Vi Sinh Vật Nội Sinh

8/10/2019 Vi Sinh Vt Ni Sinh

1/24

TRNG I HC NNG NGHIP H NI

KHOA CNG NGHSINH HC

------------

SEMINAR

TI:

Vi sinh vt ni sinh thc vt

Ging vin HD : TS. Nguyn Vn Giang

Hc Vin : Nguyn Khc Hi

Nguyn ThHu

m Thu Huyn

Lu ThPhng Tho

inh Quang TrungLp : CHCNSH-K22

H Ni2014


2/24

2

MC LC

MC LC ................................................................................................................................................2

I. MU ...........................................................................................................................................3

II. NI DUNG......................................................................................................................................5

2.1.Vi khun ni sinh thc vt.............................................................................................................5

2.1.1. Sphn lp v adng sinh hc ca vi khun endophytes..............................................5

2.1.2. Mt snhm vi khun endofphytic..................................................................................7

2.2. Nm rcng sinh................................................................................................................... 12

2.2.1 Khi nim nm rcng sinh................................................................................................. 12

2.2.2. Phn loi nm rcng sinh................................................................................................. 14

2.2.3. Cc yu tnh hng n nm rcng sinh....................................................................... 17

2.2.4. Vai tr ca nm ri vi cy trng................................................................................... 20

III. KT LUN ................................................................................................................................... 22

TI LIU THAM KHO ............................................................................................................................ 23


3/24

3

I. M U

Nhng tng tc c li gia thc vt v vi sinh vtthc y sc khe, s pht

trin ca cy trng, vn ny ang c cc nh nghin cu quan tm. Gn y, h

ang nghin cu tim nng cho gii php nng cao kh nng phn hy sinh hc ca

cc cht gy nhim trong t. Hu ht cc nghin cu ny tp trung vo cc vi khun

r (Lindow&Brandl,2003;Kuiper et al.,2004;berg et al,2005). Endophytic c th

c nh ngha l nhng vi sinh vtc tr trong ni m ca thc vt, chng khng c

biu hin ra bn ngoi v gy tc ng xu n thc vt m chng k sinh

(Holiday,1989; Schulz & Boyle, 2004), c khong 300.000 loi thc vt tn ti trn

tri t, mi loi l mt k ch cho mt n nhiu cc dng ni k sinh c tr. Ch c

mt s loi thc vt c nghin cu hon chnh v cc mi quan h ni k sinh cachng. Do c hi nghin cu v tm ra cc dng ni k sinh mi v c li trong s

a dng sinh hc ca cc h sinh thi khc nhau l ng k.

Endophytic c tr trong h sinh thi thch hp tng t nh cc chi mm

thc vt, iu ny lm cho chng tr thnh cc tc nhn kim sot sinh hc (Berg et

al., 2005). Tht vy, nhiu nghin cu ch ra rng Endophytic bacteria c kh nng

kim sot c mm bnh trn thc vt (Sturz & Matheson, 1996; Duijff et al., 1997;Krishnamurthy & Gnanamanickam, 1997), cn trng (Azevedo et al, 2000.) v c

tuyn trng ( Hallmann et al., 1997, 1998). Trong mt s trng hp chng c th y

mnh tc ny mm ca ht, thc y s hnh thnh cy con trong iu kin bt li

(Chanway,1997) v nng cao kh nng tng trng ca thc vt ( Bent &

Chanway,1998). Endophytic cn c th ngn chn mm bnh pht trin bng cch

tng hp cc cht ni sinh trung gian, qua tip tc tng hp cc cht chuyn ha

v cc hp cht hu c mi. Nghin cu c ch sn sinh cht chuyn ha mi trong s

a dng sinh hc ca Endophytic c th pht hin cc loi thuc mi iu tr c

hiu qu cc bnh ngi, thc vt v ng vt (Strobel et al., 2004).

Cng vi vic sn xut cc cht mi, nhiu Endophytic bacteria cho thykh

nng lm gim cc cht ngoi sinh (xenobiotic) hay c th hot ng nh cc vect

m u cho qu trnh . Chng c kh nng khng kim loi nng, khng khun v

lm gim cc gc hu c thng qua tip xc vi cc hp cht trong thc vt v t nichng c tr. Kh nng lm gim nhng cht xenobiotics ang c nghin cu cn


4/24

4

thn ci tin k thut kh c bng thc vt (Siciliano et al., 2001; Barac et al.,

2004; Germaine et al., 2004, 2006; Porteous-Moore et al., 2006; Ryan et al., 2007a).


5/24

5

II. NI DUNG

2.1.Vi khun ni sinh thc vt

2.1.1. Sphn lp v adng sinh hc ca vi khun endophytes

t bo v mnh trc nhng tc ng ca mi trng, Endophytic bacteria

to thnh nhng vi khun ,xm chim v k sinh trn t cy . Nhng vi khun ny

thng xm chim vo vng khng gian gia cac tbao, v chng c th c phn

lp t tt c cc b phn ca cy, bao gm c ht ging (Posada & Vega, 2005).

Endophytic bacteria c phn lp t cy mt l mm ti cy hai l mm, t nhng

loi cy thn g, nh g si v l, cy thn tho ti cy lng thc nh c ci ng

v ng. Nhng nghin cu c in v s a dng ca Endophytic bacteria tp trungvo phng php phn lp chng t cc m ni sinh sau khi kh trng b mt thc vt

vi hypochlorite natri hoc cc ha cht tng t (Miche & Balandreau, 2001). Mt

nghin cu ca et al Lodewyckx. (2002) nu ln phng php phn lp v m t c

im vi khun endophytes t cc loi thc vt khc nhau. Da trn nn tng nghin

cu ca Hallmann et al. (1997) v Lodewyckx et al. (2002),Rosenblueth & Martinez-

Romero (2006) v Berg & Hallmann (2006) cng b mt danh sch ton din ca

Endophytic bacteria c phn lp t mt lot cc b phn ca cy.

Mt nghin cu gn y ca al Porteous-Moore v cng s. (2006) m t s a

dng ca Endophytic bacteria tm thy trong cy Dng, tng trng ti mi trng

phytoremediation nhim toluene, vi mc ch xc nh cc chng tim nng tng

cng phytoremediation toluene, Ethylbenzene, v hp cht xylen (BTEX). Theom

t ca nghin cu trn, Endophytic bacteria c phn lp t hai ging Dng. Sau

khi phn lp, tin hnh phn tch, so snh trnh t ca on mi 16S RNA, BOX -PCR

ca DNA,cc trnh t mang c tnh sinh l, lin quan n cht s dng, thuc khng

sinh v kim loi nng nhy cm. Nghin cu ny v mt nghin cu ca Germaine et

al. (2004, 2006) chng minh rng trong cc cng ng a dng vi khun tm thy

trong cy Dng, mt s chng Endophytic c mt v c tim nng tng cng cc

cht hu c d bay hi phytoremediation v thuc dit c.Cng trnh nghin cu gn

y ca Frank et al. (2006) l dng k thut sinh hc phn t trong vic phn lp, tmhiu cc c tnh ca Endophytic bacteria v ng dng trong cng ng.Cc chng


