Reactive direction control for a mobile robot: A locust-like control of escape direction emerges when a bilateral pair of model locust visual neurons are integrated

Locusts possess a bilateral pair of uniquely identifiable visual neurons that respond vigorously to the image of an approaching object. These neurons are called the lobula giant movement detectors (LGMDs). The locust LGMDs have been extensively studied and this has lead to the development of an LGMD model for use as an artificial collision detector in robotic applications. To date, robots have been equipped with only a single, central artificial LGMD sensor, and this triggers a non-directional stop or rotation when a potentially colliding object is detected. Clearly, for a robot to behave autonomously, it must react differently to stimuli approaching from different directions. In this study, we implement a bilateral pair of LGMD models in Khepera robots equipped with normal and panoramic cameras. We integrate the responses of these LGMD models using methodologies inspired by research on escape direction control in cockroaches. Using ‘randomised winner-take-all’ or ‘steering wheel’ algorithms for LGMD model integration, the khepera robots could escape an approaching threat in real time and with a similar distribution of escape directions as real locusts. We also found that by optimising these algorithms, we could use them to integrate the left and right DCMD responses of real jumping locusts offline and reproduce the actual escape directions that the locusts took in a particular trial. Our results significantly advance the development of an artificial collision detection and evasion system based on the locust LGMD by allowing it reactive control over robot behaviour. The success of this approach may also indicate some important areas to be pursued in future biological research

Development and deposition of resilin in energy stores for locust jumping.

Locusts jump by using a catapult mechanism in which energy produced by slow contractions of the extensor tibiae muscles of the hind legs is stored in distortions of the exoskeleton, most notably (1) the two semi-lunar processes at each knee joint and (2) the tendons of the extensor muscles themselves. The energy is then suddenly released from these stores to power the rapid, propulsive movements of the hind legs. The reliance on the mechanical storage of energy is likely to impact on jumping because growth occurs by a series of five moults, at each of which the exoskeleton is replaced by a new one. All developmental stages (instars) nevertheless jump as a means of forward locomotion, or as an escape movement. Here, I show that in each instar, resilin is added to the semi-lunar processes and to the core of the extensor tendons so that their thickness increases. As the next moult approaches, a new exoskeleton forms within the old one, with resilin already present in the new semi-lunar processes. The old exoskeleton, the tendons and their resilin are discarded at moulting. The resilin of the semi-lunar processes and tendons of the new instar is initially thin, but a similar pattern of deposition results in an increase of their thickness. In adults, resilin continues to be deposited so that at 4 weeks old the thickness in the semi-lunar processes has increased fourfold. These changes in the energy stores accompany changes in jumping ability and performance during each moulting cycle.This is the author accepted manuscript. The final version is available from The Company of Biologists via https://doi.org/10.1242/jeb.13894

Neuromechanical Analysis of Locust Jumping

The nervous systems of animals evolved to exert dynamic control of behavior in response to the needs of the animal and changing signals from the environment. To understand the mechanisms of dynamic control, we need a means of predicting how individual neural and body elements will interact to produce the performance of the entire system. We have developed a neuromechanical application named AnimatLab that addresses this problem through simulation. A computational model of a body and nervous system can be constructed from simple components and situated in a virtual world for testing. Simulations and live experiments were used to investigate questions about locust jumping. The neural circuitry and biomechanics of kicking in locusts have been extensively studied. It has been hypothesized that the same neural circuit and biomechanics governed both behaviors, but this hypothesis was not testable with current technology. We built a neuromechanical model to test this and to gain a better understanding of the role of the semi-lunar process (SLP) in jump dynamics. The SLP are bands of cuticle that store energy for use during jumping. The results of the model were compared to a variety of published data and were similar. The SLP significantly increased jump distance, power, total energy, and duration of the jump impulse. Locust can jump precisely to a target, but also exhibit tumbling. We proposed two mechanisms for controlling tumbling during the jump. The first was that locusts adjust the pitch of their body prior to the jump to move the center of mass closer to the thrust vector. The second was that contraction of the abdominal muscles during the jump produced torques that countered the torque due to thrust. There was a strong correlation relating increased pitch and takeoff angle. In simulations there was an optimal pitch-takeoff combination that minimized tumbling that was similar to the live data. The direction and magnitude of tumbling could be controlled by adjusting abdominal tension. Tumbling also influenced jump elevation. Neuromechanical simulation addressed problems that would be difficult to examine using traditional physiological approaches. It is a powerful tool for understanding the neural basis of behavior

