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Progress in synthesis and regulation of bacterial cellulose

细菌纤维素的合成与调控进展



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ChineseJournalofBioprocessEngineering
Vol.9No.4
Jul.2011
doi:10.3969/j.issn.1672-3678.2011.04.014
OPQI
:2010-10-28
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(2005QO07)
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,Email:xinli2006@gmail.com
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:1672-3678(2011)04-0064-05
Progressinsynthesisandregulationofbacterialcelulose
LIXin,YANCailing,PANLinghong,HUANGJianzhong
(EngineeringResearchCenterofIndustrialMicrobiologyoftheMinistryofEducation,EngineeringResearchCenterof
FujianModernFermentationTechnology,FujianNormalUniversity,Fuzhou350108,China)
Abstract:Bacterialcelulose(BC)wasakindofnaturalhighpuritybiopolymercomparedwiththelignocelu
lose,theproductionandprocessingofBCwasmoreconvenientandenvironmentfriendly,thusitmadeBCbe
apromisingbiomaterial.Atpresent,GluconacetobacterwasfoundtobethehighestyieldBCproducingstrain.
Inthisreview,thesynthesisandtheregulationmechanismofGluconacetobacterBC,thegeneticalengineering
methodsandculturemethodsforhigherBCproductionwerediscussed.
Keywords:bacterialcelulose;Gluconacetobacter;regulation
  
´L?@A
(bacterialcelulose,¸
É
BC)
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,1886

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[1]。
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,´
L?@Azq¬*³çØÙTÀ
、¯
°À

€
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׶†S´Á78Cy

(Ø*+^ÄæÙG
ƒ³

ÿ³›+6¾¿.¶ùÔMB+6…;
[2]。
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l´L?@Agçi³6WÀÝ:–Ê`Ë

¾Ü
,µÍpý

VW

1»›Gw³6´L?@AR
3›,ešº?@A6*³Íp]å

1 
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E
;´
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ap,
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<
ðÛ6?@A

E
[3]。
g´L?@A6pª*¶.
,´
L?@AA
S?@ú¬Q6Œβ D 5IÃβ 1,4 
5oa‹µ¶ŸþŒ¶

ŸþüÜWìT

Q“ö
ðŒÍ

p拌

—Éùβ 1,4 ^5。
·AS?@A`Ë
,´
L?@AÝãM6
.MN
:1)
Ý!‹‡Í

!^µÍ!hÍ


}=yA½?@A
;2)
?@Ÿ3}
001~01μm


Ž’‡9Í!

…15×1010Pa,
}ðÝ]@Ý:–Êm6€!6

Òm?@A6‡
ˆN5·R3?@A612›àñfC6œ›
,

>?…

Â0

B
;3)
ÝZ96™¼N
D¼DAN

5€®!çg¿y~
60~700
±6¼
p
;4)
ÝT!6RSɍN

›U6RSá0a
N

QÀÁžŸ
[4]。
2 
Xȱ²‚cË۟p
21 
`Xȱ²‚cÈß
Ñ4´LQ5jR3?@A

i%6ݏ5`
\|Lß
(Gluconacetobacter)、
1FLß
(Rhizobi
um)、
ÝÞrLß
(Sarcina)、
O÷ÁLß
(Preudo
monas)、
Öß|Lß
(Agrobaeterium)、
·|Lß
(Achromobacrer)、
öELß
(Azotobacter)、
A|Lß
(Aerobacter)
3ÐLß
(Alcaligenes)。
5`\|

(Gluconacetobacter)
m6
Gxylinus
€SG1I

VW¶ù¶·ùÕ¾¿TùDC6?@A3RL

8áñY³4ûbSºTR:

6Íszòµ¶?
@A5‡€96¥RSLû
[5]。
22 
Xȱ²‚cŸp$%
zb<ƒN´L5`\|Lß6Íse´L?
@Aµ¶:b¾¿€46L
(Gluconacetobacter)。
¾¿
z›o­øúϵ¶¸¹5`\|LL…6´L
?@ACV

