The mechanosensory apparatus of the femoral tactile spine of the cockroach,Periplaneta americana

Summary Tactile spines are large cuticular sense organs that appear to provide insects with a sense of touch which is spatially coarse but of great sensitivity. Cockroach legs have a number of these spines on each leg and a particularly prominent spine on the end of each femur, the femoral tactile spine. The ease of recording afferent activity from this spine during mechanical stimulation has made it one of the most thoroughly studied insect mechanoreceptors and yet it has never been examined by electron microscopy.We report here the results of an examination of the femoral tactile spine by both scanning and transmission electron microscopy, as well as by light microscopy. The spine is shown to be innervated by a single sensory bipolar neuron with its soma located in the base of the spine. A canal through the wall of the spine leads to the outside and emerges just above the junction between the base of the spine and its articulating socket membrane. The sensory dendrite of the neuron passes from the soma through this canal and forms a modified ciliary sensory ending with the typical dendritic sheath and dense tubular body that is characteristic of insect mechanosensory cuticular sensilla. The tubular body is embedded in a cuticular terminal plug which closes the exterior end of the canal but appears to be fastened to the spine by a very flexible ring of cuticle. This plug is connected to the socket membrane by a specialized socket attachment which presumably serves to move the plug relative to the wall of the spine during movement of the spine within the socket. The morphology of this sensillum is discussed in terms of the possible ways in which it is stimulated by movements of the spine and also in light of the dynamic behaviour of the receptor which is now very well described.Supported by the Canadian Medical Research Council. The authors gratefully acknowledge the expert technical assistance of Sita Prasad and Rodney Gramlich     

Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion

The best supported hypothesis for the evolutionary origin of insect wings is that they evolved from articulated, leg-derived respiratory structures of aquatic ancestors. However, there are no fossils of the immediate ancestors of winged insects, and it is difficult to imagine how a functional transition from gills to wings could have occurred. Recent studies of surface-skimming locomotion in stoneflies and mayflies offer a plausible solution by showing how rudimentary wings and muscle power can be used to accomplish two-dimensional aerodynamic locomotion on the surface of water. Here we extend that line of research by examining the phylogenetic distribution and mechanistic diversity of surface skimming in stoneflies, along with a limited examination of mayflies. These investigations reveal both a broad taxonomic occurrence and a fine gradation of mechanically distinct forms. Distinct forms of wing-flapping surface skimming include (1) stoneflies that flap their wings weakly while maintaining their body in contact with the water and undulating their abdomen laterally in a swimming-like motion, (2) stoneflies that skim while elevating their body above the water and maintaining all six legs on the surface, (3) stoneflies and mayflies that skim with only four legs on the water surface, (4) stoneflies that skim with only their two hind legs on the surface, and (5) stoneflies that, beginning with a series of leg motions nearly identical to hind-leg skimmers, use their hind legs to jump from the water into the air to initiate flapping flight. Comparisons across these forms of skimming show that wing-beat amplitude, horizontal velocity, and the verticality of aerodynamic force production increase as the body orientation becomes more upright and contact with the water is minimized. These behaviors illustrate a mechanical pathway by which flying insects could have evolved from swimming ancestors via a series of finely graded intermediate stages. The phylogenetic distribution of skimming and flight in stoneflies does not indicate any clear directionality toward either greater or lesser aerodynamic abilities; however, the broad and apparently basal phylogenetic distribution of skimming taxa supports the hypothesis that the common ancestor of stoneflies was a surface skimmer. This may also be true for the common ancestor of stoneflies and mayflies, that is, the first winged insects. We combine these data with fossil evidence to form a synthetic model for the evolution of flying insects from surface skimmers.     

蜚蠊棘-钟形感器冲动发放的适应

本实验研究蜚蠊后胸足胫节棘-钾形感器冲动发放的适应特性,用探针向棘窝方向推棘以兴奋感觉神经元,用胞外电极记录神经冲动的发放,显示明显的适应现象.用计算机对冲动信号进行分析作图可得时间-瞬时频率曲线,该曲线呈负幂函数分布并可用y=B(?)公式描述,其中k为回归系数,表示该感器的适应速率,棘移动距离可影响k值,棘移动距离越长,k值越大,表示适应速率减慢.高频率移动棘可使反应发生动态和稳态两方面的变化;前者显示后续反应的脉冲个数的减少,后者则显示k值的降低,意味着适应的加快.     

