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Avaliação da interação entre Methylobacterium spp. e citros

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Methylobacterium

! " # $

% & ' ( )

* + %

(2)

,-.-Methylobacterium

/ 01)!21 03 $ 4 562

! " # " " $ " % &

(3)
(4)
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'

4 *2

Methylobacterium

" ( " " ) * + + +

" " , - , " ! "

! Methylobacterium " "

& , " .( " " +

" / " , " 0 "& *

, , , " ! "

" " " Xylella fastidiosa 1 " & " * "

Methylobacterium( " " " 2 )

Quorum Sensing 3456, * 0 7( " (& " 3 89 6 "

" 0 , , " , ,

" " " 0 " 7 ,

" + + Methylobacterium( "

" , & " ,

, Methylobacterium " , +

" " " : + ) ! " 7

" ) , + " * " " " ( " &

" "

" + + &

+ , ; ) " !

" + + " " " Methylobacterium

< , " "

& 0 " " = Methylobacterium

" " , M. extorquens > ?/@ "

" ) " 3/ 6 " "

A " + , + " * " " " 0 "

Catharanthus roseus, ) " "

" X. fastidiosa , * " 2 " X. fastidiosa,

" " & ) : , * " 0

" & " /

89 ) + + M.

mesophilicum 5> ?/?( , + * "

+ ) mxaB, " "

" " C pat, " + +

" + " 0 C acd5, + + "

" + & ) crt;

sss, + + " " , " + ,

phoD, " " " ,

89 " - + & *

Methylobacterium " : Citrus ,

" " " " " B +

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E

" * & ) " ,

Methylobacterium " ( " ) 89

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F

!4 !

7 + " 8 Methylobacterium

& " ( " " ) " & + + + "

" " & G , " & " &

Methylobacterium & " &G " " &

; " & G H & " &G & ;

" , & &G " 0 & & ( "& , & G "

& " G " " X. fastidiosa ;

I H & & " Methylobacterium( " " " G

H& ) G & 4 5 3456, H& "& 7( "G

& " 3 89 6 " ; " " H &

& , G G " " 0

8 H + , & "& Methylobacterium( " G

& , & H I H G , & " H

Methylobacterium " , + & " & " &

H & & + ) ; & " ) , H &

& " " G " ( " & " " &

" &

+ G, ; " " ) H & 2

+ + "& " H & G Methylobacterium &

" & 0 G " G

Methylobacterium " &G & " ,M. extorquens > ?/@

H " G ) " 3/ 6 0G D

" " " " G H + & & " " 0 &

" & Catharanthus roseus, ) H &

&G " " X. fastidiosa B & , H& " H & X. fastidiosa,

& " & & )G , & & " 0

& & & , & " " " 5 G & " 89

& ) + + & " H M. mesophilicum 5> ?/?(

H + & & " & " H & " +

) mxaB 3 & & &G & "

6C pat 3 + " + + &

" 0 6Cacd5 3 + + " & +

& 6 /) crt; sss, + + "

" , " + G, phoD, H & & " G H &

" 89 & + " & & G &

Methylobacterium " H & Citrus , & H " " G

H " &G " " , H + & &

" "" G H & & " " + , G H &

& " & & & )G " , & " Methylobacterium(

" ()G " G G 89

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K

. 01!42$ 9:2

/ 0 " - " (

" " , + + " "

" , * " 0 " 3" 0 " 6C

" + 3" 0 " 6C " " "

3" 0 ! " 6, *

-+ / " , " ! "

0 " " " , &

- " , * "

+ " "

/ " ! * " " " ( " /

" ( Quorum

Sensing 3456 / " +

* < " " + "

7( " (& " 3 89 6 " 0

" $ ( + , " <* * " " "

89 , " , ,

"& + + +! " : " " ,

+ " " - ! " " % " *

0 89 ! " , + "

" " " " " " "

7 L " " " ! ,

" " 0 " " "

" , " " + , .

, / + ) &

Methylobacterium ! " " *

" " " / " ,

" " , " " " !

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M

" " Methylobacterium 0 ( "& ,

) " + , "

"& + " " "

Methylobacterium "

" , + * ) "

" " + , + " " "

+ " " + , +

& A ) ! , & , , + /

* " " ; : " < " " ,

+ 0 * " ( " "

) " 2

" ) , & + " 2 + 6 +

" " Methylobacterium + " " " C 6 "

" Methylobacterium ) & +

" X. fastidiosa " " " C 6 +

(11)

N

, 42*29:2 $ 4 0* 1!2 $ 0!42 24Methylobacterium

4 %

& Methylobacterium " Citrus ,

Capsicum annuum 9 Saccharum officinarum 9 , 0

" " ( ) " 3 (" + ( Citrus limonia

"I I O Citrus sunki6, + + + " " ,

+ + ! ) " 0

" " "

(" + I , "

I 4 " 0 0 ,

+ " " ( ) 0 /)"

+ & " 3 > ?/@6,

(" + / I , +

+ + , + 0

" " + +

, + " & , * *

" * & + " " (

) , * 4 " "

, I

. * * " " 0

, & " > ?/@, > ?/ 5> ?/?,

& E/@ + " " ( )

" " " " " " I

Methylobacterium +

7 & : " + .

" " " " Methylobacterium + ;

" " ) " ! , + "

+ + " " " "

" . > > " *

Methylobacterium " 0 " ( ) "

+ " " " "

" (" + I , Methylobacterium

" & 0 " "

* Methylobacterium " 0 " & !

" " " " " " - " "

+ ("& + Methylobacterium C Citrus C ; " ( C

(12)

P

, 0!4 )42/!; 42*2!021 <Methylobacterium

"

& G &G " Methylobacterium Citrus

, Capsicum annuum 9 Saccharum officinarum 9 , " H & "I 3Citrus limonia "I Citrus sunki6, H

+ " " G " B & , & )

H + H & H " 0

" & " & & & H H &

Citrus limonia "I Citrus sunki, & " H G

H & Citrus sunki D " 0 & G, & H

& H + , & "I /)" & + "

G " 3 > ?/@6, & Q " & Citrus

limonia "I ; Citrus sunki, H + + "

7 & , " " &

"" & + , H +

& & " + H & & "I , &

& & & " Q & & Citrus sunki H

H& & " H " " G & & H& "

G & " > ?/@, > ?/ 5> ?/?, Capsicum

annuum 9 E/@ H & "I " " G

" & + " I H &, " GMethylobacterium

, H + G " 7 " H + &

G " "& " " Methylobacterium

+ ; " " ) , H& "& " &

"& + + H & H & & & "

> > G , H " & Methylobacterium

&G " G " "I + &

&G " " Citrus limonia "I Citrus sunki ,

Methylobacterium & 0 G " &

& Methylobacterium " 0 &G " G & & & +

" H & " " G "

(13)

, . 0

" " < ! "& +

& , 2 " &

+ & " "

" 38 99% 77 , NNKC 5 D>RC 7 S JC @PPP 6

B * , " " "

* " * " 0 0 3 ;99 TC 7 S J, NNK6

, " " " , + 0

* + )

" L " 35 D>RC 8>;5 ;/C 7 S J,

@PPP C S8; 5, @PP 6

" " , 2 ,

& " " + + ,

+ " + + % ) *

" + , * . + * + + & ,

38 > ;%C = 7 =/> //JC = 7 /95 5, @PPM6 / L

" = + ) " "

" - " , " &

3> 5/7 9D/ 8C % > ;7/R(> %/> , @PP?6 " ,

* " " + "

& , " " , + + , "

* + " 3 7> 8 , @PP?C 8 99 7 C 9 7$C

9 , @PP@C J9 / />C 9;B58; RC R 9 S; R, NMNC 5 D>RC 7 S J,

@PPP 6

" * " " & " ,

, , "

, " , " " ,

& A 0 3$9; J, NNFC J9 / />C 9;B58; RC R 9 S; R, NMNC

(14)

@

) ! " ! Methylobacterium " +. "

" " " =. +!

