Electrophysiological and behavioral responses of Tomicus minor (Coleoptera: Scolytidae) to host volatiles

2015·0.35 MB·

by  Yang Hua

©EntomologicaFennica.13May2015
Electrophysiological and behavioral responses
of Tomicus minor (Coleoptera: Scolytidae) to host volatiles
HuaYang+,Hong-WeiWang+,WeiYang*&Chun-PingYang
Yang,H.,Wang,H.-W.,Yang,W.&Yang,C.-P.2015:Electrophysiologicaland
behavioral responses of Tomicus minor (Coleoptera: Scolytidae) to host vola-
tiles.—Entomol.Fennica26:15–24.
Toscreenforhostvolatilesthateffectively attractthelesserpineshootbeetle,
Tomicusminor(Coleoptera:Scolytidae),animportantforestpest,watervapor
distillation extraction and gas chromatography-mass spectrometry were com-
binedtoanalyzevolatilesemittedfromuninfestedandinfestedshootsofPinus
yunnanensis(Pinaceae).ElectroantennographyandY-tubeolfactometerbehav-
ioraltestswereusedtocompareandanalyzetheresponsesofmaleandfemale
beetlestoeachofselectedeightcompounds.Thespectrometryresultsshowed
that monoterpenes are the primary chemicals emitted by P. yunnanensis. The
electroantennogramresponsesofadultT.minortosixoftheplantcompounds
peakedat10µg/µL;theexceptionswereterpinoleneandmyrcene,whichhad
maximaabove100µg/µL.(+)-(cid:1)-Pineneand(cid:2)-phellandrenewerestronglyat-
tractivetothemaleandfemaleadults.
H.Yang,H.-W.Wang,W.Yang&C.-P.Yang,KeyLaboratoryofEcologicalFor-
estry Engineering of Sichuan Province, College of Forestry, Sichuan Agricul-
tural University, Ya’an, Sichuan 625014, China; *Corresponding author’s e-
mail:[email protected].+Theseauthorscontributedequallytothisar-
ticle
Received14July2014,accepted9October2014
1.Introduction Kuzen.Likewise,Miaoetal.(2004)showedthat
Dendroctonus valens LeConte was attracted to
Volatileorganicchemicalsreleasedbyhostplants (+)-3-careneofitshostinChina.Furthermore,a
canattractinsectsandstimulatetheirfeedingand mixtureof(cid:1)-pineneandethanolwasstronglyat-
oviposition (Stanjek et al. 1997, Ruther et al. tractivetoT.piniperda(Byers1992).Thesestud-
2000, Du 2001, Inui et al. 2003, Inui & Itioka ies suggest that plant volatile chemicals can be
2007,Huetal.2009).Suchkairomonesinclude importantforattractionandhostlocationbyher-
monoterpenes, fatty acid derivatives, phenyl- bivorousinsectsand may beeffectively used to
propanoids,andisoprenoids,amongothers(Vis- monitor and control pest activities (Allison &
ser1986,Bernays&Graham1988,Bruceetal. Cardé2008,Williamsetal.2008,Ngumbietal.
2005,Zhugeetal.2010). 2009).Inmanycases,electroantennographyfol-
Zhang et al. (2007) reported that Ips lowedbyY-tubeolfactometerbehavioraltestsis
subelongatusMotschulskywaspreferentiallyat- usedtoscreenandselectcandidatevolatilechem-
tracted to (cid:1)-pinene, (cid:2)-pinene, and p-isopropyl- icalsthatelicitresponsesbyinsects(Han&Han
toluenereleasedbyitshostLarixgmelinii(Rupr.) 2007,Davidetal.2009).
16 Yangetal. (cid:127) ENTOMOL.FENNICAVol.26
Inthisstudy,compositionandamountsofvol- yunnanensis trees (age: 30 years, height: 18–20
atile chemicals were compared between un- m,diameteratbreastheight:20–25cm)into1–2
infestedandinfestedPinusyunnanensisFranchet metrelongpieces,andputthemintoseveralmesh
shoots, and electrophysiological and behavioral cages(2.5×2.5×2.0m)atroomtemperature(25
responsesofTomicusminorHartig,1834(Cole- °C± 2). Theemerging femaleand malebeetles
optera:Scolytidae)tosyntheticcompoundswere were collected and placed in individual cages
assessed.Thisworkmayleadtotheidentification (5cm×5cm×5cm,stainlesssteelmesh)atthe
ofattractantsthatcanbeusedtocontrolandmon- roomtemperature.Femaleandmalebeetleswere
itorT.minor. distinguished by the fact that male adults chirp
butfemaleadultsdonot(Ye1996).
