基于害虫—天敌关系的西花蓟马—东亚小花蝽趋光性的结构基础及其行为机理
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摘要
西花蓟马(Frankliniella occidentalis)属缨翅目(Thysanoptera)蓟马科(Thripidae)花蓟马属(Frankliniella),是严重危害蔬菜、花卉等作物的世界性害虫之一,也是近年来在我国迅速蔓延、发生危害日趋严重的重要外来入侵害虫。东亚小花蝽(Oriua sauteri)属花蝽科(Anthocoridae)小花蝽属(Orius),主要捕食蓟马、蚜虫、粉虱、叶螨等小型害虫和多种鳞翅目害虫卵等,是目前我国农林生态系统中重要的自然控制力量之一。本文立足于害虫-天敌关系,利用行为学、组织解剖学等方法系统比较研究了西花蓟马和小花蝽的趋光行为、复眼形态及其显超微结构及其影响因素等的异同及成因,以期揭示复眼视觉的结构基础和作用机理,为更好地保护利用天敌、监测和防控害虫提供依据。主要结果如下:
1.西花蓟马成虫复眼形态学和显微结构。
西花蓟马成虫复眼位于头部两侧,单个复眼约由70个小眼组成,其表面生有十余根刺状感觉毛。复眼分为背、腹两个区域,其中背区小眼排列呈半球形,腹区小眼排列成三角形。
其成虫复眼的小眼由角膜、晶锥、视杆和基膜组成,在晶锥、视杆周围分布有大量色素颗粒。从结构上看,其成虫复眼属并列像眼。
2.东亚小花蝽成虫复眼形态学和显、超微结构。
东亚小花蝽成虫复眼位于头部两侧且靠近腹面,雌雄成虫小眼数分别为230、210,呈近似半球体,感觉毛约30根。雌虫复眼直径为153.85μm,高为70.85μm;背面观,两个复眼间最近距离为168.54μm,腹面观,其间最近距离为136.84μm。雄虫复眼直径144.74μm,高60.72μm;背面观,两个复眼间最近距离为166.67μm,腹面观其最近距离为102.56μm。
其成虫复眼的小眼由角膜、晶锥、视杆和基膜组成。角膜由多层片状纤维组成,呈外部稍凸内部半圆形的双凸透镜结构,最厚处为9.41μm;晶锥无色透明,呈圆台状,长度为13.33μm,远心端直径为8.33μm,近心端直径为4.44μm;视杆长为33.33μm;基膜为栅栏状,厚度约为0.27μm。不同光强度水平下,通过光感受细胞的微绒毛排列形状和方向来调节成像,其成虫复眼为典型的并列像眼。
3.西花蓟马成虫的光谱和光强度行为反应。
在340 nm-605 nm波谱内的14个单色光波长下,雌虫趋光行为反应率较高的单色光有绿光区(498 nm和524 nm)、紫外区(380 nm)和蓝光区(440 nm),避光行为反应率较高的光谱有蓝光区(440 nm)、紫外区(340 nm)和橙色光区(582 nm);雄虫趋光行为反应率较高的单色光有绿光区(498 nm)和紫外区(380 nm),避光行为反应率较高的单色光有蓝光区(440 nm)、紫外区(340 nm和360 nm);光谱行为反应在性别之间有一定差异,雌虫敏感光谱较多(有四个单色光),但光谱敏感强度稍差(趋光行为反应率最高值为20.31%),雄虫敏感光谱相对较少(仅有两个单色光)但光敏感强度较好(趋光行为反应率最高值为27.19%)。
在一定光强度范围(Log 4.5-Log 0.0)内,不同光强度的单色光或白光刺激下,雌、雄成虫趋光行为反应率有较大变化,整体上随光强度的增强而增大;光强度对其避光行为反应率影响较小;性别对其光强度行为反应亦较小。
4.东亚小花蝽成虫的光谱和光强度行为反应。
在340 nm-605 nm波谱内的14个单色光波长下,雌虫趋光行为反应率较高的单色光有紫外区(340 nm、360 nm和380 nm)、绿光区(524 nm)和橙红光区(583 nm、605 nm);雄虫趋光行为反应率较高的单色光有紫外区(340 nm、360 nm和380 nm)、红色光区(605 nm)和绿光区(524 nm);光谱行为反应在性别之间有一定差异,雌虫敏感光谱较多,且光谱敏感强度也强于雄虫。
在一定光强度范围(Log 4.5-Log 0.0)内,不同光强度的白光刺激下,雌、雄成虫趋光行为反应率有较大变化,在log 4.5- Log 1.5光强内,随光强增强雌虫趋光反应率变化不大,在log 1.0- Log 0.0光强内随光强增强趋光反应率迅速增大,雄虫的趋光行为反应率基本上随光强增强增大;光强度对其避光行为反应率影响较小;性别对其光强度行为反应的影响亦较小。
5.西花蓟马的趋色行为。
西花蓟马对绿色、黄色和蓝色均有一定的趋性。在取食条件和非取食条件下,不同颜色背景对其趋向率影响均不大。同时,各颜色的选择性试验中(两两对比),不同颜色之间的趋向率有一定差异,但差异性均不显著。
