昆虫足的附着机制

昆虫卓越的爬行和附着能力来源于其精细的功能性黏附系统。根据形态结构的不同,昆虫的黏附系统可分为光滑型黏附垫和刚毛型黏附垫两种类型,二者在分泌液的支持下均能附着于几乎所有的光滑或粗糙的物体表面,而且这两种类型的黏附垫与界面的附着的形成均主要依赖于范德华力。本文综述了昆虫足的附着机制,包括光滑型和刚毛型两种黏附垫的结构和其形成附着的机理,以及黏附垫分泌液的功能、组成成分和释放机制,阐明了昆虫如何巧妙地解决稳定附着和快速脱附这一矛盾的问题,讨论了诸如界面的理化性质和环境湿度等环境因素对昆虫附着的影响,以期帮助人们深入地理解昆虫足的附着机制,并为其在仿生学等方面的应用提供理论依据。     

螽斯吸附足垫的构造及其吸附性能分析

许多昆虫足上有光滑吸附垫,通过二相分泌液粘附到各种表面。为理解这种基于液体的吸附系统的功能,用在螽斯身上绑细线的方法,测量其在不同表面的摩擦力和吸附力,并用高速摄像机观察足垫的构造及吸附和分离的动作,测试足垫与接触面的接触面积。结果表明螽斯的水平摩擦力大于垂直吸附力。足垫与表面接触时向身体方向拖动来增加摩擦力。分离时采用剥离的方法,但剥离方向与刚毛型足垫的相反,是从末梢端翘起分离,达到行动迅速且节省能量的目的。测试结果可用于机器人吸附足掌的仿生设计。     

昆虫抗冻蛋白的研究

抗冻蛋白是具有热滞活性,能结合并抑制冰晶生长和抑制冰的重结晶的一类蛋白质。近几年来,昆虫抗冻蛋白的研究取得了较快的发展,本文通过分析昆虫抗冻蛋白的结构特点、抗冻活性、作用机制,并讨论了抗冻蛋白在食品工业、医学、基因工程方面的应用。结果表明,昆虫抗冻蛋白虽然结构呈多样性,但有很多关键的残基具有保守性,对维持抗冻蛋白结构和功能的完整性发挥着重要的作用;抗冻蛋白是由多基因家簇编码的。其作用机制主要是吸附一抑制机制,抗冻蛋白依靠氢键吸附到冰晶格,抑制冰晶生长;昆虫抗冻蛋白的应用具有很广阔的前景。     

昆虫性别决定机制研究进展

阐述昆虫的性别决定机制是理解昆虫性别分化调控的理论基础,也为人类有效控制害虫开辟了新方向。昆虫性别决定机制存在复杂性和多样性,但主要是内因即性别决定基因级联互作调控的结果。本文对近年来基于性别决定基因级联互作的昆虫性别决定机制研究进行了综述,主要包括性别决定基因概况和重要性别决定相关基因的分子级联互作关系。目前发现昆虫重要性别决定相关基因主要集中在常染色体上,且部分基因之间存在紧密的级联互作,如Sxl,tra,dsx,csd和fem等。在这些基因中,tra/fem→dsx的调控模式在已报道的昆虫中存在共性,即tra和dsx相对较保守且tra通过性特异剪切来调控下游dsx的转录形式。目前大多数昆虫的性别决定机制还不清楚,但近年来模式昆虫性别决定机制取得了一定进展,对非模式昆虫的研究还处于起步阶段但却越来越受到重视。     

昆虫迁飞的调控基础及展望

昆虫迁飞是在长期适应多变的环境过程中进化形成的一种行为对策,也是昆虫的种类和数量繁多,以及迁飞害虫经常暴发成灾的主要原因.昆虫迁飞行为的发生不仅受到外界环境因素的影响,而且受到本身生理因素的调控.目前,国内外对此类研究主要集中在生态环境、生理因素、行为学以及种群遗传学方面的调控机制.随着分子生物学技术的发展,昆虫迁飞行为发生的分子调控机制也越来越受到重视.在对国内外主要昆虫迁飞调控机制概述的基础上,对新的分子生物学技术在昆虫迁飞调控中的应用进行了探讨与展望.     

