IAOe?IIAEUeIA AEAAeAI?ss IAOE OE?A?IE

?INOEOOO I?E?IA?IEIA?? ? A??ONIEIA??

?i. Ae. E. CAAIEIOIIAI

E?NOAIUe Ieaenaiae? A?eai?iae/

OAeE 579.841.42

Iniaeeaino? aca?iiae?? /enoeo ? ci?oaieo eoeueoo? iaoaiio?ioieo aaeoa??e

c oaa?aeeie iaoa??aeaie

03. 00. 07 — i?e?ia?ieia?y

AAOI?AOA?AO

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa a?ieia?/ieo iaoe

Ee?a – 1999

Aeena?oaoe??th ? ?oeiien

?iaioa aeeiiaia o a?aeae?e? i?e?ia?ieia?/ieo i?ioean?a ia oaa?aeeo

iiaa?oiyo ?inoeoooo i?e?ia?ieia?? ? a??onieia?? ?i. Ae. E. Caaieioiiai
IAI Oe?a?ie

Iaoeiaee ea??aiee: aeieoi?
a?ieia?/ieo iaoe

EO?AeEO ?aai Ee?eeiae/,

?inoeooo i?e?ia?ieia?? ? a??onieia??

IAI Oe?a?ie, caa?aeoaa/ a?aeae?eii

Io?oe?ei? iiiiaioe:

Iaeaoaiei TH??e ?iiaiiae/, ae. a. i., i?io., /e.-ei??. IAIO, caa?aeoaa/
a?aeae?eo a?ieia?? aaciieenethth/eo i?e?ii?aai?ci?a ?inoeoooo
i?e?ia?ieia?? ? a??onieia?? ?i. Ae. E. Caaieioiiai IAI Oe?a?ie

I?eianueea Aaeeia Ieeiea?aia, e. a. i., no. i. n. ?inoeoooo
a?ieiei?aeii? o?i?? IAI Oe?a?ie, i. Ee?a

I?ia?aeia onoaiiaa:

?inoeooo eiei?aeii? o?i?? ? o?i?? aiaee ?i. A. A. Aeoiainueeiai IAI
Oe?a?ie, i. Ee?a

Caoeno a?aeaoaeaoueny “ 17 “ ethoiai 1999 ?. i 10 aiae ia can?aeaii?

niaoe?ae?ciaaii? A/aii? ?aaee Ae 26. 233. 01 i?e ?inoeooo? i?e?ia?ieia??
? a??onieia?? ?i. Ae. E. Caaieioiiai IAI Oe?a?ie (252143, Ee?a – 143,
aoe. Caaieioiiai 154, ?inoeooo i?e?ia?ieia?? ? a??onieia?? IAI Oe?a?ie,
cae can?aeaiue).

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? ?inoeoooo

i?e?ia?ieia?? ? a??onieia?? IAI Oe?a?ie ca aae?anith: 252143, i. Ee?a,
aoe. Caaieioiiai, 154.

Aaoi?aoa?ao ?ic?neaiee “ 15 “ n?/iy 1999 ?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? A/aii? ?aaee

eaiaeeaeao a?ieia?/ieo iaoe E.
I. Io??o

Aeooaeuei?noue oaie. Iaoaiio?ioi? aaeoa??? ? aaaeeeaith eaieith
o?io?/iiai eaioethaa o ?yae? aeinenoai [Eo?aeeo e ae?., 1992; Eo?aeeo e
ae?., 1998a; ?iiaiianeay e ae?., 1991;] ? a?ae?a?athoue aeiaaeueio ?ieue
o oeeee? aoaeaoeth a a?inoa??. Oe? i?e?ii?aai?cie iathoue oi?eaeuei?
a?io?i?/i? ? o?c?ieia?/i? aeanoeaino?, ui aeicaieythoue ?i ooee?coaaoe
iaoai, a oaeiae ?iceeaaeaoe ?yae oiene/ieo niieoe: aeeaie [Strand et
al., 1988], a?iiaoe/i? nie?oe [Mountfort et al., 1990], o?ainoi?ioaaoe
aaeiaaiiio?aei? aoaeaaiaei?a [Fitch et al., 1996; Hanson et al., 1995;
Hashimoto et al., 1996; Okada et al., 1995], neioacoaaoe oe?ii?
iaoaaie?oe [Proenca, Collaco, 1987]. Oiio aea/aiiy aeaii? a?oie
i?e?ii?aai?ci?a ia? ia o?eueee ooiaeaiaioaeueia, aea e i?aeoe/ia
cia/aiiy, ine?eueee aiie iiaeooue noaoe iniiaith ?yaeo a?ioaoiieia?e a
aaeoc? ioi?iie aeiae?eey a?ae ??cieo caa?oaeiaiue [Ciaieia e ae?., 1993;
Eaaeu?iaa e ae?., 1994]. Ia?niaeoeaiei ? aeei?enoaiiy aeaieo aaeoa??e
aeey cieaeaiiy eiioeaio?aoe?? iaoaio o aoa?eueieo oaooao [Eo?aeeo e
ae?., 1980 a]. Oea aeicaiey? i?aeaeueoe aaciaeo i?iaaaeaiiy a??nueeeo
?ia?o [Iyeaiueeee, Eo?aeeo, 1991; Iyeaiueeee e ae?., 1992].

A i?e?iaeieo oiiaao a?eueo?noue i?e?ii?aai?ci?a ia?aaoaathoue o
cae??ieaiiio noai?, ui i?aeaeuo? ?o o?c?ieia?/io aeoeai?noue ? iaaea?
ee?oeiai iaai? ia?aaaae [Beveridge et al., 1997]. Oiio a
a?ioaoiieia?/ieo i?ioeanao oe?iei aeei?enoiaothoueny ?iiia?e?ciaai?
i?e?ii?aai?cie. ?o canoinoaaiiy aeicaiey? iaea?eueo iiaii ?aae?coaaoe
a?ioaoiieia?/iee iioaioe?ae i?e?iaieo iiioeyoe?e o aacia?a?aiiio ?aaeei?
eoeueoeaoaaiiy, iaaea? ?yae ia?aaaa a oi?aae?ii? i?ioeanii, a oaeiae
aeicaiey? ni?inoeoe aeae?eaiiy oe?eueiaiai i?iaeoeoo. Iaeiei c iaea?eueo
aoaeoeaieo ? aeaoaaeo niinia?a ?iiia?e?caoe?? ? cae??ieaiiy ee?oei ia
??cieo iin?yo oeyoii aaeaac?? [Ieeianeay, 1989;

Eiceye e ae?., 1991].

Ooieoe?iioaaiiy aaeoa??e, a oiio /ene?, iaoaiio?io?a a i?e?iaeieo
oiiaao aai a ooo/ieo nenoaiao iia’ycaia c ?o aca?iiae??th c
i?e?ii?aai?ciaie ?ioeo o?c?ieia?/ieo a?oi. I?i aieea inoaii?o ia
aaeaac?th iaoaiio?ioieo aaeoa??e oa eieii?caoe?th ieie oaa?aeeo
iaoa??ae?a a e?oa?aoo?? a?aeiiinoae iaia?. Io?eiaiiy aeaieo i?i
iniaeeaino? oaeiai ni?a?nioaaiiy, eiai aieea ia i?ioeane aaeaac??,
eieii?caoe?? oaa?aeeo iaoa??ae?a ? o?c?ieia?/io aeoeai?noue
iaoaiio?io?a, aacia?a/ii ? oe?eaaei ye c iaoeiai? oi/ee ci?o, oae ? aeey
canoinoaaiiy a i?aeoeoe?.

I?e ?ic?iaoe? a?ioaoiieia?e c aeei?enoaiiyi iaoaiieenethth/eo
i?e?ii?aai?ci?a, cie?aia aeey cieaeaiiy aeae?eaiiy iaoaio o aoa?eueieo
oaooao, iaiao?aeii a?aoiaoaaoe aieea ??cieo ciai?oi?o /eiiee?a ia
?ioaineai?noue eieii?caoe?? ieie oaa?aeeo iaoa??ae?a oa o?c?ieia?/io
aeoeai?noue oeeo i?e?ii?aai?ci?a. Aei oaeeo /eiiee?a, ianaiia?aae,
iaeaaeaoue aeinooii?noue iaoaio ? eenith oa aieia?noue aeiae?eey.
I?iaaaeaiiy aeine?aeaeaiue a oeueiio iai?yieo aeicaieeei a ?icoe?eoe
oyaeaiiy i?i ooieoe?iioaaiiy iaoaiio?ioieo aaeoa??e ia oaa?aeeo
iiaa?oiyo a i?e?iaeieo ? a ooo/ieo oiiaao, ? ae?iaeoe i?aeoiaee aeey ?o
aoaeoeaiiai aeei?enoaiiy.

