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“觅食”是生物生存与繁衍的基础,是生物间食物链的关键一环,也是生物学及进化生态学的核心内容[1-2]。而植物挥发物(VOCs)在寄主植物-植食性昆虫-天敌三级营养链连接中发挥着至关重要的作用,是植食性昆虫及天敌昆虫搜寻和定位寄主的重要信号物质[3-6]。据统计,在植食性昆虫与寄主互作研究中,超98%的报道是以地面昆虫(多以成虫为主)为对象,对于地下昆虫的关注较少[7-8]。相对于地面昆虫而言,在黑暗(视觉刺激缺失)、复杂的土壤环境中昆虫如何定位到寄主植物知之不多[9-11]。自Doane等[12]首次证实马铃薯叩甲(Ctenicera destructor (Brown))幼虫依靠植物根系释放的CO2定位寄主以来,胡萝卜根实蝇(Psila rosae F.)、玉米根萤叶甲(Diabrotica virgifera virgifera Leconte)及大栗鳃金龟(Melolontha hippocastani Fabricius)幼虫等多种地下昆虫被证实对CO2有显著的正趋向性[9,13],CO2一度被认为是地下植食性昆虫定位寄主的唯一信号物质[14-15]。
随着研究的深入,发现相比于CO2,根部释放的挥发物对土壤植食性昆虫定位寄主更有效[11,16-17],如大麦(Hordeum vulgare L.)根系释放的hexanal、(E)-hex-2-enal及(E)-non-2-enal等4种挥发物能显著激发金针虫(鞘翅目(Coleoptera)叩甲科(Elateridae)昆虫幼虫的统称)的定向移动[11]。植物根系挥发物在地下植食性昆虫寄主搜寻过程中发挥着重要作用,但研究报道并不多。筛胸梳爪叩甲(Melanotus cribricollis (Faldermann))是我国南方竹林金针虫的优势种[18],主要蛀食雷竹(Phyllostachys violascens (Carriere) Riviere & C. Riviere)、毛竹(Ph. edulis (Carriere) J. Houzeau)及乌哺鸡竹(Ph. vivaxs McClure)等多种笋用竹种,造成鲜笋腐烂和退笋,部分林分鲜笋受害率超70%,留养的种笋受害率高达100%,严重影响竹笋的产量和质量,威胁竹笋产业的健康发展[19]。前期研究表明,与普通小麦(Triticum aestivum L.)、玉米(Zea mays L.)等禾本科(Poaceae)作物相比,筛胸梳爪叩甲幼虫偏好取食雷竹笋,竹笋特有的挥发物可能发挥了重要作用[20],但竹笋挥发物的组成及与CO2的引诱活性差异尚未明确。本研究监测比较雷竹笋挥发物和CO2释放剂对筛胸梳爪叩甲幼虫的引诱作用,并鉴定了竹笋挥发物组分,旨在探索竹林金针虫的寄主定位及识别机制,为竹林金针虫的高效监测及防控提供科学依据。
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鲜笋诱集试验表明,雷竹鲜笋对筛胸梳爪叩甲幼虫表现出显著的吸引作用(表1),但不同质量的竹笋间差异不显著(χ2=0.576,df=2,P>0.05;图2)。释放金针虫24 h后,大多数金针虫选择在埋有竹笋的一端,聚集在竹笋周围或钻入竹笋内蛀食。供试的200头金针虫,10 g竹笋端有128头,占供试金针虫的64.00%(GP=69.43,P<0.001;GH=21.10,P=0.331;表1);20 g竹笋端有118头,占供试金针虫的59.00%(GP=61.62,P<0.001;GH=23.64,P=0.210;表1);40 g竹笋端有136头,占供试金针虫的68.00%(GP=84.17,P<0.001;GH=8.74,P=0.977;表1)。释放金针虫24 h后,不同处理均有20%左右的金针虫仍停留在中心虫室,未作出选择的金针虫在不同质量鲜笋处理中无显著差异(图2)。
表 1 筛胸梳爪叩甲幼虫对不同质量竹笋的选择
Table 1. Choice tests of M. cribricollis larvae towards different weight bamboo shoots in a dual-choice olfactometer
处理
Treatments重复
Replicates作出选择的金针虫数量(百分比)
Number of wireworms at different sides(Percentage)G检验
G test处理端
Treated side对照端
Control sideGP GH 10 g 20 128(82.05%) 28(17.95%) 69.43*** 21.10NS 20 g 20 118(81.38%) 27(18.62%) 61.62*** 23.64NS 40 g 20 136(84.47%) 25(15.53%) 84.17*** 8.74NS 注:仅对作出选择的金针虫个体数进行G检验。***表示P<0.001,NS表示差异不显著(P>0.05)。下同。
Note:G test was performed only on the number of wireworms at the treated and control sides. *** means P<0.001,NS means that the difference is not significant(P>0.05). The same followed. -
CO2对筛胸梳爪叩甲幼虫有显著的吸引作用,且不同浓度的CO2对金针虫引诱力差异显著(χ2=7.945,df=2,P<0.05;表2,图3)。放入10粒CO2释放剂颗粒时,供试的200头金针虫中161头作出了选择,CO2端有99头,占供试金针虫的49.50%(GP=8.58,P<0.01;GH=9.21,P=0.970);放入20粒CO2释放剂颗粒时,作出选择的有141头,CO2端有107头,占供试金针虫的53.50%(GP=36.70,P<0.001;GH=22.22,P=0.274);放入40粒CO2释放剂颗粒时,作出选择的有149头,CO2端有107头,占供试金针虫的53.50%(GP=29.33,P<0.001;GH=11.82,P=0.893)。20粒CO2释放颗粒剂对金针虫的引诱力最强,与10粒CO2释放颗粒剂相比有显著差异,但与40粒CO2释放颗粒剂间差异不显著(图3)。
