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Взаємозв’язок енергетичного та конструктивного обміну у штамів стафілокока, які стійкі до антибіотиків: Автореф. дис… канд. біол. наук / С.М. Козиць

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CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue i?iaeaie. C?inoaiiy e?eueeino? caoai?thaaiue,
ao?ieia?/iei oaeoi?ii yeeo ? Staphylococcus aureus neeaaea? iaeio ?c
aeooaeueieo i?iaeai i?e?ia?ieia?? oa ee?i?/ii? iaaeeoeeie. Iniaeeai?
aino?ioe iaaoaa? oey i?iaeaia a ca`yceo ?c ?iciianthaeaeaiiyi
aioea?ioeei?acenoaioieo ooai?a, ui caaaea? o?oaeiiu?a i?e i?aeai??
e?ea?nueeeo cania?a aeey ai?ioueae ?c aeaiith ?ioaeoe??th. A inoaii?e
/an, ii?yae ?c aenieith no?ee?noth aei aioea?ioee?a, ye?
aeei?enoiaoaaeeny ?ai?oa, oaeaeei a?aeaoaa?oueny i?enoinoaaiiy ? aei
iiaeo i?aia?ao?a, ui ai?iaaaeaeothoueny a e?eoaaeueio i?aeoeeo.

Oe?iea iioe?aiiy ?acenoaioieo ooai?a noao?eieiea, ca aeoieith a?eueoino?
aaoi??a, iia,ycaia ?c aeoaea ieanoe/iei iai?iii ?a/iaei, ui aea?
iiaeeea?noue aeey iiyae iaoaaie?/ieo ?aaeoe?e, ye? eaaeaoue a iniia?
a?io?i?/ieo iaoai?ci?a ?acenoaioiino?. C ae?oaiai aieo, aaiaoe/i?
aeaoa?i?iaioe, ui eiaeothoue aioea?ioeeino?ee?noue a oiae? i?ioean?a
aaiaoe/ii? ?aeiia?iaoe?? aeoaea oaeaeei iioe?ththoueny a iiioeyoe?yo
noao?eieie?a.

Iaoai?cie no?eeino? i?e?ii?aai?ci?a aeey aaaaoueio aioea?ioee?a
?icoeo?iaai?. Oaiioei?/iee i?iya no?eeino? iiaea aooe iia,ycaiee ?c
neioacii oa?iaio?a, ui ?iaeoeaothoue aioea?ioee, ?c ii?ooaiiyi
i?iieeeeaino? iiaa?oiaaeo no?oeoo?, ui caiia?aa? noai?aiith
aaeoa??inoaoe/ieo eiioeaio?aoe?e i?aia?aoa ana?aaeei? ee?oeie,
iiaeeo?eaoe??th i?oai? ae?? aioea?ioeea oa aeieeiaiith iao?aeieo
iaoaaie?/ieo oeyo?a. E??i oiai, ye iieaco? aiae?c e?oa?aoo?ieo aeaieo,
?acenoaioi?noue aei ?yaeo aioea?ioee?a iiaea aooe iia,ycaia iaeii/anii
?c ?icaeoeii aeae?eueeio a?io?i?/ieo iaoai?ci?a no?eeino?.

Ana oea na?ae/eoue i?i oa, ui no?ee?noue aei aioea?ioee?a ? neeaaeiei
a?ieia?/iei yaeuai, yea iia,ycaii c iaeiei ?c iai?yie?a aaiethoe??
iaoiaaiieo aaeoa??e oa oi?ea?oueny anueiai iaoaaie?cio i?e?iaii?
ee?oeie. Oiio iaiao?aeii eiiieaenia ii??aiyeueia aea/aiiy ?aaeoe?e
aia?aaoe/iiai oa eiino?oeoeaiiai iai?i?a o /ooeeaiai oa no?eeeo aei
aioea?ioee?a ooai?a noao?eieiea c iaoith iiooeo caaaeueieo
caeiiii??iinoae oa oeyo?a ?icaeoeo a?io?i?/ieo iaoai?ci?a
?acenoaioiino?, iaoai?ci?a aaiaoe/ii? aeaoa?i?iaoe?? oaiioei?/iiai
i?iyao no?eeino?, a oaeiae iiaeeeaeo oeyo?a ia?aaea/? aaiaoe/ieo
aeaoa?i?iaio a iiioeyoe?? noao?eieie?a.

Iaoa ? caaaeaiiy aeine?aeaeaiue. A ca,yceo ?c aeeeaaeaiei, iaoith ?iaioe
aoa ii??aiyeueiee aiae?c ?ioaineaiino? ?aaeoe?e aia?aaoe/iiai oa
eiino?oeoeaiiai iai?i?a o /ooeeaiai ooaio noao?eieiea oa ?yaeo eiai
aa??aio?a, ye? no?ee? aei oao?aoeeee?io, oei?aioai?eieo, a?eo?ii?oeeio,
iai?oeee?io oa ooceaeeio.

Aeey aeeiiaiiy ?iaioe iaiao?aeii aoei ae??oeoe oae? caaea/?:

1. Aea/eoe iniaeeaino? ooieoe?iioaaiiy iniiaieo eaoaaie?/ieo oeeee?a
ooai?a noao?eieiea a ca,yceo c no?ee?noth aei aioea?ioee?a.