2.Materialandmethods
2.2.Extractionandidentification
2.1.Lesserpineshootbeetle ofvolatilechemicalsemittedfromshoots
2.1.1.Basicbiology Branches from both infested and uninfested 2-
year-old P. yunnanensis were collected from
Thelesserpineshootbeetle,Tomicusminor,isan Yunnan pine plants in Luding County, Sichuan
oligophagous wood boring beetle that feeds on Province. Branches were then cut into 1 cm
morethan20hostspeciesofPinaceae,including pieces,and600gwereplacedintoawatervapor
Pinus yunnanensis, P. massoniana Lamb, P. ta- distillerfilledwith1200mLdeionizedwaterfor4
buliformisCarr,P.densifloraSieb.etZucc,Larix hoursasdescribedinWangetal.(2002).Thedis-
spp., and Picea spp. (Långström 1983, Ying & tilledliquidwasextractedwithanhydrousether
Huang 1984). Tomicus minor is widely distrib- (analytically pure; Chengdu Kelong Chemical,
utedinChina,Japan,Russia,andEurope(Lund- Chengdu,China)anddewateredwithanhydrous
gren 2004, Alonzo-Zarazaga 2013). The beetle sodium sulfate (analytically pure; Chengdu
undergoes one generation per year and over- Kelong Chemical). Then, the diethyl ether was
wintersasanadultinbark(Yeetal.2004,Xueet removedwithconcentratedN.Finally,2mLofa
2
al.2007).Adultsexittheirholesandfeedonfresh paleyellowtransparentgreasewithastrongaro-
shoots from April to May (Kirkendall et al. matic odor was obtained and stored in an ultra-
2008).InChina,P.yunnanensisisthepreferred coldfreezer(–80°C)(Sanyo,Osaka,Japan)and
hostforbothlarvaeandadults(Yeetal.2004). usedforchemicalidentification.
Aftereclosion,theadultsofT.minorflytothe The extracts were analyzed by gas chroma-
crownsofadjacentP.yunnanensistreesandfeed tography (GC) (Shimadzu, model 17A; Kyoto,
onshoots.Aftersexualmaturity,theymovetothe Japan)coupledwithaShimadzuQP5050Aelec-
trunkwheretheymateandoviposit.Thedamage tronionizationmassdetector(MS).TheGC-MS
tostemsmay causetreemortality by disrupting was operated in the splitless mode, and was
waterandnutrientflowbetweenthefoliageand equippedwithaDB-5(30m×0.25mm×25µm)
roots(Lietal.2006)andcanalsoresultinwith- capillary column (Agilent Technologies, Santa
eredandbrokenshoots(Yingetal.1984,Ye& Clara,CA,USA).Thecolumnoventemperature
Ding1999).Thus,controllingthevirginmaleand wasprogrammedtorisefromtheinitialtempera-
femaleadultsduringtheshoot-feedingperiodis tureof70°C(1min)to100°Cat10°Cmin–1,
critical to prevent T. minor outbreaks in P. thento250°Cat5°Cmin–1,atwhichthecolumn
yunnanensisstands. wasmaintainedfor2minutes.
Forthemassspectrometer,thevoltageofthe
2.1.2.Beetlecollectionandrearing electronimpactionsourcewas70eV,thetemper-
ature of the GC-MS connector was 250 °C, the
Lesser pine shoot beetles were collected from temperature of the ion source was 200 °C, the
YunnanpineplantsinfieldsnearLuding,Sichuan scanningspeedwas0.4s,thescanningrangewas
Province, China in May, 2013. We cut five P. mass-to-chargeratio(m/z)40–450,andtheelec-
ENTOMOL.FENNICAVol.26 (cid:127) ResponsesofTomicusminortohostvolatiles 17
triccurrentinthefilamentwas150µA.Themass vice.Whenthebaselinewasstable,theantenna
spectrogram was directly checked against the wasstimulated. Thestimulationtimewas0.5s,
standard mass spectrogram from the NIST (the andtheintervalbetweenstimuliwas30s,which
national institute of standards and technology) permittedrecoveryoftheantennalreceptors.For
database of this device to identify the chemical each compound, twelve antennae (from six dif-
component. ferentmalesandfemales)weretested,andeach
antennawasstimulatedfivetimes.Distilledhex-
ane was the standard, and the mean of the ob-
2.3.Syntheticcompounds servedfivevaluesforeachantennawasdivided
bythemeanofthetwostandardvaluestogivethe
BasedontheGC-MSanalysis,weselectedeight relativeantennalresponse(Yangetal.2013).
compoundsofthehighestrelativeabundanceto
betestedforEAGdose-response.Onlytheeight
compoundswereselectedfortheEAGdose-re- 2.5.Bioassayofbehavioralresponses
sponsetestsinordertomatchtothelimitedlabo-
ratoryconditions,expenditureetc.Thesynthetic Totestthebehavioralresponsesofadultbeetles,a
compounds of these eight chemicals were pur- glassY-tubeolfactometerwiththeinsidediame-
chasedfromTokyoChemicalIndustry Co.(To- terof0.5cmwasused.Themainarmofthede-
kyo, Japan): (+)-(cid:1)-pinene (95% pure) and (cid:2)- vicewas6cmlong,aswaseachsidearm.Thean-
caryophyllene (90% pure), and from Aladdin gle between the two side arms was 75 degrees,
Chemistry Co. (Shanghai, China): ((cid:2)-pinene andtheendsofthearmshadground-glassedges.
