全 文 ：第20卷 第4期
Vol.20 No.4
草 地 学 报
ACTA AGRESTIA SINICA
2012年 7月
Jul. 2012
PreyResponsesofPropylaeajaponicainDifferentInstars
toAcyrthosiphonpisumunderGrowthChamberCondition
WUDe-gong1,2,DUJun-li1,LIUChang-zhong1,HUGui-xin1,WANGSen-shan1,HEChun-gui1,3*
(1.KeyLaboratoryofGrasslandEcosystem,MinistryofEducation/Sino-USCenterforGrasslandandLivestockSustainable
Development,AppliedEntomologyLaboratory,ColegeofPrataculturalScience,GansuAgriculturalUniversity,Lanzhou,
GansuProvince730070,China;2.PlantScienceColegeofAnhuiScienceandTechnologyUniversity,Fengyang,
AnhuiProvince233100,China;3.GansuAcademyofAgriculturalSciences,Lanzhou,GansuProvince730070,China)
Abstract:ThefunctionalresponseofPropylaeajaponicaThunberg(Coleoptera:Coccinelidae)
tofivedensitiesofAcyrthosiphonpisum Harris(Homoptera:Aphididae)wasinvestigatedby
discmethodundergrowthchambercondition,inordertounderstandthecontrolingcapabilityof
P.japonicatoA.pisum.Plottingpreydensityagainstpreykiledbyfourlarvalinstars,adult
malesandfemalesofPropylaeajaponicafitwelwiththetypeⅡ modelofHoling’sdiscequa-
tion.Adultfemalesconsumedthehighestnumberoftheprey,folowedbyfourthinstarsanda-
dultmales.Basedonthefunctionalresponsedata,themodelpredictsamaximumof6.5,20.4,
32.3,71.4,58.8,and80.0nymphstobeconsumedperdaybyindividualfirst,second,thirdin-
star,adultmale,fourthinstar,andadultfemale,respectively.Theresultsindicatedthepreda-
tionofP.japonicaonA.pisumwasgreater.
Keywords:Propylaeajaponica;Acyrthosiphonpisum;Functonalresponse;Predation
不同龄期龟纹瓢虫对豌豆蚜的捕食功能反应
武德功1,2,杜军利1,刘长仲1,胡桂馨1,王森山1,贺春贵1,3*
(1.甘肃农业大学草业学院 草业生态系统教育部重点实验室/中-美草地畜牧业可持续发展研究中心,甘肃 兰州 730070;
2.安徽科技学院 植物科学学院,安徽 凤阳 233100;3.甘肃省农业科学院,甘肃 兰州 730070)
摘要:为了解龟纹瓢虫(Propylaeajaponica)控制豌豆蚜(Acyrthosiphonpisum)的能力,在实验室内采用圆盘法测
定了不同龄期龟纹瓢虫对豌豆蚜的捕食功能反应,以期掌握龟纹瓢虫控制豌豆蚜的能力。结果表明:龟纹瓢虫4
个龄期的幼虫和雌雄成虫捕食豌豆蚜的数量与豌豆蚜密度呈负加速曲线关系,符合 Honing-Ⅱ型功能反应模型。
龟纹瓢虫雌成虫的日捕食量最大,其次是第4龄幼虫和雄虫。模型预测了龟纹瓢虫1,2,3,4龄幼虫及雄、雌成虫
对豌豆蚜的日最大捕食量分别是6.5,20.4,32.3,71.4,58.8,80.0头。综上所述,龟纹瓢虫对豌豆蚜有较强的控制
能力。
关键词:龟纹瓢虫;豌豆蚜;功能反应;捕食
中图分类号:S476.2 文献标识码:A 文章编号:1007-0435(2012)04-0778-06
Peaaphid,Acyrthosiphonpisum Harris(Ho-
moptera:Aphididae),isoneofthemostdevasta-
tinginsectpestsofleguminousforagealoverthe
worldincludingChina[1-3].Itprefersclusteringat
thedelicatefoliageandbuds,suckingjuiceandex-
cretinghoneydew,whichaffectthequalityand
quantityofherbageproduction.Moreseriously,
peaaphidisa vectorofalfalfa mosaic virus
(AMV),adiseasethatmakesblackstemdisease
oflucernemoresevereandcausemuchmoredam-
agethantheaphiditself[4].LadybirdbeetleProp-
ylaeajaponicaThunberg(Coleoptera:Coccineli-
dae)isanativeand widespreadaphidophagous
coccinelidpredatorinChina.Ithasbeenreported
thatPropylaeajaponica preyedonSitobionavenae[5],
Aphisgosypi[6],Rhopalosiphummaidis[7],Macrosiphum
收稿日期:2011-12-12;修回日期:2012-04-11
基金项目:国家科技支撑计划项目(2006BAD04A04-01);苜蓿抗蚜新品种选育(036030)资助
作者简介:武德功(1980-),男,河南周口人,博士研究生,研究方向为草地生物多样性,E-mail:wudegong118@163.com;*通信作者 Au-
thorforcorrespondenc,E-mail:Hechungui008@qq.com
第4期 武德功等:不同龄期龟纹瓢虫对豌豆蚜的捕食功能反应
sanborni[8],Myzuspersicae[9]. Moreover,the
predatoryfunctionalresponsesofP.japonicaa-
dultswhichwereinsatiationandhungerfor24h
onA.pisuminalfalfa(MedicagosativaL.)were
determined[10].Nevertheless,noanypaperhas
coveredalstagesofP.japonicapreyingonA.