2. I?iaanoe ii??aiyeueia aea/aiiy ieenethaaeueieo i?ioean?a o /ooeeaiai
oa ?acenoaioieo ooai?a noao?eieiea.

3. Aeine?aeeoe iniiai? ?aaeoe?? a?ineioaco a?ee?a, ioeea?iiaeo eeneio oa
e?i?ae?a o no?eeeo aei aioea?ioee?a noao?eieie?a.

Iaoeiaa iiaecia ?iaioe. Aia?oa iieacaii, ui no?ee?noue aei ?yaeo ??cieo
ii iaoai?cio ae?? ia aaeoa??aeueio ee?oeio aioea?ioee?a o noao?eieiea
noi?iaiaeaeo?oueny caaaeueieie caeiiii??iinoyie ia?aaoaeiae
aia?aaoe/iiai oa eiino?oeoeaiiai iai?i?a.

Anoaiiaeaii ci?io ni?aa?aeiioaiiy iniiaieo eaoaaie?/ieo oeeee?a o
?acenoaioieo aa??aio?a noao?eieiea a ii??aiyii? ?c /ooeeaei ooaiii.
Iieacaii iineeaiiy ?ie? ae?eie?co ia oii? cieaeaiiy aeoeaiino?
iaioicioinoaoiiai oeeeeo a eaoaaie?ci? aoaeaaiae?a.

Aeyaeaii ciaioaiiy oaeaeeino? ieeneaiiy i??oaaoo oa aoeaoaoo no?eeeie
ooaiaie, ui na?ae/eoue i?i cieaeaiiy ieenethaaeueii? aeoeaiino? OeOE oa
a?eueo ?ioaineaiee a?aeoie ?ioa?iaae?ao?a oeeeeo ia a?ineioaoe/i?
i?ioeane.

No?ee?noue aei aioea?ioee?a, ye? aea/aeenue noi?iaiaeaeo?oueny
iineeaiiyi neioaco iaea?eueo aaaeeeaeo aeey ee?oeie
a?iiie?ia??a-ioeea?iiaeo eeneio, a?ee?a, e?i?ae?a.

I?aeoe/ia cia/aiiy ?iaioe. Io?eiai? ?acoeueoaoe iiaeooue aooe
aeei?enoai? aeey neeaaeaiiy a?eueo iiaii? eiiieaenii? oa?aeoa?enoeee
iaoai?ci?a no?eeino? aei aioea?ioee?a, ui a naith /a?ao cia/ii
iieaaoeoue i?aea?? e?ea?nueeeo cania?a aeey ai?ioueae c iaoiaaiieie
noao?eieieaie oa aeanoue iniiao aeey ?ic?iaee noai ?aoe?iiaeueii?
aioea?ioeeioa?ai?? caoai?thaaiue noao?eieieiai? ao?ieia??.

Ca,ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. I?aaenoaaeai?
aeine?aeaeaiiy-oea /anoeia ?iaioe, ui i?iaiaeeoueny ia eaoaae??
i?e?ia?ieia?? oa a??onieia?? Aei?i?iiao?ianueeiai aea?aeaaiiai
oi?aa?neoaoo a ?aieao aea?aeathaeaeaoii? oaie ?01-11-97 “Aeine?aeaeaiiy
iniaeeainoae iaoaaie?/ieo i?ioean?a o i?e?ii?aai?ci?a oa ?ic?iaea
oaoi?/ieo oiia ae?iaieoeoaa iiaeo a?ieia?/ii-aeoeaieo i?aia?ao?a”, yea
eii?aeeio?oueny ieaiii iaoeiai-aeine?aeieo oai I?i?noa?noaa ina?oe
Oe?a?ie.

Iniaenoee aianie caeiaoaa/a. ?iaioa aeeiiaia iniaenoi aaoi?ii i?ae
ea??aieoeoaii aeieo.a?ie.iaoe, i?ioani?a A.?.A?ii?eiaa. Iniaenoa o/anoue
aeena?oaioa iieyaaea o i?iaaaeaii? eaai?aoi?ieo aeine?aeaeaiue, aiae?c?,
ocaaaeueiaii? iaea?aeaieo aenia?eiaioaeueieo aeaieo, ni?anoaaeaii? ?o c
e?oa?aoo?ieie iaoa??aeaie. Aei??thaaiiy aeoeaiino? oa?iaio?a i?iaaaeaii
ni?eueii c eaiae.a?ie.iaoe,aeioe. A.A.Aaa?eethe oa
eaiae.a?ie.iaoe,aeioe. E.I.Aieiaeie. Aaoi? i?eiineoue ue?o aaey/i?noue
ni?a?ia?oieeai eaoaae?e a?io?i?? Ee?anueeiai iaoe?iiaeueiiai
oi?aa?neoaoo aeieo.a?ie.iaoe,i?io. A.I. Aieoeeoeueeiio oa
aeieo.a?ie.iaoe,aeioe. N.A.Oeaeiye ca aeiiiiiao a i?iaaaeaii?
aenia?eiaioo ii aeeth/aiith i?/aieo niieoe a ee?oeie noao?eieie?a.

