全 文 ：2010年 12月
第 33卷 第 4期
湖南师范大学自然科学学报
JournalofNaturalScienceofHunanNormalUniversity
Vol.33 No.4
Dec., 2010
AntifeedantandDeterrentActivityofExtractsfrom
DifferentPartsofXanthiumSibiricum(Compositae)
againstCruciferPests
ZHOUQiong1＊
①
, WEIMei-cai2 , YANYu-chen2 , WANGWen-xue2
(1.CollegyofLifeScience, HunanNormalUniversity, Changsha410081, China;
2.LaboratoryofInsectSystematicsandEvolutionaryBiology, CentralSouthUniversityofForestry＆Technology, Changsha410004, China)
Abstract Antifeedant, andovipositionorinhabiting(setling)deterentefectsofmethanolex-
tractsfromthefruit, leaf, stemandrootofXanthiumsibiricumisinvestigatedinalaboratorysetingusing
diferentbioassays, aimatdeterminingwhichpartextractsismoreefectiveonthefourkeyinsectpestsof
crucifers, includingPlutelaxylostela, Pierisrapae, MyzuspersicaeandLipaphiserysimi.Antifeedant
efectsareexaminedinthelarvaofP.xylostelaandP.rapae, usingleafdiscchoicebioassays, inaddi-
tiontoovipositiondeterentefectsofadultP.xylostelausingafreshleafdiscchoicebioassay, andin-
habitingdeterentefectsofL.erysimiandM.persicaeusinghalf-leafchoicebioassays.Theresultsshow
thattheleafextractsshowthehighestantifeedantrateoveralforP.rapae, withthemostefectivean-
tifeedantrateof99.07% , andfruitextractsprovethehighestantifeedantrateforP.xylostelawitharate
of82.87%, whilethemostefectiveovipositiondeterentrate, derivedfromleafextracts, isadeterent
rateof88.57%forP.xylostela.Themostefectiveinhabitingdeterentrateisalsorecordedfromleafex-
tractsforL.erysimi, 83.01% and62.50% forM.persicae.ExtractsfromX.sibiricumfruitpresenteda
morepotentovipositionandinhabitingdeterrentforthethreeinsects, thandideitherstemorrootex-
tracts.Basedontheirantifeedant, andovipositiondeterentandinhabitingdeterentproperties, theac-
tivecomponentsfromX.sibiricumshowthepotentialtobeusedasalternativecropprotectantagainsta
numberofpestspecies.
Keywords ExtractsfromXanthiumsibiricum;antifeedant;oviposition/inhabiting(setling)de-
terent;insectpestsofcrucifers
中图分类号 Q965.9 文献标识码 A 文章编号 1000-2537(2010)04-0099-08
苍耳不同部位提取物对十字花科蔬菜害虫的拒食和忌避活性
周 琼1＊ , 魏美才 2 , 晏毓晨2 , 王问学 2
(1.湖南师范大学生命科学学院 , 中国 长沙 410081;
2.中南林业科技大学昆虫系统与进化生物学实验室 , 中国 长沙 410004)
摘 要 为确定苍耳 Xanthiumsibiricum对昆虫活性最高的部位 ,测试了苍耳不同器官(果实 、叶 、茎和根)甲醇
提取物对几种重要十字花科蔬菜害虫(包括小菜蛾 Plutelaxylostela, 菜青虫 Pierisrapae, 桃蚜 Myzuspersicae和萝卜
蚜 Lipaphiserysimi)的拒食活性 、产卵忌避活性或忌避活性.结果表明 ,苍耳的主要活性部位为地上部分 ,尤其是叶和
果实 , 其次是茎 ,根的活性最低.拒食活性测试中 , 苍耳叶提取物对菜青虫最强 ,在所测试的浓度下 ,处理后 24 h的选
① 收稿日期:2010-03-06
基金项目:ThisprojectwassupportedbytheScientificResearchFundofHunanProvincialNaturalScienceFoundationofChina
(06JJ20099)andHunanProvincialEducationDepartment(05C336)
＊通信作者 , E-mail:zhoujoan2004@163.com
择性拒食率高达 99.07%;对小菜蛾幼虫的拒食作用以果实和叶提取物最强 , 选择性拒食率分别高达 82.87%和
73.42%.产卵忌避 /忌避活性最强的是苍耳叶提取物 ,处理后 24 h对小菜蛾成虫的选择性产卵忌避率达 88.57%,对
萝卜蚜和桃蚜的选择性忌避率分别为 83.01%和 62.50%, 同时 ,苍耳果实提取物对小菜蛾 、桃蚜和萝卜蚜的产卵忌
避 /忌避活性明显高于茎和根提取物.基于苍耳叶和果实提取物对害虫强烈的拒食 、产卵忌避或忌避活性 , 有可能作
为植物保护剂配方应用于害虫的综合治理策略中.