pisuminalfalfainChina.Therefore,thepreyre-
sponsesandeffectsofalpredaceousstagesofthis
ladybirdonthedifferentdensitiesofA.pisum
wereinvestigatedhere.Thecapacityofcontroling
peaaphidswasanalyzedtoprovideareferencefor
theuseofpestbiologicalcontrolinalfalfa.
1 MaterialsandMethods
1.1 Insectcultures
AdultsofPropylaeajaponicaoriginalycol-
lectedfromthealfalfafieldsatGansuAgricultural
UniversityinMay,2009,werefedwithA.pisum
inalfalfa(HunterRiver)incages(60cmwide×
150cmlong×50cmhigh)underlaboratorycondi-
tions(25℃±2℃,50%±10%RH,andlight/dark
=14h/10h).Adultmalesandfemaleswerepro-
videdwithA.pisuminalfalfaplantsandkeptin
cohortformating.After24h,eggswerecolected
andtransferredintocleanPetridish(7cmdiame-
ter×2cmdeep),whichcontainingasmalalfalfa
branchwithtwofoliages.Inordertomaintainhu-
midity,thebottomendofthealfalfastem was
wrappedinawater-saturatedcottonbal.Then
theseeggswereconductedinartificialintelectual
climatechamber(RXZ-300C)underabioticcondi-
tion:25℃±1℃,70%±7% RH,light/dark=14
h/10hand11000lx.P.japonicalarvaewerealso
conductedinartificialintelectualclimatechamber
whenmoltedandwerefedwithA.pisuminalfal-
fa.
AcolonyofA.pisumwasalsooriginalycol-
lectedfromalfalfafieldsandmaintainedinalfalfa
(HunterRiver)cultivatedinsmal pots(15cm
high×10cmdiameter)at20℃±3℃,50%±10%
RHand16h/8h(light/dark).
Bothladybirdandaphidindividualswerekept
inagrowthchamber(3m×4m×5m).
1.2 Methods
P.japonicalarvae(10hpost-molting),adult
malesandfemaleswerecolectedinplasticjars(11
cmlong×7cmdiameter),adultswereseparated
andsexedaccordingtothesize(maleissmalerin
sizethanfemale).Predationwasassessedbypla-
cingasinglelarva/adultinanexperimentalarena
consistingofaplasticjarcontainingasmalalfalfa
branchwithtwofoliages.Inordertomaintainhu-
midity,thebottomendofthealfalfastem was
wrappedinawater-saturatedcottonbal.
Thepredationpotentialofladybirdlarvain-
creasedaslarvaproceedstothenextstage,prey
densitieswereincreasedforlaterinstarsandadult
stages.Therefore,fivepreydensitiesofA.pisum
wereevaluated:5,10,15,20,25aphidsforthe
firstinstar;10,15,20,25,30aphidsforthesec-
ondinstar;10,20,30,40,50aphidsforthethird
instar;20,30,40,50,60aphidsforthefourthin-
star,maleandfemale,respectively.Inaddition,
therewasaplasticjarwith30aphidsofA.pisum,
whichcontainedalstagesofpeaaphid,forestima-
tingthenumberofaphidsafter24hours.A.pi-
sum weregentlytransferredtoeachexperimental
arenausingafinehairbrushputtingthemonthe
alfalfabranchdescribedabove.Tenreplicateswere
madeforeachcombinationwithcontrols.Theex-
perimentswereconductedinartificialintelectual
climatechamber(RXZ-300C)underabioticcondi-
tion:25℃±1℃,70%±7% RH,light/dark=14
h/10h,and11000lx.After24h,aldeadandliv-
ingaphidswererecorded.
