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Фенин Вадим Андреевич. Биокатализатор спиртового брожения на основе иммобилизованных клеток дрожжей Saccharomyces cerevisiae

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4
«
Saccharomyces
cerevisiae».
: 2018.
: 19.03.01
: 41-
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68
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PШаОЫ PШТЧЭ 2010
13
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4.
«
(
,
1).
.
».
,
5
...................................................................................................................... 7
1.
1.1
......................................... 10
,
..................... 10
1.2
....................................................... 12
1.3
........................................................................... 14
1.4
.............................. 24
,
1.4.1
.............................. 24
1.4.2.
............................................................................................................................. 31
1 ......................................................................................................... 38
2.
........................................... 40
2.1
........................................................................................... 40
2.2
............................................................................................ 40
2.2.1
...................................................................... 40
2.2.2
Saccharomyces cerevisiae .................................... 41
2.2.3
. ....................................................................... 42
2.2.4
. ..................................... 43
2.2.5
.......................................... 43
2.2.6
.................................................. 45
2.2.7
........................................................ 45
2.2.8
................. 48
2.2.9
............................................. 48
3.
.......................................................... 49
3.1
cerevisiae
Saccharomyces
-
................................... 49
3.2
. ........................................ 50
3.3
............................................. 51
6
3.4
..................................................... 52
3.5
-
.......................... 55
3.6
. ........................................................................................................................ 55
3.7
. ..................................................................................... 59
....................................................................................................................... 61
............................................................................................. 63
7
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Saccharomyces cerevisiae.
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.
:
Saccharomyces cerevisiae,
.
.
,
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,3
,
71 –
.
, 10
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71
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9
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10
1.
1.1
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1970–
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11
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12
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,
.
1.2
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,
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1.
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2.
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13
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3.
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4.
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6.
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14
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,
1
.
1.3
,
,
CСООЭСКЦ Д15Ж.
,
,
.
,
,
,
.
.
.
1.
,
,
,
,
Д28Ж.
,
,
Д53Ж.
Д39].
.
-
,
,
,
,
.
.
15
2.
,
,
.
,
-
Dowex-1
[27Ж
,
[23Ж.
(
2 –
2).
Saccharomyces cerevisiae
[19]
,
,
,
,
(IRA 400)
(IRA 458)
(IRA 68)
1
3.0
10,9
,
,
,
,
,
(IRA 900)
(IRA 904)
(IRA 938)
(IRA 958)
(БE 352)
1
4,0
46,8
5,3
128,0
1
/
/
. [19]
Daugulis
Saccharomyces cerevisiae,
.
,
.
,
,
,
.
,
.
,
,
,
.
16
,
.
,
.
OppОЫЦКЧЧ [48Ж.
Messing
,
, . .
,
,
,
,
,
.
,
«
»
,
,
.
.
,
,
16
,
,
,
.
,
.
,
,
.
,
,
,
,
.
,
,
,
,
.
«
» [65Ж.
17
.
,
,
.
Horisberger [30Ж,
.
3. К
(
)
ё
,
.
,
,
. JКМФ
ГКУТМ [32Ж
Micrococcus
75%
luteus.
-
.
,
,
,
)
. Navarro
(
Durand [54]
.
Saccharomyces carlsbergensis.
CШЦpОЫО
GЫТППТЭС [18Ж
,
,
,
,
,
,
,
,
,
,
,
,
(
).
(
)
18
.
.
4.
CШЦpОЫО
GЫТППТЭС [18Ж
,
,
.
,
,
.
,
,
.
,
,
,
.
Lactobacillus.
,
.
,
,
,
,
,
.
(
)
.
)
.
,
.
:
,
.
,
,
,
,
.
19
,
,
.
,
,
,
Umbilicaria
pustulate [50].
.
Rosevear
[58,59Ж
.
,
,
,
.
.
[15Ж,
,
,
;
.
)
,
,
.
,
,
,
.
,
.
-
20
.
-
-D3,6-
-
.
-D.
,
,
,
-
,
,
.