(98% pure),(+)-3-carene (90% pure), (+)-limo- Thetwosidearmswereconnectedtotwo250-mL
nene(97%pure),myrcene(90%pure),terpino- volumetricflasksbyTeflontubes.Amicro-sam-
lene(85%pure),and(cid:2)-phellandrene(99%pure). plerwasusedtoextract10µLofacompoundand
They were each dissolved in distilled hexane at solution(distilledhexane) for eachcomparison,
concentrationsof0.01,0.1,1,10,and100µg/µL and the liquid was dropped onto a filter paper
andusedfortheelectroantennogramanalysisbe- (1×1cm).
low. Filterpaperswereseparately placedintothe
twovolumetricflasks,whichwereconnectedby
Teflon tubes to a distilled water humidification
2.4.Electroantennograms bottleandacharcoalfilter.Theairflowspeedwas
controlledat0.5–0.6L/min.Thetestswererunat
Antennal recordings of the responses of female 08:00–12:00,whenthetemperatureofthelabora-
and male T. minor to the eight individual com- torywas25±2°C.
pounds were made using Syntech (Hilversum, AdultT.minorwasintroducedintotheinletof
The Netherlands) equipment comprising themainarmoftheY-tube,andtimingbeganaf-
micromanipulators,aCS-05stimulusaircontrol- tertheyhadmovedforward2cmfromtheinlet.
ler,andanIDACsignalconnectionboxfordata Adultshadtomakeachoiceatthejunctionofthe
acquisition. Electroantennogram (EAG) signals Y-tube. The tests were conducted for 5 min for
anddatawereanalyzedusingacustomizedsoft- eachadult.Ifabeetlewentforward2cmintoa
warepackage(EAGforWindowsXP,Syntech). sidearmandstayedforatleast1min,itwasre-
The antennae of T. minor were excised and cordedashavingmadeachoiceofthatodor;oth-
mountedbetweenAgandAgClglasselectrodes erwise,itwasrecordedtohavemadenochoice.
filledwithRingersolution. Aftereveryfiveadults,thetwoarmsoftheY-tube
A2-µLodorsourcesample(oneoftheeight wereinterchangedtoeliminatethepossibleinflu-
monoterpenoidcompounds)wastakenwithami- ence of individual arms on the insect behavior.
cro-sampler. It was uniformly dripped onto a After each treatment, the Y-tube olfactometer,
foldedfilterpaper(1.5×1.5cm)thatwasputinto Teflontubes,andvolumetricflaskswerewashed
a10-cmsampletube.Theendofthesampletube with ethanol and allowed to air dry. For each
wasconnectedtoanodor-stimulatingcontrolde- treatment, threereplicates of30 females and 30
18 Yangetal. (cid:127) ENTOMOL.FENNICAVol.26
Table1.Gaschromatography-massspectrometryresultsasrelativeabundances(%,mean±SE,N=3)of42
compoundsofuninfestedandinfestedshootsofPinusyunnanensis.Retentiontimesareshownbelowrelative
abundances.
Compound Uninfested Infested t
Tricyclene 0.28±0.01 0.06±0.01 10.02
3.72 3.72 P=0.001
(+)-(cid:1)-pinene 40.8±1.28 8.29±0.80 16.23
3.84 3.84 P0.001
Camphene 1.10±0.10 0.34±0.05 7.01
4.04 4.04 P=0.002
Sabinene 0.92±0.08 0.10±0.01 9.76
4.32 4.32 P=0.001
(cid:2)-pinene 10.48±0.70 12.39±0.94 –1.64
4.39 4.39 P=0.177
(cid:1)-phellandrene 0.12±0.02 0.11±0.01 0.08
4.76 4.76 P=0.448
(+)-3-carene 4.72±0.05 4.12±0.11 4.90
4.86 4.86 P=0.008
2,6-dimethylbenzenemethylether 0.16±0.01 – –
4.96 – –
(+)-limonene 6.05±0.11 5.09±0.21 4.09
5.15 5.15 P=0.015
(cid:2)-phellandrene 11.02±0.15 12.48±0.38 –3.61
5.17 5.17 P=0.023
Ocimene 0.22±0.01 0.14±0.01 9.80
5.40 5.40 P=0.001
(cid:3)-terpinene 0.28±0.02 0.16±0.03 3.52
5.63 5.63 P=0.024
Terpinolene 3.52±0.14 3.92±0.14 –2.03
6.15 6.15 P=0.112
Fenchol – 0.10±0.02 –
– 6.59 –
Camphor 0.22±0.02 0.33±0.04 –2.61
7.15 7.15 P=0.059
Camphol 0.24±0.04 0.21±0.01 0.78
7.49 7.49 P=0.478
4-terpenol 0.33±0.04 0.26±0.03 1.30
7.67 7.67 P=0.263
(cid:1)-terpilenol 0.60±0.04 2.42±0.18 –10.00
7.89 7.89 P=0.001
