光肩星天牛成虫寄主选择“记忆”效应研究
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摘要
光肩星天牛Anoplophora glabripennis Motschulsky是重要的林木蛀干害虫,危害槭属Acer、杨属Populus、柳属Salix、榆属Ulums等20多个属的树木,对我国林业造成了巨大的损失。国内外众多学者已对光肩星天牛做了大量研究,但对光肩星天牛成虫寄主选择的机制尚未明确。本研究从类似于高等动物所特有的“记忆”角度出发,在感器种类与形态特征、化学生态学、酶学、行为学等方面,较系统地开展了光肩星天牛成虫对寄主树种选择和判断的机制研究,重要结果如下:
1、在天牛成虫对不同寄主树种挥发性物质的趋向试验方面:利用Y型嗅觉仪测定了从漳河柳、箭杆杨、合作杨和新疆杨4种不同抗性等级树种上羽化的光肩星天牛雌、雄成虫对这4种寄主树种和嗜好寄主树种复叶槭枝条气味的嗅觉反应。当以空气为对照时,源自4种不同寄主树种上的天牛成虫对其原寄主树种枝条挥发性气味均有明显的正趋性反应(P < 0.05)。但从合作杨、漳河柳和箭杆杨上羽化的天牛成虫对新疆杨枝条的挥发性气味表现出不同程度的负趋性反应;来源自4种寄主树上的光肩星天牛对复叶槭枝条的挥发性气味均具有不同程度的正趋性反应,但未达显著差异(P > 0.05)。当以复叶槭枝条为对照时,4种不同寄主上羽化的光肩星天牛对原寄主气味和复叶槭气味的趋向差异不大。表明在无嗜好寄主树种(如复叶槭)的前提下,天牛成虫可识别并趋向原寄主树种气味,但如有嗜好寄主树种存在,天牛成虫就不趋向原寄主树种的气味。
2、在天牛成虫的学习行为研究方面:利用Y型嗅觉仪研究了光肩星天牛成虫在嗜好寄主复叶槭和抗性寄主新疆杨上取食不同时间后的学习行为。结果表明,天牛成虫取食复叶槭1d,对复叶槭枝条的气味选择率明显提高,并且复叶槭枝条挥发性气味就对天牛成虫产生了明显的引诱作用(P < 0.05)。取食复叶槭2d,天牛成虫对复叶槭气味的选择率继续提高。取食3d和5d,天牛成虫对复叶槭气味的选择率维持在70%左右。表明天牛成虫取食复叶槭可以出现嗜食性诱导。
没有取食新疆杨的天牛成虫,新疆杨气味对天牛成虫有明显忌避作用(P < 0.05);取食新疆杨1d,新疆杨气味对天牛成虫的忌避作用不明显(P > 0.05);取食新疆杨2d,天牛成虫对新疆杨气味的选择率超过60%,取食新疆杨3d,新疆杨的枝条的气味对天牛产生明显的引诱作用(P < 0.05),取食新疆杨5d,天牛成虫对新疆杨气味的选择率维持在取食3d的水平。表明天牛成虫取食新疆杨后可以出现习惯性反应。说明取食的经历能够改变光肩星天牛成虫对气味的选择,也就是说天牛成虫有很强的学习记忆能力。
3、在天牛成虫林间对不同寄主树种的选择试验方面:利用5年生的复叶槭、漳河柳、箭杆杨、合作杨和新疆杨,在田间的铁丝网笼中按照拉丁方配置了试验林。研究了分别从上述5种寄主树种上羽化的光肩星天牛成虫对原寄主树种选择时是否存在“记忆”效应。
从栖息、取食、交尾、刻槽和产卵的等方面分析,从5种不同寄主树种上羽化的天牛成虫选择复叶槭的比例和频次显著高于其他树种,天牛由于受到嗜好寄主复叶槭的影响,没有对原寄主树种显示出“记忆”效应,因复叶槭是光肩星天牛的嗜食寄主树种,对天牛产生了吸引作用(包括嗜食性诱导),从而推测可能掩盖了天牛成虫对原寄主树种“记忆”。
如果移除嗜好寄主复叶槭,从栖息、取食和交尾情况看,从4种不同寄主树种上羽化的光肩星天牛在漳河柳树上的比例和频次明显高于其它3种寄主树种;从产卵情况看,尽管无规律性的结果,但选择在漳河柳树上产卵的数量仍然很高。总体来看,如移除嗜好寄主复叶槭,漳河柳则表现了较强的“引诱”作用。
4、在不同树种上的天牛各虫态的消化酶活力测定方面:以上述5种寄主树种上的光肩星天牛4龄幼虫、蛹(复叶槭除外)和刚羽化的成虫为对象,测定了幼虫、蛹和成虫体内的消化酶种类和活性。结果表明,光肩星天牛的幼虫和刚羽化成虫的消化道中均具有完整的消化酶系,存在8种消化酶,即淀粉酶、果胶酶、转化酶、麦芽糖酶、半纤维素酶、类胰蛋白酶、蛋白酶和脂肪酶。在天牛蛹中只检测到脂肪酶的活性,但活性非常低,未检测到其它7种酶的活性。
5种寄主树种上刚羽化成虫的8种消化酶的活性大小顺序,没有一种和相应寄主树种上幼虫的一致,说明成虫的消化酶和幼虫的消化酶活性大小相关性不强,也就是说光肩星天牛幼虫消化酶的活性,在经过蛹期变态后不会对成虫的消化酶活性产生直接影响。
5、天牛成虫头部主要感器类型和显微特征研究:利用环境扫描电镜对光肩星天牛雌、雄成虫的触角、下唇须和下颚须上的感受器进行了观察。结果表明,天牛成虫的触角上有5类13种常见的感觉器,其中5种刺形感器、2种毛形感器、4种锥形感器、1钟形感器和1种指形感器。天牛成虫的下唇须和下颚须上有6类17种常见的感受器,其中7种刺形感器、2种毛形感器、3种锥形感器、2种栓锥感器,2种腔锥感器和1种板形感器。根据这些感器的分布和形态特征,并结合以前的文献报道,分析了各种感受器可能的功能。