îï

ðñ

Ε

áò

IôÁCº
«´Á65G5‘B>`.¶

>`
5
;6
>`5G5>`¤þBZŒ.ù

>`
5
;1
>`5Gà>`.BÁ.R¶ço]
>`5
(UDPGlc),
€f?@AµBÁ.
UDPGlc
4^.¢¶?@A

Íse
GhanseniATCC23769
SCV*6²L‹
í,ñ

VL…"
3351
ëCV

gm
3308
ëù#$y
&x
,43
ëù
tRNA
CV
,2
ëù
rRNA

[6]。
·ñ
h6¾¿‹íúS
[7],G.xylinus
m6?@AµBCV
·gçûëCV×}çÓúëÐªm
,G.hanseni
ATCC23769

G.xylinusATCC53582
6
acs
Ðªm
>"

ëCV
:acsAB、acsC

acsD[8-9],
lg
G.xylinus
13063

BRP2001
mpqµL6
bcs
Ðª>"

ë
CV
:bcsA、bcsB、bcsC

bcsD[7,10]。
ÍsÂù
G.xylinus
13063
6
bcsA

bcsB
CV&x?@AµB6

ëŸ
÷þ
[7,11],
l
G.hanseniATCC23769

G.xylinusATCC
FHK—
í5á5‘B
;GHK—
5á5‘B

GDH—
5ÙeB
;GADH—
5`ÙeB

PGI—
>`5ZŒB
;PGM—
>`5¤þB

UGP—
5à>`.B
;CSEs—
?@Aµ¶B

ASEs—
rêtµ¶B
m
1 
ŠÕÖb¾zbT&

Ë`
Xȱ²‚©,«µ
[4]
Fig.1 SimplifiedschemeofBCandacetamideproduction
usingglucoseorfructoseasthecarbonsource[4]
53582
6֑CV
acsAB
&xo?@AµB
[8]。bcsA

bcsB
4=p­·
acsAB
4=6



cÓF

acsC/bcsC
645ÍsùQ°ø
,bcsC
B·o;Ôβ
1,4
^56µ¶

h6};–wxmOQå86

[12];bcsD
B·o?@Aþ6‹‡.ÃÄ

Ââ
bcsD
OÈ:S;Ô?@Aµ¶¬Ð 0
40%[7];
‡;
‹Œ,ñ
bcsD
¢¶oúëÝ

ëÔcIj6Ý^
;
[13-14],
}
G.hanseniATCC23769
6CV*mù1I
o?–

ë
acsAB

acsC
CV

h6HëCV*m
1Iúë
acsD
CV
[6]。
}
bcs/acs
Ðª6„3

×}?úëÐª

8>"

ëCV
cmcax

ccpAx[9,15]。cmcax
&x
Ô β1,4 ^5B(endo β 1,4 glucanase,
CMCax),
¾¿1I
cmcax
}´L?@A6¢¶mŸ
^7å6ø³

8
cmcax
6UNG
?@A6
¢¶G7[16];
8
G.xylinus
};ÔÃ~Té
û
,cmcax
CV5cp\!?@A63~
[17]。ccpAx
&x?@Að@#$
(celulosecomplementingpro
tein,CcpAx),
8e?@A6µ¶Íp7å

86ë
CVNUf
,G.xylinus
ãNoµ¶?@A65
‡
[15]。
¾¿1I
CcpAx
B·o´L?@A!ÍÝ
L‹Œ6ŒÛ
,G.xylinusBPR2001
ᢶäå


·QáÚ6?@A

ÔC

lg
ccpAx
CVGæö
56 
!

