue ?o aaeaioaoe?th
aei iaoai?/ieo iaaaioaaeaiue ca ?aooiie ?nioaaiiy iiceoeaiiai
cai?ioiiai ca(yceo o eaioethc?: iaoai?/ia iaaaioaaeaiiy, ui i?eeea-aeaii
aei oeaieie – iai?i no?oeoo?ieo aeaiaio?a ?? i?aeee?oeiiiai iao?eeno –
ci?ia a(ycei-i?oaeieo aeanoeainoae oeaieie.

Aieea a?eo oaa?ei oa nai?ea?aaceaeo ia oa?aeoa? ee?oeiii? a?aeiia?ae?
i?e ae?? iaoai?/iiai iaaaioaaeaiiy a oe??? oa ai?o?. A ?icae?e?
?icaeyaea?oueny ieoaiiy i?i ca’ycie i?ae aoaeiaith eieaaaiiaiai iao?eeno
? iniaeeainoyie ia?aaea/? neaiaeo i?i iaoai?/ia iaaaioaaeaiiy aei yae?a.

Aeey c’ynoaaiiy oeueiai ieoaiiy aoei aea/aii aieea a?eo oaa?ei ia
ee?oeiio a?aeiia?aeue oe??e oa ai?oe i?e ae?? ia ieo iaoai?/iiai
iaaaioaaeaiiy in vitro.

Ia ?en. 4 iaaaaeai? aeai? i?i caeaaei?noue eiioeaio?aoe?? eieaaaio a
oe??? oa ai?o? a?ae iaoai?/iiai iaaaioaaeaiiy o 3- oa 12-i?ny/ieo uo??a.
sse aeaeii, iaeneiaeueia cia/aiiy eii-oeaio?aoe?e eieaaaio
niinoa??aa?oueny i?e iaaaaaoi iaioeo iai?oaao o 12-i?ny/ieo uo??a, i?ae

o 3-i?ny/ieo (?en. 4). Oe?e aoaeo ? ?acoeueoaoii aiaeia?/ieo ci?i a
?ioaineaiino? neioaco oeueiai a?eeo (?en. 5).

I?e oeueiio nooi?iue a?ae?ieneethaaiiy i?ie?io a eieaaai? oe??e oa ai?oe
o 12-i?ny/ieo uo??a, ye ? o 3-i?ny/ieo, ciaioo?oueny c? c?inoaiiyi
iai?oaeaiiy ? noaiiaeoue i?e iaene-iaeueieo iaaaioaaeaiiyo ((=1,8 o 105
I/i2 aeey oe??e oa (= 2,0 o 105 I/i2 aeey ai?oe) 0,35+ 0,011 oa 0,33+
0,012 a?aeiia?aeii.

_

?en. 4. Aieea iaoai?/iiai iaaaioaaeaiiy ia eiioeaio?aoe?th eieaaaio a
oe??? oa ai?o? 3- oa 12-i?ny/ieo uo??a i?ney ?ieoaaoe?? in vitro.

A?aeiii, ui iaei??th c iniiaieo ci?i o aoaeia? eieaaaiiaeo no?oeoo? c
a?eii ? ca?eue-oaiiy a ieo iiia?a/ieo eiaaeaioieo ca’yce?a, ui a cia/i?e
i??? i?aeaeuo? aei?noe?noue ? cieaeo? ?icoyaei?noue oeaieie
[Viidik,1985].

_

?en. 5. Aieea iaoai?/iiai iaaaioaaeaiiy ia ieoiio ?aae?iaeoeai?noue
3H-ienei?ie?io a caaaeueiiio eieaaai? oe??e oa ai?oe 3- oa 12-i?ny/ieo
uo??a.

Aeey ia?aa??ee i?eiouaiiy i?i oa, ui iiia?a/ia ca’ycoaaiiy eieaaaio
iiaea ci?ithaaoe a?aeiia?aeue ee?oei ia iaoai?/ia iaaaioaaeaiiy, aoei
aea/aii aieea nai?ea?aaceaeo ia eiioeaio?aoe?th oa ?ic/eii?noue eieaaaio
oe??e a i?ioean? ?? ?ieoaaoe?? in vitro c ??cieie iai?oaaie.
Nai?ea?aaceae o eiioeaio?aoe?? 20 iI caaaaea? ooai?aiith iiia?a/ieo
eiaaeaioieo ca’yce?a a eieaaai? [Neooeeee,1972].

