B CNG THNG

TRNG I HC CNG NGHIP TP. HCM

VIN KHOA HC CNG NGH & QUN L MI TRNG

N C S NGNH

C IM TNH CHT V NG DNG CA VI KHUN

KH SULPHATE

Ging vin hng dn: Nguyn Khnh Hong

Lp: DHKTMT10A
Sinh vin thc hin:
Phm Th Tm
H Th Bo Trm

14116941

Nguyn Th H Tin

14105071

Phm Th Bi Tuyn

14107601

L Qu Tuyn

14079351

TP. H Ch Minh, 2016

LI NHN XT CA GIO VIN HNG DN

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LI NHN XT CA GIO VIN PHN BIN
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LI CM N
Nhm chng ti xin chn thnh cm n n:
Ban gim hiu Trng H Cng Nghip TP.HCM ni chung v Vin Khoa Hc
Cng Ngh v Qun L Mi Trng ni ring to iu kin cho nhm chng ti lm
nn bi n bo co c s ny. GVHD thy Nguyn Khnh Hong tn tnh dy
d, gip , hng dn chng ti tm hiu r b mn n C S Ngnh. Th vin
Trng H Cng Nghip to iu kin tt nht cho chng ti tra cu ti liu lin
quan. Phng a phng tin H Cng Nghip Tp.HCM vi h thng my tnh hot
ng lin tc to iu kin cho chng ti tm kim thng tin nhanh chng. ng
cm n s gp ca tt c cc bn cung lp tn tnh gp , tham gia kin
nhm chng ti hon thnh tt bi tiu lun.
Chng ti xin chn thnh cm n!

PH LC
Chng 1: Tng quan.................................................................................................................1
1.1. Tng quan v vn c nghin cu.....................................................................1
1.2. Nhim v n...........................................................................................................4
1.3. Cu trc ti............................................................................................................5
Chng 2: NI DUNG...............................................................................................................6
2.1 c im......................................................................................................................6
2.1.1
c im hnh thi................................................................................................6
2.1.2
Nhu cu dinh dng ca SRB...............................................................................9
2.1.3
Mt s v d v hnh nh ca vi khun kh sulfate.............................................11
2.2 Tnh cht...................................................................................................................11
2.2.1
Qu trnh trao i cht v nng lng...............................................................11
2.2.1
Vn v mi......................................................................................................15
2.2.2
n mn ng ng..............................................................................................16
2.2.3
ng dng...............................................................................................................17
2.2.1
B sung sulfate tang cng s phn huy rc ti cc bi chn lp................17
3.2.2
X l rc thi giy...............................................................................................18
3.2.3
Sn xut ch phm vi sinh...................................................................................18
3.2.4 ng dng ca vi khun kh sulphate trong x l nhim du kh......................19
3.2.5
ng dng trong oxy ha cc khong vt sulfur.................................................21
Chng 3: KT LUN V KIN NGH................................................................................24
Ti liu tham kho....................................................................................................................26

DANH SCH BNG BIU

Bng 2.1. Cc c cht m vi khun kh sunphat

s dng trong sinh trng
Bng 2.2. c im mt s vi khun chnh c trong cc m qung
DANH SCH HNH V

Hnh 2.1. V tr ca SRB trong chu trnh cacbon v lu hunh.

Hnh 2.2. Cc bc kh sulfate SRB v cc enzyme tham gia
Hnh 2.3. hnh nh vi khun aceterbaxtor.
Hnh 2.4. Hnh nh vi khun desulfovibrio sp.
Hnh 2.5. S m hnh th nghim nghin cu tun hon nc r bi chn rc

Hnh 2.6 : C ch phn gii ankan

DANH SCH CC T VIT TT
SRB (Sulfate reducing bacteria) : Vi khun kh sunphat

Chng 1: Tng quan

1.1.

Tng quan v vn c nghin cu

Vi khun kh sunfate l nhm vi khun rt a dang c kh nng s dng sunfate

lm cht nhn in t cui cng. Chng l nhng vi khun k kh bt buc phn b kh
rng trn Tri t. Chng c trong t, trong cc vng trm tch, bn lng cc y ao
t, cng rnh, sng h, bin trong cc vng c iu kin sng khc nghit nh: p sut
cao, nhit cao, mn cao, hai mi trng qu kim, qu axit v c bit tn ti vi
mt s lng ln trong cc du m. S a dng cao ca vi khun kh sunfate c tm
thy cc trm tch bin, ni c nng sunfate cao.
Trong nhng cng trnh nghin cu u tin v vi khun kh sunfat ngi ta
ngh rng vi khun ny l vi khun k kh tuyt i. Tuy nhin khong 20 nm tr li
y quan im ny thay i. Cc nh khoa hc ch ra rng vi khun kh sunfat
c kh nng chng chi c vi oxy. Theo bo co ca Cypionka (2000), vi khun
kh sunfat khng ch sng c trong mi trng c oxy nhiu gi m thm ch nhiu
ngy v nhiu loi thm ch c th kh c oxy thanh nc. Qa trnh ny c th
thc s l qu trnh h hp khi kt hp vi qu trnh chuyn ha nng lng. Ngi ta
pht hin thy mt s lng ln vi khun kh sunfat trong cc vng hiu kh v
vng ranh gii gia hiu kh v k kh ca cc trm tch.

Hu ht cc loi vi khun

kh sunfatphn lp c t nhng v tr hiu kh u thuc hai chi Desulfovibrio v

Desulfomicrobium. Theo Cord- Ruwisch v cng s, kh nng sng trong mi trng
c oxy ca vi khun kh sunfat c th c gii thch nh sau: Trc ht h hp ca
cc vi khun hiu kh s tiu th oxy to iu kin thun li cho qu trnh trao i
cht ca vi khun kh sunfat. ng thi,

S do vi khun kh sunfat to ra l mt cht

kh, n s tc dng vi oxy iu kin thng lm gim nng oxy khu vc .