Ca’ycie ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie, oaiaie. Aeine?aeaeaiiy,
aeeeaaeai? a aeena?oaoe??, i?iaiaeeeenue a 1993 – 1997 ??. ca?aeii
?iai/iai ieaio a?aeae?eo i?e?ia?ieia?/ieo i?ioean?a ia oaa?aeeo
iiaa?oiyo ii athaeaeaoi?e oai? ? 21 / 163 “Caeiiii??iino? ?iiia?e?caoe??
/enoeo oa ci?oaieo eoeueoo? i?e?ii?aai?ci?a ia oaa?aeeo iaoa??aeao oa
i?eioeeie oi?aae?iiy ?o o?c?ieia?/iith aeoeai?noth”.

Iaoa ? caaea/? aeine?aeaeaiiy. Iaoa aeena?oaoe?eii? ?iaioe iieyaaea o
anoaiiaeaii? caeiiii??iinoae aaeaac?? iaoaiio?ioieo aaeoa??e aei
oaa?aeeo iiaa?oiiue i?e ?o ooieoe?iioaaii? a ci?oaieo eoeueoo?ao c
i?e?ii?aai?ciaie ?ioeo o?c?ieia?/ieo a?oi oa eieii?caoe?? ieie oeeo
iaoa??ae?a.

A ?iaio? ae??ooaaeenue oae? caaea/?:

— aeine?aeeoe iniaeeaino? ooieoe?iioaaiiy ci?oaieo eoeueoo? iaoaiio?io?a
c i?e?ii?aai?ciaie ?ioeo o?c?ieia?/ieo a?oi;

— anoaiiaeoe caeiiii??iino? aaeaac?? iaoaiio?ioieo aaeoa??e a noi?oao
eoeueoo? c i?aaenoaaieeaie i?e?ii?aai?ci?a ?ioeo o?c?ieia?/ieo a?oi aei
a?ae?io?eueieo ? a?ae?ioiaieo iaoa??ae?a;

— aeine?aeeoe iniaeeaino? eieii?caoe?? oaa?aeeo iiaa?oiiue /enoeie ?
ci?oaieie eoeueoo?aie iaoaiio?io?a oa aecia/eoe aieea oiia
ooieoe?iioaaiiy i?e?ii?aai?ci?a ia oeae i?ioean.

Iaoeiaa iiaecia io?eiaieo ?acoeueoao?a. Ia caoeno aeiinyoueny ianooii?
iia? ?acoeueoaoe.

Ooieoe?iioaaiiy iaoaiio?ioieo aaeoa??e a ci?oaieo eoeueoo?ao c
i?aaenoaaieeaie ?iaeo Bacillus noi?iaiaeaeo?oueny c?inoaiiyi ?inoiai?
aeoeaiino? iaoaiio?io?a oa ?ioaineaiino? niiaeeaaiiy ieie iaoaio. A
ci?oaieo eoeueoo?ao iaoaiio?io?a c aaeoa??yie ?iaeo Pseudomonas aeai?
iieacieee cieaeothoueny.

Aecia/aeueieie oaeoi?aie aaeaac?? aeine?aeaeaieo i?e?ii?aai?ci?a aei
oaa?aeeo iaoa??ae?a ? a?ae?ioiai? aeanoeaino? ee?oei, ui /?oei iieacaii
ia i?eeeaae? iaoaiio?ioieo aaeoa??e, a oaeiae aeaeo?ie?iaoe/i?
aeanoeaino? i?e?ii?aai?ci?a, ye? a?ae?a?aaaee i?ia?aeio ?ieue a aaeaac??
aaeoa??e ?iaeo Bacillus. I?e?ii?aai?cie i?aeoe/ii a iaeiaeiaeo
e?eueeinoyo i?ee??iethaaeenue aei a?ae?io?eueieo (neei) oa a?ae?ioiaieo
(iie?noe?ie) iaoa??ae?a. Ia aaeaac?th ie?aieo aeae?a i?e?ii?aai?ci?a,
ui, ii?yae c iaoaiio?ioaie, aoiaeyoue aei neeaaeo noi?o? eoeueoo?, ye
i?aaeei, aieeaa? eiaeai c eiiiiiaio?a oeeo eoeueoo?.

Iaoaiio?ioi? aaeoa??? a /enoeo ? ci?oaieo eoeueoo?ao ?ioaineai?oa
eieii?cothoue a?ae?io?euei? iiaa?oi? ii??aiyii c a?ae?ioiaieie.
Iaea?eueo aeoeaii oeae i?ioean i?io?ea? a aeeaeeueiiio na?aaeiaeu? ia
a?aenoai? aei 15 ii a?ae a?aieoe? iiae?eo aaciiiae?aii? ? ??aeei? oace.
E??i oiai, aiie ooai?ththoue ia oaa?aeeo iiaa?oiyo i?e?ieieii??. A
i?enooiino? Bacillus megaterium ?ioaineai?noue eieii?caoe?? oaa?aeeo
iaoa??ae?a Methylococcus capsulatus nooo?ai ia ci?ithaaeanue, o?ioe
i?aeaeuoaaeanue aeey Methylocystis parvus, a a i?enooiino? Pseudomonas
putida o ci?oaieo eoeueoo?ao aiia oiia?eueithaaeanue.

I?e iacia/ieo eiioeaio?aoe?yo iaoaio (2 – 9 %) ? aenieeo, iiiaae 29 %
eenith o aacia?e oac? aeine?aeaeai? iaoaiio?ioi? aaeoa??? neaaei
eieii?cothoue ye a?ae?io?euei?, oae ? a?ae?ioiai? iaoa??aee. I?aeaeuaiiy
ai?noo ia?oiai eiiiiiaioo o a aacia?e noi?o? aei 17 – 29 %, ye ?
iayai?noue o i?e eenith a eiioeaio?aoe?yo 5 – 29 % i?ecaiaeeoue aei
?ioaineo?eaoe?? i?ioeano eieii?caoe?? oaa?aeeo iiaa?oiiue
aeine?aeaeaieie iaoaiio?ioaie.

O?c?ieia?/ia aeoeai?noue /enoeo oa ci?oaieo eoeueoo? iaoaiio?ioieo
aaeoa??e ia oaa?aeeo iiaa?oiyo aeinyaa? iaeneiaeueieo cia/aiue ia 2 — 8
aeiao ?o eieii?caoe?? oa i?e iayaiino? ia iin?? a?coaeueii aeaeeii?
ie?aeiai? aieiae. I?e i?aeneoaii? iin?y niiaeeaaiiy iaoaio
i?e?ii?aai?ciaie nooo?ai ciaioo?oueny.

I?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoao?a. ?ic?iaeaii eoeueoeaaoi?
aeey o?eaaeiai ae?iuoaaiiy /enoeo eoeueoo? iaoaiio?ioieo aaeoa??e i?e
aeei?enoaii? aaciai? noi?o? o iai?acaieiooiio eiioo??.

Anoaiiaeai? iniaeeaino? ooieoe?iioaaiiy iaoaiio?ioieo aaeoa??e a
ci?oaieo eoeueoo?ao c i?aaenoaaieeaie i?e?ii?aai?ci?a ?ioeo
o?c?ieia?/ieo a?oi. Oea aeicaiey? i?iaiicoaaoe iniaeeaino?
ooieoe?iioaaiiy aeaieo aaeoa??e ia oaa?aeeo iiaa?oiyo i?e ?o
a?ioaoiieia?/iiio canoinoaaii?. Aecia/ai? iioeiaeuei? ia?aiao?e ?yaeo
oaeoi??a iaaeieeoiueiai na?aaeiaeua, ui caaacia/othoue ?ioaineaio
eieii?caoe?th oaa?aeeo iaoa??ae?a iaoaiio?ioieie aaeoa??yie oa ?o aenieo
o?c?ieia?/io aeoeai?noue.

Io?eiai? ?acoeueoaoe aeine?aeaeaiue iiaeooue aooe aeei?enoai? o
a?ioaoiieia?yo, ui ?ic?iaeythoueny ia iniia? ?iiia?e?ciaaieo
iaoaiio?ioieo aaeoa??e ? iai?aaeai? ia io?eiaiiy oe?iieo iaoaaie?o?a,
ciaoeiaeaeaiiy oiene/ieo iio?aeieo iaoaio ? eiai aiiieia?a, cieaeaiiy
ai?noo iaoaio a aaciaeo na?aaeiaeuao, cie?aia o ae?iaeaieo i?inoi?ao
aoa?eueieo oaoo.

Iniaenoee aianie caeiaoaa/a. Aeine?aeaiiy o?c?ieia?/ii? aeoeaiino?
iaoaiio?io?a o ci?oaieo eoeueoo?ao c oeiiaeie i?aaenoaaieeaie i?e?ioei?e
aoa?eueieo oaoo, a?ae?ioiaieo aeanoeainoae i?e?ii?aai?ci?a, /anoeia
aeine?aeaeaiue ii aaeaac?? ? eieii?caoe?? /enoeie ? ci?oaieie
eoeueoo?aie aeeiiai? o ni?aaaoi?noa? c n. i. n., e. a. i. A. I. ?ie.
Aeaeo?iiiaa?oiaa? aeanoeaino? i?e?ii?aai?ci?a aecia/ai? noi?nii c n. i.
n., e. o. i. A. N. Ai?ae?aiei. ?aooa aeine?aeaeaiue aeeiiai? iniaenoi
aaoi?ii. Aaoi? aeneiaeth? iiaeyeo an?i, ooi aeiiiiaaaa o aeeiiaii? eiai
aeena?oaoe?eii? ?iaioe.