C ?en. 6A aeaeii, ui i?e ?ieoaaoe?? c aeiaeaaaiiyi nai?ea?aaceaeo o
a?aenooiino? iai?oae eiioeaio?aoe?y eieaaaio a oe??? cieaeo?oueny ?
iaa?oue i?e iaeneiaeueieo iaaaioaaeaiiyo aiia ia ia?aaeuo? ii?ie. Ui
noino?oueny eiaaeaioii ca’ycaiiai eieaaaio, eiai e?euee?noue, noaey/e c
?ic/eiiino?, ??cei ciaioo?oueny (?en. 6A).

__

?en. 6. Aieea nai?ea?aaceaeo ia eiioeaio?aoe?th (A) ? ?ic/eii?noue (A)
eieaaaio oe??e i?e ?ieoaaoe?? in vitro:

1 — ii?ia; 2 — nai?ea?aaceae, ( = 0; 3 -nai?ea?aaceae, ( = 2,4 o 10 5
I/i2; 4 — nai?ea?aaceae, ( = 4,8 o 10 5 I/i.

Oaeei /eiii, iaea?aeai? aeai? i?aeoaa?aeaeothoue i?eiouaiiy i?i oa, ui
eieaaaiiaee iao?een niieo/ii? oeaieie aa?a o/anoue a ia?aaeaaaii?
neaiaeo i?i iaoai?/ia iaaaioaaeaiiy. I?e oeueiio nooi?iue eiai
aei?noeino?, yea aecia/a?oueny e?euee?noth o iueiio iiia?a/ieo
eiaaeaioieo ca’yce?a, aieeaa? ia aaee/eio iai?oaeaiiy, i?e yeiio aeieea?
ee?oeiia a?aeiia?aeue.

Aieea Na2+ ? oei?i?iiaceio ia iai?i, no?oeoo?o oa a(ycei-i?oaei?
aeanoeaino? oe??e oa ai?oe. ?acoeueoaoe, iaaaaeai? ia ?en. 7.1. ? 7.2,
iieacothoue, ui i?e ?ieoaaoe?? o i?e-nooiino? oei?i?iiaceio (0,1iI) aai
AAOA (5 iI), yeee iia(yco? iicaee?oeiiee eaeueoe?e, a?aeaoaa?oueny
i?aeaeuaiiy, a i?e ?ieoaaoe?? o a?ia?eaeueoe??aiio ioi/aii? (2,5 iI) –
cieaeaiiy eiioeaio?aoe?? ye caaaeueiiai, oae ? na?aeineioaciaaiiai
eieaaaio a oe??? oa ai?o?. Oeae aoaeo aaciina?aaeiuei iia(ycaiee c
a?aeiia?aeieie ci?iaie ieoiii? ?aae?iaeoeaiino? 3I-ienei?ie?io ? a
caaaeueiiio, ? a na?aeineioaciaaiiio eieaaai? iaio i?aai?a (?en. 7.3.,
7.4).

Ine?eueee iicaee?oeiiee eaeueoe?e ? iina?aaeieeii aaeeei? e?eueeino?
aioo??oiueiee?-oeiieo i?ioean?a, o oiio /ene? oeo, ui aieeaathoue ia
neioac a?ee?a [Hennigs, Holbrook, 1983; Gerday, Gilles, Bolis, 1988],
iaea?aeai? ?acoeueoaoe iiaeia aeei?enoiaoaaoe, ye iiaeaeue, ui aeicaiey?
cai?iiiioaaoe iiaeeeaee iaoai?ci ca(yceo i?ae iaaaioaaeaiiyi o oeaiei? ?
neioacii eieaaaio a ae?aiacii? ?o i?iii?oe?eii? caeaaeiino?. Iaeiei c
aaaeeeaeo eiiiiiaio?a o eaioethc? ia?aaea/? neaiaeo i?i iaoai?/ia
iaaaioaaeaiiy a?ae iiaa?oi? ee?oeie aei yae?a ? eiiieaen
Na/eaeueiiaeoe?i [Klee, Crouch, Richman, 1980].

___

_?en 7. Aieea Na2+ oa oei?i?iiaceio ia eiioeaio?aoe?th (1), ?ic/eii?noue
(2) eieaaaio oa ieoiio ?aae?iaeoeai?noue 3I-ienei?ie?io a caaaeueiiio
(3) oa na?aeineioaciaaiiio (4) eieaaai? oe??e oa ai?oe i?ney
?ieoaaoe??.

