林业科学  2015, Vol. 51 Issue (9): 134-140   PDF    
DOI: 10.11707/j.1001-7488.20150917
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文章信息

吕飞, 海小霞, 王志刚, 毕拥国, 刘炳响, 阎爱华
Lü Fei, Hai Xiaoxia, Wang Zhigang, Bi Yongguo, Liu Bingxiang, Yan Aihua
天牛视觉感受机制研究进展
Progress on the Visual Sensory Mechanism of the Longhorn Beetles (Coleoptera: Cerambycidae)
林业科学, 2015, 51(9): 134-140
Scientia Silvae Sinicae, 2015, 51(9): 134-140.
DOI: 10.11707/j.1001-7488.20150917

文章历史

收稿日期：2014-04-29
修回日期：2014-07-02

作者相关文章

吕飞
海小霞
王志刚
毕拥国
刘炳响
阎爱华

引用本文
吕飞, 海小霞, 王志刚, 毕拥国, 刘炳响, 阎爱华. 2015. 天牛视觉感受机制研究进展[J]. 林业科学, 51(9): 134-140.
Lü Fei, Hai Xiaoxia, Wang Zhigang, Bi Yongguo, Liu Bingxiang, Yan Aihua. 2015. Progress on the Visual Sensory Mechanism of the Longhorn Beetles (Coleoptera: Cerambycidae). Scientia Silvae Sinicae, 51(9): 134-140.  DOI: 10.11707/j.1001-7488.20150917
天牛视觉感受机制研究进展
吕飞, 海小霞, 王志刚 , 毕拥国, 刘炳响, 阎爱华    
河北农业大学林学院 河北省林木种质资源与森林保护重点实验室 保定 071000
收稿日期：2014-04-29；修回日期：2014-07-02
基金项目：国家自然科学基金项目(30571501)。
通讯作者：王志刚
摘要：【目的】天牛是林业重要的蛀干害虫，其中有些种类如松墨天牛、光肩星天牛等是重要的检疫性害虫，给我国林业生产和生态环境带来巨大危害。昆虫利用视觉、嗅觉、味觉、听觉信号去定位和识别寄主及配偶，在过去的几十年中，大量的工作主要集中在嗅觉及其化学生态学上；而视觉作为重要的感觉系统之一，对昆虫寄主和配偶定位及识别等多种行为活动起着至关重要的作用，然而对视觉调节天牛成虫行为的研究却相对较少，尤其是对天牛视觉系统研究的综述还未见报道。【方法】利用Web of Science，Google Scholar，CNKI等数据库，查询天牛科视觉的相关文献。对天牛成虫视觉器官(复眼)的组织解剖学结构特征、视觉对天牛成虫行为的调节作用以及灯诱在林业害虫综合治理中的应用方面研究进行综述。【结果】天牛成虫的复眼围绕在触角的基部，被触角分为上下2部，呈肾形，无晶眼或伪晶眼类型，每个小眼含有8个视网膜细胞，其中2个中心细胞被6个周缘细胞包围；日行性和夜行性天牛种类复眼解剖结构学上的主要区别在于小眼密度、感杆束占据视网膜细胞的面积比率等。虫体和寄主植物颜色、形状及大小是主要的视觉信号来源，不仅可以影响某些天牛种类的交配成功率，而且可以影响对寄主植物的定位及非寄主植物的区分。灯光诱杀是害虫综合治理(IPM)中重要的手段之一，具有延缓害虫抗药性、降低防治成本和对环境影响小等优点；诱捕光源的光谱和光强以及诱捕器的形状及颜色对害虫的诱集效果均有显著影响，目前对天牛的诱集光源(主要是黑光灯和高压汞灯一类紫外光)研究较多，而对光谱、光强以及诱捕器外形对天牛诱捕效率的影响研究较少；同样，灯光诱杀也可以用于出入境检验检疫局对检疫性的天牛害虫进行检测。【结论】视觉系统不仅在昆虫的觅食和求偶行为中起着重要的作用，而且对产卵位置识别、躲避天敌等行为均有重要的意义，但是，天牛科成虫的视觉系统及视觉生态学相关的研究却很缺乏；将来可以从寄主植物和配偶的颜色、形状、大小等特征对天牛行为活动的影响，日行和夜行性天牛复眼的解剖学特征及区别等对天牛视觉生理生态学相关内容进行深入研究，为研制优异的、环境友好型的诱捕和测报装置提供理论基础，同时也为合理调控天牛及其天敌昆虫的种群结构提供理论基础。
关键词： 蛀干害虫    天牛    视觉    复眼    灯诱    
Progress on the Visual Sensory Mechanism of the Longhorn Beetles (Coleoptera: Cerambycidae)
Lü Fei, Hai Xiaoxia, Wang Zhigang, Bi Yongguo, Liu Bingxiang, Yan Aihua    
Key Laboratory for Germplasm Resources of Forest Trees and Forest Protection of Hebei Province College of Forestry, Agricultural University of Hebei Baoding 071001
Abstract: [Objective] Longhorn beetles are important trunkborer pests of forests and can cause serious damage to forest ecosystem; some insects in cerambycidae, such as Monochamus alternatus, Anoplophora glabripennis, are even important quarantine pests. Many insects use visual, olfactory, gustatory and auditory cues to locate and recognize their preference host plants and mates. In the last few decades a great deal of work has been devoted to olfactory sense and chemical ecology. It is well known that the visual sensory system is also one of the important sensory systems of insects and plays a vital role in the behavior activities of host and mate location and recognition and so on, however, study on it is relative less compared with the research on olfactory system in the influence of behavior activities of longhorn beetle, especially the review of visual mechanism of cerambycidae is not reported. [Method] The database of Web of Science and Google Scholar, the Chinese journal full-text database of the China National Knowledge