6/24

6

Endophytic sinh sng trn thn cy, r, c c phn tch v nhn din bng cch

chy RFLP vi on mi l 16S rRNA. Nm n v phn loi trin vng nht v kh

nng tn ti c xc nh l Cellulomonas, Clavibacter, Curtobacterium,

Pseudomonas v Microbacterium c tmra thng qua trnh t gen 16S rRNA, acid

bo v s dng ngun phn tch carbon (Elvira-Recuenco & van Vuurde, 2000;

Zinniel et al, 2002)

Vic nui cy da trn mi trng nui cy c bn v mi trng nui cy c

lp tht s rt hu ch. Reiter & Sessitsch (2006) iu tra s hin din v phn loi

Endophytic bacteria cc b phn kh sinh hon ton trn cy ngh (cy ngh

albiflorus). Kt qu ca h cho thy rng tng t nh cc loi thc vt khc, cy

ngh l mt k ch cho cng ng Endophyticbacteria sinh trng v pht trin, tuy

nhin cng c loi cha c bit n. Cc bc tin hnh th nghim bao gm ly

trch, phn lp, nui cy v nhn din Endophytic bacteria bng k thut in du DNA

vi on mi 16S rRNA. Kt qu ch ra rng mt s nhm mi xut hin gp phn vo

tnh a dng ca Endophytic bacteria, ch yu l Gammaproteobacteria and

Firmicutes, chng sng ni sinh trn cy ngh.Thnh phn, phn loi cc nhm

Endophytic bacteria a dng v c s khc nhau l do s a dng ca gen 16S rRNAtrong th vin gen. Ch c ba nhm vi khun c tm thy trong mi trng nui cy,

trong c 17 loi i din. Trong khi su nhm c xc nh trong phn tch th

vin gen bao gm 38 loi. Nhm ni bt tm thy trong mi trng nui cy l nhm

Gram dng, c t l G+C thp.Trong khi cc th vin clone, cc Gammaproteo

bacteriachim u th. Nghin cu ny khng nh rng cc nhm Endophytic bacteria

trong mi trng nui cy l mt tp hp con ca a dng sinh hc tng endophyte.


7/24

7

2.1.2. Mt snhm vi khun endofphytic

2.1.2.1. Nhm vi sinh vt cnh m

Hnh 1. Chu trnh chuyn ha nit trong tnhin

Phn loi

Vi khuncnhnitc phn ra thnh ba nhm nh: nhm vi khun sngcng

sinh, nhm vi khunsngtdo v nhm vi khuntngtc vithcvtk ch.Tuy

nhin, s phn bitgiaba nhm ny, cbit l gianhm vi khuncnhm

sngtdo v nhm vi khuncnhmtngtc vithcvtth vnchacm

tmtcch r rng v mtsvi khuncxpvo nhiunhm.

Vi khun sng cng sinh:

l nhngvi khunsngtngtc vithcvtk chtheokiuhai bn cng c

li.Khi ,vi khuncngsinh s tinhnh trao ichtdinhdngvithcvtv

lm thay icutrc m thcvtnivi khunnhc,inhnhl hintngto

ntsnrcanhngcy hu.Hintngcngsinh giavikhunv cy hu

cxem l tngtc gngiavi khunv thcvt,trong ,vi khuncgil

sinh vtcng sinh. Huht cc vi khunc nhnit l vi khungram m, c kh

nnghnh thnh ntsnrcy hu,hay l nhngthnh vin xkhunthucchi

Frankial vi khungram dngc khnnghnh thnh ntsntrn cythn g,cy

hai l mmv cybi.Ban u,nhngvi khuncngsinh vicy hucphn

loi thnh chiRhizobium, do , nhng vi khun ny thng c ni n nh l

nhngvi khunntr(rhizobia). Ngy nay, nhngvi sinh vtcngsinh cy hu


8/24

8

c phn thnh nhiu chi, trong , hu ht cc loi thuc chiRhizobium,

Sinorhizobium (Ensifer), Mesorhizobium v Bradyrhizobium.

Vi khun cnh nittdo:

l nhngvi khuncnhnittngtc vithcvtm khng xm nhpvo

cthcachng theo hngcngsinh chuyn bit.Trongsny quan trngnhtl

cc loi thucchiAzotobacter, Pseudomonas v Clostridium.

Vi khun cnh nittngtc:

l nhngvi sinh vttham gia vo qu trnh traoichtdinh dngvithcvt

nhng khng lm thay i cu trc ca r thc vt. phn bit c vi sinh c

nhnit cng sinh hay tng tc vi thcvtk chth chyu da vo mc

tngtc. Scnhnit tng tc c hiu l qu trnh cnhnit bi nhng vi

sinh vt sng t do nhng li chu nh hng trc tip bi cy trng.Theo Klucas

(1991), trong mi quan htng tc, ccy trng v vi khun cnh nit u c li

nhng mi quan hny thng din ra ngu nhin nhiu hn l scng tc bt buc.

Scnh nit tng tc l mt qu trnh sinh thi trung gian gia cnh nit cng

sinh v nit tdo. V vi khun cnh nit tng tc bao gm nhng vi khun sng

tdo trong vng ln cn rcy thc vt n nhng vi sinh vt sng ni sinh trong mtbo thcvt.qu trnh cnhnitcavi sinh vttngtc c thdinra cn

nhngyu cusau y:

+ Thcvtk ch

+ Chtnnthcvt

+ Mi trngthch hpenzyme nitrogenase hotng.

+ Qu trnh chuynnitccnhtvi khunsang cy trng.

Mtsvi khuntham gia vo qu trnh cnhnittngtc inhnh nh:

Azospirillum, Burkholderia, Enterobacter, Gluconoacetobacter, Herbasspirillum v K

lebsiella

2.1.2.2 Nhm vi sinh vt sinh kch thch tsinh trng thc vt

Auxins l mt cht kch thch sinh trng c thc tng hp bi thc vt,

IAA c vai tr trong qu trnh pht trin tbo nh tng trng tbo, phn chia, hnh

thnh r(Ann Vande Broek, 1998), tuy nhin stng trng ca thc vt li chu s

tc ng khng nhtngun auxins bn ngoi v mt trong nhng ngun l auxins


9/24

9

c tng hp bi cc vi sinh vt c ch trong t. Trong cc vi sinh vt c phn

lp tvng rv trn bmt rca nhiu loi cy trng l c khnng tng hp IAA

cao (Arshad v Frankenberger, 1998). Theo Loper v Schroth (1986) th c n 80%

cc chng vi sinh vt c phn lp tvng rca nhiu loi cy trng l c khnng

tng hp auxins. Cc chng vi sinh vt thuc cc ging nh Azospirilium,

Pseudomonas, Rhizobium, Xanthomonas, Bradyrhizobium japonicum, Gluconobacter

diazotrophicus c xc nh l c khnng tng hp IAA gip kch thch ssinh

trng ca cy trng (Patten v Glick, 1996). Gn y l Ahmad (2005) phn lp

c 10 chngAzotobacterv 11 chngPseudomonastt vng rcy la m, m

tt, cy ci, u cho thy chng c khnng tng hp IAA rt cao trong mi trng c

bsung Tryptophan v khng c bsung Tryptophan.

