, 2012, 81, 3, .

306313

576.8.093.1.095.21

2012 . . . , . . , . . , . . 1
. .. ,
25.08.2011 .


. 
. 104 
,
0.22 .
.
, 
16S , .
16S
Betaproteobacteria (9499%) 
16S Herbaspirillum, Herminiimonas, Curvibacter
Burkholderia. 16S
Euryarchaeota. 
(9799%) 16S Methanobacteriales
Methanosarcinales. 16S 
(7174% ) .
: , , 
16S .
1
, 
,
XXI . 


, 
, 
.

.
,
, 
,
, 

in situ
 
[1]. 
,
, 
0.22 ,

, , .
.
1 (email: dedysh@mail.ru).

, 
,
.

1981 . 

0.3 , 


[2].
,
, 
, 

[3, 4]. 


0.1 3 [4, 5].

0.22 .

,

,
, 
[2, 3, 6].

306

307

34.1

0.64

0.04

. 0.19

1.1

4.44.6 111291

20

4.54.8

,
. /

P, /
0.03

2.5

N, /
0.34

0.02

, /

S/

5.9

pH

10

1155

, 2

,
PtCo

1.03 0.1 3.07

3.89 0.4 0.43

[1, 12].




. 
,

,
Bacteroidetes, Alphapro(
teobacteria, Betaproteobacteria, Actinobacteria Spiro(
chaetes [6]. 


120 [7].


.

, 

.

[8], 
[9],
[10]. 

[11].
, 
104
,
0.22 , 0.03
3.6%
[11].
. ,



.

81

2012

.


, 
 


( , 5810 .., 3733 ..).


, pH
.
( 4.7) 

( 40%). 
( 4.4) 


( 90% ) [12]. 

. 
( 0.20.5 ).
>
. 

(12 ) 
(Millipore,
0.22 ) .

4 , 0.5 

(4',6'2
) 10 , 
. 

Zeiss Axioplan 2 (, )
Zeiss 02, 
. 100 

2*

308

1 . 
,
0.22 , 5

0.09 .
.

>
.

, .

0.22
.
1) SB2, (/): KNO3 50,
NH4Cl 50, MgSO4 7H2O 60, aCl2 2H2O 20,
50, Na 300;
Difco 18 /.
(20 1 ) 5.2.
2) R2A
(Difco), 100 .

30 / .
2 
.

. 

( 500 ..) 
16S pPHK.

.
100

0.22 (Millipore) 

0.09
(Millipore). 
3
.

FastDNA SPIN kit for soil (Biol 101,
) 
. 

().  
(~1490 ) 16S 

9f 1492r [13]. 
(~800 )
16S 
109f 915r [14].
PE GeneAmp PCR System 9700
(PerkinElmer Applied Biosystems, ). 

1.2% 

. 

pGemT Easy Vector SystemII (Promega). 


 7 SP6.

. 

MspI RsaI,
HhI BsuRI (HaeIII). 
2.5% .

Wizard Plus
Minipreps DNA Purification System (Promega).

ABI 377A (Perkin
Elmer Applied Biosystems, ). 


SeqMan (Laser Gene 7.0; DNA Star Package).

GenBank 
Blast2 (http://
www.ebi.ac.uk/Tools/blast2/). 
16S

Bellerophon 3.0 (http://compbio.anu.edu.au/bellero
phon/bellerophon.pl).

ARB (http://www.arbhome.de).


Phylip bootstrap 
1000 .

16S 

GenBank JN825208JN825301.

.

3.89 0.40
1.03 0.10 106 1 .

0.22
1.69 0.53 3.16 0.43 104 1, 
0.4 3.1% 

().
. 
, 
,
1.68 105 1.25 105 / R2A (1 : 100)
SB2 . , 


81

2012


lone DB4 (2), DB7, DB8 (2),
DB36, DB42, DB65, MB48 (3),
MB83 (2), MB87, MB89 (2),
MB97, MB98, MB109 (2)
lone DB38 (8)
lone DB40
lone DB34 (2)
100
Herbaspirillum lusitanum, AF543312
lone DB41 (2)
lone DB45
92
lone DB60
78
Minibacterium massiliensis, CP000269
Heminiimonas glaciei, EU489741
lone MB105 (5)
lone MB111
lone MB58 (3)
100 lone DB50 (7)
100
Curvibacter gracilis, AB109889
Burkholderia fungorum, AJ544691
87
Burkholderia hospita, AY040353
90
lone MB108 (12)
lone MB115 (5)
lone MB24
99
lone MB99 (5)
Ferrovum myxofaciens, EF133508
lone DB27
Pseudomanas antarctica, FM955872
100
lone DB28
Pseudomanas fluorescens, AF506040
bacterium Ellin362, AF498744
100
99
lone MB47
lone MB125 (3)
Mesorhizobium amorphae, AF041442
100
lone MB123 (6)
Candidatus Planktophila limnetica, FJ428831

309

95

96

70

99

Betaproteobacteria

Gammaproteobacteria
Alphaproteobacteria
Actinobacteria

0.10

. 1. , 
16S , ,
Proteobacteria Actinobacteria. , ,
DB MB . 
16S Gemmata obscuriglobus (X54522), Isophaera pallida (AJ231195), Schlesneria palu(
dicola (AM162407) Planctomyces limnophilus (X62911). 0.1 .