" " " " + ,

+ & A 3J /7;7$C % >>;5C 9 , @PP@C

9// , @PP?C % 8 ;T 7 , @PP?C %/> , @PPE 6 )

! 3% 8 ;T 7 , @PPEC % 8 ;T 7 , @PPN C 5T , @PP 6

, " " " , *

" " " + 3% 8 ;T 7 , @PPEC

% 8 ;T 7 , @PP?6

" " : " " " A " ,

& , "

) " . ME' @?? & " , " " /

5 , * " " KM,E@V " 3; $/, @P P6

9 + " " N PMP KFF

2 " + @P P, "

@@ ?@K 3; $/, @P P6

+ " " +. " " " *

) 37/=/5C >D%W 7/J, @PP?6 , : "

" A " + 0 + " ,

* " ( ( , 0

+ ) & * " , +

3 % > C =; /7 /C ;5 /99 , @PP 6, * , 2 "

* " " "

) + + " " 3 5 > C J/>5 /7, NN?6 " MFV

" " ) (" + 3Citrus limonia "k6 3 % /D

WX7; >, @PPF6 % ) + " ( ) ,

" & + " * * ! " : " <

" 3 >9 5, 5 D 8;C 7 ; , NNKC % /D WX7; >,

@PP 6

& + *

(15)

'

/ . " ( ) , +

" " " 0 * + + + <

" " " 3 > XW , @PP@C 9 = , @PPE6

" " + " , Methylobacterium

" "& "

, 2 + & +

Methylobacterium " " " " "

+ + " " , " " 6 +

Methylobacterium (" +

I C 6 + Methylobacterium " "

(" + I

4 7 &

7 (7J S , C %D5 8, % C 5 8;/>5 8, W C / 7/>, % C 5 8S ,

S % " " H Methylobacterium extorquens H &

, & " , " 0 & &

6 7 = % >, ) , + FK, F, EP@F(EP'@, @PP?

8;J >;, C W 5/ 8, % C T 7$, $ C 8;99; 5, C 7/95 7, 9 % /+

" > " H & " "

" 6 7 * " >, H , + P, N ?(N@E, @PP

9 5 8D9, 5 B C $;58, S C %;99/>, S C %T/>5, / S C 9; % 7, W " "

"& 6 7 * >, 9 , + @ F, EP'(E P,

NNP

% > , C =; /7 /, % % C ;5 /99 , 5 9 " "

? , , + @@, ('K, @PP

7 >/ /, B C >7/;> , > C 5 99/5, W B C % > 7, W C 9 /, C

R/=/ , W 9 C > XW , S 9 ( & 0 (

& " & + G Methylobacterium & & 0 &

& 0 " " G * " >, 7 H T I, + FK, M@(N',

@PPN

7 >/S5, W 8 C 8 >>;5, > B & " G &G "

" 4 8 7 +> + >, , + 'M, FE( MP, @PPP

> X9 , S 9 % " 7# %

(16)

E

' 3 $ " % & 6 O / " 5

" Y9 0 4 0Z, D + 5 , " " , @PPP

> XW , S 9 C % > 7, W C % 8/> 7; WD7; >, S C = 7 /95 5, W C = 7

=DD> /, W S 9 C R/=/ , W 9 + G &G " " &

" H & Xylella fastidiosa " %

* " >, , + ?M, ENP?(EN E, @PP@

> XW , S 9 C 5 >; J;5, 8 C >> 5 , = C $D;9 >(=;9 5 , ; C

R/=/ , W 9 = G " H &G " "

" "I 6 7 * " >@

H , + EK, @@N(@'?, @PP

5$8 >, 8 7 C R 8;>, R C >58 , % C J8 9;4, > & H +

" " ) G & 0 " & H &( " + Brassica

juncea9 D > 7 , , + 'F, @' (@'K, @PP@

55D% [\ , 9 C 9 = , C ; 5, B C R/=/ , W 9 C %/7 /7, W %

+ " " " " " "

2 E F , , + EE, F, FP'(F P,

@PPN

R/=/ , W 9 % " " ; %/9 , ; 5 C R/=/ , W 9 3/ 6

% " W L /% > (% , NNM " E, K(

'K

R/=/ , W 9 C % 8/> 7; WD7; >, S C />/;> , W C > XW , S 9

/ &G " " + H " " " + " "

6 7 + >, = , + ', EP(?F, @PPP

> 5 , W C % 98/;> 5, / C 7R , ! !

. " A " = @ P W " D , NN@

>$/5, > 5 C 9%/; , B W C 5 > 7 >;, C % 8 , % C >= 98 , 5 C

9/ B;98 , 8 C $D;9 >(=;9 5 , ; ; //% > / 7 *

" + < " " +

" ? , , + @ , @PF(@@E, @PPP

>; , W % C 5 J, > % C 5;9=/>5 7/, 5 > G " "

" " G " 0 "

% * " >, , + FK, F'F(F'M, NN

>9 5, / B C 5 D 8;, / 5 C 7 ; , 9 G =

(17)

F

5 > , % C J/>5 /7, / ; ! "

2 = ! " 3Citrus sinensis39 6 "I6 + " # , " " , F', + @', NN(@P', NN?

8 7S T, C 58;58; , % C 7 ;>7, W C WD7$S;> 8, 5 C % >J8 %, W C U; ,

$ C 8 99, B $ / &G " " 0 &G "

" H & H &( & 0 " D >

* % , , + '', M (MM, @PPP

7> 8, D / J/>5, $ W % C B9 >/5, = C $ > ^ ( $D5 ^7, C W J , $ C

% D 8, B C 7/S% 7, % C ;/ />5/, % W C ;755 , C R , % W C D$;7,

C 5 8 BB> 8, D C 7, W C S/7 /8/77/, C R;%%/>9;, 9 C % D 8(% 7;,

G * G* " 0 , 5 ,

+ N, N?@( PK , @PP?

/>/;7/>, W C 9 7;, = 9 C 9 7;, W 9 % 7

" H7 % /% > O 5 ;,

NNF, + ?P

/7/ J, 5 C S/;, $ C J9 />, W S / " H &(

& 0 " G "& D , S & , + EE,

E ( E', NNM

B/>>/;> , C 4D/ ;7/, % C 9 = , C , 5 C R/=/ , W W C > XW ,

S 9 + G &G " " Eucalyptus " " 0

GPantoea agglomerans. D * * " > , , + @MK, , M( E, @PPM

B>/; 5, 5 5 C $D;9 > =;9 5 , ; > 0 " " "

" " 4 & , = , + @M, NMK(NNE,

@PPE

$9/77, > C ;9S > 8, % W + & 0 * "

, ) , + @, ? ( ?K, NMF

$9; J, > & & " H & G ( + "

6 7 * " >, H , + E , PN( K, NNF

8 99% 77 W C 4D (8 99% 77, C % 8 BB//, S B C J9 / />, W S

" &G " " 6 7 * " >, H ,

+ E', MNF(N E, NNK

8 > ;%, > C = 7 =/> //J, 9 5 C = 7 /95 5, W "

&G & H & ! * " >,

(18)

?

8/D/>, 8 C 5% 99 , J " " & 3 $$/6

" & G " " ,

; = 7 /95 5, W , S/99;7$ 7, / % 8 , >/= >5, W 3/ 6* % " > 7 H T I % " II , NNK 'F'('K'

8 99 7 , % C 9 , W B% & " &

&G G & 2 _ 4 8 7 +> >

* >, , + EF, NK(@@P, NNE

8 99 7 , % "" 0 & G "G I " G

_ +> >, 7 + T I, + F, M?F(M?M, NNK

8 99 7 , % Methylobacterium 4 4 + $ % ,

+ , + , @PK(@ ', NNK

8 99 7 , % C 9 7$, > 9 $ C 9 , W Methylobacterium &G

" + + " ( I H & & _ ; 9;77 S, 5 / C 8/ 8 ( ;7 >,

/ ; C /99; , = W 3/ 6 +> + * " >, 5 , @PP@ @F( 'F

; $/ B O $ / " " . ( $ / /; $/,

/, $> ( 9 + 5 . " " , B + @P P

+

& ]]HHH + ]& ] " ] " ] " ] ] & `

" P@ , @P P

J $ , 8 C % 7 , 8 C 7 J , B C S 7 /, C J 7/J , 5 C % >;5 J;, 8 > "

" &G " " * " % , 5 , + @E,

FE( ?@, @PPN

J8 9; , C >58 , % C R 8;>, R 5" H &( & 0 "

+ H & G H& 6 7 * " >, ) , + N?,

EK'(EMP, @PPE

J 9T /= , % C R J8 > 8/7J , 7 5 C > 7;7 , 7 = C >DJ = 5 = , / C

;= 7 = , / $ C 9/J5//= , = = , > 5/7J;, T C D>T 7 =, T ; H &

& + G " + &G & " " 4

6 7 +> >, 7 H T I, + EM, F EOF K, @PP

J9 / />, W S C 9;B58; R, > C R 9 S; R, / B ( + "

& " " " + G ! + >, , + K, 'N(E',

NMN

J /7;7$, > 9 C % >>;5, > C 9 , W >7 & " H( + (

0 " Methylobacterium 6 7 >, , + ME,

(19)

K

9 = , C > XW , S 9 C % > 7, W C % 8/> 7; WD7; >, S C R/=/ ,

W 9 ; " H &G " " " & &G & "

" Xylella fastidiosa, " " + "& * " >, ) , + 'N FF(FN, @PPE

9//, 8 5 C % 8 ;T 7, % C J;%, S C 8 ;, 5 W C 8D7$, J T C 5 , % &G " & " G H & " 3Oryza sativa9 6 G

" &G & 7@( ) &G & " > D >

7 , , + E@, EP@(EPM, @PP?