三种蝇类昆虫(双翅目)足爪垫结构的超微形态研究

利用扫描电子显微镜对双翅目3种蝇类昆虫即家蝇Musca domestica L.(蝇科)、红尾粪麻蝇Bercaea cruentata(Meigen)(麻蝇科)和肥躯金蝇Chrysomya pinguis (Walker)(丽蝇科)足的爪垫超微形态结构进行了研究,研究发现每种蝇类的前、中、后足爪垫的面积存在一定的差别,中足和后足爪垫面积较前足大;爪垫腹面均密被刚毛,每根刚毛由刚毛杆(setal shaft)和末端平板(terminal plate)组成,刚毛主要有铲状、勺状和柳叶状3种类型;后足爪垫上刚毛的密度和刚毛末端面积一般小于前、中足;位于爪垫边缘处的刚毛较长.研究还发现爪垫上的刚毛均为中空结构,且丽蝇科昆虫的刚毛末端具开口.     

非嗜食植物中的昆虫产卵驱避物及其利用

产卵是植物性昆虫生命周期中的一个重要环节,它能反映昆虫与植物相互作用的某些特点以及植食性昆虫对植物利用的策略.植物中的驱避物质在调节昆虫产卵行为过程中起着十分重要的作用.大量研究结果表明:许多非嗜食植物含有对昆虫产卵有驱避作用的次生化合物.研究植物中的昆虫产卵驱避物质不仅能在理论上加深对植食性昆虫产卵机制,植食性昆虫与植物间的相互关系以及昆虫群落构建机制等的认识,同时在害虫综合治理中有广泛的应用前景.     

植物挥发性气体(VOCs)研究进展

植物挥发性气体(VOCs)在植物一植食性昆虫-天敌三级营养关系、植物间信息传递及适应性改变上都发挥着重要作用.植物释放VOCs具特异性、系统性、时序性与节律性等特点,VOCs主要在寄主选择行为、产卵行为、求偶行为、引来昆虫夭敌干涉等方面影响植食性昆虫.VOCs-介导的植物间信息传递作用包括4个过程:"释放者"植物合成及释放气体、气体在空气中的运输、气体在植物表面的吸附及"接收者"植株对气体信号的感知.收集VOCs的方法主要有吸附-溶剂洗脱法和吸附-热脱附法.     

昆虫中肠Bt晶体蛋白受体的研究进展

苏云金芽孢杆菌Ｂａｃｉｌｌｕｓｔｈｕｒｉｎｇｉｅｎｓｉｓ杀虫作用的主要成份是胞内产生的伴孢晶体,晶体蛋白经昆虫吞食,在肠道降解为激活的毒性肽。普遍认为毒性肽的作用机制主要有两个步骤：１）与中肠表面的受体专一结合;２）在细胞膜上形成跨膜通道。杀虫晶体蛋白的专一性与中肠细胞膜表面的受体蛋白紧密相连,晶体蛋白的杀虫作用是通过昆虫中肠细胞的专一性受体而起作用。本文通过说明受体蛋白的生物学特性、分子本质及与昆虫抗性的关系,概述了近年来中肠受体蛋白的研究进展。１　昆虫中肠受体蛋白的生物学特性１１　受体蛋白…     

昆虫联想学习记忆研究的几种双通道范式

昆虫的大多数行为一直被认为是生来就有的本能行为,然而,近年来的研究已经证明大多数昆虫具有高度的学习记忆能力,并表现出对环境的适应行为。这些研究主要是采取了联想学习记忆的双通道范式。这些双通道范式,在学习记忆神经机制的研究中起到了重要的作用。本文主要综述近年来关于昆虫学习记忆研究的进展,重点介绍昆虫联想学习记忆研究的几种双通道范式。这对充分理解昆虫联想学习记忆的神经机制以及进一步深入开展学习与记忆功能的研究有重要的科学意义。     

Adhesive Contact in Animal: Morphology, Mechanism and Bio-Inspired Application

Many animals possess adhesive pads on their feet,which are able to attach to various substrates while controlling adhesive forces during locomotion.This review article studies the morphology of adhesive devices in animals,and the physical mechanisms of wet adhesion and dry adhesion.The adhesive pads are either ‘smooth' or densely covered with special adhesive setae.Smooth pads adhere by wet adhesion,which is facilitated by fluid secreted from the pads,whereas hairy pads can adhere by dry adhesion or wet adhesion.Contact area,distance between pad and substrate,viscosity and surface tension of the liquid filling the gap between pad and substrate are the most important factors which determine the wet adhesion.Dry adhesion was found only in hairy pads,which occurs in geckos and spiders.It was demonstrated that van der Waals interaction is the dominant adhesive force in geckos' adhesion.The bio-inspired applications derived from adhesive pads are also reviewed.     

