Walking on smooth or rough ground: passive control of pretarsal attachment in ants

The hymenopteran tarsus is equipped with claws and a movable adhesive pad (arolium). Even though both organs are specialised for substrates of different roughness, they are moved by the same muscle, the claw flexor. Here we show that despite this seemingly unfavourable design, the use of arolium and claws can be adjusted according to surface roughness by mechanical control. Tendon pull experiments in ants (Oecophylla smaragdina) revealed that the claw flexor elicits rotary movements around several (pre-) tarsal joints. However, maximum angular change of claws, arolium and fifth tarsomere occurred at different pulling amplitudes, with arolium extension always being the last movement. This effect indicates that arolium use is regulated non-neuronally. Arolium unfolding can be suppressed on rough surfaces, when claw tips interlock and inhibit further contraction of the claw flexor or prevent legs from sliding towards the body. To test whether this hypothesised passive control operates in walking ants, we manipulated ants by clipping claw tips. Consistent with the proposed control mechanism, claw pruning resulted in stronger arolium extension on rough but not on smooth substrates. The control of attachment by the insect claw flexor system demonstrates how mechanical systems in the body periphery can simplify centralised, neuro-muscular feedback control.     

Origin and pathway of the epidermal secretion in the damselfly head-arresting system (Insecta: Odonata)

In damselflies, the arrester system is responsible for an additional attachment of the head to the neck. It consists of a pair of mobile postcervical sclerites (SPC) covered by microtrichia. In their lateral position, SPCs can fixate the head on fields of microtrichia on the back surface of the head. The intact surface of microtrichia of the SPC is usually covered by a lipid-containing secretion. The present study provides ultrastructural data on the secretory epidermis and pore channels adapted to transport the secretion to the cuticle surface. 1) Shock-frozen preparations of the contact area show that microtrichia of co-opted surfaces do not interlock with each other. When co-opted fields are pressed to each other, deformations of flexible microtrichia of the SPC result in an increase of contact area between corresponding structures and consequently in an increase of frictional forces. 2) In the area of the SPC, only electron-lucent vesicles have been found. Histochemical procedures revealed that the material stored in vesicles and liberated on the external surface of the SPC is presumably non-volatile lipid. 3) Only a small number of large pore channels reach the surface of a microtrichium. Most of them terminate with tiny terminal channels, whose diameter is approximately six to twenty times smaller than the diameter of structures of secretory blooms occurring in SEM in shock-frozen and air-dried preparations. The terminal channels do not penetrate the epicuticular layer. It seems that the secretion reaches the epicuticle through terminal channels and diffuses through the epicuticle without any channel structures.     

Friction and adhesion in the tarsal and metatarsal scopulae of spiders

Friction and adhesion forces of the ventral surface of tarsi and metatarsi were measured in the bird spider Aphonopelma seemanni (Theraphosidae) and the hunting spider Cupiennius salei (Ctenidae). Adhesion measurements revealed no detectable attractive forces when the ventral surfaces of the leg segments were loaded and unloaded against the flat smooth glass surface. Strong friction anisotropy was observed: friction was considerably higher during sliding in the distal direction. Such anisotropy is explained by an anisotropic arrangement of microtrichia on setae: only the setal surface facing in the distal direction of the leg is covered by the microtrichia with spatula-like tips. When the leg is pushed, the spatula-shaped tips of microtrichia contact the substrate, whereas, when the leg is pulled over a surface, setae bend in the opposite direction and contact the substrate with their spatulae-lacking sides. In an additional series of experiments, it was shown that desiccation has an effect on the friction force. Presumably, drying of the legs results in reduction of the flexibility of the setae, microtrichia, spatulae, and underlying cuticle; this diminishes the ability to establish proper contact with the substrate and thus reduces the contact forces.     

