(XU-(QWRPRO±
,661
7KHUROHRISODQWRGRXUVLQWKHOHDIPLQHU/LULRP\]DVDWLYDH'LSWHUD
$JURP\]LGDHDQGLWVSDUDVLWRLG'LJO\SKXVLVDHD+\PHQRSWHUD(XORSKLGDH
2ULHQWDWLRQWRZDUGVWKHKRVWKDELWDW
<81;,$1=+$2DQG/(.$1*
6WDWH.H\/DERUDWRU\RI,QWHJUDWHG0DQDJHPHQWRI3HVW,QVHFWV5RGHQWV,QVWLWXWHRI=RRORJ\&KLQHVH$FDGHP\RI6FLHQFHV
=KRQJJXDQFXQ5RDG+DLGLDQ%HLMLQJ&KLQDHPDLO]KDR\[#SDQGDLR]DFFQONDQJ#SDQGDLR]DFFQ
.H\ZRUGV($*OHDIPLQHU/LULRP\]DVDWLYDHSDUDVLWRLG'LJO\SKXVLVDHDSODQWRGRXUV
$EVWUDFW (OHFWURDQWHQQRJUDP UHVSRQVHV RI WKH SRO\SKDJRXV OHDIPLQHU /LULRP\]D VDWLYDH DQG LWV JHQHUDOLVW SDUDVLWRLG 'LJO\SKXV
LVDHD WR KRVW DQG QRQKRVW SODQW RGRXUV RI / VDWLYDH ZHUH LQYHVWLJDWHG 7KH RGRXUV RI KHDOWK\ OHDYHV FDQ HOLFLW GLVWLQFW ($*
UHVSRQVHVLQ/VDWLYDH7KH($*UHVSRQVHVWRWKHRGRXUVRIWKHKRVWSODQWVEHDQDQGWRPDWRZHUHVWURQJHUWKDQWRQRQKRVWSODQWV
&KLQHVHURVHDQGPRUQLQJJORU\1HLWKHUKHDOWK\KRVWQRUQRQKRVWSODQWVRIWKHOHDIPLQHUHOLFLWHGGLVWLQFWLYH($*UHVSRQVHVLQWKH
SDUDVLWRLG'LVDHD2GRXUVRISK\VLFDOO\GDPDJHGOHDYHVQRPDWWHUZKHWKHURIKRVWRUQRQKRVWSODQWVLQFUHDVHGVWURQJO\WKH($*
UHVSRQVHVRIWKHOHDIPLQHUDQGLWVSDUDVLWRLG:HFRPSDUHGWKH($*UHVSRQVHVRI'LVDHDWREHDQOHDYHVRIGLIIHUHQWVWDWXV7KH
RGRXURIPLQHGOHDYHVHOLFLWHGGLVWLQFW($*UHVSRQVHVZKLFKZHUHZHDNHUWKDQWKRVHRISK\VLFDOO\GDPDJHGRQHV1RGLIIHUHQFHV
ZHUHGHWHFWHGLQWKH($*UHVSRQVHWROHDYHVZLWKHPSW\PLQHVKHDOWK\OHDYHVRUDQDLUFRQWURO7KHUROHRISODQWRGRXUVLQKRVW
ORFDWLRQRIWKHOHDIPLQHUDQGLWVSDUDVLWRLGLVGLVFXVVHG
,1752'8&7,21
7KHOHDIPLQHU/LULRP\]DVDWLYDHLVDSRO\SKDJRXVKHU
ELYRUH ZKLFK DWWDFN D YDULHW\ RI YHJHWDEOH DQG RUQD
PHQWDOFURSV3DUUHOOD.DQJ7KHOHDIPLQHU
ODUYDH DUHREOLJDWRU\ LQWHUQDOSODQW IHHGHUV7KH\ IHHGLQ
OHDI PHVRSK\OO WLVVXH DQG IRUP D VHUSHQWLQH PLQH
&KHPLFDOFRQWURORIDJURP\]LGOHDIPLQHUVKDVUHVXOWHGLQ
WKH GHYHORSPHQW RI LQVHFWLFLGH UHVLVWDQFH LQ /LULRP\]D
SRSXODWLRQ DQG WKH HOLPLQDWLRQ RI WKHLU QDWXUDO HQHPLHV
OHDGLQJWRDQLQFUHDVLQJFRQWUROSUREOHP7KHHFWRSDUDVL
WRLG'LJO\SKXVLVDHD:DONHULVXVHGLQELRORJLFDOFRQWURO
RIDJURP\]LGOHDIPLQHUVLQJUHHQKRXVHV9DQ/HQWHUHQ
:RHWV )HPDOH SDUDVLWRLGV RYLSRVLW LQ WKH ODVW
ODUYDOVWDJHVRIWKHOHDIPLQHUV0LQNHQEHUJ9DQ/HQ
WHUHQ 7KH\ XVH VHYHUDO VSHFLHV RI /LULRP\]D
LQFOXGLQJ / VDWLYDH DV KRVWV $OWKRXJK ' LVDHD FDQ
SDUDVLWH GLIIHUHQW OHDIPLQHUV RQ GLIIHUHQW FURSV WKH
XQHYHQ GLVWULEXWLRQ RI SDUDVLWRLGV DPRQJ FURSV KDV EHHQ
LQYHVWLJDWHG LQ WKH ILHOG =HKQGHU 7UXPEOH -RKQVRQ +DUD 6FKXVWHU HW DO 6DOYR 9DOODGDUHV ,W KDV EHHQ VXJJHVWHG WKDW SDUDVLWRLGV
IDYRXUDUDQJHRILQVHFWKRVWVUHODWHGUDWKHUWRDSDUWLFXODU
SODQW RU KDELWDW WKDQ WR SK\ORJHQHWLFDOO\ UHODWHG LQVHFW
KRVWV $VNHZ 6KDZ ,Q WKH WULWURSKLF LQWHUDF
WLRQV LQYROYLQJ SODQWV OHDIPLQHUV DQG ' LVDHD ERWK WKH
KRVWDQGLWVSDUDVLWRLGDUHJHQHUDOLVWV,WLVFULWLFDOWRNQRZ
ZKLFKEHVWKRVWSODQWPDWFKHVWKHQDWXUDOHQHP\XVHGLQ
ELRORJLFDOFRQWURORIWKHOHDIPLQHUV
3ODQW YRODWLOHV DUH HPSOR\HG E\ ERWK KHUELYRUHV DQG
WKHLUQDWXUDOHQHPLHVLQVHDUFKLQJIRUERWKIRRGDQGRYL
SRVLWLRQ SODFH 'H 0RUDHV HW DO 6HYHUDO VWXGLHV
KDYH VKRZQ WKDW LQVHFW KHUELYRUHV DUH DWWUDFWHG E\ WKH