Mc d c th thch nghi vi mi trng, chng c th chng chi vi oxy. Tuy nhin,
do vi khun ny thng s dng cc sn phm ln men c to thnh trong iu kin
k kh nn chng khng th c nhng thun li trong qu trnh cnh tranh vi vi sinh

vt khc pht trin trong mi trng hiu kh. Do vy, vi khun kh sunfat vn l vi
khun k kh.
Trong mi trng hiu kh, vi sinh vt d dng c th thc hin qu trnh chuyn
ha hon ton hp cht hu c thnh

. Tuy nhin trong h sinh thi k kh th s

chuyn ha li phc tp hn v i hi s tng tc ca mt lng ln cc nhm vi

khun khc nhau. Mi nhm vi khun thc hin mt qu trnh oxy ha mt phn hp
cht hu c v sn phm chuyn ha cui cng ca chng li c ng ha bi cc
thnh tip theo ca chui thc n cho n khi kt thc qu trnh oxy ha. Vi khun kh
sunfat trong iu kin k kh thng c nghin cu t trong mi quan h vi hai
nhm vi khun k kh khc nhau l vi khun metan v vi khun kh nitrat. Cc cng
trnh nghin cu trn th gii v ti Vit Nam cho thy c mi quan h hi sinh ca
vi khun kh sunfat v vi khun sinh metan, nht l trong iu kin nc bin v nc
v du m. R rng l vi khun kh sunfat sng hi sinh vi vi khun sinh metan, kh
khng hon ton cc hp cht hu c c sn trong mi trng thnh axetat v chnh
ngun axetat ny li l c cht cho cc vi khun sinh metan v cc vi khun s dng
axetat sinh trng pht trin. Cn mi quan h ca vi khun kh sunfat v vi khun
kh nitrat ang dc cc nh nghin cu rt quan tm v c tnh ng dng cao ca n.
Lysnes gn y pht hin ra chng kh nitrat a nhit phn lp t ging du
vng bin Bc s dng c cht axit bo li c kh nng sing trng trn nhng ngun
c cht a thch ca vi khun kh sunfat. iu ny to c s m ra hng ng dng
mi: s dng nhm vi khun kh nitrat cho vic c ch s pht trin ca vi khun kh
sunfat trong cc ging khoan bng phng php sinh hc thay cho ha hc nh hin
nay.

Vn gy ra bi vi khun kh sulfate:

Trong k thut, vi khun kh sulfate c th to ra cc vn khi cu kin kim loi tip

xc vi nc mui cha sulfate : Tng tc ca nc v kim loi to ra mt lp phn
t hydro trn b mt kim loi; vi khun kh sunfat sau kh hydro to ra hydrogen
sulfide, gp phn vo s n mn .

Hydrogen sulfide t vi khun kh sunfat cng ng mt vai tr trong s n mn

sulfide hu c ca b tng. N cng xy ra trong du th chua.
Mt s vi khun kh sunfat ng mt vai tr trong qu trnh oxy ha k kh metan.
CH 4 + SO 4 2- HCO3- + HS - + H 2 O
Mt phn quan trng ca metan hnh thnh bi methanogen di y bin b oxy
ha bi cc vi khun kh sunfat vng chuyn tip tch methanogenesis t cc hot
ng gim sulfate trong trm tch. Qu trnh ny cng c coi l mt bn ra chn
ln cho sulfate trong trm tch bin.
Trong thy lc ct ph cht lng c s dng frack phin hnh thu hi
kh metan (kh phin) v cc hydrocacbon, cht dit khun cc hp cht thng
c thm vo nc c ch hot ng ca vi khun kh sunfat, trnh qu trnh
oxy ha metan k kh v th h ca hydrogen sulfide. Cui cng kt qu l gim thiu
tn tht tim nng sn xut.

Tm quan trng v nh du

Sunfat xy ra rng ri trong nc bin, trm tch, hoc nc giu ang phn hy
cht hu c. Vi khun kh sunfat kh ph bin mi trng k kh l vi khun h tr
trong vic phn hy cc cht hu c.
Trong nhng mi trng ym kh, vi khun ln men ly nng lng t cc phn
t hu c ln; kt qu cc hp cht nh hn nh axit hu c v ru c tip tc b
oxy ha bi acetogens v methanogen v cnh tranh vi khun kh sunfat.
Bn thi t ao; mu en l do cc sunfua kim loi, l kt qu ca trnh phn
hy cc kim loi ca vi khun kh sunfat.
Cc c hydrogen sulfide l mt sn phm cht thi ca vi khun kh sunfat; mi
trng thi ca n cho ta thy l mt du hiu cho s hin din ca vi khun kh
sunfat trong t nhin.

1.2 Nhim v n

Tnh cp thit ca ti

Hin nay vi khun kh sulfate v ang c ng dng nhiu trong vn x

l rc thi, nc thi,.... v trong tng lai.

L do hnh thnh ti

Do ti do nhm chng em thy y l mt ti c ng dng nhiu trong

cuc sng v x l mi trng

i tng v phm vi nghin cu

i tng nghin cu ca ti l vi khun kh sulfate, phm vi nghin cu l

nghin cu v c im tnh cht v ng dng ca vi khun kh sulfate trong thc tin
v tng lai.

Phng php nghin cu

Phng php nghin cu: Nghin cu thng qua ti liu sch bo, cc bo co
khoa hc cc ti d n v ang trin khai, cc th nghim v kt qu nghin cu
ca cc nh khoa hc trong nc v trn th gii.

ngha khoa hc v thc tin

Khi khoa hc pht trin chng ta bit r hn v vai tr ca vi khun kh sulfate,

th vic ng dng trong i sng hng ngy v trong i sng sn xut cng nh trong
cng nghip v hng n tng lai.

Gii hn ti

V mt ni dung ti ch yu lm r c im, tnh cht v ng dng ca vi

khun kh sulfate trn c s l thuyt, cc chng minh khoa hc v cc bo co th
nghim lin quan ti ti.

1.3 Cu trc ti
Chng 1: TNG QUAN
1. Tng quan v vn c nghin cu
2. Nhim v n
3. Cu trc ti
Chng 2: Ni dung
1. c im
1.1.
1.2.
2. Tnh cht
2.1.
2.2.
2.3.
3. ng dng
3.1.

c im hnh thi
c dim sinh l sinh ha
Qa trnh trao i cht v nng lng
Vn v mi
n mn ng ng
B sung sulfate tang cng s phn huy rc ti cc bi
chn lp

3.2.

X l rc thi

3.3.

