DBPIA-NURIMEDIA

농약과학회지 (Korean J. Pestic. Sci.) Vol. 21, No. 2, pp. 198-213 (2017) Open Access https://doi.org/10.7585/kjps.2017.21.2.198 Online ISSN 2287-2051 Print ISSN 1226-6183 ORIGINAL ARTICLES EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 민지현 윤헌 권혜리 유용만 윤영남 * 충남대학교농업생명과학대학응용생물학과 Feeding Behavior of the Green Peach Aphid, Myzus persicae, against Several Using Electrical Penetration Graph Techniques Ji-Hyun Min, Heon Yoon, Hay-Ri Kwon, Yong-Man Yu and Young-Nam Youn* Department of Agricultural Biology, Graduate School, Chungnam National University, Daejeon, 34134, Korea (Received on May 26, 2017. Revised on June 28, 2017. Accepted on June 28, 2017) Abstract Change of feeding behavior of three different populations (Iwol, Noeun and Lab population) of the green peach aphid, Myzus persicae, were recorded and analyzed with an electrical penetration graph (EPG) against 10 insecticides. Generally, green peach aphids do tend to feed at phloem rather than xylem in pepper plant with untreated insecticides using by EPG technique. It is meaned that the green peach is phloem feeder like other aphids. Treatments of insecticides changed the feeding behavior pattern and also different feeding pattern between registered and unregistered insecticides to the green peach aphid using by EPG. In case of acetamiprid treatment, non-probing period were more increased 64, 29 and 3.2 times at standard treatment than half-concentration treatment with Iwol, Noeun and Lab population, respectively. Total period of probing behavior with standard treatment is longer than half-concentration treatment at most of treated insecticide except in the case of cyantraniliprol treatment. The total period of saliva secretion of the green peach in the phloem sheath was significantly different at half-concentration treatment of cyantraniliprol as Noeun > Lab > Iwol population. The total of phloem feeding time taken at treatment plot is significantly shorter than non-treatment. At the cyantraniliprol half-treatment, Iwol population is shorter than any other population. However, there no significantly different at xylem feeding time taken among 3 populations. Key words EPG, feeding behavior, green peach aphid, insecticide, Myzus persicae 서 론 복숭아혹진딧물 (Myzus persicae Sulzer) 은노린재목 (Hemiptera) 진딧물과 (Aphididae) 에속하며, 전세계적으로많은작물에경제적으로많은피해를주는진딧물이다 (Van Emden and Harrington, 2007). 복숭아혹진딧물의피해상은크게두가지로나뉘는데, 식물의관다발조직을섭식하여수분부족및잎말림과생장저해와같은직접적인피해를주는경우와, 여러종류의식물바이러스의매개체로서작물에바이러스병을전파함으로서간접적피해를주게된다 *Corresponding author E-mail: youngnam@cnu.ac.kr (Van Emden and Harrington, 2007). 또한감로를배설하여그을음병을유발시켜작물의광합성을저해하기도한다 (Frants et al., 2004). 흡즙형곤충의탐침및섭식행동은직접적으로관찰할수는없지만, Electrical electrical penetration graphs (EPGs) 기술을통해관찰할수있다. 이방법은 McLean and Kinsey (1964) 에의해처음으로사용되었다. EPG 기술은흡즙형구기를갖는곤충의구침이식물체에삽입되면구침의이동에따른전기적저항의차이로전압이변화되면이를증폭시킨변화신호를기록한것이며, 처음개발한 McLean과 Kinsey (1964) 는교류방식 (AC system) 을사용하였고, 이후 Tjallingii (1978) 에의해직류방식 (DC system) 으로발전되어사용되고있다. 이러한기술은대부분진딧물 198

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 199 류 (Mclean and Kinsey, 1964, 1965, 1967; Tjallingii, 1978, 1985a, b, 1988), 멸구류 (Khan and Saxena, 1984; Velusamy and Heinrichs, 1986; Kimmins, 1989; Youn and Chang, 1993; Kim et al., 2005, Seo et al., 2009a), 매미충류 (Chang, 1978; Kawabe and Mclean, 1978; Khan and Saxena, 1985; Youn, 1998), 가루이류 (Jassen et al., 1989; Lei et al., 1996; Seo et al., 2007, Yang et al., 2009, Kwon and Youn, 2014), 나무이류 (Bonani et al., 2010; Civolani et al., 2011; Youn et al., 2011) 등흡즙형곤충의섭식행동을관찰하는데이용되었다. EPG 기술은곤충의섭식행동및식물의상호작용을반영하는것으로첫째, 흡즙성곤충에대한기주식물의저항성측정과특징을정의할수있고 (Helden and Tjallingii, 1993; Alvarez et al., 2013); 둘째, 바이러스매개체로써식물병의전이메커니즘을이해할수있으며 (Nault and Gyrisco, 1966; Powell, 1991; Fereres and Moreno, 2009), 마지막으로, 한정된범위내의살충제의작용기작을이해할수있을것이다 (Harrewijn and Kayser, 1997; Costa et al., 2011). 본연구는복숭아혹진딧물세개체군내에서의 EPG 패턴을통해섭식행동을분석하고, 작용기작이다른총 10 가지의약제에대한각개체군별섭식행동반응을비교분석하였다. 재료및방법 공시충및기주 2013-14년충북지역에서재배하는오이에발생하는질병조사결과, 진천지역일대에서진딧물에의해전염되는 CMV, WMV2, ZYMV 등의바이러스병이만연된것이관찰되어 (Lee et al., 2015). 이를토대로충북진천군이월면소재의배추밭에서복숭아혹진딧물을 2016년에채집하여 이월개체군으로사용하였다. 또한매년진딧물방제를위해살충제를살포하고있는대전유성구노은동소재배추밭에서채집한개체군은노은개체군으로, 실내사육실에서장기간누대사육한개체군을실험실개체군으로하여 3가지종류의복숭아혹진딧물개체군을실험에사용하였다. 이들각개체군은실험실에서누대사육하면서실험에사용하였으며, 고추 ( 품종 : 금빛 ) 를포트 (φ6.5 6.5 cm) 에파종하여 5-6 주가경과된유묘에접종하여아크릴케이지 (30x30x50cm) 에서계대사육하였고, 온도 25 ± 1 o C, 상대습도 50-60%, 광주기 16L:8D의곤충사육실조건하에서유지하였다. Electrical Penetration Graph (EPG) EPG 기록을할때에사용한고추는포트에파종한후약 4주가경과된고추유묘를사용하였다. 고춧잎은 Garzo (2016) 의방법을참고하여신초에서 3-4번째잎위에복숭아혹진딧물을올려두었다. 복숭아혹진딧물은사육케이지에서작은붓을이용하여무시성충만을골라, Ullman and Mclean (1988) 과 Jiang et al. (1999) 의방법을참고하여약 30분을굶긴후기록을실시하였다. 한편, 진딧물은지름이 12.7 μm인금선 (Alfa Aesar, US) 을약 5cm 길이로자른후약 3cm의구리못에감은구리선끝에 xylene을용매로 silver conductive paint (CANS, ELCOAT P-100, KOREA) 를녹여금선의약 1cm 정도부착시키고, 금선의다른한쪽끝에 silver conductive paint 용액을연속적으로찍어발라충체에부착할크기의방울을만들어준비해놓은것을복숭아혹진딧물의등판에부착하였다. 복숭아혹진딧물에금선을부착한후에다른끝에달린구리못을 GIGA-8 DC EPG amplifier (Wageningen University, Netherland) 에연결하였다 (Fig. 1). 기록하는동안발생하는전기적잡음을제거하기위해 Faraday cage안에서실시하였으며, 그안에접지된구리망을둘러씌워전기적잡음을완전히제거하였 Fig. 1. The primary circuit for EPG recording and ground net covered with copper.