本文的创新点在于针对害虫测报和诱杀用灯光、色板等缺乏选择性,从保护自然界天敌和生物多样性角度出发,着眼于害虫-天敌二级营养关系,选择近年来农业生产中危害较重的外来入侵物种西花蓟马及其优势天敌东亚小花蝽为试验材料,较系统地比较研究和分析了害虫、天敌两大功能团中该两种代表性种类复眼的形态结构、光色行为反应的异同及其影响因素,试图揭示其光视觉行为生理机理,了解光色视觉信息在两者之间复眼结构、行为等细胞核种群水平上的内在联系实质及其生物学意义,为理论上深刻理解趋光性、趋色性实质及其其在协同演化和取食、繁殖等重要生命活动中的作用,实践上为改进和消除现有诱虫灯、色板的非选择性缺陷,研发具有“选择性或专一性”、“绿色化”的新灯(光)、色板及光色活性物质等提供科学依据和新思路。
Western flower thrips, Frankliniella occidentalis (Thysanoptera), is a serious worldwide pest on vegetables, flowers and other crop, but also spread rapidly in recent years, growing seriously hamful as a main alien invasive pest in China;.Orius sauteri ( Anthocoridae ) ( Orius), the predator of thrips, aphids, whiteflies, mites and other small insects and eggs of Lepidoptera pests , is a important natural control force of agro-forest ecosystem in China currently. This article is based on the Insect pest-natural enemy relationship, using behavior, histoanatomy method to study both phototactic behavior, morphology and ultrastructure of compound eye were studied, and the influencing factors of the similarities and differences and the causes were discussed in order to reveal the structural basis and action mechanism of eye's vision, and provide a basis for better protection and utilization of natural enemies to prevent and control pests. The main results are as follows:
1. The morphology and microstrcture of comopund eye of Frankliniella occidentalis The western flower thrips compound eyes located on both sides of the head, a single eye consists of about 70 small ommatidia, the surface having more than 10 thorn shape sensory hair. Compound eye is divided into a dorsal, ventral region in two, wherein the back area is arranged in the hemisphere, abdominal area is arranged into a triangle.The small eye of the adult compound eyes the cornea, crystalline cone, rod, and the basement membrane
The ommatidium of adult compouned eye was comprised of corneal, cone; rhabdom and basement membrane.Around the crystalline cone and rhabdom distributed a lot of pigment granules. Considered from the structure, the compound eye belongs to an apposition eye.
