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ABSTRACT: Unlike the falling cat, lizards can right themselves in mid-air by a swing of their large tails in one direction causing the body to rotate in the other. Here, we developed a new three-dimensional analytical model to investigate the effectiveness of tails as inertial appendages that change body orientation. We anchored our model using the morphological parameters of the flat-tailed house gecko Hemidactylus platyurus. The degree of roll in air righting and the amount of yaw in mid-air turning directly measured in house geckos matched the model's results. Our model predicted an increase in body roll and turning as tails increase in length relative to the body. Tails that swung from a near orthogonal plane relative to the body (i.e. 0-30° from vertical) were the most effective at generating body roll, whereas tails operating at steeper angles (i.e. 45-60°) produced only half the rotation. To further test our analytical model's predictions, we built a bio-inspired robot prototype. The robot reinforced how effective attitude control can be attained with simple movements of an inertial appendage.
Bioinspiration & Biomimetics 12/2010; 5(4):045001. · 1.95 Impact Factor
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ABSTRACT: The interplay between robotics and neuromechanics facilitates discoveries in both fields: nature provides roboticists with design ideas, while robotics research elucidates critical features that confer performance advantages to biological systems. Here, we explore a system particularly well suited to exploit the synergies between biology and robotics: high-speed antenna-based wall following of the American cockroach (Periplaneta americana). Our approach integrates mathematical and hardware modeling with behavioral and neurophysiological experiments. Specifically, we corroborate a prediction from a previously reported wall-following template - the simplest model that captures a behavior - that a cockroach antenna-based controller requires the rate of approach to a wall in addition to distance, e.g., in the form of a proportional-derivative (PD) controller. Neurophysiological experiments reveal that important features of the wall-following controller emerge at the earliest stages of sensory processing, namely in the antennal nerve. Furthermore, we embed the template in a robotic platform outfitted with a bio-inspired antenna. Using this system, we successfully test specific PD gains (up to a scale) fitted to the cockroach behavioral data in a "real-world" setting, lending further credence to the surprisingly simple notion that a cockroach might implement a PD controller for wall following. Finally, we embed the template in a simulated lateral-leg-spring (LLS) model using the center of pressure as the control input. Importantly, the same PD gains fitted to cockroach behavior also stabilize wall following for the LLS model.
IEEE Transactions on Robotics 03/2008; · 2.54 Impact Factor
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ABSTRACT: A musculo-skeletal structure can stabilize rapid locomotion using neural and/or mechanical feedback. Neural feedback results in an altered feedforward activation pattern, whereas mechanical feedback using visco-elastic structures does not require a change in the neural motor code. We selected musculo-skeletal structures in the cockroach (Blaberus discoidalis) because their single motor neuron innervation allows the simplest possible characterization of activation. We ran cockroaches over a track with randomized blocks of heights up to three times the animal's ;hip' (1.5 cm), while recording muscle action potentials (MAPs) from a set of putative control musculo-skeletal structures (femoral extensors 178 and 179). Animals experienced significant perturbations in body pitch, roll and yaw, but reduced speed by less than 20%. Surprisingly, we discovered no significant difference in the distribution of the number of MAPs, the interspike interval, burst phase or interburst period between flat and rough terrain trials. During a few very large perturbations or when a single leg failed to make contact throughout stance, neural feedback was detectable as a phase shift of the central rhythm and alteration of MAP number. System level responses of appendages were consistent with a dominant role of mechanical feedback. Duty factors and gait phases did not change for cockroaches running on flat versus rough terrain. Cockroaches did not use a follow-the-leader gait requiring compensatory corrections on a step-by-step basis. Arthropods appear to simplify control on rough terrain by rapid running that uses kinetic energy to bridge gaps between footholds and distributed mechanical feedback to stabilize the body.
