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).     

Simulation of a model for the coordination of leg movement in free walking insects

A computer has been used to simulate the behaviour of a model proposed to explain certain asymmetries in the free walking step patterns of adult stick insects. Complete sequences of behaviour including turns, changes in velocity, and transitions between different step patterns have been simulated and are compared with the real sequences produced by 1st instar and adult animals. The procedure for simulating the complete walking behaviour suggests that there are subtle differences in the walking system at these two stages of growth in addition to those outlined in an earlier paper. The model also provides a formal structure for expressing quantitative differences between the walking behaviour of the cockroach, locust, grasshopper, and stick insect.     

Deriving neural network controllers from neuro-biological data: implementation of a single-leg stick insect controller

This article presents modular recurrent neural network controllers for single legs of a biomimetic six-legged robot equipped with standard DC motors. Following arguments of Ekeberg et?al. (Arthropod Struct Dev 33:287?C300, 2004), completely decentralized and sensori-driven neuro-controllers were derived from neuro-biological data of stick-insects. Parameters of the controllers were either hand-tuned or optimized by an evolutionary algorithm. Employing identical controller structures, qualitatively similar behaviors were achieved for robot and for stick insect simulations. For a wide range of perturbing conditions, as for instance changing ground height or up- and downhill walking, swing as well as stance control were shown to be robust. Behavioral adaptations, like varying locomotion speeds, could be achieved by changes in neural parameters as well as by a mechanical coupling to the environment. To a large extent the simulated walking behavior matched biological data. For example, this was the case for body support force profiles and swing trajectories under varying ground heights. The results suggest that the single-leg controllers are suitable as modules for hexapod controllers, and they might therefore bridge morphological- and behavioral-based approaches to stick insect locomotion control.     

The optomotor system on the ground: on the absence of visual control of speed in walking ladybirds

Summary The strengths of the rotational and possible translational optomotor reflexes were measured over a wide range of closed-loop conditions in walking ladybirds (Coccinella septempunctata), and less thoroughly in three other species of insect. While ladybirds exhibit a strong rotational optomotor reflex, any visual control of speed there might be was found to be too feeble to be biologically significant. To see whether walking speed is instead controlled proprioceptively, changes in speed were measured when ladybirds pulled small weights. But there was no evidence of proprioceptive control either. Flying and swimming insects, on the other hand, do use visual feedback to control their translational velocity, and, unlike walking insects,must do so to cope with winds or watercurrents.     

A model for analyzing insect stage-frequency data when mortality varies with time

A model is developed for the analysis of insect stage-frequency data which may be applied to populations with age-dependent mortality. The analysis of stage-frequency data is divided into two steps. In the first step, the number of different mortality rates and their values are estimated. The second step provides estimates of developmental rates and variances for each developmental stage and in addition provides estimates of the number of recruits to each stage. The model may be used both in analysis and prediction of insect stage frequencies. Hence, in addition to estimating developmental and mortality rates from stage-frequency data, it may also be used as a simulation model for an insect population. The model is applied to two populations of Hemileuca oliviaeCockerell , a lepidopterous pest of New Mexico grasslands. The model identifies, in the two populations, different mortality rates that are related to plant productivity.     

Analysis of a distributed model of leg coordination

Using tools from discrete dynamical systems theory, we begin a systematic analysis of a distributed model of leg coordination with both biological and robotic applications. In this paper, we clarify the role of individual coordination mechanisms by studying a system of two leg oscillators coupled in one direction by each of the three major mechanisms that have been described for the stick insect Carausius morosus. For each mechanism, we derive analytical return maps, and analyze the behavior of these return maps under iteration in order to determine the asymptotic phase relationship between the two legs. We also derive asymptotic relative phase densities for each mechanism and compare these densities to those obtained from numerical simulations of the model. Our analysis demonstrates that, although each of these mechanisms can individually compress a range of initial conditions into a narrow band of relative phase, this asymptotic relative phase relationship is, in general, only neutrally stable. We also show that the nonlinear dependence of relative phase on walking speed along the body in the full hexapod model can be explained by our analysis. Finally, we provide detailed parameter charts of the range of behavior that each mechanism can produce as coupling strength and walking speed are varied. Received: 22 June 1999 / Accepted in revised form: 7 September 1999     

