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.     

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。     

昆虫足的吸附机制

自然界中有许多昆虫足上都有吸附垫,这些垫子经过进化能吸附在各种表面上,并能在运动中控制吸附力。昆虫吸附垫要么是光滑的表皮垫子,要么是密布特殊的刚毛。昆虫这种“飞檐走壁”的能力来自于粘液、复杂机械系统以及生物系统之间的相互作用。文章主要参考国内外研究昆虫吸附机制的文献,综述了能在光滑表面上行走的昆虫的基本爬行原理,对其吸附机制进行了分类。并分析目前研究的主要内容,提出目前昆虫吸附机制要解决的问题。最后对吸附机制在仿生机械应用上作了展望。     

球孢白僵菌在红火蚁体表侵染的扫描电镜观察

利用扫描电镜观察了球孢白僵菌Beauveria bassiana Bb04菌株分生孢子对红火蚁Beauveria bassiana 工蚁体壁的侵染过程。结果表明: 分生孢子多分布在红火蚁工蚁节间膜、胸部的褶皱、气门、体壁的凹陷部位、刚毛窝附近, 以及着生较密刚毛的足上。萌发的分生孢子在节间膜以及体表缝隙、刚毛窝及刚毛稀少的凹陷部位、胸部褶皱和足胫节处入侵。分生孢子在附着12 h后开始萌发, 接种后18 h附着在节间膜处的孢子首先侵入成功, 接种后24 h刚毛窝附近孢子萌发入侵, 接种后60 h胸、腹和足等部位的孢子均成功穿透侵入表皮。分生孢子可以直接以芽管侵入表皮, 也可以产生附着胞再侵入。     

蚌螨腺毛和足爪的亚显微结构观察

利用扫描电镜分别对6种蚌螨的腺毛和足爪结构进行了观察,结果表明蚌螨的腺毛是由腺体、围腺片、刚毛和腺毛板共同构成的复合结构;对弯弓蚌螨的连续切片的观察表明,腺体是从体表延伸至体腔的消化道附近,由此认为蚌螨的腺体是由体壁皮层细胞演化而来。蚌螨足末除了爪外,还观察到跗节端部背面或腹面有体壁突及刚毛,因此,蚌螨足的步行结构不是仅由爪形成的简单结构,而是由爪、跗节末端背突或腹垫和刚毛组成的复合体。     

大壁虎与无蹼壁虎脚底刚毛结构与其黏附性能的比较研究

采用扫描电子显微镜和细胞组织学技术比较大壁虎(Gekko. gecko)和无蹼壁虎(Gekko. swinhonis)脚底刚毛的形态构筑, 并观测了其与黏附性能的关系. 扫描结果显示: 壁虎脚底的刚毛排列整齐而密集、方向规则, 尖端弯曲而分支. 其中, 大壁虎脚底刚毛排列呈束状, 头端分支间距较小(小于0.2~0.3 μm), 均弯向脚心, 角度规范(约10°), 末梢呈花托状膨大; 无蹼壁虎脚底刚毛两级分支明显, 分支间距较大(第一级分支大于0.5 μm, 第二级分支大于1 μm), 末梢呈卷须状, 多角度弯曲. 组织学检测发现, 大壁虎和无蹼壁虎的刚毛均为实体, 内容物丰满, 毛底部有大量上皮细胞、脂肪细胞、色素细胞和肌肉组织, 没有腺细胞. 功能性实验证实, 壁虎脚底毛形态构筑的规范性与其黏附性能正相关, 改变脚底刚毛的形态构筑可显著降低其黏附性能. 同时, 测试结果表明2种壁虎脚底毛的切向黏附力均大于其法向黏附力, 相对体重而言, 无蹼壁虎具有更强的黏附性能, 特别是其法向黏附性能更优于大壁虎. 壁虎刚毛的形态构筑和组织学特点为制备仿壁虎刚毛群提供了重要信息.     