关键词 苍耳;拒食活性;产卵忌避 /忌避活性;十字花科蔬菜害虫
Becauseofincreasingproblemsassociatedwiththeuseofacutelytoxicsyntheticinsecticides, includingresist-
anceandtheimpactsonbothhumansandnon-targetbiologicalsystems[ 1-3] , thereisapressingneedforthedevel-
opmentofsafe, alternativecropprotectants.Amongthemostefectiveinsectpestcontrolmeasures, abotanicalreg-
ulatorofinsectpestbehaviors, suchasantifeedantsandovipositiondeterents, maybeoneofthemostpreferedop-
tions.Botanicalproductsareusefulanddesirabletoolsinmostpestmanagementprogramsbecausetheycanbeef-
fectiveandoftencomplementtheactionsofnaturalenemies[ 4-5] .Plantsecondarycompoundshavebeenthesubjectof
thoroughinvestigationsinaneforttodiscovernewsourcesofbotanicalinsectantifeedantsandovipositiondeterrents.
Thediamondbackmoth(Plutelaxylostela(Linnaeus)), cabbagewhitebuterfly(Pierisrapae(Linnae-
us)), peachaphid(Myzuspersicae(Sulzer)), andmustardaphid(Lipaphiserysimi(Kaltenbach))aredestruc-
tiveinsectsofcruciferousvegetablesinChinaandthroughouttheworld[ 6-8] .Someofthem, especialyP.xylostel-
la, havebeendificulttocontrol, andhavedevelopedaconsiderableresistancetoarangeofinsecticides, including
organophosphorus, carbamates, pyrethroids, Bacilusthuringiensis, andbenzoylphenylureas[ 1, 6, 9-13] .Thereisan
urgentneedtofindnewtypesofsafe, alternativecropprotectantsagainsttheseinsectpestsofcrucifers.
Siberiancocklebur(XanthiumsibiricumPatrin)(Compositae)isanannualdicotyledonousweedthatisex-
tremelycompetitivewithsoybeanandotheragronomiccropsinChina[ 14-16] .Itsfruitisusedasatraditionalherbal
medicineinChinafortheefectivetreatmentofsinusitis, urticaria, andarthritis, andtoregulateimmunityandre-
strainbacteria[ 17-18] .IthasbeenusedasatraditionalpesticideinChinasinceancienttimes[ 19] .Theethanolex-
tractsfromX.sibiricumprovedhighlyefectivegrowthinhibitorsandproducedstomachpoisoninP.rapae[ 20] .A-
mongthem, thealcoholextractsfromX.sibiricumprovedtohaveahighlyefectiveantifeedantefectonlaboratory
populationsofL.erysimiandM.persicae, butsafefornaturalenemiesofaphidssuchasladybirdbeetles(Menochi-
lussexmaculata(Fabricius)andCoccinelatrasversalis(Fabricius)), andparasitoids(AphidiusgifuensisandDiae-
retielarapae)[ 21-23] .ThepurposeofthecurentstudyistoexamineandcomparewhichpartextractsfromX.sibiri-
cumplanthavemoreantifeedant, ovipositionandinhabitingdeterrentefectsoninsectpestsofcruciferouscrops,
andtoserveasthefoundationformoredetailedinvestigationsofthosepartsthatprovetobemostactiveonthefeed-
ingdeterentandovipositiondeterentbehavioursofphytophagousinsects.