RGRXUVRIWKHKRVWSODQWV&RVVHHWDO/LJKWHWDO
*XHULQ 9LVVHU 3ODQWV UHOHDVH KHUELYRUH
LQGXFHG YRODWLOHV +,9V DV D UHVXOW RI IHHGLQJ E\ KHUEL
YRUHV ZKLFK DUH XVHG E\ WKH QDWXUDO HQHPLHV WR ORFDWH
KRVWV 'H 0RUDHV HW DO 7XUOLQJV HW DO 7XUOLQJVHWDO$SUHYLRXVVWXG\KDVGHPRQVWUDWHG
WKDWIHPDOH'LVDHDZHUHVWURQJO\DWWUDFWHGWRWKHRGRXUV
DULVLQJ IURP WKH LQIHVWHG EHDQ SODQWV )LQLGRUL/RJOL HW
DO 2WKHUV KDYH QRWHG WKDW WKH UHOHDVH RI SODQW
FRPSRXQGV LV KLJKO\ YDULDEOH DFURVV SODQW VSHFLHV DQG
YDULHWLHVDQG LV DOVRVHQVLWLYH WR WKHVSHFLHV RI KHUELYRUH
'LFNHHWDO7XUOLQJVHWDO7DNDED\
DVKL HW DO 'H 0RUDHV HW DO 'H 0RUDHV HW
DO GHPRQVWUDWHG WKDW SODQW HPLVVLRQV FRXOG
WUDQVPLW KHUELYRUH VSHFLHVVSHFLILF LQIRUPDWLRQ WKDW LV
GHWHFWDEOH E\ SDUDVLWRLGV %RWK WREDFFR DQG FRWWRQ SUR
GXFHG GLVWLQFW EOHQGV RI YRODWLOHV LQ UHVSRQVH WR GDPDJH
E\WZRFORVHO\UHODWHGKHUELYRUHVSHFLHV+HOLRWKLVYLUHV
FHQV )DEULFLXV DQG +HOLFRYHUSD ]HD %RGGLHV 7KH VSH
FLDOLVW SDUDVLWLF ZDVS &DUGLRFKLOHV QLJULFHSV H[SORLWV
WKHVHGLIIHUHQFHVWRGLVWLQJXLVKLQIHVWDWLRQVE\LWVKRVW+
YLUHVFHQV IURP WKRVH E\ QRQKRVWV $OWKRXJK WKHUH DUH
QXPHURXVVWXGLHVRQWKHEHKDYLRXUDOUHVSRQVHVRISDUDVL
WRLGV WR SODQW YRODWLOHV RQO\ D IHZ KDYH LQYHVWLJDWHG WKH
SK\VLRORJLFDOUHVSRQVHVWRWKHVHFKHPLFDOFXHV
,Q WKLV SDSHU ZH UHSRUW WKH ROIDFWRU\ UHVSRQVHV RI WKH
OHDIPLQHU / VDWLYDH DQG LWV SDUDVLWRLG ' LVDHD WR WKH
RGRXUVRIGLIIHUHQWSODQWVXVLQJHOHFWURDQWHQQRJUDPV:H
LQYHVWLJDWH WKH UROH RI SODQW RGRXUV LQ WKHLU ORQJUDQJH
KRVWKDELWDWSUHIHUHQFHV
&RUUHVSRQGLQJDXWKRU
7$%/(3ODQWVXVHGLQ($*WHVW
)DPLO\
6FLHQWLILFQDPH
/HJXPLQRVDH
3KDVHROXVYXOJDULV/
6RODQDFHDH
/\FRSHUVLFRQHVFXOHQWXP0LOO
&XFXUELWDFHDH
&XFXUELWDSHSR/
5RVDFHDH
5RVHFKLQHQVLV
&RQYROYXODFHDH
3KDUELWLVSXUSXUHD/9RLJW
&RPPRQ
QDPH
&RPPRQ
EHDQ
7RPDWR
6TXDVK
&KLQHVHURVH
0RUQLQJ
JORU\
0$7(5,$/6$1'0(7+2'6
3ODQWV $OO SODQWV 7DEOH H[FHSW EHDQV ZHUH JURZQ LQ DQ
H[SHULPHQWDOILHOGDWWKH,QVWLWXWHRI=RRORJ\&KLQHVH$FDGHP\
RI6FLHQFHV%HDQV3KDVHROXVYXOJDULV/ZHUHSODQWHGLQSRWV
LQWKHODERUDWRU\3ODQWVDPSOHVZHUHWDNHQIURPWKHOHDISHWLROH
VR DV WR PLQLPL]H WKH QXPEHU RI FXW VXUIDFHV WKURXJK ZKLFK
YRODWLOHV IURPZLWKLQ WKH WLVVXHVPLJKW HVFDSH &DUWULGJHVZHUH
ORDGHGZLWKDIUHVKVDPSOHRIHDFKSODQWHYHU\PRUQLQJOHIWDW
URRP WHPSHUDWXUH DQG XVHG WKURXJKRXW WKH GD\ :KHQ SK\VL
FDOO\ GDPDJHG OHDYHV ZHUH UHTXLUHG WKH\ ZHUH FXW E\ D NQLIH
EHIRUHWKHH[SHULPHQWV
/HDIPLQHU / VDWLYDH ZDV UHDUHG DV GHVFULEHG E\ =KDR .DQJ 7KH\ ZHUH UHDUHG RQ EHDQV 3KDVHROXV YXOJDULV
/ DW “ ƒ& ZLWK D SKRWRSHULRG RI / ' DQG UHODWLYH
KXPLGLW\2QO\XQPDWHGIHPDOHVZHUHXVHGLQWKHH[SHUL
PHQWV
3DUDVLWRLG ' LVDHD ZHUH FROOHFWHG DV SXSDH IURP WKH ILHOG
7KH\ZHUHVHSDUDWHGIURPWKHPLQHVDQGPDLQWDLQHGLQDFOLPDWH
FKDPEHUDW“ƒ&ZLWKDSKRWRSHULRGRI/'DQG
UHODWLYH KXPLGLW\ 2Q WKH GD\ RI HPHUJHQFH LQGLYLGXDO DGXOWV
ZHUHLVRODWHGLQWXEHVZLWKKRQH\ZDWHUDVDIRRGVRXUFH
$JDLQRQO\XQPDWHGIHPDOHVZHUHXVHGLQWKHH[SHULPHQWV
(OHFWURDQWHQQRJUDP($*
$QWHQQDSUHSDUDWLRQ$QWHQQDHRI/VDWLYDHZHUHSUHSDUHG