A self-organizing model of walking patterns of insects

Insects generate walking patterns which depend upon external conditions. For example, when an insect is exposed to an additional load parallel to the direction in which it is walking, the walking pattern changes according to the magnitude of the load. Furthermore, even after some of its legs have been amputated, an insect will produce walking patterns with its remaining legs. These adaptations in insect walking could not previously be explained by a mathematical model, since the mathemati cal models were based upon the hypothesis that the relationship between walking velocity and walking patterns is fixed under all conditions. We have produced a mathematical model which describes self-organizing insect walking patterns in real-time by using feedback information regarding muscle load (Kimura et al. 1993). As part of this model, we introduced a new rule to coordinate leg movement, in which the information is circulated to optimize the efficiency of the energy transduction of each effector orga n. We describe this mechanism as ‘the least dissatisfaction for the greatest number of elements’. In this paper, we introduce the following aspects of this model, which reflect adaptability to changing circumstances: (1) after one leg is exposed to a transient perturbation, the walking pattern recovers swiftly; (2) when the external load parallel to the walking direction is continuously increased or decreased, the pattern transition point is shifted according to the magnitude of the load increme nt or decrement. This model generates a walking pattern which optimizes energy consumption at a given walking velocity even under these conditions; and (3) when some of the legs are amputated, the model generates walking patterns which are consistent with experimental results. We also discuss the ability of a hierarchical self-organizing model to describe a swift and flexible information processing system. Received: 8 February 1993/Accepted in revised form: 12 November 1993     

Winching up heavy loads with a compliant arm: a new local joint controller

A closed kinematic chain, like an arm that operates a crank, has a constrained movement space. A meaningful movement of the chain’s endpoint is only possible along the free movement directions which are given implicitly by the contour of the object that confines the movement of the chain. Many technical solutions for such a movement task, in particular those used in robotics, use central controllers and force–torque sensors in the arm’s wrist or a leg’s ankle to construct a coordinate system (task frame formalism) at the local point of contact the axes of which coincide with the free and constrained movement directions. Motivated by examples from biology, we introduce a new control system that solves a constrained movement task. The control system is inspired by the control architecture that is found in stick insects like Carausius morosus. It consists of decentral joint controllers that work on elastic joints (compliant manipulator). The decentral controllers are based on local positive velocity feedback (LPVF). It has been shown earlier that LPVF enables contour following of a limb in a compliant motion task without a central controller. In this paper we extend LPVF in such a way that it is even able to follow a contour if a considerable counter force drags the limb away along the contour in a direction opposite to the desired. The controller extension is based on the measurement of the local mechanical power generated in the elastic joint and is called power-controlled relaxation LPVF. The new control approach has the following advantages. First, it still uses local joint controllers without knowledge of the kinematics. Second, it does not need a force or torque measurement at the end of the limb. In this paper we test power-controlled relaxation LPVF on a crank turning task in which a weight has to be winched up by a two-joint compliant manipulator.     

Mechanisms of stick insect locomotion in a gap-crossing paradigm

Locomotion of stick insects climbing over gaps of more than twice their step length has proved to be a useful paradigm to investigate how locomotor behaviour is adapted to external conditions. In this study, swing amplitudes and extreme positions of single steps from gap-crossing sequences have been analysed and compared to corresponding parameters of undisturbed walking. We show that adaptations of the basic mechanisms concern movements of single legs as well as the coordination between the legs. Slowing down of stance velocity, searching movements of legs in protraction and the generation of short steps are crucial prerequisites in the gap-crossing task. The rules of leg coordination described for stick insect walking seem to be modified, and load on the supporting legs is assumed to have a major effect on coordination especially in slow walking. Stepping into the gap with a front leg and antennal contact with the far edge of the gap provide information, as both events influence the following leg movements, whereas antennal non-contact seems not to contain information. Integration of these results into the model of the walking controller can improve our understanding of insect locomotion in highly irregular environments.Abbreviations AEP anterior extreme position - fAEP fictive anterior extreme position - PEP posterior extreme position - TOT treading-on-tarsus     