Nhiu dng vi sinh vt sng trong t c khnng tng hp nn cc cht tng

trng ca thc vt v gip cho cy trng pht trin. Trong c nhiu vi sinh vt

sng trong vng r(rhzophere) c khnng tng hp cht kch thch stng trng

ca thc vt gi l nhm (PGPR)(Trc, 2007). Vi khun vng r kch thch tng

trng thc vt ci thin stng trng thc vt v gia tng nng sut vi c chtrc

tip(cnh m, ha tan ln kh tan, sn xut ra auxin hay cytokinin) v c chgin

tip (hn ch s pht trin vi sinh vt gy hi)(Brown, 1974; Klopper v ctv, 1986,

1989; Davision, 1988; Lambert v Joos, 1989). Vi rt nhiu nhm vi sinh vt s hch

nh Pseudomonas syringae (Glickmann v ctv, 1998), Azosprillium brasilense

(Hartman v ctv, 1983), Azobacter vineelandii (Garcia-Tavares v ctv, 1987),

Agrobacterium tumefasciens (Liu v ctv, 1982), Rhizobium trifolii v Rhizobium

leguminosarum(Badenoch-Jones v ctv, 1982), Rhizobium phaseoli(Ernstsen v ctv,

1987), Bradyrhizobium japonicum (Kaneshiro v ctv, 1983), Erwinia herbola (Kogav ctv, 1991), thanh to Anabaena cylindricav Nostoc rivulare(Florenzano v ctv,

1978),Enterobacter cloacae(Koga, 1995), cc vi sinh vt trn l i din cho nhng

vi sinh vt c khnng tng hp kch thch ttng trng cho cy trng, bng 1 di

y l mt svi sinh vt c khnng tng hp kch thch ttng trng.


10/24

10

Bng 1: Cc nhm vi sinh vt tng hp IAA v cc cht dn xut

Vi sinh vt Sn phm Tc giAgrobacterium tumefasciens IAA Muller v ctv (1989)

Alcaligenes ssp. IAA Brown (1972)Arthrobacter ssp. IAA Wiknson v ctv (1994)Bacillus cereus IAA Wiknson v ctv (1994)Bacillus circuslans Tng tauxin Stzelczyl v Prokojska-

Burdziez(1984)Bacillus subtilus IAA Muller v ctv (1989)Pseudomonas spp. IAM, IAA,

IPyAMartens v Franerberger(1991)

Pseudomonas syringae pv.glycinea

IAA, Iald, ILA Fett v ctv (1987)

Streptomyces ssp. Tng tauxin Stzelczyl v Prokojska-Burdziez(1984)

Xanthomonas pv. glycine IAS, ILA, IAM Fett v ctv (1987)Xanthomonas maltophilia IAA Wiknson v ctv (1994)Rhizobium meliloti IAA Garcia-Rodriguez v

ctv(1981)Rhizobium phaseoli IAA, IM Ernsten v ctv(1987)Rhizobium trifolii IAA, IpyA, ILA Badenoch-Jones v ctv(1982)Rhizobium japonicum IAA Kaneshero v ctv (1983)

Bradyzhizobium spp. NAA Sekine v ctv (1988)Azobacter spp. IAA Mahmoud v ctv (1984)Azobacter chroococcum IAA Muller v ctv (1989)Azobacter vinelandii IAA Gonzalez-Lopez v ctv(1983)Anabaena cylindrica IBA Florenzo v ctv(1978)Nostoc rivulare IAA, IPyA Florenzo v ctv(1978)H. hiemale IAA, ICA, IAld Gay v Deband (1987)H. masophaeum IAA Stzelczyl v ctv (1992)Glomus spp. IAA Polojska v Strzelczyl (1988)Trichoderma spp. Tng tauxin Kampert v Strzelczyl

Wardomyces humicola IAA Reddy v ctv(1988)(Ngun:Trch t cc bng trong phytohormone trong t ca W.T.Frankenberger v Jr.Muhannad

Arshas, 1995.)

Cc nh khoa hc tin hnh phn lp, xc nh v nghin cu a cc vi

sinh vt ny vo phn sinh hc gp phn lm taweng nng sut cy trng v thn

thin vi mi trng sng v t c nhng kt qu kh quan trn u nnh

(Molla v ctv, 2001), la go (Rasul v ctv, 1998), trn bp lai (Chabot v ctv, 1996).

Nh vy khnng tng hp kch thch ttng trng vi sinh vt l rt ln, c th


11/24

11

chia thnh ba nhm vi sinh vt s hch l: vi khun nt r (Rhizobia), vi khun c

nh m sng tdo v vi khun vng r.

Vi khun nt r

Cc nh khoa hc pht hin ra vi sinh vt nt rtng hp c IAA ttin

cht tryptophan nhng nhm ny c thtng hp IAA khng c tryptophan , tuy nhin

nu b sung tryptopgan th chng tng hp IAA nhiu gp 7 ln (Kittell v ctv,

1989).Nhiu vi khun nt r thch hp vi loi truptophan khc nhau, nh vi khun

nt rtnt rcy u phng thch DL-tryptophan (Roy v Basu, 1988), cn vi khun

nt t cy Crotalria retusa thch D-tryptophan hn loi L-tryptophan hay DL-

tryptophan (Bhattacharyya v Basu, 1991).

Azotobacter

Sdng vi khunAzotobacter chroococcumlm phn chng cho ht ging

cung cp Nitow cho cy trng c ph bin rng ri Lin X trong nhng nm

1950 (Cooper, 1950),nhng sau ny ngi ta chng minh n c thtng hp kch

thch ttng trng thc vt (Muller v ctv, 1989) v nhiu loi khc trong ging ny

c khnng tng hp c kch thch t tng trng. Azotobacter chroococcumcthtng hp IAA trong iu kin khng c tryptophan (Elwan v El-Naggar, 1972),

nhng nu chng ta cung cp thm tryptophan th lng IAA c tng hp snhiu

hn (Apte v Sharde, 1981).

Azospirillum

Vi khun ny c phn lp tt vng rnhiu loi cy trng v loi c

vng nhit i v n i (Haahtela v ctv, 1981), trong nhng nm 1984- 1985 ccnh khoa hc pht hin nhiu ging Azospirium trong vng r ca c Kallar

(Leptochloa fusca)(Reinhoold v ctv) v nhng vi khun ny c sdng lm phn

chng cho c lm thc n gia sc nh kh nng c nh m cao (Bashan v ctv,

1989), nhng gn ay cc nh khoa hc pht hin ra thmAzospirillum c khnng

tng hp IAA cung cp cho cy trng gip tng nbawng sut cy trng (Barbieri v

Galli, 1993). Azospirillum brasilense c khnng tng hp IAA nh gen ipdC(Ann

Vande Broek,1998), Azospirillum c th tng hp tryptophan m khng c mt ca


12/24

12

tryptophan (Fallik v ctv, 1989) tuy nhin nu bsung tryptophan th chng stng

hp lng tryptophan nhiu hn (Horemans v ctv, 1986).