Verbenone 0.06±0.01 – –
8.26 – –
2-isopropyl-5-methylanisole 0.13±0.01 0.17±0.03 –1.12
8.63 8.63 P=0.325
Bornylacetate 1.78±0.19 1.02±0.09 3.59
9.65 9.65 P=0.023
(-)-(cid:1)-cubebene – 0.04±0.01 –
– 10.93 –
(cid:1)-pinene 0.10±0.01 0.36±0.05 –5.34
11.48 11.48 P=0.006
(cid:2)-elemene 0.29±0.07 0.52±0.05 –3.63
11.76 11.76 P=0.022
(cid:2)-caryophyllene 7.42±0.34 14.75±1.21 –5.85
12.32 12.32 P=0.004
Aromadendrene – 0.05±0.01 –
– 12.65 –
(E)-(cid:2)-farnesene 0.19±0.02 – –
12.82 – –
ENTOMOL.FENNICAVol.26 (cid:127) ResponsesofTomicusminortohostvolatiles 19
Table1continued
Compound Uninfested Infested t
(cid:2)-sesquiphellandrene – 0.34±0.08 –
– 12.83 –
(cid:1)-caryophyllene 1.21±0.17 2.68±0.12 –6.94
12.91 12.91 P=0.002
Myrcene 5.56±0.30 20.52±1.52 –9.67
13.38 13.38 P=0.001
(cid:1)-selinene – 0.32±0.02 –
– 13.48 –
Germacrene 0.12±0.02 0.46±0.06 –5.50
13.59 13.59 P=0.005
(cid:1)-muurolene – 2.35±0.27 –
– 13.66 –
(cid:4)-cadinene – 0.22±0.01 –
– 13.81 –
2,6-di-tert-butyl-4-methylphenol – 0.62±0.14 –
– 13.85 –
(cid:3)-cadinene 0.32±0.02 1.18±0.10 –8.27
13.94 13.94 P=0.001
(cid:4)-cadinene 1.24±0.11 – –
14.09 – –
(cid:1)-elemene – 0.15±0.02 –
– 14.14 –
(-)-isocaryophyllene – 0.30±0.05 –
– 15.29 –
t-cadinol – 1.48±0.19 –
– 16.43 –
t-muurolol – 1.95±0.18 –
– 16.69 –
a-cadinol 0.52±0.05 – –
16.70 – –
–= Compoundnotdetected.
malesofthesameeclosiontimewereperformed, ioral responses to different compounds, sepa-
andeachadultwasusedonlyonce. rately for females and males. After ANOVAs,
Duncan’s multiple range tests were used to test
thedifferencesintheattractionamongthecom-
2.6.Statisticalanalyses pounds from P. yunnanensis (Dong et al. 2000,
Liuetal.2005).Theratesofattraction,repellence
Statistical analyses were performed using the and indifference (no choice) were calculated as
SPSS18.0statisticalpackage(IBM,Chicago,IL, follows(Dingetal.1996,Yanetal.2006):
USA). Independent-sample t-tests and Bonfer-
ronicorrectionwereusedtocomparetherelative A=100×N /N (1)
t
abundances of individual compounds in un-
infested and infested plants as well as EAG re- whereAisattractionrate(%),N isthetotalnum-
t
sponses of female and male adults to the same ber of T. minor in the treatment arm of the
dose.Chi-squaredtestswereusedtotestwhether olfactometer,andNisthetotalnumberofT.mi-
femaleandmaleadultswereattractedtothese- nortested.
lectedeightcompoundsintheolfactometertrials
(Yang et al. 2013). A one-factor randomized R=100×N /N (2)
c
complete block analysis of variance (ANOVA)
wasconductedontheolfactometerdataofbehav- whereRisrepellencerate(%),N isthetotalnum-
c
20 Yangetal. (cid:127) ENTOMOL.FENNICAVol.26
Fig.1.Electrophysiologicaldose–responsecurvesofTomicusminorantennaetoeightselectedcombounds(CK
=distilledhexane,asastandardreference).Asterisksindicatesignificantdifferencesbetweenfemaleandmale
beetlesatthespecifieddose(independent-samplest-testswithBonferronicorrection,P<0.05,N=6malesand
6females).
ENTOMOL.FENNICAVol.26 (cid:127) ResponsesofTomicusminortohostvolatiles 21
ber of T. minor in the control arm of the 3.3.LaboratorybioassayofT.minor
olfactometer,andNisasinequation(1)above. adultresponsestosyntheticcompounds
Except for (+)-limonene, (cid:2)-carophyllene and
I=100–(A+R) (3)
myrcene, the remaining compounds tested had
significant attraction rates for female T. minor
whereIisrateofindifference(%),AandRasin
(Fig. 2a). Of those five compounds, (+)-(cid:1)-
equations(1)and(2).
pinene,(cid:2)-pineneand(cid:2)-phellandrenehadsignifi-
cantattractionalsotomaleadults(Fig.2b).