,
,
:
-
37-60°
.
.
,
,
.
L-
D,
,
.
.
ё
:
,
.
,
,
,
-
.
,
:
Macrocystis
Ascophyllum
.
[14,43Ж.
.
,
20
,
,
21
.
.
,
[12,43Ж.
:
SaccharomвcОs ОrОvisiaО
,
[36Ж
Erwinia rhapontici,
,
300
[12].
,
,
,
.
Cheetham
[13Ж
,
.
,
.
:
,
,
,
(
MР2+
K+.
,
[36].
[57].
. RШЬОЯОКЫ [58,59Ж
,
.
.
)
22
,
,
. RШЬОЯОКЫ [58,59Ж
,
,
,
.
,
,
,
,
,
,
-
.
,
[52].
[45Ж.
.
,
. DО RШЬК
,
. [20]
Caldariella
.
acidophila,
–
[45Ж,
.
,
-
SКФТЦКО
OЧТЬСТ [60Ж.
,
,
,
,
,
(
,
-50
,
-
-70° ).
,
,
,
.
,
.
,
23
)
,
,
,
,
.
,
,
.
,
.
,
.
,
.
3.
3–
1.
1
,
2
3
,
;
;
2.
;
,
,
;
;
24
3.
3.
)
,
,
;
,
)
-
;
,
,
;
;
;
;
)
;
;
;
,
;
;
)
:
1.4
1.4.1
,
–
,
,
.
3,5
.
,
,
,
,
.
:
25
,
.
,
–
.
,
–
1960-
,
.
,
.
AЧЭЫТЦ
Bucke [10,11Ж
. [4Ж.
4
,
,
.
4 –
-
,
Bacillus
coagulans
Streptomyces
olivaceus
Streptomyces sp.
Hemmingsen
[28]
Borglum [8]
,
.
MКЫМШЧТ MШЫТЬТ
[45]
,
( . .
),
.
,
.
,
,
,
e
.
,
.
. [53],
Narita
,
Absidia,
-
,
. Arnaud
26
Bush
[5Ж
,
Penicillium duponti
,
.
: Marconi
,
15
Morisi [45]
,
.
,
,
,
(
)
.
,
,
,
,
.
,
35%
,
,
.
,
Serratia plymuthica, Protaminobacter ruber [64Ж
Erivinia rhapontici [12].
[64Ж
Д6Ж.
Erwinia rhapontici
[12Ж
.
5
,
Erwinia rhapontici,
.
27
5 –
Erwinia
rhapontici
*
(
)
-
.
*
,
0,325
0,583
0,13
0,153
8500
400
570
40
0,263
0,01
0,34
0,10
38
25
27
8
0,60
36
,
,
(550 / )
,
E.rhapontici
.
,
25 ,
,
.
L-
L-
,
.
(
Escherichia coli
).
,
,
,
,
,
24-48
E. coli
.
.
1
.
MРCХ2,
1
,
8,5,
,
28
120
37° .
,
,
:
15.
LBrevibacterium
-
ammoniagenes,
.
.
70%
L-
1
7,0.
,
80%
.
L-
-
,
,
.
L-
-
,
-
,
NОХЬШЧ [55Ж
Rhodotorula gracilis.
,
.
,
,
90%
.
,
L-
(
.
L-
(
Pseudomonas dacunhae), L-
L-
L(
Pseudomonas putida)
L-
-
Achromobacter liquidum).
,
L-
L,
L-
.
29
Nationale de la Recherche Agronomique [31Ж
Lactobacillus
,
Casei
10
,
L-
.
.
,
11-
,
△'-
.
Curvularia lunata [49Ж:
.
С.lunata
[56Ж.
,
,
,
,
.
,
.
(2%),
(1%)
.
Arthrobacter
(Corynebacterium)
simplex,
[1Ж.
,
.
31,7
0,17
A.simplex
/
/
,
,
,
.
.
,
,
,
. VОЧФКЭКЬЮЛЫКЦКЧТКЧ
Vieth Д86Ж
30
,
Mycobacterium
800
rhodochrous,
.