9;B58; R, > C J9 / />, W S C J R9 S5J;, % C 5;% 75 7, C >95 7, W C

; ;7$, / % C R 9/5J , ; $ H & " 3 ( 6 G

Pseudomonas putida " " 6 7

* " >, H , + M, P@( P?, NMK

9 /ST JU, C = 7$> 75=/9 , W C > / D5, B C % >/, / > C $8 =;, 5 C %/R$/ T, % C = 7 /> 9/9;/, / &G " " & "

" 4 8 , , + @ , FM'(?P?, @PP@

% ;7> T, W C J9 / />, W S G " &G " "

H & H " " 6 7 * " >, H , + E ,

MNF(N , NNF

% 8 ;T 7, % C 7$DR8 9;, 5 C 5/7 8;9JD% >, % C 5/58 >;, 5 C 8D7$,

8 T C T 7$, W C 5D7 > %, 5 C 5 , % $ H & "

G " " " " + P(EK 3Oryza sativa9 6 GMethylobacterium

7 % , , + EF, E, ' F('@E, @PPE

% 8 ;T 7, % C 7$DR8 9;, 5 C 9//, 8 5 C 5D 7 > %, 5 I(

" + &G & " " "" , H & G

" " M?P'@ 3Saccharum officinarum9 6 > D > 7 ,

, + E , 'FP('FM, @PPF

% 8 ;T 7, % C 5D>/58 >/ T, = C 7 7 8 7, > C 5/7 8;9JD% >, % C

7$DR8 9;, 5 C 5D7 > %, 5 C 7$%;7 5 H &(

Methylobacterium " 3Arachis hypogaea 9 6

" H & & * " >, 7 H, T I, + F', @KP(@K?,

@PP?

% 8 ;T 7, % C J;%, T C 7$DR8 9;, 5 C JS 7, 5 S C 5 7$, % 8 C >TD,

W 8 C $ , 5 W C J , 5 C 5 , % Methylobacterium oryzae + , ", I(

, " + G &G & ", ( "G" (9(" )G

" " " 0 6 7 > %

(20)

M

% 8 ;T 7, % C 7$DR8 9;, 5 C JS 7, 5 S C 5 , % Methylobacterium

phyllosphaerae 7 + , I( , " + &G & &

&G & " 0 6 7 > % >

* " >, > L " , + FN, @@(@K, @PPN

% 8 ;T 7, % C 7$DR8 9;, 5C 5/7 8;9JD% >, % C 5D7 > %, 5 C 5 ,

7 ( H & G &G & " " "

* " 4 +, , + ?E, E( @P, @PPN

% 7 , 8 C % >;5 J;, 8 / &G " " " * "

% , 5 , + @', PN( K, @PPM

% 7 /5;7 5, / C 7 />> , C 5 , / C B> 7 15, W C 9/% 7T, W C

99 >/7 /, ; C % > $>/$ , " " & H

"& " & " 0 * " >,

" , + F, ?N( KF, @PP@

7/=/5, % B C >D%W 7/J, % 5 ?

( ,-.I @' + @PP? > " /% I /

+ aHHH " / * + b + / b / b

b @P Fb /75 ; 7/ ` " F @PPN

%/>, R 5 C % 9;7;, > C $/>8 > 5 7, " 0 G I(

" + &G & " " 3 B% 6 D * * " > >, , + E?, ' N('@?, @PPE

%/>, R 5 C % 9;7;, > C > />$, C $/>8 > 5 7, ; ('( " " "

" G I( " + &G & " " ) 8 +

4 , "& , + E', N'(N?, @PPE

;99 T, = J C 7 S J, W ; " G, , G "

+ H & &G " &G " " 0

3Lycopersicon esculentum96 " H & "

6 7 * " >, H , + E', 'FE('? , NNK

% /D WX7; >, W ( ) ; % 5 WX7; >, C / 7/$>;, W C ; ,

> % C % /D WX7; > , W ; B @PPF " E ? ( PE

% /D WX7; >, W > "I " & " " & 5 5

; 7$>/55 B 8/ ;7 />7 ; 7 9 5 ;/ T B ; >D5 7D>5/>T%/7, ? ,

@PP , > c / /) " ,

@PP KF(M@

> 7R , & " + & 0 " / 3Alnus

(21)

N

4D (8 99% 77, C J9 / />, W S ; " " " 0

& " &G Enterobacter asburiaeW%@@ "

6 7 * " >, H , + E@, EE( FE, NN?

4D/ ;7/, % " H " G G

J BSaccharum C @P P N? 3 ! " 6 O / "

5 " Y9 0 4 0Z, D + 5 , " " , @P P

> >;$D/R , 8 C B> $ , > & & 0 " &

H & + >, , + K, ' N(''N, NNN

> 5/7 9D/ 8, % C % > ;7/R(> %/> , / " &G &

" H & & * * " 0 , 5 , + N, M,

M@K(M'K, @PP?

> 55/ , 0 G G J " @P P

N? 3 $ " % & 6 O / " 5

" Y9 0 4 0Z, D + 5 , " " , @PPM

>TD, % C J;% W C 8 ;, C J;%, 5 8 C >J, 5 ; + " "

" " G Paenibacillus polymyxa/?M G

, , + 'N, @M@(@MN, @PP?

5 7 5, % 8 9 C % >; 7 , > 9 > C % > , > C 7 > /, $ C S;99 ;7 ,

9 C 9;% , $ " " " + + Heliconia

psittacorum9 B ; + , 5 , + '@, @, (M, @PPF 5/7 8;9JD% >, % C $ =;7 5 %T, = C 77 D>7 , J >

"& " G G &G Paenibacillus 8J ( F

* " >, 7 H T I, + FF, @F(@N, @PPK

5/7 8;9JD% >, % C % 8 ;T 7, % C 5D7 > %, 5 C J 77 ;T 7, 5

; " " 0 H & " Methylobacterium S &

( H & " 3Oryza sativa9 + (E'6 * " 4 +,

, + ?E, N@( PE, @PPN

5 /> , 7 C B>/; 5, 5 5 C %/9 , % C > 7;, / > 0 " "

" 0 0 " , " " " " 4

& , = , + 'P, @@F(@'E, @PP?

5 D>R, = C 7 S J, W / &G " " & 0 " &

* " G & " " H & " >,

, + F, M'( NP, @PPP

5 D>R, = C 8>;5 ;/, > C 7 S J, W " &G

+ G " " 4 8

(22)

@P

5T, C $;> D , / C W D> 7 , C $ > ; , 7 C S;99/%5, C / 9 WD ;/, C >;7, T C 7/T> , % C $;99;5, % C ;=;7(% 55 7, C >/TBD5, % &G & "

Methylobacterium " ) G H & 6

7 >, , + M', @ E(@@P, @PP

T % , 8 C 7 8 7T, C 9; 5 > %, % / "

mxa Methylobacterium extorquens % G " D *

* " > , , + ??, (K, NNM

= 7 J/7, C />/5, % C T, 5 9 C T %, W % C 5 87 >, W 9

Methylobacterium populi nov, + ", I( , " + G

&G & ", & ( 0 " 3Populus

deltoides)nigra 7'E6 0 6 7 > % >

* " >@ I , + FE, N ( N?, @PPE

= 7 9 7, 9 C JJ/>, 8 % 5 & 0 " ( " ;

J> 7, W =;55/>, W S 3/ 6 @PPE 4 > 5 = , @MK(

''P B/" " , + ?M6

=/$ , B / C = (>; 99, % C 5 , BC DT/>, W / &G " " Coffea arábica9 6 7 * " >, S & , + EF, 'K ('MP, @PPF

=/>% , 5 C 9 8 , W J C >; 8;, J /+ H &

" 0 G &G " 0 & H " 6 7

+ >, = , + N , @K( E , @PP

5 =J/9 = , / C 8/> T7 5/= , C ;7 , 5 $ C 7/ >5 =, ; "

" H & "& " " ) * "

4 +, , + ?@, ?N(K?, @PPK

S8; 5, W % % " " " & & 0 & 6 7

= % >, ) , + F@, EMK(F , @PP

R > 7 ;, % / C 7W >, 5 $ 8 C /8J /;, B C % 5 =;, 5 C B > J8;, > C

$8;$8;, 5 " " " 3Magnaporth oryzae6 G

Streptomyces sindeneusis @?' & % 6 7

(23)

@

K L 4 :2 ; ! 4M 2) $ βG.@NG 1$2) 0 1 * Methylobacterium extorquens 2 2103 9:2 1$2DO!0 $ Catharanthus roseus

4 %

Methylobacterium extorquens > ?]@ "

Citrus sinensis = , " Xylella fastidiosa. / "

" ) β( ,E( " 3/ 6

Bacillus pumilus. & , >/$9 , 0 Catharanthus

roseus, " , " " X. fastidiosa

3 & " = 6 & , +

" " A " + 3%/=6 B + * &

>/$9 " 0 " C. roseus, , " *

" & / &

>/$9 + & " 0 +

) * M. extorquens X. fastidiosa &

+ ) . " C. roseus, &

/ " * 0 Methylobacterium " =

. + .+ + < " " " " .

