长足大竹象的食物识别信号和幼虫种群变化规律研究
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摘要
竹材是造纸的主要原料和林区竹农的经济支柱,竹产业是当前发展的朝阳产业。长足大竹象(Cyrtotrachelus bugueti Guer)是当前丛生竹林的主要害虫,严重制约着竹材产量的提高和农民经济收入的增加。国家林业局2003年将长足大竹象列为我国首批林业危险性有害生物之一。
本研究采用气相色谱-质谱联用、触角电位反应及电镜扫描等多种技术手段,研究了竹林主要害虫长足大竹象的食物识别信号,明确了长足大竹象嗅觉感器的超微结构及不同类型竹笋挥发物中对长足大竹象发生作用的成分。在长足大竹象危害地调查了长足大竹象幼虫种群动态,应用SPSS16.0软件分析了长足大竹象幼虫种群变化与气候因子的关系,揭示了长足大竹象幼虫种群发生规律及其与气候、竹笋的内在统一性。本研究将长足大竹象幼虫密度的气候影响因子和食物识别信号结合起来,利用长足大竹象与竹笋间的自然化学作用在幼虫高峰期之前控制竹林主要害虫,以期为实现害虫防治、昆虫资源利用、环境无污染三者的协调统一提供理论依据。主要结果如下：
1长足大竹象头部感器的超微结构
电镜观察表明长足大竹象头部感器主要集中在触角和口器,触角感器主要集中在鞭节的末亚节,占感器的95%以上。触角上共发现了5类感器,分别为锥形感器、毛形感器、柱形感器、芽形感器、手形感器。触角以具有嗅觉功能的锥形感器最多,密度最大,其次是具有信息化合物感受功能的毛形感器。口器上分布大量的具味觉功能的长柱毛形感器。研究结果表明长足大竹象成虫对竹笋的识别主要通过鞭节末亚节上的锥形感器。
2竹笋挥发物成分测定和长足大竹象触角电位反应
气质色谱联用在竹笋挥发性物质中检测到28种物质,主要为醛类、醇类、酸类和脂肪酸的衍生物,醛类物质相对含量最高,其次是醇类和酸类物质。竹笋不同类型、同一竹笋不同部位挥发物的组分不同,丛生慈竹笋尖挥发物中以苯甲醛、芳樟醇、对羟基苯甲醛、氯己酸的含量较高,丛生慈竹笋基部挥发物以对羟基苯甲醛、2,5,8,11,14-五氧杂-16-十六烷醇、(3-甲基-环氧-2-丙烷基)-甲醇、苯甲醛的含量较高,散生苦竹笋尖挥发物中以对羟基苯甲醛、十六酸、亚油酸、(3β)-豆甾-5-烯-3-醇的含量较高,丛生慈竹笋笋尖挥发物中特有的和含量较高的成分可能是吸引长足大竹象取食产卵的重要物质。
选择含量较高的20种竹笋挥发物成分对长足大竹象成虫做触角电位测定,结果表明竹笋挥发性物质的不同组分对成虫有不同的电生理活性,其中苯甲醛、氯己酸、芳樟醇等6种成分是引起长足大竹象显著电生理活性的重要挥发物质。用此6种挥发物诱捕长足大竹象成虫的结果表明,苯甲醛诱捕长足大竹象成虫数量最多且差异达到极显著水平。研究表明长足大竹象很可能是根据以苯甲醛为主要成分构成的、具有特定成分和比例的化学图谱识别竹笋。
3长足大竹象幼虫种群变化规律研究
长足大竹象地上生活史与温度显著相关。2004-2008年于四川沐川定期调查了3个受害慈竹林内所有竹笋上的幼卵和幼虫密度变化,运用主成分、相关和通径分析及决策系数研究了影响幼虫种群变化的主要气候因子,并用8种不同的函数模型对虫口密度与旬平均气温、旬最低气温的关系进行拟合。结果表明,幼虫种群变化呈单峰型,7月中旬以前虫口数量较低,8月为高峰期,9月中旬以后虫口密度显著降低,10月上旬以后虫口密度逐渐趋向于0水平。4种分析表明,长足大竹象幼虫种群变化最主要的限制因子是旬平均气温,最主要的决策因素是旬最低气温。8种函数拟合表明平均虫口密度与旬平均气温、旬最低气温之间分别用直线函数方程Y=-0.535+0.031X、对数函数方程Y=-1.594+0.6001nX拟合的效果最好,据此建立了平均虫口密度气候预测表,提出了长足大竹象防治的温度指标：旬平均温度21.5℃或旬最低温度17.70C。
4降雨量、气温、竹笋与长足大竹象发生的内在统一性
在长足大竹象地上活动期间,7月较多的降雨量有利于成虫出土,8月较高的气温促进竹笋的生长,丰富的食源促进长足大竹象的生长繁殖,从而形成8月长足大竹象幼虫危害的高峰,此表明降雨量、气温、竹笋、长足大竹象具有内在统一性。
Bamboo was the main raw material for paper making industry and the key economic resourse for bamboo farmers, and bamboo industry had been developed into a sunrise industry. Cyrtotrachelus bugueti, the main bamboo forest pests of cluster bamboo forest, had seriously constrained the improvement of bamboo produce and the income increase of bamboo farmers. The insect had been taken as one of dangerous forestry pests by the State Forestry Administration since 2003.