2. The morphology, microstrcture and ultrastructrue of comopund eye of Orius sauteri The compound eye of Orius sauteri is located on both sides of the head and near the ventral, the facet of male and female adultswere 230 and 210 respectively, approximately hemispherical, sensory hairs ca. 30 root. The diameter of female compound eye is about 153.85μm, height about 70.85μm; in the back view, closest distance of two eye was about 168.54μm, ventral view about 136.84μm. The male eye diameter is about 144.74μm, high about 60.72μm; looks from the back, two eye has a distance of about 166.67μm, and on the ventral of about 102.56μm.
The adult eye eyes were comprised of corneal, cone, rhabdom and membrane. Corneal is composed by the multi-layer sheet fiber, present a convex lens structure as slightly convex in external and semicircular in internal, thickness of about 9.41μm; cone is colorless and transparent, shaped a circular truncated cone, the length is about 13.33μm, the distal end has a diameter of 8.33μm, the proximal end of 4.44μm; the rhabdom is about 33.33μm in length; base membrane shaped palisade, thickness of about 0.27μm. Orius sauteri adjust the imaging by microvilli shape and direction in different light intensity level is a typical apposition eye.
3. The spectral and light intensity of behavior response of Frankliniella occidentalis Under the 14 Monochromatic Wavelengths within the 340 nm -605 nm spectrum, in the female, phototactic behavior response rates were higher in the monochromatic light green ( 498 nm, 524 nm ), ultraviolet ( 380 nm ) and blue light district ( 440 nm ), photophobic behavior response rates were higher in the spectrum of blue light district (440 nm ), ultraviolet (340 nm ) and orange light ( 582 nm ); in male ,phototactic behavior response rates were higher in the monochromatic light is green ( 498 nm ) and ultraviolet ( 380 nm ), photophobic behavior response rates were higher in the monochromatic light with blue zone ( 440 nm ), ultraviolet ( 340 nm and 360 nm ); spectral behavior reaction has certain difference between the sex, the female has more sensitive spectrum, but spectral sensitive strength is slightly worse, while the male has lesssensitive spectral relatively but better light sensitive intensity.
In Log 4.5-Log 0.0 light intensity range, in stimulation of different light intensity of monochromatic light or white light ,the phototaxis behavior response rate has a big difference between female and male adult, phototaxis behavior response rate on the whole with the increasing light intensity; light avoidance behavior reaction rate also has a certain influence, but the effect is small; gender on light intensity of behavioral responses to have certain effect, but the effect is relatively small.
4. The spectral and light intensity of behavior response of Orius sauteri In the 14 monochromatic wavelength spectrum within the 340 nm -605 nm, the phototactic behavior response rates were higher in the monochromatic light with ultraviolet ( 340 nm, 360 nm and 380 nm ), green ( 524 nm ) and orange red light district ( 583 nm, 605 nm ); male phototactic behavior response rates were higher in the monochromatic light ultraviolet region ( 340 nm, 360 nm and 380 nm ), red light ( 605 nm) and green ( 524 nm ); spectral behavior of reaction between the sexes have certain difference, the female has more sensitive spectrums, and the strength of sensitive spectrum is better than male.
In Log 4.5-Log 0.0 light intensity range, in stimulation of white light in different light intensity , female, male adult phototaxis behavior response rate has significant changes, in log 4.5-log 1.5 intensity,in female, with the intensity enhancement the phototactic response rate varied little .in the log 1.0-Log 0.0 intensity, with the intensity enhancement the phototactic reaction rate increases rapidly. The phototactic behavior response rate of male increases with light intensity enhancement on the whole; the light avoidance behavior response rate has some effect with light intensity enhancement , but the effect is small; sex of Orius sauteri have certain effect on light intensity of behavioral responses, but the effect is relatively small.
5. Hasten color behavior of Frankliniella occidentalis
The western flower thrips have some taxis with the green, yellow and blue. In feeding and non-feeding conditions, western flower thrips trend rate are affected of different color background , but little effect; at the same time, comparing each two color environment, different color trend rate have certain difference, but the difference is not significant.