Ai?iaaoe?y ?acoeueoao?a aeena?oaoe??. Iaoa??aee aeena?oaoe??
aeiiia?aeaeenue ia I (VIII) Onoaiia/iio c’?cae? Oe?a?inueeiai
i?e?ia?ieia?/iiai oiaa?enoaa (Iaeana, 1993?.), ia ca?oieo
eiioa?aioe?yo-eiieo?nao iaoeiaeo ?ia?o ?inoeoooo i?e?ia?ieia?? ?
a??onieia?? IAI Oe?a?ie (1994, 1996 ??.).

Ioae?eaoe??. Ii oai? aeena?oaoe?? iioae?eiaai? 6 noaoae ? 1 oaca.

No?oeoo?a oa ia’?i aeena?oaoe??. Aeena?oaoe?y neeaaea?oueny c anooio,
iaeyaeo e?oa?aoo?e (2 ?icae?ee), aenia?eiaioaeueii? /anoeie c 4
?icae?e?a aeanieo aeine?aeaeaiue, caeeth/aiiy, aeniiae?a ? nieneo
aeei?enoaii? e?oa?aoo?e, ui i?noeoue 310 aeaea?ae. ?iaioa aeeeaaeaia ia
113 noi??ieao iaoeiiieno, ?ethno?iaaia 6 oaaeeoeyie, 22 ?enoieaie (a o.
/. 3 oioia?ao?yie).

CI?NO ?IAIOE

?icae?e 1. Iioe?aiiy oa o?c?ieia?/i? aeanoeaino? iaoaiio?ioieo

aaeoa??e

A ?icae?e? i?aaenoaaeai? e?oa?aoo?i? aeai? i?i ?iciianthaeaeaiiy
iaoaiio?ioieo aaeoa??e a i?e?iae?, ?o ii?oieiai-eoeueoo?aeuei?
aeanoeaino? oa o?io?/i? iio?aae.

?icae?e 2. Aeanoeaino? iiaa?oi? i?e?ii?aai?ci?a oa eieii?caoe?y

ieie oaa?aeeo iaoa??ae?a

?icaeyioo? iniaeeaino? aoaeiae ee?oeiii? iiaa?oi? i?e?ii?aai?ci?a c
iiceoe?e oi?ioaaiiy aeaeo?ie?iaoe/iiai ca?yaeo oa
a?ae?ioiaii-a?ae?io?eueieo aeanoeainoae. I?iaiae?ciaaii e?oa?aoo?i?
a?aeiiino? uiaei aieeao oeeo /eiiee?a ia i?ioean aaeaac??
i?e?ii?aai?ci?a aei iiaa?oiiue oaa?aeeo iaoa??ae?a. ?ice?eo? iniaeeaino?
eieii?caoe?? oaeeo iaoa??ae?a, ?? cia/aiiy c aeieia?/ii? oa i?aeoe/ii?
oi/ee ci?o. Iieacai? a?aei?iiino? ooieoe?iioaaiiy i?ae ?iiia?e?ciaaieie
oa a?eueiiaeeao/eie ee?oeiaie.

?icae?e 3. Iaoa??aee ? iaoiaee

Aeine?aeaeaiiy i?iaiaeeee ia aeaio aeaeao iaoaiio?ioieo aaeoa??e, ui
nooo?ai a?ae??ciyeeny ca aeanoeainoyie ee?oeiii? iiaa?oi? [Ai?aeeaiei e
ae?., 1997]: Methylococcus capsulatus ANA-874 ? Methylocystis parvus
OBBP; aeaio aeaeao nii?iooai?thth/eo aaeoa??e: i?ioaie?oe/ii aeoeaiiio
Bacillus megaterium ?IA-475 [Nie?iia e ae?., 1980] ? ooai? Bacillus
subtilis 9, ui oa?aeoa?ecoaaany iecueeith i?ioaie?oe/iith aeoeai?noth oa
iaeiiio ooai? ?iaeo Pseudomonas [?ie e ae?., 1990]. Eoeueoeaoaaiiy
iaoaiio?io?a cae?enithaaee o a?aeeiiii-aeieeaiiio eoeueoeaaoi??
[Eenoaiue, Eo?aeeo, 1998].

Aeaeo?ie?iaoe/iee iioaioe?ae aaeoa??e aecia/aee iaoiaeii
i?e?iaeaeo?ioi?aco [Aeiaa, Ai?aeeaiei, 1980].

A?ae?ioiai? aeanoeaino? i?e?ii?aai?ci?a ioe?ithaaee ca ?o ?iciiae?eii o
a?oaci?e nenoai? “noniaic?y i?e?ii?aai?ci?a — i-aaenaaeaeai( [Rosenberg
et al., 1980].

A aeine?aeaeaiiyo o?c?ieia?/ii? aeoeaiino? iaoaiio?io?a o ci?oaieo
eoeueoo?ao c oeiiaeie i?aaenoaaieeaie i?e?ioei?e aoa?eueieo oaoo
aeei?enoiaoaaee a?aeieo? a?ae iaoaaie?o?a noniaic?? /enoeo eoeueoo?
iaoaiio?io?a oa ?ioeo aeae?a aaeoa??e a ia’?iieo ni?aa?aeiioaiiyo 1 : 1,
10 : 1 ? 100 : 1. ?aeeiei aeaea?aeii aoaeaoeaaiai aeeaeaiiy oa aia?a??
aeey i?e?ii?aai?ci?a aoa iaoai.

O?c?ieia?/io aeoeai?noue iaoaiio?ioieo aaeoa??e ioe?ithaaee ii
niiaeeaaiith iaoaio ? eenith iaoiaeii aacio?iiaoia?ao?/iiai aiae?co
[Eo?aeeo e ae?., 1993], a oaeiae ii i?e??noo /enaeueiino? aaeoa??e o
noniaic??.

Aeey ?iiia?e?caoe?? i?e?ii?aai?ci?a aeei?enoiaoaaee /anoeiee neea /e
a??nueei? ii?iaee aoa?eueieo oaoo, aai neeyi? ? iie?noe?ieia? ieanoeie.

Aaeaac?th aaeoa??e aei iiaa?oi? aeaieo iaoa??ae?a ioe?ithaaee ca
??cieoeath ai?noo ee?oei o noniaic?? aei ? i?ney ?? aca?iiae?? c
oaa?aeei iaoa??aeii, aai ca e?euee?noth ee?oei, aaeaaciaaieo aei neeyieo
? iie?noe?ieiaeo ieanoei iaoiaeii i?yiiai i?ae?aooieo.

I?e aea/aii? i?ioeano eieii?caoe?? /enoeie ? ci?oaieie eoeueoo?aie
iaoaiio?ioieo aaeoa??e c ?ioeie i?e?ii?aai?ciaie c?acee c aaeaaciaaieie
ee?oeiaie ?ieoaoaaee a oiiaao, aeaeoeaieo aeey iaoaiio?io?a. Aieea
eiiiiiaioiiai neeaaeo aaciai? noi?o? ia i?ioean eieii?caoe?? oaa?aeeo
iaoa??ae?a iaoaiio?ioieie aaeoa??yie aeine?aeaeoaaee, aa??thth/e
ii/aoeiaith eiioeaio?aoe??th iaoaio i?e noae?e eiioeaio?aoe?? eenith,
aai iaaiaee.

Aieea aieiaino? ia niiaeeaaiiy iaoaio ?iiia?e?ciaaieie iaoaiio?ioieie
aaeoa??yie aea/aee, aeei?enoiaoth/e /anoeiee a??nueei? ii?iaee c ??ciith
ii/aoeiaith aieia?noth.

O?c?ieia?/io aeoeai?noue /enoeo ? ci?oaieo eoeueoo? iaoaiio?ioieo
aaeoa??e ? ?ioeo i?e?ii?aai?ci?a i?e eieii?caoe?? ieie oaa?aeeo
iiaa?oiiue aeine?aeaeoaaee ia ?ic?iaeai?e noaiaeia?e onoaiiaoe?,
ia??iaee/ii aecia/ath/e a eieiieao c i?e?ii?aai?ciaie ?ioaineai?noue
niiaeeaaiiy iaoaio ? eenith a i?ioean? o?eaaeiai aenia?eiaioo.

Noaoenoe/io ia?iaeo ?acoeueoao?a aeine?aeaeaiue cae?enithaaee
a?aeiia?aeieie iaoiaeaie aa??aoe?eii? noaoenoeee [Eaeei, 1968].