Anoplophora glabripennis (Motschulsky) is a major polyphagous wood-boring beetle in China that caused great losses to the forest of China. Its primary host species include Acer (maple), Populus (poplar), Salix (willow) and Ulums (elm) and is reported to feed on more than 20 genus host trees. A large number of research projects have focused on this beetle in last 20 years, but host-selection mechanism of adults is unclear. For this reason, this paper makes deep research on host-selection mechanism of adults with types and morphological characteristics of sensilla, chemical ecology, enzymology, and behavior of animal. The main results were as follows:
1. Behavioral responses of adults A. glabripennis to volatiles emanating from different host trees:
Behavioral responses of adults A. glabripennis which emerged from Salix matsudana f.lobato-glandulosa (SM), Poplus. nigra L.var. thevestina (PN), P. simonii×P. pyramidalis cv‘opera 8277’(PS)and P. alba var. pyramidalis(PA) to volatiles of these hosts twigs and preferred ash leaf maples twigs(Acer negundo) were measured with a Y-tube olfactometer in the laboratory.
With clear air as control, adults A. glabripennis emerging from four host tree species had an obvious orientation response to volatiles emanating from their natal host twigs (P < 0.05), while volatiles emanating from twigs by PA had some repellence to the adults emerging from other three tree species. In addition, the ash leaf maples twigs attracted all adults emerging from each tree species, and the attraction of volatiles emanating from ash leaf maples twigs to female adults was not significant (P > 0.05). With volatiles of ash leaf maples as control, the attraction of volatiles emanating from natal hosts and ash leaf maples to adults emerging from four different tree species had no significant difference. Therefore, A. glabripennis could recognize volatiles emanating from their host plants without existence of preferred host trees. Otherwise, A. glabripennis had no taxis to volatiles emanating from their natal host trees with existence of preferred host trees.