1 MaterialsandMethods
Plantextracts Extractsfromthefruit, stem, leaf, androotofX.sibiricumwerepulpedfromfreshplantma-
terialcolectedfromthesuburbofXiangtan(112°56′E, 27°50′N), P.R.China, inOctober-November2005.
ThefourpartsofX.sibiricumwereair-dried, thenfurtherdriedat50℃ inanovenwithaircirculationfor24 ～ 48
h, andgroundinanelectricblenderintoapowder, respectively.Eachofthefourpowders, leaf, fruit, stem, and
root, wereextractedbymethanolfor24 hinaSoxhletextractor, andtheextractedsolutionswerefiltered.Thefil-
teredsolutionswereevaporatedinarotaryevaporatorunderreducedpressuretoobtainthecrudeextracts.The
crudeextractswerestoredinarefrigeratorandtemporarilydilutedwithdistiledwaterbeforetesting.
Hostplant Pakchoicabbages(Brasicachinensis)weregrownin500mm×500mm×500mmplasticpots
containingamixtureofsandyloamsoilandpeatmosstothesixtosevenfulyextendedtrueleafstage.Thepots
werehousedinanalnicocage, whichwasplacedoutdoors, withanambienttemperatureandhumidity, enoughnat-
urallightforbioassays, andtomaintainP.xylostela, P.rapae, M.persicae, L.erysimicoloniesontheirleaves.
100 湖南师范大学自然科学学报 第 33卷
Testinsects LaboratoryP.xylostela, P.rapae, M.persicae, andL.erysimicultureswereestablishedfrom
fieldcolectionsonacabbagefarminasuburbofChangsha(111°53′E, 27°58′N), P.R.China, andmaintained
onindividualypotedPakchoicabbagesinthecage, inaroomtemperature-controledat25℃, 60% ～ 80%RH
andnaturephotoperiod.
AntifeedantchoicebioassayonP.rapaeandP.xylostela Theantifeedantactivityofleaf, fruit, stem,
androotextractsfromX.sibiricumwasassayedbyusingaleafdiscchoiceassay.FreshleavesofB.chinensiswere
excisedinto20mm×20mmsquarediscs, andthendippedinoneofthedilutedcrudeextracts(0.05g/mL, dried
powderweight)foraboutfiveseconds.Theleafdiscsthatservedascontrolsweredippedindistiledwaterand5%
methanol.Afterair-dryingfor1 h, fourpakchoileafdiscswereplacedina125mmdiameterPetridish, withtwo
treatmentsandtwocontrols, arangedalternately, ineachdishandunderlayingwithadampfilterpapertokeep
leafdiscsfreshfor24 h.Five3rdinstarlarvaewhichpre-starvedfor3 h, wereplacedatthecenterofeachPetri
dish, andalowedtofeedfor24hat25℃.Theleafdiscswerereplenishedifneeded.Eachtreatmentwasreplica-
tedsixtimes.AcleargraphpaperwasusedtomeasuretheleafareaeatenbylarvaeofP.rapaeandP.xylostela,
andtheproportionofeatenleaftothetotalleafareawascalculated.
OvipositionpreferenceofP.xylostela Inthechoiceovipositiondeterrentassays, animprovedmethodfol-
lowing[ 24] wasused.TheleafdiscsofB.chinensisexcisedinto18 mmdiameterweretreatedassameasinthe
choicefeedantdeterrentassays, andadhibitedtothebotomof80mmdiameterPetridisheswith2%agargel.The
leafsurfacefacedthebotomofthePetridish, soastokeeptheleafdiscfreshforseveraldays.Thetwotreatments
(0.05 g/mL, driedpowderweight)andtwocontrolswerealternatelyarangedinthePetridish.ThePetridishwas
thenplacedupsidedowntocoverthetopofaplasticcylinder(diam75 mm), andthebotomwaswrappedbya
pieceofgauzetomaintainventilation.Threepairsofadults, maleandfemale, emergedfortwoorthreedays, were
confinedineachcylinderforthreedaysat25℃.Adultsweresuppliedwith10% honey-watersolutionbyasoaked
smalcotonbalinthefundusofthecylinder.Thenumberofeggslaidontheleafdiscwascounteddaily.Each
treatmentwasreplicatedtwelvetimes.