XVLQJWKHPHWKRGGHVFULEHGE\=KDR.DQJD7KH
WHVWLQVHFW¶VKHDGZDVUHPRYHGDQGWKHWLSRIWKHDULVWDZDVFXW
RII ($*V ZHUH UHFRUGHG XVLQJ HOHFWURO\WHILOOHG 5LQJHU¶V
JODVVFDSLOODU\$J$J&OHOHFWURGHVWKHUHIHUHQFHHOHFWURGHZDV
SRVLWLRQHG LQWR WKH KDHPRFRHO RI WKH FUDQLDO FDYLW\ ZKLOH WKH
UHFRUGLQJHOHFWURGHZDVFRQQHFWHGWRWKHFXWDULVWD,QSUHSDULQJ
WKH DQWHQQD RI WKH SDUDVLWRLG WKH KHDG RI WKH LQVHFW ZDV
UHPRYHGDQGWKHWLSRI WKHDQWHQQDZDVFXWRII7KHUHIHUHQFH
HOHFWURGH ZDV SLHUFHG LQ WKH KDHPRFRHO RI WKH FUDQLDO FDYLW\
ZKLOH WKH UHFRUGLQJ HOHFWURGH ZDV FRQQHFWHG ZLWK WKH FXW
DQWHQQD 7KH YHQWUDO VXUIDFH RI WKH DQWHQQD ZDV RULHQWDWHG WR
IDFH WKH RGRXU VWUHDP 2QO\ WKH OHIW DQWHQQD IURP HDFK LQVHFW
ZDVXVHG
2GRXU GHOLYHU\ V\VWHP 7KH RGRXU VRXUFH FDUWULGJHV FRQ
VLVWHG RI JODVV WXEHV PP ORQJ DQG PP GLDPHWHU FRQ
WDLQLQJ J IUHVK ZHLJKW RI SODQW PDWHULDO 7KH RGRXU VWLPXOL
LQFOXGH KHDOWK\ LQWDFW OHDYHV SK\VLFDOO\ GDPDJHG
OHDYHVPLQHGEHDQOHDYHVLQIHVWHGZLWKQGLQVWDU/VDWLYDH
ODUYDHIHHGLQJDQGEHDQOHDYHVZLWKHPSW\PLQHV7KHFDU
WULGJHV ZHUH VHDOHG DW HDFK HQG ZLWK D UXEEHU VWRSSHU ZUDSSHG
LQ D SLHFH RI DOXPLQLXP IRLO $ VKRUW OHQJWK RI PP GLDPHWHU
JODVVWXEHZDVSDVVHGWKURXJKHDFKHQGRIWKHUXEEHUVWRSSHUWR
DOORZDLUWRPRYHWKURXJKWKHFDUWULGJH
$ FRQVWDQW IORZ OLWUHPLQ RI FKDUFRDOILOWHUHG DQG
KXPLGLILHGFRPSUHVVHGDLUZDVSDVVHGRYHUWKHDQWHQQDWKURXJK
DGLVSRVDEOHQR]]OHSRVLWLRQHGDERXWFPIURPWKHDQWHQQD7HVW
VXEVWDQFHV ZHUH LQWURGXFHG LQWR WKH DLU VWUHDP E\ LQVHUWLQJ WKH
WXELQJ RI WKH FDUWULGJH LQWR WKH KROH RI WKH GHOLYHU\ WXEH DQG
LQMHFWLQJ WKH FDUWULGJH FRQWHQWV TXLFNO\ DQG VPRRWKO\ 7KH
VWLPXODWLRQ GXUDWLRQ ZDV VHF (DFK VWLPXOXV ZDV IROORZHG
E\DQLQWHUYDORIDERXWVHFRQGVRIFOHDQDLU7KHVLJQDOZDV
DPSOLILHG î E\ D 8QLYHUVDO $&'& DPSOLILHU 81 DQG
YLHZHG RQ WKH FRPSXWHU VFUHHQ ($*V ZHUH UHFRUGHG ZLWK D
6\QWHFK($*UHFRUGLQJV\VWHPIRU:LQGRZV9HUVLRQ
([SHULPHQWDO SURFHGXUH )RU HDFK VWLPXOXV ($*V ZHUH
UHFRUGHG IURP DW OHDVW WHQ DQWHQQDH $IWHU HYHU\ WKLUG WR ILIWK
WHVWD³FRQWURO´VWLPXODWLRQDQHPSW\FDUWULGJHDQG³VWDQGDUG´
VWLPXODWLRQXVLQJIUHVKFDUWULGJHVLPSUHJQDWHGZLWK—ORI
KH[DQROZHUHSUHVHQWHG($*VRIWHVWSODQWVZHUHHYDOXDWHG
E\ PHDVXULQJ WKH PD[LPXP DPSOLWXGH RI QHJDWLYH GHIOHFWLRQ
±P9 HOLFLWHG E\ D JLYHQ VWLPXOXV DQG WKHQ VXEWUDFWLQJ WKH
DPSOLWXGHRIWKHUHVSRQVHRIWKHSUHFHGLQJFRQWURO7KHPLOOLYROW
UHVSRQVHV WR DOO SODQWV ZHUH FRQYHUWHG WR SHUFHQWDJH YDOXHV RI
WKHUHVSRQVHWRWKHDFFRPSDQ\LQJVWDQGDUG($*GDWDZHUHVXE
MHFWHG WR $129$ DQG PHDQV ZHUH FRPSDUHG E\ 7XNH\¶V WHVW
XVLQJ6366
5(68/76
($*UHVSRQVHVRI/VDWLYDHWRGLIIHUHQWSODQWVRGRXUV
0HDQUHVSRQVHVRIIHPDOH /VDWLYDHWRWKHKH[DQRO
VWDQGDUG —O ZHUH ±“ 6( P9 7KHUH
ZHUHGLIIHUHQFHVLQWKHPDJQLWXGHRIWKH($*UHVSRQVHV
RI/VDWLYDHWR WKHRGRXUVRI GLIIHUHQWSODQWV3 )LJ $PRQJ WKH ($* UHVSRQVHV WR KHDOWK\ OHDYHV /
VDWLYDH ZHUH VLJQLILFDQWO\ PRUH UHVSRQVLYH WR EHDQ DQG
WRPDWR 7KH ($* UHVSRQVH WR VTXDVK ZDV OHVV SUR
QRXQFHG 7KH ORZHVW PDJQLWXGH RI UHVSRQVHV ZHUH
REVHUYHGIRU&KLQHVHURVHDQGPRUQLQJJORU\$OORIWKH
KHDOWK\ SODQWV HOLFLWHG JUHDWHU ($* UHVSRQVHV WR /