Journal of Experimental Biology 03/2008; 211(Pt 3):433-46. · 3.00 Impact Factor
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ABSTRACT: Fibrillar, or "hairy," adhesives have evolved multiple times independently within arthropods and reptiles. These adhesives exhibit highly desirable properties for dynamic attachment, including orientation dependence, wear resistance, and self-cleaning. Our understanding of how these properties are related to their fibrillar structure is limited, although theoretical models from the literature have generated useful hypotheses. We survey the morphology of 81 species with fibrillar adhesives to test the hypothesis that packing density of contact elements should increase with body size, whereas the size of the contact elements should decrease. We test this hypothesis in a phylogenetic context to avoid treating historically related species as statistically independent data points. We find that fiber morphology is better predicted by evolutionary history and adhesive mechanism than by body size. As we attempt to identify which morphological parameters are most responsible for the performance of fibrillar adhesives, it will be important to take advantage of the natural variation in morphology and the potentially suboptimal outcomes it encompasses, rather than assuming evolution to be an inherently optimizing process.
Proceedings of the National Academy of Sciences 12/2007; 104(47):18595-600. · 9.68 Impact Factor
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ABSTRACT: Insects run with far greater speed and agility for their size than even the most advanced legged robots produced to date. The single-leg ground reaction forces of running insects such as the cockroach B. discoidalis provide valuable insight into the biomechanical basis for this rapid robust locomotion. To better study the running kinematics and biomechanics of these insects, a multiaxis silicon micromachined force sensor has been fabricated. The sensor consists of a 5.3-mm square plate that is supported at its corners by thin springlike beam elements. Each flexure beam is instrumented with two piezoresistive strain gauges, allowing the determination of both normal and in- plane bending force components. Typical unamplifled normal and in-plane flexure force sensitivities of 55 and 12 V/N, respectively, have been demonstrated for a sensor with 18-mum-thick flexures and a mechanical bandwidth of 1.3 kHz. Nominal normal force resolution is 2.2 nN/Hz<sup>1/2</sup> at 1 kHz. This paper details the design, fabrication, calibration, performance, and analytical modeling of the first-generation micromachined ground reaction force sensor. Preliminary data obtained from running cockroaches show that this sensor represents a marked improvement in performance over the techniques previously available for studying small-animal biomechanics.
Journal of Microelectromechanical Systems 07/2007; · 2.10 Impact Factor
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ABSTRACT: Terrestrial arthropods negotiate demanding terrain more effectively than any search-and-rescue robot. Slow, precise stepping using distributed neural feedback is one strategy for dealing with challenging terrain. Alternatively, arthropods could simplify control on demanding surfaces by rapid running that uses kinetic energy to bridge gaps between footholds. We demonstrate that this is achieved using distributed mechanical feedback, resulting from passive contacts along legs positioned by pre-programmed trajectories favorable to their attachment mechanisms. We used wire-mesh experimental surfaces to determine how a decrease in foothold probability affects speed and stability. Spiders and insects attained high running speeds on simulated terrain with 90% of the surface contact area removed. Cockroaches maintained high speeds even with their tarsi ablated, by generating horizontally oriented leg trajectories. Spiders with more vertically directed leg placement used leg spines, which resulted in more effective distributed contact by interlocking with asperities during leg extension, but collapsing during flexion, preventing entanglement. Ghost crabs, which naturally lack leg spines, showed increased mobility on wire mesh after the addition of artificial, collapsible spines. A bioinspired robot, RHex, was redesigned to maximize effective distributed leg contact, by changing leg orientation and adding directional spines. These changes improved RHex's agility on challenging surfaces without adding sensors or changing the control system.
Bioinspiration & Biomimetics 04/2007; 2(1):9-18. · 1.95 Impact Factor
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ABSTRACT: The mechanical behavior of muscle during locomotion is often predicted by its anatomy, kinematics, activation pattern and contractile properties. The neuromuscular design of the cockroach leg provides a model system to examine these assumptions, because a single motor neuron innervates two extensor muscles operating at a single joint. Comparisons of the in situ measurements under in vivo running conditions of muscle 178 to a previously examined muscle (179) demonstrate that the same inputs (e.g. neural signal and kinematics) can result in different mechanical outputs. The same neural signal and kinematics, as determined during running, can result in different mechanical functions, even when the two anatomically similar muscles possess the same contraction kinetics, force-velocity properties and tetanic force-length properties. Although active shortening greatly depressed force under in vivo-like strain and stimulation conditions, force depression was similarly proportional to strain, similarly inversely proportional to stimulation level, and similarly independent of initial length and shortening velocity between the two muscles. Lastly, passive pre-stretch enhanced force similarly between the two muscles. The forces generated by the two muscles when stimulated with their in vivo pattern at lengths equal to or shorter than rest length differed, however. Overall, differences between the two muscles in their submaximal force-length relationships can account for up to 75% of the difference between the two muscles in peak force generated at short lengths observed during oscillatory contractions. Despite the fact that these muscles act at the same joint, are stimulated by the same motor neuron with an identical pattern, and possess many of the same in vitro mechanical properties, the mechanical outputs of two leg extensor muscles can be vastly different.