SettlingdeterrentchoicebioassayonM.persicaeandL.erysimi Animprovedhalf-leafdiscbioassay[ 21, 25]
wasapplied.TheleavesofB.chinensiswereexcisedtoformroundpieces(diam40 mm)withveininthecerter.
Thentheleafdiscswereadhibitedon2% agargelasmentionedabove.Thedilutedcrudeextracts(0.05 g/mL,
driedpowderweight)werespreaduniformlyononesideofthebackoftheleafastreatment, andtheisodosealco-
holanddistiledwaterwasspreadontheothersideasthecontrol.Folowingthespreadextractleafdiscsbeingair-
dried, 16 ～ 22 3rdinstaraphidswerecarefulyplacedontotheeachleaf-discbackinequalnumberonbothsidesof
theleafvein.ThePetridishwassealedbyaparafilmprickedwithairholesbyaninsectneedletokeepitventila-
ted, turnedupsidedownandheldat25 ℃.Thenumbersofaphidsthatsetledoneitherthetreatedorcontrolsides
ofeveryleafdiscwererecordedat24h, 48h, and72 haftertreatment.Eachtreatmentwasreplicatedthreetimes.
Dataanalyses Anon-choice/ choicefeeding(oviposition)deterenceratewascalculatedusingtheformula:
Choicefeeding(orinhabiting/oviposition)deterencerate(%)=100% [ (C-T)/(C+T)] , whereCand
Tarethecontrolandtreatedleafareasconsumedbytheinsects(Ismanetal.1990), ornumbersofaphidssetled
inthecontrolandtreatedleaf-halfdisc[ 21] , ornumbersofeggsdepositedonthecontrolandtreatedleafdiscbythe
diamondbackmoth[ 25] .
DataanalyseswerecariedoutusingSPSS(StatisticalProductandServiceSolutions, StatisticsPackageforthe
SocialSciences)onthebasisofactualnumbersobserved.Paired-samplesTTestandDuncansMultipleRangeTest
(DMRT)wereusedtodeterminethesignificantdiferencesbetweenthecontrolgroupsandtreatedgroups.
2 Results
AntifeedantchoiceassayonP.rapaelarvae Theantifeedantefectofcrudeextractsfromdiferentpartsof
X.sibiricumonP.rapaewasshowninTable1.ThePakchoicabbageleafareaseatenbyP.rapaelarvaeforthecon-
trolandthetreatmentspreadedbyleafandftruitcrudeextractsshowedasignificantdiference.Theproportionof
101第 4期 周 琼等:苍耳不同部位提取物对十字花科蔬菜害虫的拒食和忌避活性
PakchoicabbageleafareaeatenbyP.rapaelarvaeforthecontrolsamplewas99.53%, whereastheleafextract
treatedsamplewas0.47%.Forthefruitextractexperiment, thefigureswere98.06%forthecontroland1.94%
forthetreatment.Thehighestantifeedantrateof99.07% wasachievedfromtheleafextractofX.sibiricum.The
fruitextractfolowedwithantifeedantrateof96.13%.Theremainingstemandrootextractsproducedanantifeed-
antrateof45.60% and32.57%, respectively, butthetreatedpakchoicabbageleafareaseatenbyP.rapaelarvae
didnotshowanysignificantdiferencefromthecontrolgroups.
Tab.1 AntifeedanteffectofmethanolextractsfromfourpartsofX.sibiricumonP.rapaeandP.xylostellainleafdiscchoicebioassays
Testedinsects Extractedparts
Meanleafareas(±SE)offeeding(mm2 perinsect)
Control Treatment
Choiceantifeedant
rate/%
P.rapae
Leaf 641.00±79.73 a 3.00±0.58 b 99.07
Fruit 573.67±136.43a 11.33±5.50 b 96.13
Stem 475.75±87.90 a 177.75±165.11 a 45.60
Root 504.00±103.47a 256.33±111.34 a 32.57
P.xylostela
Fruit 151.67±24.56 a 14.21±8.67 b 82.87
Leaf 142.08±17.78 a 21.78±10.68 b 73.42
Stem 86.42±23.68 a 39.50±29.01a 37.26
Root 177.32±25.19 a 100.84±35.70a 27.49
Note:MeansfolowedbycommonletersinarowarenotsignificantlydiferentatP=0.05 byPaired-samplesTTest.