A a?ia?eaeueoe??aiio na?aaeiaeu?, eiee eaeueoe?e iinooia? aei ee?oeie ca
?aooiie eiioeai-o?aoe?eiiai a?aae??ioo, iiaeiia ooai?thaaoeny aeinoaoiy
e?euee?noue aeoeaieo eiiieaen?a Na/eaeueiiaeoe?i, ui noi?iaiaeaeo?oueny
cieaeaiiyi neioaco eieaaaio (?en. 7.3, 7.4).

O a?aenooiino? ?ii?a eaeueoe?th (i?e aeiaeaaaii? AAOA) ??cei
cieaeo?oueny iiaeeea?noue ooai?aiiy oaeeo aeoeaieo eiiieaen?a, oiio ui
/anoea aioo??ee?oeiiiai eaeueoe?th aeoiaeeoue o i?aeee?oeiiee i?ino??.
Oei?i?iiacei, ui o aea/ai?e eiioeaio?aoe?? aoiaeeoue o ee?oeio ? yaey?
niaith eeane/iee aioeeaeueiiaeoe?iiaee aaaio [Kuo, 1992; Klimowski,
1983], oaae caaaaea? ae-ieeiaiith aeinoaoiuei? e?eueeino? eiiieaen?a
Na/eaeueiiaeoe?i. sse aeaeii, a iaio oeeo aeiaaeeao i?e ciaioaii? /enea
aeoeaieo eiiieaen?a Na/eaeueiiaeoe?i a?aeaoaa?oueny i?aeaeuaiiy neioaco
eieaaaio (?en. 7.3., 7.4.).

I?e ae?? iaoai?/iiai iaaaioaaeaiiy ia aea/ai? i?aaie a ieo oaeiae
niinoa??aa?oueny ?iaeoeoe?y neioaco eieaaaio (?en. 2). A?aeiii, ui i?e
aeaoi?iaoe?? ee?oei (iai?eeeaae o?a?iaeano?a) ci?ith?oueny no?oeoo?a ?o
oeeoineaeaoo, ui aaaea aei ia?aaoaeiae no?oeoo?ii-ooie-oe?iiaeueiiai
ca(yceo i?ae eiiiiiaioaie eaioethao ia?aaea/? neaiaeo a?ae
i?aeee?oeiiiai iao-?eeno /a?ac iiaa?oith ee?oeie aei ?? yae?a [Butt,
Bishop, 1997].

Oaeei /eiii, iaeiei c iaoai?ci?a, ui aecia/a? ca?eueoaiiy neioaco
eieaaaio i?e iaoai?/iiio iaaaioaaeaii?, iiaea aooe ci?ia
no?oeoo?ii-ooieoe?iiaeueiiai eaioethao ia?aiino neaiaeo o oeaiei?.
Iaea?aeai? ?acoeueoaoe na?ae/aoue, ui iaeiei c aeaiaio?a oeueiai
eaioethao, yeee aaciina?aaeiuei aieeaa? ia neioac eieaaaio iiaeeeai ?
eiiieaen Na/eaeueiiaeoe?i.

O ii??aiyii? c i?oei?noth ? ?icoyaei?noth oe??e (( = (3,3+0,44) ( 105
I/i2, ( =16 %), yeo ?ieoaoaaee i?e cae/aeieo oiiaao, i?ney ?? ?ieoaaoe??
o a?ia?eaeueoe??aiio ioi/aii? oe? iaoa-i?/i? aeanoeaino? i?aeoe/ii ia
ci?iththoueny (( = (4,0 + 0,56) o 106 I/i2, ( = 17,5 %), a i?ney
?ieoaaoe?? o i?enooiino? AAOA i?oei?noue cia/ii cieaeo?oueny (( = (0,69+
0,015) o 106 I/i2), a ?? ?icoyaei?noue i?aeaeuo?oueny (( = 27%).