The contralateral coordination of walking legs in the crayfish Astacus leptodactylus

The coupling mechanisms which coordinate the movement of ipsilateral walking legs in the crayfish have been described in earlier investigations. Concerning the coupling between contralateral legs it was only known that these influences are weaker than those acting between ipsilateral legs. The nature of these coupling mechanisms between contralateral legs of the crayfish are investigated here by running left and right legs on separate walking belts at different speeds. The results show that coordination is performed by a phase-dependent shift of the anterior extreme position of the influenced leg. This backward shift leads to a shortening of both the return stroke and the following power stroke. As the coupling influence is only weak, several steps might be necessary to retain normal coordination after a disturbance. This corresponds to v. Holst's relative coordination. The influences act in both directions, from left to right and vice versa. However, one side may be more or less dominant. A gradient was found in the way that anterior leg pairs show less strong coordination than posterior legs. In some cases the coupling between diagonally neighbouring legs was found to be stronger than between contralateral legs of the same segment. The interpretation of this result is still open.     

The control of walking movements in the leg of the rock lobster

1. Experiments with rock lobsters walking on a treadmill were undertaken to obtain information upon the system controlling the movement of the legs. Results show that the position of the leg is an important parameter affecting the cyclic movement of the walking leg. Stepping can be interrupted when the geometrical conditions for terminating either a return stroke or a power stroke are not fullfilled. 2. The mean value of anterior and posterior extreme positions (AEP and PEP respectively) of the walking legs do not depend on the walking speed (Fig. 1). 3. When one leg is isolated from the other walking legs by placing it on a platform the AEPs and PEPs of the other legs show a broader distribution compared to controls (Figs. 2 and 3). 4. Force measurements (Fig. 4) are in agreement with the hypothesis that the movement of the leg is controlled by a position servomechanism. 5. When one leg stands on a stationary force transducer this leg develops forces which oscillate with the step rhythm of the other legs (Fig. 5). 6. A posteriorly directed influence is found, by which the return stroke of a leg can be started when the anterior leg performs a backward directed movement. 7. Results are compared with those obtained from stick insects. The systems controlling the movement of the individual leg are similar in both, lobster and stick insect but the influences between the legs seem to be considerably different.     

Mechanical properties of the isolated catch apparatus of the sea urchin spine joint: muscle fibers do not contribute to passive stiffness changes

The catch apparatus (CA) is the collagenous ligament at the spinal joint of sea urchins. It maintains spine posture by stiffening and allows spine movement by softening. A CA preparation, which was isolated from ossicles, was used to test the hypothesis that frictional forces between collagen fibers and ossicles are the source of stiffness changes. Isolated preparations of the CA changed in stiffness, thus falsifying the hypothesis. Another hypothesis proposes that muscle fibers, which represent a relatively small component of the CA, cause stiffening of the CA by contraction. Chemicals that evoked contraction in spine muscles did not always stiffen the CA: the CA of Heterocentrotus mammillatus softened in response to artificial seawater with potassium concentration elevated to 100 mM. This provided evidence against the muscle-based hypothesis. The present results suggest that the stiffness changes of the CA are based on changes in the mechanical properties of the extracellular components of the connective tissue and are therefore related to the connective tissue catch that is widespread in other echinoderms.     

Duplication of a peripheral sensory neuron in the cockroach Periplaneta americana

Summary We have recently examined the electrophysiology and ultrastructure of approximately 100 tactile spines from the metathoracic legs of adult cockroaches. In only one animal the single sensory neuron that innervates the spine was replaced with a pair of apparently identical neurons which we believe were both functional. As far as we are aware this is the first reported study of unprovoked duplication in a peripherally-located insect sensory neuron.Supported by the Canadian Medical Research Council and the Alberta Heritage Foundation for Medical Research     

Control of body position of a stick insect standing on uneven surfaces

When a multi-legged animal walks over uneven surfaces, each leg has to span a different distance between body and ground. Thus the animal has to solve the problem of how to control the body height, i.e. to coordinate the movement of the legs in such a way that the vertical projections of leg lengths match these distances. For the standing animal, this is investigated here by testing twelve different substrate configurations and measuring body height and forces applied by the legs on the substrate. The results are consistent with the hypothesis that the legs can be considered to represent independent height controllers. They can be understood as proportional controllers with nonlinear characteristics. The mechanical coupling between the leg is sufficient to explain the experimental results. Thus, no neuronal coupling has to be assumed to exist between these controllers. This agrees with a hypothesis proposed earlier for walking animals (Cruse 1976).     