AntifeedantchoiceassayonP.xylostela IntheantifeedantchoiceassayonP.xylostela, therewasalsoa
significantdiferenceinthecontrolandfruitandleafextracttreatmentsonthepakchoicabbageleafareaseatenby
P.xylostelalarvae(Table1).TheproportionofPakchoicabbageleafareaseatenbyP.xylostelalarvaeonthe
controlsamplewas91.43%, whereasthefruitextracttreatmentproduced8.75%.Thecorrespondingmeasuresfor
theleafextractspreadexperimentwere86.71% forthecontroland13.29%forthetreatment.Thefruitextractex-
hibitedastrongantifeedantcapability, withantifeedantrateof82.87%, folowedbytheleafextracttreatmentwith
antifeedantrateof73.42%.Theantifeedantratefromthestemandrootextractsindicatedalowerantifeedantefect
thaneitherfruitorleafextracts, witharateof37.26% and27.49% respectively.Onceagain, thestemandroot
extractassaysshowednosignificantdiferencebetweencontrolandtreatmentforthetreatedpakchoicabbageleafar-
easeatenbyP.xylostelalarvae.
Deterrentchoicebioassay ThecrudeextractsofX.sibiricumshowedanobviousovipositiondeterentefect
onP.xylostela, andaninhabitingdeterentefectonL.erysimiandM.persicaeinthedeterentchoicebioassay.
Theovipositiondeterrentactivityforthefourextractswascalculatedbyovipositiondeterentratevalues.Theinhab-
itingdeterrentefectonaphidswasalsovariedforthefourextractsandwascalculatedbyinhabiting(setling)de-
terentratevalues.
OvipositiondeterrentchoiceassayonP.xylostela Thecrudeextractsfromtheleaf, fruit, andstemofX.
sibiricumshowedobviousovipositiondeterrentefectsonP.xylostela(Table2).P.xylostelafemalesdepositedan
averageof22.77eggs(94.29%)onthecontrolsand1.38eggs(5.71%)ontheleafextracttreatments24hafter
spread.Theratewasfolowedby43.69 eggs(81.25%)onthecontrolsand10.08eggs(18.75%)onleafdiscs
treatedwithfruitextract.Thestemextractexperimentresultedinfemalesdepositing, anaverage, 34.67 eggs
(72.06%)onthecontrolsand13.44 eggs(27.94%)onthetreatmentspreadwithstemextract.Asaresult, the
crudeextractwithhighestovipositiondeterrentactivityonP.xylostelawasleafextractfromX.sibiricum, withtheo-
vipositiondeterentratesof88.57%(24 h), 78.91%(48h), and69.88%(72 h)afterspread, folowedbythe
fruitextractwithadeterrentrateof62.51% (24 h), 50.67% (48 h), and42.70% (72 h)afterspread.The
stemextracttreatmentovipositiondeterrentratewas44.13% (24h), 29.19% (48 h), and32.46%(72h)af-
terspread.Therootextractexperimentshowednoovipositiondeterentefectonthediamondbackmoth.Theaccu-
mulativeovipositiondeterrentratereducedastimeelapsed.
102 湖南师范大学自然科学学报 第 33卷
Tab.2 OvipositiondeterrentefectofextractsfromfourpartsofX.sibiricumonP.xylostelainleafdiscchoicebioassays
Treatmentpairs
AccumulativeMean(±SE)numbersofeggsdepositedbyDiamondback
mothfemalesonPakchoicabbageleaf-discsprayedwithextracts
Ovipositiondeterrent
rate/%
24h 48 h 72 h 24h 48 h 72 h
Leafextracts/Control 1.38±0.72B/22.77±6.57A 7.00±2.16B/59.38±10.36A 13.54±3.28B/76.38±9.63A 88.57 78.91 69.88
Fruitextracts/
Control
10.08±2.75B/
43.69±6.88A
24.23±4.38B/
74.00±9.13A
38.92±5.54B/
96.92±8.44A 62.51 50.67 42.7
Stemextracts/Control 13.44±4.03B/34.67±7.84A 28.56±5.45B/52.11±7.77A 46.11±10.50B/90.44±18.01A 44.13 29.19 32.46
Rootextracts/
Control
5.00±3.15A/
3.34±2.98A
13.80±4.21A/
10.90±4.32A
17.20±5.70A/
15.60±4.70A -19.9 -11.74 -4.88
Note:MeansfolowedbycommonletersinapairarenotsignificantlydifferentatP=0.01byPaired-samplesTTest.