The jumping mechanism of cicada Cercopis vulnerata (Auchenorrhyncha, Cercopidae): skeleton-muscle organisation, frictional surfaces, and inverse-kinematic model of leg movements

In Auchenorrhyncha, jumping is achieved by metathoracic muscles which are inserted into the trochanter of the hind leg. The synchronisation of movements of the hind legs is a difficult problem, as the leg extension that produces the jump occurs in less than 1 ms. Even slight asynchrony could potentially result in failure of a jump. Both the synchronisation of the movements of a pair of jumping legs, and their stabilisation during a jump, seem to be important problems for small jumping insects. The present study was performed in order to clarify some questions of the functional morphology of the leafhopper jumping mechanism. It is based on skeleton-muscle reconstruction, high-speed video recordings, transmission (TEM) and scanning electron microscopic (SEM) investigations of the cuticle, together with 3D inverse-kinematic modelling of angles and working zones of hind leg joints of cicada Cercopis vulnerata (Cercopidae). The complete extension of the hind leg takes less than 1 ms, which suggests that the jump is powered not only by the muscle system, but also by an elastic spring. Histological staining and fluorescence microscopy showed resilin-bearing structures, responsible for elastic energy storage, in the pleural area of the metathorax. Synchronisation of hind leg movements may be aided by microtrichia fields that are located on the medial surface of each hind coxa. In Auchenorrhyncha, hind coxae are rounded in their anterior and lateral parts, whereas medial parts are planar, and contact each other over a rather large area. The inverse-kinematic model of propulsive leg movements was used to draw the surface outlined by the medial surface of the coxa, during the jump movement. This is a cone surface, faced with its bulged-in side, medially. Surfaces outlined by the movements of both right and left coxae overlap in their anterior and posterior positions. In both extreme positions, coxae are presumably connected to each other by coupled microtrichia fields. Thus, in extreme positions, both coxae can be moved synchronously.     

仿爱夜蛾成虫在莫高窟模拟壁画表面的运动行为及其损害机理

【目的】探明莫高窟优势病害昆虫仿爱夜蛾Apopestes spectrum (Esper)成虫在模拟壁画表面的运动行为,以揭示壁画损坏过程及其机理。【方法】本研究依据古代敦煌壁画制作工艺制作模拟壁画。利用扫描电镜观察了仿爱夜蛾成虫爬行足跗节的微观形态,定量分析了成虫翅面鳞粉量及其对壁画的污染程度,观测动物运动行为并系统记录了病害昆虫在90°模拟壁画表面的运动行为特征,分析了仿爱夜蛾成虫运动参数与模拟壁画表面粗糙度间的关系。【结果】仿爱夜蛾成虫主要依靠胸足的侧爪、中垫、掣爪片和微刺的协调作用在壁画表面黏附和爬行,在90°模拟壁画表面依然具有很强的爬行能力。仿爱夜蛾成虫通过调整步态实现在不同粗糙度表面的稳定爬行,壁画粗糙度决定了其是否需要扇动翅膀,以增大在垂直壁画表面稳定爬行的前进动力。【结论】仿爱夜蛾成虫爬行时的爪部接触冲击和拍翅行为均易造成壁画损坏,对于带有起甲、酥碱等病害的脆弱质壁画危害的风险更大。     

Mechanical,sensory and glandular structures in the tarsal unguitractor apparatus of Chironomus riparius (Diptera,Chironomidae)

Summary The legs of chironomid midges from a laboratory colony were examined in the region of the joint between the fifth tarsal segment and the pretarsus, especially the surface of the unguitractor and the manner in which the unguitractor fits into a ventromedian groove in the edge of the tarsus when the joint is flexed. The region was reconstructed from serial sections to clarify the spatial relations of the internal structures to one another and to the external structures. Ultrastructural characteristics of the cells and cuticle suggest a secretory function of the unguitractor. An amphinematic scoloparium is suspended between the point at which the unguitractor attaches to its tendon and the transverse diaphragm within the tarsus. This mechanoreceptor could serve as a proprioceptive sensor of the position of unguitractor and tarsus; it could also be an exteroceptor, sensing vibration in the substrate and/or the air. In the context of functional morphology, the clamping of the unguitractor in the ventral hollow in the tarsus could have the effect of resetting the sensitivity of the sensor. On the other hand, this arrangement could also simply act as an energy-saving means of fixing the claws in the grasping position for long periods.Abbreviations aj adhering junction - bl basal labyrinth - bm basement membrane - cc cuticular cap - cd cellular diaphragm - cth cuticular thickening - dp matrix with dense particles - ec enveloping cells - f filaments - fot funnel-like opening of tendon - ic intracellular canaliculi - lp lateral process of ec - pAc process of A cell - pv pulvillus - r rootlets of cilia - sc scolopale cell - sco scoloparium - sd septate desmosomes - t tendon - tb tubular body - tu tubules - u unguis, or claw - upl unguitractor plate     