Structural correlates of increased adhesive efficiency with adult size in the toe pads of hylid tree frogs

Tree frogs are able to climb smooth, vertical substrates using specialised toe pads which adhere via an area-based wet adhesive mechanism. Although the link between pads and arboreality in frogs is well-established, few studies have investigated the influence of morphology on adhesion. Trinidadian tree frogs from the genus Hyla are geometrically similar. There is a tendency towards comparatively reduced mass in larger species, but toe pad area increases as expected with isometry. As adhesion is area-dependent, forces are affected directly by the increase in mass relative to pad area, and there is a decrease in the ability of larger species to adhere to smooth rotation platforms. However, there is an increase in force per unit area that suggests larger species have more efficient toe pads. Toe pad structure is very similar though there are variations in the details of a number of features. Crucially, although differences in morphology appeared small they had demonstrable effects on adhesive efficiency of the pads. Epithelial cell area correlates positively with frog length and adhesive efficiency, related features of cell density and intercellular channel length correlate negatively. These findings are discussed in relation to the different forces involved in the tree frogs’ wet adhesive system.     

Adhesive and frictional properties of tarsal attachment pads in two species of stick insects (Phasmatodea) with smooth and nubby euplantulae

In the present study, the tarsal attachment pads (euplantulae) of two stick insect species (Phasmatodea) were compared. While the euplantulae of Cuniculina impigra (syn. Medauroidea extradentata) are smooth, those of Carausius morosus bear small nubs on their surfaces. In order to characterize the adhesive and frictional properties of both types of euplantulae, adhesion and friction measurements on smooth (Ra=0.054 μm) and rough (Ra=1.399 μm) substrates were carried out. The smooth pads of C. impigra generated stronger adhesion on the smooth substrate than on the rough one. The adhesive forces of the structured pads of C. morosus did not differ between the two substrates. Friction experiments showed anisotropy for both species with higher values for proximal pulls than for distal pushes. In C. impigra, friction was stronger on the smooth than on the rough surface for both directions, whereas in C. morosus friction was stronger on the smooth surface only for pushes. This shows that smooth attachment pads are able to generate relatively stronger adhesion and friction on a flat smooth surface than on a rough one. In contrast, nubby pads have similar adhesion on both substrates, and also show no difference in friction in the pulling direction. This leads to the conclusion that smooth pads are specialized for rather smooth substrates, whereas nubby pads are better adapted to generate stronger forces on a broader range of surfaces.     

Self-Drying: A Gecko's Innate Ability to Remove Water from Wet Toe Pads

When the adhesive toe pads of geckos become wet, they become ineffective in enabling geckos to stick to substrates. This result is puzzling given that many species of gecko are endemic to tropical environments where water covered surfaces are ubiquitous. We hypothesized that geckos can recover adhesive capabilities following exposure of their toe pads to water by walking on a dry surface, similar to the active self-cleaning of dirt particles. We measured the time it took to recover maximum shear adhesion after toe pads had become wet in two groups, those that were allowed to actively walk and those that were not. Keeping in mind the importance of substrate wettability to adhesion on wet surfaces, we also tested geckos on hydrophilic glass and an intermediately wetting substrate (polymethylmethacrylate; PMMA). We found that time to maximum shear adhesion recovery did not differ in the walking groups based on substrate wettability (22.7±5.1 min on glass and 15.4±0.3 min on PMMA) but did have a significant effect in the non-walking groups (54.3±3.9 min on glass and 27.8±2.5 min on PMMA). Overall, we found that by actively walking, geckos were able to self-dry their wet toe pads and regain maximum shear adhesion significantly faster than those that did not walk. Our results highlight a unexpected property of the gecko adhesive system, the ability to actively self-dry and recover adhesive performance after being rendered dysfunctional by water.     