Ai?iaaoe?y ?iaioe. Iaoa??aee aeena?oaoe?eii? ?iaioe aoee i?aaenoaaeai?
ia I I?aeia?iaei?e iaoeiai-i?aeoe/i?e eiioa?aioe?? /Aei?i?iiao?ianuee
1995/; II Oe?a?inueeiio ?aae?ia?ieia?/iiio c`?cae? /Aei?i?iiao?ianuee,
1995/; Eaenoiinueeiio neiiiceoi? c iieaeoey?ii? oa ee?oeiii? a?ieia??
/NOA,1996, 1997/; VII c,?cae? Oe?a?inueeiai a?io?i?/iiai oiaa?enoaa
/Ee?a,1997/; II Oe?a?inueeiio a?io?ce/iiio c,?cae? /Oa?e?a, 1998/; I
I?aeia?iaei?e iaoeiai-i?aeoe/i?e eiioa?aioe?? “Iaoea oa ina?oa”
/Aei?i?iiao?ianuee, 1998/, VII I?aeia?iaei?e eiioa?aioe?? “Aeueyin
O?ainac” (Aei?i?iiao?ianuee, 1998).

Ioae?eaoe??. Ii oai? aeena?oaoe?? iioae?eiaaii 8 iaoeiaeo ?ia?o.

no?oeoo?a oa ia’?i aeena?oaoe??. Aeena?oaoe?eia ?iaioa neeaaea?oueny
c anooio, iaeyaeo e?oa?aoo?e, aenia?eiaioaeueii? /anoeie, iaaiai?aiiy,
aeniiae?a ? nieneo oeeoiaaii? e?oa?aoo?e, yeee aeeth/a? 227
iaeiaioaaiue. ?iaioa aeeeaaeaia ia 140 noi??ieao iaoeiiieniiai oaenoo,
?ethno?iaaia 17 ?enoieaie oa 19 oaaeeoeyie.

CI?NO ?IAIOE

Iaeyae e?oa?aoo?e

Iaeyae e?oa?aoo?e neeaaea?oueny ?c 3 ?icae?e?a, a yeeo aena?oeai?
ieoaiiy eaoaaie?/iiai iai?io ? aeoioeyoe?? aia?a?? o noao?eieie?a,
ioa?aeoa?eciaai? iniaeeaino? a?ineioaco iniiaieo eean?a niieoe – a?ee?a,
ioeea?iiaeo eeneio, e?i?ae?a oa aoaeaaiae?a, a oaeiae i?eaaaeai? iniiai?
a?io?i?/i? iaoai?cie no?eeino? aei ?yaeo iaea?eueo aaaeeeaeo
aioea?ioee?a. Iieacaii, ui i?ioeane iaoaaie?cio o noao?eieie?a aea/ai?
iaaeinoaoiuei ? oiio iaiao?aeia iiaeaeueoa a?eueo aeaoaeueia aea/aiiy ye
ie?aieo aia?aaoe/ieo oa a?ineioaoe/ieo ?aaeoe?e, oae ? iaoaaie?cio a
oe?eiio, ui ni?eyoeia a?eueo aoaeoeai?e ai?iouea? ?c aeaiith
?ioaeoe??th.

iaoa??aee ? iaoiaee aeine?aeaeaiue

Ia,?eoii aeine?aeaeaiue aoee ooaie Staphylococcus aureus: 8325-4 oa eiai
aa??aioe, ye? i?noyoue ieaci?aee ?acenoaioiino? aei ??cieo aioea?ioee?a.
Ooaie aoee i?aaeinoaaeaii c eieaeoei? iinoeoooo aiiaeaiiieiai? oa
iie?iaiieiai? ii. Ae.E.Aaiaea? AII ?ini?.

Aeey iaea?aeaiiy noniaici? “nii/eaath/eo” eeioei i?iaiaeeee iaeiie/aiiy
aiiiane ooai?a noaoieieiea ia oaa?aeiio aeeaeeueiiio na?aaeiaeui, ui
iinoeoue IIA, ?I 7,0-7,2 i?e t =37i N aei na?aaeeie eiaa?eoii/ii? oace
?inoo. Aeey iaea?aeaiiy aaceeioeiiiai aiiiaaiaoo ?oeioaaiiy eeioei
noaoieieiea i?iaiaeeee ia oeueo?acaoeiaiio aeaceioaa?aoi?i OCAeI-1 c
aeniiiaioeiaeueiei aei?iiiithaa/ai. Icao/oaaiiy i?iaiaeeee ia i?ioyci 10
oaeeei i?a?eaenoi c iioa?aaeii a 30 nae. Iac?oeiiaai? eeioeie oa
eeioeiii noiiee inaaeaeoaaee oeaio?eooaoaaiiyi i?e 20 000 ia/oa. ia
i?ioyci 60 oa.

I?e ae?iuoaaiii ooai?a noaoieieiea a aaceeniaaiio na?aaeiaeui aiaa?iaii
oiiae noai?thaaeenue a aiaa?inoaoi i?e caiiii eiai aoiinoa?e ia
aaciiiaeiaiee acio.

Aeoeaiinoi eaoaaie?/ieo oa?iaioia aeieieico, iaioicioinoaoiiai oeyoo oa
oeeeeo o?eea?aiiiaeo eeneio aecia/aee a?aeiia?aeieie caaaeueiii?eeiyoeie
iaoiaeaie Aeoeai?noue aaenie?iace, eaeoaoaeaa?ae?iaaiace oa
ooia?aoaiae?aoace aecia/aee ca iaoiaeaie i?eaaaeaieie o Ei/aoiaa /1980/,
6-oinoio?oeoie?iace, aeueaeieace o?oeoici-1-6-aeeoinoaoo, i??oaaoe?iace,
IAAeI-aeaaiae?iaaiace, icioeeo?aoaeaaiae?iaaiace – ca iaoiaeaie
i?eaaaeaieie a Methods in enzymol./1982/,
aetheici-6-oinoaoaeaaiae?iaaiace oa 6-oinoiaetheiiaoaeaaiae?iaaiace ca
iaoiaeii Kornberg, Horecker /1955/, o?aineaoieace – ca iaoiaeii Dische a
iiaeeoieaoei? Brownstone, Denstedt /1961/, oeeo?aoneioace – ca iaoiaeii
Muller-Kraft, Babel /1983/, iaeaoaeaaiae?iaaiace – ca iaoiaeii E? E?i
Aaia /1970/.