,
,
AФЭТОЛШХКРОЭ
Fermenta [1Ж,
CСООЭСКЦ [15].
. FЮЫЮСКЬСТ
, 1-
, 1-
. [22Ж
,
, 1-
,
1,15-
,
.
,
0,05%
.
Nocardia corallina,
(
115
/ )
N.butanica
N.paraffinica .
.
,
.
,
,
-6Achromobacter butyri,
.
,
.
+
+
Д48Ж.
Brevibacterium ammoniagenes,
Klibanov
Puglisi [38Ж
Alcaligenes eutrophus
,
,
+,
,
,
,
,
,
2,6.
31
.
1.4.2.
,
.
6
(
,
)
,
,
,
.
,
,
,
.
,
,
.
[36Ж
Saccharomyces
,
erevisiae,
.
,
,
.
,
,
[36].
Kluyveromyces marxianus
32
NКЯКЫЫШ
DЮЫКЧН [54Ж
Saccharomyces carlsbergensis
,
.
.
15
.
,
,
( . .
),
,
,
.
,
.
,
DКЮРЮХТЬ
,
. [19]
S.cerevisiae
.
,
,
120
/ ..
.
200
.
(
)
АКНК
3×106
,
. [68].
S.cerevisiae
:
5×106
,
.
-
,
,
100
3
/
Д69Ж.
,
.
,
.
[70Ж
,
250 / .
,
S.cerevisiae
-
,
33
,
,
,
,
100
100
250 / ,
100 /
8×109
(
,
.
-
).
250 /
,
150, 200
250 / ,
114
/
96%
2
,
,
.
,
10
,
, АКНК
. Д90Ж
.
0,86
0,67
:
,
.
2
. CСШ
. [17Ж
Saccharomyces cerevisiae,
.
6
,
,
,
6–
100 / .
[17] (
(Vt))
( /( × ))
7
14
10
21
34
:
2
65%
CO2. SСТШЭКЧТ
ВКЦКЧО [62Ж
,
.
АТХХТКЦЬ
MЮЧЧОМФО [71Ж
.
.
15 / .
(
4
3-8):
5.
,
.
7
.
1
Kierstan
Bucke [36]
2
S.cerevisiae
CК
Wada et al.
[88]
S.cerevisiae
GЫШЭО
[24]
.
Z.mobilis
3
(
(
(
EЭOH, /( × )
,D
,D
/
,%
,
7 –
100
4
80
5
0,1
6
Vt
7
4,3
N)
100
86
1,0
Vb
43
85
0.8
Vl
53
)
150
)
35
7.
1
GЫШЭО
[24]
2
Z.mobilis Ca
.
Linko
Linko [42]
АТХХТКЦЬ
Munnecke
[71]
MКЫРКЫТЭТЬ
. [47]
3
S.cerevisiae
CК
S.cerevisiae
CК
(
Z.mobilis Ca
Vt,
150
4
75
5
0,85
6
Vl
7
44
)
175
83
0,3
–
21
127
63
4,6
Vl
54
87
97
2,4
1,6
Vl
Vl
102
71
N)
100
; Vb,
,
Vb
.
. Vt
Vt
–
(127 / )
0,05
85%
53,8
/( × ),
CaCl2,
.
. LТЧФШ
Laminaria
.
; Vl,
Vl.
LТЧФШ [42Ж
Saccharomyces cerevisiae
LТЧФШ
-
LТЧФШ
80 / .
,
,
2
50
5×10 9
/( × ). Veliky
АТХХТКЦЬ [67]
.
,
. BТЫЧЛКЮЦ
. [7]
,
,
.
40-80%
.
)
[63] –
.
(
36
,
,
,
,
.
HШХМЛОЫР
Margalith [29].
,
Saccharomyces erevisiae,
,
.
-
,
,
,
,
.
.
,
,
5 / .
,
)
300 /
,
14,5% (
20° . HШХМЛОЫg
MКЫРКХТЭС Д29Ж
.
:
,
.
,
,
.
