20
*Corresponding Author E-mail: [email protected] © 2002 Association of Applied Biologists Ann. appl. Biol. (2002), 140:109-127 Printed in Great Britain 109 Breeding for resistance to whitefly-transmitted geminiviruses By MOSHE LAPIDOT 1 * and MICHAEL FRIEDMANN 2 1 Department of Virology, Agricultural Research Organization, The Volcani Center, Bet Dagan 50250, Israel 2 Department of Plant Genetics, Agricultural Research Organization, The Volcani Center, Bet Dagan 50250, Israel (Accepted 11 March 2002; Received 27 December 2001) Summary Geminiviruses comprise a large and diverse family of viruses that infect a wide range of important monocotyledonous and dicotyledonous crop species and cause significant yield losses. The family Geminiviridae is divided into three genera, one of which is Begomovirus. Species of this genus are transmitted by the whitefly Bemisia tabaci in a persistent, circulative manner and infect dicotyledonous plants. Severe population outbreaks of B. tabaci are usually accompanied by a high incidence of begomoviruses. During the last two decades, there has been a worldwide spread of the B biotype of B. tabaci, accompanied by the emergence of whitefly-transmitted geminiviruses. Control measures in infected regions are based mainly on limitation of vector populations, using chemicals or physical barriers. However, under conditions of severe whitefly attack, none of these control measures has sufficed to prevent virus spread. Thus, the best way to reduce geminivirus damage is by breeding crops resistant or tolerant to the virus, either by classical breeding or by genetic engineering. A number of begomoviruses have been the subject of much investigation, due to their severe economic impact. This review considers the most severe viral diseases of four major crops (tomato, bean, cassava and cotton). The approaches taken to breed for resistance to these viral diseases should provide a perspective of the issues involved in breeding for begomovirus resistance in crop plants. Key words: Geminivirus, resistance, breeding, cassava, bean, tomato, cotton, Bemisia tabaci Introduction Geminiviruses are a large and diverse family of viruses that infect a wide range of important crop species, including both monocotyledonous and dicotyledonous plants, and cause significant yield loses. These viruses are characterised by the geminate morphology of the capsid and by circular single-stranded DNA (ssDNA) genomes. The family Geminiviridae is divided into three genera, based on genome structure, plant host and insect vector (Fauquet et al ., 2000). Species of the genus Mastrevirus ( Maize str eak virus as type member) are transmitted by leafhoppers (Homoptera: Cicadellidae), have a monopartite genome, and generally infect monocotyledonous plants. Species of Curtovirus (Beet curly top virus, BCTV, as type member) are also transmitted by leafhoppers, have a monopartite genome, but with a different genetic organisation from that of the mastreviruses, and infect dicotyledonous plants. Species of Begomovirus ( Bean golden mosaic virus, BGMV, as type member) are transmitted by the whitefly Bemisia tabaci (Gennadius) and infect dicotyledonous plants. Most begomoviruses of the Old World and all New World begomoviruses have bipartite genomes (A and B components); however, some Old World begomoviruses, including Tomato yellow leaf curl virus (TYLCV) have a monopartite genome. All begomoviruses are transmitted by their whitefly vector in a persistent, circulative manner. That is, once the vector feeds on an infected host plant and acquires the virus, transmission of the virus can then occur within hours, and may continue for the life span of the vector (Cohen & Harpaz, 1964). However, as shown for TYLCV, transmission efficiency declines with time (Cohen & Harpaz, 1964). Control measures in infected regions have traditionally emphasised vector control (Cohen & Antignus, 1994; Polston & Anderson, 1997; Hilje et al ., 2001; Palumbo et al ., 2001), mainly by pesticides or physical barriers. Chemical control methods have been only partially effective, since whitefly populations can reach very high numbers, leading to intensive pesticide use (sometimes twice daily) in attempts to eliminate the vector before it transmits the virus. Furthermore, there are concerns that the vector may develop pesticide resistance and the intense application of pesticides may have deleterious effects on the environment (Pico et al., 1996; Palumbo et al., 2001). Physical barriers such

breeding for resistance to whitefly-transmited geminivirus .pdf

Embed Size (px)

Citation preview

  • !

    !"

    #$$%&'#()$

    !"# $% &$"& '('&!')%""''""!"'$"" !"'$"" !"# #! ! %&!'$ $" *( &$"'# '"'(%+""&)(!( '*#! "&'( % ' '','()('&$)# ''+!!$'&!'!"'$"" #$' * #"#$'""',- "&* ) $$ !!"#,(%( !!"&,%"" *%'($ '')"! +'(( ,)"$) #"&'(,"'#"&*)+!!"#,'(%!"&)('&$.' ''% * "'"$ &!' %" , $" $''""&!'"#"#$'" + %!(!$ "#( !$, * ")+!"'" "& )('&$''!-+""& '( !"'"$ ( &&! '"#' #* ( + '(, ') '" !% % ,,%!"# ''"'"$''"'( +'(,!$ !$,%",%'!%%*,"&,%"" (,'( ,/!'"&!( '%'"+'"'( !""!#!'*( )!" '(" ' $ "&&"/"!"# '"'"+,+! !"''"*(##"!( '-'",&" '!'"'( $ ("$#"# #!'"&'( "$,%&",%"" '!!"##$' *