Aeaii niaeo?ioioiiao?e/ieo aeii?thaaiue iaea?aeoaaee ia “Specord uv vis”
(I?ia//eia) i?e 300 N. Aeoeaiinoue oa?iaioia ae?aaeaee a iiieue
ia?aoai?aiiai noano?aoo ca 1 oaeeeio ia 1 ia aieea.

Iioaineaiinoue ieeneaiiy noano?aoia aeeoaeueiei eaioethaii aea/aeinue
iiey?ia?aoi/iei iaoiaeii ia iiey?ia?aoi LP-7 (*NN?), c aeei?enoaiiyi
cae?eoiai aeaeo?iaea oeia Eea?ea.

Aaia?aoe?th o?ainiaia?aiii? ??cieoe? aeaeo?e/ieo iioaioe?ae?a (( ia
oe?eeo ee?oeiao noao?eieiea aea/aee ca iaoiaeii, yeee ?ic?iaeaiee
Liberman, Skulachev/1970/ ?c canoinoaaiiyi neioaoe/ieo e?iio?eueieo
i?iieeath/eo ?ii?a oao?aoai?eoinoii?th (O??+). E??i oiai, aaia?aoe?th ((
aei??thaaee ii ?ioaineaiino? oeoi?anoeaioe??
8-ai?e?iiiaooae?i-1-noeueoiiaoo (AIN-).

Aecia/aiiy iaeiie/aiiy eeth/iaeo iaoaaieioia eaoaaieicio aetheice:
iiei/ii? eeneioe, ioeoiai? eeneioe, ii?iaeiia?aaeii? eeneioe i?iaiaeeee
ca iaoiaeaie i?eaaaeaieie o Anao?ai? /1961/. Eiioeaio?aoe?th NI2
aecia/aee iaiiiao?e/iei iaoiaeii a aia?ao? Aa?ao?aa. Aecia/aiiy
iaeiie/aiiy iaoaaieioia a aiaa?iaieo oiiaao i?iaiaeeee i?e iieoaoaaiii a
i?iai?eao Ooiaa?aa (aaeooi -4 ii ?o. noiaioey).

A aenia?eiaioao ii aei? iiaiaioi?a NaF ia iioaineaiinoue aeieieico
aeacaiee iiaiaioi? aiineee a iieoaaoeieia na?aaeiaeua a eiioeaio?aoei?
10-3 I.

Aecia/aiiy aeoeaiinoi a?ineioaoe/ieo oa?iaio?a ania?oao- oa
aeaiiiaiiiio?ainoa?ac i?iaiaeeee ca iaoiaeii Oaieaethi oa ?i. /1986/,
aoeaoee-NiA-ea?aieneeace – ca iaoiaeii Olsen and Merrik /1968/.

A aeine?aeao ii aeeth/aiith ii/aieo iiia?aaeieeia aieea oa ioeea?iiaeo
eeneio ee?oeie noao?eieie?a ae?iuoaaee aei ?aiiuei? aeniiiaioe?aeueii?
oace ia IIA i?e 370N. I?ney aecia/aiiy eiioeaio?aoe?? a?eea a ee?oeiao a
aeine?aei? i?ia??ee aiineee noniaic?th, ui i?noeea 1-2 ia a?eea.

Aeey oa?aeoa?enoeee neioaco a?eea aiineee: DL- [4- 14N ] ania?aa?iiao
eeneioo, DL –[iaoee-14 N] – iao?ii?i, [14 N] – na?ei, [14N] –
?cieaeoeei, [2,3-3I] -o?eioioai. Aeey aeine?aeaeaiiy neioaco ioeea?iiaeo
eeneio aiineee [8- 14N] – aaeai?i, [2-14N] – o?aoeee, [5-iao?e-3I] –
oei?i. Eiio?ieai neoaeeee noniaic?? ee?oei noao?eieiea, ui aoee
?iaeoeaiaai? i?ia??aaiiyi ia i?ioyc? 10 oa.

O?aeoe?iioaaiiy a?iiane i?iaiaeeee ca iaoiaeii Park, Hancock /1960/.
?aaeiiaeoeaiinoue aecia/aee ca aeiiiiiaith noeeioeeeyoeieiiai
ei/eeueieea oi?ie “ Intertechnigue” (O?aioeiy). ?ioaineai?noue
aeeth/aiiy i?/aieo iiia?aaeiee?a ioe?ithaaee a ?iioeuenao ca 1 oa ia 1
ia a?eea.

Aecia/aiiy aieea a aaceeioeiiiio aiiiaaiaoi i?iaiaeeee ca iaoiaeii Lowry
et al. /1951/. Noaoenoe/io ia?iaeo aenia?eiaioaeueieo aeaieo i?iaiaeeee
ii Eae?io /1990/

?acoeueoaoe aeine?aeaeaiue oa ?o iaaiai?aiiy

1.Aeoeai?noue eaoaaie?/ieo oa?iaio?a o ooai?a noao?eieiea, ui
aea/aeenue.