?icae?e 4. O?c?ieia?/ia aeoeai?noue iaoaiio?ioieo aaeoa??e o /enoeo ?

ci?oaieo eoeueoo?ao

Anoaiiaeai? aca?iia?aeiineie a ci?oaieo eoeueoo?ao, ui i?aaenoaaeai?
oeiiaith i?e?ioei?ith aoa?eueieo oaoo. Aei ?o neeaaeo, ii?yae c
iaoaiio?ioaie, aoiaeeee aaoeeee ? inaaaeiiiiaaee. Aea/aii aieea aeaio
ooai?a ?iaeo Bacillus — i?ioaie?oe/ii aeoeaiiai B. megaterium oa B.
subtilis, yeee oa?aeoa?ecoaaany neaaeeie e?oe/ieie aeanoeainoyie, ia
?inoiao aeoeai?noue ? niiaeeaaiiy iaoaio iaoaiio?ioaie. Iieacaii, ui
i?enooi?noue oeeo aaoa?io?io?a ni?e/eiyea noeioethth/ee aieea ia aeai?
iieacieee. A oaeeo oiiaao e?euee?noue aaoeee iaeaea ia ci?ithaaeanue.

?ioa ea?oeia niinoa??aaeanue o ci?oaieo eoeueoo?ao iaoaiio?io?a c

P. putida. A aeaiiio aeiaaeeo a?aeaoaaeinue cieaeaiiy ?inoiai?
aeoeaiino?

M. capsulatus ? M. parvus, a oaeiae ciaioaiiy e?eueeino? niiaeeoiai ieie
iaoaio. Iaea?aeai? ?acoeueoaoe aeicaieythoue i?iaiicoaaoe iniaeeaino?
ooieoe?iioaaiiy aeaieo aaeoa??e ia oaa?aeeo iiaa?oiyo i?e ?o
a?ioaoiieia?/iiio canoinoaaii?.

?icae?e 5. Aaeaaceai? aeanoeaino? /enoeo iaoaiio?ioieo eoeueoo? ? ?o

noi?oae c ?ioeie i?e?ii?aai?ciaie

Aea/ai? iniaeeaino? aaeaac?? /enoeo eoeueoo? aei a?ae?io?eueieo ?
a?ae?ioiaieo iaoa??ae?a. Iaea?eueoith caeaoi?noth aei aaeaac??
oa?aeoa?ecoaaany

M. parvus (?en. 1).

?en. 1. Caeaaei?noue aaeaac?? (a) i?e?ii?aai?ci?a aei /anoeiie neea
?ici??ii

1–2 ii a?ae oa?i?io ?o aca?iiae?? (t):

1 — M. parvus, 2 — M. capsulatus, 3 —

B. megaterium, 4 — P. putida, 5 —

B. subtilis. Oiiae aeine?aeo: aeenia?n?eia na?aaeiaeua — oinoaoiee aooa?
(0,15 M, pH – 7,0), t — 20 (C

I?e aca?iiae?? oeeo i?e?ii?aai?ci?a c iiaa?oiath neea ca 1 aiaeeio
i?ee??iethaaeinue 65 % ee?oei. A cia/ii iaioeo e?eueeinoyo
aaeaacoaaeenue aei aeaiiai iaoa??aeo M. capsulatus ? B. megaterium —
40,9 ? 35,0 % ca 1 aiaeeio a?aeiia?aeii.

Anoaiiaeaii, ui aeyaeai? a?aei?iiino? a aaeaac?? aeine?aeaeaieo
i?e?ii?aai?ci?a iaoiiaeai? ??cieoeath o aeanoeainoyo iiaa?oi? ?oi?o
ee?oei. sse a?aeiii [Eiceye e ae?., 1991; Busscher, Weerkamp, 1987;
Hermansson et al.,

1982 ], iaea?eueo ?noioii ia aaeaac?th aieeaathoue a?ae?ioiai? ?
aeaeo?iiiaa?oiaa? aeanoeaino? aca?iiae?th/eo iiaa?oiiue. Iniaeeai
iii?oiei a aaeaac?? aaeoa??e M. parvus aoa aianie a?ae?ioiaiino?
iiaa?oi? ?o ee?oei (oaae. 1). Ia aeeaey/enue ia aenieee iaaaoeaiee
ca?yae (– 28 iA), oe? i?e?ii?aai?cie caaaeyee ?o aenie?e a?ae?ioiaiino?
iaea?eueo ?ioaineaii i?ee??iethaaeeny aei oaa?aeiai iaoa??aeo. Aaeoa???
M. capsulatus oa?aeoa?ecoaaeenue a?ae?io?eueiith iiaa?oiath. Ia
caaaeath/e ia cia/ii iaioee iaaaoeaiee ca?yae (– 14 iA), oe?
i?e?ii?aai?cie i?ee??iethaaeenue aei neea a cia/ii iaioeo e?eueeinoyo.
Oae? ?acoeueoaoe na?ae/aoue i?i nooo?ao ?ieue oaeoi?o a?ae?ioiaiino? a
i?ioean? aaeaac?? iaoaiio?io?a aei oaa?aeeo iiaa?oiiue.

Oaaeeoey 1

Oa?aeoa?enoeea aeayeeo aeanoeainoae iiaa?oi? i?e?ii?aai?ci?a

Aeae Aeanoeaino? iiaa?oi?

aaeoa??e A?ae?ioiai?noue, % z-iioaioe?ae, iA

M. capsulatus 9,5 ( 1,0 –14

M. parvus 86,6 ( 2,3 –28

B. megaterium 6,2 ( 2,9 –15

B. subtilis 7,8 ( 1,1 –31

P. putida 3,0 ( 1,1 0

I?ei?oee: 1. aeenia?n?eia na?aaeiaeua oinoaoiee aooa? (0,15 M, pH 7,0)

2. “0”— i?e aeaieo oiiaao a?eueo?noue ee?oei ca?yaeo ia iathoue.

?io? aaeoa??? — B. megaterium, B. subtilis ? P. putida oaeiae
oa?aeoa?ecoaaeenue a?ae?io?eueiith iiaa?oiath. Ia i?eeeaae? oeeo
i?e?ii?aai?ci?a /?oei i?iaeyaeaeanue ?ieue aeaeo?ie?iaoe/iiai
iioaioe?aeo a i?ee??ieaii? ee?oei aei oaa?aeeo iiaa?oiiue: aaeoa???, ui
iaee iaioee ca?yae ee?oei, a a?eueoeo e?eueeinoyo aaeaacoaaeenue aei
oaa?aeeo iaoa??ae?a.

Oaeei /eiii, aeniea aaeaac?eia caeaoi?noue aeine?aeaeaieo
i?e?ii?aai?ci?a, ianaiia?aae, iaoiiaeaia a?ae?ioiai?noth iiaa?oi?, a i?e
iecueeeo cia/aiiyo oeueiai iieacieea i?ee??ieaiiy ee?oei aecia/a?oueny
aaee/eiith ?oiueiai ca?yaeo. A oiiaao aenia?eiaio?a a?ae?ioiai?noue
oaa?aeeo iaoa??ae?a i?aeoe/ii ia aieeaaea ia aaeaac?th ??cieo aeae?a
aaeoa??e: iaeaea an? aiie i?aeoe/ii a iaeiaeiaeo e?eueeinoyo
i?ee??iethaaeenue ye aei a?ae?ioiaieo, oae ? aei a?ae?io?eueieo
iaoa??ae?a.

Aea/ai? caeiiii??iino? aaeaac?? iaoaiio?ioieo aaeoa??e i?e ?o
ooieoe?iioaaii? o i?enooiino? ?ioeo i?e?ii?aai?ci?a (oaae. 2). Oae, ia
aeeaey/enue ia oa, ui o noi?oao eoeueoo? M. capsulatus ? B. megaterium
e?euee?noue iaoaiio?io?a caaeaaaee ia ii?yaeie a?eueoa, i?ae aaoeee,
/enaeuei?noue aaeaaciaaieo aei oaa?aeeo iaoa??ae?a ee?oei oeeo aeaio
aeae?a a?ae??ciyeanue iacia/ii. Iiae?aia ea?oeia niinoa??aaeanue ? a
noi?oao eoeueoo? M. capsulatus ? B. subtilis. A aeaiiio aeiaaeeo,
iiaeeeai, ia? i?noea iiaeeo?eaoe?y iiaa?oi? ee?oei aaoeee iaoaaie?oaie
i?e?ii?aai?ci?a, cie?aia, iaoaiio?ioa. Oea i?ecaiaeeoue aei ci?ie ??
aeanoeainoae oa ni?ey? i?aeaeuaiith aaeaac?? aaoeee.

Oaaeeoey 2

E?euee?noue iaoaiio?io?a (/enaeueiee, ee?oei / ii2) oa ?ioeo
i?e?ii?aai?ci?a (ciaiaiiee, ee?oei / ii2), aaeaaciaaieo aei oaa?aeeo
iaoa??ae?a ?c noi?oae

noniaic?e eoeueoo?