Biomechanics of smooth adhesive pads in insects: influence of tarsal secretion on attachment performance

Many insects possess smooth adhesive pads on their legs, which adhere by thin films of a two-phasic secretion. To understand the function of such fluid-based adhesive systems, we simultaneously measured adhesion, friction and contact area in single pads of stick insects (Carausius morosus). Shear stress was largely independent of normal force and increased with velocity, seemingly consistent with the viscosity-effect of a continuous fluid film. However, measurements of the remaining force 2 min after a sliding movement show that adhesive pads can sustain considerable static friction. Repeated sliding movements and multiple consecutive pull-offs to deplete adhesive secretion showed that on a smooth surface, friction and adhesion strongly increased with decreasing amount of fluid. In contrast, pull-off forces significantly decreased on a rough substrate. Thus, the secretion does not generally increase attachment but does so only on rough substrates, where it helps to maximize contact area. When slides were repeated at one position so that secretion could accumulate, sliding shear stress decreased but static friction remained clearly present. This suggests that static friction which is biologically important to prevent sliding is based on non-Newtonian properties of the adhesive emulsion rather than on a direct contact between the cuticle and the substrate.     

An integrative study of insect adhesion: mechanics and wet adhesion of pretarsal pads in ants

Many animals that locomote by legs possess adhesive pads. Suchorgans are rapidly releasable and adhesive forces can be controlledduring walking and running. This capacity results from the interactionof adhesive with complex mechanical systems. Here we presentan integrative study of the mechanics and adhesion of smoothattachment pads (arolia) in Asian Weaver ants (Oecophylla smaragdina).Arolia can be unfolded and folded back with each step. Theyare extended either actively by contraction of the claw flexormuscle or passively when legs are pulled toward the body. Regulationof arolium use and surface attachment includes purely mechanicalcontrol inherent in the arrangement of the claw flexor system. Predictions derived from a ‘wet’ adhesion mechanismwere tested by measuring attachment forces on a smooth surfaceusing a centrifuge technique. Consistent with the behavior ofa viscid secretion, frictional forces per unit contact arealinearly increased with sliding velocity and the increment stronglydecreased with temperature. We studied the nature and dimensions of the adhesive liquidfilm using Interference Reflection Microscopy (IRM). Analysisof ‘footprint’ droplets showed that they are hydrophobicand form low contact angles. In vivo IRM of insect pads in contactwith glass, however, revealed that the adhesive liquid filmnot only consists of a hydrophobic fluid, but also of a volatile,hydrophilic phase. IRM allows estimation of the height of theliquid film and its viscosity. Preliminary data indicate thatthe adhesive secretion alone is insufficient to explain theobserved friction and that rubbery deformation of the pad cuticleis involved.     

Functional demands of dynamic biological adhesion: an integrative approach

Climbing organisms are constantly challenged to make their way rapidly and reliably across varied and often novel terrain. A diversity of morphologically and mechanically disparate attachment strategies have evolved across widely distributed phylogenetic groups to aid legged animals in scaling these surfaces, notable among them some very impressive adhesive pads. Despite the differences between, for example, the dry fibrillar pads of geckos and the smooth, secretion-aided pads of stick insects, I hypothesize that they face similar functional demands in their environment. I outline three broad criteria defining dynamic biological adhesion: reusability, reversibility, and substrate tolerance. Organismal adhesive pads must be able to attach repeatedly without significant decline in performance, detach easily at will, and adhere strongly to the broadest possible range of surfaces in their habitat. A survey of the literature suggests that evidence for these general principles can be found in existing research, but that many gaps remain to be filled. By taking a comparative, integrative approach to biological dynamic adhesion, rather than focusing on a few model organisms, investigators will continue to discover new and interesting attachment strategies in natural systems.     

Underwater locomotion in a terrestrial beetle: combination of surface de-wetting and capillary forces

For the first time, we report the remarkable ability of the terrestrial leaf beetle Gastrophysa viridula to walk on solid substrates under water. These beetles have adhesive setae on their feet that produce a secretory fluid having a crucial role in adhesion on land. In air, adhesion is produced by capillary forces between the fluid-covered setae and the substrate. In general, capillary forces do not contribute to adhesion under water. However, our observations showed that these beetles may use air bubbles trapped between their adhesive setae to walk on flooded, inclined substrata or even under water. Beetle adhesion to hydrophilic surfaces under water was lower than that in air, whereas adhesion to hydrophobic surfaces under water was comparable to that in air. Oil-covered hairy pads had a pinning effect, retaining the air bubbles on their feet. Bubbles in contact with the hydrophobic substrate de-wetted the substrate and produced capillary adhesion. Additional capillary forces are generated by the pad's liquid bridges between the foot and the substrate. Inspired by this idea, we designed an artificial silicone polymer structure with underwater adhesive properties.