A ?iaio? aoei i?iaaaeaii ii??aiyeueia aea/aiiy aeoeaiino? eeth/iaeo
oa?iaio?a iniiaieo eaoaaie?/ieo oeyo?a oa iaeiie/aiiy oeaio?aeueieo
iaoaaie?o?a o /ooeeaiai oa ?acenoaioieo aei ?yaeo aioea?ioee?a ooai?a
noao?eieiea.

Ia ia?oiio aoai? aoei i?iaaaeaii aea/aiiy iaeiie/aiiy e?ioeaaeo
i?iaeoeo?a ae?eie?co. Anoaiiaeaii, ui a?eueoa /anoeia aetheice, yea
ooee?co?oueny ii noai? Aiaaeaia-Iaea?aioa -Ia?iana, ia?aoai?th?oueny a
iiei/io eeneioo. Oaeaee?noue iaeiie/aiiy iiei/ii? eeneioe ciaoiaeeoueny
a iaaeao 0,210-0,270 ieiieue ia 1 ia a?eea ca 1 aiae.
Aioea?ioeei?acenoaioi? ooaie noao?eieie?a oa?aeoa?ecothoueny a?eueoeie
ia 14-29% oaeaeeinoyie iaeiie/aiiy eaeoaoo. Ii?yae c oeei, a?aei?/aii
iaeiie/aiiy a iaaaeeeeo e?eueeinoyo aoeaoaoo oa NI2 (oaae.1).

Oaaeeoey 1

Iaeiie/aiiy e?ioeaaeo i?iaeoeoia aeieieico oeieeie eeioeiaie ooai?a
noaoieieiea a aiaa?iaieo oiiaao.

Ooaie E?euee?noue, ie iieue ia 1 ia a?eea ca 1 aiae.

noao?eieiea Eaeoao Aoeaoao NI2

8325-4 0,210+0,030 0,011+0,002 0,010+0,001

NO-4 0,270+0,040 0,014+0,003 0,012+0,002

NN-4 0,250+0,050 0,013+0,003 0,011+0,001

Ii??aiyeueia aea/aiiy iaeiie/aiiy eaeoaoo a aa?iaieo oa aiaa?iaieo
oiiaao o ooai?a noao?eieiea, ui aea/aeenue, aeaco? ia iacia/io ??cieoeth
a oaeaeeino? ae?eie?co a aeaieo oiiaao. Aoaeo Ianoa?a, ?ic?aoiaaiee ii
oi?ioe? A Eeaeioeaeea?a /1965/, aeinoeaaa 25,2-28,0% (oaae.2).

Oaaeeoey2

.Iaeiie/aiiy eaeoaoo oeieeie eeioeiaie ooai?a noaoieieiea a aa?iaieo oa
aiaa?iaieo oiiaao.

Ooaie E?euee?noue eaeoaoo, ieiieue ia 1ia a?eea ca 1 aiae Aoaeo

noao?eieiea aa?iai? oiiae aiaa?iai? oiiae Ianoa?a,

%

8325-4 0,157+0,021 0,210+0,030 25,2

NO-4 0,197+0,025 0,270+0,040 27,0

NN-4 0,180+0,035 0,250+0,050 28,0

Aeey c’ynoaaiiy ?aaoeyoi?ieo iaoai?ci?a ae?eie?co o noao?eieiea aoei
aea/aii iaeiie/aiiy eaeoaoo i?e aianaii? ?yaeo ?ioa?iaae?ao?a ia oii?
i?enooiino? a na?aaeiaeu? aetheice. Oaeaee?noue ae?eie?co c?inoa? i?e
aeiaeaaaii? aetheici-6-oinoaoo, o?oeoici-1,6-aeeoinoaoo oa i??oaaoo.
Iaie aoee c?iaeai? i?eiouaiiy, ui o noao?eieiea e?i?ooaaiiy ae?eie?co
iiaea a?aeaoaaoeny ia ??ai? aaenie?iace, oinoio?oeoie?iace,
i??oaaoe?iace. Aeieacii oeueiio iineoaeeee aecia/ai? iaie iecuee?
aeoeaiino? aeacaieo o?ueio oa?iaio?a a ii??aiyii? ?c aeueaeieacith
o?oeoici-1,6-aeeoinoaoo, yea ia a?aeiineoueny aei oa?iaio?a, ui
?aaoeththoue oaeaee?noue ae?eie?co.

A ?icaeoie io?eiaieo ?acoeueoao?a aoei i?iaaaeaii ioe?ieo ?ie? ae?eie?co
a caaaeueiiio eaoaaie?ci? aetheice ii aeoeaiino? aaenie?iace oa
eaeoaoaeaa?ae?iaaiace, a oaeiae ii ??aith iaeiie/aiiy i??iaeiia?aaeii?
eeneioe oa aieeao ia oeae i?ioean ?ia?a?oi?a ooi?enoiai iao??th o
aeine?aeaeo?ieo ooai?a noao?eieiea.

Aeoeai?noue aaenie?iace oa eaeoaoaeaa?ae?iaaiace o ooai?a, ye? ianooue
ieaci?aee ?acenoaioiino? aeua, i?ae o aeo?aeiiai ooaio ia 15-69%.
??aaiue aeoeaiino? a aiaa?iaieo oiiaao aeuee i?ae a aa?iaieo aeey an?o
ooai?a (oaae.3).