表 2 筛胸梳爪叩甲幼虫对不同浓度CO2的选择
Table 2. Choice tests of M. cribricollis larvae towards CO2 of different concentrations in a dual-choice olfactometer
处理
Treatments重复
Replicates作出选择的金针虫数量(百分比)
Number of wireworms at different sides(Percentage)G检验
G-test处理端
Treated side对照端
Control sideGP GH T1 20 99(61.49%) 62(38.50%) 8.58** 9.21NS T2 20 107(75.89%) 34(24.11%) 36.70*** 22.22NS T3 20 107(71.81%) 42(28.19%) 29.33*** 11.82NS 注:T1、T2、T3分别表示10、20和40粒CO2释放剂颗粒,下同。**表示P<0.01。
Note: T1, T2 and T3 presents 10, 20 and 40 granules of CO2 resource, respectively. The same followed. ** means P<0.01. -
在竹笋和CO2释放剂颗粒之间,筛胸梳爪叩甲幼虫显著偏好竹笋,竹笋的吸引力强于CO2,但不同的时间处理(24 h和48 h)间差异不显著(χ2=0.314,df=1,P>0.05)。供试的200头金针虫中,24 h后竹笋端有139头(占供试金针虫的69.5%),CO2端有34头(GP=68.37,P<0.001;GH=15.40,P=0.697;表3),而48 h后,竹笋端有142头(占供试金针虫的71.0%),CO2端有35头(GP =47.69,P<0.001;GH=15.40,P=0.749;表3)。
表 3 筛胸梳爪叩甲幼虫在竹笋和CO2释放物间的选择
Table 3. Choice tests of M. cribricollis larvae between bamboo shoots and CO2 resource in a dual-choice olfactometer
处理
Treatments重复
Replicates作出选择的金针虫数量(百分比)
Number of wireworms at different sides(Percentage)G检验
G-test竹笋
Bamboo shoots二氧化碳
CO2GP GH 24 h 20 139(80.35%) 34(19.65%) 68.37 *** 15.40 NS 48 h 20 142(80.23%) 35(19.77%) 47.69 *** 14.58 NS -
固相微萃取法收集、鉴定雷竹笋挥发物17种,其中萜烯类化合物7种(均是倍半萜),羧酸类化合物2种,醇类、酯类及酚类化合物各1种,还有1种烷烃和1种烯烃类化合物(表4)。17种化合物中,相对含量最高的为烯烃类化合物,相对含量占比达66.62%,其次是倍半萜(27.98%),而7种倍半萜中以α-葎草烯含量最高,占16.49%(表4)。
表 4 雷竹笋挥发物的成分及其相对含量
Table 4. Relative content (%) and volatile components of Ph. violascens shoot collected by HS-SPME
物质峰编号
No. of
peak保留时间
Retention
time/min保留指数
Retention
indices/iu分子式
Molecular
formula化合物名称
Compounds相对含量±标准误
Relative content ± SE/%1 9.12 2705 C20H40O 1-(乙烯氧基)-十八烷 1-(ethenyloxy)-octadecane 0.68±0.21 2 10.11 1180 C9H10O 4-(1-甲基乙烯基)-苯酚 4-(1-methylethenyl)-phenol 0.19±0.03 3 10.65 1471 C11H22O2 十一烷酸 undecanoic acid 0.20±0.05 4 10.96 1620 C16H32 (Z)-7-十六碳烯 (Z)-7-hexadecene 2.58±0.42 5 11.04 1413 C14H30 tetradecane 十四烷 0.30±0.06 6 11.28 1430 C15H24 α-香柠檬烯 α-bergamotene 0.88±0.19 7 11.39 1494 C15H24 β-石竹烯 β-caryophyllene 3.07±1.87 8 11.60 1440 C15H24 (Z)-β-farnesene (Z)-β-法呢烯 0.42±0.11 9 11.64 1446 C15H24 β-倍半水芹烯 β-sesquiphellandrene 1.16±0.08 10 11.71 1579 C15H24 α-葎草烯 α-humulene 16.49±3.25 11 11.88 1304 C13H26 1-十三碳烯 1-tridecene 64.04±8.23 12 12.11 1500 C15H24 β-双油烯 β-bisabolene 2.84±0.98 13 12.23 1435 C15H24 γ-杜松烯 γ-cadinene 3.12±1.01 14 12.84 1790 C16H34O 2-己基-1-癸醇 2-hexyl-1-decanol 0.41±0.03 15 12.91 1910 C19H40 十九烷 nonadecane 0.84±0.92 16 17.43 1908 C16H22O4 邻苯二甲酸二异丁酯 diisobutyl phthalate 1.96±0.97 17 18.56 1968 C16H32O2 十六烷酸 hexadecanoic acid 0.82±0.31