The food identification signal of C. bugueti, including its ultrastructure of touch sensilla and smell sensilla, as well as the volatiles effecting on C. bugueti from different varieties and types of bamboo shoots, were studied by using the scanning electron microscopy, the gas chromatography-mass spectrometry (GC-MS), the elctroantennogram (EAG) technique respectively. The larvae population dynamics of C. bugueti and its relationship to climate factors were analyzed too. The study combined climate factor effecting on larvae density with food identification signal of C. bugueti for controlling the bamboo pest by the nature chemistry between C. bugueti and bamboo shoot.This study provided theoretical basis for harmony among control of bamboo pest, utilization of insect resource and protection of environment. The results were as follow:
1 Antenna ultrastructure and food identification sensilla of C. bugueti
Abservation using scanning electron microscopy showed that the head sensilla of C. bugueti mainly distributed in antenna and mouthpart, and the antenna consists of the scape, pedicel and seven flagellomeres. The sensillaes of flagellomeres in the end of antenna were over 95%of antenna sensilla. In antenna, one type of sensilla basiconic, two type of sensilla trichodea, one type of sensilla column, one type of bud-shaped sensilla and one type of hand-shaped sensilla were distinguished with the shapes and distribution of sensilla. The sensilla basiconic with olfactory function were the largest density sensilla, the second was the sensilla trichodea with the pheromones feeling function. Mouthparts also distributed a large number of long columns sensilla trichodea with taste function. The results showed that volatiles from bamboo shoots were apperceived mainly through the sensilla basiconica in flagellomeres of antenna.
2 Study on relationship between volatile components of bamboo shoots and C. bugueti
28 bamboo shoot volatiles were discovered by GC-MS, and the results showed that the volatiles from the tine of cluster bamboo shoot were with high content of benzaldehyde, linalool,4-hydroxy-benzaldenhyde and 8-chlorocapric acid, the volatiles from the bottom of cluster bamboo shoot were with high content of 4-hydroxy-benzaldenhyde,2,5, 8,11,14-pentaoxahexadecan, (3-methyl-oxiran-2-yl)-methanol, benzaldehyde, and the volatiles from the tine of scattered bamboo shoot were with high content of 4-hydroxy-benzaldenhyde, hexadecanoic acid, linoleic acid and stigmast-5-en-3-ol (3.beta.24s). The results suggested that the high content or the special volatiles of the tine of cluster bamboo shoot might be the important substance attracting C. bugueti.
EAG reaction suggested that the antenna of C. bugueti imago reacting to the different volatiles from bamboo shoots showed different electrophysiological reaction, and benzaldehyde, linalool and 8-chlorocapric acid and so on could caused C. bugueti imago significant electrophysiological reaction. With them trapping C. bugueti imago, the results showed that benzaldehyde could attracted the most imagoes. The study suggested that C. buqueti might identified the bamboo shoot by the chemistry finger map with primary benzaldehyde and special proportion of components.
3 Overground propagatation of C. buqueti was markedly impacted by temperature. From 2004 to 2008, the eggs and larvae of C. buqueti in all bamboo shoots of three bamboo forests were regularly investigated in the Muchuan Sichuan. The principal component analysis, as well as correlation and path analysis and decision coefficient, was exerted to study the effect of main climate factors on larvae population dynamics, and eight functions were used to analyze the relationship among C. buqueti larval density and mean temperature and minimum temperature. The results showed that the larvae population fluctuation was single peaked, with the peak in August. The larvae population was low before mid-July and evidently reduced after mid-September and gradually tended to zero level after early October. The results of four analyzing methods suggested that the most important limiting-factor and the most important decision factor of larvae population fluctuation was respectively the mean temperature and the minimum temperature, two forecasting models for the larvae population fluctuation were established, i.e., C. buqueti larvae density (Y) and mean temperature(X) follow the equation of Y=-0.535+0.031X, and C. buqueti larvae density(Y) and minimum temperature (X) follow the equation of Y=-1.594+0.6001nX. Based on these results, the forecast tables for C. buqueti larvae density were provided, and for controlling larvae density of C. buqueti, mean temperature 21.5℃of ten days and minimum temperature 17.7℃often days were suggested too.