A?aeiii, ui Na2+ ? ?i?oe?aoi?ii ooai?aiiy iiia?a/ieo ca(yce?a i?ae
iaeieaaaiiaeie a?e-eaie a i?aeee?oeiiiio iao?een? [Aeueaa?on, A?ae,
Euethen e ae?., 1994]. Oiio ca?eueoaiiy eiai eiioeaio?aoe?? o oeaiei?
aaaea aei i?aeaeuaiiy ?? i?oeiino?, a cieaeaiiy – aei ciaioaiiy oeueiai
iieacieea.

sse aeaeii c ?en.7, i?ney ?ieoaaoe?? i?e i?aeaeuai?e eiioeaio?aoe?? Na2+
ai?no eieaaaio a oeaiei? cieaeo?oueny, a i?e eiai a?aenooiino? –
ca?eueoo?oueny.

Oeae aoaeo iiaeia ?icaeyaeaoe, ye aaeaioeaiee i?ioean, ui ni?yiiaaiee ia
caa?aaeaiiy ia?aeiieo iaoai?/ieo aeanoeainoae oeaieie.

AENIIAEE.

In vivo oa in vitro i?iaaaeaii ii??aiyeueia aeine?aeaeaiiy aieeao
iaoai?/iiai iaaaioaaeaiiy ia neioac eieaaaio, aeanoeio,
iinoo?aineyoe?ei? oa iinoneioaoe/i? iiaeeo?eaoe?? eieaaaio a oe???,
ai?o?, nooiaeeeeyo oa a?aeae?eao na?oey, a oaeiae ia a(ycei-i?oaei?
aeanoeaino? oeeo i?aai?a o 3- oa 12-i?ny/ieo uo??a.

Iieacaii, ui in vivo a i?aaiao, ye? caciathoue o ii??aiyii? c ?ioeie
a?eueoeo iaoai?/ieo iaaaioaaeaiue, i?aeaeuo?oueny eiioeaio?aoe?y
eieaaaio oa ?ioaineai?noue eiai neioaco, nooi?iue a?ae?ieneethaaiiy o
iueiio i?ie?io oa eiai iiia?a/iiai ca(ycoaaiiy. ??aaiue ?ioaineo?eaoe??
eiaeiiai c oeeo i?ioean?a caeaaeeoue a?ae oeio oeaieie.

Ia i?eeeaae? oe??e oa ai?oe in vivo iieacaii, ui eiioeaio?aoe?y a ieo
aeanoeio ei?aeth? ia c aaee/eiaie iai?oaeaiue, yeeo aiie caciathoue, a
c? nooiaiai iaeneiaeueieo cai?ioieo aeaoi?iaoe?e oeeo i?aai?a.

Iieacaii, ui in vivo a ae?aiacii? o?c?ieia?/ieo iaaaioaaeaiue
ni?aa?aeiioaiiy eiioeaio?aoe?e aeanoeio oa eieaaaio, a oaeiae ??aaiue
iiaeeo?eaoe?e inoaiiueiai oi?iothoue iioeiaeuei? ii a?aeiioaiith aei
aaee/eie oa ?iciiae?eaiiy iaoai?/ieo iaaaioaaeaiue a(ycei-i?oaei?
aeanoeaino? niieo/ii? oeaieie i?aai?a.

Iieacaii, ui in vitro, i?ae ae??th iaoai?/iiai iaaaioaaeaiiy
?ioaineaiino? neioaco ? eiioeaio?aoe?? eieaaaio oa aeanoeio a oe??? oa
ai?o? uo??a nii/aoeo ca?eueoothoueny i?ney /iai ii/eiathoue
cieaeoaaoeny. I?e oeueiio iaeneioie ?ioaineaiinoae oeeo caeaaeiinoae o
12-i?ny/ieo uo??a niinoa??aathoueny i?e iaioeo aaee/eiao iaaaioaaeaiue,
i?ae o 3-i?ny/ieo.

In vitro iieacaii, ui a oe??? oa ai?o? oaa?ei ??ciiai a?eo i?ae ae??th
iaoai?/iiai iaaai-oaaeaiiy a?aeaoaa?oueny iino?eia cieaeaiiy nooiaiy
a?ae?ieneethaaiiy i?ie?io a eieaaai? – aioo??oiueiee?oeiii? noaae?? eiai
i?ioean?iao. O oie aea /an ?ioaineai?noue iicaee?oeiii? noaae??
i?ioean?iao eieaaaio – nooi?iue eiai iiia?a/iiai ca(ycoaaiiy –
caeaaeeoue a?ae a?eo oaa?ei. O 3-i?ny/ieo uo??a c? c?inoaiiyi
iaaaioaaeaiiy aiia nii/aoeo cieaeo?oueny, a iio?i – i?aeaeuo?oueny; o
12-i?ny/ieo – oey caeaaei?noue ia? i?ioeeaaeiee oa?aeoa?.