Infrastructure (CNKI) were used to systematically search for publication relevant to visual system of cerambycidae. This paper not only introduces the characteristic anatomy structure of the visual organ of cerambycidae, but also reviews roles the visual system plays for the host and mate location and recognition of cerambycidae, as well as the utilization of research results on visual mechanism in the integrated pest management of forests. [Result] The adult of longicorn beetle possess two compound eyes, each of which is divided into two parts by antenna. The eyes occupy lateral position on either side of the head and surround the basal of antenna and the shape is kidney type and belongs to acone eye and pseudocone eye. Every ommatidium has eight retinula cells, and two central retinula cells are surrounded by six peripheral cells. The distinctions of diurnal and nocturnal longicorn beetles are mainly based on ommatidium density and the ratio of area of rhabdoms and retinula cells and so on in the anatomical structures. The visual cues of host plants and mates, including color, shape and size, are often the three key components, and those cues not only can affect mating success ratio of some longicorn beetles but also can influence host-plant location and non-host plant distinction. Light trap is one of important methods of Integrated Pest Management, and it has the merits of delaying insect resistance development, reducing control cost and little effect on environment. The trapping effect of pests was influenced significantly by the spectrum and intensity of light resource, as well as shape and color of trapping apparatus, but the light resource to lure longicorn beetles is mainly ultraviolet light, including black light and high-pressure mercury discharge lamp. The researches of spectrum and intensity of trapping light resource and shape of trapping apparatus to influence mean catch of target pests are insufficient up to now. The light trap may also be used to detect quarantine longicorn beetles in the entry-exit inspection and quarantine. [Conclusion] Visual systems play an important role in the forage searching and mating of insects, and are also important for the recognition of oviposition sites, avoidance natural enemy and so on. However, the researches of visual systems and visual ecology of longicorn beetles remain scarce. In the future, the visual physiological and ecological relevant contents should be further researched, including how to influence activity behavior of longicorn beetles by the color, shape and size of potential host plants and mates, the anatomical characteristics and distinction of diurnal and nocturnal longicorn beetles and so on. The researches can supply a theoretical basis for the development of outstanding and environmentally friendly trap and prediction and forecast device and regulate and control reasonably population structure of longicorn beetles and their nature enemy.
Key words: Trunkborers    cerambycids    visual    compound eye    light trapping    

天牛作为一类重要的林木蛀干害虫，以幼虫蛀干危害为主，轻者造成树势衰弱，重者造成树木死亡，严重损害了森林生态系统的经济和生态效益(魏建荣等，2007)。目前全世界已经记录的天牛大约有4 000个属35 000种以上(Hanks，1999)；在我国的记录大约有2 700种(嵇保中等，2002)。种类繁多的天牛科昆虫，其成虫对寄主及配偶的定位和识别过程必然呈现多样化特征，多数研究认为是寄主挥发物及性信息素起着主要作用，在配偶定位上大体分为长距离、短距离和接触性信息素3种(江望锦等，2005)，但也有研究认为视觉在天牛对寄主及配偶的定位和识别过程中起关键作用(李德家等，1999; 刘博等，2012)。

