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。     

Locomotion and adhesion: dynamic control of adhesive surface contact in ants

Tarsal adhesive pads of insects are highly dynamic organs that play an important role in locomotion. Many insects combine fast running performance with strong resistance to detachment forces. This capacity requires an effective control of attachment forces at the tarsus and pretarsus. Here we investigate mechanisms of attachment control in Asian weaver ants (Oecophylla smaragdina) by measuring the dynamics of the adhesive contact area and the claws during locomotion. O. smaragdina ants walking upside down on a smooth substrate used only a fraction (approx. 14%) of their maximum possible contact area. When these ants were loaded with 30 mg weights (corresponding to approx. 6 times their own body weight), however, they employed much larger (but still submaximal; approx. 60%) contact areas. The increase of contact area was accompanied by a stronger flexion of the claws, which demonstrates the participation of the claw flexor muscle in the control of adhesive contact. However, only part of the contact area dynamics could be explained by the action of the claw flexor. During the stance phase, adhesive contact area changed while the claws remained motionless. Even when corrected for the effects of claw flexion, adhesive contact areas differed by a factor of 2.1 between loaded and unloaded ants. Our findings give evidence that running ants control their adhesive contact area by a combination of active movements of the claw flexor muscle and passive reactions of the mechanical system.     

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.     

Mechanics of adhesion through a fibrillar microstructure

Many organisms have evolved a fibrillated interface for contactand adhesion as shown by several of the papers in this issue.For example, in the Gecko, this structure appears to give themthe ability to adhere and separate from a variety of surfacesby relying only on weak van der Waals forces. Despite the lowintrinsic energy of separating surfaces held together by vander Waals forces, these organisms are able to achieve remarkablystrong adhesion. To help understand adhesion in such a case,we consider a simple model of a fibrillar interface. For it,we examine the mechanics of contact and adhesion to a substrate.It appears that this structure allows the organism, at the sametime, to achieve good, ‘universal’ contact and adhesion.Due to buckling, a carpet of fibrils behaves like a plasticsolid under compressive loading, allowing intimate contact inthe presence of some roughness. As an adhesive, we conjecturethat energy in the fibrils is lost upon decohesion and unloading.This mechanism can add considerably to the intrinsic work offracture, resulting in good adhesion even with only van derWaals forces. Analysis of the mechanics of adhesion throughsuch a fibrillar interface provides rules for the design ofthe microstructure for desired performance as an adhesive.     

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。     

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.     

Ground reaction forces in vertically ascending beetles and corresponding activity of the claw retractor muscle on smooth and rough substrates

We measured ground reaction forces in fore–aft and normal directions of single hind and front legs in vertically ascending Sagra femorata beetles (Coleoptera, Chrysomelidae) on a smooth and a rough substrate. Simultaneously, we performed electromyographic recordings (EMGs) of the hind leg claw retractor muscle that partly controls the attachment structures. On both substrates, hind legs produced upward- as well as downward-directed forces during one stance phase. Forces were equivalent in both directions. Front legs generated only upward-directed forces. The main function of hind legs in ascending beetles in the second half of the stance thus probably prevented the animals from tilting away from the substrate. The EMGs of hind legs showed an early spike during stance with large amplitude. It was mostly followed by few additional spikes with large amplitude and in some cases of spikes with smaller amplitude distributed throughout the stance phase. We found significantly more spikes on the rough substrate than on the smooth one. This is probably due to the more important role of pretarsal claws than tarsal hairy attachment pads on the rough substrate or to the reduced adhesive forces on the rough substrate that have to be compensated by additional muscle activity.     

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。     

Scaling and biomechanics of surface attachment in climbing animals

Attachment devices are essential adaptations for climbing animals and valuable models for synthetic adhesives. A major unresolved question for both natural and bioinspired attachment systems is how attachment performance depends on size. Here, we discuss how contact geometry and mode of detachment influence the scaling of attachment forces for claws and adhesive pads, and how allometric data on biological systems can yield insights into their mechanism of attachment. Larger animals are expected to attach less well to surfaces, due to their smaller surface-to-volume ratio, and because it becomes increasingly difficult to distribute load uniformly across large contact areas. In order to compensate for this decrease of weight-specific adhesion, large animals could evolve overproportionally large pads, or adaptations that increase attachment efficiency (adhesion or friction per unit contact area). Available data suggest that attachment pad area scales close to isometry within clades, but pad efficiency in some animals increases with size so that attachment performance is approximately size-independent. The mechanisms underlying this biologically important variation in pad efficiency are still unclear. We suggest that switching between stress concentration (easy detachment) and uniform load distribution (strong attachment) via shear forces is one of the key mechanisms enabling the dynamic control of adhesion during locomotion.     