Journal of Experimental Biology 10/2006; 209(Pt 17):3370-82. · 3.00 Impact Factor
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ABSTRACT: The American cockroach, Periplaneta americana, is reported to follow walls at a rate of up to 25 turns s(-1). During high-speed wall following, a cockroach holds its antenna relatively still at the base while the flagellum bends in response to upcoming protrusions. We present a simple mechanosensory model for the task-level dynamics of wall following. In the model a torsional, mass-damper system describes the cockroach's turning dynamics, and a simplified antenna measures distance from the cockroach's centerline to a wall. The model predicts that stabilizing neural feedback requires both proportional feedback (difference between the actual and desired distance to wall) and derivative feedback (velocity of wall convergence) information from the antenna. To test this prediction, we fit a closed-loop proportional-derivative control model to trials in which blinded cockroaches encountered an angled wall (30 degrees or 45 degrees ) while running. We used the average state of the cockroach in each of its first four strides after first contacting the angled wall to predict the state in each subsequent stride. Nonlinear statistical regression provided best-fit model parameters. We rejected the hypothesis that proportional feedback alone was sufficient. A derivative (velocity) feedback term in the control model was necessary for stability.
Journal of Experimental Biology 06/2006; 209(Pt 9):1617-29. · 3.00 Impact Factor
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ABSTRACT: Legs of sprawled-posture, quadrupedal trotting geckos (Hemidactylus garnotii) each functioned differently during constant average-speed locomotion. The center of mass decelerated in the first half of a step and accelerated in the second half, as if geckos were bouncing in fore-aft and side-to-side directions. Forelegs decelerated the center of mass only in the fore-aft direction. Hindlegs provided all the acceleration in the latter half of the step. Lateral ground reaction forces were always directed toward the midline and exceeded the magnitude of fore-aft forces. The differential leg function of sprawled-posture geckos resembled sprawled-posture hexapods more than upright-posture quadrupeds. The pattern of leg ground reaction forces observed may provide passive, dynamic stability while minimizing joint moments, yet allow high maneuverability. Integrating limb dynamics with whole body dynamics is required to resolve the trade-offs, if any, that result from stable sprawled-posture running with differential leg function.
Journal of Experimental Biology 02/2006; 209(Pt 2):249-59. · 3.00 Impact Factor
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ABSTRACT: Geckos with adhesive toe pads rapidly climb even smooth vertical surfaces. We challenged geckos (Hemidactylus garnotii) to climb up a smooth vertical track that contained a force platform. Geckos climbed vertically at up to 77 cm s(-1) with a stride frequency of 15 Hz using a trotting gait. During each step, whole body fore-aft, lateral and normal forces all decreased to zero when the animal attached or detached its toe pads. Peak fore-aft force was twice body weight at mid-step. Geckos climbed at a constant average velocity without generating decelerating forces on their center of mass in the direction of motion. Although mass-specific mechanical power to climb was ten times the value expected for level running, the total mechanical energy of climbing was only 5-11% greater than the potential energy change. Fore- and hindlegs both pulled toward the midline, possibly loading the attachment mechanisms. Attachment and detachment of feet occupied 13% and 37% of stance time, respectively. As climbing speed increased, the absolute time required to attach and detach did not decrease, suggesting that the period of fore-aft force production might be constrained. During ascent, the forelegs pulled toward, while hindlegs pushed away from the vertical surface, generating a net pitching moment toward the surface to counterbalance pitch-back away from the surface. Differential leg function appears essential for effective vertical as well as horizontal locomotion.