天牛成虫对寄主和配偶的定位及识别机制是一种复杂的行为反应，涉及视觉、嗅觉和触觉等多种感觉系统。目前的研究主要集中于嗅觉以及相关的化学生态学领域，视觉仅有少量研究。江望锦等(2005)、周琳等(2006)对天牛性信息素组成、感受器官(触角)及其神经传导等相关研究做了详细综述，Allison等(2004)对天牛的化学生态学进行了综述，高瑞桐等(2001)对光肩星天牛行为学、寄主选择性、造成的经济损失及其预测预报和综合防治等方面进行了总结，Haack等(2010)对全球光肩星天牛(Anoplophora glabripennis)和星天牛(A. chinensis)的入侵管理做了相关综述，Harks(1999)对幼虫寄主植物与天牛生殖策略的相互关系做了综述。但是，视觉对天牛行为活动的影响、视觉感受器官的组织解剖等天牛视觉相关领域研究综述未见报道。本文评述天牛复眼组织解剖学、视觉对成虫行为的调节以及视觉机制在天牛综合治理中的应用等方面的研究进展，为深入研究提供参考。

1 天牛成虫视觉感受器官研究进展

复眼是昆虫重要的视觉感受器官。天牛成虫复眼一般环绕在触角基部，被触角分为上下2个部分。Koyama等(1975)对产于日本的多种天牛复眼内部显微结构进行观察，发现其复眼呈肾形，属无晶眼类型(acone type)，由角膜、角膜形成的伪晶锥细胞、虹膜色素细胞、视杆细胞、小网膜细胞和视网膜色素细胞6部分组成，每个小眼包含8个视网膜细胞，感杆束的形状和长度因种类而异；研究中尽管也包括锯天牛(Prionus insularis)，但未对其复眼结构进行详细描述。之后，Gokan等(1979a)对锯天牛的复眼内部超微结构进行了细致观察，发现该天牛每个小眼包含8个视网膜细胞，其中2个中心细胞被6个周缘细胞包围；半融合类型(semi-fused type)的感杆束(rhabdom)占据整个视网膜细胞体积的绝大部分，推测该结构也许是夜出性种类的典型构造。Gokan等(1979b)对5种日出性和2种夜出性天牛复眼组织结构解剖研究表明，日出性天牛形成感杆束的感杆相互分离，长度要较夜出性的短；夜出性种类的感杆束占视网膜细胞的比例要比日出性的大；不同光暗条件下，内部结构有明显的变化；但是对外部结构的描述不详细，仅对花天牛属(Leptura)复眼表面结构具有角膜乳突进行了简单描述。嵇保中等(1991)对35种天牛小眼密度及林下灯诱情况做了对比研究，并根据灯诱情况及小眼密度将天牛的活动类型分为夜出性活动、昼夜均活动和日出性活动3类。夜出性活动种类，其小眼密度值为70.86~240.75个 ·mm-2，对灯光的趋性较明显；昼夜均能活动的种类，其小眼的密度值一般为312.38~585.00个 ·mm-2，表现出较弱的趋光性；日出性活动种类，一般在703.13~2 687.63个 ·mm-2，对于灯光的趋性很弱。

总体来说，天牛成虫复眼环绕在触角基部，呈肾形，属于无晶眼或伪晶眼(Gokan et al.，1979a; Koyama et al.，1975)，小眼密度随着活动类型的区别具有明显差异。但是对天牛成虫复眼外部结构、内部超微结构、光感受器类型等方面的研究仍然很缺乏。