InhabitingdeterrentchoiceasayonL.erysimi Intheinhabitingdeterentchoiceassay, theinhabiting
deterrentactivityonL.erysimivariedaccordingtotheextractofX.sibiricumapplied(Table3).Theaveragenum-
bersofL.erysimiinhabitingonthePakchoicabbageleafweresignificantlydiferentbetweenthecontrolsectionand
thesectiontreatedwithleaf, fruit, orstemcrudeextracts24 hlater.Themostactivecrudeextractwasfromthe
leavesofX.sibiricumwithachoicedeterentrateof83.01% (24h), 54.99% (48 h), and43.54%(72h)af-
tertreatment.Thefruitextracttreatmentreturnedachoicedeterentrateof53.52% (24 h), 44.98 % (48h),
and27.83% (72h)aftertreatment.Thestemextractsprovedtobemostactive24haftertreatmentwithaninhab-
itingdeterentrateof62.50%, buttherewasnodeterentefect48hand72haftertreatment.Therootextractac-
tivitywasthelowestat24 h(17.33%), andnodeterentefectwasshownafter48 hor72h.Theinhabitingde-
terentefectofalextractsfromX.sibiricumwasreducedastimeelapsed, butthedegreevariedaccordingtothe
particularextract.Thestemextractdecreasedfastestfrom62.50% at24 hto-5.91% at48haftertreatment.
Tab.3 InhabitingdeterenteffectofextractsfromfourpartsofX.sibiricumonL.erysimiandM.persicaeinhalf-leafchoicebioassays
Tested
insects
Treatment
pairs
Mean(±SE)numbersofsettlingofaphidsinhabitingonPakchoicabbagehalf-leafsprayedwithextracts Inhabitingdeterrentrate/%
24h 48 h 72 h 24h 48 h 72h
L.erysimi
Leafextracts/Control 1.67±0.67b/14.33±1.33a 3.00±1.15b/10.33±1.20a 3.67±1.76a/9.33±1.86a 83.01 54.99 43.54
Fruitextracts/Control 3.33±0.67b/11.00±1.53a 3.67±1.67b/9.67±2.03a 4.33±1.76a/7.67±1.86a 53.52 44.98 27.83
Stemextracts/Control 3.00±0.58b/13.00±0.58a 6.00±0.58a/5.33±0.33a 6.33±3.28a/5.33±2.03a 62.50 -5.91 -8.58
Rootextracts/Control 6.20±1.59a/8.80±1.50a 7.40±0.51a/6.20±1.24a 5.20±0.80a/5.00±1.05a 17.33 -8.82 -1.96
M.persica
Leafextracts/
Control
3.00±1.52b/
13.00±2.08a
4.00±0.58b/
11.00±1.15a
3.00±1.15b/
10.67±0.67a 62.50 46.67 56.11
Fruitextracts/
Control
4.00±1.52b/
13.33±2.84a
6.33±0.88a/
9.67±0.88a
7.67±0.88a/
9.00±1.00a 53.84 20.88 7.98
Rootextracts/
Control
5.00±0.63b/
9.40±0.51a
3.40±0.40a/
5.00±0.32a
7.20±0.66a/
6.00±0.63a 30.56 19.05 -9.09
Stemextracts/
Control
7.33±0.88a/
8.67±0.88a
6.33±0.67a/
8.33±1.20a
8.33±0.67a/
6.67±0.88a 8.38 13.64 -11.07
Note:MeansfollowedbyacommonleterinapairmeantherearenotsignificantlydiferenceatP=0.05byPaired-samplesTTest.