+ ("& + Methylobacterium extorquensCEglAC ; " ( C " CXylella fastidiosa

K ; ! 42 2)2 L 4 021 2D βG.@NG 1$2) 1 01

Methylobacterium extorquens 1$ 1$2 ;<!0 2 2103 !021 2D Catharanthus roseus

"

Methylobacterium extorquens > ?]@ H &G " "&

Citrus sinensis & H G = , " G Xylella fastidiosa &

" H " G ) & G "

3/ 6 Bacillus pumilus. & , >/$9 , H "

Catharanthus roseus, & " H & X. fastidiosa 3

" " G = 6 & & H + G "

" " " G 3%/=6 ; H + & & >/$9 " 0 &

" C. roseus, H& & , " & & H G G

G & & & , & H +

H & & )G + & " & M. extorquens X. fastidiosa " & & & )G C. roseus, H& & G " " H & & &

& " & G Methylobacterium "

(24)

@@

J GH Methylobacterium extorquensC / C " ( " C 5G " " CXylella fastidiosa

K . 0

0 + " *

" & " , 2 " /

" ( +.

& " " / " 7 " ) ,

" " ! Methylobacterium " "

" " 3 > XW , @PP C > XW , @PP@C 8 99 7 C

9 , NNEC %/>C % 9;7;C $/>8 > 5 7, @PPE6, +

" " + / " 3% 8 ;T 7 , @PP?C 9// ,

@PP?6 / " , Methylobacterium " " !

" " ( ) 3 > XW , @PP 6

Citrus , " = , "

" X. fastidiosa 3 > XW , @PP@6 , + 0

" " ! X. fastidiosa,

" " = "

5 & " "

" X. fastidiosa, = & "

Methylobacterium & / &

. + Methylobacterium , !

* . " 3 > XW , @PP@C 9 = ,

@PPEC 5 7 5, @PPF6 ,

Methylobacterium X. fastidiosa 0 in vitro, * ,

" , " "

Methylobacterium 39 = , @PPE6 , 3@PP?6

(25)

@'

" 0 C. roseus Nicotina clevelandii, "

Methylobacterium X. fastidiosa

) & " , + "

+! + + " "

) " * + " " "

0 " 3B 8/T , NN C 9 % /9 , NNEC

% 5;7 , NNF6 > " , % ) 9 3@PP 6 + + , "

M. extorquens % , + ) *

" mxaB 3 * " " 6,

" " * +

& " " 3 7 8 7TC $8 58C

9 J/ , NNEC 9; 5 > %C %D>>/99C 9 7, NN@C = >8 9 , @PP@6,

" Methylobacterium &

3 7 (J7 S , @PP?C 5T , @PPF6, " )

& Methylobacterium , " M. extorquens

" " " ,

" + @ 3 > 5/7J C ; 7 = C > 7;7 , @PP 6, & )

3 8 6 3 D>4D/C %/>9/ DC $> 9/ D, NNFC J > J = C

8;5 5/> > C 9; 5 > %, @PP@6, " 3= 7 ;/7 , @PP'6

& A 3J /7;7$C % >>;5C 99 , @PP@6 , "

Methylobacterium &

" , " " * " " "

& " , & 3@PPM6 "

) cry B. thuringienses M. extorquens "

Bombyx mori 7 , " " +

" * "

/ 0 " * " " * &

" " " " , * "

"& & " +

(26)

@E

X. fastidiosa " $ ( + )

& , * " = " 3 8 7$ , NN'6 "

+ 38 J;75, NMN6 1 + ,

" < " " , * * X. fastidiosa

+ ) " " " 3 D> /99C

B;79 T, NKNC D> /99C 8 J;75, NN?C > 55/ ; , NNP6 &

" " X. fastidiosa

) " 3/ 5 6 3 5;9= , @PP C 5;% 5 7 , @PPP6

+ . % " " / A " = 3B 6,

/ 5, 3 5;9= , @PP 6,

" " * , + < +

) . " * , " " + "

, " " , + ,

" , = , " &

& 3% 8 , NN@C % 8 , NNEC > 55/ ; ,

NNP6

B ( / " X. fastidiosa " + ) Citrus B / S J 2 3/5 94/D5 6

" +. " "

! " / 5 3 5 /> 7C 5 /S > C

$>//7 />$, NNN6, " " "

(27)

@F

/ 5 " "

" ! Bacillus ! " " " "

/ 5 7 I 3 NNN6 " + "

0 ) Bacillus & $9 , *

0 , ( ,E ( " , * "

0 9 3@PPF6, +

" " ! Bacillus 3 L2 , @PP 6 * <

" , " B. pumilus 9 ? " " +

, , , " 3egl 6 "

5 0 + L2 3@PP'6, " 0 "& * X.

fastidiosa " , Methylobacterium " "

0 & ) 0

" ) " " , $ 3@PPN6

" M. mesophilicum 5> ?/? )

" + 3$B 6, 0 " "

C. roseus, ) " 0 "

Bucephalogonia xanthophis, + X. fastidiosa pauca

* " +

& , " "& * C.

roseus , " " "

" " X. fastidiosa pauca / & 0

, M. extorquens > ?]@, & " , "

" ) $B C. roseus

* , " + " M. mesophilicum 5> ?/? 3$ ; , @PPN6, M.

extorquens > ?]@ " 0 & 2 + + M.

extorquens > ?]@ " " " " " X. fastidiosa,

" " )

(28)

@?

4 7 &

7 (7J S , C %D5 8, % C 5 8;/>5 8, W C / 7/>, % C 5 8S ,

S % " " H Methylobacterium extorquens H &

, & " , " 0 & &

6 7 = % >, ) , + FK, F, EP@F(EP'@, @PP?

7 >/ /, B C 9 = , C $ ;, 5 C > XW , S 9 C % 8/> 7;, S W> C

= 7 =/> //J, 9 5 C = 7 /95 5, W C R/=/ , W 9 % G &

" H Methylobacterium mesophilicum Xylella fastidiosa

6 7 * " >, H , + F@, E N(E@?, @PP?

7 8 7T, C $8 58, % C 9 J/ , B & " " &

&G * " G * *

+ % 6 , 9 , + 'PE, ??F(?KE, NNE

> XW , S 9 C% 8/> 7;, S W C $D;9 >(=;9 5 , ; C >> 5 , = C

5 >; J;5, 8 C R/=/ , W 9 = G " H &G "

" " "I 6 7

* " >, H , + EK, @@N−@'?, @PP

> XW , S 9 C % > 7, W C % 8/> 7; WD7; >, S C = 7 /95 5, W C = 7

=DD> /, W S 9 C R/=/ , W 9 + G &G " " &

" H & Xylella fastidiosa " %

* " >, , + ?M, ENP?(EN E, @PP@

R/=/ , W 9 > XW , S 9 $ " G " + &

& & " " 4 8 * 4 +, , + FEE, @@'(@'',

@PP'

D>4D/, C %/>9/ D, T C $> 9/ D, 8 &(" ( G " G(

β(&G )G G 3 8 6 & GMethylobacterium extorquens "

& &( " ( 8 * " > + >, , + EE,

'?K('K?, NNF

8 7$, W C $ >7;/>, % C R>/;J, 9 C > 55/ ;, = C =1, W %

" " & )G ( " " " + "&

" Xylella fastidiosa * " >, 7 H T I,

+ @K, 'K( E@, NN'

8 ;, T W C 9 S>/7 / $>;$ > /7, W C 19 7$/>, 9 C % >/9, 9 C D>4D/,

C % 55 7, 9 C $> 9/ D, C %^$D/R, " " " " G

Bacillus thuringiensis G & &G &Methylobacterium extorquens

(29)

@K

> /, S C >8//%, 5 /" G & &G & " " +

" D * * " > >, , + ?@, @E'(@FP, NMN

5 /> 7, W S C 5 /S > , 5 C $>//7 />$, / "