Vi khun Pseudomonas syringae pv. Savastanoi

Pseudomonasl vi khun gram m, vi khun ny to ra cc bu hay nt trn

nhiu loi vy g (Smidt v Kosuge, 1978), nhng sau ny nhiu nh khoa hc

chng minh loi ny c khnng tng hp IAA t tryptophan nhhai gen iaaM v

iaaH trong plasmid (White v Ziegler, 1991), gn y, Gardan v ctv (1992) tm thy

131 dng t loi ny c phn lp tt vng r ca nhiu loi cy trng c kh

nng tng hp IAA.

Vi khun Guconacetobacter

Vi khun Guconacetobacter v Guconobacterthuc hAcetobacteraceae, y

l nhng vi khun sinh trng tt mi trng hiu kh, nhit 300v pH khong

5.5-6. C nhiu loi khc nhau trong loi Guconacetobacter diazotrophicus

c xc nh l hin din rt nhiu trong ma v cc loi cy ha bn, chng c kh

nng c nh m, ha tan ln, tng hp kch thch t tng trng auxin cho cy

(Mthukumarasamy v ctv, 2002; Madhaiyan, 2004).

Vi khun Burkholder ia

Vi khunBurkholderia thu c nh m vi khun gram m, c hnh dng xon,

chng c thdi chuyn nhcc chim mao phn u(Jesus and al, 2004). Chng

sinh trng trong iu kin kkh hoc hiu kh, nhng trong mi trng kkh spht

trin tt hn(Paulina and al, 2001). Vi khunBurkholderia sng vng rca rt

nhiu loi cy nh: ng, ma, c ph, hho bnc khnng tng hp cc cht tngtrng cho cy trng.

2.2. Nm rcng sinh

2.2.1 Khi nim nm rcng sinh

Nm rcng sinh (Mycorrhiza) l mt thut ngc Frank sdng ln u tin

vo nm 1885 khi pht hin mi lin hgia si nm v rtrn cy thng v mt s

cy l rng. Mycorrhiza c ngun gc t ting Latinh, myco ngha l nm v


13/24

13

rhizac ngha l r, mycorrhizal (cng sinh cng c li) mi quan hgia hr

ca thc vt v nm.

a. b. c.

Hnh 1.1. Mt shnh nh vnm r:

a.Si nm rphn nhnh, b.u chp si nm r,

c.si nm rang vn ra trn bmt gc cy

Ngi ta pht hin ra cch y 370 triu nm c nm cng sinh trong cc cy h

quyt. n gia thkXX th cc hiu bit vnm rcng sinh cng nhiu v su sc

hn. Hin nay, cc nh khoa hc chuyn ngnh u nhn thc c rng, tuyt i b

phn thc vt u c nm rcng sinh v l mt hin tng phbin trong tnhin.

Theo thng k ca Trappe (1962) c khong 535 loi nm thuc 81 chi 30 h10 b

nm c thcng sinh vi trn 15.000 loi thc vt khc nhau.