Ia i?aenoaa? aea/aiiy aieeao Na2+ oa oei?i?iiaceio ia ?ioaineai?noue
neioaco oa eiioeaio?aoe?th eieaaaio a oe??? oa ai?o? 3-i?ny/ieo uo??a in
vitro aenoiooi i?eiouaiiy, ui ca?eueoaiiy e?eueeino? eiiieaen?a
Na/eaeueiiaeoe?i i?ecaiaeeoue aei ?ia?a??oaaiiy , a ?? ciaioaiiy – aei
?ioaineo?eaoe?? neioaco eieaaaio. I?eiouaii, ui ?ioaineo?eaoe?y neioaco
eieaaaio a oe??? oa ai?o? i?e ae?? iaoai?/iiai iaaaioaaeaiiy
a?aeaoaa?oueny ca ?aooiie cieaeaiiy aioo??oiueiee?oeiii? eiioeaio?aoe??
?ii?a Na i?e ?o aeoiae? c ee?oeie a ?acoeue-oao? iiyae a ?? iaia?ai?
aeiaeaoeiaeo ii? o i?ioean? aeaoi?iaoe??.

Iieacaii, ui aei?noe?noue i?aeee?oeiiiai iao?eeno niieo/ii? oeaieie, ui
caeaaeeoue a?ae nooiaiy iiia?a/iiai ca(ycoaaiiy a iueiio eieaaaio,
aecia/a? aaee/eio iai?oaeaiiy, i?e yeiio a?aeaoaa?oueny ee?oeiia
a?aeiia?aeue ia iueiai.

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. — ?. 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 a eiaea iicaiii/iuo
aeeaioiuo i?e oaiia?aoo?iie aaeaioaoeee // I?iaeaiu e?eiaeieiaee. —
1997. — ? 3. — N. 27-31.

Aa?aocaiei I.A., Aiaoe A.A., Ia?neee A.Y. Aeeyiea aeaoi?iaoeee ia iaiai
aaeeia e iaoaie/aneea naienoaa ai?ou e eiaee e?un in vitro // Aanoi.
i?iae. aeie. e iaae. — 1997. — ? 25. — N. 12-18.

Aa?aocaiei I.A., Ia?neee A. Y. Aeeyiea ioeaaeaeaiey ia iaiai eieeaaaia e
yeanoeia a eiaea e?un i?e ?acee/iuo iai?yaeaieyo // I?iaeaiu
e?eiaeieiaee. — 1998. — ? 1. — N. 66-67.

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? // XIV
c(?cae Oe?a?inueeiai o?c?ie. oia. — Oac. aeii., Ee?a. — 1994. — N. 59.

Aa?aocaiei I.A., Eociaoeia ?.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 oae-oi??a // ?? eiia?an iaoio?c?ieia?a Oe?a?ie. — Oac.
aeii., Ee?a. — 1996. — N. 148.

Aa?aocaiei I.A., Iaaeia I.A., Ooaanueea E.A., Ia?nueeee ?.A. Iieaeoey?i?
iaoai?cie ci?i no?oeoo?e i?aeee?oeiiiai iao?eeno niieo/ii? oeaieie i?e
iaoai?/iiio iaaaioaaeaii? // VII Oe?a?inueeee a?io?i?/iee c(?cae. — Oac.
aeii., Ee?a. — 1997. — O. ??. — N. 136-137.

Aa?aocaiei I.A., Eocuei?n A.A., I?e?o?ia I.A., Ia?nueeee ?.A. Aieea
aeaoi?iaoe?? ia iai?i a?ee?a oa iaoai?/i? aeanoeaino? ai?oe uo??a in
vitro // Iao. XV c(?caeo Oe?a?inueeiai o?c?ie. oia. — O?c?ieia?/iee
aeo?iae. — 1998. — O. 44, ? 3. — N. 148-149.

Ia?neee A.Y., Aa?aocaiei I.A., Eocueien A.Y., Aaneeueaaa E.A. Aic?anoiua
iniaaiiinoe ooieoeeiiaeueiuo aiciiaeiinoae eieeaaaineioace?othueo eeaoie
// ??? Iaaeaeoia?iaeiue neiiiceoi “Aeieiae/aneea iaoaieciu noa?aiey”. —
Oac. aeiee., Oa?ueeia. — 1998. — N. 26.