1.3 Dataanalysis
Therelationshipsbetweenmeanrateofcon-
sumption(preyconsumed/preyoffered×100)by
adultmales,femalesanddifferentlarvalinstarsre-
latedtopreydensitywereanalyzedusinggeneral
regressionequation(SPSS13.0forwindowsand
Excel2003).HolingcurvilineartypeⅡ equa-
tion[11]wasusedtofitthedataforfunctionalre-
sponse.Inthismodel,thenumberofpreycon-
sumed(Na)isafunctionofpreydensity(N)as
folows:
Na= (a’TtN)/(1+a’ThN),
Wherea’istheattack(discovery)rateofthe
prey,Ttisthetotaltimeavailable(1dor24hin
thisexperiment),andThisahandlingtimeforone
prey.
MakeTt=1,B=1/a’,A=Th,TransformNa
= (a’TtN)/(1+a’ThN)into1/Na=B(1/N)+
A,calculatevalueofa’andThbyusingtheleast
squaremethod.
977
草 地 学 报 第20卷
2 Results
Thepercentageofpreyconsumedbyeach
predatorystageisnegativelycorrelatedwiththe
preydensitiesoffered(Fig.1)withsubsequentr2
valuesbeinghighest(0.956)forsecondinstarand
lowest(0.759)foradultfemales.Therateofprey
consumptiondeclinedwiththeincreasingpreyden-
sityforalpredatorystages,suggestingthatthese
stagesexhibitatypeⅡfunctionalresponse(Fig.
2).ThetypeⅡfunctionalresponsesofalstages
ofPropylaeajaponicaarefolowed:
Firstinstar:Na=0.229N/(1+0.022579N);
Secondinstar:Na=0.512N/(1+0.025037N);
Thirdinstar:Na=0.656N/(1+0.020336N);
Fourthinstar:Na=0.870N/(1+0.01218N);
Adultmale:Na=0.568N/(1+0.009826N);
Adultfemale:Na=0.779N/(1+0.009738N).
The fourth instars showed the highest
(0.870)rateofattack(Table1).However,the
firstinstarsexhibitedthelowest(0.338)rateof
attack.Handlingtimedecreasedaspredatorylarva
proceededtothenextstage,andtheadultfemales
hadtheshortesthandlingtime(18min).Subse-
quently,themodelpredictsamaximumconsump-
tionof6.5,20.4,32.3,71.4,58.8,80.0A.pi-
sumaphidsbyfirst,second,thirdandfourthin-
stars,adultmalesandfemales,respectivelyin24
hours(Table1).
Fig.1 PercentpredationofP.japonicatoA.pisumonalfalfa
Note:PointsshowaveragenumberofaphidseatenorkiledbyP.japonicaateachlevelofpreyavailability.
Errorbarsshowstandarderrors
087
第4期 武德功等:不同龄期龟纹瓢虫对豌豆蚜的捕食功能反应
Fig.2 FunctionalresponseofP.japonicatoA.pisumonalfalfa
Note:PointsshowaveragenumberofaphidseatenorkiledbyP.japonicaateachlevelofpreyavailability.
Errorbarsshowstandarderrors.ThelineshowstheHoling’smodelforatypeⅡfunctionalresponse
Table1 TypeⅡfunctionalresponseparametersofPropylaeajaponicaatdifferentdensitiesofAcyrthosiphonpisum
Stage/instar n a’±SE Th±SE/min Tt/Th R
Firstinstar 10 0.338±0.015 0.1533±0.0279(220.75) 6.5 0.985**
Secondinstar 10 0.512±0.032 0.0489±0.0112(70.42) 20.4 0.970**
Thirdinstar 10 0.656±0.041 0.0310±0.0090(44.64) 32.3 0.959**
Fourthinstar 10 0.870±0.056 0.0140±0.0040(20.16) 71.4 0.961**
Male 10 0.568±0.022 0.0173±0.0029(24.91) 58.8 0.990**
Female 10 0.779±0.045 0.0125±0.0042(18.00) 80.0 0.962**
Note:Risthecorrelationcoefficientbetween1/Nand1/Na
3 DiscussionandConclusion
ThedataindicatesthataltestedstagesofP.
japonicaexhibitedtypicaltypeⅡ functionalre-
sponse(Fig.2).Similarreponsewereearlierre-
portedforotheraphidophagous,suchas,Sitobion
avenae[5],Aphisgossypii[6],Rhopalosiphum
maidis[7],Macrosiphumsanborni[8],Myzuspersi-
cae[9],orothercoccinelidssuchasHippodamia
(Adonia)variegate[12-16],Coccinellaseptempunc-
tata[17-18],Adoniavariegate[19],Hippodamiatre-
decimpunctata[13],etc.