200 민지현 윤헌 권혜리 유용만 윤영남 Table 1. Characterization of treated insecticide in experiments Category Insecticide Chemical group Registered for M. persicae Unregistered for M. persicae Formulation, A.I. (%) Mode of action Systemic action neonicotinoids WP, 8% Nicotinic acetylcholine receptor (nachr) competitive modulators O diamides DC, 5% Ryanodine receptor modulators X pyridine azomethine WP, 25% Chordotonal Organ TRPV Channel Modulators pyridine azomethine SC, 6.5% Chordotonal Organ TRPV Channel Modulators X tetramic acid SC, 22% Inhibitors of acetyl CoA carboxylase O pyrazole EC, 5% Uncouplers of oxidative phosphorylation via disruption of the proton gradient neonicotinoids SG, 50% Nicotinic acetylcholine receptor (nachr) competitive modulators O Gamma-cyhalothrin pyrethroids CS, 1.4% Sodium channel modulators X benzoylureas SC, 10% spinosyns WG, 10% Inhibitors of chitin biosynthesis, type 0 (IGR) Nicotinic acetylcholine receptor (nachr) allosteric modulators O X X X 다. 이러한 EPG 시스템에서복숭아혹진딧물의섭식행동은총 6시간동안기록하였다. EPG waveform은 EPG data acquisition (daq) (W.F. Tjallingii, Wageningen University, The Netherlands) 을통하여 IBM personal computer에저장되었으며, 저장된자료는 EPG signal analysis (ana) (W.F. Tjallingii, Wageningen University, The Netherlands) 로분석하였다. 그리고 EPG 매개변수는 Sarria et al. (2009) 에의해개발된 EPG-Exel Data Workbook을이용하여파형별소요시간및횟수를파악하였다. EPG 파형을 EPG signal analysis로분석할때에는 Tjallingii (1978) 와 Prado와 Tjallingii (2007) 에서정의한파형을참고하여 8가지매개변수를기준으로결과를분석하였는데, 각매개변수의특징은다음과같다. 1 어떠한탐침행동도하지않은시간 (total duration of non-probe time); 2 총탐침소요시간 (total duration of pathway phase); 3 체관부섭식시간 (total duration of feeding phloem); 4 물관부섭식시간 (total duration of feeding xylem); 5 섭식을하다가구침을빼고다음섭식때까지소요된시간 (total duration of derailed stylet); 6 6시간동안구침을식물체에찔러넣은횟수 (total number of probes); 7 EPG 셋팅후맨처음구침을찔러넣을때까지소요된시간 (time to the first probe); 8 EPG 셋팅후맨처음구침을찔러체관부흡즙을하기까지소요되는시간 (time to first feeding phloem) 등이다. 또한, 복숭아혹진딧물의섭식행동을관찰한 EPG 기록은복숭아혹진딧물이 EPG 시스템에서이탈하거나혹은죽은경우를제외하고정상적으로 6시간동안고추잎에서흡즙을하는경우만모아 3반복을실시하였으며, 이러한기록을모아통계분석하였다. 시험약제약제는단일유효성분으로이루어져있는약제를선택하였으며, 2016 작물보호제지침서 ( 발행처 : 한국작물보호협회 ) 상에복숭아혹진딧물이적용병해충으로등록되어있는 5 가지약제와등록되어있지않은 5가지약제를선택하였다. 등록약제선발기준은작용기작및계통이다른성분중에서새로운작용기작을갖거나, 저항성진딧물방제에효과를갖는약제를임의로선발하였으며, 비등록약제선발기준은작용기작및계통이다른성분중에서흡즙성해충 ( 총채벌레류, 노린재류, 가루이류등 ) 을적용해충으로갖는약제를임의로선발하였다. 각약제의처리는추천농도와추천농도의 1/2배로나누어처리하였었다 (Table 1). 약제처리약제에대한반응실험은개체군별복숭아혹진딧물의섭식행동에어떠한영향을미치는지알아보기위해진행하였다. 고추유묘에실험농도의각약제를 hand spray (Furupla Co. LTD, Japan) 에담고, 일정거리에서포트당약 5번의펌프를통해기주에약제를충분히노출한후음지에서약 30 분간건조하여 EPG 기록에이용하였으며, 3반복수행후분석하였다. 통계분석모든약제를각농도별로처리한처리구와무처리구는총 6시간동안 3회반복실험을수행하였고, EPG 기록으로분석한결과는 SPSS (IBM SPSS Statistics 22.0) 일원배치분산분석 (one way analysis of variance, ANOVA) 을통해각기록결과간 P<0.05 범위에서유의성검정을수행하였으

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 201 Table 2. Summary of aphids waveforms EPG waveform Tissue location of stylet tips Activities NP Non probing surface Walking and external salivation Pathway phase (C) Epidermis/mesophyll/ all tissue Cuticle penetration/sheath salivation/many activities during pathway Potential drop (pd) All living cells Stylet puncture; salivation, ingestion E1 Sieve elements Phloem salivation E2 Sieve elements Phloem ingestion G Xylem Xylem active ingestion F All tissues Derailed stylet Æpenetration difficulties Ø Fig. 2. Characteristics of EPG waveforms recorded from Myzus persicae.

202 민지현 윤헌 권혜리 유용만 윤영남 며, 사후검정은 Duncan 분석을통해약제간의유의차를확인하였다. 결과 복숭아혹진딧물의약제별섭식행동비교복숭아혹진딧물세개체군의무처리구에서 6시간동안섭식하는데소요한시간의양상은탐침하지않는 (NP) 시간보다는탐침하는 (C) 시간이더많았고, 물관부섭식 (G) 보다는체관부를섭식하는 (E1, E2) 시간이더많은것으로확인하였다. 따라서약제별섭식행동을비교할때, 위의양상과달리나타날경우섭식행동의저해를받은것으로판단하였다. 복숭아혹진딧물의각개체군별섭식행동을비교하기에앞서, 실험에서얻은복숭아혹진딧물 EPG 파형의종류와파형에따른섭식행동의의미는 Table 2와같다. 또한 Fig. 2는총 6 시간동안복숭아혹진딧물이섭식할때나타나는전형적인패턴을나타낸것으로, 좀더자세한각파형은 Fig. 2와같다. 어떠한탐침행동도하지않은시간탐침하지않은총기간은이월개체군의경우, 약제를기준량으로처리했을때, 유의한차이를보였다 (P=0.037). 등록된약제 5가지를기준량으로처리했을때, 기준량의 1/2 배처리구보다 4-65배정도비탐침기간이증가하였고, 등록되지않은약제 5가지를기준량으로처리했을때, 기준량의 1/2배처리구보다 2-11배정도감소하였다 (Table 3). 또한등록된약제 5가지를기준량의 1/2배처리한결과무처리보다더적은비탐침기간을가졌으며, 대조적으로, 같은조건에서등록되지않은약제 5가지에서는무처리보다더많은비탐침기간을가졌다. 노은개체군의경우약제를기준량의 1/2배처리한결과에서매우유의한차이를보였는데 (P=0.002), pymetrozine과 pyrifluquinazon을기준량의 1/ 2배처리한결과, 무처리보다약 6배더많은비탐침기간을나타냈으며, 다른약제들에비해가장많은비탐침기간을가졌다. 실험실개체군의경우, 모든약제를기준량의 1/2배처리한결과무처리보다더많은비탐침기간을가졌다. Table 3. Average running seconds depending on total duration of non-probe (NP) time using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration of 10 different insecticides for 6 hours Gamma-cyhalothrin 1/2 124.1 ± 7.7a a) 225.3 ± 235.7a 901.6 ± 794.1a 0.343 1 7,996.4 ± 8,027.3ab 6,640.0 ± 3,212.1ab 2,971.5 ± 79.1a 0.277 1/2 408.6 ± 403.5a 1,528.7 ± 744.2a 925.9 ± 243.6a 0.236 1 1,698.3 ± 1,744.3a 4,242.5 ± 4,920.0a 5,127.6 ± 84.0a 0.564 1/2 1,226.5 ± 1,523.6a 12,290.3 ± 5,734.3c 6,729.7 ± 6,630.0a 0.246 1 5,941.5 ± 8,297.2ab 11,933.6 ± 9,223.5ab 4,830.5 ± 5,902.7a 0.667 1/2 1,049.1 ± 291.5a 12,690.4 ± 1,878.5c 7,639.3 ± 10,661.8a 0.315 1 12,150.5 ± 5,878.4b 15,073.4 ± 569.8b 3,773.5 ± 4,244.5a 0.146 1/2 319.5 ± 14.6a 473.4 ± 180.7a 5,276.9 ± 2,801.3a 0.089 1 2,337.9 ± 138.2a 9,379.4 ± 10,196.7ab 5,648.5 ± 3,521.1a 0.587 1/2 10,012.4 ± 8,170.4b 2,901.8 ± 3,674.0ab 3,965.3 ± 3,385.6a 0.308 1 909.5 ± 1,020.3a(a) b) 3,361.9 ± 1,762.3a(ab) 4,767.5 ± 1,090.0a(b) 0.032 * 1/2 3,965.3 ± 3,385.6ab 7,983.5 ± 5,035.1bc 3,717.2 ± 3,969.5a 0.428 1 1,498.5 ± 802.5a 2,273.6 ± 2,598.7a 2,267.0 ± 1,489.0a 0.835 1/2 5,035.1 ± 5,154.3ab 3,881.8 ± 1,498.0ab 2,439.9 ± 1,883.0a 0.646 1 2,591.4 ± 1,983.9a 5,688.2 ± 8,291.7ab 5,431.2 ± 7,011.4a 0.810 1/2 5,460.8 ± 4,494.4ab 4,480.5 ± 2,608.7ab 6,179.2 ± 665.3a 0.795 1 3,927.3 ± 1,464.2a 1,298.5 ± 282.9a 1,887.4 ± 2,370.8a 0.194 1/2 2,050.9 ± 524.0ab 985.2 ± 556.2a 2,449.8 ± 2,129.4a 0.331 1 816.8 ± 414.5a 3,602.7 ± 3,105.1a 3,057.9 ± 2,204.9a 0.419 Control - 1,912.1 ± 1,373.7aba 1,874.1 ± 2,786.4aba 561.3 ± 663.9aa P (1/2 fold) ab 0.186 0.002** 0.468 P (1 fold) ab 0.037 * 0.130 0.752 ; Concentration. Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations.