2. Learning behavior of adult A. glabripennis:
The learning behavior of A.glabripennis provided A. negundo and P. alba var. pyramidalis (PA) as food for different days was studied using Y-tube olfactometer . After adults were provide ash leaf maple as food for 1 day, response rate of adults to volatiles emanating from ash leaf maple twigs had obviously increased and the attraction of volatiles emanating from A. negundo to adult a significant difference(P < 0.05). Adults were provided ash leaf maple as food for 2 days, and response rate of adults to volatiles of ash leaf maple increased continuously. As for the adults that were providing ash leaf maple as food for 3and 5days, response rate of adults to volatiles of ash leaf maple maintained up to 70%. It was that demonstrated adults feeding on ash leaf maple induced their preference. Volatiles of PA had a strong repellent effect to adults that were not fed PA (P < 0.05). After adults were provide PA as food for 1 day, the repellent of volatiles emanating from PA twigs to female adults was not significant (P > 0.05). Adults were provided PA as food for 2 days, response rate of adults to volatiles of PA reached to 60%. Adults that were provided PA as food for 3 days were significantly attracted to volatiles emanating from PA twigs (P < 0.05). Adults that were provided PA as food for 5 days, response rate of adults to volatiles of PA maintained the same level with those adults were provided PA as food for 3 days. It demonstrated adults that were feeding on PA induced their habituation. Our experiments demonstrated that feeding experience of adult A.glabripennis was changed response to odour. That is to say, adults A.glabripennis have the ability of learning.
3. Behavior of host-selection of A. glabripennis in mixed forest:
A mixed forest plot with five tree species of 5-year-old were planted in the cage in the field and arranged in a Latin-square design. The tree species were SM, PN, PS, PA and AN. Whether adult A. glabripenni emerging from these five species had“memeory”effect to its natal host tree were researched and explored. Through observing and analyzing inhabiting, feeding, mating, and oviposition of beetles on these trees, the percentage and frequency of observations on AN was higher than on other tree species. Adult A.glabripenni had no showing“memory”effect to their larval host tree species. This is probably due to adults feeding on ash leaf maple induced their preference and the“memory”effect of adult to their larval host was enshrouded.
If there was no AN, the percentage or frequency of observations of adult on SM was higher than on other 3 tree species by observing and analyzing inhabiting, feeding, mating, and oviposition of beetles on these trees. As for oviposition of beetles on these trees, it had no a role. On the whole, SM trees showed strong“attractant”effects if there was no AN.
4. Components and activity of digestive enzymes in A. glabripennis
Larvae feeding on five different tree species, pupae in four different trees, and adults emerging from five host species were selected as experimental materials. The components and activity of digestive enzymes extracted from these materials of A. glabripennis were analyzed and detected .The results revealed that 8 digestive enzymes were detected in larvae and adult A. glabripennis: amylase, pectinase, invertase, maltase, hemicellulase, tryptase, pepsin, and lipase. In addition, it was reported there were endo-β-1,4-glucanase (Cx-cellulase), exdo-β-1,4-glucanase (C1-cellulase) andβ-1,4-glucosidase (β-glycosidase) in larvae and adult. So the larvae and adult A. glabripennis have integrated digestive enzymes. The activity of amylase, pectinase, invertase, maltase, hemicellulase, tryptase and pepsin was not detected in pupae. The activity of lipase in pupae was detected, but its activity was very low. The order of each digestive enzymes activity of larvae of five different tree species was not consisted with the adult emerging from the same tree species. The digestive enzymes activities of adults had no directly correlation with larvae. From larvae to adults by holometamorphosis, the digestive enzymes activities of larvae had no directly effected on adults.
5. Ultra-structure and types of sensilla on the head of A. glabripennis
Ultra-structure, number and distribution of sensilla on the antennae, labial palpi and maxillary palpi of A. glabripennis were observed with an environmental scanning electron microscope. The results indicated that there were 5 types and 13 kinds of sensilla on antennae, including of 5 kinds of sensilla chaeticas, 2 kinds of sensilla trichodeas, 4 kinds of sensilla basiconcas, 1 kind of sensilla campullacea, and 1 kind of finer sensilla.