Aa?aocaiei I.A., Iaaeia I.A., I?e?o?ia I.A., Ia?nueeee ?.A. Iai?ii?
iaoai?cie ci?oeiaiiy niieo/ii? oeaieie i?ae ae??th iaoai?/iiai
iaaaioaaeaiiy // ?? c(?cae Oe?a?inueeiai a?io?c. oia. — Oac. aeii.,
Oa?e?a.- 1998. — N.120.

AIIOAOe?ss

Aa?aocaiei I.A. Aieea iaoai?/iiai iaaaioaaeaiiy ia neioac eieaaaio,
aeanoeio oa a(yc-ei-i?oaei? aeanoeaino? niieo/ii? oeaieie. – ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe
c niaoe?aeueiino? 03.00.04 — a?io?i?y — Oa?e?anueeee aea?aeaaiee
oi?aa?neoao, Oa?e?a, 1999.

Aeine?aeaeaii aieea iaoai?/iiai iaaaioaaeaiiy ia neioac aeanoeio,
eieaaaio, a?ae-?ieneethaaiiy i?ie?io ? nooi?iue iiia?a/iiai ca(ycoaaiiy
eieaaaio o oe???, ai?o?, na?oeaaiio i(yc? oa nooiaeeeeyo 3- ?
12-i?ny/ieo uo??a in vivo oa in vitro.

In vivo iieacaii, ui oi?ioaaiiy a(ycei-i?oaeieo aeanoeainoae niieo/ii?
oeaieie ? ?acoeueoaoii niaoeeo?/ieo aeey eiaeiiai i?aaia ni?aa?aeiioaiue
?ioaineaiinoae neioaco aean-oeio, eieaaaio oa aea/aieo noaae?e eiai
i?ioean?iao. In vitro anoaiiaeai? a?eia? iniaeeaino? ae?aiacii?a iai?oa,
aea a?aeaoaa?oueny ?iaeoeoe?y neioaco eieaaaio oa eiai i?ioean?iao.
Aia?oa iieacaii, ui iaoai?/ia iaaaioaaeaiiy aeeeeea? ?iaeoeoe?th neioaco
aeanoeio. Iieacaii, ui a aaceaeueoe??aiio na?aaeiaeu?, niinoa??aa?oueny
ca?eueoaiiy, a o a?ia?eaeueoe??aiio – ciaioaiiy ?ioaineaiino? neioaco
eieaaaio. Anoaiiaeaii, ui aei?noe?noue i?aeee?oeiiiai iao?eeno aiee-aa?
ia ia?aaea/o neaiaeo i?i iaoai?/ia iaaaioaaeaiiy ia ee?oeie.

Eeth/ia? neiaa: eieaaai, aeanoei, neioac, i?aeee?oeiiee iao?een,
a(ycei-i?oaei? aeanoeaino?, iaoai?/ia iaaaioaaeaiiy.

AIIIOAOeEss

Aa?aocaiei I.A. Aeeyiea iaoaie/aneiai iai?yaeaiey ia neioac eieeaaaia,
yeanoeia e aycei-oi?oaea naienoaa niaaeeieoaeueiie oeaie. — ?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, 1999.

Enneaaeiaaii aeeyiea iaoaie/aneiai iai?yaeaiey ia neioac yeanoeia,
eieeaaaia, aeae?ieneee?iaaiey i?ieeia, e iiia?a/iia naycuaaiea
eieeaaaia a eiaea, ai?oa, na?aea/iie iuooea e nooiaeeeeyo 3- e
12-iany/iuo e?un in vivo e in vitro.

In vivo iieacaii, /oi oi?ie?iaaiea iioeiaeueiuo aycei-oi?oaeo naienoa
niaaeeieoa-eueiie oeaie yaeyaony ?acoeueoaoii niaoeeoe/aneeo aeey
eaaeaeiai i?aaia niioiioaiee eioaineaiinoae neioaca yeanoeia, eieeaaaia
e eco/aiiuo noaaeee aai i?ioeanneiaa. In vitro onoaiiaeaiu aic?anoiua
iniaaiiinoe aeeaiaciiia iai?yaeaiee, a eioi?uo i?ienoiaeeo eiaeoeoeey
neioaca eieeaaaia e aai i?ioeanneiaa. Aia?aua iieacaii, /oi iaoaie/aneia
iai?yaeaiea aucuaaao eiaeoeoeeth neioaca yeanoeia. Iieacaii, /oi a
aaneaeueoeeaaie n?aaea i?ienoiaeeo oaaee/aiea, a a aeia?eaeueoeeaaie —
nieaeaiea neioaca eieeaaaia, a aeanoeinoue iaaeeeaoi/iiai iao?eena
aeeyao ia ia?aaea/o neaiaea i iaoaie/aneii iai?yaeaiee ia eeaoee.