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 203 의경우기준량의 1/2배에서보다기준량을처리했을때, 세개체군에서비탐침기간이 64( 이월 ), 29( 노은 ), 3.2( 실험실 ) 배정도더길게나타났다. 의경우도마찬가지로, 기준량처리시세개체군에서각각약 4( 이월 ), 2( 노은 ), 3( 실험실 ) 배더많이나타났고, spirotetramat도기준량에서각각 7, 19, 1.1배더많이나타났다. 이와대조적으로, dinotefuran을기준량처리시세개체군에서각각약 2, 3, 1.6배더적게나타났고, novaluron도기준량처리시 1.3, 3, 3배더적게나타났다. 의경우기준량을처리했을때, 세개체군간에유의한차이를나타냈는데 (P=0.032) 실험실 > 노은 > 이월개체군순으로비탐침기간을가졌으며실험실개체군은이월개체군보다약 5배더많은비탐침기간을나타냈다. 총탐침소요시간탐침활동의총기간은이월개체군의경우, 약제를기준량으로처리했을때, 유의차를보였다 (P=0.039). 기준량의 1/2배를처리하였을때 gamma-cyhalothrin과 novaluron을 제외한나머지약제에서무처리보다더적은탐침활동을보였지만, cyantraniliprol을제외한나머지등록약제는기준량을처리했을때, 탐침활동이더욱많이나타나무처리구와차이를보였다. 특히 acetamiprid, spirotetramat을기준량으로처리했을때, 무처리보다약 2배많은탐침활동을가졌다. 비등록약제는 novaluron을제외하고, 기준량을처리했을때, 탐침활동이더적게나타났다. 노은개체군의경우, pymetrozine, novaluron, spinosad를처리하였을때, 위의이월개체와같이탐침활동은기준량에서더욱많이나타났지만, 나머지약제에서는기준량의 1/2배처리구보다적은탐침활동을보였다. 실험실개체군의경우, 세개체군중에서모든약제에서기준량처리시더많은구침활동을하거나기준량의 1/2배처리구결과와비슷한수준을나타냈다. 은기준량을처리했을때, 세개체군에서탐침활동은 5( 이월 ), 1.6( 노은 ), 1.4( 실험실 ) 배더많이나타났다. 대조적으로 gamma-cyhalothrin은각각 1.1, 1.8, 1.4배더적게나타났다. 은기준량의 1/2배배처리시실험실 > 노은 > 이월개체군순의탐침활동을, 기준량처리 Table 4. Average running seconds depending on total duration of pathway phase (C) using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration of 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 1,736.2 ± 1,202.6a a) 4,914.5 ± 2,675.6ab 2,738.8 ± 1,229.8a 0.342 1 11,771.1 ± 6,884.4bc 3,198.3 ± 306.6a 10,281.8 ± 6,597.1ab 0.375 1/2 4,218.2 ± 1,944.8ab 12,279.5 ± 6,315.4b 6,188.7 ± 478.1ab 0.237 1 1,390.2 ± 1,225.6a 7,103.2 ± 4,300.3a 12,032.9 ± 5,673.0b 0.177 1/2 1,609.2 ± 943.0a(a) b) 5,724.6 ± 1,201.2 ab(b) 2,918.0 ± 1,365.1 a(ab) 0.084 1 9,608.9 ± 2,860.4bc 9,416.0 ± 8,944.1a 4,187.8 ± 3,146.7ab 0.613 1/2 5,309.4 ± 1,655.7ab 7,175.3 ± 514.4ab 3,459.1 ± 4,000.2a 0.441 1 7,993.0 ± 6,915.3abc 5,567.9 ± 1,709.0a 4,436.2 ± 5,049.6ab 0.787 1/2 3,723.2 ± 1,005.3 ab(a) 7,395.3 ± 1,208.3 ab(ab) 11,698.5 ± 2,426.0b(b) 0.039* 1 12,454.0 ± 3,368.5 c(a) 3,759.6 ± 387.5a(b) 11,598.7 ± 825.6b(a) 0.040* 1/2 6,507.4 ± 2,001.1ab 7,983.5 ± 5,526.8ab 6,357.8 ± 4,824.3ab 0.884 1 4,332.6 ± 3,443.0ab 6,234.9 ± 843.7a 9,428.9 ± 3,601.2ab 0.178 1/2 6,357.8 ± 4,824.3ab 7,125.7 ± 3,146.5ab 4,466.9 ± 2,333.3a 0.665 1 5,624.0 ± 740.5abc 6,730.0 ± 2,939.9a 5,457.2 ± 2,341.8ab 0.756 1/2 7,792.0 ± 2,660.8ab 9,293.2 ± 1,574.8ab 6,433.8 ± 4,443.1ab 0.565 1 7,183.5 ± 4,110.2abc 4,929.3 ± 2,679.3a 4,581.2 ± 2,961.1ab 0.605 1/2 5,342.7 ± 4,316.9ab 4,151.2 ± 2,306.3a 7,554.2 ± 3,073.5ab 0.491 1 8,570.1 ± 4,222.8abc 6,531.7 ± 3,255.5a 8,959.8 ± 7,132.8ab 0.829 1/2 9,557.0 ± 5,894.4b 6,601.4 ± 4,249.0ab 11,221.9 ± 3,459.2b 0.466 1 4,563.8 ± 807.1ab 8,943.6 ± 6,388.9a 8,223.9 ± 3,942.5ab 0.508 Control - 6,702.6 ± 2,048.8ab abc 4,699.2 ± 4,928.3aba 1,207.2 ± 908.1aa P(1/2fold)ab 0.333 0.526 0.026* P(1fold)ab 0.039* 0.850 0.159 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations.