Eiiiiiaioe Ni?aa?ae- *enaeuei?noue i?e?ii?aai?ci?a

noi?oae

iioaiiy ia’?i?a aaeaaciaaieo aei iiaa?oi?,

ee?oei / ii2 a noniaic?? ia

ii/aoeo,

noniaic?e Neei Iie?noe?ie ee?oei / ie

M. capsulatus

+ 10 : 1

(2,22 ( 0,47) ( 103

(1,25 ( 0,21) ( 103

(7,70 ( 1,60) ( 102

(7,40 ( 1,40) ( 102

(2,73 ( 0,40) ( 108

(1,58 ( 0,19) ( 107

B. megaterium

100 : 1

(2,25 ( 0,52) ( 103

(2,35 ( 0,39) ( 102 (5,40 ( 1,30) ( 102

(2,20 ( 0,42) ( 102 (3,79 ( 0,57) ( 108

(7,90 ( 0,83) ( 105

M. capsulatus

+ 10 : 1

(3,22 ( 0,62) ( 103

(1,71 ( 0,33) ( 103

(5,62 ( 1,09) ( 102

(5,20 ( 1,03) ( 103

(2,27 ( 0,31) ( 108

(1,95 ( 0,21) ( 107

B. subtilis

100 : 1

(1,85 ( 0,38) ( 103

(3,98 ( 1,12) ( 102 (2,80 ( 0,83) ( 102

(4,22 ( 1,47) ( 103 (3,33 ( 0,46) ( 108

(3,34 ( 0,51) ( 106

M. capsulatus

+ 10 : 1

(3,95 ( 0,82) ( 103

(8,70 ( 1,40) ( 102

(2,84 ( 0,58) ( 103

(5,45 ( 1,05) ( 102

(1,52 ( 0,23) ( 108

(3,74 ( 0,30) ( 108

P. putida

100 : 1

(5,67 ( 1,18) ( 103

(5,20 ( 0,80) ( 102 (4,58 ( 0,97) ( 103

(4,40 ( 0,85) ( 102 (4,40 ( 0,65) ( 108

(8,90 ( 0,74) ( 107

M. parvus

+ 10 : 1

(4,72 ( 1,03) ( 104

(5,50 ( 0,90) ( 102

(4,24 ( 0,94) ( 104

(7,34 ( 1,10) ( 102

(8,80 ( 1,03) ( 107

(2,75 ( 0,29) ( 106

B. megaterium

100 : 1

(5,60 ( 1,12) ( 104

(55 ( 20) (6,03 ( 1,31) ( 104

(1,02 ( 0,14) ( 102 (9,68 ( 1,12) ( 107

(6,00 ( 0,93) ( 105

I?ei?oee: 1. Iioe/ia aonoeia noniaic?? M. parvus neeaaeaea 0,05 iae;
?ioeo i?e?ii?aai?ci?a — 0,8 iae.

2. Oiiae aeine?aeo: aeeaeeueia na?aaeiaeua aeey iaoaiio?io?a, t — 20( C,
3 aiae.

O noi?oao eoeueoo? M. capsulatus ? P. putida iaaiaee, /enaeuei?noue
ee?oei inoaiiuei? eoeueoo?e, i?ee??ieaieo aei neea ? iie?noe?ieo aoea
cia/ii ieae/ith o ii??aiyii? c oaeith aeey M. capsulatus. A oie aea /an,
inoaii?e e?aua aaeaacoaaany aei oeeo iaoa??ae?a, iniaeeai, iie?noe?ieo.
Ine?eueee a /eno?e eoeueoo?? M. capsulatus a a?eueoeo e?eueeinoyo
i?ee??iethaaany aei a?ae?io?eueii? iiaa?oi?, oaea yaeua iiaea aooe
aeeeeeaia ni?aoe??th iaoaaie?o?a inaaaeiiiiaaee ia iiaa?oi? iie?noe?ieo
oa ni?eyoe i?aeaeuaiith eiai a?ae?io?eueiino?, ?, ye iane?aeie,
iaoiiaethaaoe c?inoaiiy aaeaac?? M. capsulatus aei iueiai.

Cia/i? a?aei?iiino? niinoa??aaee a aaeaac?? M. parvus ? B. megaterium a
noi?oao ?o eoeueoo?. Oae, e?euee?noue ee?oei M. parvus, aaeaaciaaieo aei
oaa?aeeo iaoa??ae?a, ia?aaeuoaaea iaeaea ia 2 ii?yaeee /enaeuei?noue
ee?oei aaoa?io?ioa. Iiaeeeai, oea iaoiiaeaii aenieith nii??aeiai?noth M.
parvus aei oaa?aeeo iiaa?oiiue, ui a ia?oo /a?ao caeiaa a?eueo aeinooii?
aeey aaeaac?? ae?eyiee ia iiaa?oi? oaa?aeeo iaoa??ae?a, iia??ooth/e
iiaeeeaino? aeey i?ee??ieaiiy ?ioeo i?e?ii?aai?ci?a, o aeaiiio aeiaaeeo,
aeey B. megaterium.

Oaeei /eiii, ia aaeaac?th i?e?ii?aai?ci?a a ci?oaieo eoeueoo?ao, ye
i?aaeei, aieeaa? eiaeai c eiiiiiaio?a oeeo eoeueoo?.

?icae?e 6. Eieii?caoe?y oaa?aeeo iaoa??ae?a /enoeie oa ci?oaieie

eoeueoo?aie iaoaiio?ioieo aaeoa??e

Iieacaii, ui /eno? eoeueoo?e iaoaiio?ioieo aaeoa??e o a?aeiia?aeieo
oiiaao aeoeaii eieii?cothoue oaa?ae? iaoa??aee c ??ciith
a?ae?ioiai?noth. I?ney 6 ae?a ?ieoaoaaiiy ia oaa?aeeo iaoa??aeao aeai?
i?e?ii?aai?cie ooai?ththoue e?oii? aa?aaaoe ? i?e?ieieii??.
*enaeuei?noue M. capsulatus i?e oeueiio aeinyaa? aeanyoe?a oeny/ (?en.
2, a), a M. parvus — nioaiue oeny/ ee?oei ia ii2 iiaa?oi? c?ace?a.

A oie aea /an i?e ?ieoaoaaii? B. megaterium (?en. 2, a) ? P. putida a
oiiaao, aeaeoeaieo aeey iaoaiio?ioieo aaeoa??e, a?aei?/aee cieaeaiiy ?o
/enaeueiino? ye ia iiaa?oi? c?ace?a, oae ? a aeeaeeueiiio na?aaeiaeu?,
ui iiynith?oueny a?aenooi?noth aeaea?ae aeeaeaiiy a oaeeo oiiaao aeey
aaoa?io?io?a.

I?ioean eieii?caoe?? neea aaeoa??yie M. capsulatus a ci?oaieo
eoeueoo?ao c B. megaterium aiia?eueithaaany, a oie aea /an
?ioaineai?noue eieii?caoe?? iie?noe?iea oeei iaoaiio?ioii ia
ci?ithaaeanue ii??aiyiii c /enoith eoeueoo?ith (?en. 3, a). M. parvus o
ci?oaieo eoeueoo?ao c aeaiei aaoa?io?ioii aeaui e?aua eieii?coaaa
oaa?ae? iaoa??aee, iniaeeai iie?noe?ie. A oie aea /an /enaeuei?noue
i?ee??ieaieo ee?oei B. megaterium oaeaeei cieaeoaaeanue ? a e?ioe?
oa?i?io ?ieoaoaaiiy aiie i?e?ineii?/ii ia aeyaeyeenue. Oea iia’ycaii, ia
iaoo aeoieo, c ianoa/ath iaiao?aeii? e?eueeino? iaoaaie?o?a
iaoaiio?io?a, iaiao?aeieo aeey i?aeo?eiaiiy aeeoo?ae?yeueiino? aaoeee.
Ooieoe?iioaaiiy

P. putida o ci?oaieo eoeueoo?ao c iaoaiio?ioaie, iaaiaee,
aiia?eueithaaei i?ioean eieii?caoe?? iiaa?oiiue oeeie aaeoa??yie (?en.
3,a). Iaeiae e?euee?noue ee?oei aeaiiai aaoa?io?ioa iinooiiai c?inoaea,
ui iiaea na?ae/eoe i?i niiaeeaaiiy iei iaoaaie?o?a, ui neioacothoueny a
i?ioean? iaoaaie?cio iaoaio iaoaiio?ioieie aaeoa??yie.

?en. 2. *enaeuei?noue /enoeo eoeueoo? aaeoa??e M. capsulatus (a) ?