Sn xut ch phm vi sinh

3.4.

ng dng ca vi khun kh sulphate trong x l nhim

du kh

3.5.

ng dng trong oxy ha cc khong vt sulfur

Chng 3: Tng kt

Chng 2: NI DUNG
2.1c im
2.1.1 c im hnh thi
Vi khun kh sunfat l nhm vi khun c sinh l phc tp nhng c c im
ni bt l chng s dng sunfat

lm cht nhn in t cui cng. Cc loi vi

khun kh sunfat khc nhau th hin hnh thi t bo, kh nng di ng, kh nng
hnh thnh bo t, hm lng GC cha trong ADN, desulfovibrin, h thng
cytochrome, nhit ti u cho sinh trng v kh nng trao i cht (oxy hon ton
hay khng hon ton c cht) .
Nhm vi khun kh sunfat c hnh dng t bo rt a dng ty theo tng chi
( vi kch thc t bo khong 0.3 1.5

m x 0.8 10 m ): Hnh que nh

Desulfobulbus, Desulfotomaculum, Thermodesulfobacterium, Thermodesulfohabdus,

Thermodesulfatator; hnh du phy nh Desulfovibiro, Thermodesulfovibrio; hnh
ovan nh Desulfobacter, Desulfobacterium; hnh cu nh Desulfococcus; hnh ti
Desulfosarcina; hnh si nh Desulfonema.
Trong t nhin, vi khun kh sunfat bi t nhin hoc bm vo gi th no ,
nht l cc mu st, ng st. Ty loi m chng c th ng n c hay kt i, hoc
xp chui ngn, cng c khi chng tp hp la thnh mt m ln xn .
Mt s loi vi khun kh sunfat chuyn ng nh tim mao hoc mt chm
tim mao, trong khi mt s khc chi di ng trong giai on u ca qu trnh pht
trin sau kh nng gim dn v mt hn. Mt s khc li khng c kh nng di
chuyn.
Cho n thi im ny, c rt nhiu loi vi khun kh sunfat c pht hin
v phn loi. Vi s pht trin ca k thut phn tch trnh t gen 16S rARN kt hp
vi cc c im sinh l trn xy dng c cy pht sinh loi ca vi khun ny.
Cy pht sinh loi cho thy s a dng rt cao s lng loi ca nhm vi khun kh
sunfat.
6

Da trn trnh t phn tch rARN, vi khun kh sunfat c chia thnh bn

nhm chnh: vi khun kh sunfat Gram m a m; vi khun kh sunfat Gram dng
sinh bo t; vi khun kh sunfat a nhit v vi khun kh sunfat c a nhit.2.1.1 c
im sinh l, sinh ha
Cc vi khun kh sunfat c coi l mt nhm kiu hnh, cng vi cc vi
khun kh lu hunh khc, cho mc ch nhn dng. Chng c tm thy trong mt
vi dng pht sinh loi khc nhau. Tnh n nm 2009, 60 chi c cha 220 loi vi
khun kh sunfat c bit n.
Nhm vi khun kh sunfat Gram m a m gm hai h ln l
Desulfovibrioaceae (gm hai chi Desulfovibrio v Desulfomicrobium) v
Desulfobacteriaceae. H Desulfobacteriaceae phong ph v s chi hn vi khong
gn 23 chi nh Desulfobulbus, Desulfobacter, Desulfobotulus, Desulfobacterium... .
iu th v ca h Desulfobacteriaceae l kh nng hnh thnh cm vi khun
gip chng li nhng iu kin bt li ca mi trng nh quan st thy
Desulfosarcina hay kh nng di ng cho php chng di chuyn n nhng vng c
nhiu cht dinh dng hn.
Ngc li vi nhm vi khun kh sunfat Gram m a m, nhm vi khun kh
sunfat Gram dng sinh bo t c mt s loi thch nghi c vi mi trng sng c
s thay i nhit t m ln cao do chng c c ch hnh thnh bo. Tuy vy, nhit
ti u cho sinh trng ca nhm vi khun ny l t 25 40 , mt vi loi c th l
t 40 -65

. n nay chng ta mi ch khm ph ra mt chi Gram dng c kh nng

sinh bo t duy nht l chi Desulfotomaculum. Vi kh nng hnh thnh bo t khi gp

iu kin sng bt li nn chi Desulfotomaculum thng c tm thy trong cc mi
trng sng m khi bt gp cc chi vi khun kh sunfat a m khc (nh cc
cnh ng la nc ni c s xen k ca iu kin c oxy v thiu oxy do s thay i
ca ma nc ln.
Hai nhm vi khun kh sunfat cn li l nhm vi khun kh sunfat a nhit v
nhm vi khun c a nhit (Archaea) u c phn lp ti cc mi trng c nhit
cao nh cc ging khoan su di mt t c nhit ln ti 80 . Cc vi khun kh
7

sunfat a nhit ch y l vi khun Gram m, thuc cc chi Thermodesulfovibrio. Mc

d nhm ny c nhit ti u cho sinh trng 55 - 75

nhng chng vn c th

sng dc nhit thp hn. Cc loi thuc chi Archaeoglobus, Caldivirga v

Thermocladium (u c phn lp ti cc h thng bin nng) l nhng i din duy
nht ca vi khun kh sunfat trong nhm vi khun c Archaea.
Vi khun kh sulfate (SRB) l cc vi khun sinh trng k kh, s dng sulfate
lm cht nhn in t cui cng oxy ha hydro hay cc hp cht hu c v tn thu
nng lng cho mc ch sinh trng (phn ng 1.10).
2CH2O + SO42- + H+
H2S + Me2+

H2S + 2HCO3-

MeS + 2H+

(1.10)
(1.11)

SRB ph bin trong mi trng k kh, ni chng c vai tr quan trng trong c
chu trnh lu hunh v chu trnh cacbon (hnh 2.1).

Hnh 2.1. V tr ca SRB trong chu trnh cacbon v lu hunh.

2.1.2 Nhu cu dinh dng ca SRB

Hu ht SRB c nhu cu dinh dng n gin v sinh trng tt trong mi
trng c ngun cacbon/nng lng n nh. Ngun cacbon v in t thch hp i
vi SRB bao gm cc axit hu c mch ngn nh acetate, lactate, pyruvate v ru.
Ph thuc vo cch oxy ha cht hu c m SRB c th c phn chia thnh
hai nhm trao i cht nh sau:
-

Nhm oxy ha khng hon ton: oxy ha cc hp cht hu c n acetate.

Thuc nhm ny ch yu l cc loi thuc chi Desulfovibrio spp.