筛胸梳爪叩甲幼虫对竹笋挥发物及CO2的行为反应
Larval Behavioural Responses of Melanotus cribricollis (Coleoptera: Elateridae) to the Volatiles Emitted from Bamboo Shoot and Carbon Dioxide
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摘要:
目的 探析南方竹区重要地下害虫筛胸梳爪叩甲幼虫定位寄主的行为及化学机制,为竹林金针虫的高效监测及防控提供科学依据。 方法 利用双向选择仪测定筛胸梳爪叩甲幼虫对竹笋和CO2释放剂颗粒的行为反应。采用固相微萃取气质联用法(SPME-GC-MS)收集鉴定雷竹鲜笋的挥发性组分,并测定其相对含量。 结果 竹笋和CO2释放剂均能显著引发筛胸梳爪叩甲幼虫定向运动。筛胸梳爪叩甲幼虫对不同质量鲜笋的取食选择无显著差异(P>0.05),而不同浓度的CO2释放剂对筛胸梳爪叩甲幼虫的定向行为有显著影响(P<0.05)。在竹笋和CO2释放剂之间,筛胸梳爪叩甲幼虫显著偏好竹笋(P<0.05)。收集鉴定雷竹笋挥发物17种,相对含量较高为烯烃类化合物(66.62%)和倍半萜(27.98%),倍半萜中以α-葎草烯含量最高。 结论 筛胸梳爪叩甲幼虫对新鲜雷竹笋和CO2释放剂颗粒均有显著趋向性,筛胸梳爪叩甲幼虫更偏好新鲜竹笋,相比CO2释放剂,竹笋萌发时所释放的挥发物对竹林金针虫搜寻寄主更为有效。本研究分离鉴定7种可能对筛胸梳爪叩甲幼虫具有吸引作用的萜烯类化合物,为竹林金针虫新型防控技术研发提供了重要依据。 Abstract:Objective To understand the mechanism of food foraging below ground of Melanotus cribricollis larva which is the dominant and most destructive soil-dwelling insect of bamboo shoots in south China and provide important basic information for the development of effective monitor and control techniques. Method We tested the orientation behavior of M. cribricollis larvae towards bamboo shoot and CO2-releasing capsules by using a dual-choice olfactometer. The volatiles emitted from Phyllostachys violascens shoot were tentatively identified by solid phase micro-extraction (SPME) coupled with GC-MS, and their relative content were determined as well. Result Both bamboo shoot and CO2 source could significantly induce the directional movement of the larvae of M. cribricollis. There was no significant difference in the larval behavior among different weights of bamboo shoots (P>0.05), whiel larval orientation behavior was significantly correlated with CO2 concentration (P<0.05). Between bamboo and CO2-releasing capsules, wireworms preferred significantly to the bamboo shoot (P<0.05). A total of 17 compounds emitted from Ph. violascens shoot were tentatively identified. The higher relative contents were olefins (66.62%) and sesquiterpenes (27.98%), and the sesquiterpene compound with the highest content was α-humulene. Conclusion M. cribricollis larvae shows significant stronger orientation towards bamboo shoot and CO2 source, and prefers bamboo shoot significantly. Compared with CO2 source, the volatile organic compounds (VOCs) emitted from bamboo shoot show to be more effective in food foraging of bamboo shoot wireworm. In addition, 7 terpene compounds with potential attraction to wireworms are identified in this study, which provids important basic information for the development of attract-and-kill formulations for the control of wireworms. -
Key words:
- wireworm
- / root herbivore
- / carbon dioxide
- / VOCs
- / attractant.