The innovation of this paper is to focus on the pest-natural enemy two trophic relationships, systematically studies and analyzes the pest, natural enemy, two functional group in the representation of the category - western flower thrips and Orius sauteri of their compound eye structure, physiological mechanism of light, color behavior and similarities, differences and the causes, to reveal the inherent essence and biological significance of the color, light visual information in compound eye structure, behavior and other aspects between them , providing scientific basis for the lighting and optical activity measurement and trap and kill technical for practice use. At the same time, initially revealed the physiological mechanism and causes of Orius sauteri and western flower thrips' photoreception and phototactic behavior at the cellular and population levels.
1.西花蓟马成虫复眼形态学和显微结构。
西花蓟马成虫复眼位于头部两侧,单个复眼约由70个小眼组成,其表面生有十余根刺状感觉毛。复眼分为背、腹两个区域,其中背区小眼排列呈半球形,腹区小眼排列成三角形。
其成虫复眼的小眼由角膜、晶锥、视杆和基膜组成,在晶锥、视杆周围分布有大量色素颗粒。从结构上看,其成虫复眼属并列像眼。
2.东亚小花蝽成虫复眼形态学和显、超微结构。
东亚小花蝽成虫复眼位于头部两侧且靠近腹面,雌雄成虫小眼数分别为230、210,呈近似半球体,感觉毛约30根。雌虫复眼直径为153.85μm,高为70.85μm;背面观,两个复眼间最近距离为168.54μm,腹面观,其间最近距离为136.84μm。雄虫复眼直径144.74μm,高60.72μm;背面观,两个复眼间最近距离为166.67μm,腹面观其最近距离为102.56μm。
其成虫复眼的小眼由角膜、晶锥、视杆和基膜组成。角膜由多层片状纤维组成,呈外部稍凸内部半圆形的双凸透镜结构,最厚处为9.41μm;晶锥无色透明,呈圆台状,长度为13.33μm,远心端直径为8.33μm,近心端直径为4.44μm;视杆长为33.33μm;基膜为栅栏状,厚度约为0.27μm。不同光强度水平下,通过光感受细胞的微绒毛排列形状和方向来调节成像,其成虫复眼为典型的并列像眼。
3.西花蓟马成虫的光谱和光强度行为反应。
在340 nm-605 nm波谱内的14个单色光波长下,雌虫趋光行为反应率较高的单色光有绿光区(498 nm和524 nm)、紫外区(380 nm)和蓝光区(440 nm),避光行为反应率较高的光谱有蓝光区(440 nm)、紫外区(340 nm)和橙色光区(582 nm);雄虫趋光行为反应率较高的单色光有绿光区(498 nm)和紫外区(380 nm),避光行为反应率较高的单色光有蓝光区(440 nm)、紫外区(340 nm和360 nm);光谱行为反应在性别之间有一定差异,雌虫敏感光谱较多(有四个单色光),但光谱敏感强度稍差(趋光行为反应率最高值为20.31%),雄虫敏感光谱相对较少(仅有两个单色光)但光敏感强度较好(趋光行为反应率最高值为27.19%)。
在一定光强度范围(Log 4.5-Log 0.0)内,不同光强度的单色光或白光刺激下,雌、雄成虫趋光行为反应率有较大变化,整体上随光强度的增强而增大;光强度对其避光行为反应率影响较小;性别对其光强度行为反应亦较小。
4.东亚小花蝽成虫的光谱和光强度行为反应。
在340 nm-605 nm波谱内的14个单色光波长下,雌虫趋光行为反应率较高的单色光有紫外区(340 nm、360 nm和380 nm)、绿光区(524 nm)和橙红光区(583 nm、605 nm);雄虫趋光行为反应率较高的单色光有紫外区(340 nm、360 nm和380 nm)、红色光区(605 nm)和绿光区(524 nm);光谱行为反应在性别之间有一定差异,雌虫敏感光谱较多,且光谱敏感强度也强于雄虫。
在一定光强度范围(Log 4.5-Log 0.0)内,不同光强度的白光刺激下,雌、雄成虫趋光行为反应率有较大变化,在log 4.5- Log 1.5光强内,随光强增强雌虫趋光反应率变化不大,在log 1.0- Log 0.0光强内随光强增强趋光反应率迅速增大,雄虫的趋光行为反应率基本上随光强增强增大;光强度对其避光行为反应率影响较小;性别对其光强度行为反应的影响亦较小。
5.西花蓟马的趋色行为。
西花蓟马对绿色、黄色和蓝色均有一定的趋性。在取食条件和非取食条件下,不同颜色背景对其趋向率影响均不大。同时,各颜色的选择性试验中(两两对比),不同颜色之间的趋向率有一定差异,但差异性均不显著。