3.Results Amongthefivecompoundsthatattractedfe-
males, there were significant differences in the
rateofattraction(P=0.030,df =4,df =10,F=
3.1.Comparisonofchemicalcompositions
2.85). (+)-(cid:1)-pinene had a high1er attra2ction rate
oftissuesfromuninfestedandinfested
P.yunnanensis forfemalesthan(cid:2)-pineneandterpinolene,while
(+)-3-carene and (cid:2)-phellandrene did not differ
fromanyoftheothercompounds(Fig.2a).How-
FromuninfestedandinfestedP.yunnanensis,30
ever, among the three compounds that attracted
and 37 compounds were identified by GC-MS,
males,therewerenosignificantdifferencesinthe
respectively. Monoterpenes are the primary
rateofattraction(P=0.127,df =2,df =6,F=
chemicalsemittedbyP.yunnanensis.Thereinto,
1 2
1.71).
eightofthosecompoundshadthehighestrelative
abundancesinbothtypesofplants:(+)-(cid:1)-pinene,
(cid:2)-pinene, (+)-3-carene, (+)-limonene, terpino-
lene, (cid:2)-carophyllene, (cid:2)-phellandrene, and myr- 4.Discussion
cene.Theretentiontimesandrelativeproportion
oftotalpeakareasofallidentifiedcompoundsare Host volatiles identified in infested and
listedinTable1. uninfestedshootsofP.yunnanensiswerediffer-
ent.Feedingby T.minorinducednewchemical
compoundsinthetissues,suchasfenchol,(-)-(cid:1)-
3.2.EAGactivityofsyntheticcompounds cubebene, aromadendrene, (cid:2)-sesquiphellandre-
ne,(cid:1)-selinene,(cid:1)-muurolene,(cid:4)-cadinene,(cid:1)-ele-
Therewasaclearelectrophysiologicaldose–re- mene, (-)-isocaryophyllene, t-cadinol, t-muuro-
sponserelationshipforalleightcompounds.The lol, and 2,6-di-tert-butyl-4-methylphenol. Pro-
EAGresponsesofT.minoradultspeakedwhen duction of novel compounds due to herbivory
thecompoundconcentrationswere10µg/µL,ex- feeding is in parallel to the findings of earlier
ceptforterpinoleneandmyrcene,forwhichthe studiesthatthetypesandquantitiesofchemicals
responseswerestillrisingat100µg/µL(Fig.1). released by plants chewed by phytophagous in-
In most cases, female and male beetles sectsaredifferentfrom“constitutive”chemicals
showednodifferencesintheirEAGresponsesto (Mamiya&Enda1972)andareaffectedby the
thecompounds(Fig.1).TheEAGresponseoffe- feedingactivitiesofinsects(Lou&Cheng2000,
maleadultswassignificantlyhigherthanthoseof Degenhardt & Lincoln 2006). Paré and Tum-
maleadultsfor(+)-3-careneat1µg/µL(t=3.844, linson(1997)usedCO-impulselabelingtoprove
2
df=5,P=0.018),10µg/µL(t=3.733,df=5,P= that only insect feeding induced production of
0.020), and 100 µg/µL (t = 2.761, df = 5, P = newvolatileorganiccompoundsthatarenotre-
0.044),whiletheEAGresponseofmaleadultsto leasedbyundamagedplants.Likewise,Loughrin
myrcene was significantly higher than those of etal.(1995)showedthatcrabappleleavescanbe
femaleadultsat0.01µg/µL(t=-2.739,df=5,P= inducedbytheJapanesebeetletoreleaseagreat
0.045). dealofterpenoids.
Different host plant volatile compounds can
havevaryingimpactsoninsectactivities,because
thephysiologicalactivityofacompoundcanvary
22 Yangetal. (cid:127) ENTOMOL.FENNICAVol.26
Fig.2.Y-tubeolfactometerresponsesofTomicusminoradultsto10µg/µLofdifferentodors.