B. megaterium (a) (N1) ia iiaa?oi? neea (1) ? iie?noe?ieo (2), a oaeiae
a noniaic?? (N2), (3) a caeaaeiino? a?ae /ano ?ieoaoaaiiy. Oiiae
aeine?aeo: iaoaii-iia?o?yia noi?o (3 : 7), t – 37 (C

?en. 3. *enaeuei?noue aaeoa??e M. capsulatus (1–3) ? B. megaterium (4–6)
(a), oa M. parvus (1–3) ? P. putida (4–6), (a) a caeaaeiino? a?ae /ano
?ieoaoaaiiy: ia oaa?aeeo iaoa??aeao (N1) – nee? (1,4) ? iie?noe?ie?
(2,5); a oaeiae a noniaic?? (N2) – (3,6). Ni?aa?aeiioaiiy ia’?i?a acyoeo
noniaic?e 10 : 1. Oiiae aeine?aeo: iaoaii-iia?o?yia noi?o (3 : 7), t –
37 (C (a) ? 30 (C (a)

Aea/ai? caeiiii??iino? eieii?caoe?? iaoaiio?ioieie aaeoa??yie oaa?aeeo
iaoa??ae?a a caeaaeiino? a?ae eiiiiiaioiiai neeaaeo aaciai? noi?o?.
Anoaiiaeaii, ui i?ioean eieii?caoe?? aeine?aeaeaieie iaoaiio?ioaie ye
a?ae?io?eueieo, oae ? a?ae?ioiaieo noano?ao?a ?ioaineaii i?io?eaa o
oe?ieiio ae?aiacii? eiioeaio?aoe?e iaoaio ? eenith (?en. 4, 5). I?e
ai?no? a aacia?e

?en. 4. Eieii?caoe?y iiaa?oi? neea (1) ? iie?noe?ieo (2) aaeoa??yie

M. parvus o aacia?e oa ??aee?e oac? a caeaaeiino? a?ae eiioeaio?aoe??
CH4 a aacia?e noi?o? ? a?aenoai? (h), o ii, aei a?aieoe? iiae?eo
“??aeeia – aac” (0), “+” — iaae ??aiai ??aeeie, “–” — caio?aia /anoeia.
Oiiae aeine?aeo: ii/aoeiaa eiioeaio?aoe?y eenith — 14 %, t – 30 (C

noi?o? iacia/ieo (2 – 5 %) eiioeaio?aoe?e iaoaio ? eenith eieii?caoe?y
oaa?aeeo iaoa??ae?a iaoaiio?ioieie aaeoa??yie a?aeaoaaeanue ii??aiyiii
neaaei. Oea, eiia??ii, iaoiiaeaii e?i?ooaaiiyi i?ioeano aeaieie
noano?aoaie. I?aeaeuaiiy ?o ai?noo iiiaae 9 % noi?iaiaeaeo?oueny a?eueo
?ioaineaiith eieii?caoe??th oaa?aeeo iaoa??ae?a iaoaiio?ioaie. Iaeiae,
i?e eiioeaio?aoe?yo eenith, ye? ia?aaeuoaaee 28 %, niinoa??aaee
?ia?aoaaiiy aeaiiai i?ioeano.

Anoaiiaeaii, ui eieii?caoe?y oaa?aeeo iaoa??ae?a iaoaiio?ioieie
aaeoa??yie i?io?ea? ye ia iiaa?oiyo, ui caio?ai? a aeeaeeueia
na?aaeiaeua, oae /anoeiai ? ia oeo, ui ciaoiaeyoueny a aacia?e oac?.
Nooo?ai? ??cieoe? a eieii?caoe?? neea ? iie?noe?ieo oeeie
i?e?ii?aai?ciaie ia niinoa??aaeinue. Iaea?eueo ?ioaineaii oeae i?ioean
i?io?eaa ia /anoei? c?ace?a, caio?ai?e o aeeaeeueia na?aaeiaeua ia
a?aenoai? ia a?eueoa 15 ii a?ae a?aieoe? iiae?eo “??aeeia – aac”.

I?e canoinoaaii? ?iiia?e?ciaaieo iaoaiio?ioieo aaeoa??e o
a?ioaoiieia?/ieo i?ioeanao iaeiei c iaeaaaeeea?oeo oaeoi??a, ui
aecia/aoeiooue o?c?ieia?/io aeoeai?noue oeeo i?e?ii?aai?ci?a, ?
aieia?noue ioi/oth/iai na?aaeiaeua. Iaie anoaiiaeaii, ui iaea?eueo
?ioaineaii oeae i?ioean a?aeaoaaany

?en. 5. Eieii?caoe?y iiaa?oi? neea (1) ? iie?noe?ieo (2) aaeoa??yie

M. capsulatus o aacia?e oa ??aee?e oac? a caeaaeiino? a?ae
eiioeaio?aoe?? O2 a

aacia?e noi?o? ? a?aenoai? (h), o ii, aei a?aieoe? iiae?eo “??aeeia –
aac” (0),

“+” — iaae ??aiai ??aeeie, “–” — caio?aia /anoeia. Oiiae aeine?aeo:
ii/aoeiaa eiioeaio?aoe?y iaoaio — 30 %, t – 37 (C

ia cii/ai?e iiaa?oi? ia i?ioyc? ia?oeo aeaio ae?a. Ciaioaiiy aieiaino?
iin?y, yeei neoaeeea a??nueea ii?iaea aoa?eueieo oaoo, i?ecaiaeeei aei
ciaioaiiy ia’?i?a niiaeeoiai iaoaio aeine?aeaeaieie eoeueoo?aie
iaoaiio?io?a.

I?e canoinoaaii? ?iiia?e?ciaaieo iaoaiio?ioieo aaeoa??e c iaoith
cieaeaiiy iaoaiiaeae?eaiiy c ae?iaeaieo i?inoi??a aoa?eueieo oaoo
i?aeoe/ii a?aenooi? iiaeeeaino? aianaiiy aeiaeaoeiaeo aeic aeeaeeueiiai
na?aaeiaeua aeey oeeo i?e?ii?aai?ci?a. Iieacaii, ui a oiiaao
iaeii?aciaiai aianaiiy aeeaeeueiiai na?aaeiaeua iaea?eueo aeoeaia
niiaeeaaiiy iaoaio ?iiia?e?ciaaieie eoeueoo?aie iaoaiio?io?a oa ?o
ci?oaieie eoeueoo?aie c aaeoa??yie ?iaeo Bacillus a?aeaoaaeinue ia
ae?oao —ainueio aeiao eoeueoeaoaaiiy (?en. 6). C /anii ?ioaineai?noue
niiaeeaaiiy iaoaio ?iiia?e?ciaaieie i?e?ii?aai?cia-

?en. 6. ?ioaineai?noue niiaeeaaiiy iaoaio /enoith eoeueoo?ith M.
capsulatus (1) ? a ci?oaieo eoeueoo?ao c B. megaterium (2) oa B.
subtilis (3) i?e eieii?caoe?? ieie oaa?aei? iiaa?oi?

ie cieaeoaaeanue ? noaiiaeea ca 14 aeiao 42 % aeey /enoi? eoeueoo?e

M. capsulatus ? 12 oa 60 % aeey ci?oaieo eoeueoo? c B. megaterium oa B.
subtilis, a?aeiia?aeii, o ii??aiyii? c 1th aeiaith ?o eoeueoeaoaaiiy.
Io?eiai? ?acoeueoaoe aeicaieythoue i?iaiicoaaoe i?ioean ieeneaiiy CH4 o
ae?iaeaieo i?inoi?ao aoa?eueieo oaoo iaoaiio?ioieie aaeoa??yie i?e ?o
ooieoe?iioaaii? c ?ioeie i?aaenoaaieeaie oeiiai? i?e?ioei?e oeeo aeii?o.

AENIIAEE

1. ?ic?iaeai? eaai?aoi?i? onoaiiaee c eiioo?ii oee?eoeyoe?? aaciai?
noi?o?, ui aeicaieythoue cieceoe iiaeeea?noue eiioai?iaoe??
iaoaiio?ioieo aaeoa??e i?e ?o eoeueoeaoaaii? a ??aeeeo ? aaa?eciaaieo
na?aaeiaeuao.

2. Aeey iiaeaethaaiiy oiia ?nioaaiiy iaoaiio?io?a o aoa?eueieo oaooao
Aeiiaano ceiiiiiiaai? ci?oai? eoeueoo?e, aei neeaaeo yeeo e??i
iaoaiio?ioieo aaeoa??e aaaaeai? i?e?ii?aai?cie ?iae?a Bacillus /e
Pseudomonas. O?c?ieia?/ia aeoeai?noue iaoaiio?ioieo aaeoa??e o ??cieo
i?e?iaeieo oa a?ioaoiieia?/ieo i?ioeanao aoaea caeaaeaoe a?ae oeo aeae?a
i?e?ii?aai?ci?a, c yeeie aiie ni?a?niothoue. Ooaie ?iaeo Bacillus o
ci?oaieo eoeueoo?ao ia ?ia?aothoue o?c?ieia?/io aeoeai?noue
iaoaiio?io?a. A oie aea /an Pseudomonas putida o ci?oai?e eoeueoo??
?ia?ao? ?icaeoie iaoaiio?io?a.

3. Aeniea a?ae?ioiai?noue M. parvus ? aieiaiei oaeoi?ii, ui iaoiiaeth?
?o ?ioaineaio aaeaac?th aei a?ae?io?eueieo ? a?ae?ioiaieo iaoa??ae?a, ia
aeeaey/enue ia cia/iee iaaaoeaiee ca?yae iiaa?oi? ee?oei. Aaeaac?y
aaeoa??e, ee?oeie yeeo oa?aeoa?ecothoueny a?ae?io?eueiith iiaa?oiath, a
cia/i?e i??? aecia/a?oueny ?o aeaeo?iiiaa?oiaaeie aeanoeainoyie ?
c?inoa? c? ciaioaiiyi iaaaoeaiiai ca?yaeo ee?oei. O noi?oao noniaic?e
aaeoa??e ?iae?a Bacillus ? Pseudomonas oa iaoaiio?io?a nooo?ai
ci?ith?oueny aaeaac?y eiaeiiai c ie?aieo aeae?a i?e?ii?aai?ci?a.