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表 1 筛胸梳爪叩甲幼虫对不同质量竹笋的选择
Table 1. Choice tests of M. cribricollis larvae towards different weight bamboo shoots in a dual-choice olfactometer
处理
Treatments重复
Replicates作出选择的金针虫数量(百分比)
Number of wireworms at different sides(Percentage)G检验
G test处理端
Treated side对照端
Control sideGP GH 10 g 20 128(82.05%) 28(17.95%) 69.43*** 21.10NS 20 g 20 118(81.38%) 27(18.62%) 61.62*** 23.64NS 40 g 20 136(84.47%) 25(15.53%) 84.17*** 8.74NS 注:仅对作出选择的金针虫个体数进行G检验。***表示P<0.001,NS表示差异不显著(P>0.05)。下同。
Note:G test was performed only on the number of wireworms at the treated and control sides. *** means P<0.001,NS means that the difference is not significant(P>0.05). The same followed.表 2 筛胸梳爪叩甲幼虫对不同浓度CO2的选择
Table 2. Choice tests of M. cribricollis larvae towards CO2 of different concentrations in a dual-choice olfactometer
处理
Treatments重复
Replicates作出选择的金针虫数量(百分比)
Number of wireworms at different sides(Percentage)G检验
G-test处理端
Treated side对照端
Control sideGP GH T1 20 99(61.49%) 62(38.50%) 8.58** 9.21NS T2 20 107(75.89%) 34(24.11%) 36.70*** 22.22NS T3 20 107(71.81%) 42(28.19%) 29.33*** 11.82NS 注:T1、T2、T3分别表示10、20和40粒CO2释放剂颗粒,下同。**表示P<0.01。
Note: T1, T2 and T3 presents 10, 20 and 40 granules of CO2 resource, respectively. The same followed. ** means P<0.01.表 3 筛胸梳爪叩甲幼虫在竹笋和CO2释放物间的选择
Table 3. Choice tests of M. cribricollis larvae between bamboo shoots and CO2 resource in a dual-choice olfactometer
处理
Treatments重复
Replicates作出选择的金针虫数量(百分比)
Number of wireworms at different sides(Percentage)G检验
G-test竹笋
Bamboo shoots二氧化碳
CO2GP GH 24 h 20 139(80.35%) 34(19.65%) 68.37 *** 15.40 NS 48 h 20 142(80.23%) 35(19.77%) 47.69 *** 14.58 NS 表 4 雷竹笋挥发物的成分及其相对含量
Table 4. Relative content (%) and volatile components of Ph. violascens shoot collected by HS-SPME
物质峰编号
No. of
peak保留时间
Retention
time/min保留指数
Retention
indices/iu分子式
Molecular
formula化合物名称
Compounds相对含量±标准误
Relative content ± SE/%1 9.12 2705 C20H40O 1-(乙烯氧基)-十八烷 1-(ethenyloxy)-octadecane 0.68±0.21 2 10.11 1180 C9H10O 4-(1-甲基乙烯基)-苯酚 4-(1-methylethenyl)-phenol 0.19±0.03 3 10.65 1471 C11H22O2 十一烷酸 undecanoic acid 0.20±0.05 4 10.96 1620 C16H32 (Z)-7-十六碳烯 (Z)-7-hexadecene 2.58±0.42 5 11.04 1413 C14H30 tetradecane 十四烷 0.30±0.06 6 11.28 1430 C15H24 α-香柠檬烯 α-bergamotene 0.88±0.19 7 11.39 1494 C15H24 β-石竹烯 β-caryophyllene 3.07±1.87 8 11.60 1440 C15H24 (Z)-β-farnesene (Z)-β-法呢烯 0.42±0.11 9 11.64 1446 C15H24 β-倍半水芹烯 β-sesquiphellandrene 1.16±0.08 10 11.71 1579 C15H24 α-葎草烯 α-humulene 16.49±3.25 11 11.88 1304 C13H26 1-十三碳烯 1-tridecene 64.04±8.23 12 12.11 1500 C15H24 β-双油烯 β-bisabolene 2.84±0.98 13 12.23 1435 C15H24 γ-杜松烯 γ-cadinene 3.12±1.01 14 12.84 1790 C16H34O 2-己基-1-癸醇 2-hexyl-1-decanol 0.41±0.03 15 12.91 1910 C19H40 十九烷 nonadecane 0.84±0.92 16 17.43 1908 C16H22O4 邻苯二甲酸二异丁酯 diisobutyl phthalate 1.96±0.97 17 18.56 1968 C16H32O2 十六烷酸 hexadecanoic acid 0.82±0.31 -
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