There were 6 types and 17 kinds of sensilla on labial palpi and maxillary palpi, including of 7 kinds of sensilla chaeticas, 2 kinds of sensilla trichodeas, 3 kinds of sensilla basiconcas, 2 kinds of sensilla styloconica, 2 kinds of sensilla coeloclnicas, and 1 kind of sensilla placoclea. The potential functions of sensilla were dicussed on the basis of its distribution, shape characters as well as correlative references.
1、在天牛成虫对不同寄主树种挥发性物质的趋向试验方面:利用Y型嗅觉仪测定了从漳河柳、箭杆杨、合作杨和新疆杨4种不同抗性等级树种上羽化的光肩星天牛雌、雄成虫对这4种寄主树种和嗜好寄主树种复叶槭枝条气味的嗅觉反应。当以空气为对照时,源自4种不同寄主树种上的天牛成虫对其原寄主树种枝条挥发性气味均有明显的正趋性反应(P < 0.05)。但从合作杨、漳河柳和箭杆杨上羽化的天牛成虫对新疆杨枝条的挥发性气味表现出不同程度的负趋性反应;来源自4种寄主树上的光肩星天牛对复叶槭枝条的挥发性气味均具有不同程度的正趋性反应,但未达显著差异(P > 0.05)。当以复叶槭枝条为对照时,4种不同寄主上羽化的光肩星天牛对原寄主气味和复叶槭气味的趋向差异不大。表明在无嗜好寄主树种(如复叶槭)的前提下,天牛成虫可识别并趋向原寄主树种气味,但如有嗜好寄主树种存在,天牛成虫就不趋向原寄主树种的气味。
2、在天牛成虫的学习行为研究方面:利用Y型嗅觉仪研究了光肩星天牛成虫在嗜好寄主复叶槭和抗性寄主新疆杨上取食不同时间后的学习行为。结果表明,天牛成虫取食复叶槭1d,对复叶槭枝条的气味选择率明显提高,并且复叶槭枝条挥发性气味就对天牛成虫产生了明显的引诱作用(P < 0.05)。取食复叶槭2d,天牛成虫对复叶槭气味的选择率继续提高。取食3d和5d,天牛成虫对复叶槭气味的选择率维持在70%左右。表明天牛成虫取食复叶槭可以出现嗜食性诱导。
没有取食新疆杨的天牛成虫,新疆杨气味对天牛成虫有明显忌避作用(P < 0.05);取食新疆杨1d,新疆杨气味对天牛成虫的忌避作用不明显(P > 0.05);取食新疆杨2d,天牛成虫对新疆杨气味的选择率超过60%,取食新疆杨3d,新疆杨的枝条的气味对天牛产生明显的引诱作用(P < 0.05),取食新疆杨5d,天牛成虫对新疆杨气味的选择率维持在取食3d的水平。表明天牛成虫取食新疆杨后可以出现习惯性反应。说明取食的经历能够改变光肩星天牛成虫对气味的选择,也就是说天牛成虫有很强的学习记忆能力。
3、在天牛成虫林间对不同寄主树种的选择试验方面:利用5年生的复叶槭、漳河柳、箭杆杨、合作杨和新疆杨,在田间的铁丝网笼中按照拉丁方配置了试验林。研究了分别从上述5种寄主树种上羽化的光肩星天牛成虫对原寄主树种选择时是否存在“记忆”效应。
从栖息、取食、交尾、刻槽和产卵的等方面分析,从5种不同寄主树种上羽化的天牛成虫选择复叶槭的比例和频次显著高于其他树种,天牛由于受到嗜好寄主复叶槭的影响,没有对原寄主树种显示出“记忆”效应,因复叶槭是光肩星天牛的嗜食寄主树种,对天牛产生了吸引作用(包括嗜食性诱导),从而推测可能掩盖了天牛成虫对原寄主树种“记忆”。
如果移除嗜好寄主复叶槭,从栖息、取食和交尾情况看,从4种不同寄主树种上羽化的光肩星天牛在漳河柳树上的比例和频次明显高于其它3种寄主树种;从产卵情况看,尽管无规律性的结果,但选择在漳河柳树上产卵的数量仍然很高。总体来看,如移除嗜好寄主复叶槭,漳河柳则表现了较强的“引诱”作用。
4、在不同树种上的天牛各虫态的消化酶活力测定方面:以上述5种寄主树种上的光肩星天牛4龄幼虫、蛹(复叶槭除外)和刚羽化的成虫为对象,测定了幼虫、蛹和成虫体内的消化酶种类和活性。结果表明,光肩星天牛的幼虫和刚羽化成虫的消化道中均具有完整的消化酶系,存在8种消化酶,即淀粉酶、果胶酶、转化酶、麦芽糖酶、半纤维素酶、类胰蛋白酶、蛋白酶和脂肪酶。在天牛蛹中只检测到脂肪酶的活性,但活性非常低,未检测到其它7种酶的活性。
5种寄主树种上刚羽化成虫的8种消化酶的活性大小顺序,没有一种和相应寄主树种上幼虫的一致,说明成虫的消化酶和幼虫的消化酶活性大小相关性不强,也就是说光肩星天牛幼虫消化酶的活性,在经过蛹期变态后不会对成虫的消化酶活性产生直接影响。
5、天牛成虫头部主要感器类型和显微特征研究:利用环境扫描电镜对光肩星天牛雌、雄成虫的触角、下唇须和下颚须上的感受器进行了观察。结果表明,天牛成虫的触角上有5类13种常见的感觉器,其中5种刺形感器、2种毛形感器、4种锥形感器、1钟形感器和1种指形感器。天牛成虫的下唇须和下颚须上有6类17种常见的感受器,其中7种刺形感器、2种毛形感器、3种锥形感器、2种栓锥感器,2种腔锥感器和1种板形感器。根据这些感器的分布和形态特征,并结合以前的文献报道,分析了各种感受器可能的功能。