Grip and detachment of locusts on inverted sandpaper substrates

Locusts (Locusta migratoria manilensis) are characterized by their strong flying and grasping ability. Research on the grasping mechanism and behaviour of locusts on sloping substrates plays an important role in elucidating the mechanics of hexapod locomotion. Data on the maximum angles of slope at which locusts can grasp stably (critical angles of detachment) were obtained from high-speed video recordings at 215 fps. The grasping forces were collected by using two sensors, in situations where all left legs were standing on one and the right legs on the other sensor plate. These data were used to illustrate the grasping ability of locusts on slopes with varying levels of roughness. The grasping morphologies of locusts' bodies and tarsi were observed, and the surface roughness as well as diameters of their claw tips was measured under a microscope to account for the grasping mechanism of these insects on the sloping substrate. The results showed that the claw tips and part of the pads were in contact with the inverted substrate when the mean particle diameter was in the range of 15.3-40.5 μm. The interaction between pads and substrates may improve the stability of contact, and claw tips may play a key role in keeping the attachment reliable. A model was developed to explain the significant effects of the relative size of claw tips and mean particle diameter on grasping ability as well as the observed increase in lateral force (2.09-4.05 times greater than the normal force during detachment) with increasing slope angle, which indicates that the lateral force may be extremely important in keeping the contact reliable. This research lays the groundwork for the probable design and development of biomimetic robotics.     

CORRECTIONS, VOLUME 29

Part 1, under the frontispiece portrait of Dr. N. B. Eales,the words ‘President 1948–1951’ should havebeen added. Page 103, line 49, for ‘Newton Collection’ read‘Norman Collection (Canon Norman)’. 185, line 37, for ‘capillaris’ read ‘capillacca’. 188, Table 1, for ‘bemoralis’. read ‘nemoralis’. 188, Table 2, for ‘Cochlicella acuta (Müll)? ventrosa(Fér.)’ read ‘Cochlicella ventrosa (Fér.)’. 191, line 24, for ‘araheo-’ read ‘archeo-’.     

Adhesion forces measured at the level of a terminal plate of the fly's seta

The attachment pads of fly legs are covered with setae, each ending in small terminal plates coated with secretory fluid. A cluster of these terminal plates contacting a substrate surface generates strong attractive forces that hold the insect on smooth surfaces. Previous research assumed that cohesive forces and molecular adhesion were involved in the fly attachment mechanism. The main elements that contribute to the overall attachment force, however, remained unknown. Multiple local force-volume measurements were performed on individual terminal plates by using atomic force microscopy. It was shown that the geometry of a single terminal plate had a higher border and considerably lower centre. Local adhesion was approximately twice as strong in the centre of the plate as on its border. Adhesion of fly footprints on a glass surface, recorded within 20 min after preparation, was similar to adhesion in the centre of a single attachment pad. Adhesion strongly decreased with decreasing volume of footprint fluid, indicating that the layer of pad secretion covering the terminal plates is crucial for the generation of a strong attractive force. Our data provide the first direct evidence that, in addition to Van der Waals and Coulomb forces, attractive capillary forces, mediated by pad secretion, are a critical factor in the fly's attachment mechanism.     

The Nitrogen Content of Plants and the Self-thinning Rule of Plant Ecology: A Test of the Core-skin Hypothesis

The ‘core-skin’ hypothesis postulates that secondarilythickened plants behave energetically as an inert ‘core’covered by an active ‘skin’, the ‘skin’being two-imensional, the ‘core’ three-dimensional.This would explain the ‘self-thinning ‘or‘–3/2’ rule of plant ecology, that is, the tendencyfor log (dry weight per plant) and log (number of plants perunit area) to progress along a straight line relationship, withslope = – 3/2’. The hypothesis was tested as follows. Plant nitrogen contentwas used as an estimate of the mass of ‘skin’ perplant, and dry weight as an estimate of the mass of the ‘core’.As plants mature the slope of the relationship between y = log(mass of nitrogen per plant) and x = log (mass of dry matterper plant) is expected to decline from an initial value of 1.0towards a final value of 0.66. The intercept of the relationshipis expected to reflect the intrinsic content of ‘skin’per unit of ‘core’. Genotypic variation in thisparameter should cause genotypic differences in the maximumattainable yield of biomass per unit area. The expectations were investigated by fitting the function y= p+qx+r exp – x to 30 sets of data on plant nitrogencontent, plant weight and time in 18 different vegetables. Simplelinear regressions of y on x were fitted to more limited setsof data on weights and nitrogen contents of mature trees. Theexpectations were, with some minor exceptions, confirmed. Nitrogen, yield, plant competition, self-thinning     