" " " , S & , + ME, ' M( '@@,

NNN

5;9= , B > C =/ >/, 9 C J/% />, / 9 C 9/; /, C >>D , B & Xylella fastidiosa ) G ""& G + + "

& " G D * * " > , , + @P', ?F( K , @PP

=;5, % W C B>/7 8, S W C 5 8 , 7 S ) " " " " H &

& G "& * " >, 7 H T I, 'PN('@E,

NM

/ 5 DR , C J; , % C />/;> , / C 9/ (B;98 , 8 C % 8 ,

% /) & " G( Xylella fastidiosa +

* " >, 7 H T I, + FP, @@'(@@M, @PPF

B 8/T, W S C ;% J, % > C % 5;7 , 5 B C T9 >, W % C >;5 7, 5

$ " G &G " " G G ;

7 >/S5, W 8 C 8;> 7 , 5 5 3/ 6 * " > 7 9 5 (

= , NN EP (E

B/>>/;> B;98 , 5 C 4D/ ;7/, % C $ 5, C > 55/ , C 9 = ,

C R/=/ , W 9 C > XW , S 9 / &G "Methylobacterium extorquens

) & G β( ,E( " Catharanthus roseus

= @P P

B;$D/;> , % % C 9 > %//, 9 C %D>>/99, W C $> 9/ D, %;$D/R,

" " G & &G & " "

Methylobacterium extorquens D * * " > , , + N', NFO@PP, @PPP

$ ;, 5 C 9 = , C 4D/ ;7/, % C D>; , % C 9 /5, W > 5 C > XW ,

S 9 C %;99/>, C R/=/ , W 9 Methylobacterium mesophilicum

GBucephalogonia xanthophis " " G =

& !+ 6 7 * " >, 5 , + EK, E, EEM(EFE, @PPN

8 99% 77, W , 4D ( 8 99% 77, , % 8 BB//, S B , J9 / />, W S

" &G " " 6 7 * " >, H ,

+ E', MNFON E, NNK

8 99 7 , % C 9 , W B% & " &

&G G & 2 _ 4 8 7 +> >

(30)

@M

8 J;75, 9 Xylella fastidiosa)G O " &

4 8 7 +> + >, , + @K, @K (@NP, NMN

J /7;7$, > 9 C % >>;5, > C 9 , W >7 & " H( + (

0 " Methylobacterium 6 7 >, , + ME,

M'@( ME@, @PP@

J > J = , 7 C 8;5 5/> = , 9 C 9; 5 > %, % / G( (&G )G G G & & " + &G &Methylobacterium extorquens %

" gap ,gap@P, pha> 6 7 >,

, + ME, ? KE(? M , @PP@

9 = , C > DW , S 9 C % 8/> 7; W> , S C R/=/ , W 9 > " " G Xylella fastidiosa, " " +

"& * " >, ) , + '', 'P@('P?, @PP

9 = , C > XW , S 9 C % > 7, W C % 8/> 7; WD7; >, S C R/=/ ,

W 9 ; " H &G " " " & &G & "

" Xylella fastidiosa, " " + "& * " >, ) , + 'N FF(FN, @PPE

9 = , C 9;, S C R/=/ , W 9 C 8 > D7$, W 5 > , " "

* + G & " & &G " " Methylobacterium

mesophilicum 6 7 * " > * + , , + ?F, F'F(

FE , @PP?

9 = , C > XW , S 9 C R/=/ , W 9 /+ &G " " 0

Citrus sinensis Catharanthus roseus G &G " " !+ 6 7 * " >, 5 , + EF, ( E, @PPK

9 % /9, W 5 C 7 />, W 9 C ;% J, % C J/99T, W 9 C 7 />5 7, W W C D> 7;, C B D9J/ W>, W 5 C D>7/>, W ; + " , )

G & cry 3"6 Bacillus thuringiensis Kurstaki "

Clavibacter xyli cynodontis % * " >,

S & , + ?P, FP (FPM, NNE

9//, 8 5 C % 8 ;T 7, % C J;%, S C 8 ;, 5 W C 8D7$, J T C 5 ,

% &G " & " G H & " 3Oryza sativa9 6 G

" &G & 7@( ) &G & " D > 7 , , + E@, EP@(EPM, @PP?

9;, S C >; W S C /;U/;> , C %;> 7 , = 5 C T>/5, W C B> 7 ,

B C 5 , % $ " "& " G Xylella fastidiosa

" , $ , 5 , + MF, F, FP (FPF, @PP

9; 5 > %, % / C %D>>/99, W C 9 7, 8 3/ 6 % & & 0

(31)

@N

9;% , C 4D/ ;7/, % C BD7$ > , % 8 C 7 >/ /, B C > XW , S 9 C 5;9= B;98 , % C ;RR;> 7;J9/;7/>, C R/=/ , W 9 % " " " 0

+ ( ,E " &G " Bacillus pumilus

+ >, , + ?M, FK(?F, @PPF

% 8 , % C >D5 , / C 4D $$; , W C 9 $ , % % C ; /99;, % C

BD>9 7;, > " - & " 2 "

= 4 D A , 9 , + ?, F'(

FK, NNE

% 8 , % C 5;9=1>; , W 9 C ;5 , > C >;5 B 7;, % C / B;9 (

5 >;78 , W + + < " +

" 3 = 6 ? , , + ', F F(F' , NN@

% 8 ;T 7, % C 5D>/58 >/ T, = C 7 7 8 7, > C 5/7 8;9JD% >, % C

7$DR8 9;, 5 C 5D7 > %, 5 C 7$%;7 5 H &(

Methylobacterium " 3Arachis hypogaea 9 6

" H & & * " >, 7 H T I, + F', E, @KP(@K?,

@PP?

% 8 ;T 7, % C 7$DR8 9;, 5C 5/7 8;9JD% >, % C 5D7 > %, 5 C 5 ,

7 ( H & G &G & " " "

* " 4 +, , + ?E, E( @P, @PPN

% >U, W C 9; 5 > %, % / + + + (& (

+ " &G & & $ ( + " * " >, 7 H

T I, + EK, E E(E N, @PP

%; 8/9;, B " &G " H 0G H &

&G G ! , J , + ?, E E(E N, @PP

%;99/>, 5G " " " " >%" , > & + , + E@, ?KOKE, @PPK

% 7;/>, W % C 9;7 S, 5 / + + G "

" " 7 + 1

%> 7 7 7 % @S & , + PP, FNKK(

FNM@, @PP'

% 7 /;> , C >/7 D ;7, W C W $ D/;U(/=/;9 > , 5 C T>/5, W C $ >7;/>,

% C =1, W % Catharanthus roseus, ) & & "

Xylella fastidiosa $ , 5 , + MF, @E?(@F , @PP

%D> 58;$/, C 5J $, B + H & G H &

(32)

'P

7 7J ;, 8 C 8 58;% , S C %;J;, 8 C J S ;, 5 C J D5 JD, % % " G

G ""& G 0 0G " ) & G 0 G

Bacillus 5 $9 % * " >, , + ?F,

?, @F@P(@F@?, NNN

7/%/ / J(% >58 99, % C % " 7 9 , > C B> 7R/7, W W C

S W ;/ 8 S5J;, 9 C B 99, > % & + / 0G " "

( & & & ) " " "

+ +> >, > "I+ , + PM, 'FN( '?M, NNF

7/S% 7, J 9 , 9%/; , > , D> /99, 8 , 9;7 S, 5 / D

" G Xylella fastidiosa" 0 Vitis vinifera

% * " >, , + ?N, K' NOK'@K, @PP'

%/>, R 5 C % 9;7;, > C > />$, C $/>8 > 5 7, ; ('( " " "

" G I( " + &G & " " ) 8 +

4 , "& , + E', N'(N?, @PPE

D> /99, 8 C B;79 T, 8 /+ " " " " d

" G & & & +> + >, 9 " , + ?N, 'N'(

'NF, NKN

D> /99, 8 C 8 J;75, 9 B )G ( " &

4 8 7 +> + >, , + 'E, ' ( F , NN?

4D (8 99% 77, C J9 / />, W S ; " " " 0

& " &G Enterobacter asburiae W%@@ "

6 7 * " >, H , + E@, EE( FE, NN?

4D (8 99% 77, C /78 % D, 7 C J9 / />, W S " &G

" "& & 6 7 * " >, H ,

+ E', FKK(FM@, NNK

> 55/ ;, = C $ >7;/>, % C =1, W % C />/ , % W $ C /U/;> , > > C

4D $$; W C / 7/$>;, W " " )G d

"& + , + %

4 0P % , , + ' P, 'EF('EN, NNP

>TD, % C J;% W C 8 ;, C J;%, 5 8 C >J, 5 ; + " "

" " G Paenibacillus polymyxa/?M G

, , + 'N, @M@(@MN, @PP?