本文的创新点在于针对害虫测报和诱杀用灯光、色板等缺乏选择性,从保护自然界天敌和生物多样性角度出发,着眼于害虫-天敌二级营养关系,选择近年来农业生产中危害较重的外来入侵物种西花蓟马及其优势天敌东亚小花蝽为试验材料,较系统地比较研究和分析了害虫、天敌两大功能团中该两种代表性种类复眼的形态结构、光色行为反应的异同及其影响因素,试图揭示其光视觉行为生理机理,了解光色视觉信息在两者之间复眼结构、行为等细胞核种群水平上的内在联系实质及其生物学意义,为理论上深刻理解趋光性、趋色性实质及其其在协同演化和取食、繁殖等重要生命活动中的作用,实践上为改进和消除现有诱虫灯、色板的非选择性缺陷,研发具有“选择性或专一性”、“绿色化”的新灯(光)、色板及光色活性物质等提供科学依据和新思路。
Western flower thrips, Frankliniella occidentalis (Thysanoptera), is a serious worldwide pest on vegetables, flowers and other crop, but also spread rapidly in recent years, growing seriously hamful as a main alien invasive pest in China;.Orius sauteri ( Anthocoridae ) ( Orius), the predator of thrips, aphids, whiteflies, mites and other small insects and eggs of Lepidoptera pests , is a important natural control force of agro-forest ecosystem in China currently. This article is based on the Insect pest-natural enemy relationship, using behavior, histoanatomy method to study both phototactic behavior, morphology and ultrastructure of compound eye were studied, and the influencing factors of the similarities and differences and the causes were discussed in order to reveal the structural basis and action mechanism of eye's vision, and provide a basis for better protection and utilization of natural enemies to prevent and control pests. The main results are as follows:
1. The morphology and microstrcture of comopund eye of Frankliniella occidentalis The western flower thrips compound eyes located on both sides of the head, a single eye consists of about 70 small ommatidia, the surface having more than 10 thorn shape sensory hair. Compound eye is divided into a dorsal, ventral region in two, wherein the back area is arranged in the hemisphere, abdominal area is arranged into a triangle.The small eye of the adult compound eyes the cornea, crystalline cone, rod, and the basement membrane
The ommatidium of adult compouned eye was comprised of corneal, cone; rhabdom and basement membrane.Around the crystalline cone and rhabdom distributed a lot of pigment granules. Considered from the structure, the compound eye belongs to an apposition eye.
2. The morphology, microstrcture and ultrastructrue of comopund eye of Orius sauteri The compound eye of Orius sauteri is located on both sides of the head and near the ventral, the facet of male and female adultswere 230 and 210 respectively, approximately hemispherical, sensory hairs ca. 30 root. The diameter of female compound eye is about 153.85μm, height about 70.85μm; in the back view, closest distance of two eye was about 168.54μm, ventral view about 136.84μm. The male eye diameter is about 144.74μm, high about 60.72μm; looks from the back, two eye has a distance of about 166.67μm, and on the ventral of about 102.56μm.