204 민지현 윤헌 권혜리 유용만 윤영남 시이월 > 실험실 > 노은개체군순의탐침활동으로유의차를보였다 (Table 4). 체관부에서섭식행동체관부를섭식할때두가지의다른섭식행동 (E1, E2) 이수반되는데, 첫번째는체관부로의 gelling 타액을분비하여구기를고정시킬수있는구기집형성후 watery 타액을분비하여세포내구멍을뚫고, 섭식전까지분비하는행동과두번째는반드시 E1 이후에나타나는것으로체관부를섭식 (E2) 하는행동이있다 (Tjallingii, 2006; Prado and Tjallingii, 2007; Bos et al., 2010). E1는독립적으로발생할수있지만, E2는독립적으로발생할수없으므로섭식행동분석시두파형을개별적으로분석하였다. 체관부에서타액분비시간 (E1) 체관부를섭식하기전타액을분비하는데총소요시간은세개체군에서농도에상관없이유의한차이는나타나지않았다 (Table 5). 이월개체군의경우, 모든약제를기준량으 로처리했을때 pyrifluquinazon, spinosad를제외하고체관부타액분비기간을더많이가졌다. 특히 spirotetramat, pymetrozine은기준량으로처리했을때무처리대비각각 6 배, 5배더많이나타났으며, 가장많은체관부타액분비기간을갖는다. 노은개체군의경우, 모든약제를기준량으로처리했을때 gamma-cyhalothrin, novaluron, spinosad를제외하고체관부타액분비기간을더적게가졌다. 그리고기준량의 1/2배를처리했을때와비교하여기준량처리시 gamma-cyhalothrin를제외하고모든약제에서무처리대비체관부타액분비가감소한것을확인하였다. 특히 pymetrozine과 pyrifluquinazon을기준량으로처리했을때, E1 파형은나타나지않았으며, 이는체관부를섭식하지않았음을나타내는결과이다. 실험실개체군은모든약제를기준량으로처리했을때 cyantraniliprole에서가장많은타액분비기간을나타내었다. 복숭아혹진딧물의구침을체관부내에침투하여타액을분비하는총소요시간에서약제당개체군별유의차는 cyantraniliprole을기준량의 1/2배처리하였을때, 개체군간 Table 5. Average running seconds depending on total duration of phloem salivation (E1) time using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration of 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 397.3 ± 523.3ab a) 53.0 ± 14.7a 166.7 ± 186.9a 0.605 1 750.0 ± 348.9abc 50.0 ± 15.8a 884.0 ± 560.6a 0.209 1/2 59.6 ± 33.1a(a) b) 1,807.7 ± 139.2b(b) 79.4 ± 38.9a(a) 0.000 ** 1 541.9 ± 725.5abc 203.4 ± 235.9a 2,948.8 ± 3,903.7b 0.510 1/2 430.9 ± 146.3ab 26.3 ± 37.2a 423.1 ± 302.4a 0.205 1 1,130.8 ± 1,371.7bc 0.0 ± 0.0a 89.5 ± 126.6a 0.403 1/2 641.6 ± 558.7ab 808.3 ± 1,143.1ab 72.8 ± 4.3a 0.625 1 251.4 ± 309.8ab 0.0 ± 0.0a 88.8 ± 14.1a 0.461 1/2 55.3 ± 4.2a 1,223.6 ± 1,689.7ab 249.9 ± 261.1a 0.525 1 1,489.7 ± 493.8c 384.5 ± 491.6a 384.9 ± 514.3a 0.177 1/2 16.5 ± 28.5a 280.1 ± 293.2a 267.4 ± 286.8a 0.371 1 70.8 ± 105.4a 254.8 ± 386.0a 558.1 ± 590.8a 0.400 1/2 267.4 ± 286.8a 652.0 ± 792.5ab 177.8 ± 151.5a 0.500 1 275.9 ± 255.9ab 29.5 ± 41.9a 495.6 ± 393.1a 0.191 1/2 0.0 ± 0.0a(a) 927.9 ± 473.2 ab(ab) 918.8 ± 772.2a(b) 0.118 1 337.2 ± 329.6ab 1,031.7 ± 1,711.8a 119.3 ± 141.3a 0.547 1/2 112.4 ± 130.3a(a) 390.9 ± 309.6 ab(ab) 670.4 ± 156.5a(b) 0.051 1 442.7 ± 339.1ab 552.7 ± 540.5a 204.5 ± 296.2a 0.590 1/2 1,172.1 ± 966.2b 135.4 ± 129.1a 1,704.3 ± 2,778.5a 0.550 1 443.4 ± 345.0ab 150.7 ± 197.9a 562.5 ± 534.2a 0.451 Control - 213.7 ± 268.3aab 628.6 ± 1,025.0aa 313.9 ± 383.6aa P(1/2fold) ab 0.084 0.301 0.755 P(1fold) ab 0.171 0.842 0.304 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations.

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 205 에노은 > 실험실 > 이월개체군순으로유의차가나타났다. 도기준량의 1/2배처리하였을때, 개체군간에유의값에가까운차이를나타냈는데실험실 > 노은 > 이월개체군순으로나타났다. 를기준량의 1/2배처리하였을때, 무처리와대비하여세개체군의체관부타액분비기간이모두감소한것을확인하였다. 체관부에서섭식시간 (E2) 체관부를섭식한총소요시간은이월개체군의경우, 기준량의약제를처리하였을때유의차를보였으며, 무처리구와 cyantraniliprole, dinotefuran, spinosad 기준량처리구는나머지약제에비해많은섭식시간을나타냈다 (Table 6). 등록약제 5가지를기준량의 1/2배로처리했을때, 무처리대비모두더많은시간을나타냈으며, 대조적으로비등록약제 5가지를동일하게처리했을때는더적은시간을나타내며유의차를확인하였다. 노은개체군은약제를기준량의 1/ 2배로처리했을때, spinosad를제외하고나머지약제에서무처리대비더적은시간동안체관부를섭식하였으며, 기 준량으로처리했을때는모든약제에서무처리대비체관부섭식의감소를보였다. 그리고앞서 E1 파형이나타나지않은 pymetrozine과 pyrifluquinazon의경우 E2 역시관찰할수없었다. 특히 pyrifluquinazon에서가장적은 E2 시간을가졌다. Pyfluquinazon 기준량의 1/2배처리구에서는앞서 E1은나타냈지만, E2는관찰할수없었다. 실험실개체군의경우, spirotetramat을기준량의 1/2배처리하였을때나머지개체군에비해 4( 이월 ), 6( 노은 ) 배더적은체관부섭식을보이며유의차를나타냈지만, 기준량을처리하였을때에는나머지개체군과달리체관부섭식의감소가나타나지않고오히려증가하였다. 또모든약제에서농도관계없이무처리구에비해적은 E2시간을가졌다. E2 소요시간의약제당개체군별유의차는 cyantraniliprole을기준량처리했을때, 이월 > 노은 > 실험실개체군순으로유의차를나타냈고, spirotetramat을기준량의 1/2배처리했을때, 이월 > 노은 > 실험실개체군순으로유의차를나타냈고, gamma-cyhalothrin을기준량의 1/2배처리하였을때, 체관부섭식소요시간은실험실 > 노은 > 이월개체군순으로유의차를나타냈 Table 6. Average running seconds depending on total duration of phloem ingestion (E2) time using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration of 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 16,165.5 ± 3,794.3b a) 7,080.6 ± 3,247.2ab 15,839.5 ± 523.8ab 0.084 1 1,051.3 ± 1,486.8ab 10,329.2 ± 4,804.5a 2,291.4 ± 208.4a 0.089 1/2 16,897.9 ± 2,310.7b 4,476.5 ± 6,330.8ab 14,352.8 ± 713.8ab 0.097 1 17,948.7 ± 3,700.4b(b) b) 5,121.5 ± 3,688.7a(a) 162.4 ± 229.7a(a) 0.021* 1/2 18,309.6 ± 439.4b(b) 3,264.2 ± 4,616.2ab(ab) 11,499.7 ± 4,959.0ab(b) 0.069 1 4,847.5 ± 6,855.4ab 0.0 ± 0.0a 7,210.3 ± 10,196.9ab 0.632 1/2 14,566.5 ± 2,536.5b 0.0 ± 0.0a 10,364.9 ± 14,658.2ab 0.349 1 125.9 ± 130.1a 0.0 ± 0.0a 11,464.0 ± 11,848.8ab 0.299 1/2 17,377.2 ± 852.4b(b) 12,441.1 ± 3,080.5b(b) 2,591.8 ± 2,175.6a(a) 0.015* 1 1,775.3 ± 1,403.6a(a) 5,093.8 ± 6,977.5 a(ab) 3,728.7 ± 3,789.6a(b) 0.079 1/2 4,992.8 ± 8,647.7ab 8,159.5 ± 7,223.4ab 10,259.2 ± 8,908.6ab 0.747 1 10,893.8 ± 10,166.6ab 3,801.4 ± 6,584.1a 5,980.2 ± 5,294.2ab 0.542 1/2 10,259.2 ± 8,908.6ab 3,011.5 ± 5,216.0ab 13,133.0 ± 6,329.2ab 0.265 1 12,629.2 ± 3,534.8ab 4,729.0 ± 8,108.2a 13,319.0 ± 3,615.9ab 0.187 1/2 0.0 ± 0.0a(a) 5,757.7 ± 2,168.8ab(ab) 11,351.1 ± 6,747.7ab(b) 0.040* 1 8,429.1 ± 8,888.8ab 9,205.4 ± 9,218.0a 11,166.0 ± 9,819.0ab 0.934 1/2 7,714.7 ± 10,120.0ab 9,476.4 ± 4,124.3ab 6,923.2 ± 3,134.8ab 0.890 1 5,267.3 ± 7,743.5ab 10,663.8 ± 3,802.4a 6,083.5 ± 10,536.9ab 0.681 1/2 7,909.1 ± 6,184.7ab 13,339.2 ± 3,647.6b 4,870.9 ± 5,534.1a 0.214 1 15,179.9 ± 1,276.3b 6,171.2 ± 8,393.2a 8,768.3 ± 7,712.9ab 0.268 Control - 11,185.2 ± 4,211.6abab 12,513 ± 8,127.5ba 19,510.1 ± 1,983.6bb P(1/2fold) ab 0.068 0.131 0.203 P(1fold) ab 0.033* 0.537 0.226 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations.