Anoplophora glabripennis (Motschulsky) is a major polyphagous wood-boring beetle in China that caused great losses to the forest of China. Its primary host species include Acer (maple), Populus (poplar), Salix (willow) and Ulums (elm) and is reported to feed on more than 20 genus host trees. A large number of research projects have focused on this beetle in last 20 years, but host-selection mechanism of adults is unclear. For this reason, this paper makes deep research on host-selection mechanism of adults with types and morphological characteristics of sensilla, chemical ecology, enzymology, and behavior of animal. The main results were as follows:
1. Behavioral responses of adults A. glabripennis to volatiles emanating from different host trees:
Behavioral responses of adults A. glabripennis which emerged from Salix matsudana f.lobato-glandulosa (SM), Poplus. nigra L.var. thevestina (PN), P. simonii×P. pyramidalis cv‘opera 8277’(PS)and P. alba var. pyramidalis(PA) to volatiles of these hosts twigs and preferred ash leaf maples twigs(Acer negundo) were measured with a Y-tube olfactometer in the laboratory.
With clear air as control, adults A. glabripennis emerging from four host tree species had an obvious orientation response to volatiles emanating from their natal host twigs (P < 0.05), while volatiles emanating from twigs by PA had some repellence to the adults emerging from other three tree species. In addition, the ash leaf maples twigs attracted all adults emerging from each tree species, and the attraction of volatiles emanating from ash leaf maples twigs to female adults was not significant (P > 0.05). With volatiles of ash leaf maples as control, the attraction of volatiles emanating from natal hosts and ash leaf maples to adults emerging from four different tree species had no significant difference. Therefore, A. glabripennis could recognize volatiles emanating from their host plants without existence of preferred host trees. Otherwise, A. glabripennis had no taxis to volatiles emanating from their natal host trees with existence of preferred host trees.