Eeth/aaua neiaa:, eieeaaai, yeanoei, neioac, iaaeeeaoi/iue iao?een,
aycei-oi?o-aea naienoaa, iaoaie/aneia iai?yaeaiea.

SUMMARY

Garbuzenko O.B. The influence of the mechanical load on collagen and
elastin synthesis ane visko-elastic properties of the connective tissue.
— Manuskript.

Thesis for a scientific degree of candidate of biological sciences by
speciality 03.00.04 — biochemistry. — Kharkov State University,
Kharkov,1999.

The investigation of the mechanical load influence on collagen and
elastin synthesis, the degree of proline hydroxylation in collagen and
level of collagen cross-linking in skin, aorta, heart and tendon of
3-month rats in vivo and in skin and aorta of 3- and 12-month rats in
vitro has been done.

In vivo it was shown that in each of investigated organs the synthesis
intensity of both proteins and studied steps of collagen metabolism are
not the same. Achilles and tail tendons have the same collagen
concentration, but the mechanical loads which Achilles tendons are
exposed to are bigger. So the level of proline hydroxylation in collagen
and the degree of its cross-linking in this tendons are also higher. As
for the level of collagen metabolism it is higher in tail tendons which
speaks about the less structural and metabolic stability of its
supramolecular formations. In skin that is subjected to considerably
bigger mechanical loads in comparison to aorta collagen concentration
and the degree of its cross-linking are also higher. From the other side
the aorta wall ability to large backing deformations depends on high
enough elastin concentration in it. The distribution and the level of
collagen formation and structural stability in heart sections are back
proportional to the loads they have, because in heart the main
resistance to the load do myofibrillar elements.

So, all these facts are the main condition for the formation of
connective tissue visko-elastic properties, that are optimal to the
quantity and distributions of the mechanical loads these organs have.

In vitro for the first time it was established that mechanical load
leads to the increasing of elastin concentration in skin and aorta in
basis of which according changes in its synthesis lying. Also it was
determined the diapason of loads in which the intensification of
collagen and elastin synthesis occurs in animals of different ages. It
was shown that cells answer of skin and aorta connective tissue depends
on the rats age. In 3-month animals the maximums of collagen
concentration and synthesis are observed at higher mechanical loads than
in 12-month rats. Besides the degree of collagen cross-linking of aorta
and skin in younger rats with the increasing of load begins to
decreasing and than increase but dose not reach the original level; in
one-year rats this dependence has opposite character. Probably such
differences in cell answer on the mechanical load connect with the
increasing of the connective tissue extracellular matrix cruelty with
age. For the confirming of such proposition there was investigated the
influence on the mechanical load on collagen concentration and
cross-linking in skin at the presence of semicarbazid – the blockator of
cross-linking in collagen formations. In this case the cruelty of
extracellular matrix brings down. It was shown the decreasing of
collagen concentration.

It was suggested, that lowering of matrix cruelty leads to the breaking
in chain of mechanical load signal translation to the cell membrane.

For the first time it was obtained, that in noncalcium environment (with
the presence of EGTA) without mechanical load the increasing of collagen
synthesis occurs. At the presence of calcium (2,5mM) the decreasing of
collagen synthesis is occur. At the incubation with chlorpromasin which
is the anticalmodulin agent the increasing of collagen synthesis was
observed. It was supposed that the complex Ca/calmodulin takes place in
collagen metabolism.

Key words: collagen, elastin, synthesis, extracellular matrix,
visko-elastic properties, mechanical load.

I?aeienaii aei ae?oeo 27.08.99 ?. Cai. 527.

Oe?aae – 100 i?ei. Oiia. ae?oe. a?e. 1.1. Oi?iao 60o841

OIA “Ciaiea”, 310051, e. Oa?e?a, aoe. A?oaia, 32

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