Cc loi nm cng sinh phn ln thuc cc b: nm tn (Agaricales), nm tn

(Russulales), nm gan b (Boletales), nm m (Tricholomatales), nm bng

(Hymenogastales), nm c nga v cng (Sclerodermatales), nm bng cao

(Gautieriales) v nm phi phin (Aphyllophorales) trong ngnh ph nm m

(Basidiomycotina), cc bnm mng mm (Helotiales), nm a, nm cc. Nhng b

nm ny hnh thnh nm ngoi cng sinh, cn mt s loi nm b ti trong

(Endogonales), trong ngnh phnm tip hp (Zygomycotina) c thhnh thnh nm

ni cng sinh.Rnm l hin tng cng sinh thc vt phbin trong tnhin. Nm cng sinh

nhn c tc ththc vt hp cht cacbon, cc cht dinh dng m thc vt cng

nhn c dinh dng v nc cn thit gip nhau cng c li.N va c c trng

ca brcy thng thng li va c c tnh ca nm chuyn tnh. Rv nm cng

tn ti l kt quca stin ha chung. Stn ti ca chng c li cho ssng cn

ca cy, tng khnng khng vi iu kin bt li, tng khnng sinh trng v

cng c li cho ssng ca nm. Mi quan hny pht trin n mc rt kh phn

bit.
http://aa.wrs.yahoo.com/_ylt=A0S0zu6VIxNLL0EBDh9tUwx.;_ylu=X3oDMTBqaTFoaGxvBHBvcwMxNwRzZWMDc3IEdnRpZAM-/SIG=1k3s2j9v9/EXP=1259631893/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhiza+and+plant&b=1&ni=20&ei=utf-8&y=T%C3%ACm+ki%E1%BA%BFm&pstart=1&fr=yfp-t-101&w=496&h=409&imgurl=www.laspilitas.com/advanced/pictures/roots.jpg&rurl=http://www.laspilitas.com/advanced/advroots.htm&size=30k&name=roots+jpg&p=mycorrhiza+and+plant&oid=15b38395c62bdbfc&fr2=&no=17&tt=564&sigr=11fpc04pv&sigi=11e4rbkjc&sigb=147dp6a45&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zuzOIRNLglUACPptUwx.;_ylu=X3oDMTBpY2Y5NXNiBHBvcwM2BHNlYwNzcgR2dGlkAw--/SIG=1ktc0q2nl/EXP=1259631438/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhizal&b=1&ni=20&ei=UTF-8&pstart=1&fr=yfp-t-101-s&fr2=tab-web&w=699&h=361&imgurl=botit.botany.wisc.edu/images/130/Root/Mycorrhizal_root_130_.jpg&rurl=http://botit.botany.wisc.edu/images/130/Root/Mycorrhizal_root_130_.php?highres=true&size=38k&name=Mycorrhizal+root...&p=mycorrhizal&oid=be6b14ea0d36bcf2&fr2=tab-web&no=6&tt=8567&sigr=12jo5ok4m&sigi=11v6bi0p7&sigb=13kjmooqg&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zu7zIBNLEmQAloBtUwx.;_ylu=X3oDMTBqbjgzbGMzBHBvcwM5OQRzZWMDc3IEdnRpZAM-/SIG=1ir8dpdbk/EXP=1259631219/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhizal&b=81&ni=20&ei=UTF-8&pstart=1&fr=yfp-t-101-s&fr2=tab-web&w=110&h=144&imgurl=www.canadiangardening.com/photos/mycorrhizal.jpg&rurl=http://www.canadiangardening.com/gardens/regions_que3.shtml&size=7k&name=mycorrhizal+jpg&p=mycorrhizal&oid=f198d00c88e651ae&fr2=tab-web&no=99&tt=8567&sigr=11r7puuni&sigi=11g4a8js7&sigb=13ljoh85g&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zu6VIxNLL0EBDh9tUwx.;_ylu=X3oDMTBqaTFoaGxvBHBvcwMxNwRzZWMDc3IEdnRpZAM-/SIG=1k3s2j9v9/EXP=1259631893/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhiza+and+plant&b=1&ni=20&ei=utf-8&y=T%C3%ACm+ki%E1%BA%BFm&pstart=1&fr=yfp-t-101&w=496&h=409&imgurl=www.laspilitas.com/advanced/pictures/roots.jpg&rurl=http://www.laspilitas.com/advanced/advroots.htm&size=30k&name=roots+jpg&p=mycorrhiza+and+plant&oid=15b38395c62bdbfc&fr2=&no=17&tt=564&sigr=11fpc04pv&sigi=11e4rbkjc&sigb=147dp6a45&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zuzOIRNLglUACPptUwx.;_ylu=X3oDMTBpY2Y5NXNiBHBvcwM2BHNlYwNzcgR2dGlkAw--/SIG=1ktc0q2nl/EXP=1259631438/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhizal&b=1&ni=20&ei=UTF-8&pstart=1&fr=yfp-t-101-s&fr2=tab-web&w=699&h=361&imgurl=botit.botany.wisc.edu/images/130/Root/Mycorrhizal_root_130_.jpg&rurl=http://botit.botany.wisc.edu/images/130/Root/Mycorrhizal_root_130_.php?highres=true&size=38k&name=Mycorrhizal+root...&p=mycorrhizal&oid=be6b14ea0d36bcf2&fr2=tab-web&no=6&tt=8567&sigr=12jo5ok4m&sigi=11v6bi0p7&sigb=13kjmooqg&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zu7zIBNLEmQAloBtUwx.;_ylu=X3oDMTBqbjgzbGMzBHBvcwM5OQRzZWMDc3IEdnRpZAM-/SIG=1ir8dpdbk/EXP=1259631219/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhizal&b=81&ni=20&ei=UTF-8&pstart=1&fr=yfp-t-101-s&fr2=tab-web&w=110&h=144&imgurl=www.canadiangardening.com/photos/mycorrhizal.jpg&rurl=http://www.canadiangardening.com/gardens/regions_que3.shtml&size=7k&name=mycorrhizal+jpg&p=mycorrhizal&oid=f198d00c88e651ae&fr2=tab-web&no=99&tt=8567&sigr=11r7puuni&sigi=11g4a8js7&sigb=13ljoh85g&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zu6VIxNLL0EBDh9tUwx.;_ylu=X3oDMTBqaTFoaGxvBHBvcwMxNwRzZWMDc3IEdnRpZAM-/SIG=1k3s2j9v9/EXP=1259631893/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhiza+and+plant&b=1&ni=20&ei=utf-8&y=T%C3%ACm+ki%E1%BA%BFm&pstart=1&fr=yfp-t-101&w=496&h=409&imgurl=www.laspilitas.com/advanced/pictures/roots.jpg&rurl=http://www.laspilitas.com/advanced/advroots.htm&size=30k&name=roots+jpg&p=mycorrhiza+and+plant&oid=15b38395c62bdbfc&fr2=&no=17&tt=564&sigr=11fpc04pv&sigi=11e4rbkjc&sigb=147dp6a45&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zuzOIRNLglUACPptUwx.;_ylu=X3oDMTBpY2Y5NXNiBHBvcwM2BHNlYwNzcgR2dGlkAw--/SIG=1ktc0q2nl/EXP=1259631438/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhizal&b=1&ni=20&ei=UTF-8&pstart=1&fr=yfp-t-101-s&fr2=tab-web&w=699&h=361&imgurl=botit.botany.wisc.edu/images/130/Root/Mycorrhizal_root_130_.jpg&rurl=http://botit.botany.wisc.edu/images/130/Root/Mycorrhizal_root_130_.php?highres=true&size=38k&name=Mycorrhizal+root...&p=mycorrhizal&oid=be6b14ea0d36bcf2&fr2=tab-web&no=6&tt=8567&sigr=12jo5ok4m&sigi=11v6bi0p7&sigb=13kjmooqg&type=JPGhttp://aa.wrs.yahoo.com/_ylt=A0S0zu7zIBNLEmQAloBtUwx.;_ylu=X3oDMTBqbjgzbGMzBHBvcwM5OQRzZWMDc3IEdnRpZAM-/SIG=1ir8dpdbk/EXP=1259631219/**http:/vn.images.search.yahoo.com/images/view?back=http://vn.images.search.yahoo.com/search/images?p=mycorrhizal&b=81&ni=20&ei=UTF-8&pstart=1&fr=yfp-t-101-s&fr2=tab-web&w=110&h=144&imgurl=www.canadiangardening.com/photos/mycorrhizal.jpg&rurl=http://www.canadiangardening.com/gardens/regions_que3.shtml&size=7k&name=mycorrhizal+jpg&p=mycorrhizal&oid=f198d00c88e651ae&fr2=tab-web&no=99&tt=8567&sigr=11r7puuni&sigi=11g4a8js7&sigb=13ljoh85g&type=JPG


14/24

14

2.2.2. Phn loi nm rcng sinh

Ngi ta chia nm rra lm 3 loi: ngoi sinh, ni sinh v ni ngoi sinh.

2.2.2.1. Nm rngoi cng sinh (Ectomycorrhiza)

Nm rngoi cng sinh l si nm bao quanh rdinh dng cha ha g, khngxuyn qua m tbo m chko di gia cc vch tbo. c trng c bn ca chng

l:

Trn bmt dinh dng hnh thnh mt mng nm (mantle) do cc si nm

an cho nhau.

Gia cc tbo tng v rhnh thnh mt mng li do thsi nm sinh

trng m thnh gi l li Hartig (Hartig net).

Do tc dng ca nm r, brngn, to, gin v c mu sc khc nhau, tn

rv biu b khng c lng ht, bmt mng c nhiu si nm ko di ra.

R nm ngoi cng sinh ni chung khng c hnh dng v mu sc nht nh

nhng rt dnhn bit bng mt thng. Tnh a dng thhin trn loi cy chv

nm rkhc nhau.

Hnh 1.2.Hnh thi ca rnm ngoi cng sinh: 1.Dng n trc, 2.Dng lng chim,

3.Dng cha nng, 4.Dng thp, 5.Dng khng quy tc, 6.Dng c,

7.Dng sn si

Con ng xm nhp vo rca nm rngoi cng sinh:


15/24

15

Bc 1: Si nm tip xc, nhn bit v

bm cht cc t bo biu b r gn nh

ca r non, tch cc pht trin.