Frictional surfaces of the elytra-to-body arresting mechanism in tenebrionid beetles (Coleoptera : Tenebrionidae) : design of co-opted fields of microtrichia and cuticle ultrastructure

In beetles, the system responsible for an attachment of forewings (elytra) to the thorax consists of interlocking fields of microtrichia (MT) located between thorax and body and between left and right elytra. The present study provides comparative data about microtrichia design on the thorax and elytra in three species of tenebrionid beetles (Tribolium castaneum, Tenebrio molitor, Zophobas rugipes) (Coleoptera : Tenebrionidae), which are different in their size. The length, width, density and directionality of microtrichia in 13 MT fields (4 on the thorax, 1 on the abdomen, 7 on the elytra, and 1 on the costal vein of the hindwing) were quantified. (1) Parameters studied are dependent on the dimension of an insect. The length of the microtrichia of most fields compared increases with an increase in body size. The MT width in the majority of fields increases with an increase in the elytra length. The MT density decreases with an increase in the elytra length. (2) Both width and length of microtrichia increase with an increase in the distance between single MT. The density of outgrowths increases with an increase in their length and width. (3) The fields oriented along the same spatial axis constitute functional groups responsible for a particular direction. Co-opted fields can be oriented in the same or opposite directions. (4) The design of MT correlates in co-opted surfaces. There are 3 field groups, which were stated as functionally corresponding to one another : the medial, anterio-lateral, and posterio-lateral. The lengths and widths of microtrichia from fields of these functional groups were quite similar in corresponding fields. Length-to-width ratios of MT in elytral fields were usually weakly correlated with those of thoracic fields. The distances between microtrichia on the elytra surface directly depended on those of the thorax. Distance-to-width ratio of MT of one surface slightly increased with an increase in this parameter on the co-opted surface. The MT densities on co-opted fields were usually quite different. (5) The ultrastructure of the cuticle suggests differences in the material properties of the cuticle between MT fields. The thoracic fields usually consist of elastic cuticle, whereas elytral fields are much harder. Usually, a MT field of elastic cuticle corresponds to the field composed of hard cuticles. The study also provides information about the ultrastructure of epidermal cells and about the design of pore channels, which are presumably responsible for production and transport of an adhesive secretion into the area of contact between lateral fields. Sensory organs monitoring contact between co-opted binding sites were also studied. The results of this study may aid in understanding the morphological basis of cuticular microsculptures acting as frictional devices.     

Structure and mechanics of the tarsal chain in the hornet, Vespa crabro (Hymenoptera: Vespidae): implications on the attachment mechanism

Two combined mechanisms on the hornet tarsus are adapted to attachment to the substrate: a friction-based (claws and spines) and an adhesion-based one (arolium). There are two ranges of substrate roughness optimal for attachment, either very smooth or very rough. There is an intermediate range of substrate grains of small but non-zero size, where both of these mechanisms fail. The optimal size of substrate grains for hornet grasping was 50-100 microm. Maximal hold to the substrate was achieved when surface irregularities were clamped between the claws of opposite legs. In such a position, the insect could withstand an external force which was almost 25 times larger than its own weight. The tarsal chain is an important part of the entire attachment mechanism. The articulations in the kinematic chain of tibia-tarsus-pretarsus are monocondylar. Three tarsal muscles and one head of the claw retractor muscle originate in the tibia. On pull to the retractor tendon, the tarsus bends in a plane. All elements of the tarsal kinematic chain have one active degree of freedom. The distance between the intertarsomeric articulation point and the tendon of the claw retractor (75-194 microm) corresponds to an efficiency of 1 degrees per 1-3 mircom of pulling distance travelled by the tendon. The claw turns about 1 degrees per 4.3-5.0 microm of pulling distance travelled by the unguitractor. The arolium turns forward and downward simultaneously with flexion of the claws. The kinematic chain of the arolium lacks real condylar joints except the joint at the base of the manubrium. Other components are tied by flexible transmissions of the membranous cuticle. The walking hornet rests on distal tarsomeres of extended tarsi. If the retractor tendon inside the tarsus is fixed, passive extension of the tarsomeres might be replaced by claw flexion. Tarsal chain rigidity, measured with the force tester, increased when the retractor tendon was tightened. Probably, pull to the tendon compresses the tarsomeres, increasing friction within contacting areas of rippled surfaces surrounding condyles within articulations.     