2 视觉对天牛成虫行为的调节 2.1 视觉在天牛配偶定位和识别中的作用

昆虫配偶定位和识别被多种感觉器官调节，如视觉(Gorb，1998; Szentesi et al.，2002)、嗅觉(Ginzel et al.，2003; Johansson et al.，2007; Lopes et al.，2005)、听觉(Cocroft et al.，2005)和触觉(Bonduriansky，2001)。天牛配偶定位机制较复杂，王广利等(2007)认为天牛的交配行为多数由短距离或接触性信息素启动，视觉、虫体颜色、形状、大小以及虫体体表化合物均可影响其交配成功率。不同天牛种类对配偶定位的作用方式有显著区别，大致分为3类： 1)通过雄虫的长距离性信息素吸引，再由雌虫的短距离性信息素进行刺激共同完成(江望锦等，2005)；通过该种方式进行配偶定位的天牛种类很多，如葡萄巨虎天牛(Xylotrechus pyrrhoderus)雌虫首先受到雄虫远距离的性信息素刺激而接近雄虫，接着雄虫受到雌虫的性信息素刺激产生兴奋，进而产生交尾行为(Iwabuchi，1985)。柳杉纹虎天牛(Anaglyptus subfasciatus)(Leal et al.，1995)、北美家天牛(Hylotrupes bajulus)(周琳等，2006)、苎麻双脊天牛(Paraglenea fortunei)(Wang et al.，1991)等均是依靠雌虫或者雄虫释放性信息素来吸引异性定位而产生交尾行为。2)由短距离的性信息素刺激雌雄虫产生交尾行为的天牛种类(江望锦等，2005)；如胸斑星天牛(Anoplophora malasiaca)(Fukaya，2003)、北美断条黑天牛(Plectrodera scalator)(Ginzel，2003)、柳杉天牛(Semanotus japonicus)(Kim et al.，1993)等均依靠雌雄虫短距离性信息素吸引完成。3)通过寄主植物气味的引诱雌雄成虫向寄主植物聚集，近距离相遇后由视觉或其他感觉系统相互调节进一步完成，如光肩星天牛(贺萍等，1993)、松墨天牛(Monochamus alternatus)(樊建庭等，2007; 刘博等，2012)等。

对于视觉或嗅觉在天牛配偶定位和识别中所起作用研究，主要是通过对天牛复眼致盲或切除触角后，观察试虫交尾的行为学反应，然后分析视觉和嗅觉在配偶定位和识别行为中的作用。贺萍等(1993)对光肩星天牛的求偶识别机制进行研究发现，远距离存在性信息素吸引现象，但是在近距离时，复眼被涂黑的情况下，交尾成功率显著下降，从而证明了光肩星天牛的配偶定位和识别是由视觉和嗅觉共同作用完成的。而李德家等(1999)则认为光肩星天牛不是依靠嗅觉信息来寻找和识别配偶的，而是由雌虫对雄虫的视觉刺激作用下引起的。在研究视觉和嗅觉对天牛的配偶定位和识别所起的关键作用时，视觉起着关键作用的种类有眉斑并脊天牛(Glenea cantor)(Lu et al.，2007)、星天牛(Fukaya et al.，2004; 2005; Yasui，2009)、松墨天牛(樊建庭等，2007; 刘博等，2012)等；但是也有种类如云斑白条天牛(Batocera lineolata)(潘翔宇等，2012)、栗山天牛(Massicus raddei)(魏建荣等，2013)复眼致盲后仍可以进行正常的配偶搜寻和交尾行为，说明视觉在配偶定位和识别行为中起的作用可能不大。

2.2 视觉在天牛觅食行为中的作用

视觉如何调节天牛觅食行为的研究报道较少。杨雪彦等(1995)、王福贵等(2000)通过观察混交林中黄斑星天牛(A.nobilis)对寄主的选择行为，发现其主要通过视觉、触角摆动和胫节磨擦逐步选择到达合适寄主。树皮粗糙程度、叶片和枝条特征及混交林密度对该天牛选择寄主过程中的爬行、飞翔和栖息行为有一定影响。刘博等(2012)发现松墨天牛在远距离寄主定位过程中，视觉信号起到了重要的作用；王健敏等(2012)对该虫进行嗜好颜色分析，发现雌、雄成虫均偏好选择棕褐色系，对应于林间衰弱和濒死寄主针叶呈现的颜色，说明该虫通过视觉判断偏好攻击长势衰弱的林木，属于次期性蛀干害虫；一系列的研究证明了视觉在松墨天牛成虫的寄主选择过程中发挥着重要作用。

同时，昆虫对寄主植物的定位和识别中还表现在雌虫对产卵场所选择性上。光肩星天牛雌虫产卵具有明显的寄主选择性(Morewood et al.，2003)，复眼具有一定的感光性(嵇保中等，1991; 张波等，1991)，但视觉在光肩星天牛觅食行为及将要产卵的雌虫对寄主的定位和识别中起的作用仍未见报道，视觉在天牛对寄主植物的定位和识别是否起着关键作用有待于深入研究。