Hnh 1.3.Cc si nm trn bmt r1-2

ngy sau khi tip xc u tin vi gc

Bc 2:Giai on sau ca sxm nhp

r: cc si nm hnh thnh lp ph dy

c trn bmt r.

Hnh 1.4.Mng lp phHartig sau 2-4

ngy

2.2.2.2. Nm rni cng sinh (Endomycorrhiza)

Nm rni cng sinh l thsi nm c thxuyn qua tbo v rcy ch, khng

bin i hnh thi, bmt rkhng hnh thnh mng nm chc cc si la tha, lng

ht vn ginguyn. Tuy nhin, thsi nm vn ko di gia gian bo, nhng khng

hnh thnh mng li Hartig. Si nm xuyn qua vch tbo vo trong hnh thnh vi

ht. Nhng loi ny rt kh nhn bit bng mt thng.

Cn cvo kt cu si nm c vch ngn v vi ht, chia ra 2 loi: khng c vch

ngn (Aseptate - endotrophic mycorrhiza) v c vch ngn (Septate - endotrophic

mycorrhiza).

Loi khng c vch ngn thng c dng ti bng (Vesicular) v dng chm

(Arbuscular), gi l rnm dng ti chm (Vesicular- Arbuscular mycorhiza) v gi

tt l VA.

Cc loi c vch ngn li cn cvo cy chv hnh dng si nm trong tbo m

chia ra : r nm dng quyn (Ericaceous mycorhiza) si nm trong t bo dng


16/24

16

xon vng (coil); rnm loi hLan (Orchidaceous mycorhiza) si nm trong tbo

dng kt tht nt (knot) hoc cc (peloton).

Con ng xm nhp vo rca nm rni cng sinh:

1. Thit lp im tip xc v pht trin dc theo bmt ca r.

2.

Tip theo, mt hoc nhiu si nm hnh thnh chphng gia cc tbo biu

b. Sxm nhp rxy ra khi si nm tchsng phng thm nhp vo tbo

biu b hoc tbo vvo r.

1 2 3

Hnh 1.5.Con ng xm nhp ca nm rni cng sinh: 1.Si nm t hnh thnh 2

chphng gia cc tbo biu b (mi tn), 2.Si nm ti mt im vo (E) xm nhp

cc tbo v(mi tn) khong 1 ngy sau khi tip xc vi gc, 3.Cc tbo di (L) vngn (S) ca rthm nhp vo tbo ngn (mi tn).

2.2.2.3. Nm rni ngoi cng sinh (Ectoendo mycorrhiza)

Nm rni ngoi cng sinh c c trng ca chai loi trn. Chng thng c r

cc cy thng, cng l, quyn qumng v cy thuc hlan thy tinh.


17/24

17

Hnh 1.6.c trng bngoi ca rc nm cng sinh thsi nm v bo tngoi r:1.Nhn, 2.Dng li, 3.Dng ht, 4.Dng lng nhung, 5.Dng bng, 6.Dng

lng, 7.Dng gai ngn, 8.Dng gai di.

2.2.3. Cc yu tnh hng n nm rcng sinh

2.2.3.1. Tnh chuyn ha ca r

Do schn lc v tnh thch nghi khc nhau, phm vi tn ti cy chkhc nhau.

Mt sloi nm hnh thnh trn nhiu loi cy. Mt sloi chcng sinh trn vi loicy. Theo Wu renjian (1983) chia nhng loi sng trn 5 loi cy gm 10 loi thuc

Amanita, Cantharenlus; nhng loi sng trn 2-3 loi cy nh Boletus, Ramaria; ch

sng trn mt loi cy Suilus, Russula.

Cng mt loi nm nhng chng khc nhau cng c tnh chuyn ha khc nhau.

Nm Lactarius delicisus c 3 kiu chuyn ha khc nhau (thng, vn sam v lnh

sam).Nhng c loi phm vi cy chrt hp nh nm bng mu trng chc cy

bch n.

Nhng loi cy khc nhau cng la chn nhng loi nm thch hp. Mt s loi

cy c thdng nhiu loi nm khc nhau hnh thnh rnm nh thng 5 l; bch

n khng c tnh chuyn ha i vi nm r. Bch n c tnhin c 400 loi nm

cng sinh.

Ngc li c nhiu loi nm rc tnh chuyn ha rt mnh, nh nm ngoi sinh

trn cy long no, nm cnga ht u cy ln cy non gim hom khng hnh thnh

nm cng sinh.


18/24

18

Nhng loi cy khc nhau, cc giai on pht trin khc nhau loi nm cng sinh

cng khng hon ton nh nhau. Ni chung nhng cy rng non lng rnm rt t,

cy rng trng thnh sloi chim nhiu nht v rng qu gi li rt t. Khng nhng

thchng cn biu hin sgiao thoa nhau v dtrn cy bch n v cy dng, lc

cy cn non ssut hin ca nm ni sinh l chnh, nhng n tui ln nm ngoi sinh

chim u th, v cui cng nm ngoi cng sinh thay th. Nguyn nhn ca hin tng

ny c th l do c tnh sinh l ca cy cc giai on khc nhau.

2.2.3.2. c tnh sinh l nm r

i vi nm rngoi cng sinh

a. Ngun carbon

Ngun cacbon chyu l lignin v cenllulose. Nhng loi nm khc nhau c nhucu vngun cacbon khng ging nhau. Phn ln nm ngoi cng sinh cn ng n

glucose, maltose, xenlulose v mt st loi cn ng a nh tinh bt.

Cng trong mt chi nm, nhu cu vhp cht cacbon cng khng ging nhau nh

Tricholoma cn ng n, nhng Tricholoma fumosum cn lignin v xenlulose,

Tricholoma decorum cn xenlulose, Tricholoma matsutake cn glucose, tinh bt v

pectin.

Khng nhng th, cng mt giai on sinh trng khc nhau, nhu cu dinh dng

cng khc nhau.

Nm ngoi cng sinh sdng ngun cacbon do brcung cp. Mt sloi c th

tit ra enzyme phn gii ng. Cho nn hp cht cacbon trong rc hay khng, t

hay nhiu c lin htrc tip n sinh trng pht trin ca r.

b. Ngun Nit

Ni chung nm khng sdng nit v c. Nhng trong mt strng hp c thsdng nit v c tng hp thnh cht hu c. Nm rngoi sinh vc bn cng

nh vy.

Phn ln cc loi nm rngun m amon dhp thu hn so vi m nitrat. Theo

Guo Xiuzhen (1989) th nghim 14 loi nm cng sinh trn mi trng MMN cho

rng bt men, cao tht b v pepton, nm d s dng, si nm sinh trng rt tt.

Nhng i vi m ur th khng sdng c.


19/24

19

Cc loi nm khc nhau nhu cu vngun nit cng khc nhau. Mt s kin cho

rng nm cng sinh c tc dng cnh nit nhng cha xc nh c hot tnh c

nh nit r rt.

c. Dinh dng khong

Trong dinh dng, P l thnh phn quan trng nht. Trong nucleotide v

nucleotide phosphoride l thnh phn quan trng khng ththiu c. Nhu cu vP

quan trng hn ccc cht khc. Khng chvy, nm rc thlm cho cy hp th

c P trong khng gian m rkhng thhp thu c, n tham gia vo vng tun

hon P trong c ththc vt. V vy, P l mt loi dinh dng v cng quan trng

hnh thnh rnm.