Armored cuticular membranes in Brachycera (Insecta,Diptera)

This study describes and quantifies the ultrastructural design of some cuticular protuberances in the Diptera-Brachycera that appear to have a specialized mechanical function. The microsculpture of membranes in the transitional areas of prothorax-neck, neck-head, head-mouthparts, and in the articulations thorax-coxa and coxa-trochanter was investigated in eight species from six families (Tabanidae, Stratiomyidae, Calliphoridae, Syrphidae, Muscidae and Drosophilidae) by means of scanning electron microscopy. The membrane armatures are single or grouped microtrichia, often located on microplates in the flexible membrane. The different membranes were compared using the following details: microtrichia length, microplate length and width, the shortest distance between microplates, and the number of microtrichia per microplate. Five different types of microstructure were observed: (1) single short papilla-like microtrichia directly arising from the flexible membrane, (2) single elongated microtrichia directly arising from the flexible membrane, (3) single microtrichia located on small areas of the inflexible cuticle, here called microplates, (4) microtrichia groups located on the microplates, and (5) microplates without microtrichia in the flexible membrane. The study reveals the differences in the surface ultrastructure of membranes between different parts of body and between different taxa of flies. The prothorax-neck transition is less specialized in membrane armature as compared with the head-neck articulation, and especially with that of the head-proboscis. The head-proboscis transition has the most complex membrane armature of those species studied. The membrane of the basal joints of the legs has larger plates and a higher number of microtrichia per plate than that of the more distal joints. The complexity of the membrane armature depends on the dimensions of the animal: the smallest species have short ungrouped microtrichia or micropapillae, whereas large species have the highest degree of grouping, the longest microtrichia, and the largest microplates. Three main mechanical functions served by the different armored membranes are suggested, based on the specific characteristics of the microsculptural design: (1) fixation of the folds, (2) prevention of folding, and (3) determination of direction of folding. J. Morphol. 234:213–222, 1997. © 1997 Wiley-Liss, Inc.     

Comparative morphology of pretarsal scopulae in eleven spider families

Many wandering spiders bear attachment pads (scopulae) on their tarsi, consisting of hierarchically-branching adhesive setae. Amongst spider families and even species, these show remarkable differences in morphology. Using scanning electron microscopy, the scopula microstructure of sixteen spider species was described, with the focus on pretarsal scopulae (claw tufts). Area and shape of the claw tuft, seta and setule density, as well as seta and spatula dimensions were analysed and compared. Claw tufts of the majority of species studied show a similar gradient in size and shape from anterior to posterior legs: the dimension of pads increases, while setal density decreases. Commonly, there is also a gradient of both the seta and spatula size within the claw tuft: Setae become larger from the proximal to the distal part of the pad, and spatulae size increases in the same direction at the level of individual seta. Often, different hierarchical levels of claw tuft organisation are differently expressed in different species: Species with lower setal density usually have broader setae. Smaller spatula size often implicates higher setule density. Evolutionary and ecological aspects of the scopula origin are discussed.     

Mineral in skeletal elements of the terrestrial crustacean Porcellio scaber: SRμCT of function related distribution and changes during the moult cycle

Terrestrial isopods moult first the posterior and then the anterior half of the body, allowing for storage and recycling of CaCO₃. We used synchrotron-radiation microtomography to estimate mineral content within skeletal segments in sequential moulting stages of Porcellio scaber. The results suggest that all examined cuticular segments contribute to storage and recycling, however, to varying extents. The mineral within the hepatopancreas after moult suggests an uptake of mineral from the ingested exuviae. The total maximum loss of mineral was 46% for the anterior and 43% for the posterior cuticle. The time course of resorption of mineral and mineralisation of the new cuticle suggests storage and recycling of mineral in the posterior and anterior cuticle. The mineral in the anterior pereiopods decreases by 25% only. P. scaber has long legs and can run fast; therefore, a less mineralised and thus lightweight cuticle in pereiopods likely serves to lower energy consumption during escape behaviour. Differential demineralisation occurs in the head cuticle, in which the cornea of the complex eyes remains completely mineralised. The partes incisivae of the mandibles are mineralised before the old cuticle is demineralised and shed. Probably, this enables the animal to ingest the old exuviae after each half moult.     