Environmental rugosity, body size and access to food: a test of the size-grain hypothesis in tropical litter ants

In terrestrial walking organisms, long legs help to decrease the cost of running, allowing animals to step over environmental interstices rather than walking through them. However, long legs can complicate the infiltration of these interstices, which may contain food sources and refugia. Since the number of environmental interstices perceived by an organism (rugosity) increases as it body size decreases (size-grain hypothesis, SGH), natural selection should favor proportionally smaller legs with decreasing body size. Recent work demonstrated that ants fit this hypothesis. We experimentally tested the assumption of the SGH that small ants, which have proportionally smaller legs than larger ants, are more successful in exploring environmental interstices because they can easily penetrate them. We examined the ability of tropical litter ant species with different body sizes to access food baits in 'landscapes' (=plots) with different levels of rugosity and food exposure. In the first experiment, three levels of landscape rugosity were defined by manipulating the density of leaf litter placed on the ground plots: a) plain landscape: no litter fall, b) intermediate rugosity (∼0.5 kg of litter fall) and c) high rugosity (∼1 kg). In a second experiment, food baits were in plain landscapes, exposed or covered by leaf litter. The body lengths of ants that first accessed food baits ranged from 1.5 to 12 mm. Ants that first reached food baits in the most rugose landscapes were ∼40% smaller than ants that first found baits in plain landscapes. Smaller ants were also the first to access covered food. The application of a phylogenetic comparative method suggested the same patterns. We conclude that these results support the size grain hypothesis. Environmental rugosity might have operated as a selective force to shape the morphological characteristics of litter ant species.     

Effects of natural and simulated herbivory on spine lengths of Acacia drepanolobium in Kenya

We present experimental evidence supporting the hypothesis that increased spine length in acacia species is a defense induced by herbivory. Acacia drepanolobium is the dominant tree over large areas of East Africa. Each individual tree is occupied by one of four ant species at our study site. Using two types of electric fences, we have effectively controlled herbivory by megaherbivores (elephants and giraffes) and other large mammalian herbivores at a field site in Laikipia, Kenya since 1995. Mean spine lengths of new spines on trees occupied by the most abundant ant species (presumed to be a defensive mutualist) have shown a slow and steady decline over the first five years of the experiment on branches protected from these herbivores. This reduction has been 35–40%, or approximately half of the reduction in spine length that we anticipate will eventually occur, based on trees that have been protected from herbivory for many years. In contrast, trees occupied by a resident ant species that systematically prunes shoots have shown no reduction in spine length associated with herbivore exclusion treatments. Experimental pruning of shoots similar to that carried out by this ant species resulted in longer spines on seedlings in a greenhouse setting. Simulated large mammal browsing in the field rapidly (re-)induced greater spine lengths on trees that had been protected from large mammals for five years. The slow relaxation of spine length in the absence of herbivory, contrasted with its rapid induction after simulated browsing, suggests that there is a difference in the reliability of these two signals. Spine length responses to herbivory were extremely local (limited to individual branches). These branch-specific responses are consistent with the hypothesis that induced defense in this system evolved in the context of within-tree spatial variation in herbivore pressure, in particular variation in branch height.     

Morphological evolution of spiders predicted by pendulum mechanics

Background

Animals have been hypothesized to benefit from pendulum mechanics during suspensory locomotion, in which the potential energy of gravity is converted into kinetic energy according to the energy-conservation principle. However, no convincing evidence has been found so far. Demonstrating that morphological evolution follows pendulum mechanics is important from a biomechanical point of view because during suspensory locomotion some morphological traits could be decoupled from gravity, thus allowing independent adaptive morphological evolution of these two traits when compared to animals that move standing on their legs; i.e., as inverted pendulums. If the evolution of body shape matches simple pendulum mechanics, animals that move suspending their bodies should evolve relatively longer legs which must confer high moving capabilities.