. LТЧФШ
. [43Ж
Kluyveromyces fragilis
. MКЫРКЫТЭТЬ
BКУpКТ [46Ж
K.fragilis,
. D-
,
,
,
.
. MКХОЬгФК
[44Ж
Pachysolen tannophilus
.
37
,
,
«
»
.
,
,
,
[36Ж,
. CСТКЧР
. [16Ж
S.cerevisiae
,
,
,
.
,
,
.
HКРОЫНКХ
MШЬЛКМС [25Ж
,
.
,
Zymomonas mobilis. Grote
. [24]
MКЫРКЫТЭТЬ
. AЦТЧ
,
Г.mobilis
Verachtert [3Ж
(
Saccharomyces bayanus,
,
. Д47Ж
)
,
Z.mobilis
.
,
Clostridium acetobutylicum,
HКРРЬЭЫШЦ
MШХТЧ [26Ж.
,
,
,
,
.
,
.
, KЫШЮаОХХ
. [41Ж
,
Clostridium butylicum,
.
,
38
,
Bacillus,
.
,
,
(
)
[40].
,
Bacillus amyloliquefaciens
-
[61Ж.
,
,
,
.
1
,
.
,
.
,
.
,
,
:
.
:
,
.
.
,
,
,
-
,
,
,
39
.
,
,
.
40
2.
2.1
:
–
.
Saccharomyces cerevisiae (
,
–
«
-
»,
);
–
(
–
«
», .
).
2.2
2.2.1
(
125
500
).
30
48-50° ,
– 55-58°
.(
.
).
.
5-10
,
. (
).
10-20%
,
.
( )
.
,
,
8.
41
8–
,%
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
22
24
26
28
30
0
999,9
1003,9
1008,0
1012,1
1016,2
1020,3
1024,5
1028,7
1032,9
1037,1
1041,4
1045,6
1049,9
1054,3
1058,7
1063,1
1067,5
1071,9
1076,4
1080,9
1085,5
1094,7
11104,0
1113,4
1123,0
1132,7
10
999,7
1004,7
1007,6
1011,6
1015,6
1019,6
1023,7
1027,7
1031,9
1036,0
1040,1
1044,4
1048,6
1052,9
1057,1
1061,5
1065,8
1070,2
1074,6
1079,1
1083,5
1092,6
1101,8
1111,1
1120,6
1130,1
15
999,1
1003,0
1006,9
1010,9
1014,8
1018,8
1022,9
1026,9
1031,0
1035,1
1039,3
1043,4
1047,6
1051,9
1056,1
1060,4
1064,7
1069,1
1073,5
1077,9
1082,3
1091,3
1100,5
1109,7
1119,1
1128,6
,
20
998,2
1002,1
1006,0
1009,9
1013,9
1017,9
1021,9
1025,9
1029,9
1034,0
1038,1
1042,3
1046,5
1050,7
1054,9
1059,2
1063,5
1067,8
1072,2
1076,5
1081,0
1089,9
1099,0
1108,2
1117,5
1127,0
25
997,1
1000,9
1004,8
1008,7
1012,7
1016,6
1020,6
1024,6
1028,6
1032,7
1037,8
1040,9
1045,1
1049,3
1053,5
1057,7
1062,0
1066,3
1070,6
1075,0
1079,4
1088,3
1097,3
1106,5
1115,8
1125,2
30
995,7
999,5
1003,4
1007,3
1011,2
1015,2
1019,2
1023,2
1027,2
1031,2
1035,3
1039,4
1043,4
1047,7
1051,9
1056,1
1060,4
1064,6
1069,0
1073,3
1077,7
1086,5
1095,5
1104,6
1113,9
1123,2
– 6-8%
115°
0,5
-
30
.
.
2%.
,
.
50-60
2.2.2
.
Saccharomyces cerevisiae
Saccharomyces cerevisiae
-
42
,
,
,
1.
1–
.
.
t = 30°C
-
5-7
.
,
.
30°C.
2.2.3
.
.