A Dynamics and Stability Framework for Avian Jumping Take-off

Jumping take-off in birds is an explosive behaviour with the goal of providing a rapid transition from ground to airborne locomotion. An effective jump is predicated on the need to maintain dynamic stability through the acceleration phase. The present study concerns understanding how birds retain control of body attitude and trajectory during take-off. Cursory observation suggests that stability is achieved with relatively little cost. However, analysis of the problem shows that the stability margins during jumping are actually very small and that stability considerations play a significant role in selection of appropriate jumping kinematics. We use theoretical models to understand stability in prehensile take-off (from a perch) and also in non-prehensile take-off (from the ground). The primary instability is tipping, defined as rotation of the centre of gravity about the ground contact point. Tipping occurs when the centre of pressure falls outside the functional foot. A contribution of the paper is the development of graphical tipping stability margins for both centre of gravity location and acceleration angle. We show that the nose-up angular acceleration extends stability bounds forward and is hence helpful in achieving shallow take-offs. The stability margins are used to interrogate simulated take-offs of real birds using published experimental kinematic data from a guinea fowl (ground take-off) and a diamond dove (perch take-off). For the guinea fowl the initial part of the jump is stable, however simulations exhibit a stuttering instability not observed experimentally that is probably due to absence of compliance in the idealised joints. The diamond dove model confirms that the foot provides an active torque reaction during take-off, extending the range of stable jump angles by around 45{\deg}.Comment: 21 pages, 11 figures; supplementary material: https://figshare.com/s/86b12868d64828db0d5d; DOI: 10.6084/m9.figshare.721056

Impact of Different Developmental Instars on Locusta migratoria Jumping Performance

Ontogenetic locomotion research focuses on the evolution of locomotion behavior in different developmental stages of a species. Unlike vertebrates, ontogenetic locomotion in invertebrates is poorly investigated. Locusts represent an outstanding biological model to study this issue. They are hemimetabolous insects and have similar aspects and behaviors in different instars. This research is aimed at studying the jumping performance of Locusta migratoria over different developmental instars. Jumps of third instar, fourth instar, and adult L. migratoria were recorded through a high-speed camera. Data were analyzed to develop a simplified biomechanical model of the insect: the elastic joint of locust hind legs was simplified as a torsional spring located at the femur-tibiae joint as a semilunar process and based on an energetic approach involving both locomotion and geometrical data. A simplified mathematical model evaluated the performances of each tested jump. Results showed that longer hind leg length, higher elastic parameter, and longer takeoff time synergistically contribute to a greater velocity and energy storing/releasing in adult locusts, if compared to young instars; at the same time, they compensate possible decreases of the acceleration due to the mass increase. This finding also gives insights for advanced bioinspired jumping robot design

Resilin and chitinous cuticle form a composite structure for energy storage in jumping by froghopper insects