Nhm oxy ha hon ton: Oxy ha cc hp cht hu c (bao gm c

acetate) hon ton thnh CO2.. Trong nhm ny c a dng cc loi SRB khc
nhau, nh Desulfobacter spp., Desulfobaterium spp., Desulfosarcina spp.
SRB thc hin trao i cht oxy ha cc c cht hu c s dng sulfate lm

cht nhn in t cui cng. S kh sulfate thnh sulfide tiu th 8 in t v cc qu

trnh sinh ha thng qua nhiu bc trung gian vi s tham gia ca nhiu enzyme .

Hnh 2.2. Cc bc kh sulfate SRB v cc enzyme tham gia

Phn ng c th c tm tt nh sau:
SO42- SO32- HSO3- HS- S2-

10

Bng 2.1. Cc c cht m vi khun kh sunphat s dng trong sinh trng:

Loi hp cht
Cht v c
Axit monocarboxylic

Cc hp cht c s dng
;
Focmat;axetat; propionat; butyrat; isobutyrat; 2-

Axit monocarboxylic

metylbutyrat; 3-metylbutyrat; 3-metylvalerat; pyrurat; lacrat

Succinal; fumarat; malat; oxalat; malenat; glutarat
Metanol; etanol; propanol; butanol; pentanol; isobutanol;

Ru

etylen glycol (mono-,di-, v tetra); 1,2-propandiol; 1,3-

Axit amin

propandiol; glyxerol
Glyxin; serin; alanin; cystein;cystin; treonin; valin; leuxin;

Cacbonhydrat
Hydrocacbon

isoleuxin; aspartat; glutamat; phenyl alanin

Fructoz; glucoz; mannoz; xyloza; rhamnoz; tinh bt
Hexadecan; hexadecanoat; benzoat; 2-,3-, v 4hydroxybenzoat; phenol; p-cresol;catechol; resorcinol;
hydroquinon; phlorglucinol; pyrogallol; axit nicotinic;
indol; quiolin; phenylacet; naphthalen, 2-methylnaphthalen;

Cc hp cht khc

tetralin; xylen; toluene; ethylbenzen; ethylcyclopentan

Cholin; betain; oxamat; axeton; cyclohexanol;v mt s cc
hp cht vng no khc

Nh vy ta c th thy vi khun kh sunphat s dng cc loi c cht

khc nhau t nhng hp cht hu c n gin nh hydro, lactat cho n nhng hp
cht phc tp hn nh cc hydratcarbon, hydrocarbon.
c bit l trong nhng nm gny, cc nh nghin cu ngy cng pht hin ra
nhiu hp cht hydrocarbon du m m vi khun kh sunphat c th lm ngun c
cht

2.1.3

Mt s v d v hnh nh ca vi khun kh sulfate.

11

Hnh 2.3: Hnh nh vi khun aceterbaxtor

Hnh 2.4. Hnh nh vi khun desulfovibrio sp.

2.2 Tnh cht

2.2.1 Qu trnh trao i cht v nng lng
Vi khun kh sunfat thc hin qu trnh trao i cht v nng lng ca chng
trong iu kin k kh nghim ngt v c th oxy ha kh Eh< -150mV. S c mt ca
oxy nh hng tiu cc n s pht trin ca chng. Nu trong mi trng c oxy th
ch mt s t loi c th tip tc sinh trng. Trong iu kin k kh, da vo qu trnh
trao i cht vi khun kh sunfat c chia thnh hai nhm chnh. Cc loi thuc
nhm th nht tin hnh oxy ha khng hon ton cc hp cht hu c vi axetat l
sn phm cui cng. i din ca nhm ny c th k n chi Desulfovibrio,
Desulfobulbus, Archaeoglobus...Cc loi thuc nhm th hai oxy ha hon ton cht
hu c k c axetat thnh

nh Desulfobacterium, Desulfococcus, Desulfonema,

Thermodesulfobacterium... . Hu ht cc loi thuc nhm mt sinh trng nhanh hn

12

nhm hai, trong khi cc loi thuc nhm hai cn c th hp tc vi cc loi thuc
nhm mt bng cch s dng axetat do cc loi nhm mt thi ra trong qu trnh trao
i cht.
Vi khun kh sunfat c th s dng c nhiu loi c cht khc nhau. Mt s
cht ny c nhiu chng s dng trong khi mt s cht khc li ch c th l ngun
nng lng cho duy nht mt chng. Chi Desulfovibrio v cc chi khc thuc nhm vi
khun kh sunfat ta nhit Gram m nh ni l nhng loi hp v c cht ( ch gm
hydro, lactat, pyruvat), tri li, chi Desulfotomaculum li c bit n l chi c kh
nng s dng ngun c cht rt linh ng. N s dng rt nhiu cht lm cht in t
cho qu trnh sinh trng v pht trin ca mnh
Cc c cht vi khun kh sunfat s dng trong qu trnh sinh trng: axit
monocarboxylic
Focmat: axetat; propionat; butyrat; lactat
Succinat: fumarat; malat;oxalat; malenat
Metanol: etanol; propanol; butanol
Glyxin: alanin; cystein;toluen; cc hp cht khc
Cholin; betain; oxamat; axeton; v mt s cc hp cht vng no khc
nh hng ca cc yu t mi trng n sinh trng ca vi khun kh sunfat.
Hot ng ca vi sinh vt trong t nhin chu nh hng nhiu ca cc yu t vt l,
ha hc. Nhng yu t ny nh hng khng nhng n ln v thnh phn loi ca
cc qun th vi sinh vt m cn nh hng n hnh thi v sinh l ca chng. mt
s loi, nu cc yu t nh nhit , nng mui v pH cao hn hoc thp hn
mc ti u th c th dn ti s thay i ng k v trao i cht, hnh dng t bo v
qu trnh sinh sn. Tuy rng, trong cc mi trng sng t nhin c rt nhiu cc yu
t khc nhau tc ng ln c th sng, song mt yu t trong chng bao gi cng c
ngha c bit. l nhng yu t gii hn kh nng sng ca cc vi sinh vt mt
mc nht nh.
Trong ba yu t chnh c coi l ng vai tr quan trng trong vic
kim sot sinh trng ca vi khun kh sunfat l nhit , mn v pH

13

 Nhit :

Nhit nh hng n phn ng kh sunfat, s to thnh

S v s oxy ha

c cht cng nh hnh thi t bo. Theo bo co ca Mueller v Nielsen, mt s

chng vi khun kh sunfat qua nhit, phn ng kh sunfat quan st c nhit
35-75

v trong khong nhit 50-60

s kh sunfat xy ra vi tc cao nht.