5 7 5, 9 " H7

7 Curtobacterium flaccumfaciens 7#

" " @PPF @ 3 $ " % & 6 ( / "

(33)

'

5/7 8;9JD% >, % C $ =;7 5 %T, = C 77 D>7 , J >

"& " G G &G Paenibacillus 8J ( F

* " >, 7 H T I, + FF, @F(@N, @PPK

5;% 5 7, W $ C >/;7 8, B C >>D , C >/D, B C /7 ; , % C 9= >/7$ , > C 9=/5, 9 % C > T , W / C ; , $ 5 C ;5 5 C

>> 5, % 8 C 7 >5;, / C > ;7, 5 C =1, W % C >; 7/5, % > 5 C D/7 , % > C % >$ , C % >$ , 9 / C >> > , % C >>/>, 8 C

9 D , 7 C 9 % , C 5 , B B C 5 , % > C 5 (7/ , % C

D ;78 , 9 9 C >;5 B 7;, % C ; 5(7/ , / C /7 , C /;( >>T, 8 C B ;7 7;, C B/>>/;> , W 5 C B/>>/>; , = C B/>> , W C B> $ , W 5 C B> 7[ , 5 C B> 7 , % C B> 8%/, % C BD>9 7, 9 > C $ >7;/>, % C

$ 9 % 7, $ 8 C $ 9 % 7, % 8 5 C $ %/5, 5 9 C $>D />, C 8 , 9 C

8 8/;5/, W C WD74D/;> , % 9 C J/% />, / 9 C J; W;% , W C J>;/$/>, W / C JD> % /, / / C 9 ;$>/ , B C 9 % ;5, % > C 9/; /, 9 C 9/% 5, / $ % C

9/% 5, % = B C 9 /5, 5 C 9 /5, > C % 8 , W C % 8 , % C

% /;> , % 7 C % /;> , 8 % B C % >;7 , 9 C % >4D/5, % = C % > ;75,

/ 9 C % > ;75, / % B C % 5DJD% , T C %/7 J, B % C %;> , / C

%;T J;, T C % 75 /;> (=; >/99 , C % 7, 8 C 7 $ ;5, % C

7 5 ;%/7 , 9 C 7/ , 9 / 5 C 78 7; W , C 7 >/$ , B $ C 7D7/5, 9 C

9;=/;> , % C 9;=/;> , % C 9;=/;> , > C 9%;/>;, C >;5, C /;U , > C />/;> , $ C />/;> W , 8 C /54D/> , W C 4D $$; , > C > /> , $ C > >;$D/5, = C % > 5 , W C > 5 W , = / C 5#, > $ C 5 7 /99;, > = C 5 S 5 J;, 8 / C 5;9= , > C 5;9= , % C 5;9= , B > C 5;9= , S C 5;9=/;> , W B C 5;9=/5 >;, % 9 R C 5;4D/;> , S W C

5 DR , C 5 DR , C />/7R;, % B C >DBB;, C 5 ;, 5 % C 5D8 J ,%

8 C = 99 , 8 , = 7 59DT5, % C =/>W =5J;( 9%/; , 5 C =/ >/, 9 C

R $ , % C R R, % C %/; 7;5, W C 5/ D 9, W & * " &

& Xylella fastidiosa.1 , 7 H T I, + EP?, F ( FK, @PPP

5 > 9, W J % " 7# 7# ? BGlycine

maxC H7 G @PP' KE 3

$ " % & 6 O / " 5 " Y9 0

4 0Z, D + 5 , " " , @PP'

5T, C ;%%/>5, W C J7;/B, C = >8 9 , W % &G & "

+ Methylobacterium extorquens " 0 Medicago

truncatula " + " % * " >,

, + K , K@EF(K@F@, @PPF

(34)

'@

% 5;7 , 5 B C 9/;5 />, > C ;% J, % C / 8, > % C J/99T, W 9 B " Clavibacter xyli cynodontis ) & " "

cry 3"6 Bacillus thuringiensis / " "

, "& , + F, EE@(EEM, NNF

T % , 8 C 7 8 7T, C 9; 5 > %, % / "

mxa Methylobacterium extorquens % G " D *

* " > , , + ??, (K, NNM

> 5/7J , T C ; 7 = , / $ C > 7;7 , 7 = " &G & " "

&G G * " >, 7 H T I, + KP, ?@'(?'@, @PP

= 7 ;/7, 5 W C % >U, W C d >;/7, 7 C 9; 5 > %, % / $ " "& " 0 & " G & " & H G Methylobacterium

extorquens % " %

* " >, , + ?N, KF?'(KF??, @PP'

= >8 9 , W " ( &H G &G )

(35)

''

N 9:2 $ 1G 0 G;2*2 401 !21 1 L 4 :2 $ ) 1

1A2 A0$2 1 01! 4 9:2 1!4 Methylobacterium mesophilicum 4. Q////QG 1! ;2 $ 04

4 %

Methylobacterium " " " + *

" " 0 " " / "

* " * . " ,

" " " , " .(

" " + / " ; Methylobacterium

, " " " " , " 2

" Quorum sensing 3456 ,

" " + " ,

+ " " " , "

" ( 45 0 7( " (& " 3 89 6

" " 0 , " " " $ ( +

* + + " " , " !

Methylobacterium " ) , & + , >

* + , 356(7( " (859 )

+ + M. mesophilicum 5> ?/? &

, " + + " "

" " 3mxaB6,

" 3crt;6, + + " + " 0

3pat6, " + & 3acd56,

" " & " 3sss6, " " 3phoD6 7

" - + , + * ) mxaB, acd5 pat

+ 89, : " "

+ " " ( 7 , 89 "

! " ) crt;, sss phoD

" - 89

Methylobacterium

(36)

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N 1G < G;2*2 401 !21 !021 21 !; ) 1 L 4 021 $ 401) Methylobacterium mesophilicum 4. Q////QG 1! 01! 4 !021

"

Methylobacterium &G & " " + " & "

& " " & " " G

* " + , G " " H & " "

G & & " & ( H & / " &G &

Methylobacterium & &G & , " "

" " , H& " "" & & 4 5

G 3456 & , & " " " " &G " +

"& & , + " " + , "&

" ( " 5 45 G 7( "G (& "

3 89 6 " , " G $ ( + " & +

" H & , " & Methylobacterium S & &

" ) , & H I + , G * + >, & " 356(7(

" (859 & ) & " H M.

mesophilicum 5> ?/?( B & , H " &G & "

& & 3mxaB6, G & "

" 3crt;6, + " + + &

" 0 3pat6, " & + &

3acd56C " " 3sss6C & " G 3phoD6 D

& + " , H + & mxaB,acd5 patH + )

& " 89, & " & " & " (

" 8 H + , & 89 " & + " &

) crt;, sss phoD & +

& " G 89 Methylobacterium

(37)

'F

N . 0

Methylobacterium " $ ( + " < "

∝O@ " , " " + , " 0 " "

" " " 3 T % C 7 8 7TC

9; 5 > %, NNM6 / " * 0 , , *

" , * . " 3= 7 J/7 , @PPE6 "

" " +. " + 3 > XW , @PP@C % 8 ;T 7 ,

@PPN C % 8 ;T 7 , @PPN C 5/7 8;9JD% > , @PPNC 5T , @PP 6

4 " " , Methylobacterium +

" " + , +

& A ) ! 39// , @PP?C % 8 ;T 7 , @PPFC

%/>C % 9;7;C $/>8 > 5 7, @PPEC 5/7 8;9JD% > , @PPNC 5T ,

@PP 6 3B/>>/;> B;98 C

> XW C JD9;75JT( 5 > 9, @PP C % 8 ;T 7 , @PPEC % 8 ;T 7 ,

@PP?6 Methylobacterium "

3 > /C >8//%, NMN6, * " "

35T , @PPF6,

" Quorum Sensing 3456 & : "

" (

45 " + * " " 0 <

" " " "

+ " ; " "

" ) " * " " "

+ , , , < " " +

" " .( " 3 D/>C

% 8/5;D5, @PPEC W ;7 , @PP?C 5 7 8/5( 7 >/> 5 , @PPKC 5 C

5 7WD#7C 9;= >/5, @PP?6 ) + +

" ( " , 45

+ + " ( 3 >7 > , @PPKC 5 7 8/5(

(38)

'?