InhabitingdeterrentchoiceasayonM.persicae Theinhabitingdeterrentactivityvariedaccordingtothe
particularextractofX.sibiricumagainstM.persicae(Table3).TheaveragenumberofM.persicaeinhabitingonthe
pakchoicabbageleaf24haftertreatmentwassignificantlydiferentbetweenthecontrolsectionandthesectiontrea-
tedbyleafextract.Howeveritshowedasmalerornodiferencebetweenthecontrolandthesectiontreatedwith
thefruit, stem, androotextracts.ThemostactiveextractwasfromtheleafofX.sibiricumwithachoiceinhabiting
deterentrateof62.50% (24h), 46.67% (48 h), and56.11% (72h)aftertreatment, folowedbythefruitex-
103第 4期 周 琼等:苍耳不同部位提取物对十字花科蔬菜害虫的拒食和忌避活性
tractwith53.84%(24h), 20.88%(48h), and7.98% (72h)aftertreatment.Therootandstemextractsshowed
anevensmalerinhabitingdeterrentactivityrateat24hand48haftertreatment, andnodeterrentefectat72h.
ComparativeactivityofthefourextractsofX.sibiricum Theresultsoftheleafdiscchoicebioassayson
P.xylostelaandP.rapaesuggestthatthehighestantifeedantactivecomponentsprincipalyexistintheleafandfruit
extractsofX.sibiricum(especialyonP.rapaewithachoiceantifeedantrateover96% atthetestedconcentra-
tion), folowedbythestemextracttreatment.Thelowestantifeedantactivitywasreturnedfortherootextractsolu-
tion.Intheoviposition/inhabitingdeterrentbioassaysonP.xylostela, L.erysimi, andM.persicae, themostactive
componentswerealsoextractedfromtheleavesandfruits, inparticulartheleaves, folowedbythestemextract,
andtherootextractlastofal.However, therootextractdidshowahigherinhabitingactivitythanthestemextract
againstM.persicae.
3 Discussion
X.sibiricumisanon-hostplantforthefourherbivorestested.Ourresultsdemonstratedthattheefectofan-
tifeedantandoviposition/ inhabitingdeterentsonthetestinsectsvariedaccordingtotheparticularextractofthe
plant.Themostactivetreatmentswerefromleaves, fruitsandstemextracts, whichindicatesthattheinsect-suscep-
tibleactivecomponentsexistedmostlyintheabovegroundpartsofX.sibiricum.Theleavesandfruitscontained
moreantifeedantandoviposition/inhabitingdeterrentactivecomponentsthaneitherthestemsorroots.Theactive
chemicalsintheabovegroundpartsoftheplantcouldinterferewiththehost-locationofphytophagousinsects, and
deterthemfrominhabitingordepositingeggsontheplantleaf.Theresultsalsoindicatethattheextractspreventthe
insectsfromingestingtheplantandthusimpedethesepestsbuilding-uptheirpopulationsonX.sibiricum.Moreo-
ver, thetreatmentsfromtheleafandfruitextractsmightpossessintensiveinsecticidalactivitytolarvaP.rapaeby
inducingraisedlarvalmortality, abnormalpupaeorpupamortality, orcrimpledwingsofemergedadultsinthisex-
periment.Basedontheirantifeedantandoviposition/ inhabitingdeterentproperties, theefectiveactivecompo-
nentsfromX.sibiricumhavepotentialforuseasalternativecropprotectantagainstanumberofpestspecies.
ItwasreportedthatXanthiumspeciesplantscouldproducevariantcombinationsofsesquiterpenelactonesin
theirleavesandfruits, suchasxanthinin, xanthanol, xanthatin, xanthumin, xanthumanolanddeacetoxylxanthu-
min[ 26-29] .Xanthanolides, isolatedfromXanthiumspecies, wereusedasananti-atachingrepelentagainstBlue
mussel[ 30] , andactedasinsectdevelopmentinhibitors[ 31] .Further, moredetailedstudiesarewarantedtodeter-
minewhetherornottheantifeedantandoviposition/inhabitingdeterentactivecomponentsaresesquiterpenelac-
tones, whetheroneormoreingredientsaremostactive, andhowinsectsareafected.Furtherinvestigationmayalso
determinehowtheextractsfromdiferentstagesofthedevelopmentofX.sibiricummightprotectplantsfromsome
herbivoreinsectsandhowitmightvarywiththeseasons.
