Feeding choice behavior and mechanism of adult Meliboeus ohbayashii primoriensis
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摘要:目的 通过比较核桃缘吉丁成虫对核桃、苹果和山楂这3种植物在取食选择和取食量上的差异，并分析植物挥发物的组成与含量，该研究为进一步开发核桃缘吉丁植物源引诱剂提供了依据。方法 测定核桃缘吉丁成虫对核桃、苹果和山楂叶片的取食选择和取食量，采用动态顶空法收集这3种植物的枝叶挥发物组分及相对含量的测定，通过卡方检验分析比较核桃缘吉丁对植物挥发物的行为反应。结果 核桃缘吉丁成虫对不同植物的取食选择和取食量存在显著差异，成虫对核桃的取食选择性最高且取食核桃叶片，对苹果和山楂的选择性较低，且存在不取食现象。从核桃、苹果和山楂的枝叶挥发物中，共收集到45种植物挥发性化合物，包括烷烃类5种、酯类8种、芳香烃类3种、萜类18种、醛类2种、醇类4种和酮类5种。在行为反应测定中，莰烯对核桃缘吉丁雌雄成虫均具有明显的趋向反应，而乙酸冰片酯、癸醛和桉叶油醇这3种单体化合物仅对雄虫具有一定的引诱效果。结论 核桃可作为核桃缘吉丁成虫补充营养的寄主植物，该成虫不取食苹果和山楂。莰烯、乙酸冰片酯、癸醛和桉叶油醇对核桃缘吉丁成虫具有一定的吸引效果，需要重点关注，这些枝叶挥发物成分在核桃缘吉丁成虫寄主植物识别与选择的阶段中可能发挥着重要作用。Abstract:Objective By comparing differences in the selection and consumption of Meliboeus ohbayashii primoriensis to 3 kinds of plants (Juglans regia, Malus pumila and Crataegus pinnatifida), and by analyzing the composition and content of plant volatiles, this research provides a solid basis for a further development of the plant source attractant for M. ohbayashii primoriensis.Method The leaves of three plants (J. regia, M. pumila and C. pinnatifida) were used to determine host plant choice and consumption by this insect pest. Volatile components in the branches and leaves of these three plants were collected using the dynamic headspace method and were analyzed by gas chromatography-mass spectrometry. Furthermore, Chi-square test was used to analyze and compare the differences in behavioral responses of M. ohbayashii primoriensis adults in the volatiles emitted by these plants.Result Significant differences were found in the selection and consumption of the three plants by M. ohbayashii primoriensis. Adults displayed high feeding selectivity for J. regia and consumed J. regia leaves. Whereas they showed low feeding selectivity for M. pumila and C. pinnatifida, without consuming their leaves. A total of 45 plant volatile compounds were identified from the branches and leaves of these three plants, including five alkanes, eight esters, three aromatic hydrocarbons, 18 terpenes, two aldehydes, four alcohols and five ketones. In the experiment of behavioral responses, both male and female adults of M. ohbayashii primoriensis were attracted by camphene in obvious tendency responses, while only males were attracted by bornyl acetate, decanal and eucalyptol in a certain attraction.Conclusion J. regia can be used as a host plant to supplement the nutrition of adult M. ohbayashii primoriensis, whereas the leaves of M. pumila and C. pinnatifida are not suitable for consumption. Camphene, bornyl acetate, decanal and eucalyptol had a certain attraction effect on the adults M. ohbayashii primoriensis. These volatile components may play an important role during the identification and selection of host plants by adult M. ohbayashii primoriensis. These volatile components of branches and leaves may play an important role during the identification and selection of host plants by adult M. ohbayashii primoriensis.
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图 1 核桃缘吉丁成虫在3种植物叶片上的着落数量
不同小写字母表示不同植物间存在极显著差异(P < 0.001)，χ2 = 81.750，df = 2，渐进显著性 = 0.000。Different lowercases indicate highly significant differences between plants (P < 0.001), χ2 = 81.750, df = 2, Asymp. Sig. = 0.000.
Figure 1. Number of Meliboeus ohbayashii primoriensis adults observed on foliage of three plants
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图 2 3种植物叶片面积与质量间的线性关系
Figure 2. Regression of leaf area and leaf mass among three plants
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图 3 无竞争条件下核桃缘吉丁成虫(雌虫和雄虫)对3种植物叶片的取食面积和质量
不同小写字母表示不同植物间存在极显著差异(P < 0.001)，χ2 = 16.129，df = 2，渐进显著性 = 0.000。Different lowercases indicate highly significant differences between plants (P < 0.001), χ2= 16.129, df = 2, Asymp. Sig. = 0.000.
Figure 3. Leaf area and leaf mass consumed by M. ohbayashii primoriensis adults (female and male) on three plants under non-choice conditions
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图 4 两项选择条件下核桃缘吉丁成虫(雌虫和雄虫)对3种植物叶片的取食面积
*表示不同植物间存在显著差异(P < 0.05)，Z = −2.201，渐进显著性(双侧) = 0.028。下同。* indicates significant differences between plants (P < 0.05), Z = −2.201, Asymp.Sig. (2-tailed) = 0.028. The same below.