Trong mi trng tng hp P v c lm ngun P nh KH2PO4, K2HPO4. Phn ln

P hu c trong t l ngun dinh dng P cho nm r.

Lng cc cht khc nh Ca, S, K, Mg rnm cn khng nhiu lm. Tng nng

l 4M/1000 lt. Cc cht vi lng nh Cu, Zn, Fe, Mn, B, Mo c thcung cp cho r

nm hp thnhng nhu cu mt lng rt b, khong 0,001- 7mg/lit.

d. Cc cht sinh trng

Trong qu trnh sinh trng pht trin nm r yu cu cht sinh trng nh

vitamin, cht kch thch v nhn tsinh trng khc. Cc cht c nhn tmitrng m nm khng tng hp c.

i vi nm rni cng sinh

a. Ngun carbon

Nm ni cng sinh cn dinh dng tngoi vo. Nh nm ngoi cng sinh nhn

c hp cht cacbon tbr, thng qua nguyn tnh du C14chng minh c

nm ly hp cht cacbon do cy quang hp, chyu l tp trung vo git du, vch t

bo v trong protein, acid hu c v acid amin ca tbo nm r.

Trn thc t, nm rsinh trng pht trin cn hp cht cacbon, b rnon cng

cn hp cht dinh dng. Thi ku xm nhim nm ni cng sinh: rnm v cy

tranh ginh nhau hp cht cacbon, do khng dinh dng cacbon nm r tm thi

ngng li, khi hai bn sinh trng hin tng ny dn c ci thin, s tch ly

cacbon trong cy dn tng ln xc tin chai cng sinh trng (Guo Xiuzhen, 1989).

Sau khi hp thhp cht cacbon do cy quang hp, si nm bin chng thnh loiesterza hoc acid hu c. Nhng cht ny cy kh hp thu nhng theo schnh lch


20/24

20

nng , hp cht cacbon khng ngng trcy i vo tbo nm gip qu trnh trao

i ca bn thn nm rv lm cho nm v cy chcng pht trin.

b. Dinh dng P

Nm rni cng sinh cng nh nm ngoi sinh u c thht P trc tip tt,

sau khi chuyn ha cung cp cho cy. Mc d cc quan im sdng cht P kh tan

trong t vn cha ng nht, nhng cy c r nm ht P nhng khu vc m r

khng ht c l khng th khng tha nhn. Hot tnh ca enzyme phosphoraza

tng cao r rt (Allen, 1981), thsi nm cng cha phosphoraza u ly cht hu c

trc tip t t. Tc ht P ca si nm hn lng ht ca r gp 6 ln (Sander,

1923).

2.2.4. Vai tr ca nm ri vi cy trng

2.2.4.1. Mrng din tch hp thca rcy

Si nm cng sinh l c quan hp thu chyu ca rnm, t bit l vng thiu

photpho (P). Si nm ngoi cng sinh c thko di ra xung quanh rlm tng tc

ht P ln gp 6 ln. Tui thca thsi nm trong t tng i cao hn so vi lng

ht ca rcy. Slng thsi nm cng nhiu hn slng lng ht ca r.

Nhiu nghin cu cho thy si nm ngoi cng sinh v b nm hnh rca nm c

nga vcng c thko di 510 cm. Trong 1g t rng tng chiu di ca si nm

c thn 40 m. Cho nn trong gii tnhin san cho nhau ca thsi nm hnh

thnh mng li, hp cht cacbon ca cy nh m c sphn phi li, shp thu P

cng thng qua si nm m phn phi u cho cy rng.

2.2.4.2. Tng khnng hp thu P v dinh dng ca cy ch

Trong t u c lng P nht nh, lng P kh tan chim tlkh cao c thln

n 9599%, chc mt lng rt t P ha tan m cy c thhp thc.

Nm rngoi cng sinh tit enzyme phosphorase chuyn P khng tan thnh P ha

tan, cung cp cho cy. Hot tnh ca enzyme tng gp my ln so vi cy khng c r

nm. Ngoi ra, nm rngoi cng sinh c thsn sinh mui oxalate kt hp vi Fe,

Al, mui P khng tan trong t, t m lm tng khnng ht P ca rcy.

2.2.4.3. Shnh thnh cht kch thch sinh trng ca nm r

Trong qu trnh cng sinh vi r cy nm hnh thnh nhiu cht kch thch sinh

trng nh cht sinh trng tbo (auxin), cht phn chia tbo (cytokinin), vitamin


21/24

21

B1, indol-3acetic acid (IAA). Nhiu loi nm cng sinh u tit ra trc v sau khi

cng sinh vi cy.

2.2.4.4. Nm rnng cao sc chng chu ca cy

Nhiu nghin cu chng minh, sau khi nhim nm cho cy, cy chc khnng

chng kh hn, chng chu mn, nhit , m v pH cc oan, chng li iu kin

kim loi nng.

Khi trng rng trn ni vi t lsng ca cy con nhim nm cng sinh tng

14% trong iu kin kh hn tuyt i. Khnng chng chu cc iu kin khc cng

tng ln.

2.2.4.5. Nm rci thin mi trng quanh r

Kt quca nhiu nghin cu cho thy xung quanh rcy thng xut hin tng kt

dnh rng gp nhiu ln so vi cy khng c nm r, tng to ra khu trao i ion b

rtng khnng hp thu v vn chuyn cy ch, c li cho sinh trng pht trin ca

cy trng.

2.2.4.6. Nm rlm tng khnng khng bnh ca cy trng

Nm 1942, Davis pht hin nm rngoi cng sinh c thlm gim bt bnh hi

r. Cc nm 1968, 1982, 1994, nhiu tc giu cp n nm cng sinh c thgim

bnh thi cr thng xung 25%, chng khng chphng trbnh m cn tng khnng sinh trng ca cy con.

Theo Tang Ming (1994), pht hin sau khi bn nm cng sinh cho cy bch n

bnh kh xanh do vi khun (Pseudomonas solanacerum) t pht sinh, trong khi cy

khng c nm cng sinh t l bnh ln n 25%. Cc bnh tuyn trng, bnh mc

sng, bnh bu ru gim i r rt.


22/24

22

III. KT LUN

Vi sinh vt ni k sinh thc vt (Endophytic) c tm thy trong hu ht cc

loi thc vt, chng c tr trong ni m ca thc vt k ch v gia chng hnh

thnh mt lot cc mi quan h khc nhau nh cng sinh tng h, cng sinh dinh

dng, hi sinh Hu ht cc dng ni k sinh ny bt u xut hin t vng r hay

b mt l, tuy nhin, mt s loi c th k sinh trn ht. Endophytic thc y thc vt

tng trng, tng nng sut v ng vai tr l mt tc nhn iu ha sinh hc.