nhit thp hn hoc cao hn, phn ng kh sunfat xy ra vi tc thp hn v

pha lag ko di hn. Hnh thi t bo rt khc nhau cc nhit khc nhau. Th tch
t bo cng tng khi nhit tng. i vi cc mu ly t vng Alaska v bin Bc
nhit 35 . Cc t bo hnh que c kch thc trong khong 0.3 -0.6 m 1.5 3
m chim u th trong khi 50
60

cc t bo dng si t 5-50 m li chim u th.

cc t bo chim u th di t 2 4m v c ng knh t 0.3 0.7 m cn t

bo chim u th 75

li l t bo hnh cu nh vi ng knh trong khong 0.2

0.5 m.
Trong t nhin vi khun kh sunfat sng c iu kin nhit khc nhau t
0-100 .Mt s chng thuc chi Desulfovibrio c kh nng pht trin nhit 0 .
Do vic ly mu cng nh nui cy vi khun kh sunfat a lnh gp nhiu kh khn
nn nhm vi khun ny chua c quan tm nghin cu nhiu. Cc vi khun kh
sunfat a m c th sng dc nhit 4-65

trong nhit ti u cho s sinh

trng l trong khong 29-42 . Vi khun kh sunfat a nhit li c th sng trong

khong nhit 40-89

v nhit ti u cho s sinh trng ca chng l 55- 76.

Cn i vi vi khun kh sunfat siu a nhit, nhit ti u cho s sinh

trng ca chng l 80-92

ng thi nu nhit nh hn 46

hoc ln hn 110

th chng khng th sng c.

c trng v nhit :

14

Cc phn ng nhit ca qun x vi khun kh sunfat c s dng nh l c

trng nhit ca qun x vi khun ti
Nhit lm nh hng n tc tng trng ca cc cng ng vi khun
nhit di nc
Trm tch c thu thp t mt b ven bin, thm lc a, v dc trm tch t
pha ty nam, ng nam i Ty Dng v Cc Bc su 4327 m. Nhit dao
ng ti t -1oC 8oC Bc Cc. c im nhit ny qun x vi khun kh
sunfat c th hot ng c l incubations ngn hn vi 35S-sulfate trong mt khi
gradient bao trm mt phm vi nhit 0-40 C.
Nhit ti u (Topt) gia 27 C v 30 C cho cc trm tch thm Nam i Ty
Dng l biu hin ca mt cng ng mesophilic psychrotolerant / sulfate-kh,
trong khi Topt 20 C Nam i Ty Dng dc v thm Bc Cc trm tch cho thy
mt cng ng ch yu a lnh.
Cn ti trm tch lc a dc cho thy iu kin nhit thun li l 30 C
mn (NaCl)
Vi khun kh sunfat c tm thy trong mi trng nc t nhin c nng
mui NaCl t 0% n bo ha. Ty tng loi m nhu cu NaCl khc nhau. C nhng
loi khng cn cung cp NaCl trong mi trng nui cy nh cc vi khun kh sunfat
nc ngt. Chng c th b c ch nu nng NaCl >2-3%. Ngc li, nhiu loi
phn lp c t nc bin rt cn NaCl trong mi trng sng. Chng khng th sinh
trng c trong mi trng nc ngt m i hi 10-30 gNaCl/l v thnh thong
cn c mui magie cho qu trnh sinh trng.
Trong mi trng mn t nhin nh h mui NaCl, ngi ta thy vi khun kh
sunfat vn hot ng nng mui NaCl gn bo ha. Tuy nhin cha c bo co
no khng nh vi khun kh sunfat a mn i hi nng mui NaCl cao cho sinh
trng ti u. Mt vi vi khun kh sunfat c th chu ng c nng mui NaCl
cao hn mc d hot ng ca chng b gim bt khi nng NaCl tng ln n 510%.

15

Vi khun a m chu mn cao, chng hn Desulfovibrio vietnamensis c kh nng

chu mn n nng 30% NaCl. Tuy nhin, loi ny cng c th sng nng 0%
NaCl v thch hp nht cho s sinh trng 5% NaCl.
i vi vi khun a nhit th khong nng NaCl m chng c th sinh trng
c l 0-6% v thch hp nht l 1-2%. Trong khi nng NaCl i vi vi khun
siu a nhit la 1-2% NaCl
PH
S sinh trng v sinh sn ca vi khun kh sunfat b nh hng mnh bi pH mi
trng. Khong pH thch hp ca vi khun kh sunfat nm trong khong t 6-9. Tuy
nhin, i vi tng loi vi khun kh sunfat li c pH thch hp ring. Chng hn vi
khun kh sunfat a m c pH thch hp nm trong khong 6.8-7.5. Tuy vy chng c
th sng c nu pH gim xung 5 hay tng l n 10. Cn i vi vi khun a nhit
chng c th sng c trong khong pH 5.5-9.1 v thch hp trong khong 68.-7.5 ,
trong khi pH vi khun rt a nhit l 6-7.
Trong mi trng c pH khng thch hp vi khun kh sunfat vn sng c do
chng vn c th sinh trng trong cc sinh thi hp c pH trung tnh c to nn
bi cc sn phm trao i cht ca chng.

2.2.1 Vn v mi
Khi khng c s hin din ca oxy, sulfate c coi nh l cht cung cp
oxy (chnh xc hn l cht nhn in t) cho qu trnh oxy ha sinh ha ca vi
khun k kh. Trong iu kin k kh, sulfate b kh thnh S2-. Ion S2- s kt hp
vi ion H+ vi hng s phn ly KA1 = 9,1.10-8. Quan h gia cc dng H2S, HSv S2- ti cc pH khc nhau ca dung dch cha 10-3 M H2S (hay 32 mg/L H2S)
Ti pH = 8 trong dung dch tn ti ch yu hai dng HS- v S2-, H2S ch tn
ti mt lng rt nh, v vy p sut ring phn ca n rt thp. Do , vn
mi khng xy ra. Ti pH < 8 cn bng hng ti s hnh thnh H2S, ti pH = 7,
16

80% S2- dng H2S. Khi mt lng ln sulfate b kh thnh ion sulfide, p sut
ring phn ca H2S gy ra vn v mi. Do c tnh ca kh H2S, trong
khng kh hm lng ca H2S nn nh hn 20 ppm.