VDWLYDH WKDQ GLG FRQWUROV 3 0XFK KLJKHU ($*
UHVSRQVHVZHUHGHWHFWHGWRSK\VLFDOO\GDPDJHGOHDYHVIRU
DOOWKHWHVWSODQWV7KH($*UHVSRQVHVWRSK\VLFDOO\GDP
DJHGOHDYHV ZDV VLPLODUIRU PRUQLQJ JORU\&KLQHVH URVH
DQGVTXDVK'DPDJHGOHDYHVRIWRPDWRHOLFLWHGDUHVSRQVH
RIKLJKHUPDJQLWXGH
($*UHVSRQVHVRI'LVDHDWRGLIIHUHQWSODQWVRGRXUV
7KHPHDQUHVSRQVHRIIHPDOH'LVDHDWRWKHKH[DQRO
VWDQGDUG —O ZDV >± “ 6( P9@
$OWKRXJK WKH PDJQLWXGH RI WKH ($* UHVSRQVHV WR WKH
KHDOWK\OHDYHVRIWRPDWRZDVKLJKHUWKDQWRRWKHUKHDOWK\
OHDYHVWKH\ZHUHQRKLJKHUWKDQWKHFRQWUROV)LJ3
+RZHYHU PHFKDQLFDOO\ GDPDJHG OHDYHV HOLFLWHG
KLJK ($* UHVSRQVHV IURP WKH SDUDVLWRLGV )LJ ZLWK
GDPDJHG EHDQ OHDYHV HOLFLWLQJ UHVSRQVHV RI WKH KLJKHVW
PDJQLWXGH 7KH ($* UHVSRQVHV WR GDPDJHG WRPDWR DQG
&KLQHVH URVH ZHUH KLJKHU WKDQ WKRVH WR WKH GDPDJHG
VTXDVK3
($*UHVSRQVHVRI'LVDHDWREHDQOHDYHVLQGLIIHUHQW
VWDWXV
)LJVKRZVWKH($*UHVSRQVHVRIIHPDOH'LVDHDWR
EHDQOHDYHVRIGLIIHUHQWVWDWXV7KH($*UHVSRQVHVRIWKH
SDUDVLWRLG WR WKH KHDOWK\ OHDYHV DQG OHDYHV ZLWK HPSW\
PLQHVVKRZHGQRVLJQLILFDQWGLIIHUHQFHWRWKHDLUFRQWURO
,QIHVWHG OHDYHV ZLWK QGLQVWDU / VDWLYDH ODUYDH IHHGLQJ
ZLWKLQWKHPHOLFLWHGGHWHFWDEOHUHVSRQVHVWR'LVDHDEXW
Fig. 1. EAG responses of L. sativae to the odours of different plants (n > 10). The vertical line indicates standard deviation. Letters within the brackets represent significant differences of EAG responses between healthy plants and controls. Letters without
brackets represent significant difference of EAG responses to physically damaged plants. Means with the same letter on the bar are
not significantly different (P < 0.05).
the magnitude was less than that elicited by the mechanically damaged leaves (P < 0.01).
DISCUSSION
This EAG analysis of the response of L. sativae to different plants demonstrates that the antennae of the leafminers respond selectively to host plant odours. They are
more sensitive to host plants, such as bean and tomato,
than they are to non-host plants such as Chinese rose and
morning glory. Behavioural studies in the olfactometer
produced analogous results (Zhao & Kang, 2002b).
Odours of host plants, such as bean and tomato, had
similar levels of attraction to the leafminer, and were
much more attractive than were the non-host plants. All
the healthy plants, no matter whether host plants or nonhost plants, elicited higher magnitudes of response than
did controls (Fig. 1). The green leaf volatiles (GLVs),
which generally exist in green plants, may attract the
polyphagous leafminer. When the plants were damaged
mechanically, greater amount of plant volatiles, especially
the green leaf volatiles, were released (Visser, 1979),
which may contribute to the high EAG responses of L.
sativae (Fig. 1). Previous studies have shown that L.
sativae were more sensitive to green leaf volatiles and
other characteristic volatiles of the host plants (Zhao &
Kang, 2002a). We suggest that the leafminer, L. sativae,