Methodology/Principal Findings

We tested this hypothesis in spiders, a group of diverse terrestrial generalist predators in which suspensory locomotion has been lost and gained a few times independently during their evolutionary history. In spiders that hang upside-down from their webs, their legs have evolved disproportionately longer relative to their body sizes when compared to spiders that move standing on their legs. In addition, we show how disproportionately longer legs allow spiders to run faster during suspensory locomotion and how these same spiders run at a slower speed on the ground (i.e., as inverted pendulums). Finally, when suspensory spiders are induced to run on the ground, there is a clear trend in which larger suspensory spiders tend to run much more slowly than similar-size spiders that normally move as inverted pendulums (i.e., wandering spiders).

Conclusions/Significance

Several lines of evidence support the hypothesis that spiders have evolved according to the predictions of pendulum mechanics. These findings have potentially important ecological and evolutionary implications since they could partially explain the occurrence of foraging plasticity and dispersal constraints as well as the evolution of sexual size dimorphism and sociality.     

Capillary filtration rate, venous compliance, and blood flow in arms and legs do not change during bed rest for 20 days

It has been generally accepted that pooling of the blood in the legs is one reason for the orthostatic intolerance experienced after space flights. This is also the reasoning behind the application of anti-G suits during reentry after space flights. Fighter pilots also use the anti-G suit, the hypothesis being that this prevents the pooling of blood in the legs. In order to investigate if immobilization during bed rest would induce peripheral cardiovascular deconditioning we measured capillary filtration rate, venous compliance, and blood flow in arms and legs during bed rest.     

A new species of Metacyclops Kiefer, 1927 (Copepoda,Cyclopidae, Cyclopinae) from the Chihuahuan desert,northern Mexico

A new species of the freshwater cyclopoid copepod genus Metacyclops Kiefer, 1927 is described from a single pond in northern Mexico, within the binational area known as the Chihuahuan Desert. This species belongs to a group of Metacyclops species with a 3443 spine formula of swimming legs. It is morphologically similar to Metacyclops lusitanus Lindberg, 1961 but differs from this and other congeners by having a unique combination of characters, including a caudal rami length/width proportion of 3.5–3.8, a innermost terminal seta slightly longer than the outermost terminal seta, intercoxal sclerites of legs 1-4 naked, a strong apical spine of the second endopodal segment of leg 1 and one row of 6-8 small spinules at the insertion of this spine. The finding of this species represents also the first record of the genus in Mexico and the third in North America, where only two other species, Metacyclops gracilis (Lilljeborg, 1853)and Metacyclops cushae Reid, 1991 have been hitherto reported. This is also the first continental record of a species of Metacyclops from an arid environment in the Americas. This species appears to be endemic to the Chihuahuan Desert, thus emphasizing the high endemicity of this area.     

Effect of strength training and the practice of Alpine skiing on bone mass density, growth, body composition, and the strength and power of the legs of adolescent skiers

This work examines the influence of practicing strength training and Alpine skiing over 2 years on bone mineral density (BMD), growth, body composition, and the strength and power of the legs of adolescent skiers. The study subjects were 20 adolescent skiers (10 girls and 10 boys) and 19 sedentary adolescents (9 girls and 10 boys), all 13-16 years of age. The BMDs of the lumbar column (L2-L4) and hip (neck of the femur, trochanter, and Ward's triangle) were determined by dual x-ray photon absorptiometry at the beginning and end of the experimental period. The increase in height and the percentage fat and muscular masses of the subjects were also recorded, as was their ability to jump (countermovement jump [CMJ]), their leg strength and power (squat test), and their leg anaerobic power (continuous jump test [CMJ15″]). No significant differences were seen in the increase in height, body weight, or percentage fat mass between the skiers and sedentary subjects, although the boy skiers showed a significant increase in percentage muscular mass (p < 0.05) compared to the sedentary boys. The improvement in the values of the different CMJ variables was significantly greater among the boy skiers than among the sedentary boys (p < 0.001-0.01). The same was true for the girls (p < 0.001), except for CMJ15″. The skiers experienced a significantly greater increase in L2-L4 BMD than the sedentary subjects (boys p < 0.05; girls p < 0.01). These results suggest that Alpine skiing combined with rational strength training involves no special risk for the physical development of young people, has a positive effect on the power and the percentage of muscle mass in the legs, and helps to have a higher bone density in the lumbar spine (L2-L4).     