Endophytic sn xut hng lot cc sn phm t nhin c li cho thc vt k ch m ta

c th khai thc nhng tc nhn ng dng trong y hc, nng nghip hay cng

nghip. Ngoi ra n cn c tim nng loi b cc cht gy nhim trong t bng cch

tng cng kh nng kh c trn thc vt v lm cho t tr nn mu m thng quachu trnh photphat v c nh m. Ngy cng c nhiu quan tm trong vic pht trin

cc ng dng tim nng cng ngh sinh hc ca Endophytic pht trin cc ging

cy trng c kh nng kh c ng thi c kh nng sn xut sinh khi v nhin liu

sinh hc.


23/24

23

TI LIU THAM KHO1. Trn VnMo, 2004. Sdng vi sinh vt c ch. Nh xut bn Nng nghip, H Ni

2. Azevedo JL, Maccheroni J Jr, Pereira O & Ara WL (2000) Endophytic microorganisms: a review on

insect control and recent advances on tropical plants. Electr J Biotech 3: 4065.

3.

Barac T, Taghavi S, Borremans B, Provoost A, Oeyen L, Colpaert JV, Vangronsveld J & Van DerLelie D (2004) Engineered endophytic bacteria improve phyto-remediation of water-soluble, volatile,

organic pollutants.Nat Biotechnol22: 583588.

4. Bent E & Chanway CP (1998) The growth-promoting effects of a bacterial endophyte on lodgepole

pine are partially inhibited by the presence of other rhizobacteria. Can J Microbiol44: 980988.

5. Berg G & Hallmann J (2006) Control of plant pathogenic fungi with bacterial endophytes.Microbial

Root Endophytes(SchulzBJE,BoyleCJC & SieberTN, eds), pp. 5369. Springer-Verlag, Berlin.

6. Berg G, Eberl L & Hartmann A (2005) The rhizosphere as a reservoir for opportunistic human

pathogenic bacteria.Environ Microbiol7: 16731685.

7. Chanway CP (1997) Inoculation of tree roots with plant growth promoting soil bacteria: an emerging

technology for reforestation. Forest Sci 43: 99112.

8. Duijff BJ, Gianinazzi-Pearsonand V & Lemanceau P (1997) Involvement of the outer membrane

lipopolysaccharides in the endophytic colonization of tomato roots by biocontrol Pseudomonas

fluorescensstrain WCS417r.New Phytol135: 325334.

9. Elvira-Recuenco M & Van Vuurde JWL (2000) Natural incidence of endophytic bacteria in pea

cultivars under field conditions. Can J Microbiol46: 10361041.

10.Germaine K, Keogh E, Borremans B et al. (2004) Colonisation of poplar trees bygfpexpressing

bacterial endophytes.FEMS Microbiol Ecol48: 109118.

11.Germaine K, Liu X, Cabellos G, Hogan J, Ryan D & Dowling DN (2006) Bacterial endophyte-

enhanced phyto-remediation of the organochlorine herbicide 2,4-dichlorophenoxyacetic acid.FEMS

Microbiol Ecol57: 302310.12. Habte M., N. W. Osorio, 2001. Arbuscular Mycorrhizas: Producing and applying Arbuscular

Mycorrhizal Inoculum, University of Hawaii at Manoa.

13.

14.Hallmann J, Quadt-Hallmann A, Mahaffee WF & Kloepper JW (1997) Bacterial endophytes in

agricultural crops. Can J Microbiol43:895914.15.Hallmann J, Quadt-Hallmann A, Rodrguez-Kbana R & Kloepper JW (1998) Interactions

betweenMeloidogyne incognitaand endophytic bacteria in cotton and cucumber. Soil Biol

Biochem30: 925937.

16.Krishnamurthy K & Gnanamanickam SS (1997) Biological control of sheath blight of rice: induction

of systemic resistance in rice by plant-associated Pseudomonas spp. Curr Sci 72: 331334.

17.Kuiper I, Lagendijk EL, Bloemberg GV & Lugtenberg BJ (2004) Rhizoremediation: a beneficial

plantmicrobe interaction. Mol Plant Microbe Interact 17: 615.

18.

Lindow SE & Brandl MT (2003) Microbiology of the phyllosphere.Appl EnvironMicrobiol69: 18751883.


24/24

19.Lodewyckx C, Vangronsveld J, Porteous F, Moore ERB, Taghavi S, Mezgeay M & Van Der Lelie

D (2002) Endophytic bacteria and their potential applications. Crit Rev Plant Sci21: 583606.

20.Miche L & Balandreau J (2001) Effects of rice seed surface sterilization with hypochlorite on

inoculatedBurkholderia vietnamiensis.Appl Environ Microbiol67: 30463052.

21.Porteous-Moore F, Barac T, Borremans B, Oeyen L, Vangronsveld J, Van Der Lelie D, Campbell

D & Moore ERB (2006) Endophytic bacterial diversity in poplar trees growing on a BTEX-

contaminated site: the characterisation of isolates with potential to enhance phytoremediation. Sys

App Micro 29: 539556.

22.Posada F & Vega FE (2005) Establishment of the fungal entomopathogenBeauveria

bassiana(Ascomycota: Hypocreales) as an endophyte in cocoa seedlings (Theobroma

cacao).Mycologia97: 11951200.

23.Reiter B & Sessitsch A (2006) Bacterial endophytes of the wildflower Crocus albiflorusanalyzed by

characterization of isolates and by a cultivation-independent approach. Can J Microbiol52: 140149.

24.

Robert P. Ryan, Kieran Germaine,Ashley Franks, David J. Ryan and David N., 2008 Dowling

Bacterial endophytes: recent developments and applications,FEMS Microbiology Letters, Vol 278-1

25.Siciliano S, Fortin N, Himoc N et al. (2001) Selection of specific endophytic bacterial genotypes by

plants in response to soil contamination.Appl Environ Microbiol67: 24692475.

26.Strobel G, Daisy B, Castillo U & Harper J (2004) Natural products from endophytic

microorganisms.J Nat Prod67: 257268.

27.Sturz AV & Matheson BG (1996) Populations of endophytic bacteria which influence host-resistance

toErwinia-induced bacterial soft rot in potato tubers.Plant Soil184: 265271.

28.

Zinniel DK, Lambrecht P, Harris BN et al. (2002) Isolation and characterization of endophyticcolonizing bacteria from agronomic crops and prairie plants.Appl Environ Microbiol68: 21982208.

29.http://www.bio-organics.com/mycorrhizae-products/

30.http://s4.zetaboards.com/BioFood_Tech/topic/8391205/1/

31.

http://www.mycoroot.com/research.html

32.http://www.soilmoist.com/products/mycorrhizal.php

33.http://www.rodaleinstitute.org/20101206_a-complete-how-to-on-farm-am-fungus-inoculum-

production

Documents

Vi Sinh Vật Nội Sinh