Journal of Experimental Biology 02/2006; 209(Pt 2):260-72. · 3.00 Impact Factor
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ABSTRACT: A multi-axis sensor with micro-Newton force resolution has been designed and fabricated for the purpose of studying the biomechanics of insect locomotion. The sensor comprises a rigid central plate supported at each corner by thin beams with piezoresistive transducers sensitive to both normal and in plane forces. The sensor is configured to measure the instantaneous ground reaction force components produced by insects as large as cockroaches when they step onto the sensor plate during walking or running. This paper describes the results of analytical modeling and finite element simulations used to characterize the performance and mechanical behavior of the sensor microstructure.
TRANSDUCERS, Solid-State Sensors, Actuators and Microsystems, 12th International Conference on, 2003; 07/2003
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ABSTRACT: The individual muscles of a multiple muscle group at a given joint are often assumed to function synergistically to share the load during locomotion. We examined two leg extensors of a running cockroach to test the hypothesis that leg muscles within an anatomical muscle group necessarily manage (i.e. produce, store, transmit or absorb) energy similarly during running. Using electromyographic and video motion-analysis techniques, we determined that muscles 177c and 179 are both active during the first half of the stance period during muscle shortening. Using the in vivo strain and stimulation patterns determined during running, we measured muscle power output. Although both muscles were stimulated during the first half of shortening, muscle 177c generated mechanical energy (28 W x kg(-1)) like a motor, while muscle 179 absorbed energy (-19 W x kg(-1)) like a brake. Both muscles exhibited nearly identical intrinsic characteristics including similar twitch kinetics and force-velocity relationships. Differences in the extrinsic factors of activation and relative shortening velocity caused the muscles to operate very differently during running. Presumed redundancy in a multiple muscle group may, therefore, represent diversity in muscle function. Discovering how muscles manage energy during behavior requires the measurement of a large number of dynamically interacting variables.
Journal of Experimental Biology 03/2002; 205(Pt 3):379-89. · 3.00 Impact Factor
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ABSTRACT: Geckos are exceptional in their ability to climb rapidly up smooth vertical surfaces. Microscopy has shown that a gecko's foot has nearly five hundred thousand keratinous hairs or setae. Each 30-130 microm long seta is only one-tenth the diameter of a human hair and contains hundreds of projections terminating in 0.2-0.5 microm spatula-shaped structures. After nearly a century of anatomical description, here we report the first direct measurements of single setal force by using a two-dimensional micro-electromechanical systems force sensor and a wire as a force gauge. Measurements revealed that a seta is ten times more effective at adhesion than predicted from maximal estimates on whole animals. Adhesive force values support the hypothesis that individual seta operate by van der Waals forces. The gecko's peculiar behaviour of toe uncurling and peeling led us to discover two aspects of setal function which increase their effectiveness. A unique macroscopic orientation and preloading of the seta increased attachment force 600-fold above that of frictional measurements of the material. Suitably orientated setae reduced the forces necessary to peel the toe by simply detaching above a critical angle with the substratum.
Nature 07/2000; 405(6787):681-5. · 36.28 Impact Factor
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ABSTRACT: Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.
Science 05/2000; 288(5463):100-6. · 31.20 Impact Factor
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ABSTRACT: Locomotion results from complex, high-dimensional, non-linear, dynamically coupled interactions between an organism and its environment. Fortunately, simple models we call templates have been and can be made to resolve the redundancy of multiple legs, joints and muscles by seeking synergies and symmetries. A template is the simplest model (least number of variables and parameters) that exhibits a targeted behavior. For example, diverse species that differ in skeletal type, leg number and posture run in a stable manner like sagittal- and horizontal-plane spring-mass systems. Templates suggest control strategies that can be tested against empirical data. Templates must be grounded in more detailed morphological and physiological models to ask specific questions about multiple legs, the joint torques that actuate them, the recruitment of muscles that produce those torques and the neural networks that activate the ensemble. We term these more elaborate models anchors. They introduce representations of specific biological details whose mechanism of coordination is of interest. Since mechanisms require controls, anchors incorporate specific hypotheses concerning the manner in which unnecessary motion or energy from legs, joints and muscles is removed, leaving behind the behavior of the body in the low-degree-of-freedom template. Locating the origin of control is a challenge because neural and mechanical systems are dynamically coupled and both play a role. The control of slow, variable-frequency locomotion appears to be dominated by the nervous system, whereas during rapid, rhythmic locomotion, the control may reside more within the mechanical system. Anchored templates of many-legged, sprawled-postured animals suggest that passive, dynamic self-stabilization from a feedforward, tuned mechanical system can reject rapid perturbations and simplify control. Future progress would benefit from the creation of a field embracing comparative neuromechanics.