Aioea?ioeei?acenoaioi? ooaie oa?aeoa?ecothoueny ? a?eueoeie aaee/eiaie
iaeiie/aiiy i??oaaoo ia 29-57% (?en.1). NaF ?ia?ao? iaeiie/aiiy i??oaaoo
noao?eieieaie a aa?iaieo oiiaao ia 11-62%, a aiaa?iaieo oiiaao ia
35-61%. I?ioeaio ?ia?aoaaiiy aeey ?acenoaioieo aa??aio?a cia/ii aeuee a
ii??aiyii? ?c /ooeeaei ooaiii, ui na?ae/eoue i?i ia?aaaao ae?eie?oe/iiai
oeyoo a eaoaaie?ci? aoaeaaiae?a o ?acenoaioieo ooai?a.

Aeniea iioaineaiinoue aa?iaiiai aeieieico o noaoieieieia, ui oaeiae
aeyaeaia a ?yaei aeineiaeaeaiue (Aa?oaiiae/ /1962,1964/), iiaeeeai,
iiynith?oueny oei, ui noaoieieiee – oaeoeueoaoeaii aiaa?iaii
iie?ii?aaiicie.

Oaaeeoey 3

Aeoeaiinoue oa?iaio?a ae?eie?co o ooaiia noao?eieiea, /ooeeaiai oa

noieeeo aei aioeaiioeeia.(iiieue ca1oa. ia 1ia aieea).

Ooaie

noaoieieiea Aaenie?iaca Eaeoaoaeaa?ae?iaaiaca

Aa?iaii oiiae Aiaa?iaii oiiae Aa?iaii oiiae Aiaa?iai? oiiae

8325-4 15,0(1,2 16,8(0,4 ?(0,05 30,2(3,1 40,4(2,5 ?(0,05

NO – 1 13,1(0,3 ?(0,05 12,9(0,8 ?(0,05 48,2(5,2 ?(0,05 51,3±4,5
?(0,05

NO – 4 17,9(0,6 ?(0,05 19,6(0,7 ?(0,05 39,4±3,4 ?(0,05 50,3(4,2 ?(0,05

NN – 1 17,3(0,9 ?(0,05 19,4(0,6 ?(0,05 34,7(2,3 ?(0,05 49,3(4,1 ?(0,05

NN – 4 15,8(0,4 ?(0,05 18,1(0,7 ?(0,05 43,2(1,6 ?(0,05 46,4(2,5 ?(0,05

994 18,1(0,8 ?(0,05 17,4(0,9 ?(0,05 47,1(3,2 ?(0,05 52,4(2,7 ?(0,05

NA – 1 17,1(0,4 ?(0,05 20,0(0,6 ?(0,05 24,6(3,7 ?(0,05 24,3(3,4 ?(0,05

NF – 8 18,9(0,4 ?(0,05 20,3(0,4 ?(0,05 51,1(4,4 ?(0,05 64,3(6,2 ?(0,05

NA – 5 17,6(0,7 ?(0,05 19,5(0,6 ?(0,05 42,8(3,4 ?(0,05 54,3(2,7 ?(0,05

NF – 2 18,0(0,6 ?(0,05 19,3(0,5 ?(0,05 39,6(4,8 ?(0,05 44,3(5,2 ?(0,05

Aeueoa?iaoeaiei ae?eie?co ? iaioicioinoaoiee oeyo, yeee iinoaaey?
?ioa?iaae?aoe aeey i?ioean?a a?ineioaco. I?e aiae?c? ?acoeueoao?a ii
aeoeaiino? oa?iaio?a iaioicioinoaoiiai oeeeeo iaie aoei anoaiiaeaii, ui
aeoeai?noue oa?iaio?a ieenethaaeueiiai aoaio aeaa?ae?iaaiac
aetheici-6-oinoaoo oa 6-oinoiaetheiiaoo cieaeaia o ?acenoaioieo aei
aioea?ioee?a aa??aio?a a na?aaeiueiio ia 10-38 % a ii??aiyii? ?c
/ooeeaei ooaiii (?en2).

Aeoeai?noue oa?iaioa a?ineioaoe/ii? /anoeie -o?aineaoieace oaeiae ieae/a
o no?eeeo aa??aio?a ia 14-48%. Aeey an?o ooai?a oa?aeoa?ii iacia/ia
ca?eueoaiiy aeoeaiino? oa?iaio?a iaioicioinoaoiiai oeeeeo a aa?iaieo
oiiaao.

Oaeei /eiii, aeey ?acenoaioieo aei ??cieo aioea?ioee?a aa??aio?a
oa?aeoa?ia ci?ia ni?aa?aeiioaiiy iniiaieo eaoaaie?/ieo i?ioean?a:
i?aeaeuaiiy ae?eie?oe/iiai oeyoo ia oii? cieaeaiiy ?ioaineaiino?
iaioicioinoaoiiai oeeeeo.