臭蜣螂体壁表面结构及其与减粘脱附功能的关系

采用扫描电镜技术观察并描述了臭蜣螂Copris ochus头部、胸腹部和足部体表上的凹陷和刚毛的结构。发现有4种简单的凹陷,其三维结构、边缘类型和位于凹陷中央的带孔刚毛的长度在身体不同部位均有不同。体壁上还有中央具隆起的凹陷,其中心具带孔刚毛;或缺中心刚毛但在其半环状外边沿上具一小刚毛的凹陷。简单凹陷广泛分布于除腹部以外的各部分,但具中央隆起的凹陷仅出现在前胸背板,具半环状外边沿的凹陷出现在腿节上。在鞘翅纵沟纹的念珠状结中有管状开口。体壁上还有大小和形状不同的刚毛,它们不同于生于凹陷的刚毛。凹陷和刚毛的分布及形态构成体表的非光滑表面,作者探讨了这些结构与体表减粘脱附功能的关系。     

荒漠拟步甲科鳖甲族三种幼虫头壳色素区的观察

荒漠拟步甲科鳖甲族昆虫的幼虫具有相似的圆柱状体形,从形态上不易区别。以往对荒漠拟步甲幼虫的形态研究注重于上唇和足等部位的刚毛和小刺的特征,少有对头壳颜色的记载。通过在体视显微镜下观察室内饲养的鳖甲族昆虫(鞘翅目:拟步甲科)小胸鳖甲Microdera punctipennis Kaszab、光滑鳖甲Anatolica polita borealis Kaszab和细颈露颚甲Colposcelis microderoides microderoides Reitter的大龄幼虫,发现它们的头壳色素区呈现出不同的形状和大小,可方便地用于区分这3种幼虫。同时,测量结果显示幼虫头壳宽与前胸背板宽之比在这3种幼虫有显著差异。     

浙江省洞头拟裸长角虫兆属一新种（长角虫兆科：弹尾纲）

记述浙江省洞头岛拟祼长角虫兆属1无眼新种,岛屿拟裸长角虫兆Coecobrya islandica Shi&Pan sp.nov.。此新种的鉴定特征为无眼,弹器基具"光滑"刚毛,胫胕节内侧无"光滑"刚毛,下唇MREL1L2为光滑刚毛,X和X4为光滑刺状小刚毛,上唇基刚毛,腹部第II–IV节的感觉毛以及背部毛序。该新种与短毛拟裸长角虫兆Coecobrya brevis Xu et al.,2012最相似。本文给出了该新种的特征图及与相似种的详细特征比较。模式标本保存于台州学院生命科学学院和南京农业大学植保学院昆虫系。     

用扫描电镜观察链霉菌的孢子形态

用扫描电镜观察了分属10个类群、30个种的80株链霉菌的孢子形态。其表面结构可分为光滑、刺状、毛发和鳞片状类型。孢子的表面结构类型与孢子丝形态间具有一定的相关性。直形孢子丝所形成的孢子,其表面结构多为光滑型。螺旋形的孢子丝所形成的抱子,其表面结构,除光滑型外,尚有其他几种类型。气生菌丝体的颜色与孢子表面结构之间也存在一定的相关性。气生菌丝体呈灰色者,孢子表面结构的类型比较多。气生菌丝体呈其他颜色者,其孢子表面结构的类型则比较单一。     

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.     

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 design and biomechanics of friction-based releasable attachment devices in insects

Design of attachment devices in insects varies enormously inrelation to different functional loads. Many systems, locatedon different parts of the body, involve surfaces with particularfrictional properties. Such systems evolved to attach partsof the body to each other, or to attach an insect to the substratumby providing fast and reversible attachment/detachment. Amongthese systems, there are some that deal with predefined surfaces,and others, in which one surface remains unpredictable. Thefirst type of system occurs, for example, in wing-locking devicesand head-arresting systems and is called probabilistic fasteners.The second type is mainly represented by insect attachment padsof two alternative designs: hairy and smooth. The relationshipbetween surface patterns and/or mechanical properties of materialsof contact pairs results in two main working principles of thefrictional devices: mechanical interlocking, or maximizationof the contact area. We give an overview of the functional designof two main groups of friction-based attachment devices in insects:probabilistic fasteners and attachment pads.     