2.2.2 n mn ng ng
S n mn nh cng (crown) ca ng bton l c bit nghim trng khi
m nc thi sinh hot c nhit cao, thi gian lu trong cng di v nng
sulfate cao, iu ny xy ra nhiu vng ca M, c bit nhng vng pha
nam nc ny. Nguyn nhn ca s n mn c cho l do qu trnh kh sulfate
t H2S thnh H2SO4. Thc ra H2S, hay H2S acid, l mt acid yu, yu hn c
H2CO3 v t nh hng n bton c cht lng cao. Tuy nhin, trong h thng
cng thot nc t chy, H2S l nguyn nhn gin tip gy ra s n mn nh
cng.
i vi cng thot nc t chy thng t dng trong nhng mi trng c
s hin din ca sulfate v c nhng bin i sinh hc. H thng cng l mt
phn ca h thng x l v trong qu trnh vn chuyn nc thi lun xy ra cc
bin i sinh hc. Nhng bin i ny i hi c mt ca oxy, nu lng oxy
khng do qu trnh thng gi t nhin ca khng kh trong cng, qu trnh kh
sulfate thnh sulfide s xy ra. pH thng thng ca nc thi, hu ht S2-
dng H2S v mt phn ca n bay vo lp khng kh trn lp nc thi trong
cng. Nu h thng cng c thng gi tt v thnh cng v nh cng kh ro,
vic hnh thnh ca H2S khng gy ra s n mn cng. Tuy nhin, trong trng
hp thng gi km, thnh v nh cng m t, H2S s ha tan vo lp nc trn
thnh v nh cng tng ng vi p sut ring phn ca n trong khng kh hin
din trong cng. iu ny hu nh khng gy nguy hi no.
Vi khun c kh nng oxy ha H2S thnh H2SO4 c mt khp ni trong t nhin
v trong nc thi. V d nhin l loi vi khun ny cng c mt trn thnh v
nh cng ti nhng lc lu lng ln hay theo mt s cch khc. Do iu kin
hiu kh l lun tn ti trong h thng cng, nhng vi khun hiu kh oxy ha H2S
thnh H2SO4 v sau tr nn m c v n mn bton. Vi khun Thiobacillus,
17

c kh nng oxy ha H2S thnh H2SO4 pH = 2, c cho l loi vi khun chnh

gy ra vn ny. Qu trnh hnh thnh H2SO4 c bit nghim trng nh cng
do ti qu trnh rt nc l nh nht.

2.2.3 ng dng
2.2.4 B sung sulfate tang cng s phn huy rc ti cc bi chn lp
Ti cc bi chn lp rc nguyn nhn gy nhim trm trng cho nc mt v
nc ngm l nc r rc. Do trong nc r rc c hm lng cc cht hu c kh
phn hy, nn trong iu kin t nhin s phn hy rc cc bi chn lp din ra
chm gy ra nhim.
tng cng s phn hy rc ti cc bi chn lp ngi ta s dng bin
php ti tun hon nc r rc ch nhiu oxy t do hoc lin kt di dng sunfat
(SO42-), nitrat (NO3-), ...cao, thng qua cc qu trnh kh sunfat hoc kh nitrat vi
khun s ly oxy t y phn hu hiu kh hoc thiu kh cc cht hu c trong rc
thi. Bng cch ny s hn ch c lng nc r rc x ra mi trng bn ngoi,
ng thi rt ngn c thi rt ngn thi gian rc, gim din tch bi chn lp.
Thng qua mt ti l nghin cu qu trnh ti tun hon nc rc trn m hnh
phng th nghim. Kt qu cho thy Mi trng sunfat biu hin vai tr tch cc trong
phn hu thnh phn cht hu c th rn trong rc vo th lng trong nc rc, v
c ho thnh phn hu c th lng th hin qua vic to kh nng oxy ho mnh hn
cho qu trnh phn hu rc bng vi sinh. Bng cch m tng kh nng thi ra ca
ng rc v kh nng x l thnh phn hu c kh phn hu vi sinh trong nc rc.

18

Hnh 2.5. S m hnh th nghim nghin cu tun hon nc r bi chn rc

3.2.2 X l rc thi giy

Trong qu trnh x l rc thi giy bng phng php ym kh. Ngi ta s
dng vi khun kh sulfate oxi ha cc axit hu c bc cao thnh H2S v CO2.
Cc vi khun kh sunphat ny s dng sunphat v sunfit nh l nhng cht nhn in
t trong qu trnh trao i cht ca cc hp cht hu c to ra H2S, CO2.
S qu trinh x l ym kh
Cc cht hu c phc tp
v ho tan

Cc hp cht hu c n
gin

Cc axit hu c bc cao

H2 v CO2

axetat

CH4 v CO2
19

3.2.3 Sn xut ch phm vi sinh

Cc phng php ch yu c ng dng x l nc thi ni chung v nc
nhim phn st ni ring l cc phng php ha l v ch phm vi sinh. Trong
phng php x l bng cc cc ch phm vi sinh vt c s quan tm hn.
Mt trong nhng chng vi khun c ng dng trong cng ngh x l nc nhim
phn st l chng vi khun kh sulfate Desulfovibrio sp. y l vi khun Gram m, k
kh, khng hnh thnh bo t, t bo l n vibrios vi c kch thc 0,5-0,7 m x 1,5-3
m, vi khun c hnh du phy, duy chuyn nh roi, c th tm thy trong bn y ao,
trm tch bin, trong cc ging khoan khai thc du kh, m nc ngm, trong rut
ng vt, trong phn...
Phng php ny da trn kh nng kh ion sulfate (SO42-) ng thi oxi ha cc
hp cht hu c (lactate, acetate, ethanol, methanol) to ion sulfide (H 2S, HS-, S2-) ca
vi khun Desulfovibrio sp. Ion sulfide kt hp vi ion st ha tan trong nc to kt
ta di dng sulfide bn vng
Phn ng loi b st ca vi khun Desulfovibrio sp. :

2CH3CHOHCOOH + 3 SO42- 3H2S + 6 HCO3Fe2+ + H2S FeS + 2H+

3.2.4 ng dng ca vi khun kh sulphate trong x l nhim du kh.

Vi khun kh sunfat (SRB) phn b rng ri trong t nhin, c bit l trong
ging khai thc du kh v trm tch y thy vc. Khi nhc n vi khun kh sulfate
trong du kh ngi ta ngh ngay n tc hi ca ca chng nh gy ra cc hin tng
n mn kim loi trong cc ging du v cc cng trnh ngm hay kh nng phn hy
du lm nh hng n cht lng du m. Nhng trong mt s trng hp nh v
ng ng dn du hay s c trn du gy nhim t du kh th th dng lm tc
nhn x l nhim.