    + '!+,%+! +,+'"'"+!"''"+)

    $% &$"& '(' &!') %"& #"''!"# #! + !$% ,"'(""!"'$"" !"'$"" #$' + ! %&!'$$" * ( !(!' , '(%'"#("$"%"&'(!# ,!!$ %$. ' %" *(&$"'# '" '(%+ , "%" '!'+ #$'(" ' !' !'"0' *+ 1222* #! "& '( % %' % + ' '#, ' '' , $&("## ""#'3##+ ( ""#'' %"+ %$$&!'""!"'$"" #$' *#! "&''+4+ '#,$ "' '',$&("## +(""#''%"+,')'(&&'%'!"% '" &" '(' "& '( ' + &!' !"'$"" #$' * #! "&'#'"% '#, ' '' , '()('&$ ) &!'!"'$"" #$' *" ',%"" "&'(5$6"$7$$5)6"$7,%""

    ( ,#'' %" !"#"' 8(")+ " 5$6"$7 ,%"" +!$%, ' 94(""#''%"*$$,%"" ' '','()('&$!'"# ''+!!$'* (' +"!'(!'"& " &!' (" ' #$' !0 '( +' ""&'( !'("!!)'((" + !"' &" '( $& # "& '( !'""(:#;+?* ")+ (")&"94+' "&&!!!$ )'('"(:#;+?*

    "'"$ &!' %" (''"$$ #( !'" !"'"$ "(:

    '% +

  • &. ( ! ( , '(' '" #"'!' !"# "(:

    '% +

  • &"*+

  • !!"!#$$%!!&'$(!)*)+,*,(

    #(!(!!#$-(- !# ! ! !! !!"(.$(!!/$(%!-+!((!+#(!!+.!(%!%!0%(

    +1!(!(%!!+!

  • "

    &!'#$' 8 &"'")(!( ! &"'% '(&&!'"&94&!'""'($* '($ + !(#&!' ' ' !"$,!"'"'(" '#" %)' + '($%'(!" '$*

    "&'(/"", '!$ '($"#'"&94. '! '( '"&'( '!$$ #$,'(#$'*$'%''('"&&!'"+"!$'"# %")'(!"'" $$(/"&&!'"'( '"& #'" !,'( !"*+

  • #

    !(!$$+ %% '% '( "! "&' "+"(,'""&'$'"&'( '"#$'!$$ !( "& '!*+

  • $

    ''' * (')"/"%"# "&!"",%#$ +'($!'( %"# + )(!( ##'$ )" '!' #'$+ &"&&')$,#"#$'" *!(%"# '"'(! *" '($!%"#+%"+ H$ !" "!! !(!' , $$ ,$!-+ $$ + #-+)('#'"'# *"'(%"#+'(($+! !(!' ,$% -,+"''$ #,'# *, "&'(%"#!,'!" +'%'!, C ' "! +! ''%'( "&,%%$ "$ :%(+2F "& '( #("'#! '" &" '!*

    '"$+$'("%( ''!$' ( , $"# &" %$ "! "& '!+ '( "' 0' ("!$# *!' ' ( (")'(', #% '! % &"&&'! !*'+(%($ ''$ ,$"#A*+84$;*+.>+)(" '!) &"'(#" $$"#$?1=+)(!( (") '! '" !($"" '" '( %$ ! %)*-) +$=1?G.=?+)(" '!) &"I2I+) (")'"!'(! % !"'"$$% '! '" !($"" +)14$;*+

  • %

    (&"+'( ( !',%"'(,B $'""&'($$"& '!'(' ##'%&$* " ! ) #'("%("',.(&!+"'(- '(" '#"''&!'"$'%! $ $"!'" 0':%''+

  • &

    ) %'$!"$')'('," ""',)%('*

    % &",%"" '! ! 0 '( ' '""&"!$'"#"'"!"$ '(')$$ '(#("""& "+ " '" #"#$ '( '! $$ "& '('$ ,% !* ( $"#' "&"$!$- $-'" '!% )"$,C'$$,$+ ')"$$$") !%"& #'"$ #"#%'"'$'" !''!" &" '' "!*

    ( ')"" ' #"')('&$.' '' "&!"''"!"''"$&!#$ +!"''"$&!$ *!(%(!! "&,"'( "!')'((%()('&$#"#$'" + ,%"" ( , /"!' '(! $%' "&'( *+ ' 4+ ,#'',%"" *+

  • '

    '(&" , ,$ '" $"# #"'"!"$ "&'&!$"!$'" %)('&$ !%$$'(!"#"' "&4*( ' ("$," '"% "!$'" # !" ''"'$ !"'" *

    !" "##

    ')"$ %'$ &!$'' '( $"#' "& ''%#$ &"$!$- +, "#"$"#( "& 0! + $- '" '!% )'&'"'( $!'"#"! % '( ,% &&"' * 6'("'"$!$- +,%&"$ '! $''"#("'#! !"&'( ''#$' +**+ $!'% #$' "$$ " '( , "& '(# !" , !"& #'" "" #'" '* !'(,%"" ' '',)('&$ + '( "!$'" !% $''"'() " "&'(*""+'( "&&$"!$'"+"&(%(!''"&$&" )('&$ + &&!'!, $%4 -*++ ,' '" $"! &&'&"23$(%*+

  • !