206 민지현 윤헌 권혜리 유용만 윤영남 다. 노은개체군에서비등록약제 chlorfenapyr에대한결과는등록약제와마찬가지로농도에상관없이무처리와대비했을때모두적은 E2 시간을가졌으며, 농도가높을때, 약 2배적은체관부섭식시간나타냈다. 과 novaluron의경우, 세개체군에서농도가증가해도각각유사한수준의 E2를나타냈으며, spinosad는노은개체군에서만농도증가에따른체관부섭식의감소를나타냈다. 물관부에서섭식시간 (G) 물관부를섭식할때나타나는파형 (G) 은세개체군내에서약제별유의차가나타나지않았으며, 약제에따른개체군별분석결과도유의차는없었다 (Table 7). 그러나유의차에가깝게나타난것은실험실개체군에서약제를기준량으로처리하였을때, novaluron의경우물관부섭식시간이가장많은것으로나타났다. 그리고노은개체군에 pyrifluquinazon을 처리하였을때, 두농도에대한물관부섭식시간이유사하게나타났다. 대부분의경우무처리구와같이물관부섭식이거의일어나지않거나짧게일어나기때문에섭식행동의저해유무를판단하기어렵다. 관다발조직이외에서섭식시간 (F) 진딧물의구침이정상적인경로를이탈하였을때를나타내는파형 (F) 은세개체군내에서약제별유의차가나타나지않았으며, 약제에따른개체군별분석결과 pymetrozine 을기준량처리하였을때실험실 > 노은, 이월개체군순으로유의차가나타났다 (Table 8). 본파형의경우약제의농도가증가함에따라발생기간도증가하는경향을보이는데, acetamiprid를제외하고각개체군에서살충효과를보인것으로확인된약제의경우에서그수치가증가하는경향을보였다. Table 7. Average running seconds depending on total duration of feeding xylem (G) using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration of 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 0.0 ± 0.0a a) 0.0 ± 0.0a 0.0 ± 0.0a - 1 0.0± 0.0a 0.0 ± 0.0a 1,023.2 ± 792.7a 0.465 1/2 0.0 ± 0.0a 517.3 ± 686.3ab 0.0 ± 0.0a 0.429 1 0.0 ± 0.0a 0.0 ± 0.0a 1,075.6 ± 1,521.1ab 0.465 1/2 0.0 ± 0.0a 0.0 ± 0.0a 0.0 ± 0.0a - 1 0.0± 0.0a 60.9 ± 86.2a 0.0 ± 0.0a 0.465 1/2 0.0 ± 0.0a 849.9 ± 1,201.9b 0.0 ± 0.0a 0.465 1 0.0 ± 0.0a 853.3 ± 1,206.8a 36.4 ± 51.5a 0.477 1/2 0.0 ± 0.0a 0.0 ± 0.0a 266.5 ± 376.9a 0.465 1 1,023.2 ± 792.7a 0.0 ± 0.0a 121.9 ± 172.4a 0.465 1/2 0.0 ± 0.0a 0.0 ± 0.0a 725.8 ± 1,257.0a 0.422 1 1,636.9 ± 2,835.2a 1,817.4 ± 3,147.8a 0.0 ± 0.0a 0.628 1/2 725.8 ± 1,257.0a 0.0 ± 0.0a 0.0 ± 0.0a 0.422 1 108.3 ± 187.5a 2,049.8 ± 3,550.3a 25.5 ± 38.5a 0.443 1/2 3,661.7 ± 6,308.8a 0.0 ± 0.0a 0.0 ± 0.0a 0.419 1 0.0± 0.0a 0.0 ± 0.0a 0.0 ± 0.0a - 1/2 412.1 ± 713.8a 0.0 ± 0.0a 0.0 ± 0.0a 0.422 1 952.0 ± 889.7a 0.0 ± 0.0a 2,382.5 ± 2,072.1b 0.158 1/2 0.0 ± 0.0a 0.0 ± 0.0a 485.3 ± 840.5a 0.422 1 0.0± 0.0a 197.4 ± 341.8a 0.0 ± 0.0a 0.422 Control - 0.0 ± 0.0aa 239.1 ± 414.1aba 0.0 ± 0.0aa P(1/2fold) ab 0.706 0.305 0.752 P(1fold) ab 0.589 0.796 0.062 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides.

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 207 Table 8. Average running seconds depending on total duration of derailed stylet (F) using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 0.0 ± 0.0a a) 9,296,7 ± 6,195.0b 1,933.6 ± 2,734.5b - 1 0.0± 0.0a 1,349.1 ± 1,907.9a 4,271.8 ± 5,849.7ab 0.542 1/2 0.0 ± 0.0a 690.4 ± 294.9a 0.0 ± 0.0a 0.429 1 0.0± 0.0a 4,847.9 ± 5,801.5a 175.8 ± 92.1a 0.382 1/2 0.0 ± 0.0a 96.0 ± 135.8a 0.0 ± 0.0a - 1 0.0± 0.0a(a) b) 135.0 ± 191.0a(a) 5,181.0 ± 1,204.0b(b) 0.008 ** 1/2 0.0 ± 0.0a 0.0 ± 0.0a 32.4 ± 45.9a 0.465 1 1,054.7 ± 1,491.5a 60.9 ± 86.2a 1,779.3 ± 2,516.3ab 0.639 1/2 0.0 ± 0.0a 26.5 ± 37.4a 1,470.9 ± 1,877.2ab 0.465 1 2,279.9 ± 3,224.3a 2,919.5 ± 4,128.8a 0.0 ± 0.0a 0.542 1/2 0.0 ± 0.0a 1,801.8 ± 3,032.2a 0.0 ± 0.0a 0.404 1 3,750.3 ± 3,385.3a 6,108.0 ± 5,347.4a 826.8 ± 1,432.1a 0.297 1/2 0.0 ± 0.0a 2,651.5 ± 4,592.6a 0.0 ± 0.0a 0.422 1 1,233.7 ± 2,136.8a 5,773.9 ± 6,678.6a 0.0 ± 0.0a 0.261 1/2 5,073.8 ± 8,788.1a 1,696.4 ± 2,904.5a 319.3 ± 553.0ab 0.566 1 2,902.7 ± 3,980.7a 558.6 ± 967.5a 21.0 ± 36.3a 0.349 1/2 2,525.4 ± 4,374.2a 3,064.5 ± 4,605.9a 0.0 ± 0.0a 0.580 1 2,392.4 ± 3,426.7a 2,354.8 ± 4,078.6a 2,069.8 ± 3,431.9ab 0.993 1/2 831.7 ± 720.3a 490.7 ± 849.8a 431.5 ± 747.4ab 0.799 1 0.0± 0.0a 2,452.7 ± 4,160.5a 768.2 ± 1,260.6a 0.512 Control - 1,547.3 ± 1,401.4aa 1,618.9 ± 2,804.0aa 0.0 ± 0.0aa P (1/2fold) ab 0.782 0.258 0.307 P (1fold) ab 0.570 0.697 0.165 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations. 총탐침횟수복숭아혹진딧물이섭식을시도한총횟수를나타내는 (total number of probes) 결과는 (Table 9), 무처리에서나타난횟수를기준으로많을경우, 약제에대한반응으로섭식에적합하지않을때계속해서적합한먹이를찾고자하는시도로해석된다. 노은개체군에서농도에관계없이약제간차이를확인할수있었지만, 나머지개체군에서는유의차를확인할수없었다. 노은개체군에서탐침수는 pymetrozine 과 pyrifluquinazon에서가장많이나타났다. 약제에따른개체군별분석결과 pyrifluquinazon을기준량으로처리하였을때노은 > 이월 > 실험실개체군순으로탐침수의유의차를관찰할수있었으며, spirotetramat을기준량의 1/2배로처리하였을때실험실 > 노은 > 이월개체군순으로탐침수의유의차가나타났고, novaluron을기준량의 1/2배로처리하였을때실험실 > 노은 > 이월개체군순으로세개체군간 에유의차를확인할수있었는데이는 novaluron에대한살충효력시험결과실험실개체군에서 60% 미만이지만최대방제가를나타낸것과연관지을수있을것으로보인다. 최초탐침하는데소요된시간복숭아혹진딧물의첫탐침시간 (time to first probe) 은먹이에반응하는진딧물의행동을보는것으로, 섭식에기피를일으키는약제의경우첫탐침시간이지연되어나타나기도한다. 따라서첫탐침까지걸리는시간이길수록섭식행동을저해하는것으로판단하여분석하였다. 그결과무처리구와처리구간의유의차를확인할수없었지만각약제의농도에따른반응을통해섭식행동의변화를살펴보았다 (Table 10). 각개체군별로살펴보면, 실험실개체군의경우약제농도가증가했을때, 모든약제에대한반응은무처리에비해첫탐침시간이지연되었으며, 이월개체군은