employ both the GLVs and other characteristic volatiles
of host plants in orientating to their host habitats. Behav-
Fig. 2. EAG responses of female D. isaea to the odours of different plants (n = 10). The vertical line indicates standard deviation.
Letters within the brackets represent significant difference of EAG responses between healthy plants and controls. Letters without
brackets represent significant difference of EAG responses to physically damaged plants. Means with the same letter on the bar are
not significantly different (P < 0.05).
447
Fig. 3. EAG responses of D. isaea to the odours of bean in different status (n = 10). The vertical line indicates standard deviation.
Means with the same letter on the bar are not significantly different ( P < 0.01).
ioural assays are needed to determine the effect of the
GLVs and the characteristic volatiles of host plants on the
leafminer.
In locating their hosts, parasitoids must search for
potential hosts living on different plants growing in
diverse habitats. Chemical stimuli act as the cues which
direct many parasitoids to their hosts (Dicke & Minkenberg, 1991). Some parasitoids use volatiles emitted by
undamaged plants to locate the habitat and, possibly, the
microhabitat of their host (Elzen et al., 1984; Ngi-Song et
al., 1996). Two parasitoids Cotesia flavipes and Cotesia
sesamiae were strongly attracted to odours emanating
from several gramineous plants (maize, sorghum, and a
wild host, napier grass), which are host plants of stemborers. Both of them had showed preferences for the host
plants of their prey. C. flavipes showed a preference for
maize over sorghum, while C. sesamiae was more
attracted to sorghum odours (Ngi-Song et al., 1996). In
the EAG analysis of D. isaea, none of the EAG responses
to the odours of healthy plants differed from the controls,
regardless of whether they were hosts or non-hosts of the
leafminer (Fig. 2). The odours of mechanically damaged
leaves elicited higher EAG responses than did intact
leaves, which may be due to the higher quantities of the
“green leaf volatiles” (GLVs) released by damaged leaves
(Visser, 1979). No preferences of D. isaea to particular
plants were detected in the EAG responses to the odours
of healthy and physically damaged leaves. An early bioassay of D. isaea showed that the parasitoids were
slightly attracted by the bean odour (Finidori-Logli et al.,
1996). Larval and adult conditioning of D. isaea affected
slightly the females’ selection of plant type on which the
host larvae were located (Coaker & Cheah, 1993). In this
study, D. isaea pupae were removed from the plants
before their emergence. The adults had thus no experience with the plants, and no distinct EAG responses to
odours of beans were detected. Whether they would have
448
more sensitive EAG responses to host plants after conditioning remains to be investigated.