The adult eye eyes were comprised of corneal, cone, rhabdom and membrane. Corneal is composed by the multi-layer sheet fiber, present a convex lens structure as slightly convex in external and semicircular in internal, thickness of about 9.41μm; cone is colorless and transparent, shaped a circular truncated cone, the length is about 13.33μm, the distal end has a diameter of 8.33μm, the proximal end of 4.44μm; the rhabdom is about 33.33μm in length; base membrane shaped palisade, thickness of about 0.27μm. Orius sauteri adjust the imaging by microvilli shape and direction in different light intensity level is a typical apposition eye.
3. The spectral and light intensity of behavior response of Frankliniella occidentalis Under the 14 Monochromatic Wavelengths within the 340 nm -605 nm spectrum, in the female, phototactic behavior response rates were higher in the monochromatic light green ( 498 nm, 524 nm ), ultraviolet ( 380 nm ) and blue light district ( 440 nm ), photophobic behavior response rates were higher in the spectrum of blue light district (440 nm ), ultraviolet (340 nm ) and orange light ( 582 nm ); in male ,phototactic behavior response rates were higher in the monochromatic light is green ( 498 nm ) and ultraviolet ( 380 nm ), photophobic behavior response rates were higher in the monochromatic light with blue zone ( 440 nm ), ultraviolet ( 340 nm and 360 nm ); spectral behavior reaction has certain difference between the sex, the female has more sensitive spectrum, but spectral sensitive strength is slightly worse, while the male has lesssensitive spectral relatively but better light sensitive intensity.
In Log 4.5-Log 0.0 light intensity range, in stimulation of different light intensity of monochromatic light or white light ,the phototaxis behavior response rate has a big difference between female and male adult, phototaxis behavior response rate on the whole with the increasing light intensity; light avoidance behavior reaction rate also has a certain influence, but the effect is small; gender on light intensity of behavioral responses to have certain effect, but the effect is relatively small.
4. The spectral and light intensity of behavior response of Orius sauteri In the 14 monochromatic wavelength spectrum within the 340 nm -605 nm, the phototactic behavior response rates were higher in the monochromatic light with ultraviolet ( 340 nm, 360 nm and 380 nm ), green ( 524 nm ) and orange red light district ( 583 nm, 605 nm ); male phototactic behavior response rates were higher in the monochromatic light ultraviolet region ( 340 nm, 360 nm and 380 nm ), red light ( 605 nm) and green ( 524 nm ); spectral behavior of reaction between the sexes have certain difference, the female has more sensitive spectrums, and the strength of sensitive spectrum is better than male.
In Log 4.5-Log 0.0 light intensity range, in stimulation of white light in different light intensity , female, male adult phototaxis behavior response rate has significant changes, in log 4.5-log 1.5 intensity,in female, with the intensity enhancement the phototactic response rate varied little .in the log 1.0-Log 0.0 intensity, with the intensity enhancement the phototactic reaction rate increases rapidly. The phototactic behavior response rate of male increases with light intensity enhancement on the whole; the light avoidance behavior response rate has some effect with light intensity enhancement , but the effect is small; sex of Orius sauteri have certain effect on light intensity of behavioral responses, but the effect is relatively small.
5. Hasten color behavior of Frankliniella occidentalis
The western flower thrips have some taxis with the green, yellow and blue. In feeding and non-feeding conditions, western flower thrips trend rate are affected of different color background , but little effect; at the same time, comparing each two color environment, different color trend rate have certain difference, but the difference is not significant.
The innovation of this paper is to focus on the pest-natural enemy two trophic relationships, systematically studies and analyzes the pest, natural enemy, two functional group in the representation of the category - western flower thrips and Orius sauteri of their compound eye structure, physiological mechanism of light, color behavior and similarities, differences and the causes, to reveal the inherent essence and biological significance of the color, light visual information in compound eye structure, behavior and other aspects between them , providing scientific basis for the lighting and optical activity measurement and trap and kill technical for practice use. At the same time, initially revealed the physiological mechanism and causes of Orius sauteri and western flower thrips' photoreception and phototactic behavior at the cellular and population levels.
引文
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