–a.Females.–b.Males.Meanpercentages(±SE)ofadultschoosingthetestcompound,con-
trol,orwithnochoicearereported(N=90).Totheleftofthebars,significantdifferencesbe-
tweentreatmentandcontrolaredenoted;Chi-squaredtests:**(P<0.01),*(P<0.05),NS
(P(cid:5)0.05).Forthecompoundsshowingsignificantattraction,differencesintherateofattraction
amongthecompoundsareshowntotherightofthebars;compoundswithdifferentlettersdiffer
significantly(P<0.05,Duncan’smultiplecomparisontest).Beetleswereconsideredtomakeno
choicewhentheyspentthewholetestperiodinmainarmoftheolfactometerorvisitedeither
testarmforlessthan1min.
withconcentration(Fanetal.2004).Thereisan suggeststhattheantennaeoffemaleandmaleT.
optimum dose range for the insect response to minoradultsmightdifferinthetypeandnumber
eachvolatilecompound.Withinthisrange,there- ofreceptorsorinphysiologicalfunctioning(Lu
sponse rate rises as the dose increases. Beyond 2008).
this range, insect attraction declines as the dose Westudiedtheantennalresponseforonlythe
changes(Fanetal.2003).Wefoundthatmaleand eightcompoundswiththehighestabundancesin
femaleT.minorresponsesreachedmaximumat theGC-MSanalyses.However,therewerecon-
10 µg/µL for (+)-(cid:1)-pinene, (cid:2)-pinene, (+)-3- siderable differences between infested and
carene, (+)-limonene, (cid:2)-carophyllene, and (cid:2)- uninfestedplantsalsoinsomelessabundantcom-
phellandrene,whileformyrcenethey peakedat pounds (Table 1). Insects can be responsive to
100µg/µL.Interestingly,insomecases,theEAG very low concentrations of some compounds,
responsesofmaleandfemaleantennaeweredif- meaningthatitisworthtoincludelow-concentra-
ferent,e.g.forterpinolene,theEAGresponsesof tioncomboundsinfuturestudiesofantennalre-
femalesweremaximumat10µg/µL,whilethose sponsesandattractiontoinsects.
ofmaleswerehighestat100µg/µL.Thisstrongly Plant chemicals are of vital importance in
ENTOMOL.FENNICAVol.26 (cid:127) ResponsesofTomicusminortohostvolatiles 23
host-plantselectionbybarkbeetles.Inthisstudy, cationbyGC-EADofthetwo-componenttrail-follo-
bothsexesshowedbasicallysimilarresponsesto wingpheromoneofProrhinotermessimplex(Isoptera,
Rhinotermitidae, Prorhinotermitinae). — Journal of
thetestedcompoundsintheselectiontestsinaY
InsectPhysiology55:751–757.
tube,inspiteofthefactthattheEAGresponsesof
Degenhardt,D.C.&Lincoln,D.E.2006:Volatileemis-
femaleandmaleantennaeweredifferentforsome sionsfromanodorousplantinresponsetoherbivory
compounds. (+)-(cid:1)-pinene, (cid:2)-pinene, and (cid:2)- andmethyljasmonateexposure.—JournalofChemi-
phellandrenewereattractivetobothmaleandfe- calEcology32:725–743.
Ding,H.J.,Guo,Y.Y.&Wu,C.H.1996:Designand
malebeetles,while(+)-3-careneandterpinolene
applicationoffourarmsolfactometrywiththestudyof
wereattractiveonlytofemalebeetles.Somebark
olfactionbehaviorofinsect.—EntomologicalKnow-
beetlespecieshavebeenpreviouslyshowntobe ledge33:241–243.[InChinese.]
attractedtohostplantvolatiles,including(+)-(cid:1)- Dong,W.X.,Wang,R.&Zhang,Z.N.2000:Electroanten-
pinene,(+)-3-carene,and(cid:2)-pinene(Hobsonetal. nalresponsesofaparasitoid(Microplitismediator)to
cottonplantvolatiles.—ActaEntomologicaSinica43
1993).Likewise,Miaoetal.(2004)reportedthat
(Supplement):119–125.[InChinese.]
(+)-(cid:1)-pinene, (+)-3-carene, and (cid:2)-pinene were
Du,J.W.2001:Plant–insectchemicalcommunicationand
repellenttoDendroctonusvalenswhenreleased itsbehaviorcontrol.—ActaPhytophysiologicaSinica
at150mg/day.Theseresultsmayimplynomajor 27:193–200.[InChinese.]
differencesinolfactionbetweenmaleandfemale Fan,H.,Li,J.Q.&Jin,Y.J.2003:Electrophysiological
and behavioral responses of adult Anoplophora
beetles,butfurtherstudiesarewarranted.How-
glabripennis(Motschulsky)tovolatilecomponentsof
ever, our electrophysiological and behavioral
host–plant.—ForestryStudiesinChina5:25–29.[In
dataindicatethatfuturefieldstudiesmayyieldre- Chinese.]
sultsthatareusefulforthedevelopmentoftraps Fan,H.,Jin,Y.J.,Li,J.Q.&Chen,H.J.2004:Advances
tomonitorelesserpineshootbeetlepopulations. onplantvolatilesemiochemicalsattractingherbivo-
rousinsects.—JournalofBeijingForestryUniversity
26:76–81.[InChinese.]