Physically damaged bean leaves elicited the most distinctive EAG responses in D. isaea, with mined leaves
second. The odours of the mined leaves was more attractive behaviourally to the parasitoid than were the
mechanically damaged leaves (Findori-Logli et al., 1996).
The greater amount of GLVs in the mechanically damaged bean leaves may have little attraction to D. isaea,
whereas the particular volatiles released by the mined
leaves were attractive to the parasitoid. Plants may be
induced to emit attractants utilized by insect parasitoids
by the host saliva (Turlings et al., 1990). The volatile
blends that are released in response to herbivore feeding
are not identical to those elicited by artificial damage
done by mechanical means (Dick & van Loon, 2000).
Two components, cis-3-hexen-1-ol and 4-hydroxy-4methyl-2-pentanone, were present in significantly larger
quantities in the leaf extracts from mined or damaged
bean plants than in those from healthy ones (FindoriLogli et al., 1996). The compound 1-Octen-3-ol was
emitted only by the mined leaves (Findori-Logli et al.,
1996). It is understood that the chemosensory system of
arthropods is able to detect volatile compounds at concentrations that are so low that they escape attention in gaschromatographic mass spectrometric analyses (Pickett,
1990; Pickett et al., 1998). 1-Octen-3-ol may be only one
of many herbivore-induced volatile (HIV) components.
Whether 1-Octen-3-ol itself or the whole HIV blend has
the attractive function needs further research. The EAG
response of D. isaea to the odour of leaves with empty
mines is similar to the healthy ones. This may be due to
the cessation of HIV released when the herbivore stopped
feeding. Although some parasitoids use volatiles emitted
by undamaged plants to locate the habitat and possibly
the microhabitat of their host, the HIVs were more important in guiding D. isaea to their hosts. We suggest that for
VXFFHVVIXO HVWDEOLVKPHQW WKH SDUDVLWRLG VKRXOG EH
UHOHDVHGDIWHUWKHLQIHVWDWLRQRIWKHSODQWE\WKHOHDIPLQHU
7KH UHOHDVH RI SODQW FRPSRXQGV LV KLJKO\ YDULDEOH
DFURVVSODQWVSHFLHVDQGYDULHWLHVDQGLVDOVRVHQVLWLYHWR
WKHVSHFLHVRIWKHKHUELYRUHLQYROYHG'LFNHHWDO
7XUOLQJV HW DO 7DNDED\DVKL HW DO 5|VHHWDO'H0RUDHVHWDO,WLVWKHKHUEL
YRUHVWKDWSURYLGHWKHSDUDVLWRLGVZLWKWKHGHWHFWDEOHDQG
UHOLDEOHFXHVLQKRVWORFDWLRQ'H0RUDHV/HZLV
GHPRQVWUDWHGWKDWWZRSDUDVLWRLGV&DUGLRFKLOHVQLJULFHSV
9LHUDQG0LFURSOLWLVFURFHLSHV&UHVVRQERWKGHSHQG
SULPDULO\ RQ KHUELYRUHLQGXFHG VLJQDOV DQG WKHVH WZR
VSHFLHVGHPRQVWUDWHDVLJQLILFDQWSODQWVSHFLHVSUHIHUHQFH
&QLJULFHSVIRUWREDFFRDQG0FURFHLSHVIRUFRWWRQ7KH
OHDIPLQHULV DSRO\SKDJRXVKHUELYRUH$UH WKHUHDQ\GLI
IHUHQFHV LQ WKH OHDIPLQHULQGXFHG YRODWLOHV DPRQJ WKH