An integrated biomechanical analysis of normal stair ascent and descent

Three normal males of similar height and weight ascended and descended a five step staircase with a riser height of 22 cm and a tread of 28 cm. EMG, force plate and cine data were collected for the stride over the second to fourth step during each mode. Kinematic and kinetic analyses were integrated with EMG to yield an interpretation of the mechanics of normal stair walking. Movement from one step to the next involved simultaneous lifting and horizontal translation of the body, and each stride showed specific phases for progression. The extensor muscles about the knee played a dominant role in progression from one step to the next in both modes coupled with the ankle plantar flexors. The total lower limb extensor pattern, called the support moment, was highly correlated between subjects and to level walking. Intra- and inter-subject variability of the motor patterns were also determined. The greatest variability was seen at the hip, while stereotypic kinetic patterns emerged at the ankle and knee for all subjects across the 24 trials of each mode.     

A Computer-Controlled Video System for Real-Time Recording of Insect Flight in Three Dimensions

A computer-controlled video system for real-time recording of insect flight in three dimensions is described. The flight paths of moths were recorded in a flight tunnel using two CCD cameras placed adjacent to each other at angles of 45 and 135° to the flight tunnel axis and separated by a distance of 120 cm. They were connected to two 2⁸-level gray-scale frame grabbers via two external synchronizers. The two-dimensional coordinates of the flying insect were obtained from the two cameras at 40-ms intervals and transferred to host computer for processing and monitor for real-time display. Due to speed limitation in the image acquisition hardware, construction of the three-dimensional file was carried off-line. The flying insect was rendered as a dark spot in a bright background using a homogeneous light source. As the insect enters into the field of view of the two cameras, the light distribution changes, and the frame grabber detects only those variation in the light distribution which results from a flying insect. The target insect can be as small as 3 pixels and can be tracked in a stereoscopic field of view 60 cm long and 50 cm high. A method was developed that allowed for scalar scoring of various pheromone sources to assess their attractiveness using vector flight parameters. This method was applied successfully for optimization of pheromone blend of the grapevine moth, Lobesia botrana.     

The applicability of the plant vigor and resource regulation hypotheses in explaining Epiblema gall moth–Parthenium weed interactions

A basic question in insect–plant interactions is whether the insects respond to, or regulate plant traits, or a complex mixture of the two. The relative importance of the directions of the influence in insect–plant interactions has therefore been articulated through both the plant vigor hypothesis (PVH) and the resource regulation hypothesis (RRH). This study tested the applicability of these hypotheses in explaining the interactions between Parthenium hysterophorus L. (Asteraceae) and its stem‐galling moth, Epiblema strenuana Walker (Lepidoptera: Tortricidae). Parthenium plants exposed to galling were sampled at three sites in north Queensland, Australia, over a 2‐year period, and the relationship between gall abundance and plant vigor (plant height, biomass, flowers per plant, and branches per plant) was studied. To test the predictions of PVH and RRH, the vigor of parthenium plants protected from galling using insecticides was compared to galled plants and plants that escaped from galling. The vigor of ungalled plants was less than the vigor of galled plants. The higher plant vigor in galled plants was not due to galling, as was evident from insecticide exclusion trials. The insect seemed to preferentially gall the more vigorous plants. These findings support the predictions of the PVH and are contrary to those of RRH. Since gall abundance is linked to plant vigor, galling may have only a limited impact on the vigor of parthenium. This has implications for weed biological control. If the objective of biological control is to regulate the population of a plant by a galling insect, a preference for more vigorous plants by the insect is likely to limit its ability to regulate plant populations. This may explain the paucity of successes against biocontrol of annual weeds using gall insects.     

Coastal insect herbivore communities are affected more by local environmental conditions than by plant genotype

Abstract.