Acknowledgements.WeacknowledgethestaffoftheFor-
Han,B.Y.&Han,B.H.2007:EAGandbehavioralre-
estryBureauofLuding,SichuanProvince,China,fortheir
sponsesofthewinglessteaaphidToxopteraaurantii
kindhelpwithourfieldinvestigation.Wealsothankthe
(Homoptera:Aphididae)toteaplantvolatiles.—Acta
anonymous reviewers for valuable comments on the
EcologicaSinica27:4485–4490.[InChinese.]
manuscript.ThisworkwasfundedbytheFinancingPro-
Hobson,K.R.,Wood,D.L.,Cool,L.G.,White,P.R.,
jectoftheConstructionofEcologicalForestryEngineer-
Ohtsuka,T.,Kubo,I.&Zavarin,E.1993:Chiralspeci-
ingfortheUpstreamYangtzeRiver,China.
ficityinresponsesbythebarkbeetleDendroctonus
valens to host kairomones. — Journal of Chemical
Ecology19:1837–1846.
References Hu,J.,Angeli,S.,Schuetz,S.,Luo,Y.&HajkeA.E.2009:
Ecology and management of exotic and endemic
AsianlonghornedbeetleAnoplophoraglabripennis.
Allison,J.D.&Cardé,R.T.2008:Malepheromoneblend
—AgriculturalandForestEntomology11:359–374.
preferencefunctionmeasuredinchoiceandno-choice
Inui,Y.,Miyamoto,Y.&Ohgushi,T.2003:Comparisonof
windtunneltrialswithalmondmoths,Cadracautella.
volatileleafcompoundsandherbivorousinsectcom-
—AnimalBehaviour75:259–266.
munitiesonthreewillowspecies.—PopulationEco-
Alonzo-Zarazaga,M.A.2013:FaunaEuropaea:Tomicus logy45:41–46.
minor. Fauna Europaea version 2.6. URL http:// Inui, Y. &Itioka, T. 2007:Species-specificleafvolatile
www.faunaeur.org.(Sitevisitedon14October,2014) compounds of obligate Macaranga myrmecophytes
Bernays,E.A.&Graham,M.1988:Ontheevolutionof andhost-specificaggressivenessofsymbioticCrema-
hostspecificityinphytophagousarthropods.—Eco- togaster ants. — Journal of Chemical Ecology 33:
logy69:886–892. 2054–2063.
Bruce,T.J.A.,Wadhams,L.J.&WoodcockC.M.2005: Kirkendall,L.R.,Faccoli,M.&Ye,H.2008:Description
Insecthostlocation:avolatilesituation.—Trendsin oftheYunnanshootborer,TomicusyunnanensisKir-
PlantScience10:269–274. kendall&Faccolisp.n.(Curculionidae,Scolytinae),
Byers,J.A.1992:Attractionofbarkbeetles,Tomicuspini- anunusuallyaggressivepineshootbeetlefromsout-
perda,HylurgopspalliatusandTrypodendrondomes- hernChina,withakeytothespeciesofTomicus.—
ticum and other insects to short-chain alcohols and Zootaxa1819:25–39.
monoterpenes. — Journalof ChemicalEcology 18: Långström,B.1983:Lifecyclesandshoot-feedingofthe
2385–2402. pineshootbeetles.—StudiaForestaliaSuecica163:
David,S.D.,Blanka,K.,Pavel,J.,Anna,B.,Josef,C.,Ro- 1–29.
bertH.,Jan,S.,Christian,B.&Irena,V.2009:Identifi- Li,L.S.,Liu,H.P.,Chen,P.,Li,H.R.,Långström,B.&
24 Yangetal. (cid:127) ENTOMOL.FENNICAVol.26
Francoies,L.2006:Comparisonstudyonecological Wang,Y.Y.,Dong,G.&Liu,W.B.2002:Improvementof
characteristicsofTomicuspiniperdaandT.minor.— distillationtestequipmentformakingwatervapor.—
JournalofWestChinaForestryScience35:1–5.[In JilinForestryScienceandTechnology31:51–52.[In
Chinese.] Chinese.]
Liu,Y.,Guo,G.X.,Chen,J.L.&Ni,H.X.2005:Behavio- Williams,L.,Rodriguez-Saona,C.,Castle,S.C.&Zhu,S.
ralandelectrophysiologicalresponsesoffourpredato- 2008: EAG-active herbivore-induced plant volatiles
ryinsectspeciestosemiochemicalsofwheat.—Acta modifybehavioralresponsesandhostattackbyanegg
EntomologicaSinica48:161–165.[InChinese.] parasitoid.—JournalofChemicalEcology34:1190–
Lou,Y.G.&Cheng,J.A.2000:Herbivoreinducedplant 1201.
volatiles:Primarycharacteristics,ecologicalfunctions Xue,Y.G.,Ding,Q.H.,Hua,Z.J.,Xia,W.J.&Ga,Z.J.M.
anditsreleasemechanism.—ActaEntomolgicaSini- 2007:PreliminarystudyonlifehistoryandharmofTo-
ca20:1097–1106.[InChinese.] micusminor.—AnhuiAgriculturalScienceBulletin
Loughrin, J. H., Potter, D. A. &Hamilton-Kemp, T. R. 13:154–155.[InChinese.]