C ch phn gii Ankan

Cc ankan mch thng l nhng thnh phn d b phn hu nht trong cc hp cht
ca du m.
20

Hnh 2.6: C ch phn gii ankan

C ch ca qu trnh phn gii metan

CH4 CH3OH HCHO HCOOHCO2

Phn gii cac hp cht thm:

Cc hydrocacbon thm c chuyn thnh cc dn xut octo_ hay para_

dyoxitphenyl. Di tc dng ca h thng enzyme cm ng (enzyme oxygennaza), cc
vng thm s b ct t. Cui cng, cc vng thm b ct t trn c hydroxyl ha to
thnh cc axit hu c

Bng c ch phn gii cc hidrocacbon c trong hn hp du m nn vi khun

kh sulfate c th dng nh tc nhn x l nhim t du m

21

3.2.5

ng dng trong oxy ha cc khong vt sulfur

Vn nghin cu v phong ho trn cc m qung ni chung v cc m qung

sulfur ni ring ang l mt vn thi s c nhiu nh a cht c bit quan tm.
Kt qu nghin cu nhiu khu vc khc nhau trn Tri t chng minh rng hin
nay c rt nhiu khong sn v ang c khai thc t cc i biu sinh ca cc m
qung.
Trong nghin cu v phong ho, mt trong nhng nhim v quan trng cn gii
quyt cho mt vng lnh th v mt i tng m c th l cc tc nhn gy phong
ho. Bng con ng ho hc thun tu hin nay ngi ta xc nh nhng tc nhn
ho hc quytnh cho qu trnh phong ho gm: nc, oxy, kh carbonic v cc dung
dch t nhin khc. Tuy nhin gn y ngi ta xc nh rng cc vi sinh vt cng
l mt trong cc tc nhn rt quan trng thc ynhanh qu trnh oxy ho - mt trong
nhng qu trnh ch yu xy ra trong s thnh to i biu sinh cc m qung, c bit
l cc m qung sulfur
Cc loi vi khun ny v tng th c chia thnh 2 nhm. Nhm cc vi khun
c tc dng oxy ho cc vt cht nn (cc khong vt sulfur) v nhm cc vi khun c
tc dng kh cht nn.
Bng 2.2. c im mt s vi khun chnh c trong cc m qung
Tn vi khun

Thiobacillus

Thiobacillus
Thiooxidans

Hnh dng v kch thc

Khong

Cht nn

(m m)

pH ca

b oxy ho ti

dung dch

hoc kh

sng
Cc vi khun oxy ho cht nn
Dng a c mt di tt 1 - 5
Lu
0,4 x (1-0,5)

hunh S2-,

Dng a c di tt xon

SO32Lu

1-5

(0,5 - 0,8) x 1

M qung
lu hunh
Ch yu

hunh

m lu

SO32Lu

hunh
Ch yu

Thiobacillus

Dng a vi ui trn

Thioparus

v di tt phn cc -

hunh,

m lu

Thiobacillus

di n 1,5
Dng a c mt di

H2 S
Lu

hunh
M than

tt

hunh,

0,5 x 1

H2S,

pyrit ho

neapolitanus

3 - 10

Kiu m tn

3-7

22

Thiobacillus

Dng a vi ui trn

Fe2+
Cc

organoparus

c 1-2 di tt ct (0,5-

cht

Leptospilliu

0,8)x(1-1,5)
Dng a xon 0,5 x

hu c
Fe2+,

hunh
Qung

3,5

S2_ ,S22_

ho

1-4

ferrooxidans

Qung
lu

sulfur
(pyrit

Stibiobacter

Dng a c di tt ct

senarmonti
Cc vi khun

0,5 x (0,5 - 1,8)

5-8

Sb

3+

ho)
Qung
antimonit

kh cht nn
Khun kh
sulfat

Cc vi khun kh cht nn
Dng a
5 - 10

S6+, vt

Qung

cht hu

sulfur v

lu hunh

Trong cc m qung, cc vi khun ch yu sinh tn trong cc dung dch lu

chuyn trong m, trong cc l hng v khe nt ca khi qung. S sinh tn nh vo
s thm nhp ng thi ca vt cht hu c v v c vo su trong lng t. Vi khun
sinh tn trong iu kin ca s phong ho cc khong vt qung (cc sulfur v
arsenur) do chng b kch thch pht trin vi tc tng trng nhanh (t mt c th
phn ra thnh 60 -70 c th trong mt ngy m), cho nn hot ng sng ca vi
khun lm cho nng lng v c cu ca i tng chuyn ho mnh. Ni cch
khc, s tp trung vi mt ln cc c th sng ca vi khun s chuyn thnh mt
nhn t gy ra s bin i biu sinh ca cc m qung - tc l nhn t ca s hnh
thnh i biu sinh trn cc m qung.
Qu trnh ch yu, trong c s tham gia c lc ca Thiobacillus
ferrooxidans, l qu trnh oxy ho cc sulfur. Qu trnh ny xy ra mnh, nhanh hn
nhiu so vi qu trnh oxy ho bng con ng ho hc thun tu. Trong tin trnh
sinh tn vi vt cht hu c, cc qu trnh sinh ho trong t bo ca vi khun xy ra c
s tham gia ca cc in t do phn ng oxy ho trong cc cht v c to ra. T bo vi
khun m n kch thch s oxy ho khong vt ng vai tr ca mt cht oxy ho,
23

cn vi vai tr ca mt m hnh in ho ca s oxy ho t nhin cc khong vt (cc

qung sulfur) sn c trong dung dch th n ng vai tr ca mt "catod sng" c o.
Cc dung dch t nhin c cha Thiobacillus ferrooxidans thng c c trng
th oxy ho cao (0,6 - 0,65 volt) c kh nng oxy ho cc khong vt sulfur khc nhau
theo mc bn vng ca chng.
S oxy ho cc khong vt qung bng con ng vi khun khc vi
con ng ho hc khng ch cng cao m c v c im ca dung dch to ra,
m v sau dung dch c th kt tinh thnh nhng khong vt biu sinh. Ngi ta
cng nhn ra mt c im ca s oxy ho cc sulfur bng con ng vi khun l s
ph thuc vo nng v trng thi ca cc nguyn t c to ra trong dung dch.
Do tc "chy qua" l hng v khe nt ca khong vt cao nn qu trnh m bo
dung dch b pha long t, v do gi cho nng ca chng mc tng i cao
cng trng thi c trng ca cc nguyn t. Trong iu kin , trong s cc ion
phc c to ra trong dung dch trc ht phi c mt cc cation kiu (Cu HSO 4)+ v
( Fe HSO4)2+ - cc cation ny b thu phn s to c s cho s kt tinh ca cc khong
vt biu sinh t cc nhm sulfat baz.