45 + + 7( " (& " 3 89 6 "

" / " " " $ ( + * + +

" " , " " 0

& 3 >7 > , @PPKC 8 , NNMC %;99;C 559/>, @PP?C 9 8

, @PP@C 5 7 8/5( 7 >/> 5 , @PPKC S8; /C S;7 75, @PPK6 ,

* 89 " + ,

" , & + ! " 3 D/>C

% 8/5;D5, @PPE6

89 " !

Methylobacterium + 3@PP?6 " M. extorquens %

45 + 89 "

" " " - 0& , % & G 5 3@PPK6

" " 0 89 Methylobacterium .

.+ , 0 > " , 3@PPN6 0 * "

89 0 M. mesophilicum5> ?/?, " 3 > XW ,

@PP@6 " - ? 89 , 356(7(

3@/,KR6( " (859 356(7(3@/6( " (859, 0

+ 0 0 " 89

" " " $ ( + " ( " " M.

mesophilicum 5> ?/?, + " * " "

" " " " , " * , " + ! "

M. mesophilicum 5> ?/? " "

" 89 ) X. fastidiosa

7 , +. & ! 0 / " 2

" Pseudomonas aeruginosa, " + 89, * ? ?

" + ) " " ,

@@@ 3S $7/> C @PP'6 . A " " Sinorhizobium

meliloti + * 89 ) " +

) 3 8/7 , @PP'6 5"&H 0 3@PPK6 * >,

* + 0 P.

(39)

'K

/ Methylobacterium, +. & : "

" " 39// , @PP?C % 8 ;T 7 , @PPN C % 8 ;T 7

, @PPN C ;> ;9 , @PPPC 5T , @PP 6, " " & "

89 ) + + /

* " 89

3 /7\9=/> , @PP?6, *

" 0 " Methylobacterium 3 7 >/ / , @PP?6

7 , " - +

& * " & " "

" ( + " )

" ! " 3 $ J;C 9 8C S;99; 75, @PPMC

S 7$ , @P P6 5 , " " & "

) " " 7

" ) , 3@P P6 + ) + +

M. mesophilicum 5> ?/?( + " * +

& < , + " /

" , " " )

" < " , * "

" + +

= - " " ) M.

mesophilicum 5> ?/?, 2 + + , * >,

356(7( " (859 ) mxaB, acd5,crt;,pat, sss phoD,

+ + " "

4 7 &

7 />5 7, W C % >>;5, W C 7D77, 7 C 7 8 7T, C 9; 5 > %, % /

7 " * " & Methylobacterium extorquens % moxB moxW

+ + & ) $ NP ; 7 8 7T, !+ + % > 7

(40)

'M

7 >/ /, B C 9 = , C $ ;, 5 C > XW , S 9 C % 8/> 7;, S C = 7

=/> //J, 9 5 C = 7 /95 5, W C R/=/ , W 9 % G &

" H Methylobacterium mesophilicum Xylella fastidiosa

6 7 * " >@ H , + F@@ E N(E@?, @PP?

7 8 7T, C $8 58, % C 9 J/ , B & " " &

&G * " G * *

+ % 6 , 9 , + 'PE, ??F(?KE, NNE

> XW , S 9 C % > 7, W C % 8/> 7; WD7; >, S C = 7 /95 5, W C = 7

=DD> /, W S 9 C R/=/ , W 9 + G &G " " &

" H & Xylella fastidiosa " %

* " >, , + ?M, ENP?(EN E, @PP@

7/>W;, 5 C B9;/$/>, Patatin(9 I H G G " 0G

" _* " >, > , + FP, F@@(F@F, @PPE

7/>W;, 5 C D> 55, C B9;/$/>, & & & Legionella

pneumophilaO " ) , , & I " + "

0 6 7 * * " >, 8 , + @NM, ?N( M , @PPM >7 > , % 9 C S S7, 5 C D>>, C D9 8D>5 , 5 W C % 75 7, > / C

5 9% 7 , $ 4 , + " " G "

( " + + ! + 4 > > , 9 ,

+ '?@, ?F( M', @PPK

D/>, S C % 8/5;D5, D " *

2 >, 9 , + K, E@N(E'', @PPE

$ J;, C 9 8, J 8 C S;99; %5, J W /)

" H & & G(Pyrenophora "

" * + >, 9 , + N, ?EF(??P, @PPM

%;99;, C 559/>, 9 " ( " &H G ,

S & , + ' , '( ?, @PP?

8 , C $ , C 8/7, T C 58 S, C B >> 7 , 5 J " "G (

& " * ( G $ ( + ( "

" * * " 0 , 5 , + , N( @N, NNM

8/7$, C /77 7 , 5 % C RR > ;, J ; C /77/ (S , = C 8 > 9 7 ,

/ 9 C > ;75( > S7/, > % C D5 8/J, % & pst(phoD

" & " G G " & "Escherichia coli + "

(41)

'N

8/7, 8 C / 9; 5J;, % C > ;75 7, W C > 9B/, $ C D/>, S " G H ( G Sinorhizobium meliloti N( "G & "

* ( & G & 6 7

>, , + MF, FP@N(FP'?, @PP'

8;77 D> ;, C 9 8 7 >, C 5D7 > %, 5 & " 0 (

"G" ( (" )G " &G "

&G & " & &G / 6 7

* " > + >, ) , + @F, EP'( E , @PPN

> /, S C >8//%, 5 /" G & &G & " " +

" D * * " > >, , + ?@, @E'(@FP, NMN

D> , % 7 * +> " % @P P '@

3% $ " % & 6 O / " 5 "

Y9 0 4 0Z, D + 5 , " " , @P P

B/>>/;> B;98 , 5 C > XW , S 9 C JD9;75JT( 5 > 9, W " "

Methylobacterium " & ; 5;% e5;

;7 />7 ; 7 9 / ;7; ; [\ ;/7 ^B; D5 , " " 4 % R5

D5 , @PP

$9; J, > C > =; , C R >7T, W C 8/7$, R C D 7, W C %" 7J/T,

H & G " 4 8

, " > , + @?, @@K(@E@, @PPK

$ D>; 7, C B> 7 /R( 8 >9 , C = >8 9 , W &G> ; + + &

Methylobacterium extorquens % 6 7 >, , + ', P@K( P'F, @PPM

$>;5 S , C B;7/> 7, C /=/>5 7, 9 C S;99; %5 7, 7 > C 5 9% 7 ,

$ & & > H (" G " G & Serratia

& & * * " >, 9 , + @, MP( @, @PPN

8 > ;%, > C = 7 =/> //J, 9 5 C = 7 /95 5, W "

&G & H & ! * " >,

, + ?, P, E?'(EK , @PPM

W ;7 , ; " " + I , + 7/>

" % & & > G 5 " G K " @PPF 6 7 4 >

> 0 7 , 9 , + ', EFNOE?', @PP?

W D> 7 , C >/7;/>, C > ; >, 5 C B >; , 5 % C >;7, T C $ 9; 7 , C

$;> D , / C >/TBD5, > &G &G G H

Methylobacterium nodulans Crotalaria podocarpa * G* "

0 , 5 , + M, P? ( P?M, @PPF

(42)

EP

W;% , 5 C 5D$ 7D% , 7 C 5DRDJ;, C J;, C 8 T 58;, % C T J T % , C

8T % , C 5 %;RD, / C JDS , C 58; , C , 5 9 ( "

G ) G

,Lotus japonicus $1 4 +, IG , + , @?'(@KE, @PPE 9 = , C 9;, S C R/=/ , W 9 C 8 > D7$, W 5 > , " "

* + G & " & &G " " Methylobacterium

mesophilicum 6 7 * " > * + , , + ?F, F'F(

FE , @PP?

9 % > 8/, % $ C S 77/>, 9 C >1 ;75, 5 C 8 >/9, W & & & " + " H I " " & & &

& D * * " > 4 8, , + '@, E? (EK', @PPM

9//, 8 5 C % 8 ;T 7, % C J;%, S C 8 ;, 5 W C 8D7$, J T C 5 , %

&G " & " G H & " 3Oryza sativa9 6 G

" &G & 7@( ) &G & " > D >

7 , , + E@, EP@(EPM, @PP?