Figure 4. Leaf area consumed by M. ohbayashii primoriensis adults (female and male) on three plants under two-choice conditions
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图 5 两项选择条件下核桃缘吉丁成虫(雌虫和雄虫)对3种植物叶片的取食质量
Figure 5. Leaf mass consumed by female M. ohbayashii primoriensis adults (female and male) on three plants under two-choice conditions
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图 6 核桃缘吉丁成虫(雌虫A和雄虫B)对植物挥发物的行为反应
*表示不同植物挥发物与对照之间存在显著差异(P < 0.05)。* indicates significant differences between plant volatiles and control (P < 0.05).
Figure 6. Behavioral responses of Meliboeus ohbayashii primoriensis adults (female A and male B) to plant volatiles
表 1 3种植物枝叶的主要挥发物成分及相对含量
Table 1 Composition and relative contents of main volatile released from the branch and foliage of three plants
种类 Type 植物挥发物 Plant volatile 相对含量 Relative content/% 核桃 J. regia 苹果 M. pumila 山楂 C. pinnatifida 烷烃类 Alkanes 十二烷 Dodecane 0.20 ± 0.03 0.10 ± 0.06 十四烷 Tetradecane 0.07 ± 0.01 0.18 ± 0.02 0.11 ± 0.04 十五烷 Pentadecane 0.11 ± 0.03 0.17 ± 0.04 0.10 ± 0.03 十六烷 Hexadecane 0.08 ± 0.25 十七烷 Heptadecane 0.11 ± 0.02 0.09 ± 0.01 酯类 Esters 碳酸二乙酯 Diethyl carbonate 0.58 ± 0.07 1.36 ± 0.70 甲酸叶醇酯 cis-3-hexenyl formate 1.43 ± 0.11 3.42 ± 0.40 乙酸叶醇酯 Leaf acetate 5.37 ± 0.45 9.92 ± 3.17 丙烯酸异辛酯 2-ethylhexyl acrylate 1.53 ± 0.17 7.40 ± 1.48 4.60 ± 1.95 乙酸冰片酯 Bornyl acetate 0.39 ± 0.11 异戊酸香叶基酯 Geranyl isovalerate 0.52 ± 0.09 苯乙酸叶醇酯 (Z)-hex-3-enyl phenylacetate 1.17 ± 0.21 棕榈酸甲酯 Methyl hexadecanoate 0.80 ± 0.05 芳香烃类 Aromatic hydrocarbons 邻二甲苯 o-xylene 0.21 ± 0.04 精萘 Naphthalene 0.77 ± 0.34 2.48 ± 0.34 1,6-己内酰胺 1, 6-caprolactams 2.28 ± 0.30 萜类 Terpenes α-蒎烯 α-pinene 23.43 ± 0.76 10.62 ± 0.10 14.12 ± 1.38 莰烯 Camphene 0.34 ± 0.10 桧稀 Sabenene 1.90 ± 0.38 月桂烯 β-myrcene 1.39 ± 0.17 β-蒎烯 β-pinene 2.81 ± 0.26 樟脑 (+)-2-bornanone 3.84 ± 0.43 右旋萜二烯 D-limonene 9.15 ± 0.09 长叶环烯 Longicyclene 0.63 ± 0.13 2.30 ± 0.43 2.57 ± 0.82 β-波旁烯 (-)-β-bourbonene 2.64 ± 0.38 榄香烯 (-)-b-elemene 0.63 ± 0.19 长叶烯 Longifolene 1.24 ± 0.16 10.41 ± 0.22 7.42 ± 1.23 β-石竹烯 β-caryophyllen 6.70 ± 0.43 3.76 ± 0.12 0.73 ± 0.18 (E)-β-金合欢烯 (E)-β-farnesene 11.89 ± 0.89 大根香叶烯 Germacrene D 1.26 ± 0.37 α-法尼烯 (E,E)-α-farnesene 5.27 ± 1.42 5.54 ± 1.55 法尼醇 Farnesanol 0.79 ± 0.07 反式角鲨烯 Squalene 0.85 ± 0.14 1.89 ± 1.01 氧化石竹烯 (-)-caryophyllene oxide 1.93 ± 0.35 醛类 Aldehydes 壬醛 1-nonmanal 1.33 ± 0.34 5.44 ± 0.56 3.99 ± 1.48 癸醛 Decanal 0.42 ± 0.09 2.87 ± 0.94 2.97 ± 0.86 醇类 Alcohols 桉叶油醇 Eucalyptol 6.42 ± 1.05 2-乙基己醇 2-ethylhexanol 11.91 ± 1.19 16.93 ± 3.11 4-萜烯醇 Terpinen-4-ol 0.28 ± 0.09 1.51 ± 0.39 1.37 ± 0.34 alpha-松油醇 alpha-terpineol 2.17 ± 0.29 酮类 Ketones 松香芹酮 Pinocarvone 1.15 ± 0.11 反式松莰酮 (E)-pinocamphone 0.30 ± 0.07 香叶基丙酮 Geranylacetone 14.99 ± 1.89 对乙基苯乙酮 Ethylacetophenone 1.90 ± 0.21 异佛尔酮 Isophorone 12.28 ± 2.11 
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