RIGHTS : This article is licensed under the BioMed Central licence at http://www.biomedcentral.com/about/license which is similar to the 'Creative Commons Attribution Licence'. In brief you may : copy, distribute, and display the work; make derivative works; or make commercial use of the work - under the following conditions: the original author must be given credit; for any reuse or distribution, it must be made clear to others what the license terms of this work are.Abstract Background Many insects jump by storing and releasing energy in elastic structures within their bodies. This allows them to release large amounts of energy in a very short time to jump at very high speeds. The fastest of the insect jumpers, the froghopper, uses a catapult-like elastic mechanism to achieve their jumping prowess in which energy, generated by the slow contraction of muscles, is released suddenly to power rapid and synchronous movements of the hind legs. How is this energy stored? Results The hind coxae of the froghopper are linked to the hinges of the ipsilateral hind wings by pleural arches, complex bow-shaped internal skeletal structures. They are built of chitinous cuticle and the rubber-like protein, resilin, which fluoresces bright blue when illuminated with ultra-violet light. The ventral and posterior end of this fluorescent region forms the thoracic part of the pivot with a hind coxa. No other structures in the thorax or hind legs show this blue fluorescence and it is not found in larvae which do not jump. Stimulating one trochanteral depressor muscle in a pattern that simulates its normal action, results in a distortion and forward movement of the posterior part of a pleural arch by 40 μm, but in natural jumping, the movement is at least 100 μm. Conclusion Calculations showed that the resilin itself could only store 1% to 2% of the energy required for jumping. The stiffer cuticular parts of the pleural arches could, however, easily meet all the energy storage needs. The composite structure therefore, combines the stiffness of the chitinous cuticle with the elasticity of resilin. Muscle contractions bend the chitinous cuticle with little deformation and therefore, store the energy needed for jumping, while the resilin rapidly returns its stored energy and thus restores the body to its original shape after a jump and allows repeated jumping

Antibody Labelling of Resilin in Energy Stores for Jumping in Plant Sucking Insects

The rubbery protein resilin appears to form an integral part of the energy storage structures that enable many insects to jump by using a catapult mechanism. In plant sucking bugs that jump (Hemiptera, Auchenorrhyncha), the energy generated by the slow contractions of huge thoracic jumping muscles is stored by bending composite bow-shaped parts of the internal thoracic skeleton. Sudden recoil of these bows powers the rapid and simultaneous movements of both hind legs that in turn propel a jump. Until now, identification of resilin at these storage sites has depended exclusively upon characteristics that may not be specific: its fluorescence when illuminated with specific wavelengths of ultraviolet (UV) light and extinction of that fluorescence at low pH. To consolidate identification we have labelled the cuticular structures involved with an antibody raised against a product of the Drosophila CG15920 gene. This encodes pro-resilin, the first exon of which was expressed in E. coli and used to raise the antibody. We show that in frozen sections from two species, the antibody labels precisely those parts of the metathoracic energy stores that fluoresce under UV illumination. The presence of resilin in these insects is thus now further supported by a molecular criterion that is immunohistochemically specific

Insects

In this thematic series, engineers and scientists come together to address two interesting interdisciplinary questions in functional morphology and biomechanics: How do the structure and material determine the function of insect body parts? How can insects inspire engineering innovations

The jumping mechanism of flea beetles (Coleoptera, Chrysomelidae, Alticini), its application to bionics and preliminary design for a robotic jumping leg

Flea beetles (Coleoptera, Chrysomelidae, Galerucinae, Alticini) are a hyperdiverse group of organisms with approximately 9900 species worldwide. In addition to walking as most insects do, nearly all the species of flea beetles have an ability to jump and this ability is commonly understood as one of the key adaptations responsible for its diversity. Our investigation of flea beetle jumping is based on high-speed filming, micro- CT scans and 3D reconstructions, and provides a mechanical description of the jump. We reveal that the flea beetle jumping mechanism is a catapult in nature and is enabled by a small structure in the hind femur called an ‘elastic plate’ which powers the explosive jump and protects other structures from potential injury. The explosive catapult jump of flea beetles involves a unique ‘high-efficiency mechanism’ and ‘positive feedback mechanism’. As this catapult mechanism could inspire the design of bionic jumping limbs, we provide a preliminary design for a robotic jumping leg, which could be a resource for the bionics industry