3.Iniaeeaino? i?ioean?a ieeneaiiy o no?eeeo aei aioea?ioee?a

noao?eieie?a

A ?acoeueoao? ii??aiyeueiiai aeine?aeaeaiiy /ooeeaiai oa ?acenoaioieo
aa??aio?a ii ieeneaiith ?yaeo noano?ao?a iaie anoaiiaeaii, ui
noao?eieiee aeoeaii ieeneththoue aetheico, i??oaao oa aoeaoao (oaae.4).
Niiaeeaaiiy eenith i?e ieeneaii? aetheice ciaoiaeeoueny a iaaeao
47,1-83,6 iiieue ca 1oa. ia 1ia a?eea. No?ee? aei aioea?ioee?a aa??aioe
oa?aeoa?ecothoueny a?eueoeie ia 25-77% aaee/eiaie ieeneaiiy aetheice.
Aea aaea i?e ieeneaii? i??oaaoo ??cieoey a oaeaeeino? ieeneaiiy i?ae
aeo?aeiei ooaiii oa ?acenoaioieie aa??aioaie cia/ii ciaioo?oueny. A
ii??aiyeueiee aiae?c oaeaeeino? ieeneaiiy aoeaoaoo aeaco? ia cieaeaiiy
aeoeaiino? ieenethaaeueieo oa?iaio?a OeOE o ?acenoaioieo aa??aio?a oa
a?eueo ?ioaineaia oieinoa ieeneaiiy.

Oaaeeoey 4

Ieenethaaeueia aeoeai?noue oe?eeo ee?oei ooai?a
noao?eieiea

Ooaie Oaeaee?noue iiaeeiaiiy eenith iiieue I2 oa-1. ia-1 a?eea

noao?eieiea Aetheica I??oaao Aoeaoao

8325-4 47,1 ( 3,0 31,1 ( 3,4 15,2 ( 2,7

NT-1 75,8 (6,3 ?(0,05 25,3 ( 4,3 ?(0,05 12,4 ( 2,5 ?(0,05

NC-1 83,6 ( 7,1 ?(0,05 27,5 ( 5,6 ?(0,05 12,5 ( 3,5 ?(0,05

994 71,5 ( 6,8 ?(0,05 21,8 ( 3,7 ?(0,05 11,1 ( 2,8 ?(0,05

NF-8 59,0 ( 4,0 ?(0,05 20,8 ( 3,1 ?(0,05 9,9 ( 1,9 ?(0,05

NF-2 69,3 ( 5,7 ?(0,05 20,3 ( 4,6 ?(0,05 9,8 ( 1,5 ?(0,05

?acoeueoaoe aeine?ae?a ii ieeneaiith ?yaeo ai?iieeneio ooaiaie
noao?eieiea iieacaee iaio ?ioaineaia ieeneaiiy a ii??aiyii? c
aetheicith.

Oaaeeoey 5

?ioaineai?noue ieeneaiiy ?yaeo ai?iieeneio oe?eeie ee?oeiaie ooai?a
noao?eieiea.

Ooaie Oaeaee?noue ieeneaiiy, iiieue I2 oa-1. ia a?eea-1

noao?eieiea Ania?aa?iiaa eeneioa Iao?ii?i ?cieaeoeei O?eioioai Na?ei

8325-4 14,0( 3,6 12,3(3,5 11,4(3,0 10,5(3,1 15,3(3,5

NT-1 11,8( 2,7 10,9(2,2 10,7(3,2 9,6(2,0 12,5(2,9

NC-1 11,5( 2,4 11,0(2,9 10,9(2,8 9,5(2,1 12,0(1,8

994 11,0( 2,0 10,5(1,7 10,0(3,1 9,3(1,9 11,9(2,4

NF-8 10,2 (2,8 10,4(2,1 9,0(2,0 9,0(1,5 10,1(2,5

NF-2 10,4( 2,2 10,1(1,8 9,5(1,9 9,2(2,3 10,2(2,3

Aaee/eie ieeneaiiy a na?aaeiueiio neeaee 9-15 iiieue I2 ca oa. ia ia
a?eea (oaae.5). Aeacoth/e ia iaaaeeeo ??cieoeth i?ae ?acenoaioieie
aa??aioaie oa /ooeeaei ooaiii, iiaeia a?aei?oeoe oeaio?aeueio
oaiaeaioe?th aei cieaeaiiy ?ioaineaiino? ieeneaiiy ai?iieeneio o no?eeeo
ooai?a, ui i?aeoaa?aeaeo? ?ai?oa c?iaeaiee aeniiaie, i?i caaaeueia
cieaeaiiy ieenethaaeueieo oa?iaio?a o aeaieo ooai?a.

C iaoith c’ynoaaiiy, ye a?aeaoaa?oueny ieeneaiiy ai?iieeneio o ooai?a
noao?eieiea /a?ac aeeoaeueiee eaioetha, /e oea iia,ycaii c oieinoei
ieeneaiiyi, iaie i?iaaaeaii aecia/aiiy aaia?aoe?? o?ainiaia?aiii?
??cieoe? aeaeo?e/ieo iioaioe?ae?a ia nii?yaath/?e iaia?ai? noao?eieie?a
i?e aeei?enoaii? a yeino? noano?aoa – ai?iieeneio. Iaea?aeai?
?acoeueoaoe na?ae/aoue i?i iinooiaiiy a?aeiiaethth/eo aea?aaeaio?a a
aeeoaeueiee eaioetha i?e ieeneaii? ai?iieeneio.

I?i cieaeaiiy ieenethaaeueii? aeoeaiino? o ?acenoaioieo ooai?a
noao?eieie?a iiaeia noaa?aeaeoaaoe ? ia iniia? ciaioaiiy aeoeaiino?
eeth/iaiai oa?iaioo OeOE -oeeo?aoneioace ia 20-37% a ii??aiyii? c
/ooeeaei aa??aioii. Aea, a oie aea /an, i?e aecia/aii? aeoeaiino?
oa?iaio?a, iia’ycaieo c i?ioeanaie a?ineioaco – cai?ioiuei?
iaeaoaeaa?ae?iaaiace oa ooia?aoa?ae?aoace ie aeyaeee ca?eueoaiiy
aeoeaiino? oa?iaio?a o no?eeeo ooai?a a na?aaeiueiio ia 15-86% (?en3).