2. Learning behavior of adult A. glabripennis:
The learning behavior of A.glabripennis provided A. negundo and P. alba var. pyramidalis (PA) as food for different days was studied using Y-tube olfactometer . After adults were provide ash leaf maple as food for 1 day, response rate of adults to volatiles emanating from ash leaf maple twigs had obviously increased and the attraction of volatiles emanating from A. negundo to adult a significant difference(P < 0.05). Adults were provided ash leaf maple as food for 2 days, and response rate of adults to volatiles of ash leaf maple increased continuously. As for the adults that were providing ash leaf maple as food for 3and 5days, response rate of adults to volatiles of ash leaf maple maintained up to 70%. It was that demonstrated adults feeding on ash leaf maple induced their preference. Volatiles of PA had a strong repellent effect to adults that were not fed PA (P < 0.05). After adults were provide PA as food for 1 day, the repellent of volatiles emanating from PA twigs to female adults was not significant (P > 0.05). Adults were provided PA as food for 2 days, response rate of adults to volatiles of PA reached to 60%. Adults that were provided PA as food for 3 days were significantly attracted to volatiles emanating from PA twigs (P < 0.05). Adults that were provided PA as food for 5 days, response rate of adults to volatiles of PA maintained the same level with those adults were provided PA as food for 3 days. It demonstrated adults that were feeding on PA induced their habituation. Our experiments demonstrated that feeding experience of adult A.glabripennis was changed response to odour. That is to say, adults A.glabripennis have the ability of learning.
3. Behavior of host-selection of A. glabripennis in mixed forest:
A mixed forest plot with five tree species of 5-year-old were planted in the cage in the field and arranged in a Latin-square design. The tree species were SM, PN, PS, PA and AN. Whether adult A. glabripenni emerging from these five species had“memeory”effect to its natal host tree were researched and explored. Through observing and analyzing inhabiting, feeding, mating, and oviposition of beetles on these trees, the percentage and frequency of observations on AN was higher than on other tree species. Adult A.glabripenni had no showing“memory”effect to their larval host tree species. This is probably due to adults feeding on ash leaf maple induced their preference and the“memory”effect of adult to their larval host was enshrouded.
If there was no AN, the percentage or frequency of observations of adult on SM was higher than on other 3 tree species by observing and analyzing inhabiting, feeding, mating, and oviposition of beetles on these trees. As for oviposition of beetles on these trees, it had no a role. On the whole, SM trees showed strong“attractant”effects if there was no AN.
4. Components and activity of digestive enzymes in A. glabripennis
Larvae feeding on five different tree species, pupae in four different trees, and adults emerging from five host species were selected as experimental materials. The components and activity of digestive enzymes extracted from these materials of A. glabripennis were analyzed and detected .The results revealed that 8 digestive enzymes were detected in larvae and adult A. glabripennis: amylase, pectinase, invertase, maltase, hemicellulase, tryptase, pepsin, and lipase. In addition, it was reported there were endo-β-1,4-glucanase (Cx-cellulase), exdo-β-1,4-glucanase (C1-cellulase) andβ-1,4-glucosidase (β-glycosidase) in larvae and adult. So the larvae and adult A. glabripennis have integrated digestive enzymes. The activity of amylase, pectinase, invertase, maltase, hemicellulase, tryptase and pepsin was not detected in pupae. The activity of lipase in pupae was detected, but its activity was very low. The order of each digestive enzymes activity of larvae of five different tree species was not consisted with the adult emerging from the same tree species. The digestive enzymes activities of adults had no directly correlation with larvae. From larvae to adults by holometamorphosis, the digestive enzymes activities of larvae had no directly effected on adults.
5. Ultra-structure and types of sensilla on the head of A. glabripennis
Ultra-structure, number and distribution of sensilla on the antennae, labial palpi and maxillary palpi of A. glabripennis were observed with an environmental scanning electron microscope. The results indicated that there were 5 types and 13 kinds of sensilla on antennae, including of 5 kinds of sensilla chaeticas, 2 kinds of sensilla trichodeas, 4 kinds of sensilla basiconcas, 1 kind of sensilla campullacea, and 1 kind of finer sensilla.
There were 6 types and 17 kinds of sensilla on labial palpi and maxillary palpi, including of 7 kinds of sensilla chaeticas, 2 kinds of sensilla trichodeas, 3 kinds of sensilla basiconcas, 2 kinds of sensilla styloconica, 2 kinds of sensilla coeloclnicas, and 1 kind of sensilla placoclea. The potential functions of sensilla were dicussed on the basis of its distribution, shape characters as well as correlative references.
引文
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