Saccharomyces cerevisiae
250
50
2
5
1500
/
,
0,9%-
NaCl.
.
43
2.2.4
.
.
(1:5000).
2-3
40 .
,
,
.
,
200.
2.2.5
.
(
–
2)
,
,
.
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( )
,
.
(
).
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(
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1/400
,
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.
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44
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.
.
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.
.
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).
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100).
ё
.
1
(1):
N
N–
a  1000
n
hS
(1)
1
;
a–
h–
S–
n–
;
,
;
,
2
;
.
45
2.2.6
,
:
–
2%-
.
500
10
.
,
;
–
:
1-
:
:
–1:1
2-
:
:
:
:
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:
ё
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:
ё
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–
ё
ё
10-
Х2.
5%-
ё
2+
.
з1
,
.
.
CaCl2
.
.
3
1%, 0,5%
0,9%2.2.7
.
,
0,25%
NaCl
.
46
.
10%1
.
,
,2
.
,
.
(8
1
20
).
(
8,89%).
:
50
1)
(
–
)
з 10
;
2)
15
,
;
25
3)
;
15
4)
5)
UNICO-1200
5
.
(
2,28 / .
3)
;
47
3–
( )
(m)
,
, m,
,
:
A = -0,1262m+2,2286
(2)
, С,
/ ,
:
С 
m–
,
N–
V–
m N
V
(3)
;
;
,
,
.
,
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.
48
2.2.8
ё
,
.
,
:
,
.
.
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,
.
,
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,
,
,
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.
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.
.
,
.
,
.
С, / ,
С =С
С
. .
–
, /
. .–
:
С
(4)
,
49
3.
3.1
Saccharomyces cerevisiae
-
4
Saccharomyces
cerevisiae.
,
,
,
;
– 4,5-5,0
.
,
.
4–
-
Saccharomyces cerevisiae,
5
Saccharomyces cerevisiae
-
.
50
5–
Saccharomyces cerevisiae,
,
,
.
1500
/
.
100
3.2
.
.
,
,
6.
51
6–
(
800)
,
,
,
–
.
3.3
ё
(
7–
(
800)
7).
52
10
.
– 249.
,
6,225×108
.
3.4
8
,
.
CaCl2,
,
.
8–
3
.
,
(
3
(
9-12).
)
53
9 –
1%-
-
Saccharomyces cerevisiae
10 –
0,5%-
Saccharomyces cerevisiae
54
11 –
0,25%-
Saccharomyces cerevisiae
12 –
-
55
3.5
-
.
9.
9–
1%
,
/
0,5%
0,25%
0
1,75
1
1,88
2
1,84
24
0,17
0
1,69
1
2
1,78 1,76
24
0,26
0
1,65
1
1,68
2
1,71
24
0,23
37,9
27,6
30,8
16,3
42,7
35,6 37,1
15,6
45,8
43,5
41,1
15,8
,
1%.
3.6
.
4
(
,
2
(
16,17,18).
13,14,15).
800
56
13 –
1%-
(
14 –
0,5%(
4)
-
4)
57
15 –
0,25%(
16 –
-
4)
,
1% (
800)
58
17 –
,
18 –
0,5% (
800)
0,25% (
800)
,
,
,
,
1%,
59
,
0,25%-
.
,
,
0,5%-
,
ё
,
.
– 1%.
3.7
Saccharomyces cerevisiae
19 –
19.
60
,
,
,
2
10.
10 –
1
№
1
2
2
3
4
1
2
3
4
5
6
7
24,0 34,0
38,0
40,0
26,0
35,0
37,0
40,0
50,0
54,0
50,0
14,2 18,2
22,5
23,3
17
20,2
29,3
32,8
38
45,5
37,2
9,8
15,5
16,7
9
14,8
7,7
7,2
12
8,5
12,8
, /
, /
, /
15,8
61
,
.
.
,
,
-
–
.
.
,
,
.
,
,
,
,
.
,
,
.
.
,
,
.
.
,
,
,
.
.
62
«
»
,
1 %.
.
,
(
4
).
63
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