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.     

Functional morphology of the glandula prostatica,ejaculatory valve,and ductus ejaculatorius of the silkworm Bombyx mori

The penis of the silkmoth, Bombyx mori, consists of two parts covered with cuticle, the corpus penis and crus penis, and a third part, the radix penis, without a cuticle but surrounded by a thick sphincter. The radix penis is divisible into anterior and posterior parts. The ductus (d.) ejaculatorius passing through the penis has no secretory cells. In the anterior radix penis, the wall of the d. ejaculatorius is thin and without folds; in the posterior section, it is thick, with folds in its lumen. The glandula (g.) prostatica is divisible into anterior and posterior parts according to differences in the histological and morphological characteristics of the cells and their secretions, which contain many heterogeneous substances. In the anterior g. prostatica, secretions accumulate separately in the anterior and posterior sections before ejaculation. Unlike the posterior region, the anterior region displays a large mass(es) at the periphery of the lumen along the secretory cell layer. Judging from staining properties, the pearly body and the first layer of the spermatophore wall, which, after copulation, form in the female bursa copulatrix, seem to be derived from the secretions of the anterior and posterior regions of the g. prostatica, respectively. The secretion of the posterior g. prostatica contains initiatorin, which acts as a sperm-activating factor in the inner and outer matrices of the spermatophore. An ejaculatory valve is found between the radix penis and the g. prostatica. The opening of this valve is regulated by the surrounding sphincter, thus impeding the back-flow of secretions and seminal fluid in the radix penis and resulting in their transport outwards during ejaculation. The musculature of the d. ejaculatorius and the corpus penis promotes further transport of these secretions into the female bursa copulatrix.     

Ultrastructure of the contact surfaces of Passalurus ambiguus (Rudolphi, 1819) (Nematoda)

The ultrastructure of the contact surfaces (integument and intestinal wall) of the nematode Passalurus ambiguus has been studied. The integument is composed according to the scheme common for all nematodes and includes a cuticle, hypodermis and a muscular layer. The specificity is with regard to the epicuticle, the different number of the cuticular sublayers in the anterior, central and the posterior parts of the worm body and the absence of a basal cuticular membrane. The intestinal wall consists of epithelial cells with microvilli. The ultrastructural characteristics of both contact surfaces indicate their main functions--absorption, secretion, transport, protection, movement, etc.     

The adhesive devices in larvae of Lepidoptera (Insecta, Pterygota)

Adhesion to smooth surfaces by means of thin fluid lipid film was studied on living larvae of 71 species of Lepidoptera by a simple ”light reflection method”. The method made it possible to localize exactly the sites of adhesion and to estimate roughly the film thickness, within a certain range. Furthermore, it revealed the general presence of mobile lipid on the entire insect surface. The observations on living larvae were complemented by comparative structural studies of the adhesive parts with light and scanning electron microscopes on preserved specimens of 161 species. Specialized adhesive devices were found in great diversity on larval legs and prolegs, especially in larvae living in the open air on their food plants. Two main surface types of adhesive cuticle were found: (1) cuticle with a flexible smooth surface and (2) cuticle with very numerous small projections (microtrichia) with spatulate and recurved apices. Both the functional implications of the adhesive cuticular structure and the role of the adhesive fluid as well as the evolution of the adhesive devices are discussed. The adhesive effect is due to ”capillary” or meniscus forces. Accepted: 29 July 1999     

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.     

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.     