4 The emergence of C. bugueti had the intrinsic unity with rainfall, temperature and bamboo shoots. During overground propagatation of C. buqueti, the growth of bamboo shoot was facilitated by the more rainfall of July and the higher temperature of August, and provided abundand food for C. buqueti. Therefore, the high larvae population came into being.
本研究采用气相色谱-质谱联用、触角电位反应及电镜扫描等多种技术手段,研究了竹林主要害虫长足大竹象的食物识别信号,明确了长足大竹象嗅觉感器的超微结构及不同类型竹笋挥发物中对长足大竹象发生作用的成分。在长足大竹象危害地调查了长足大竹象幼虫种群动态,应用SPSS16.0软件分析了长足大竹象幼虫种群变化与气候因子的关系,揭示了长足大竹象幼虫种群发生规律及其与气候、竹笋的内在统一性。本研究将长足大竹象幼虫密度的气候影响因子和食物识别信号结合起来,利用长足大竹象与竹笋间的自然化学作用在幼虫高峰期之前控制竹林主要害虫,以期为实现害虫防治、昆虫资源利用、环境无污染三者的协调统一提供理论依据。主要结果如下：
1长足大竹象头部感器的超微结构
电镜观察表明长足大竹象头部感器主要集中在触角和口器,触角感器主要集中在鞭节的末亚节,占感器的95%以上。触角上共发现了5类感器,分别为锥形感器、毛形感器、柱形感器、芽形感器、手形感器。触角以具有嗅觉功能的锥形感器最多,密度最大,其次是具有信息化合物感受功能的毛形感器。口器上分布大量的具味觉功能的长柱毛形感器。研究结果表明长足大竹象成虫对竹笋的识别主要通过鞭节末亚节上的锥形感器。
2竹笋挥发物成分测定和长足大竹象触角电位反应
气质色谱联用在竹笋挥发性物质中检测到28种物质,主要为醛类、醇类、酸类和脂肪酸的衍生物,醛类物质相对含量最高,其次是醇类和酸类物质。竹笋不同类型、同一竹笋不同部位挥发物的组分不同,丛生慈竹笋尖挥发物中以苯甲醛、芳樟醇、对羟基苯甲醛、氯己酸的含量较高,丛生慈竹笋基部挥发物以对羟基苯甲醛、2,5,8,11,14-五氧杂-16-十六烷醇、(3-甲基-环氧-2-丙烷基)-甲醇、苯甲醛的含量较高,散生苦竹笋尖挥发物中以对羟基苯甲醛、十六酸、亚油酸、(3β)-豆甾-5-烯-3-醇的含量较高,丛生慈竹笋笋尖挥发物中特有的和含量较高的成分可能是吸引长足大竹象取食产卵的重要物质。
选择含量较高的20种竹笋挥发物成分对长足大竹象成虫做触角电位测定,结果表明竹笋挥发性物质的不同组分对成虫有不同的电生理活性,其中苯甲醛、氯己酸、芳樟醇等6种成分是引起长足大竹象显著电生理活性的重要挥发物质。用此6种挥发物诱捕长足大竹象成虫的结果表明,苯甲醛诱捕长足大竹象成虫数量最多且差异达到极显著水平。研究表明长足大竹象很可能是根据以苯甲醛为主要成分构成的、具有特定成分和比例的化学图谱识别竹笋。
3长足大竹象幼虫种群变化规律研究
长足大竹象地上生活史与温度显著相关。2004-2008年于四川沐川定期调查了3个受害慈竹林内所有竹笋上的幼卵和幼虫密度变化,运用主成分、相关和通径分析及决策系数研究了影响幼虫种群变化的主要气候因子,并用8种不同的函数模型对虫口密度与旬平均气温、旬最低气温的关系进行拟合。结果表明,幼虫种群变化呈单峰型,7月中旬以前虫口数量较低,8月为高峰期,9月中旬以后虫口密度显著降低,10月上旬以后虫口密度逐渐趋向于0水平。4种分析表明,长足大竹象幼虫种群变化最主要的限制因子是旬平均气温,最主要的决策因素是旬最低气温。8种函数拟合表明平均虫口密度与旬平均气温、旬最低气温之间分别用直线函数方程Y=-0.535+0.031X、对数函数方程Y=-1.594+0.6001nX拟合的效果最好,据此建立了平均虫口密度气候预测表,提出了长足大竹象防治的温度指标：旬平均温度21.5℃或旬最低温度17.70C。
4降雨量、气温、竹笋与长足大竹象发生的内在统一性
在长足大竹象地上活动期间,7月较多的降雨量有利于成虫出土,8月较高的气温促进竹笋的生长,丰富的食源促进长足大竹象的生长繁殖,从而形成8月长足大竹象幼虫危害的高峰,此表明降雨量、气温、竹笋、长足大竹象具有内在统一性。
Bamboo was the main raw material for paper making industry and the key economic resourse for bamboo farmers, and bamboo industry had been developed into a sunrise industry. Cyrtotrachelus bugueti, the main bamboo forest pests of cluster bamboo forest, had seriously constrained the improvement of bamboo produce and the income increase of bamboo farmers. The insect had been taken as one of dangerous forestry pests by the State Forestry Administration since 2003.