4. A aeaeoeaieo aeey iaoaiio?io?a oiiaao, oe? aaeoa??? ?ioaineaii
eieii?cothoue a?ae?io?euei? ? a?ae?ioiai? iaoa??aee, ooai?thth/e ia 6o
aeiao i?e?ieieii?? a /enoeo ? a a?ia?ieo eoeueoo?ao c i?aaenoaaieeaie
?iaeo Bacillus. I?e oeueiio /enaeuei?noue aaoeee ia oaa?aeeo iaoa??aeao
iinooiiai cieaeoaaeanue.

5. A ci?oaieo eoeueoo?ao c P. putida i?ioean eieii?caoe?? oaa?aeeo
iiaa?oiiue iaoaiio?ioieie aaeoa??yie oiia?eueithaaany, a /enaeuei?noue
inaaaeiiiiaae c?inoaea.

6. ?iiia?e?ciaai? iaoaiio?ioi? aaeoa??? caeaoi? eieii?coaaoe
a?ae?io?euei? ? a?ae?ioiai? iaoa??aee o oe?ieiio ae?aiacii?
eiioeaio?aoe?e iaoaio oa eenith o aacia?e noi?o?. Iaea?eueo ?ioaineaii
i?ioean eieii?caoe?? oaa?aeeo iaoa??ae?a i?io?ea? i?e ai?no? o noi?o? 17
– 28 % iaoaio oa 5 – 17 % eenith. Nooo?aeo a?aei?iiinoae o eieii?caoe??
a?ae?io?eueieo (neei) oa a?ae?ioiaieo (iie?noe?ie) iiaa?oiiue ??cieie ca
a?ae?ioiai?noth eoeueoo?aie iaoaiio?io?a ia a?aei?/aii.

7. Iaoaiio?ioi? aaeoa??? aeoeaii ieenethaaee iaoai ca iayaiino? ia iin??
(a??nueea ii?iaea aoa?eueieo oaoo) a?coaeueii aeaeeii? aieiae. I?e
ciaioaii? aieiaino? iin?y ?ioaineai?noue aeaiiai i?ioeano iinooiiai
cieaeoaaeanue.

8. Iaea?eueo aeoeaia niiaeeaaiiy iaoaio /enoeie oa ci?oaieie eoeueoo?aie
iaoaiio?io?a c aaeoa??yie ?iaeo Bacillus a?aeaoaaeinue ia ae?oao —
ainueio aeiao eieii?caoe?? ieie oaa?aeeo iaoa??ae?a, ui aeicaiey?
i?iaiicoaaoe i?ioean ieeneaiiy CH4 o ae?iaeaieo i?inoi?ao aoa?eueieo
oaoo iaoaiio?ioieie aaeoa??yie i?e ?o ooieoe?iioaaii? c ?ioeie
i?aaenoaaieeaie oeiiai? i?e?ioei?e oeeo aeii?o.

NIENIE ?IA?O, IIOAE?EIAAIEO CA OAIITH

AeENA?OAOe??

1. Eenoaiue A. A., Eeaaeue I. O., Eo?aeeo E. E. e ae?. Aeeyiea iaeioi?uo
oeceei-oeie/aneeo oaeoi?ia n?aaeu ia aaeaaceth iaoaiio?ioiuo aaeoa?ee //
Iee?iaeie. aeo?i. — 1996. — 58, ( 3. — N. 62 — 70.

2. Ai?aeeaiei A. N., Eenoaiue A. A., Eo?aeeo E. E. Iiaa?oiinoiua
naienoaa e aaeaacey iaoaiio?ioiuo aaeoa?ee // Oeiey e oaoiieiaey aiaeu —
1997. — 19, ? 2. — N. 216 — 221.

3. Eenoaiue A. A., Eo?aeeo E. E. Aeeyiea aeaaeiinoe ia iio?aaeaiea
iaoaia eiiiaeeeciaaiiuie iaoaiieeneythueie aaeoa?eyie // I?e?ia?ie.
aeo?i. — 1998. — 60, ( 2. — N. 79 — 85.

4. ?ie A. A., Eenoaiue A. A., Eo?aeeo E. E. Oeceieiae/aneay aeoeaiinoue
iaoaiio?ioia a niaoaiiuo eoeueoo?ao n oeie/iuie i?aaenoaaeoaeyie
iee?ioei?u oaieueiuo oaoo // I?e?ia?ie. aeo?i. — 1998. —60, ( 3. — N. 24
— 30.

5. Eo?aeeo E. E., Ai?aeeaiei A. N., Eeaaeue I. O., Eenoaiue A. A.
N?aaieoaeueiue aiaeec aaeaacee ?acee/iuo aeaeia iaoaiio?ioiuo aaeoa?ee e
oaa?aeui iaoa?eaeai // I?eee. aeioei. e iee?iaeie. — 1998. — 34, ? 3. —

N. 245 — 250.

6. Eenoaiue A. A., ?ie A. A., Eo?aeeo E. E., Oaa/aiei O. A. Aaeaacey
niaoaiiuo eoeueoo? iaoaiio?ioia e oaa?aeui iaoa?eaeai // I?eee. aeioei.
e iee?iaeieiaey. — 1998. — 34, ? 4. — N. 414 — 418.

7. Eenoaiue A. A., Eeaaeue I. O., Eo?aeeo E. E. ?ieue oeceei-oeie/aneeo
oaeoi?ia a aaeaacee iaoaiio?ioiuo aaeoa?ee // Iaoa??aee I onoaiia/iai
(VIII) c’?caeo oe?a?inueeiai i?e?ia?ieia?/iiai oiaa?enoaa. Oac. aeiee. —
I?e?ia?ie. aeo?i. — 1994. — 56, ? 2. — N. 64.

AIIOAOe??

E?noaiue I. A. Iniaeeaino? aca?iiae?? /enoeo ? ci?oaieo eoeueoo?
iaoaiio?ioieo aaeoa??e c oaa?aeeie iaoa??aeaie. — ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaith eaiaeeaeaoa a?ieia?/ieo
iaoe ii niaoe?aeueiino? 03. 00. 07 — i?e?ia?ieia?y. — ?inoeooo
i?e?ia?ieia?? ? a??onieia?? IAI Oe?a?ie, Ee?a, 1998.

Aeena?oaoe?y i?enay/aia aea/aiith iniaeeainoae ooieoe?iioaaiiy ci?oaieo
eoeueoo?, ui i?aaenoaaeai? oeiiaith i?e?ioei?ith aoa?eueieo oaoo, aei
neeaaeo yei?, ii?yae c iaoaiio?ioaie, aoiaeyoue aaoeeee ?
inaaaeiiiiaaee. Aea/ai? aeanoeaino? iiaa?oi? ee?oei aeacaieo
i?e?ii?aai?ci?a ? aecia/aii ?o aieea ia aaeaac?th aei a?ae?io?eueieo
(neei) ? a?ae?ioiaieo (iie?noe?ie) iaoa??ae?a. Iieacaii, ui
ooieoe?iioaaiiy iaoaiio?ioieo aaeoa??e o a?ia?ieo eoeueoo?ao c
i?aaenoaaieeaie ?iae?a Bacillus ? Pseudomonas nooo?ai aieeaa? ia
aaeaac?th eiaeiiai c oeeo aeae?a i?e?ii?aai?ci?a.

Aea/ai? iniaeeaino? eieii?caoe?? oaa?aeeo iaoa??ae?a /enoeie oa
ci?oaieie eoeueoo?aie i?aaenoaaiee?a oeiiai? i?e?ioei?e aoa?eueieo oaoo
a oiiaao, aeaeoeaieo aeey iaoaiio?io?a. Aeine?aeaeaia caeaaei?noue
?ioaineaiino? eieii?caoe?? neea oa iie?noe?ieo iaoaiio?ioaie c ??ciei
nooiaiai a?ae?ioiaiino? iiaa?oi?, a?ae eiiiiiaioiiai neeaaeo aaciai?
noi?o?. Aecia/ai? iioeiaeuei? aeey oeueiai i?ioeano eiioeaio?aoe??
iaoaio oa eenith o aacia?e noi?o?.

Aeine?aeaeaii aieea aieiaino? iin?y (a??nueea ii?iaea aoa?eueieo oaoo)
ia ?ioaineai?noue ieeneaiiy iaoaio ?iiia?e?ciaaieie iaoaiio?ioieie
aaeoa??yie, a oaeiae oa?i?io eoeueoeaoaaiiy ia o?c?ieia?/io aeoeai?noue
/enoi? ? ci?oaii? eoeueoo? iaoaiio?io?a.