Journal of Experimental Biology 01/2000; 202(Pt 23):3325-32. · 3.00 Impact Factor
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ABSTRACT: We tested the hypothesis that fast-running hexapeds must generate high levels of kinetic energy to cycle their limbs rapidly compared with bipeds and quadrupeds. We used high-speed video analysis to determine the three-dimensional movements of the limbs and bodies of cockroaches (Blaberus discoidalis) running on a motorized treadmill at 21 cm s-1 using an alternating tripod gait. We combined these kinematic data with morphological data to calculate the mechanical energy produced to move the limbs relative to the overall center of mass and the mechanical energy generated to rotate the body (head + thorax + abdomen) about the overall center of mass. The kinetic energy involved in moving the limbs was 8 microJ stride-1 (a power output of 21 mW kg-1, which was only approximately 13% of the external mechanical energy generated to lift and accelerate the overall center of mass at this speed. Pitch, yaw and roll rotational movements of the body were modest (less than +/- 7 degrees), and the mechanical energy required for these rotations was surprisingly small (1.7 microJ stride-1 for pitch, 0.5 microJ stride-1 for yaw and 0.4 microJ stride-1 for roll) as was the power (4.2, 1.2 and 1.1 mW kg-1, respectively). Compared at the same absolute forward speed, the mass-specific kinetic energy generated by the trotting hexaped to swing its limbs was approximately half of that predicted from data on much larger two- and four-legged animals. Compared at an equivalent speed (mid-trotting speed), limb kinetic energy was a smaller fraction of total mechanical energy for cockroaches than for large bipedal runners and hoppers and for quadrupedal trotters. Cockroaches operate at relatively high stride frequencies, but distribute ground reaction forces over a greater number of relatively small legs. The relatively small leg mass and inertia of hexapeds may allow relatively high leg cycling frequencies without exceptionally high internal mechanical energy generation.
Journal of Experimental Biology 08/1997; 200(Pt 13):1919-29. · 3.00 Impact Factor
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ABSTRACT: Stability is fundamental to the performance of terrestrial locomotion. Running cockroaches met the criteria for static stability over a wide range of speeds, yet several locomotor variables changed in a way that revealed an increase in the importance of dynamic stability as speed increased. Duty factors (the fraction of time that a leg spends on the ground relative to the stride period) decreased to 0.5 and below with an increase in speed. The duration of double support (i.e. when both tripods, or all six legs, were on the ground) decreased significantly with an increase in speed. All legs had similar touch-down phases in the tripod, but the shortest leg, the front one, lifted off before the middle and the rear leg, so that only two legs of the tripod were in contact with the ground at the highest speeds. Per cent stability margin (the shortest distance from the center of gravity to the boundaries of support, normalized to the maximum possible stability margin) decreased with increasing speed from 60% at 10 cms-1 to values less than zero at speeds faster than 50 cms-1, indicating instances of static instability at fast speeds. The center of mass moved rearward or posteriorly with respect to the base of support as speed increased. Moments about the center of mass, as shown by the center of pressure (the equivalent of a single 'effective' leg), were variable, but were balanced by opposing moments over a stride. Thus, hexapods can exploit the advantages of both static and dynamic stability. Static or quasi-static assumptions alone were insufficient to explain straight-ahead, constant-speed locomotion and may hinder discovery of behaviors that are dynamic, where kinetic energy and momentum can act as a bridge from one step to the next.