The food identification signal of C. bugueti, including its ultrastructure of touch sensilla and smell sensilla, as well as the volatiles effecting on C. bugueti from different varieties and types of bamboo shoots, were studied by using the scanning electron microscopy, the gas chromatography-mass spectrometry (GC-MS), the elctroantennogram (EAG) technique respectively. The larvae population dynamics of C. bugueti and its relationship to climate factors were analyzed too. The study combined climate factor effecting on larvae density with food identification signal of C. bugueti for controlling the bamboo pest by the nature chemistry between C. bugueti and bamboo shoot.This study provided theoretical basis for harmony among control of bamboo pest, utilization of insect resource and protection of environment. The results were as follow:
1 Antenna ultrastructure and food identification sensilla of C. bugueti
Abservation using scanning electron microscopy showed that the head sensilla of C. bugueti mainly distributed in antenna and mouthpart, and the antenna consists of the scape, pedicel and seven flagellomeres. The sensillaes of flagellomeres in the end of antenna were over 95%of antenna sensilla. In antenna, one type of sensilla basiconic, two type of sensilla trichodea, one type of sensilla column, one type of bud-shaped sensilla and one type of hand-shaped sensilla were distinguished with the shapes and distribution of sensilla. The sensilla basiconic with olfactory function were the largest density sensilla, the second was the sensilla trichodea with the pheromones feeling function. Mouthparts also distributed a large number of long columns sensilla trichodea with taste function. The results showed that volatiles from bamboo shoots were apperceived mainly through the sensilla basiconica in flagellomeres of antenna.
2 Study on relationship between volatile components of bamboo shoots and C. bugueti
28 bamboo shoot volatiles were discovered by GC-MS, and the results showed that the volatiles from the tine of cluster bamboo shoot were with high content of benzaldehyde, linalool,4-hydroxy-benzaldenhyde and 8-chlorocapric acid, the volatiles from the bottom of cluster bamboo shoot were with high content of 4-hydroxy-benzaldenhyde,2,5, 8,11,14-pentaoxahexadecan, (3-methyl-oxiran-2-yl)-methanol, benzaldehyde, and the volatiles from the tine of scattered bamboo shoot were with high content of 4-hydroxy-benzaldenhyde, hexadecanoic acid, linoleic acid and stigmast-5-en-3-ol (3.beta.24s). The results suggested that the high content or the special volatiles of the tine of cluster bamboo shoot might be the important substance attracting C. bugueti.
EAG reaction suggested that the antenna of C. bugueti imago reacting to the different volatiles from bamboo shoots showed different electrophysiological reaction, and benzaldehyde, linalool and 8-chlorocapric acid and so on could caused C. bugueti imago significant electrophysiological reaction. With them trapping C. bugueti imago, the results showed that benzaldehyde could attracted the most imagoes. The study suggested that C. buqueti might identified the bamboo shoot by the chemistry finger map with primary benzaldehyde and special proportion of components.
3 Overground propagatation of C. buqueti was markedly impacted by temperature. From 2004 to 2008, the eggs and larvae of C. buqueti in all bamboo shoots of three bamboo forests were regularly investigated in the Muchuan Sichuan. The principal component analysis, as well as correlation and path analysis and decision coefficient, was exerted to study the effect of main climate factors on larvae population dynamics, and eight functions were used to analyze the relationship among C. buqueti larval density and mean temperature and minimum temperature. The results showed that the larvae population fluctuation was single peaked, with the peak in August. The larvae population was low before mid-July and evidently reduced after mid-September and gradually tended to zero level after early October. The results of four analyzing methods suggested that the most important limiting-factor and the most important decision factor of larvae population fluctuation was respectively the mean temperature and the minimum temperature, two forecasting models for the larvae population fluctuation were established, i.e., C. buqueti larvae density (Y) and mean temperature(X) follow the equation of Y=-0.535+0.031X, and C. buqueti larvae density(Y) and minimum temperature (X) follow the equation of Y=-1.594+0.6001nX. Based on these results, the forecast tables for C. buqueti larvae density were provided, and for controlling larvae density of C. buqueti, mean temperature 21.5℃of ten days and minimum temperature 17.7℃often days were suggested too.
4 The emergence of C. bugueti had the intrinsic unity with rainfall, temperature and bamboo shoots. During overground propagatation of C. buqueti, the growth of bamboo shoot was facilitated by the more rainfall of July and the higher temperature of August, and provided abundand food for C. buqueti. Therefore, the high larvae population came into being.
引文
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