TherelativerateofpreyconsumptionbyP.
japonicawashigheratlowdensities,whichindi-
catedthatthispredatorwouldbemoreeffectiveat
controlingtheA.pisumpopulationatlowdensi-
ties.ThefailurecausesofP.japonicaincontrol-
lingtheaphidathigherpopulationdensitiescould
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草 地 学 报 第20卷
be:(i)Honeydewexcretedbyaphidsaffectsthe
searchingbehaviorofthepredators[20];(i)More
importantly,ahighrateofpreyconsumptionisnot
afeature ofaphidophagouscoccinelids[21] that
leadstothefailureofaphidbiologicalcontrolpro-
gramsusingonlypredatorycoccinelids.There-
fore,forconsiderationofA.pisumbiocontrol,the
coccinelidshavetobereleasedearlybeforethea-
phidsreachhighdensity.
Thereleasestageofpredatorsinfluencesits
effectivenessandpotentialeconomicbenefitsbe-
causethesearchratesandhandlingtimesofpreda-
torsaredifferentindifferentdevelopmentalstages
[12,14].Therefore,thedataprovideinsightstoas-
sesswhichstagesforthereleaseofP.japonicaa-
gainstA.pisum.ThefirstinstarslarvaeofP.ja-
ponicabecomeeasilythepreyofotherladybirddue
toitssmalsizewhichshouldbecautioustobere-
leased.ThefourthinstarsandadultstagesofP.
japonicacouldbeefficienttosuppressA.pisum
densitybecauseofhigherattackratesandshorter
handlingtimes.Thethirdinstarslarvaemaybe
cost-effectivestagesbecauseofshortening mass
rearingperiods.
Theelevatedfunctionalresponsecurveofa-
dultfemaleoverthatoftheadultmaleindicatesthe
possibilityofdelayedsatiation,fasterdigestion
[22],morelargebodysizethanmales;hencegrea-
terfoodrequiredandhighernutritionaldemands,
particularlyforeggformation.
Innaturalconditions,ladybeetlewilpreyon
alstagesofaphidsencountered,ratherthanjust
higherinstarsnymphs.Ifonlyusinghigherinstars
nymphs,itwouldnotreflectthereality.Itisalso
difficultto distinguish between higherinstars
nymphsandadultaphids.Ifonlyadultaphidswere
used,itwouldreduceconsumptionofladybug.
Forreactionofnaturalenemypredationonaphids
inthefield,alstagesofA.pisum wereusedin
theexperiment.
Thestudyoffunctionalresponseinthelabora-
torycouldbeusedtoinferbasicmechanismsun-
derlyingnaturalenemy-preyinteraction[23].Such
studiesprovidevaluableinformationforbiological
controlprograms.Duringbiologicalcontrolevalu-
ationprocesses,comparisonsofparametervalues
oftwoormorepredatorsmaybemoremeaningful
andconvenientthansimilarcomparisonsinvolving
functionalresponsecurves[24].
However,therelationsbetweenfunctionalre-
sponsesobservedinthelaboratoryandfieldper-
formanceofnaturalenemiesisnotidentical,some
studiesshowedasignificantdifferencebetweenthe
responsesobservedinlaboratoryandfieldenviron-
ment[23].Itisrecognizedthatfunctionalresponses
derivedfromlaboratoryresearchmayhavelittlere-
semblancetothosethatcouldbemeasuredinthe
field.Forinstance,otherfactors,suchasintrinsic
growthrates,hostpatchiness,predationandcom-
petition,hosttraits,andenvironmentalcomplexi-
ties(abioticandbioticfactors)alsohaveamajor
influenceontheefficiencyofpredatorinmanaging
thepreypopulation.Thedataprovidedhereindi-
catehowthesepredatorswilrespondtoincreasing
preydensityundersimplifiedexperimentalcondi-
tions.Forconclusiveestimationsoftheirbiocon-
trolpotential,furtherfieldstudiesareneeded.
Recentyears,peaaphidpopulationissteadily
increasingandhasbeenbeingadominancepopula-
tioninalfalfafieldatLanzhou,Gansuprovince.
Thereforeitisneededtomakeproperprevention
andcontrolofpeaaphidinordertoavoidalarge
yieldloss.
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(责任编辑 李美娟)
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