KRVW SODQWV" &DQ WKH SDUDVLWRLG ' LVDHD VHOHFW SODQWV E\
GLVFULPLQDWLQJ DPRQJ WKHVH YRODWLOHV" &RPSDUDWLYH
UHVHDUFKRQKHUELYRUHLQGXFHGYRODWLOHVRIGLIIHUHQWSODQWV
DQG WKH EHKDYLRXUDO UHVSRQVHV RI WKH SDUDVLWRLG WR WKHP
QHHGWREHFRQWLQXHG
$&.12:/('*(0(176 :H DUH JUDWHIXO WR 'U &KDR 'RQJ
=KX ,QVWLWXWH RI =RRORJ\ &KLQHVH $FDGHP\ RI 6FLHQFHV IRU
LGHQWLI\LQJ WKH SDUDVLWRLG 'LJO\SKXV LVDHD :H DOVR H[WHQG RXU
WKDQNVWRWKHDQRQ\PRXVUHYLHZHUVIRUWKHLUKHOSIXOFRPPHQWV
7KLVUHVHDUFKZDVVXSSRUWHGE\JUDQWVIURPWKH6WDWH.H\%DVLF
5HVHDUFK DQG 'HYHORSPHQW 3ODQ *UDQW 1R *
&KLQHVH$FDGHP\RI6FLHQFHV.=%DQG$JULFXO
WXUH'HSDUWPHQW RI3HRSOH¶V 5HSXEOLFRI &KLQD*UDQW 1R
5()(5(1&(6
$6.(:556+$:05$FFRXQWRI&KDOFLGRLGHDSDUD
VLWL]LQJOHDIPLQLQJLQVHFWVRIGHFLGXRXVWUHHVLQ%ULWDLQ%LRO
-/LQQ6RF±
&2$.(5 7+ &+($+ &$ &RQGLWLRQLQJ DV D IDFWRU LQ
SDUDVLWRLG KRVW SODQW SUHIHUHQFH %LRFRQWURO 6FL 7HFK ±
&266( $$72'' -/ 0,//$5 -*0$57,1(= /$ %$5.(5
7& (OHFWURDQWHQQRJUDSKLF DQG FRXSOHG *DV &KURPD
WRJUDSKLF(OHFWURDQWHQQRJUDSKLF UHVSRQVHV RI WKH 0HGLWHUUD
QHDQ IUXLW IO\ &HUDWLWLV FDSLWDWD WR PDOHSURGXFHG YRODWLOHV
DQGPDQJRRGRXU-&KHP(FRO±
'( 025$(6 &0 /(:,6 :- 780/,1621 -+ ([DP
LQLQJ SODQWSDUDVLWRLG LQWHUDFWLRQV LQ WULWURSKLF V\VWHPV $QQ
6RF(QWRPRO%UDVLO±
'( 025$(6 &0 /(:,6 :- $QDO\VHV RI WZR SDUDVL
WRLGVZLWKFRQYHUJHQWIRUDJLQJVWUDWHJLHV-,QVHFW%HKDY
±
'(025$(6&0/(:,6:-3$5(3:$/%251+7780
/,1621-++HUELYRUHLQIHVWHGSODQWVVHOHFWLYHO\DWWUDFW
SDUDVLWRLGV1DWXUH±
',&.(09$1/221--$0XOWLWURSKLFHIIHFWVRIKHUEL
YRUHLQGXFHGSODQWYRODWLOHVLQDQHYROXWLRQDU\FRQWH[W(QWR
PRO([S$SSO±
',&.( 0 0,1.(1%(5* 23-0 5ROH RI YRODWLOH LQIR
FKHPLFDOV LQ IRUDJLQJ EHKDYLRU RI WKH OHDIPLQHU SDUDVLWRLG
'DFQXVDVLELULFD'LSWHUD$JURP\]LGDH-,QVHFW%HKDY
±
',&.(06$%(/,60:7$.$%$<$6+,-%58,1-3267+8086
0$ 3ODQW VWUDWHJLHV RI PDQLSXODWLQJ SUHGDWRUSUH\
LQWHUDFWLRQVWKURXJKDOOHORFKHPLFDOVSURVSHFWVIRUDSSOLFDWLRQ
LQSHVWFRQWURO-&KHP(FRO±
(/=(1*: :,//,$06 +- 9,1621 6% ,VRODWLRQ DQG
LGHQWLILFDWLRQ RI FRWWRQ V\QRPRQHV PHGLDWLQJ VHDUFKLQJ
EHKDYLRU E\ WKH SDUDVLWRLG &DPSROHWLV VRQRUHQVLV - &KHP
(FRO±
),1,'25,/2*/, 9 %$*1(5(6 $* &/(0(17 -/ 5ROH
RI SODQW YRODWLOHV LQ WKH VHDUFK IRU D KRVW E\ WKH SDUDVLWRLG
'LJO\SKXV LVDHD +\PHQRSWHUD (XORSKLGDH - &KHP (FRO
±
*8(5,1309,66(5-+(OHFWURDQWHQQRJUDPUHVSRQVHV
RI WKH FDUURW IO\ 3VLOD URVDH WR YRODWLOH SODQW FRPSRQHQWV
3K\VLRO(QWRPRO±
-2+1621 0: +$5$ $+ ,QIOXHQFH RI KRVW FURS RQ
SDUDVLWRLGV+\PHQRSWHUDRI/LULRP\]DVSS'LSWHUD$JUR
P\]LGDH(QYLURQ(QWRPRO±
.$1* / (FRORJ\ DQG 6XVWDLQDEOH &RQWURO RI 6HUSHQWLQH
/HDIPLQHUV6FLHQFH3UHVV%HLMLQJSS>LQ&KLQHVH@
/,*+7 '0 -$1* (% ',&.(16 -& (OHFWURDQWHQQR
JUDPUHVSRQVHVRIWKHPHGLWHUUDQHDQIUXLWIO\&HUDWLWLVFDSL
WDWD WR D VSHFWUXP RI SODQW YRODWLOHV - &KHP (FRO ±
0,1.(1%(5*23-09$1/(17(5(1-&7KHOHDIPLQHU