A aiaa?iaieo oiiaao aeoeai?noue oa?iaio?a OeOE i?iyaey?oueny, aea
cieaeaia a na?aaeiueiio ia 10-37%.

I?eaaaeai? aeai? na?ae/aoue i?i cieaeaiiy ieenethaaeueii? aeoeaiino?, o
?acenoaioieo aa??aio?a oa i?i iai?aaeai?noue ?ioa?iaae?ao?a OeOE ia
a?ineioaoe/i? i?ioeane.

4.A?ineioaoe/i? i?ioeane noao?eieie?a

A?aeiii, ui aia?aaoe/iee iai?i o?nii aca?iiiia,ycaiee c a?ineioaoe/ieie
?aaeoe?yie. Iaie aoee aecia/ai? aeoeaiino? oa?iaio?a neioaco
ai?iieeneio- ania?oao- oa aeai?iai?iio?ainoa?ac o eieaeoe?? ooai?a
noao?eieiea (oaae.6). Anoaiiaeaii, ui ?aaeoe?y ia?aai?ioaaiiy

Oaaeeoey 6

Aeoeaiinoue ania?oao- oa aeai?iaiiiio?ainoa?ac o ooaiia noaoieieiea,

yei noieei aei aioeaiioeeia (iiieue noano?aoo ca 1oa. ia 1 ia aieea
)

Ooaie noaoieieiea Ania?oaoai?iio?ainoa?aca Aeai?iai?iio?ainoa?aca

Aa?iaii oiiae Aiaa?iaii oiiae Aa?iaii oiiae Aiaa?iaii oiiae

8325-4 23,4(2,4 29,5( 1,3 ?(0,05 10,6(0,7 11,9(0,3 ?(0,05

NO – 1 30,4(1,6 ?(0,05 34,4( 0,9 ?(0,05 12,5(0,4 ?(0,05 14,0( 0,7 ?(0,05

NO – 4 36,0(2,1 ?(0,05 41,9(0,7 ?(0,05 16,0(1,2 ?(0,05 17,0 ( 0,9 ?(0,05

NN – 1 28,0(0,6 ?(0,05 31,8, (1,3 ?(0,05 15,3(0,2 ?(0,05 14,6 ( 0,4
?(0,05

NN – 4 38,4(0,7 ?(0,05 39,3(1,8 ?(0,05 12,7(0,6 ?(0,05 14,9 ( 0,5 ?(0,05

994 39,8(2,3 ?(0,05 41,9 (0,3 ?(0,05 14,8(0,9 ?(0,05 16,7 ( 0,3 ?(0,05

NA – 1 36,5(0,5 ?(0,05 37,4(1,1 ?(0,05 18,0(0,7 ?(0,05 19,5 ( 0,4 ?(0,05

NF – 8 25,7(1,3 ?(0,05 30,4(1,6 ?(0,05 12,1(0,9 ?(0,05 12,7 (1,1 ?(0,05

NA – 5 34,4(0,4 ?(0,05 36,0(1,2 ?(0,05 14,4(0,2 ?(0,05 15,4 ( 0,6
?(0,05

NF – 2 30,4(2,0 ?(0,05 34,2(0,8 ?(0,05 17,8(0,5 ?(0,05 19,1 ( 0,3
?(0,05

ai?iieeneio a?aeaoaa?oueny ?ioaineai?oa o no?eeeo ooai?a ia 20-70%. A
aiaa?iaieo oiiaao aeoeai?noue oa?iaio?a aeua a ii??aiyii? ?c aa?iaieie
aeey an?o ooai?a, aeeth/ath/e aeo?aeiee, ui iiaeeeai iia,ycaii ?c
cieaeaiiyi ieenethaaeueieo i?ioean?a oa iineeaiiyi ?aaeoe?e
a?ineioaco.Aiaeia?/i? ?acoeueoaoe aoee iaea?aeai? ? ii aeeth/aiith
ai?iieeneio a ee?oeie noao?eieie?a (oaae.7). ?acenoaioi? aa??aioe
oa?aeoa?ecothoueny a?eueoei ia 14-31% aaee/eiaie aeeth/aiiy
ania?aa?iiai? eeneioe, iao?ii?io, ?cieaeoeeio, o?eioioaio oa na?eio. Oea
iiaea na?ae/eoe i?i iineeaiiy neioaco a?eea o ooai?a noao?eieie?a, ye?
?acenoaioi? aei aioea?ioee?a.

Oaaeeoey 7

Aeeth/aiiy ii/aieo aiiiieeneio a ee?oeie /ooeeaiai oa
noieeeo ooaiia noaoieieiea (?ii.ca oa-1.ia ia a?eea-1)

Ooaie noaoieieiea Ania?aaiii-aa eeneioa O?eioi-oai Na?ei Iaoiiiii
?cieaeoeei

8325-4 8340(317 5910(400 7120(560 6810(340 7380(506

NT-1 7839(405 ?(0,05 6737(210 ?(0,05 6194(310 ?(0,05 7491(420 ?(0,05
9077(114 ?(0,05

NC-1 9924(470 ?

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