3 灯诱在天牛综合治理中的应用

灯光诱杀在害虫综合治理(IPM)中占有重要地位，是农林业害虫物理防治中重要的手段之一(李志刚等，2012; 张纯胄等，2007)。在天牛的综合防治中也有报道，由于松墨天牛是松材线虫(Bursa-phelenchus xylophilus)的重要传播媒介(宁眺等，2004)，而且具有明显的趋光性，因此，对其进行灯光诱杀的报道相对较多，如梁细弟等(2000)利用高压汞灯和黑光灯对松墨天牛进行诱杀研究，得出高压汞灯对松墨天牛的诱集效果较好。然而，周成枚等(2002)却认为高压汞灯和黑光灯对松墨天牛的诱杀效果都较好。其他种类也有报道，如Mcintosh等(2001)设计了4种不同形状的诱捕器，林间诱捕结果表明： 在诱集的27 336只木材蛀干害虫中，有79%属于天牛科害虫、15%属于吉丁虫科害虫；其中最多的属为墨天牛属、脊虎天牛属；并发现交叉叶片类型的诱捕器效果是最好的；于永浩等(2009)发现蔗根土天牛(Dorysthenes granulosus)成虫对诱虫灯光源选择不严格，但上灯率和光源有关；对天牛(Arhopalus ferus)的灯光诱集研究中，发现紫外和黄色光源对该虫的诱集效果较好(Pawson et al.，2009a)。对栗山天牛成虫的研究，也发现其具有趋光性和在树干上聚集交尾习性，可以利用诱虫灯对其进行诱捕(高纯等，2008; 高国平，2001)。

灯光诱杀不仅在田间可以应用，同样也可以应用到出入境检验检疫测报上，如： Pawson等(2009b)为了降低熏蒸剂的使用，研究一种基于视觉“推-拉”策略的新型诱杀天牛的诱虫灯，经测试黄色和紫外灯可以有效的吸引天牛。因此，利用天牛对不同光谱的趋性反应特性，开发专一、高效的物理防治方法和测报用工具，可以为林业害虫的综合治理提供新策略和新方法，从而减少化学农药对生态环境的污染。

4 展望

目前，关于视觉对天牛行为调节的研究主要集中在求偶和觅食行为上，但昆虫的视觉功能不仅表现在求偶和觅食行为中，在其他行为中也起着不可忽视的作用，比如： 在躲避天敌方面，无翅桃蚜(Myzus persicae)不仅可以识别龟纹瓢虫(Propylaea japonica)的挥发物，而且可以识别龟纹瓢虫的体色(李为争等，2010)；在方向定位上，蜣螂(Scarabaeus zambesianus)在取得食物后为了快速的离开，利用黄昏时分天空辐射的偏振光进行导航，从而尽可能呈直线爬行以逃避其他蜣螂的竞争(Dacke et al.，2003)，一些沙漠蚁、蜜蜂(Apis)、家蝇(Musca domestica)等也可以利用偏振光进行导航(Labhart et al.，1999)。对于光周期影响昆虫滞育的研究报道则更多。但是，视觉是如何影响天牛避敌、定向等行为活动的研究未见报道，因此，该方面的研究需要进一步深化。

生境管理是近几年提出的一种保护性生物防治方法，主要是通过合理的搭配不同种植结构来提高生物多样性，增加害虫与天敌之间的相互作用从而提高整个生态系统的控害保益功能(Landis et al.，2000; 戈峰，1998; 尤民生等，2004; 赵紫华等，2013)。了解天牛及其天敌的嗜好和排斥颜色，在行道树或者混交林中搭配不同颜色的树种或者树干涂刷不同颜色(Goyer et al.，2004)，利用“排斥的颜色”将害虫排斥出去或者利用“吸引的颜色”将天敌吸引进来(Cook et al.，2006)，从而既达到了美观又起到防治害虫的作用，为调控天牛及其天敌的种群结构，综合管理害虫及其天敌打下理论基础。 同时，对天牛嗜好颜色的了解，还可以用于诱捕器外形的制作上，从而增强性信息素对天牛的诱集效果，如伍苏然等(2010)林间诱捕松墨天牛发现形状和颜色均能显著影响诱捕效果，柱形诱捕器显著优于棱形的，褐色诱捕器显著高于其他颜色。因此，对天牛视觉系统的研究既可以丰富感觉系统的理论知识，也可以为林业害虫综合治理提供理论支持和技术指导。

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