208 민지현 윤헌 권혜리 유용만 윤영남 Table 9. Average number depending on total number of probes using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 6.0 ± 4.2a a) 5.0 ± 4.2a 13.0 ± 7.1a 0.387 1 44.5 ± 47.4a 39.0 ± 12.7abc 27.5 ± 12.0a 0.846 1/2 10.5 ± 4.9a 45.0 ± 26.9ab 19.5 ± 4.9a 0.230 1 8.5± 4.9a 19.5 ± 4.9a 25.0 ± 14.1a 0.148 1/2 14.5 ± 9.2a 57.5 ± 27.6bc 30.0 ± 7.1a 0.181 1 53.0 ± 62.2a 79.5 ± 2.1c 31.0 ± 35.4a 0.567 1/2 18.5 ± 6.4a 95.0 ± 8.5c 42.5 ± 57.3a 0.213 1 19.0 ± 7.1a(a) b) 71.0 ± 1.4bc(b) 7.5 ± 4.9a(a) 0.002 ** 1/2 7.0 ± 2.8a(a) 16.0 ± 7.1ab(a) 41.0 ± 2.8a(b) 0.011 * 1 30.0 ± 2.8a 43.5 ± 6.4abc 165.0 ± 162.6b 0.402 1/2 50.0 ± 44.5a 18.7 ± 21.1ab 35.3 ± 33.9a 0.569 1 10.0 ± 7.0a 17.7 ± 4.0a 46.3 ± 30.2a 0.104 1/2 35.3 ± 33.9a 44.3 ± 28.4ab 21.0 ± 22.6a 0.627 1 18.0 ± 7.5a 26.3 ± 13.2ab 11.7 ± 4.0a 0.220 1/2 17.7 ± 15.0a 32.3 ± 8.7ab 23.0 ± 14.0a 0.423 1 36.0 ± 25.7a 39.7 ± 46.1abc 29.7 ± 23.7a 0.934 1/2 17.0 ± 10.6a(a) 18.7 ± 3.1ab(a) 55.7 ± 20.0a(b) 0,019 * 1 60.7 ± 55.9a 15.0 ± 2.6a 22.0 ± 26.2a 0.311 1/2 28.0 ± 7.5a 15.3 ± 14.0ab 25.7 ± 13.5a 0.442 1 27.7 ± 17.2a 41.7 ± 22.5abc 39.3 ± 20.0a 0.675 Control - 17.0 ± 9.2aa 29.0 ± 34.7abab 11.3 ± 11.8aa P (1/2fold) ab 0.447 0.014* 0.504 P (1fold) ab 0.523 0.099 0.096 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations. cyantraniliprole, spinosad를제외하고모두지연되었고, 노은개체군은 cyantraniliprole, chlorfenapyr, spinosad를제외하고모두지연되었다. 각약제별로반응을살펴보면먼저, acetamiprid의농도에따른각개체군의첫탐침시간은노은개체군에서는큰차이없었지만이월개체군에서는기준량처리시약 300초더늦게나타났으며, 실험실개체군에서는오히려약 290초더빠르게나타났다. 그리고실험실개체군의경우, acetamiprid을기준량의 1/2배를처리하였을때첫탐침까지약 3-70초미만으로소요된다른약제에서와달리 300초대에이르며유의값에가까운수치를나타냈다. 살충효과가나타나지않은 cyantraniliprole의경우, 세개체군에서두농도에따른유의한차이는나타나지않았으며, 실험실개체군을제외한이월, 노은개체군에서무처리와유사한수치를보였다. 을기준량의 1/2배로처리하였을때에는세개체군간의차이가적게나타났지만, 기준량에서이월개체군은무처리와유사한첫탐침을보였으며노은개체군은약20초더늦은첫탐침을보였고, 실험실개체군은농도에따른차이는나타나지않았다. 의경우, 모든개체군에서농도에따른첫탐침시간의변화가가장뚜렷하게나타났다. 기준량의 1/2배를처리하였을때와비교해서기준량을처리했을때, 이월개체군의경우약 1,550초, 노은개체군은약 190초, 실험실개체군은약 180초정도첫탐침행동이많이지연된것을관찰할수있었다. 2016 작물보호제지침서에서표기된내용에따르면, 약제저항성을갖는해충에효과적으로작용하며빠른섭식억제작용및약효지속이길게나타나는약제특성을가지며이에따라섭식기피행동으로나타난것으로볼수있다. 또한기준량의 1/2배처리하였을때이월 > 실험실, 노은개체군순으로첫탐침시간의유의차를관찰할수있었다. 는노은개체군을제외하고두개체군에