Structure and function of the arolium of Mantophasmatodea (Insecta)

All species of the insect order Mantophasmatodea characteristically keep the 5th tarsomere and pretarsus (arolium plus two claws) turned upwards and off the substrate. The unusually large arolium was studied in two species of Mantophasmatodea using bright field light microscopy, reflection microscopy, fluorescence microscopy, TEM, SEM, and Cryo‐SEM. It contains an epithelial gland, numerous tracheoles, and nerves. The gland consists of enlarged epithelial cells with large nuclei, mitochondria, RER, golgi complexes, microtubules, and numerous secretion vesicles. Evidence for exocytosis of the vesicles into the gland reservoir between the epithelial gland and the thick cuticle could be observed. Cryo‐SEM revealed that the ventral side of the arolium and distal part of its dorsal side are covered with a liquid film. Fluid footprints of arolia of individuals walking on a glass plate also indicate the presence of secretory fluid on the arolium surface. Behavioral experiments using animals with ablated arolia showed that representatives of Mantophasmatodea do not need their arolia to detect and respond to vibratory communication signals nor to catch small to medium‐sized prey. Individuals with ablated arolia were not able to move upside down on a smooth glass plate. We conclude that Mantophasmatodea use their arolia for attachment when additional adhesion force is required (e.g. windy conditions, handling large prey, mating). They can bring their arolia in contact with the surface in a very fast reflex (18.0 ± 9.9 ms). The secretory fluid found on the surface is produced by the gland and transported to the outside, presumably through small pore channels, to enhance adhesion to the substrate. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Ultrastructure of the thoracic dorso‐medial field (TDM) in the elytra‐to‐body arresting mechanism in Tenebrionid Beetles (Coleoptera: Tenebrionidae)

Beetles with flying ability lock their elytra (forewings) to the thorax or/and abdomen using complex locking devices. These structures are often supplemented with microtrichia fields of the inner surface of the elytra and adjacent parts of the pterothorax. The present study provides information about the ultrastructure of microtrichia of the dorso‐medial fields of the thorax (TDM) in tenebrionid beetles (Tribolium castaneum, Zophobas rugipes). Epidermal cells located under the TDM field contain large electron‐lucent vesicles connected to rough endoplasmic reticulum. Microtrichia and underlying cuticle of the TDM have a high density of pore channels, which are responsible for transport of an epidermal secretion onto the TDM surface. In order to show the presence of the secretion on the cuticle surface, TDM fields of air‐dried specimens were compared with those in specimens after two treatments, such as (1) dehydration in ethanol and acetone, and (2) dental‐wax‐cast technique applied to living beetles. This revealed the presence of the nonvolatile film on the intact microtrichial surface. Possible functions of this film are suggested to be (1) the increase of adhesive forces in the contact area and (2) providing soft coupling and release of two corresponding parts of the elytra‐locking device. J. Morphol. 240:101–113, 1999. © 1999 Wiley‐Liss, Inc.     

The problem of the number of tarsomeres in the regenerated cockroach leg.

There are 5 tarsomeres in the normal cockroach leg, but this number is often reduced in regenerated legs. In order to examine this complicated situation, fore-, mid-, and hindlegs of German cockroaches were amputated at 11 different tarsal levels and at 18 different times during the last instar. When tarsi were amputated at or proximal to the 3rd tarsomere, 4-segmented tarsi regenerated. When legs were amputated distal to the 3rd tarsomere, the regenerated tarsi had 5 segments. Three-segmented tarsi rarely regenerated when legs were amputated proximal to 3rd tarsomere and in the latter half of the instar period. The lengths of all tarsomeres of regenerated tarsi were measured together with those of unoperated contralateral tarsomeres, and the ratios of the former to the latter were calculated. The ratios ranged from 28 to 138% for the various tarsomeres and levels of amputation. From a comparison of the ratios and morphological observations, it was suggested that the 3rd tarsomere of the normal 5-segmented tarsus has disappeared in the regenerated 4-segmented tarsus. Pads and disto-lateral spines of tarsomeres were observed on unoperated and regenerated tarsi. It was of interest that double spines were often found on the 4-segmented tarsi, mostly on the 2nd tarsomere, just proximal to the position of the missing 3rd tarsomere. This observation supported the idea that the 3rd tarsomere has not simply disappeared, but has probably fused with the 2nd tarsomere.