Journal of Experimental Biology 01/1995; 197:251-69. · 3.00 Impact Factor
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ABSTRACT: Despite impressive variation in leg number, length, position and type of skeleton, similarities of legged, pedestrian locomotion exist in energetics, gait, stride frequency and ground-reaction force. Analysis of data available in the literature showed that a bouncing, spring-mass, monopode model can approximate the energetics and dynamics of trotting, running, and hopping in animals as diverse as cockroaches, quail and kangaroos. From an animal's mechanical-energy fluctuation and ground-reaction force, we calculated the compression of a virtual monopode's leg and its stiffness. Comparison of dimensionless parameters revealed that locomotor dynamics depend on gait and leg number and not on body mass. Relative stiffness per leg was similar for all animals and appears to be a very conservative quantity in the design of legged locomotor systems. Differences in the general dynamics of gait are based largely on the number of legs acting simultaneously to determine the total stiffness of the system. Four- and six-legged trotters had a greater whole body stiffness than two-legged runners operating their systems at about the same relative speed. The greater whole body stiffness in trotters resulted in a smaller compression of the virtual leg and a higher natural frequency and stride frequency.
Journal of Comparative Physiology 01/1993; 173(5):509-517. · 2.01 Impact Factor
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ABSTRACT: Most animals move intermittently, yet many proposed performance limitations of terrestrial locomotion are based on steady-state measurements and assumptions. We examined the effect of work-rest transitions by exercising the ghost crab, Ocypode quadrata (28.1 +/- 8.1 g), intermittently on a treadmill at 0.30 m/s, a supramaximal speed [i.e., greater than the speed that elicits the maximal rate of oxygen consumption (VO2)]. Duration of the exercise and pause periods, ratio of exercise to pause, and speed during the exercise period were varied to determine the effect on performance. Crabs fatigued after 7.5 min of continuous running, a distance capacity (i.e., total distance traveled before fatigue) of 135 m. When the task was done intermittently with 2-min exercise and 2-min pause periods, the crabs fatigued after 87 min (a total distance of 787 m), representing an 5.8-fold increase in distance capacity compared with continuous exercise at the same absolute speed (0.30 m/s) and a 2.2-fold increase in distance capacity compared with continuous exercise at the same average speed (0.15 m/s). Pause periods less than 30 s did not result in greater distance capacity compared with continuous exercise at the same average speed. Longer (3-5 min) and shorter exercise periods (less than or equal to 30 s) decreased distance capacity. Leg muscle lactate increased 10-fold to 15 mumol/g leg during intermittent exercise. However, significant amounts of lactate were cleared from the leg during the brief pause periods.(ABSTRACT TRUNCATED AT 250 WORDS)
The American journal of physiology 06/1992; 262(5 Pt 2):R852-9.
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ABSTRACT: Many-legged animals, such as crabs and cockroaches, utilize whole-body mechanics similar to that observed for running bipeds and trotting quadrupedal mammals. Despite the diversity in morphology, two legs in a quadrupedal mammal, three legs in an insect and four legs in a crab can function in the same way as one leg of a biped during ground contact. To explain how diverse leg designs can result in common whole-body dynamics, we used a miniature force platform to measure the ground reaction forces produced by individual legs of the cockroach Blaberus discoidalis. Hexapedal runners were not like quadrupeds with an additional set of legs. In trotting quadrupedal mammals each leg develops a similar ground reaction force pattern that sums to produce the whole-body pattern. At a constant average velocity, each leg pair of the cockroach was characterized by a unique ground reaction force pattern. The first leg decelerated the center of mass in the horizontal direction, whereas the third leg was used to accelerate the body. The second leg did both, much like legs in bipedal runners and quadrupedal trotters. Vertical force peaks for each leg were equal in magnitude. In general, peak ground reaction force vectors minimized joint moments and muscle forces by being oriented towards the coxal joints, which articulate with the body. Locomotion with a sprawled posture does not necessarily result in large moments around joints. Calculations on B. discoidalis showed that deviations from the minimum moments may be explained by considering the minimization of the summed muscle forces in more than one leg. Production of horizontal forces that account for most of the mechanical energy generated during locomotion can actually reduce total muscle force by directing the ground reaction forces through the leg joints. Whole-body dynamics common to two-, four-, six- and eight-legged runners is produced in six-legged runners by three pairs of legs that differ in orientation with respect to the body, generate unique ground reaction force patterns, but combine to function in the same way as one leg of a biped.
Journal of Experimental Biology 08/1991; 158:369-90. · 3.00 Impact Factor