1995:Volatilecompoundsinducedbyherbivoryactas Visser,J.H.1986:Hostodorperceptioninphytophagous
aggregationkairomonesfortheJapanesebeetle(Po- insects.—AnnualReviewofEntomology31:121–
pillia japonica Newman). — Journal of Chemical 144.
Ecology21:1457–1467.
Yan,S.C.,Cheng,H.,Yang,H.,Yuan,H.E.,Zhang,J.&
Lu,R.C.2008:StudyontheecologyofTomicusyunna-
Chi,D.F.2006:EffectsofplantvolatilesontheEAG
nensis and Tomicus minor (Coleoptera: Scolytidae)
responseandbehaviorofthegreytigerlongicorn,Xy-
andtheirkairomones.—BeijingForestryUniversity,
lotrechusrustius(L.)(Coleoptera:Cerambycidae).—
Beijing.16pp.[InChinese.]
ActaEntomologicaSinica49:759–767.[InChinese.]
Lundgren,J.G.2004:ExoticForestPestInformationSys-
Yang,H.,Yang,W.,Yang,C.P.,Zhu,T.H.,Huang,Q.,
temforNorthAmerica:Tomicusminor.NorthAmeri-
Han,S.&Xiao,J.J.2013:Electrophysiologicaland
canForestCommission.URLhttp://spfnic.fs.fed.us/
behavioralresponsesofthewhite-stripedlonghorned
exfor/data/pestreports.cfm?pestidval=115&langdis.
beetle, Batocera lineolata, to the diurnal rhythm of
(Sitevisitedon14October,2014)
host plant volatiles of holly, Viburnum awabuki. —
Mamiya,Y.&Enda,N.1972:TransmissionofBursaphe-
JournalofInsectScience85:1–11.
lenchuslignicolus(Nematoda:Aphelenchoididae)by
Ye,H.1996:StudiesonthebiologyofTomicuspiniperda
Monochamusalternatus(Coleoptera:Cerambycidae).
(Col.,Scolytidae)intheshoot-feedingperiod.—Acta
—Nematologica18:159–162.
EntomolgicaSinica39:58–62.[InChinese.]
Miao,Z.W.,Zhang,Z.N.,Wang,P.X.,Guo,Y.Y.&Sun,
Ye,H.&Ding,X.S.1999:ImpactsofTomicusminoron
J.H.2004:Responseoftheredturpentinebeetle,Den-
distribution and reproduction of Tomicus piniperda
droctonusvalensLeConte(Coleoptera:Scolytidae)to
(Col.,Scolytidae)onthetrunkofthelivingPinusyun-
hostsemiochemicalsanditsimplicationinmanage-
nanensis trees. — Journal of Applied Entomology
ment.—ActaEntomologicaSinica47:360–364.[In
123:330–333.
Chinese.]
Ye,H.,Lu,J.&Francois,L.2004:OnthebionomicsofTo-
Ngumbi,E.,Chen,L.&Fadamiro,H.Y.2009:Comparati-
micusminor(Hartig)(Coleoptera:Scolytidae)inYun-
veGC-EADresponsesofaspecialist(Microplitiscro-
nanProvince.—ActaEntomologicaSinica47:223–
ceipes)andageneralist(Cotesiamarginiventris)para-
228.[InChinese.]
sitoid to cotton volatiles induced by two caterpillar
Ying,H.F.&Huang,F.S.1984:Economicinsectfaunaof
species.—JournalofChemicalEcology35:1009–
China.Fascicle23,Coleoptera:Scolytidae.—Scien-
1020.
cePress,Beijing.53pp.[InChinese.]
Paré,P.W.&Tumlinson,J.H.1997:Inducedsynthesisof
plantvolatiles.—Nature385:30–31. Zhang,Q.H.,Schlyter,F.,Chen,G.F.&Wang,Y.2007:
Ruther,J.,Reinecke,A.,Thiemann,K.,Tolasch,T.,Fran- ElectrophysiologicalandbehavioralresponsesofIps
cke,W.&Hilker,M.2000:Matefindingintheforest subelongatustosemiochemicalsfromitshosts,non-
cockchafer, Melolontha hippocastani, mediated by hosts,andconspecificsinChina.—JournalofChemi-
volatiles from plants and females. — Physiological calEcology33:39–404.
Entomology25:172–179. Zhuge,P.P.,Luo,S.L.,Wang,M.Q.&Zhang,G.2010:
Stanjek,V.,Herhaus,C.,Ritgen,U.,Boland,W.&Städler, ElectrophysiologicalresponsesofBatocerahorsfiedi
E. 1997: Changes in the leaf surface chemistry of (Hope)adultstoplantvolatiles.—JournalofApplied
Apiumgraveolens(Apiaceae)stimulatedbyjasmonic Entomology134:600–607.
acidandperceivedbyaspecialistinsect.—Helvetica
ChimicaActa80:1408–1420.