Ultrastructural Aspects of Graft Incompatibility Between Pear and Quince In vitro

Callus cells of pear (Pyrus communis cv. ‘Bartlett’)underwent lethal cellular senescence in response to graftingwith callus cells of quince (Cydonia oblonga cv. ‘VanDeman’). Similar responses occurred when the cells werein direct contact and when they were separated by a porous membranefilter. These results indicate that direct cellular contactis not necessary to elicit graft incompatibility between pearand quince. Cydonia oblonga, grafting, incompatibility, pear, Pyrus communis, quince     

Factors Affecting the Abscission of Reproductive Organs in Yellow Lupins (Lupinus luteus L.): III. ENDOGENOUS GROWTH SUBSTANCES IN VIRUS-INFECTED AND HEALTHY PLANTS AND THEIR EFFECT ON ABSCISSION

Extracts of small and mature-size lupin pods yielded four substancesaffecting the growth of wheat-coleoptile sections: one acidpromotor (A), two acid inhibitors(B and X), and one neutralinhibitor(Y). Inhibitor B was extremely active, however, coleoptile sectionsshowed no signs of toxic effects; they resumed growth at a rapidrate after rinsing them and adding ß-indolylaceticand (IAA) to the medium. 1 µg of IAA was required to counteractthe effect of ‘B’ extracted from 230 mg. Of tissue.On an equal fresh weight basis the inhibiting action of ‘B’in lupin pods was 500–1,500 times more potent than thatof ‘inhibitor ß’ in etiolated pea seedlings. Small pods of plants infected with pea-mosaic virus yielded3 times the amount of ‘A’ of healthy plants (equivalentto 1 µg. IAA 0.3 µg. IAA per 25 g. of tissue respectively),and approximately the amount of ‘B’. Mature podsof virus-infected plants again yielded more‘A’,but also 2? times more ‘B’ than pods of healthyplants. Healthy pods yielded more ‘A’ than virus-infectedpods, and there was no difference in ‘X’. A lupin abscission test was developed and the effects of proximaland distal application of -naphthyl acetic acid (NAA) are presented,and discussed with respect to results of other abscission tests. ‘A’ accelerated abscission when applied proximally,and delayed or prevented it when applied distally. ‘B’strongly accelerated abscission when applied in either way.A possible mechanism explaining the abscission-inducing effectof developing pods on later flowers is discussed in terms ofthe substances ‘A’ and ‘B’. The partlyprevented abscission observed on virus-infected plants was foundto agree well with the proposed mechanism.     

Developmental Genetics and Morphological Evolution of Flowering Plants, Especially Bladderworts (Utricularia): Fuzzy Arberian Morphology Complements Classical Morphology

This review compares new developmental models on flowering andother vascular plants with evolutionary hypotheses formulatedby Agnes Arber (1879–1960) and like-minded botanists.Special emphasis is laid on philosophical basics such as perspectivism,pluralism about evolutionary modelling, continuum way of thinking,and fuzzy logic. Arber's perspective is best labelled as F uzzyA rberian M orphology (FAM Approach). Its proponents (‘FAMmers’)treat structural categories (e.g. ‘roots’, ‘shoots’,‘stems’, ‘leaves’, ‘stipules’)in vascular plants as concepts with fuzzy borderlines allowingintermediates (including transitional forms, developmental mosaics).The FAM Approach complements Cla ssical Plant M orphology (ClaMApproach), which is the traditional approach in botany. ClaMproponents (‘ClaMmers’) postulate that the structuralcategories of vascular plants are regarded as concepts withclear-cut borderlines and without intermediates. However, duringthe evolution of vascular plants, the root-shoot distinctionand the stem-leaf distinction have become blurred several timesdue to developmental changes, resulting in organs with uniquecombinations of features. This happened, for example, in thebladderworts (Utricularia, Lentibulariaceae). When focusingon the ‘leaf’, the FAM Approach is identical toArber's ‘partial-shoot theory of the leaf’ and Sinha's‘leaf shoot continuum model’. A compound leaf canrepeat the developmental pathway of the whole shoot, at leastto some degree. For example, compound leaves of Chisocheton(Meliaceae)with indeterminate apical growth and three-dimensional branchingmay be seen as developmental mosaics sharing some growth processeswith whole shoots! We focus here on the FAM Approach becausethis perspective is especially promising for developmental geneticistsstudying flowering and other vascular plants. Copyright 2001Annals of Botany Company Review, body plan, developmental mosaics, leaf development, history of botany, homeosis, homeotic genes, Lentibulariaceae, morphological evolution, process morphology, stipules, Utricularia, flowering plants     