9;, T C R8 7$, T PhoD " H & + +

" " Escherichia coli % "

+ % + >, S & , + F , @PN@(@PNN, @PPK

9; 5 > %, % / C 8;5 5/> = , 9 & I G I " G

Methylobacterium 6 7 >, , + ME, M M, @PP'

9 8, W C >95 7, > S C T >J, S 5 C 5 /T, $ G ) , * "& "

+ + G " 1 , S & , + NN, EE O E?,

@PP@

9 9J/% , W 5 C 59 %, W & 2 " G &

" " 7 f( " 7 f( " *

* %" >, , + @F, F?K(FKP, @PPM

% " 7 9 , > C B 99, > " &

& & % + % ) ! , ) ,

+ @K, KPN( K ', NN'

% 8 ;T 7, % C 7$DR8 9;, 5 C 5/7 8;9JD% >, % C 5/58 >;, 5 C 8D7$,

8 T C T 7$, W C 5D7 > %, 5 C 5 , % $ H & "

G " " " " + P(EK 3Oryza sativa9 6 GMethylobacterium

7 % , , + EF, E, ' F('@E, @PPE

% 8 ;T 7, % C 7$DR8 9;, 5 C 9//, 8 5 C 5D 7 > %, 5 I(

" + &G & " " "" , H & G

" " M?P'@ 3Saccharum officinarum9 6 > D > 7 ,

(43)

E

% 8 ;T 7, % C 5D>/58 >/ T, = C 7 7 8 7, > C 5/7 8;9JD% >, % C

7$DR8 9;, 5 C 5D7 > %, 5 C 7$%;7 5 H &(

Methylobacterium " 3Arachis hypogaea 9 6

" H & & * " >, 7 H T I, + F', E, @KP(@K?,

@PP?

% 8 ;T 7, % C 7$DR8 9;, 5 C JS 7, 5 S C 5 , % Methylobacterium

phyllosphaerae 7 + , I( , " + &G & &

&G & " 0 6 7 > % >

* " >@> L " , + FN, @@(@K, @PPN

% 8 ;T 7, % C 7$DR8 9;, 5C 5/7 8;9JD% >, % C 5D7 > %, 5 C 5 ,

7 ( H & G &G & " " "

* " 4 +, , + ?E, E( @P, @PPN

7/%/ / J(% >58 99, % C % " 7 9 , > C B> 7R/7, W W C

S W ;/ 8 S5J;, 9 C B 99, > % & + / 0G " "

( & & & ) " " "

+ +> >, > "I+ , + PM, 'FN( '?M, NNF

7D77, 7 C T, C 7 8 7T, & " & &G

Methylobacterium extorquens % + % 6 , 9 , + @?P, MFK(

M?@, NMN

%/>, R 5 , % 9;7;, > , $/>8 > 5 7, " 0 G I(

" + &G & " " 3 B% 6 D * * " > >, , + E?, ' NO'@?, @PPE

/7\9=/>, $ 7 C % >;7, C 7 / , B C 5 D>/9, C %;9 7, C = >8 9 , W Methylobacterium extorquens % " + G "G &

" H & "& &G & " H & "

D , , + FMP, F? (F?K, @PP?

;> ;9 , % C 9 DJJ 7/7, 8 C 5 ;/ 8, 8 C %T99T9 , > C 8 8 9 ,

" " " & " "& 3 5G + 9 6 G

&G 0 % * " >, , + ??,

'PK'('PKK, @PPP

B BB9, % S H & " + * " ( > (

> 1 4 +, 9 , + @N, EF, @PP

7$DR8 9;, 5 C % 8 ;T 7, % C 5 , " "G (& "

* ( H ( + Methylobacterium 6 7

(44)

E@

%;7;, % C >DR, 9 > C $ ;, C > XW , S 9 C % >5 ; 9;, W 9 ( & "G (8 9 " Methylobacterium mesophilicum:5G &

6 7 1 , &, + K@, @ @ 'P(

@ 'E, @PPN

> % J/>5, C >D;W />, W % C 9/J 77/ / >/R, > 8 C % >% 7, B %

( G * + ( > 1 ,

, + ''N, ?@(??, @PP'

5 7 8/5( 7 >/> 5, % C D/>, S C $ , % C > ;75 7, W C S7;/,

4 ( & 0 G " " H &

+ + ! 7 !+ 4 > > , 9 , + '?@,

EN( ?', @PPK

5 7 % 77, $ /+ " & " "

&H G + 7 + % > +> , 7 H T I, + EM', ?N( KE,

@PPN

5 8S > R, C S 9 />, 5 C % > /7, 5 % C J;>5 88 B/>, B C 7D55/>, % C

5 , D D * + ( > ( > G & )

* ( Pseudomonas aeruginosa. >

> + % >, 8 , + 'MK, F '(F@ , @PPK

5 ;/>, % 8 + "& "

; 9/, > J 3/ 6 % " +> > 9 " " ,

NNM M ( '?

5/=/>;, / C 8 5;/, 8 B C 8 SJ8/ , W C 8 % 5, $ 8 & " 0

+ " " & G 35556 G + +

" H & I H " " " D * * " > ,

, + 'PE, EK(FE, @P P

5/7 8;9JD% >, % C % 8 ;T 7, % C 5D7 > %, 5 C J 77 ;T 7, 5

; " " 0 H & " Methylobacterium H &

( H & " 3Oryza sativa9 + (E'6 * " 4 +,

, + ?E, N@( PE, @PPN

5 , % W C 5 7WD#7, W C 9;= >/5, W >& 0 ( & " "

" " H * " >, 7 H T I, + F@, ' ?KO' KE, @PP?

5T, C $;> D , / C W D> 7 , C $ > ; , 7 C S;99/%5, C / 9 WD ;/, C >;7, T C 7/T> , % C $;99;5, % C ;=;7(% 55 7, C >/TBD5, % &G & "

Methylobacterium " ) G H & 6

(45)

E'

5T, C ;%%/>5, W C J7;/B, C = >8 9 , W % &G & "

+ Methylobacterium extorquens " 0 Medicago

truncatula " + " % * " >,

, + K , K@EF(K@F@, @PPF

T % , 8 C 7 8 7T, C 9; 5 > %, % / "

mxa Methylobacterium extorquens % G " D *

* " > , , + ??, (K, NNM

= 7 J/7, C />/5, % C T, 5 9 C T %, W % C 5 87 >, W 9

Methylobacterium populi nov, + ", I( , " + G

&G & ", & ( 0 " 3Populus

deltoides)nigra 7'E6 0 6 7 > % >

* " >@ I , + FE, N ( N?, @PPE

S $7/>, = / C D587/99, C 55 >, 9 C > J5, ; C ;$9/S5J;, 8

% " G G Pseudomonas aeruginosa* ( "

H & & + 6 7 >, , + MF, @PMP(

@PNF, @PP'

S 7$, U C 9;D, S C 8/7, U C 7$, C 7$, T C 8D 7$, U C S/;, $ C 8 7, 4 C

8D 7$, 9 C J 7$, R ) " " H

H& + G " 7 ( B9 " "

" * >, 9 , + P @P P +

a& //HHH " " / EK (@@@N/ P/N` " @@ @P P

S8; /, / S C S;7 75, 5 (" " " & &

Agrobacterium tumefaciens + + ! + 4 > > ,

9 , + '?@, 'F( EM, @PPK

UD, R C ; 7, C 5D7, R C 9;7, W C 8D , T ; " " G

&G & ) G " Deinococcus

radiodurans * " >, > , + F', ?F@( ?F@, @PPK

R8 7$, % C 9; 5 > %, % / " + + &

(46)

EE

I 21 0$ 4 9S D01 0

- " ( ,

) " & " " + + - ,

2 "

" " ! " " "

+ , " + +

+ " " / "

, ! Methylobacterium, * !

" " " A " *

" " " "

"& * X. fastidiosa " , Methylobacterium

* 0 < " " " , + *

+ " ,

Methylobacterium " , : " +

A "

7 & + * Methylobacterium

& + " " + ( )

" " - " " , " 4

0 " 0 , + *

+ + : "

+ " " " +

( ) 0 B + " * & + "

* * & + +

4 " " , +

" " . " " &

" ( , " + & +

"& " " + +

; ) " ! Methylobacterium <

" , * +

(47)

EF

Methylobacterium ! " " "

" " , + & + 0

" , : "

0 " , " " & &

".( & +

B + * M. extorquens > ?/@ )

" 3/ 6 " 0 " 0 & * " (

" " X. fastidiosa. , + *

0 , )

+ & Methylobacterium " ,

+ " " "

" "

" " * Methylobacterium

" & , + * ) mxaB,

acd5, crt;, pat, sss phoD M. mesophilicum 5> ?/?

356(7( " (859 7 " ) , + * ,

" - + , 89 " +

mxaB, " " " "

C pat, * " " + + " +

" 0 C acd5, "

+ & , * "

+ " " + + " @, M.

mesophilicum 5> ?/? + " " " , , +

* " " < + acd5

& ) crt;,phoD sss 89

" - + , * " " +

" "

" +

-89 Methylobacterium

+ " , &. - " (

(48)

E?

+ - , " & +

" + + Methylobacterium( , "

" - . " " " , + * "

< " " ,

" + " Methylobacterium " "

" " + , " , ,

Referências

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