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 209 Table 10. Average running seconds depending on Time to first probe using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 0.4 ± 0.1a a) 13.3 ± 18.5a 315.6 ± 338.7b 0.327 1 388.5 ± 478.6a 12.6 ± 5.6a 24.2 ± 1.2a 0.415 1/2 16.6 ± 19.1a 16.0 ± 3.3a 33.1 ± 4.0a 0.365 1 1.8± 2.5a 6.4 ± 8.7a 36.4 ± 51.4a 0.535 1/2 24.3 ± 27.2a 33.1 ± 30.9a 24.3 ± 27.2a 0.940 1 8.7± 12.3 51.5 ± 54.4ab 21.5 ± 4.4a 0.486 1/2 43.1 ± 5.1a(b) b) 0.9 ± 0.4a(a) 3.5 ± 4.7a(a) 0.003 ** 1 1,594.9 ± 2,067.0b 193.7 ± 272.8b 285.8 ± 401.3a 0.521 1/2 55.2 ± 77.9a 4.3 ± 5.5a 16.3 ± 22.6a 0.585 1 226.1 ± 255.0a 66.8 ± 64.2ab 25.9 ± 35.1a 0.478 1/2 79.8 ± 26.3a 179.9 ± 226.5b 29.0 ± 26.8a 0.420 1 101.2 ± 156.9a 8.1 ± 13.2a 51.5 ± 7.0a 0.497 1/2 29.0 ± 26.8a 684.9 ± 782.9b 68.5 ± 78.1a 0.221 1 36.7 ± 52.5a 16.1 ± 27.0a 32.5 ± 37.0a 0.808 1/2 213.9 ± 302.5a 35.8 ± 41.5a 14.6 ± 13.3a 0.377 1 33.2 ± 57.0a 47.3 ± 47.3ab 84.8 ± 116.7a 0.728 1/2 114.0 ± 164.6a 17.7 ± 24.5a 52.0 ± 23.5a 0.508 1 17.9 ± 15.7a(a) 49.0 ± 60.6ab(ab) 116.0 ± 35.1a(b) 0.067 1/2 35.9 ± 23.5a 91.8 ± 145.7a 6.9 ± 8.7a 0.505 1 5.0± 4.5a 3.7 ± 2.4a 174.5 ± 301.1a 0.436 Control - 5.8 ± 5.8aa 8.6 ± 9.6aa 6.3 ± 2.1aa P(1/2fold) ab 0.678 0.241 0.078 P(1fold) ab 0.162 0.383 0.685 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations. 서는높은농도일수록첫탐침이늦춰지는것을보였다. 은노은개체군에서가장늦은탐침을보였다. 그러나추천농도에서는이와유의한결과를볼수없었다. Gamma-cyhalothrin은이월개체군을제외하고두개체군에서는높은농도일수록첫탐침이늦춰지는것을보였다. 체관부에서처음으로섭식하는데소요된시간복숭아혹진딧물의첫체관부섭식시간 (Time to first feeding phloem) 은탐침활동을거쳐실제섭식을갖기까지걸리는시간을통해, 약제반응으로인한저해여부를유추할수있다 (Table 11). 모든약제를기준량의 1/2배로처리하였을때, 각개체군별반응으로이월개체군의경우 chlorfenapyr, gamma-cyhalothrin, novalruon, spinosad를제외한약제에서무처리보다더빠른시간내에체관부를섭식한것으로나타났다. 다음노은개체군은동일한농도에서모든약제에 서무처리에비해체관부첫섭식시간이지연된것으로나타났으며, 마지막으로실험실개체군도마찬가지로나타났다. 또모든약제를기준량으로처리하였을때에는, 이월개체군에서 acetmiprid, pymetrozine, spirotetramat 은낮은농도보다더욱지연된섭식시간을보였으며, 비등록약제의경우낮은농도보다더빠른섭식시간을나타내며유의한결과는없었다. 마찬가지로노은개체군에서 pymetrozine, chlorfenapyr, spinosad가낮은농도에서보다더욱지연된섭식시간을보였지만나머지약제에서는모두낮은농도에서보다더빠른섭식시간을나타냈다. 그러나실험실개체군의경우 pyrifluquinazon, dinotefuran을제외하고모든약제에서낮은농도보다더욱지연된섭식시간을나타냈다. 특히 pymetrozine은농도가증가했을때모든개체군에서체관부첫섭식시간이더욱지연되는것을관찰하였고, 약제에따른개체군별차이는 pyrifluquinazon을기준량으로처

210 민지현 윤헌 권혜리 유용만 윤영남 Table 11. Average running seconds depending on Time to first feeding phloem using EPG techniques for Myzus persicae on pepper leaf treated 2 different concentration 10 different insecticides for 6 hours Gamma- cyhalothrin 1/2 5,434.1 ± 3,794.2a a) 14,506.1 ± 3,228.7ab 4,186.7 ± 2,641.9a 0.090 1 14,420.3 ± 10,082.8a 11,258.2 ± 4,810.1abc 8,614.5 ± 7,799.4a 0.778 1/2 4,020.0 ± 3,270.9a 11,694.5 ± 13,982.6ab 5,535.4 ± 1,656.3a 0.662 1 832.8 ± 285.6a 7,870.8 ± 8,427.5abc 13,951.0 ± 10,714.5a 0.374 1/2 3,266.1 ± 466.5a 11,854.1 ± 13,705.2 ab 5,213.0 ± 1,945.4a 0.589 1 11,647.2 ± 14,050.7a 21,548.5 ± 54.4c 14,368.2 ± 10,192.5a 0.639 1/2 3,053.7 ± 3,036.1a 21,599.1 ± 0.4b 11,231.6 ± 14,662.9a 0.247 1 1,234.2 ± 208.3a(a) b) 21,406.3 ± 272.8c(b) 1,955.1 ± 1,084.7a(a) 0.000 ** 1/2 3,091.7 ± 2,451.8a 9,154.6 ± 3,086.0ab 10,309.1 ± 10,081.2a 0.537 1 3,550.6 ± 1,918.5a(a) 2,484.7 ± 1,719.4 a(a) 17,845.4 ± 3,824.7a(b) 0.017 * 1/2 16,527.4 ± 8,673.9ab 10,500.3 ± 9,831.9ab 9,277.1 ± 10,879.0a 0.648 1 10,605.0 ± 10,017.6a 17,790.6 ± 6,577.7bc 11,444.8 ± 10,119.9a 0.596 1/2 9,277.1 ± 10,879.0ab 17,457.7 ± 5,475.0ab 5,948.3 ± 2,428.1a 0.210 1 8,934.0 ± 3,557.9a 13,033.7 ± 7,521.5abc 4,821.1 ± 2,514.8a 0.215 1/2 21,386.1 ± 302.5b(a) 12,718.1 ± 4,629.1 ab(b) 7,086.6 ± 2,393.2a(b) 0.003 ** 1 8,361.0 ± 10,280.9a 9,695.6 ± 10,302.9abc 10,349.2 ± 9,842.2a 0.971 1/2 13,208.9 ± 9,555.5ab 12,105.8 ± 4,101.7ab 8,429.7 ± 5,442.4a 0.681 1 10,205.9 ± 10,079.3a 10,571.3 ± 4,317.3abc 15,400.6 ± 10,505.7a 0.732 1/2 11,716.4 ± 6,439.8ab 8,169.0 ± 3,785.8ab 9,522.3 ± 10,461.2a 0.844 1 5,423.5 ± 1,956.5a 11,477.2 ± 7,911.7abc 9,903.9 ± 10,574.4a 0.631 Control - 9,612.2 ± 5,581.4aba 13,342.9 ± 5,004.5aab 9,239.7 ± 4,337.6aa P(1/2fold) ab 0.080 0.453 0.934 P(1fold) ab 0.583 0.095 0.474 Values represent by means ± SD, *P<0.05, **P<0.01; Completely randomized one-way analysis of variance, ANOVA, Post Hoc Tests by Duncan in SPSS version 22.0. a) indicates significantly different among insecticides; ( ) b) indicates significantly different among populations. 리하였을때노은개체군에서가장늦게나타났으며, 실험실개체군, 이월개체군순으로유의차를나타냈다. 역시기준량으로처리하였을때실험실개체군에서가장늦게나타났으며, 이월개체군, 노은개체군순으로유의차를나타냈다. 또, Gamma-cyhalothrin을기준량의 1/2배로처리하였을때세개체군간에유의차를확인할수있었다. 고찰 본연구에서는 EPG 기술을이용하여복숭아혹진딧물에작용기작이다른계통의약제를처리하였을경우살충효력과함께섭식행동에미치는영향을알아보고자하였다. Rawat et al. (2013) 에따르면, 복숭아혹진딧물의저항성발달에맞춰실험실및현장, 온실조건에서진딧물에대해다 른그룹에속한몇가지새로운화합물을포함한다양한살충제의효율성과경제적중요성이보고되고있지만, 이러한살충제대부분은대상이아닌생물체에높은독성과높은잔류성및환경에유해한다른영향때문에내성이생기게된다. 따라서적은잔류성, 안정성, 효율적인방제방법의대안책으로진딧물에대한다른살충제의독성평가가필요하다고보고하였다. 따라서본연구에서는복숭아혹진딧물의체내작용기작이다른 8 계통의 10개약제를선발하였으며, 이중에서복숭아혹진딧물에등록되어있는 acetamiprid, cyantraniliprole, pymetrozine, pyrifluquinazon, spirotetramat와비등록약제인, chlorfenapyr, dinotefuran, gamma-cyhalothrin, novaluron, spinosad를처리하였을경우에복숭아혹진딧물에미치는영향을관찰하고자하였다. EPG 기술은위의다양한살충제에대한복숭아혹진딧물의반응을관찰하기위해사용한기술로, EPG기술을활용

EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 211 하여관찰한복숭아혹진딧물의섭식행동패턴은 6가지로분류하였다. 섭식을하지않을때나타나는파형인 Np, 섭식을시작하여구기가기주식물에닿고식물체에침투할때나타나는파형인 C, 체관부에섭식시타액을분비할때나타나는파형인 E1, 체관부섭식할때나타나는파형인 E2, 물관부를섭식할때나타나는 G, 구기가경로를이탈하였을때나타나는파형인 F를포함하여관찰하였다. 본실험결과, cyantraniliprole의섭식행동변화는실험실개체군에서농도증가에따른섭식행동의변화를관찰했는데첫탐침시간의지연, 총탐침수의증가, 총탐침활동증가, 비탐침활동증가, 체관부첫섭식시간지연, 체관부섭식시간의감소, 구침이탈시간의증가를확인하였으며, 이는약제에의한반응으로섭식행동의저해작용을확인한것으로볼수있다. 이를뒷받침하는 Jacobson and Kennedy (2014) 의연구에따르면, 복숭아혹진딧물에대한 imidacloprid 와 cyantraniliprole의섭식행동의저해효과를비교하였으며결과적으로 imidacloprid보다는섭식행동의저해효과가늦게나타나지만, 유의한효과가있는것으로결론지었다. Naeun (1995) 이보고한바에의하면저농도의 pirimicarb가섭식행동에영향을주지않는다고하였지만, 본실험에서사용한약제를기준량의 1/2배로처리하였을때복숭아혹진딧물의 EPG 분석결과처리구와무처리구사이에유의한차이를확인할수있었고, 기준량농도처리시더욱뚜렷한차이를확인하였다. 이러한기초적인자료는복숭아혹진딧물의더욱효과적이고새로운방제방법을모색하기위한자료로서사용될것으로사료된다. Literature Cited Alvarez, A. E., V. G. Broglia, A. M. Alberti D Amato, D. Wouters, E. van der Vossen, E. Garzo and B. Vosman (2013) Comparative analysis of Solanum stoloniferum responses to probing by the green peach aphid Myzus persicae and the potato aphid Macrosiphum euphorbiae. Insect Sci. 20(2):207-227. Bass, C., A. M. Puinean, C. T. Zimmer, I. Denholm, L. M. Field, S. P. Fosterand and M. S. Williamson (2014) The evolution of insecticide resistance in the peach potato aphid, Myzus persicae. Insect Biochem. Mol. Biol. 51:41-51. Blackman, R. L. and V. F. Eastop (2000) Aphids on the world s crops an identification and information guide. New York: Willey and Son. pp. 466. Bonani, J. P., A. Fereres, E. Garzo, M. P. Miranda, B. Appezzato-Da-Gloria and J. R. S. Lopes (2010) Characterization of electrical penetration graphs of the Asian citrus psyllid, Diaphorina citi, in sweet orange seedlings. Entomol. Exp. Appl. 134:35-49. Bos, J. I., D. Prince, M. Pitino, M. E. Maffei, J. Win and S. A. Hogenhout (2010) A functional genomics approach identifies candidate effectors from the aphid species Myzus persicae (green peach aphid). PLoS Genet.6(11):e1001216. Chang, V. C. S. (1978) Feeding activities of the sugarcane leafhopper: Identification of electronically recorded waveforms. Ann. Entomol. Soc. Am. 71:31-36. Civolani, S., M. Leis, G. Grandi, E. Garzo, E. Pasqualini, S. Musacchi, M. Chicca, G. Castaldelli, R. Rossi and W. F. Tjallingii (2011) Stylet penetration of Cacopsylla pyri; an electrical penetration graph (EPG) study. J. Insect Physiol. 57:1407-1419. Costa, R. R., J. C. Moraes and R. R. DaCosta (2011) Feeding behaviour of the greenbug Schizaphis graminumon wheat plants treated with imidacloprid and/or silicon. J. Appl. Entomol. 135(12):115-120. Fereres, A. and A. Moreno (2009) Behavioural aspects influencing plant virus transmission by homopteran insects. Virus Res. 141(2):158-168. Garzo, E., A. Moreno, S. Hernando, V. Mario, M. Torne, E. Santamaria and A. Fereres (2016) Electrical penetrationgraph technique as a tool to monitor the early stages of aphid resistance to insecticides. Pest Manag. Sci. 72(4):707-718. Gaspari, M., D. Lykouressis, D. Perdikis and M. Polissiou (2007) Nettle extract effects on the aphid Myzus persicae and its natural enemy, the predator Macrolophus pygmaeus (Hemiptera: Miridae). J. Appl. Entomol. 131:652 657. Harrewijn, P. and H. Kayser (1997), a fastacting and selective inhibitor of aphid feeding. insitu studies with electronic monitoring of feeding behaviour. Pest. Sci. 49(2):130-140. Helden, M. and W. F. Tjallingii (1993) Tissue localization of lettuce resistance to the aphid Nasonovia ribisnigri using electrical penetration graphs. Entomol. Exper. Appl. 68(3): 269-278. Jacobson, A. L. and G. G. Kennedy (2014) Electrical penetration graph studies to investigate the effects of cyantraniliprole on feeding behavior of Myzus persicae (Hemiptera: Aphididae) on Capsicum annuum. Pest Manag. Sci. 70(5):836-840. Jassen, J. A. M., W. F. Tjallingii and J. C. van Lenteren (1989) Electrical recording and ultrastructure of stylet penetration by the greenhouse whitefly. Entomol. Exp. Appl. 52:69-81. Jiang, Y. X., H. Lei, J. L. Collar, B. Martin, M. Muiz and A. Fereres (1999) Probing and feeding behavior of two distinct biotypes of Bemisia tabaci (Homoptera: Aleyrodidae) on tomato plants. J. Econ. Entomol. 92(2):357-366. Kasprowicz, L., G. Malloch, J. Pickup and B. Fenton (2008) Spatial and temporal dynamics of Myzus persicae clones in fields and suction traps. Agricul. For. Entomol. 10:91-100. Kawabe, S. and D. L. McLean (1978) Electronically recorded wave forms associated with salivation and ingestion behaviour of the Aster leafhopper, Macrosteles fascifrons

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EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 213 EPG 기술을이용한복숭아혹진딧물 (Myzus persicae) 의몇가지살충제에대한섭식행동 민지현 윤헌 권혜리 유용만 윤영남 * 충남대학교농업생명과학대학응용생물학과 요약 3 곳의지역에서채집된복숭아혹진딧물세개체군 ( 이월, 노은, 실험실개체군 ) 의 EPG 패턴을통해섭식행동을분석하고, 작용기작이다른총 10 가지의약제에대한각개체군별섭식행동반응을비교분석하였다. 약제를처리하지않은무처리구에서복숭아혹진딧물의섭식행동성향은체관부에서섭식하는시간이물관부에서보다더길었다. 탐침하지않은총기간은 의경우반량처리보다기준량을처리했을때, 3 개의개체군에서비탐침기간이무처리구보다이월개체군에서는 64 배, 노은개체군에서는 29 배, 실험실개체군에서는 3.2 배더길게나타났다. 탐침활동은 cyantraniliprol 을제외한나머지등록약제는기준량을처리했을때더욱많이나타났다. 복숭아혹진딧물의구침을체관부내에침투하여타액을분비하는총소요시간에서약제당개체군별유의차는 cyantraniliprole 을기준량의반량을처리하였을때, 노은 > 실험실 > 이월개체군순으로유의차가나타났다. 모든약제처리구에서농도와상관없이무처리구에비해체관부에서섭식하는시간이짧았으며, 특히 cyantraniliprole 을기준량처리했을때, 이월 > 노은 > 실험실개체군순으로유의차를나타냈었다. 물관부에서섭식하는시간은세개체군모두약제별유의차가나타나지않았다. 따라서각약제별로서로다른개체군에서의감수성차이를확인하고저항성발달여부살펴저항성관리를수행해야할것으로판단된다. 색인어 복숭아혹진딧물, EPG, 살충제, 섭식행동