# x
 
ÕG
:´
L?@A6µ¶·¸¹º»
6L…á3RøÔ

½·mnç6ÔC

8M6?
@A
I、
?@A

¼¢?@A6ŠµS
[18]。
?
@AµBÐª 3×}úë
bglxA
CV

8&x
Ý β 5oB(βglucosidase,G3ase)UN6
#$
[19]。cmcax

bglxA
6×}TL´L?@A6
µ¶×}^ú뵶0a6†…ÃÄ

23 
Xȱ²‚Ÿpc#z{[
G.xylinus
6?@Aµ¶K
cdiGMP
6¸¹

cdiGMP
øù´LI³6!]F³
,¸
¹^èîR
SC¢¶
、´
Á.†

Á–4563Rñf4´Á
_ü6`ðø³
[20]。Ryjenkov
G
[21-22]
6¾¿1
I
:bcsA
4=„6
PilZ
‹Œ†á56
cdiGMP
6
‹µþ;
,cdiGMP
I˵L?@AµB
bcsA
„
J¸¹?@A63R
。G.xylinus´
ÁÔ
cdiGMP
ÓÍIÃ"
GGDEF
‹Œ†6]éo`ž.B
(diguanylatecyclase,DGC)
"
EAL
‹Œ†6>`
]”B
A(phosphodiesteraseA,PDEA)
6`ÿø³
J@™
。DGC
Á.±pª6
GTP
¢¶
cdiGMP,
l
PDEA
·
PDEB
ú‡Á.
cdiGMP
0a¶
5′
GMP[23]。Christen
G
[24]
¾¿1I

¿~4©
DGC
#$6
GGDEF
‹Œ†ÔcQ×}^
cdiGMP
­Œ
‹µþ;
(Isite),
Vþ;}
cdiGMP
F­ê
´Áâ…ÃÄmŸ^€1é6ø³

8
cdiGMP
‹
µL]éo`ž.B
DGC
6
Isite
þ;û
,DGC
6
BUGh ¡Nóÿë
}
G.xylinus
m
,dgc
CV
pdea
CVþçÓú
ë
cdg
Ðªm

gm
pdea
þç
dgc
6„3

m
G.xylinus
×}

ë!ÍÓF6
cdg
Ðª
———
cdg1、cdg2

cdg3,
gm
cdg1
ìõo
80%
6´Á
DGC

PDEA
UN

l
cdg2

cdg3
p­ìõo
15%

5%
6UN
[25]。
3 
ä¥Xȱ²‚`‡cxZ
31 
K2¢xZ
ñ5í5ù


,G.xylinus
äå3S
ÒQ6?@A

l65H`

ï
1)。
5H
`È0k78_6
pH,
gl:S´Á6R:K»

ºú9:S?@A63~ 0
。Shigematsu
G
[26]
IÃCVæö ¡ŒÛo
G.xylinusBPR2001

5ÙeBÂíEL…
GDI,
VL…Q3R5
`

}ݏ9: ñ5ù

F

ª.5ù
?@A65‡Ëè•L…\!o
17
±

G.xylinus
Qh5pîh¼Ú6?@A

Óûù
5pîrêt
———
úû4€6¼ÚN45


t63R©ªo?@Aµ¶6s;
———
ço]>
`5
(UDPGlc)(
ï
1)。Ishida
G
[27]
g
G.xyli
nusBPR2001
L…ø1µLo

…rêtÂíE„
¤…
EP1。
Ül·#í6`ÿ
,EP1
·è•L…
G.xylinusBPR2001`
Ë

lÜzð9: 63~·
è•L…~™úS

WX9: 6?@A3~ÿl
0ko
。EP1
78_¤¶Q[‹6Š§_

gm>
"o~cp6´Á?@A^¯S

6SLá56
Œçrêtl8Âo78_64N

»Õo´Á
?@A6S‹

gl\!o?@A3~

}
EP1
6
78ÃÄm«rêtf

g?@A3~ïÒLo
è•L…6¼ì

8«ك5û

˟Lorê
t6Þí
[28]。
VW¨80k78_6؆NË6
\!´L?@A3~6úûÎ3

!GASm×}

뢶ço]>`5
(UDPGlc)
6?@Aµ¶C)

<|5µBC)
UD
PG
à>`.BC)

l´LmÝfúû
。Nakai
G
[29]
}
A.xylinumsubsp.sucrofermentansBPR2001
mT
éoY˜6|5µB

VB5jÁ.|5ço]
>`
(UDP)
¢¶ço]>`5
(UDPGlc)
í
56á·ÿ*

ñ
G.xylinus
Té|5µB

QK5
jY³,ù›G6

F|5µ¶?@A

lm5j
\!?@A63~

}
G.xylinus
m*¶ETéDLðLß2[#
$
(VHb)
CV

5:S´LR:¬Í²
50%,
VHb+
}zð789: 6?@A3~Ëè•L…
\!o

±

ºú9¾¿1I}zð789: 0
kA`m6Aˆ‡áñ,~[Íó\!
VHb+
?@A63~
[30-31]。
öo
G.xylinus,
gç´LËݵ¶?@A6