• 1 We compared the effects of plant genotype and local environment on population densities of a community of coastal insect herbivores in west-central Florida. Reciprocal transplants of four genotypes of three species of coastal plants, Borrichia frutescens, Iva frutescens and Limbricata, were made in July 1992 between a series of off-shore islands.
• 2 For each plant species, phytophagous insects with a wide range of feeding modes including gall-makers, stem borers, leaf miners and sap suckers were affected more by local environment than by plant genotype. Whereas host genotype had a significant effect on the population densities of gall-makers on B.frutescens in the spring of 1993, no significant effect on the denrities of any other insect species was found and the effect on the gall-makers on Borrichia disappeared in the summer, 1 year after the transplants had been made. In our study, local environment had by far the greatest effect on insect population densities among islands. This is an unusual result because in other studies over 80% of the insect species examined have been affected by plant genotype (Karban, 1992). This result is consistent with that reported by Stiling (1994), who censused populations of two phytophagous insects on reciprocal transplantr ot Borrichia in north Florida.
• 3 Local environment also had an effect on insect population densities within islands. This result contrasts with similar studies performed in north Florida (Stiling, 1994), where population densities did not differ within areas, and underlies how some biotic processes may change with in the same community even over relatively small changes in species range.

     

Three-phase DNA-origami stepper mechanism based on multi-leg interactions

Nanoscale stepper motors such as kinesin and dynein play a key role in numerous natural processes such as mitotic spindle formation during cell division or intracellular organelle transport. Their high efficacy in terms of operational speed and processivity has inspired the investigation of biomimetic technologies based on the use of programmable molecules. In particular, several designs of molecular walkers have been explored using DNA nanotechnology. Here, we study the actuation of a DNA-origami walker on a DNA-origami track based on three principles: 1) octapedal instead of bipedal walking for greater redundancy; 2) three pairs of orthogonal sequences, each of which fuels one repeatable stepping phase for cyclically driven motion with controlled directionality based on strain-based step selection; 3) designed size of only 3.5 nm per step on an origami track. All three principles are innovative in the sense that earlier demonstrations of steppers relied on a maximum of four legs on at least four orthogonal sequences to drive cyclic stepping, and took steps much larger than 3.4 nm in size. Using gel electrophoresis and negative-stain electron microscopy, we demonstrate cyclic actuation of DNA-origami structures through states defined by three sets of specific sequences of anchor points. However, this mechanism was not able to provide the intended control over directionality of movement. DNA-origami-based stepper motors will offer a future platform for investigating how increasing numbers of legs can be exploited to achieve robust stepping with relatively small step sizes.     

Identification of genes encoding N‐glycan processing β‐N‐acetylglucosaminidases in Trichoplusia ni and Bombyx mori: Implications for glycoengineering of baculovirus expression systems

Glycoproteins produced by non‐engineered insects or insect cell lines characteristically bear truncated, paucimannose N‐glycans in place of the complex N‐glycans produced by mammalian cells. A key reason for this difference is the presence of a highly specific N‐glycan processing β‐N‐acetylglucosaminidase in insect, but not in mammalian systems. Thus, reducing or abolishing this enzyme could enhance the ability of glycoengineered insects or insect cell lines to produce complex N‐glycans. Of the three insect species routinely used for recombinant glycoprotein production, the processing β‐N‐acetylglucosaminidase gene has been isolated only from Spodoptera frugiperda. Thus, the purpose of this study was to isolate and characterize the genes encoding this important processing enzyme from the other two species, Bombyx mori and Trichoplusia ni. Bioinformatic analyses of putative processing β‐N‐acetylglucosaminidase genes isolated from these two species indicated that each encoded a product that was, indeed, more similar to processing β‐N‐acetylglucosaminidases than degradative or chitinolytic β‐N‐acetylglucosaminidases. In addition, over‐expression of each of these genes induced an enzyme activity with the substrate specificity characteristic of processing, but not degradative or chitinolytic enzymes. Together, these results demonstrated that the processing β‐N‐acetylglucosaminidase genes had been successfully isolated from Trichoplusia ni and Bombyx mori. The identification of these genes has the potential to facilitate further glycoengineering of baculovirus‐insect cell expression systems for the production of glycosylated proteins. © 2009 American Institute of Chemical Engineers Biotechnol. Prog., 2010     

A note on energy expenditure in walking on level ground and uphill

In walking, energy is wasted in the process of up-and-down movement of the center of gravity of the body during each step, as well as in the kinetic energy involved in the swinging forward of each extrèmity. In this paper the frictional loss in muscles is not considered. It is shown that for a prescribed available amount of metabolic power expenditure there exists an optimal size of the step and an optimal (maximal) speed of walking for the size of the step. Calculated values are of the correct order of magnitude. In walking uphill there exists a type of step for which there is no “lost” up-and-down motion of the center of gravity of the body. This step is optimal for walking up a hill of a given incline.     