,
0.93 102 2.00 102 /, . 
, 0.51.2%


.
, 
,
11 . 
16S 
(9799%) 

Mesorhizobium, Bradyrhizo(
bium, Pseudomonas, Sphingomonas Agrobacterium.

, 0.30.5 ,
, 
.

16S Pseudomonas,

81

2012

DSB4 DSB11, ,

(. ).

.

16S 
86 , 25
, 61
. 74 

Betaproteobacteria, 4 Alphaproteo(
bacteria, 2 Gammaproteobacteria
Actinobacteria (. 1).
,
,
Oxalobacteraceae, Comamonadaceae and
Burkholderiaceae. 
16S 
(9499%) 

310

Herbaspirillum, Herminiimonas, Curvibacter

Burkholderia.

DB27 DB28, Gammapro(
teobacteria, (99%) 
16S pPHK 
Pseudomonas, 
[15].

16S
DSB4 DSB11,
.
,
Alphaproteobacteria, 
(~90%
16S )
. 
(9597%) 
16S 
Ellin362, [16].
, Actino(
bacteria,
(95%) 16S 

, Candidatus Planktophila
limnetica [17].

. 
16S 
,
, 120 
Euryarchaeota, 48 
72 
(. 2).

(9799%) 16S 

Methanobacteriales Methanosarci(
nales. 34

Methanobacterium, ,

[18, 19]
[20]. 3 4 , 
,

16S Metha(
nobrevibacter Methanosarcina .

(7174%
16S ) 
.
(8599%)
, 

, ,
, [21],
[2225].


, 

.

, 


[11].



.
,
0.22 , 

,
[11].

, 
(
).

(0.51.2%) 

.

, 
.
Pseudomonas, 

, 
16S . , 
Mesorhizobium,
Bradyrhizobium, Sphingomonas Agrobacterium

( ) .
.
, 

,

.

, 
, Beta(
proteobacteria.

[1],
[2628]. , 


16S Herminiimonas glaciei 

0.043 3, [27].

81

2012

95

311

lone DA 7 (2), DA 17 (3), DA 86,

DA 91, MA 14 (18), MA 17, MA 20,
MA 24, MA 30, MA 62, MA 63,
Mackenzie river archeon, EF014514

lone DA 36
lone DA 66
Mackenzie river archeon, DQ310430
100 lone DA 74
73
lone DA 85
contaminaled aquifer archaeon, AF050612
100
lone DA 3 (4), DA 62 (2), DA 65,
DA 69, DA 71 (2), DA 90 (2),
95
MA 29, MA 41, MA 45 (4)
lone DA 46 (2)
lone MA 54
lone DA 83
lone MA 47
94
Mackenzie river archeon, EU244266
63
arctic peat archaeon, AM712544
arctic lake archaeon, EU782019
lone DA 57
84
lone MA 25
99 lone DA 16 (2), DA 87,
96
MA 32, MA59
66
hydrote rmal vent archaeon, AB019752
freshwater sediment euryarchaeote, GU257043
lone DA 15
Mackenzie river archeon, DQ310396
94
86
lone DA 55
100
hydrotermal vent archaeon, AB019753
lone DA 26, DA 42
97
MA 18, MA 31, MA 64
63
Mackenzie river archeon, DQ310418
lone MA 11
100 lone MA 15
lone MA 4
lone DA 63 (2)
lone DA 34, DA 43, DA 80, DA 88, MA 1 (2), MA 5, MA 13 (18), MA 22, MA 27, MA 28, MA 33, MA 39, MA 40
minerotrophic fen archaeon, EU155988, arctic peat archaeon, DQ089005, subarctic permafrost archaeon, DQ859923
Methanobacterium bryantii, AF28688
Methanobacterium formicicum, AF028689
97

94
100

lone DA 33, DA 39, DA 89

Methanobacterium thermoautotrophicus, AE000930
lone MA 21 (2)
100 lone DA 25
Methanobrevibacter smithii, U55235
lone DA 47 (4)
100
Methanosarcina lacustera, AF432127
Methanosarcina mazeii, AF028691

LDS
cluster

Methano(
bacteriales

Methano(
sarcinales

0.10

. 2. , 
16S , , 
Methanosarcinales Methanobacteriales, ,
. , , 
D M .
16S Archaeoglobus fulgidus (AE000965), Archaeoglobus profundus (AF297529), Nanoarchaeum equitans (AJ318041) Sul(
folobus solfataricus (X03235). 0.1 .

Minibacterium massiliensis,

, 
[26].

Herbaspirillum Curvibacter,

0.30.5 [29, 30]. 
, , 

,
.

81

2012

16S 
Burkholderia,
.

[3, 8, 31].
,

.
, 


312

, 
[17, 32, 33]. ,
, 
,
.
,
Ellin362, 

[16],
.

, 
, 
, Methanobacteriales Methanosar(
cinales.
Methano(
bacterium, ,
, .

16S 
, 

Euryarchaeota. 
, ,

(AF050612) [34] 
(AB019752) [35].

16S 
Dagow,
[21]. 

LDS (Lake Dagow Sediment).
,

Euryarchaeota.

, 
, ,
,
.. [20, 2225].



, 
Euryarchaeota.

.
,

,
0.22 .



.





: , 
02.740.11.0023.

1. .., .., ..,
.., .. 


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