Diurnal Phototropism in Leaves of Lavatera cretica L. under Conditions of Simulated Solar-Tracking

Diurnal laminar reorientation was followed in solar-trackingleaves of Lavatera cretica L. under simulated conditions. Asimulated ‘sun’ was moved over the lamina in a 180?arc in the vertical plane of the mid-vein, at an angular velocityof 15? h^–1 in a regime of 12-h photoperiods. In one groupof plants the petioles of the experimental leaves were arrangedto face ‘sunrise’, while in the other they werearranged to face ‘sunset’. At ‘sunrise’,the laminae in both groups, which were inclined towards theanticipated direction of ‘sunrise’, changed theirelevation towards the rising ‘sun’, resulting inprogressive reduction in the angle of incidence (AI) of lighton the laminar surface (AI= differential between laminar and‘solar’ elevation). As a result, laminar and ‘solar’elevation converged, and laminar reorientation gradually ceased,until the ‘solar’ elevation had passed the normalto the laminar surface (AI=0?). laminar reorientation was thenresumed, but its direction was reversed to follow the directionof ‘solar’ reorientation. During most of the remaining‘day’, laminar elevation (LE) trailed that of the‘sun’ by an average of 11?-14?. Laminar reorientationthen anticipated ‘sunset’ by starting to slow down60 to 90min in advance. During the 12-h dark period, the laminareoriented towards the anticipated direction of the subsequent‘sunrise’. The time-course of nocturnal reorientationwas qualitatively different in the two groups of experimentalplants. The time-course of diurnal phototropism under naturaland simulated conditions is analysed and compared and differencesand similarities between them are discussed. Key words: Diurnal phototropism, solar-tracking, vectorial excitation     

Relative Sensitivity and Tolerance of some Gladiolus Cultivars to Sulphur Dioxide

Five Gladiolus cultivars, namely ‘Aldebaran’, ‘BrightEye’, ‘Illusion’, ‘Manisha’ and‘Manmohan’, were exposed to 1 and 2 µg l^–1sulphur dioxide to test their relative-sensitivity toleranceto the pollutant Plants were fumigated experimentally for 2h daily Foliar injury symptoms were observed first in ‘Manisha’followed by ‘Aldebaran’ and ‘Illusion’at the higher dose Photosynthetic pigments and leaf extractpH were significantly decreased, particularly in ‘Manisha’and ‘Illusion’ Overall disturbances in the plantmetabolism due to SO₂ treatment led to retarded growth of plants,as evident from decreased shoot length and phytomass valuesThe order of sensitivity of the five Gladiolus cultivars toSO₂ was as follows, with the greatest first Manisha, Illusion,Aldebaran, Bright Eye, Manmohan Cultivars, Gladiolus, sensitivity, sulphur dioxide, tolerance     

Investigations on the Control of Lettuce Seed Germination at High Temperatures

The activity of sodium hypochlorite solution in relieving thermo-inhibitionof germination of lettuce seed is shown to be due to its chlorinecontent. Results of experiments in which the pericarp, and pericarpand endosperm were removed, together with direct measurementsof penetration forces, suggest that the relief of thermo-inhibitionresults from weakening of the pericarp by the hypochlorite.Differences between the cultivars ‘Cobham Green’and ‘Grand Rapids’ in the contributions made bypericarp and endosperm to germination control at 35 °C aredemonstrated. Key words: Lactuca sativa L, Chlorine, Thermo-inhibition     

Friction ridges in cockroach climbing pads: anisotropy of shear stress measured on transparent,microstructured substrates

The contact of adhesive structures to rough surfaces has been difficult to investigate as rough surfaces are usually irregular and opaque. Here we use transparent, microstructured surfaces to investigate the performance of tarsal euplantulae in cockroaches (Nauphoeta cinerea). These pads are mainly used for generating pushing forces away from the body. Despite this biological function, shear stress (force per unit area) measurements in immobilized pads showed no significant difference between pushing and pulling on smooth surfaces and on 1-μm high microstructured substrates, where pads made full contact. In contrast, on 4-μm high microstructured substrates, where pads made contact only to the top of the microstructures, shear stress was maximal during a push. This specific direction dependence is explained by the interlocking of the microstructures with nanometre-sized “friction ridges” on the euplantulae. Scanning electron microscopy and atomic force microscopy revealed that these ridges are anisotropic, with steep slopes facing distally and shallow slopes proximally. The absence of a significant direction dependence on smooth and 1-μm high microstructured surfaces suggests the effect of interlocking is masked by the stronger influence of adhesion on friction, which acts equally in both directions. Our findings show that cockroach euplantulae generate friction using both interlocking and adhesion.