/LULRP\]D EU\RQLDH DQG / WULIROLL 'LSWHUD $JURP\]LGDH
WKHLUSDUDVLWHVDQGKRVWSODQWVDUHYLHZ$JULFXOWXUDO8QLYHU
VLW\ :DJHQLQJHQ 3DSHUV $JULFXOWXUDO 8QLYHUVLW\
:DJHQLQJHQ7KH1HWKHUODQGV9ROSS
1*,621* $- 29(5+2/7 :$ 1,$*, 3*1 ',&.( 0
$<(57(< -1 /:$1'( : 9RODWLOH LQIRFKHPLFDOV
XVHG LQ KRVW DQG KRVW KDELWDW ORFDWLRQ E\ &RWHVLD IODYLSHV
&DPHURQ DQG &RWHVLD VHVDPLDH &DPHURQ +\PHQRSWHUD
%UDFRQLGDHODUYDOSDUDVLWRLGVRIVWHPERUHUVRQ*UDPLQDH-
&KHP(FRO±
3$55(//$ 03 %LRORJ\ RI /LULRP\]D $QQX 5HY (QWR
PRO±
3,&.(77 -$ *DV FKURPDWRJUDSK\PDVV VSHFWURPHWU\ LQ
LQVHFWSKHURPRQHLGHQWLILFDWLRQWKUHHH[WUHPHFDVHKLVWRULHV
,Q 0F&DIIHU\ $5 :LOVRQ ,' HGV &KURPDWRJUDSK\
DQG ,VRODWLRQ RI ,QVHFW +RUPRQHV DQG 3KHURPRQHV 3OHQXP
3UHVV1HZ<RUNSS±
3,&.(77 -$ :$'+$06 /- :22'&2&. &0 ,QVHFW
VXSHUVHQVH 0DWH DQG KRVW ORFDWLRQ E\ LQVHFWV DV PRGHO V\V
WHPVIRUH[SORLWLQJROIDFWRU\LQWHUDFWLRQ7KH%LRFKHPLVW
$XJ±
5g6(8650$18.,$1$+($7+55780/,621-+
9RODWLOH VHPLRFKHPLFDOV UHOHDVHG IURP XQGDPDJHG FRWWRQ
OHDYHV $ V\VWHPLF UHVSRQVH RI SODQWV WR FDWHUSLOODU GDPDJH
3ODQW3K\VLRO±
6$/92 $ 9$//$'$5(6 *5 $Q DQDO\VLV RI OHDIPLQHU
DQG SODQW KRVW UDQJHV RI WKUHH &KU\VRFKDULV VSHFLHV +\P
(XORSKLGDHIURP$UJHQWLQD(QWRPRSKDJD±
6&+867(5 '- *,/5($7+ -3 :+$5721 5$ 6(<0285 35
$JURP\]LGDH'LSWHUDOHDIPLQHUVDQGWKHLUSDUDVLWRLGV
LQ ZHHGV DVVRFLDWHG ZLWK WRPDWR LQ )ORULGD (QYLURQ
(QWRPRO±
7$.$%$<$6+, - ',&.( 0 3267+8086 0$ 9DULDWLRQ
LQ FRPSRVLWLRQRI SUHGDWRUDWWUDFWLQJDOOHORFKHPLFDOV HPLWWHG
E\ KHUELYRUHLQIHVWHG SODQWV UHODWLYH LQIOXHQFH RI SODQW DQG
KHUELYRUH&KHPRHFRORJ\±
785/,1*7&-780/,1621-+(//(5)-/(:,6:-
/DUYDOGDPDJHG SODQWV 6RXUFH RI YRODWLOH V\QRPRQHV WKDW
JXLGH WKH SDUDVLWRLG &RWHVLD PDUJLQLYHQWULV WR WKH PLFUR
KDELWDWRILWVKRVWV(QWRPRO([S$SSO±
785/,1* 7&- 780/,1621 -+ /(:,6 :- ([SORLWD
WLRQRIKHUELYRUHLQGXFHGSODQWRGRXUVE\KRVWVHHNLQJSDUD
VLWLFZDVSV6FLHQFH±
9$1 /(17(5(1 -& :2(7 - %LRORJLFDO DQG LQWHJUDWHG
SHVW FRQWURO LQ JUHHQKRXVHV $QQX 5HY (QWRPRO ±
9,66(5-+(OHFWURDQWHQQRJUDPUHVSRQVHVRIWKHFRORUDGR
EHHWOH /HSWLQRWDUVD GHFHPOLQHDWD WR SODQW YRODWLOHV (QWRPRO
([S$SSO±
=(+1'(5 *: 7580%/( -7 +RVW VHOHFWLRQ RI /LULR
P\]D VSHFLHV 'LSWHUD $JURP\]LGHD DQG DVVRFLDWHG SDUD
VLWHV LQ DGMDFHQW SODQWLQJV RI WRPDWR DQG FHOHU\ (QYLURQ
(QWRPRO±
=+$2 <; .$1* / D 5ROH RI SODQW YRODWLOHV LQ KRVW
SODQW ORFDWLRQ RI WKH OHDIPLQHU /LULRP\]D VDWLYDH 'LSWHUD
$JURP\]LGDH3K\VLRO(QWRPRO±
=+$2 <; .$1* / E 2OIDFWRU\ UHVSRQVHV RI WKH OHDI
PLQHU /LULRP\]D VDWLYDH 'LSWHUD $JURP\]LGDH WR WKH
RGRXUV RI KRVW DQG QRQKRVW SODQWV - $SSO (QWRPRO LQ
SUHVV
=+$2 <; .$1* / ,PSURYHPHQW RI HOHFWURDQWHQQR
JUDP WHFKQLTXH IRU GLSWHUDQ IOLHV (QWRPRO .QRZOHGJH ±>LQ&KLQHVH@
=+$2 <; .$1* / &ROG WROHUDQFH RI WKH OHDIPLQHU
/LULRP\]D VDWLYDH 'LSW $JURP\]LGDH - $SSO (QWRPRO
±
5HFHLYHG1RYHPEHUUHYLVHG-DQXDU\DFFHSWHG0D\