24

Chng 3: KT LUN V KIN NGH

Vi khun kh sunphat l mt nhm vi khun phn b ra rng trong nhin c

bit nhng ni c hm lng sunphat cao nh trong nc bin hay trong trm tch
y ao, sng, bin. c trng quan trng nht ca vi khun kh sunphat l to ra kh
H2S trong qu trnh sinh dng, cng chnh iu no gy nh hng khng tt cho
h sinh thi c mt s xut hin nhm vi khun ny, d trong iu kin khc nghit
nh: p sut cao, nhit cao, mn cao hay trong mi trng ha km, ha axit...th
vi khun sunphat vn pht trin.
Vi khun kh sunphat khng ch nh hng n cc ngnh cng ngh khai thc
qung m, cc ngnh cng ngh x l rc thi Vi khun kh sunphat thi ra mt
lng ln kh H2S nh hng n cnh quan, sinh thi, mi trng v con ngi. Th
nhng vi khun kh sunphat cng c tm quan trng khc c bit. Vi khun kh
sunphat ph bin nht trong mi trng k kh n h tr trong vic phn hy cht hu
c; trong mi trng ym kh, n ln men ly nng lng t cht hu c ln; kt qu
cc hp cht nh hn nh axit hu c v ru c tip tc b oxy ha
bi acetogens v methanogen v cnh tranh vi khun sunfat gim.
Hin nay trn th gii c nhiu nghin cu v vi khun kh sunphat m ch
yu nghin cu v c im, tnh cht, hn ht l nhng ng dng ca n hin ti v
tng lai.
Sau mt thi gian nghin cu th nhm chng em thy c vi khun kh
sunphat c mt s c im tiu biu v hnh thi nh: th hin hnh thi t bo, kh
nng di ng, kh nng hnh thnh bo t, hm lng GC cha trong ADN,
desulfovibrin, h thng cytochrome, nhit ti u cho sinh trng v kh nng trao
i cht (oxy hon ton hay khng hon ton c cht) [Castro v cs, 2000]. Nhm vi
khun kh sunfat c hnh dng t bo rt a dng ty theo tng chi ( vi kch thc t
bo khong 0.3 1.5

mx0.8 10 m): Hnh que nh Desulfobulbus,

Desulfotomaculum, Thermodesulfobacterium, Thermodesulfohabdus,

25

Thermodesulfatator, hnh du phy nh Desulfovibiro, Thermodesulfovibrio, hnh

ovan nh Desulfobacter, Desulfobacterium, hnh cu nh Desulfococcus, hnh ti
Desulfosarcina, hnh si nh Desulfonema. V c im v sinh l, sinh ha nh: a
m, a nhit.
Tm hiu cc tnh cht ca vi khun kh sunfate nh: Qu trnh trao i cht v
nng lng, cc yu t nh hng nh: nhit , mn, pH; vn v mi; n mn
ng ng v qua cc tnh cht trn ta c th bit c tnh cht ha l ng dng
ca vi khun ny vo i sng.
Vi khun kh sunfate c rt nhiu ng dng trong vic bo v mi trng nh:
Nghin cu tun hon nc r c b sung sunphat cho cc bi chn lp rc thi th
nh; x l rc thi giy bng phng php ym kh; Nghin cu sn xut ch phm vi
sinh c hot tnh x l sulfate t chng vi khun Desulfovibriosp. c phn lp t
phn tru, b v ng dng ca vi khun kh sunfate trong vic x l i nhim du kh
cng nh trong vic x l nc thi axit t hot ng khai thc khong sn.

26

Ti liu tham kho

Ting Vit
1. Cng ty c phn tin hc, cng ngh, mi trng, TCT Than & Khong sn Vit

Nam (2012), Kt qu phn tch nc thi m than.

2. H S Giao, Mai Th Ton (2010), Nhng im nng mi trng trong hot

ng khai thc m Vit Nam, Hi ngh khoa hc k thut m quc t 2010.

3.

Bi Cng Quang (2011), Tc ng ca cc hot ng khai thc m n ngun

nc v h sinh thi, Chuyn bo v mi trng trong khai thc khong sn,
H Thy Li.

4. Nguyn Danh Sn (2011), Mi trng v pht trin bn vng trong qun l khai

thc ti nguyn khong sn Vit Nam, Chuyn bo v mi trng trong khai

thc khong sn, Vin Khoa hc x hi Vit Nam.
5. http://123doc.org/document/78981-kha-nang-phan-huy-dau-mo-va-cac-san-pham-

dau-mo.htm (20 March, 2016)

Ting Anh
1. Muyzer, G. and stam, A.J.(June 2008). the ecology and biotechnology of
sulphate-reducing bacteria .
2. Ernst-Detlef Schulze, Harold A. Mooney (1993), Biodiversity and ecosystem
function, Springer-Verlag, pp.88-90.
3. Barton, Larry L.and Fauque, Guy D. (2009). Biochemistry, Physiology and
Biotechnology of Sulfate-Reducing Bacteria. Advances in Applied Microbiology
68: 41-98.
4. Larry Barton (ed.) (1995), Sulfate-reducing bacteria, Springer.
5. Kasper U.Kjeldsen, Catherine Joulian, and Kjeld Ingvorsen (2004). Oxygen
Tolerance of Sulfate Reducing Bacteria in Activated sludge. Environmental
Science and Technology 38 (7): 2038-2043.

27

6. Pfennig N. and Biebel H. (1986), "The dissimilatory sulfate-reducing bacteria", in

Starr et al., The Prokaryotes: a handbook on habitats, isolation and identification
of bacteria, Springer.
7. Bahr M, Crump BC, Ceraj VK, Teske A, Sogin ML, Hobbie JE (2005),
Molecular chacterization of sulfate-reducing bacteria in a New England salt
marsh, Environ. Microbiol., 7, pp.11751185.
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