Eeth/ia? neiaa: iaoaiio?ioe, ci?oai? eoeueoo?e, aeanoeaino? iiaa?oi?,
aaeaac?y, eieii?caoe?y, ciai?oi? oaeoi?e, o?c?ieia?/ia aeoeai?noue.

Eenoaiue A. A. Iniaaiiinoe acaeiiaeaenoaey /enouo e niaoaiiuo eoeueoo?
iaoaiio?ioiuo aaeoa?ee n oaa?aeuie iaoa?eaeaie. — ?oeiienue.

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie eaiaeeaeaoa aeieiae/aneeo
iaoe ii niaoeeaeueiinoe 03. 00. 07 — iee?iaeieiaey. — Einoeooo
iee?iaeieiaee e ae?onieiaee IAI Oe?aeiu, Eeaa, 1998.

Aeenna?oaoeey iinayuaia eco/aieth iniaaiiinoae ooieoeeiie?iaaiey
niaoaiiuo eoeueoo?, eioi?ua i?aaenoaaeaiu oeie/iie iee?ioei?ie oaieueiuo
oaoo, a ninoaa eioi?ie, ia?yaeo n iaoaiio?ioaie, aoiaeyo aaoeeeeu e
inaaaeiiiiaaeu. Ia iiaeaeyo aeia?iuo eoeueoo? iieacaii, /oi i?enoonoaea
nii?iia?acothueo aaeoa?ee Bacillus megaterium EIA-476 e Bacillus
subtilis 9 noeioee?iaaei ?inoiaoth aeoeaiinoue e iio?aaeaiea iaoaia
Methylocystis parvus OBBP e Methylococcus capsulatus ANA-874,
/eneaiiinoue aaoeeee i?e yoii i?aeoe/anee ia eciaiyeanue. A i?enoonoaee
Pseudomonas putida 11, iaiai?io, iaaethaeaee, nieaeaiea oeceieiae/aneie
aeoeaiinoe iaoaiio?ioia, oiaaea eae /eneaiiinoue yoiai aaoa?io?ioa
aic?anoaea.

Eco/aia naycue naienoa iiaa?oiinoe e aaeaacee enneaaeiaaiiuo
iee?ii?aaieciia e aeae?ioeeueiui (noaeei) e aeae?ioiaiui (iieenoe?ie)
iaoa?eaeai. Iieacaii, /oi eioaineaiia i?ee?aieaiea M. parvus e yoei
iaoa?eaeai iaoneiaeaii aunieei iieacaoaeai aeae?ioiaiinoe, ianiio?y ia
cia/eoaeueiue io?eoeaoaeueiue ca?yae yoeo aaeoa?ee. Inoaeueiua
iee?ii?aaieciu oa?aeoa?eciaaeenue aeae?ioeeueiie iiaa?oiinoueth, a aa
ca?yae nouanoaaiii aeeye ia aaeaaceth, eioi?ay aic?anoaea n oiaiueoaieai
io?eoeaoaeueiiai ca?yaea eeaoie. Aaeoa?ee ?aciuo ?iaeia i?aeoe/anee a
iaeeiaeiauo eiee/anoaao i?ee?aieyeenue eae e aeae?ioeeueiui, oae e
aeae?ioiaiui iaoa?eaeai.

Enneaaeiaaiu iniaaiiinoe aaeaacee iaoaiio?ioia e i?aaenoaaeoaeae ?iaeia
Bacillus e Pseudomonas a aeia?iuo eoeueoo?ao e onoaiiaeaii, /oi ia
i?ee?aieaiea iee?ii?aaieciia aeeyao eaaeaeue ec eiiiiiaioia oaeeo
eoeueoo?.

Eco/aiu iniaaiiinoe eieiiecaoeee oaa?aeuo iaoa?eaeia a oneiaeyo,
yeaeoeaiuo aeey iaoaiio?ioia yoeie aaeoa?eyie, a oaeaea eo niaoaiiuie
eoeueoo?aie n oeacaiiuie aaoa?io?ioaie. Iieacaii, /oi iaoaiio?iou
eioaineaii eieiiece?otho aeae?ioeeueiua e aeae?ioiaiua iaoa?eaeu a
/enouo e a niaoaiiuo eoeueoo?ao n i?aaenoaaeoaeyie ?iaea Bacillus.
*eneaiiinoue aaoeeee ia oaa?aeuo iaoa?eaeao i?e yoii nieaeaeanue. A
i?enoonoaee P. putida i?ioeann eieiiecaoeee oaa?aeuo iiaa?oiinoae
iaoaiio?ioaie caiaaeeyeny, a eiee/anoai inaaaeiiiiaae aic?anoaei.

Enneaaeiaai i?ioeann eieiiecaoeee iaoaiio?ioiuie aaeoa?eyie oaa?aeuo
iaoa?eaeia a caaeneiinoe io eiiiiiaioiiai ninoaaa aaciaie niane.
Onoaiiaeaii, /oi aeaiiua iee?ii?aaieciu niiniaiu eieiiece?iaaoue
oeacaiiua iaoa?eaeu a oe?ieii aeeaiaciia eiioeaio?aoeee iaoaia e
eenei?iaea, a iaeaieaa eioaineaii — i?e niaea?aeaiee yoeo aacia a niane
17 – 28% e 5 – 17%, niioaaonoaaiii. Nouanoaaiiuo ioee/ee a eieiiecaoeee
aeae?ioeeueiuo e aeae?ioiaiuo iaoa?eaeia iaoaiio?ioaie ia ioia/aii.

Enneaaeiaaii aeeyiea aeaaeiinoe iineoaey (ai?iay ii?iaea oaieueiuo
oaoo) ia eioaineaiinoue ieeneaiey iaoaia eiiiaeeeciaaiiuie
iaoaiio?ioiuie aaeoa?eyie. Iieacaii, /oi iaeaieaa aeoeaii yoio i?ioeann
i?ioaeaao i?e iaee/ee ia iineoaea aecoaeueiie aeaae.

Eco/aia oeceieiae/aneay aeoeaiinoue /enoie e niaoaiiuo eoeueoo?
iaoaiio?ioiuo e nii?iia?acothueo aaeoa?ee i?e eieiiecaoeee ieie oaa?aeie
iiaa?oiinoe (/anoeoe noaeea) a oneiaeyo aeeeoaeueiiai eoeueoeae?iaaiey
aac caiaiu ieoaoaeueiie c?aaeu. Onoaiiaeaii, /oi iaeaieaa aeoeaiia
iio?aaeaiea iaoaia eiiiaeeeciaaiiuie eoeueoo?aie iaoaiio?ioia eae a
/enoie, oae e a niaoaiiuo eoeueoo?ao n aaoeeeeaie i?ienoiaeeo ia 2 – 7
nooee eoeueoeae?iaaiey. Iieo/aiiua ?acoeueoaou iicaieytho
i?iaiice?iaaoue eioaineaiinoue i?ioeanna ieeneaiey CH4 a au?aaioaiiuo
i?ino?ainoaao oaieueiuo oaoo eiiiaeeeciaaiiuie iaoaiio?ioiuie aaeoa?eyie
e eo niaoaiiuie eoeueoo?aie n ae?oaeie i?aaenoaaeoaeyie yoeo yeiieo n
oeaeueth nieaeaiey auaeaeaiey iaoaia a ai?iua au?aaioee.

Eeth/aaua neiaa: iaoaiio?iou, niaoaiiua eoeueoo?u, naienoaa
iiaa?oiinoe, aaeaacey, eieiiecaoeey, aiaoiea oaeoi?u, oeceieiae/aneay
aeoeaiinoue.

Kisten O. G. Peculiarities of pure and mixed cultures of methanotrophic
bacteria interaction with solid materials. — Manuscript.

Thesis for a candidate degree by speciality 03. 00. 07 — microbiology.
— Institute of Microbiology and Virology of National Academy of Science
of Ukraine, Kyiv, 1998.

Dissertation is devoted to study of features of functioning of mixed
cultures, which are submitted by a typical microflora of coal mines,
contenting methanotrophs, Bacillus and Pseudomonas species. Properties
of a cells surface of these microorganisms are investigated and is
determined of these properties influence on adhesion to hydrophilic
(glass) and hydrophobic (polystyrene) materials. Is shown, that
functioning of methanotrophic bacteria in mixed cultures with the
representatives of Bacillus and Pseudomonas genus essentially influenced
on adhesion of each of these species of microorganisms.

Features colonization of solid materials by pure and mixed cultures of
the representatives of a typical microflora of coal mines in elective
conditions for methanotrophs are investigated. Dependence of
colonization intensity of a glass and polystyrene by methanotrophs with
a different cells surface hydrophobicity degree from a different gas mix
contents is investigated. Methane and oxygen concentration optimum in a
gas mix for this process are determined.

Influence of humidity of a carrier (rock particles of coal mines) on
methane oxidation intensity of immobilized methanotrophic bacteria as
well as time of cultivation on physiological activity of pure and mixed
cultures of methanotrophs is investigated.

Key words: methanotrophs, mixed cultures, cells surface properties,
adhesion, colonization, external factors, physiological activity.

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