Theefectofplantdefensivecompoundsisdiversewithdiferentinsects, eventhoughtheirfoodpreferencesare
similar.Theefectofdefensivecompoundsshowedmoreactivityforspecialiststhanforgeneralists.Inthisstudy,
amongthefourtestedvegetableinsectpests, M.persicaeisageneralist, withhostrangeofmorethan400plantspe-
ciesfrommorethan40 families[ 32] .Theotherthreeinsects, P.xylostela, P.rapae, andL.erysimiarespecialist
herbivoreswithlimitedfeedingchoicesofthefamilyCruciferae[ 6] .Ourresultsfoundthattheleafandfruitextracts
fromX.sibiricumhadthehighestantifeedantefectonP.rapae, withanantifeedantrateof99.07% and96.13%
respectively.TheefectonP.xylostelawas73.42% (leaf)and82.87% (fruit).Theleafextracthadhigherin-
habitingdeterentefectonL.erysimi(83.01%)thanonM.persicae(62.50%).Thisresultisconsistentwith
Bernays(2000)whoconcludedthatspecialistshavegreatersensitivitytodeterentsthangeneralists[ 33] .Thehigh
abilityofP.xylostelatodevelopresistancetoalmostalgroupsofinsecticides[ 6] mightbeonereasonwhythefeed-
ingdeterentefectofX.sibiricumextractswaslesssensitiveonP.xylostelathanonP.rapae.
Theearliestformsoflandplantsandinsectshavecoexistedforaslongas350milionyears[ 34] .Onthebasisof
thislong-standingrelationship, thestrategiesemployedbyplantstoresistorevadetheirinsectherbivoresarevery
104 湖南师范大学自然科学学报 第 33卷
diverse.Accumulatinghighlevelsofcompoundsthatfunctionasbiochemicaldefensesthroughtheirtoxicityisone
oftheimportantstrategiesthatplantsseektominimiseherbivoredamage.Asaresult, aninsectherbivoreonly
feedsonalimitedrangeofplantspecies, anddamagearelimitedbythenumberinsectherbivoresatractedtoapar-
ticularspecies[ 35] .Moreover, theplantsecondarycompoundsaregenus-orspecies-specific[ 36] .Thesensitivityofa
rangeofphytophagousinsectswithsimilarhostpreferenceisdiverse.Thisrestrictstheexcessingestingofhost
plantsbytheinsectherbivores.Consequently, theplantpopulationavoidsbeingwipedout, andthephytophagous
insectpopulationsalsosurvive, whichaccountsforthedefensivecompoundsfoundinplantsplayingasignificant
roleinregulatingtherelationshipbetweeninsectsandplants, andforthesurvivalandevolutionofplants.
Isman(2006)suggestedthattoproduceabotanicalinsecticideonacommercialscale, thesourceplantbio-
massmustbeobtainableonanagriculturalscale, andpreferably, notonaseasonalbasis.Unlesstheplantinques-
tionisextremelyabundantinnature, oralreadygrownforanotherpurpose, itmustbeamenabletocultivation[ 37] .
X.sibiricummeetsthiscriterionbecauseofitshighlyadaptabletoarangeofenvironments, andeasilyspreadingand
planting[ 14] .Itmayserveasapotentialnewtypeofbotanicalbehaviorregulativeprotectantagainstinsectpests,
anditissafefornaturalenemies, humanbeings, andtheenvironment.Theantifeedantandovoposition/inhabiting
activecompoundsinX.sibiricumisefectiveagainstinsectherbivores, althoughthetypeandstrengthofthedefen-
sivecompoundsfromdiferentstagesofgrowth, andfromvariouspartsoftheplantneedtobeinvestigatedfurtherin
ordertohelpustounderstandthenaturalmechanismsofnon-hostplantsagainstinsectherbivores.Suchelucidation
mayleadtothedevelopmentofnewtypesofbehaviorregulatorsininsectpestmanagement.
Acknowledgements WeexpressourappreciationtoourAustraliancoleague, Mr.BrianHilyar, forrevie-
wingearlierdraftsofthemanuscript.
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(编辑 王 健)
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