    '"48 "*+1222*' $-$ '(' !",%% &"&&'

    "! "& '!!$'" #"$$ "& '!'",%"" *" '!+$") -*

  • (,' &")'('(94!%$ (") $ #'"$"#' !" &" $ &!'")'( '* ( '!) & '"$,C# ""&'(' %!#"'+ !' %!#$' C# %'(! % "$ ' '( $$ (") " '! '" $ &!'" L- *+

  • )''7 8

    (& '" ''""&&"% ) '(C# ""&&!''&%

    "&4 ' %!6*) #$' '$+

  • %'$ #"'(# !"&""4*"$ '":,' 122

  • "

    &!''(!"#)$$ "!"& '!% * '"+ '!% '"% &")$ #! !"#$''(" ' %!#$' )'('('#" "$#'$ '!*""+ $ 0! ( , ,%"" ,'$ &"'($"#'"&)'(). #!'"& '!*

    '#

    "','" &" '(%!$'$ !(%;'"+'(4"$!'+'%+ $*,GI?P2

  • #

    61 0 !". ,. B):9.:)9 . "! #

    02!-@? & '

    6+8 +@%!!#

    ;-+):9""&" "

    '$75/';2!!?;-

    !" !# *(!?"'#'

    1866!!?))99). "! #%

    186 %345!--59)9:99:C)56:?!#!!$

    @' >,% +8=; +*!!?:56 !.*3--%& %&$

    75/';7) +@%

    $A!!?,)9):8*&($"'

    **)*+

  • $

    $' '

    6 @ !!"+ $ :9 < > )$! !$

    6 % B9'&% ;?:99:9; ? )+ $!&"'

    !!.(!.

    =' >=!!; *)$*)# $% $!

    =@!!#*9568 : F -!#% #%$

    =82 5% 45 3 4

    +0'186938@9$5*)8@ ;7=+ :)9 A!$$"*-$"$)

    8*,9 3@

    4!!" 3G;=:!" "## "$

    $ :

    $'$'"#

    $ *= ; )9=

    ."# $##

    $ 0 !-- 99):? "

    $*!!A: =# " &

    $*!!:="?$!%"

    ,3/;9). ? ))? ""#

    5;4%!!? ) 9 .* EE))):8EE !!&E% E

    8=() * !!- 9 9)("$%$$'"1.@//1/C)3,,3,

    )+8,28'&%22@ )9)'

    "

    "!"#!. %# %&

    3/+1/51!!!;9)(:3-!"

    3*+8@,445186!!; 9) )9) -# " %

    ,++'*2311%

    2311 !!"@ 9) $ ) . " "'

    @7+;'&%!!.;)56?"!$!"

    75/';2 5);!!-;

    *# $#"

    *5 )@ $A !!! ,:):9B:"# '$

    *5=,%@&+8,* !!-; :9)# '""#

    *5&6)1%*&8(

    $&"!&%%

  • %

    *+ 11$!!"+ ((;9) H )!-"? $ !%

    *+ 11$ !!- )9 B *)9)$!&

    *+ $11$!!!5:9)

    -$"&$#

    *)8@!!;9) *

    ?.#'$

    *)8@ 3 ))9:9)

    "$ "$"

    *)8@$556!!?;7 &)9)("4.(.$

    *$=8%$AA:9 $ $

    *$975/';7) $A @:. :989 *&%-%$$%%$

    *3B1+%0 $#!$

    +;3% 625%*, + !!" ( )9 * ((," ""45667""$&"%&5A.1.*/(

    *$26@: 9 :A-@ A-@$!. '"%

    *$8*263: :!. $%$ $&

    *261'285 @9:' '#!. '"% '"&

    C6*,&16&,!!!;:):)-!& &%

    @!!?5(56) (,

    "" )#'&!$

    @66!"++(

    &!"#"!

    )6 !!.,.:(!. $$

    25*,8'0!!-;9:)

    99-!&%#&%$

    275/';!!.3

    $"%

    275/';$!!-;9EI("%&%&&1.@//1/C)3,

    ,3,

    2 75/'; $ !!- ;9'!#

    275/';28

    @

    9!!-;( :9)9: ) '

    ??!"$!#$

    B8 * 3 %@6 !!"99568 : :&

  • &

    !-! "$ "!93%8*@0( ' =-#"

    91 3!!-(::& ! - !%'%"

    938@$186!!-;: 9) *--! ! #

    938@$33%&)+6=

    186 !!-@ : .* . 9 :9 9)? %$ %!

    35@!!(:9:9-?$$$$%#

    ;8@'!-#6)78D9("%#

    ;$ )=!!( &--$'$!

    48@34=%&)+6

    186 45 !! 3 * 9) 56 9

    ?#&!'

    45,8/+048@34

  • '