Load along the tibial shaft during activities of daily living

External load at the tibia during activities of daily living provides baseline measures for the improvement of the design of the bone–implant interface for relevant internal and external prostheses. A motion analysis system was used together with an established protocol with skin markers to estimate three-dimensional forces and moments acting on ten equidistant points along the tibial shaft. Twenty young and able-bodied volunteers were analysed while performing three repetitions of the following tasks: level walking at three different speeds, in a straight-line and with sudden changes of direction to the right and to the left, stair ascending and descending, squatting, rising from a chair and sitting down. Moment and force patterns were normalised to the percentage of body weight per height and body weight, respectively, and then averaged over all subjects for each point, about the three tibial anatomical axes, and for each task. Load patterns were found to be consistent over subjects, but different among the anatomical axes, tasks and points. Generally, moments were higher in the medio/lateral axis and influenced by walking speed. In all five walking tasks and in ascending stairs with alternating feet, the more proximal the point was the smaller the mean moment was. For the remaining tasks the opposite trend was observed. The overall largest value was observed in the medio/lateral direction at the ankle centre in level walking at high speed (9.1% body weight * height on average), nearly three times larger than that of the anterior/posterior axis (2.9) during level walking with a sidestep turn. The present results should be of value also for in-vitro mechanical tests and finite element models.     

Stable bipedal walking with a swing-leg protraction strategy

In bipedal locomotion, swing-leg protraction and retraction refer to the forward and backward motion, respectively, of the swing-leg before touchdown. Past studies have shown that swing-leg retraction strategy can lead to stable walking. We show that swing-leg protraction can also lead to stable walking. We use a simple 2D model of passive dynamic walking but with the addition of an actuator between the legs. We use the actuator to do full correction of the disturbance in a single step (a one-step dead-beat control). Specifically, for a given limit cycle we perturb the velocity at mid-stance. Then, we determine the foot placement strategy that allows the walker to return to the limit cycle in a single step. For a given limit cycle, we find that there is swing-leg protraction at shallow slopes and swing-leg retraction at steep slopes. As the limit cycle speed increases, the swing-leg protraction region increases. On close examination, we observe that the choice of swing-leg strategy is based on two opposing effects that determine the time from mid-stance to touchdown: the walker speed at mid-stance and the adjustment in the step length for one-step dead-beat control. When the walker speed dominates, the swing-leg retracts but when the step length dominates, the swing-leg protracts. This result suggests that swing-leg strategy for stable walking depends on the model parameters, the terrain, and the stability measure used for control. This novel finding has a clear implication in the development of controllers for robots, exoskeletons, and prosthetics and to understand stability in human gaits.     

On-line monitoring and control of substrate concentrations in biological processes by flow injection analysis systems

Concentrations of substrates, glucose, and ammionia in biological processes have been on-line monitored by using glucose-flow injection (FIA) and ammonia-FIA systems. Based on the on-line monitored data the concentrations of substrates have been controlled by an on-off controller, a PID controller, and a neural network (NN) based controller. A simulation program has been developed to test the control quality of each controller and to estimate the control parameters. The on-off controller often produced high oscillations at the set point due to its low robustness. The control quality of a PID controller could have been improved by a high analysis frequency and by a short residence time of sample in a FIA system. A NN-based controller with 3 layers has been developed, and a 3(input)-2(hidden)-1(output) network structure has been found to be optimal for the NN-based controller. The performance of the three controllers has been tested in a simulated process as well as in a cultivation process ofSaccharomyces cerevisiae, and the performance has also been compared to simulation results. The NN-based controller with the 3-2-1 network structure was robust and stable against some disturbances, such as a sudden injection of distilled water into a biological process.