,Nobles
G
[32]
tjo¤
G.xylinus
6cp?@
Aµ¶BÐª
(acsABΔC),}÷´Áj« Syne
chococcusleopoliensisUTCC100
L…mºTTé

³
µj«5jµ¶h‡¹6?@A

6Ï3SRÑ
áñøùR3RS‰i6è–Ê

Q—åàR
3L…lmáñY³,ùFn786L…JR3
ð6´L?@A3Ù

IÃWH^Ä/¯

³/¯f6
Gxylinus
}
gaµ¶?@A6ÃÄmñ«

rê5,
(Glc
NAc)
ÈC

qµ6?@AûIyO^SQKÝ´
L?@A›U6RS`FN‡ˆN5

lmaG
o?@AËñ0a6pÌ

?@AûIyO^S
66
R
 
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Â
 
^
 
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Ä
  
!

"
 
FnGÚLB?@AB¼a

}š};Ô6wx
‹í1I
:20d
ÔO^SáñGRS0a

³?@
Aqµo,U6*³st
[33]。
32 
de23
öoY³CV^Ä/¯L…–

ùÝÑ4}7
8^_„ºTòœ6tj
。Park
G
[34]
1I

}CÔ
78CmJÂ
1%
6rJáñ45Q3?@A6
G.hanseni
6´Á©Í

³?@A63~\!
17
±
。Bae
G
[35]
Y³5~78C}šE12†mR
3?@A

1I³
H2SO4Yυœ65~78C
ËQœ65~78C6?@A3~\!o
76%,
lmTk65~ÓÍe!3?@AÍp7å
。Seto
G
[36]
1IgBÖßmpqµL63?@AL
|

áñ‘›ø>"
G.xylinus
óí¶|L
Lacto
bacilusmali
6

û´L

ÒmO78

û´LµL
6?@A3~6֋78
G.xylinus
6

±

6ùI
ôÁ_ü6`ðø³\!?@A3~\ˆou
6;2
。Hu
G
[37]
}
G.xylinus
}~78mJÂA
SR:
‰CžG?
(1methylcyclopro
pene,1MCP),
1I·e—`Ë

lÜ´Á6R:K
Loú¼
h6?@A63~8Âo
254%。
öoe78CºT-.–

QÓ678›àË
È})´L?@A63~My

z…7812
i#:

«ó+¨~

ô†9Í!

Q¨µ~op6
^À.R3

l†…125ÝÞ\!R3ÞÐ

0
kR3¶é

ÒmRSÿ*¯m6RSµ¶ô†9
ÍË`eTk

I}zÝ4ûQÓxE6ÿ*¯
G*³ç†…78

7îWX

}~õ

A0àÿ*
¯

ªöÿ*¯/º6A0RSÿ*¯
[38]。
4 
*
´L?@A3À}è

ìG1éí‘z”9¢
¶$3H„˜ì26 Ú

Òzº«ØÙ

ƒÿ

®
+
、¯
°

.^

i

|÷GTÀ

hÍs´L?
@A6›GáTNùäå]Gç?@A6R3Ð

yzµ6ÿ*¯

îA0ÿ*¯}~ÿ*¯

¤±v‹µ

³g/786›àRÑ5aG6ë


?úëaG6Î36*³CV^Ä/¯G
|° ¡J\!3~

îãj´L?@AR3L…
63?@Aûü

òí´L?@A3~k

R3¶é!

äå`
a‹ž612¼ìk

,wI3À.

VW

 
ú96^ø*‹µCV^č!íÍ78G ¡
J\!´L?@A612µÐ

ÓûÂ9´L?@
Aµ¶†‡ˆ6¾¿

yzµ6RSÿ*¯

Ò
ºú9e´L?@A6·ø

*+‹ŒGºT¾
¿

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42.*³

M¬6q

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76 
!

# x
 
ÕG
:´
L?@A6µ¶·¸¹º»
[15] StandalR,IversenTG,CoucheronDH,etal.Anewgenere
quiredforceluloseproductionandageneencodingcelulolytic
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