Function of the spine

In spite of the considerable effort which has been invested in attempts to understand the mechanism of human spines, substantial controversy remains, particularly in connection with assumptions which have to be made by those engaged in biological modelling.The hypothesis presented here is that the living joint has stress sensors driving a feedback mechanism, an arrangement which could react to imposed loads by modifying muscular action in such a way as to minimize stress at the joints and therefore the risk of injury. A theory of this kind gives an image of the spine not in terms of a spatial picture, as would a CAT scan, but in terms of stresses, forces and moments acting at the intervertebral joints. Calculations show that the erectores spinae alone cannot support more than about 50 kg; there must be some other mechanism to explain man's ability substantially to exceed that load.It is suggested that the interaction between the erectores spinae and the abdominals are of fundamental importance in the function of the spine; how they are co-ordinated during the lifting of weights is examined in detail. The theory resulting from this hypothesis is used to relate spinal injury and an injured subject's posture and behaviour.A mathematical formulation permits an objective evaluation of the spine, and a procedure for determining an automatic diagnosis of lumbar spine disabilities is proposed.     

A cockroach that jumps

We report on a newly discovered cockroach (Saltoblattella montistabularis) from South Africa, which jumps and therefore differs from all other extant cockroaches that have a scuttling locomotion. In its natural shrubland habitat, jumping and hopping accounted for 71 per cent of locomotory activity. Jumps are powered by rapid and synchronous extension of the hind legs that are twice the length of the other legs and make up 10 per cent of the body weight. In high-speed images of the best jumps the body was accelerated in 10 ms to a take-off velocity of 2.1 m s(-1) so that the cockroach experienced the equivalent of 23 times gravity while leaping a forward distance of 48 times its body length. Such jumps required 38 μJ of energy, a power output of 3.4 mW and exerted a ground reaction force through both hind legs of 4 mN. The large hind legs have grooved femora into which the tibiae engage fully in advance of a jump, and have resilin, an elastic protein, at the femoro-tibial joint. The extensor tibiae muscles contracted for 224 ms before the hind legs moved, indicating that energy must be stored and then released suddenly in a catapult action to propel a jump. Overall, the jumping mechanisms and anatomical features show remarkable convergence with those of grasshoppers with whom they share their habitat and which they rival in jumping performance.     

Sexual selection for male mobility in a giant insect with female-biased size dimorphism

Female-biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female-biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males.     

新鳋属两个种的分化在超微结构上的表达

本文对寄生桡足类鳋科的二种新鳋首次作了扫描电镜观察,发现游泳足上的细微结构以及胸板的形态在种间很不相同,这些形态上的变化,对两个种来说是分化,对整个生物界来说,就是生物多样性的表达。     

Comparative strengths and structural properties of the upper and lower cervical spine in flexion and extension

The purpose of this study is to test the hypothesis that the upper cervical spine is weaker than the lower cervical spine in pure flexion and extension bending, which may explain the propensity for upper cervical spine injuries in airbag deployments. An additional objective is to evaluate the relative strength and flexibility of the upper and lower cervical spine in an effort to better understand injury mechanisms, and to provide quantitative data on bending responses and failure modes. Pure moment flexibility and failure testing was conducted on 52 female spinal segments in a pure-moment test frame. The average moment at failure for the O-C2 segments was 23.7+/-3.4Nm for flexion and 43.3+/-9.3Nm for extension. The ligamentous upper cervical spine was significantly stronger in extension than in flexion (p=0.001). The upper cervical spine was significantly stronger than the lower cervical spine in extension. The relatively high strength of the upper cervical spine in tension and in extension is paradoxical given the large number of upper cervical spine injuries in out-of-position airbag deployments. This discrepancy is most likely due to load sharing by the active musculature.     

Leg coordination and swimming in an ant, Camponotus americanus

ABSTRACT. Leg movements of Camponotus americanus workers during straight swimming and turning are described herein. Thrust is generated through the different speeds and drag control between power v. return strokes in the forelegs. During the power stroke, femur, tibia and tarsus are straightened and thereby increase resistance; they bend backward during the return stroke. These thrusting legs move in a vertical plane which is similar to their position during walking. The backward stretching mesothoracic and metathoracic legs act, in conjunction with the gaster, as a rudder. Swimming in ants can be derived from walking; the major transformation being a suppression of the rhythmic movements of the middle and hind legs.