Microstructure of the tarsal attachment devices in the earwig Timomenus komarovi

The biological attachment device on the tarsal appendage of the earwig, Timomenus komarovi (Insecta: Dermaptera: Forficulidae) was investigated using field emission scanning electron microscopy to reveal the fine structural characteristics of its biological attachment devices to move on smooth and rough surfaces. They attach to rough substrates using their pretarsal claws; however, attachment to smooth surfaces is achieved by means of two groups of hairy tarsal pads. This biological attachment device consists of fine hairy setae with various contact sizes. Three different groups of tenent setae were distinguished depending on the cuticular substructure of the endplates. Two groups of setae commonly had flattened surfaces, and they were covered with either spoon‐shaped or spatula‐shaped endplates, respectively. While the flattened tip setae were distributed at the central region, the pointed tip setae were characteristically found along the marginal region. There were no obvious gender‐specific differences between fibrillar adhesive pads in this insect mainly because the forceps‐like pincers are used during copulation to grasp the partner.     

Functionally Different Pads on the Same Foot Allow Control of Attachment: Stick Insects Have Load-Sensitive “Heel” Pads for Friction and Shear-Sensitive “Toe” Pads for Adhesion

Stick insects (Carausius morosus) have two distinct types of attachment pad per leg, tarsal “heel” pads (euplantulae) and a pre-tarsal “toe” pad (arolium). Here we show that these two pad types are specialised for fundamentally different functions. When standing upright, stick insects rested on their proximal euplantulae, while arolia were the only pads in surface contact when hanging upside down. Single-pad force measurements showed that the adhesion of euplantulae was extremely small, but friction forces strongly increased with normal load and coefficients of friction were 1. The pre-tarsal arolium, in contrast, generated adhesion that strongly increased with pulling forces, allowing adhesion to be activated and deactivated by shear forces, which can be produced actively, or passively as a result of the insects'' sprawled posture. The shear-sensitivity of the arolium was present even when corrected for contact area, and was independent of normal preloads covering nearly an order of magnitude. Attachment of both heel and toe pads is thus activated partly by the forces that arise passively in the situations in which they are used by the insects, ensuring safe attachment. Our results suggest that stick insect euplantulae are specialised “friction pads” that produce traction when pressed against the substrate, while arolia are “true” adhesive pads that stick to the substrate when activated by pulling forces.     

Insect walking techniques on thin stems

The attachment ability of insects on surfaces are associated not only with the micro- and nanostructure of the adhering part of an attachment device, but also with the global scale kinematics responsible for contact formation and release. In the present study, the locomotory techniques of several representatives of insects from four different orders (Orthoptera, Heteroptera, Coleoptera, and Hymenoptera), possessing different types of attachment structures, are described. The study is based on video recordings of insects walking on a flat surface and on cylindrical rods of various thickness, imitating plant stems. Attachment devices of tarsi and pretarsi were visualized using Scanning Electron Microscopy. The results show a different manner in the use of adhesive structures on substrates with various curvatures. Insects bearing attachment pads on proximal tarsomeres usually touch flat and curved substrates using all tarsomeres, whereas insects with their attachment devices on the distal tarsomeres usually walk on flat surfaces using the distal tarsomeres of the overextended tarsus. On substrates, with diameters comparable to or larger than the tarsus length, insects walk above the stem by clasping the stem with the bent tarsi. On thin stems, insects clasp the stem between their tarsi and hang under the stem. Thus, on thin and thick rods, forces applied to attachment organs act in opposite directions. There are two methods of leg positioning for walking on a rough flat substrate. In the first case, the tarsus is straightened and the rough substrate is gripped between the claws and the proximal complex of attachment devices (tarsal euplantulae, fossulae spongiosa, and terminal spurs of tibiae). In the second case the tibia does not touch the substrate; the insect is supported only by distal tarsomeres. The tarsus is in an overextended condition. On rods, with diameters comparable to or larger than the tarsus length, insects walk by clasping the stem with the bent tarsi. This posture is characteristic for the majority of insects independent of the tarsal position they normally use while walking on a plane. If the rod’s diameter is smaller than the tarsus length, walking insects usually clutch it between contralateral tarsi. Using such a posture they are supported by interlocking or by strong friction, generated by attachment devices of the proximal tarsomeres, and do not use attachment devices of the pretarsus. Contact with the substrate is reinforced due to the coordinated contralateral clutch using all supporting legs. It is concluded that the use of different types of attachment structures correlates with locomotory techniques. Handling Editor: Heikki Hokkanen     

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.     

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.