Biomedical subjects

R J Full

Publications and source records attributed to R J Full.

At least 19 recordsLinked to original sources

PubMed2006

Differential leg function in a sprawled-posture quadrupedal trotter.

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.

Analysis of Variance↗

PubMed2006

Dynamics of geckos running vertically.

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.

Adhesiveness↗

PubMed2006

Task-level control of rapid wall following in the American cockroach.

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.

Animals↗

PubMed2006

In situ muscle power differs without varying in vitro mechanical properties in two insect leg muscles innervated by the same motor neuron.

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.

Analysis of Variance↗

PubMed2002

Dynamics and stability of legged locomotion in the horizontal plane: a test case using insects.

Motivated by experimental studies of insects, we propose a model for legged locomotion in the horizontal plane. A three-degree-of freedom, energetically conservative, rigid-body model with a pair of compliant virtual legs in intermittent contact with the ground allows us to study how dynamics depends on parameters such as mass, moment of inertia, leg stiffness, and length. We find periodic gaits, and show that mechanics alone can confer asymptotic stability of relative heading and body angular velocity. We discuss the relevance of our idealized models to experiments and simulations on insect running, showing that their gait and force characteristics match observations reasonably well. We perform parameter studies and suggest that our model is relevant to the understanding of locomotion dynamics across species.

Animals↗

PubMed2002

A motor and a brake: two leg extensor muscles acting at the same joint manage energy differently in a running insect.

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.

Animals↗

PubMed2000

Adhesive force of a single gecko foot-hair.

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.

Adhesiveness↗

PubMed2000

How animals move: an integrative view.

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.

Animals↗

PubMed1999

Templates and anchors: neuromechanical hypotheses of legged locomotion on land.

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.

Animals↗

PubMed1997

Three-dimensional kinematics and limb kinetic energy of running cockroaches.

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.

Animals↗

PubMed1994

Dynamic and static stability in hexapedal runners.

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.

Animals↗

PubMed1992

Intermittent exercise alters endurance in an eight-legged ectotherm.

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)

Aerobiosis↗

PubMed1991

Mechanics of a rapid running insect: two-, four- and six-legged locomotion.

To examine the effects of variation in body form on the mechanics of terrestrial locomotion, we used a miniature force platform to measure the ground reaction forces of the smallest and, relative to its mass, one of the fastest invertebrates ever studied, the American cockroach Periplaneta americana (mass = 0.83 g). From 0.44-1.0 ms-1, P. americana used an alternating tripod stepping pattern. Fluctuations in gravitational potential energy and horizontal kinetic energy of the center of mass were nearly in phase, characteristic of a running or bouncing gait. Aerial phases were observed as vertical ground reaction force approached zero at speeds above 1 ms-1. At the highest speeds (1.0-1.5 ms-1 or 50 body lengths per second), P. americana switched to quadrupedal and bipedal running. Stride frequency approached the wing beat frequencies used during flight (27 Hz). High speeds were attained by increasing stride length, whereas stride frequency showed little increase with speed. The mechanical power used to accelerate the center of mass increased curvilinearly with speed. The mass-specific mechanical energy used to move the center of mass a given distance was similar to that measured for animals five orders of magnitude larger in mass, but was only one-hundredth of the metabolic cost.

Analog-Digital Conversion↗

PubMed1991

Leg design in hexapedal runners.

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.

Animals↗

PubMed1991

Animal motility and gravity.

1. Specific resistance to locomotion will remain constant if changes in the amount of force produced or in the cost of force production are directly proportional to changes in gravitational acceleration. 2. If changes in gravitational acceleration result in a disproportionate decrease of the effective mechanical advantage of limbs or reduction in contact time for developing force, then specific resistance may increase. 3. If changes in gravitational acceleration disproportionately increase effective mechanical advantage or the contact time for developing force, then specific resistance may decrease. 4. Finally, changes in the direction and magnitude of gravitational acceleration will affect the energetics and mechanics of animal motility even for small runners that appear to defy gravity.

Animals↗

PubMed1990

Mechanics of six-legged runners.

Six-legged pedestrians, cockroaches, use a running gait during locomotion. The gait was defined by measuring ground reaction forces and mechanical energy fluctuations of the center of mass in Blaberus discoidalis (Serville) as they travelled over a miniature force platform. These six-legged animals produce horizontal and vertical ground-reaction patterns of force similar to those found in two-, four- and eight-legged runners. Lateral forces were less than half the vertical force fluctuations. At speeds between 0.08 and 0.66 ms-1, horizontal kinetic and gravitational potential energy changes were in phase. This pattern of energy fluctuation characterizes the bouncing gaits used by other animals that run. Blaberus discoidalis attained a maximum sustainable stride frequency of 13 Hz at 0.35 ms-1, the same speed and frequency predicted for a mammal of the same mass. Despite differences in body form, the mass-specific energy used to move the center of mass a given distance (0.9 J kg-1m-1) was the same for cockroaches, ghost crabs, mammals, and birds. Similarities in force production, stride frequency and mechanical energy production during locomotion suggest that there may be common design constraints in terrestrial locomotion which scale with body mass and are relatively independent of body form, leg number and skeletal type.

Animals↗

PubMed1990

Energetics of ascent: insects on inclines.

Small animals use more metabolic energy per unit mass than large animals to run on a level surface. If the cost to lift one gram of mass one vertical meter is constant, small animals should require proportionally smaller increases in metabolic cost to run uphill. To test this hypothesis on very small animals possessing an exceptional capacity for ascending steep gradients, we measured the metabolic cost of locomotion in the cockroach, Periplaneta americana, running at angles of 0, 45 and 90 degrees to the horizontal. Resting oxygen consumption (VO2rest) was not affected by incline angle. Steady-state oxygen consumption (VO2ss) increased linearly with speed at all angles of ascent. The minimum cost of locomotion (the slope of the VO2ss versus speed function) increased with increasing angle of ascent. The minimum cost of locomotion on 45 and 90 degrees inclines was two and three times greater, respectively, than the cost during horizontal running. The cockroach's metabolic cost of ascent greatly exceeds that predicted from the hypothesis of a constant efficiency for vertical work. Variations in stride frequency and contact time cannot account for the high metabolic cost, because they were independent of incline angle. An increase in the metabolic cost or amount of force production may best explain the increase in metabolic cost. Small animals, such as P. americana, can easily scale vertical surfaces, but the energetic cost is considerable.

Animals↗

PubMed1990

Effect of variation in form on the cost of terrestrial locomotion.

The mass-specific minimum cost of terrestrial locomotion (Cmin) decreases with an increase in body mass. This generalization spans nearly eight orders of magnitude in body mass and includes two phyla. The general relationship between metabolic cost and mass is striking. However, a significant amount of unexplained interspecific variation in Cmin exists at any given body mass. To determine how variation in morphology and physiology affects metabolic energy cost, we measured the oxygen consumption of three comparably sized insects running on a miniature treadmill; the American cockroach Periplaneta americana, the caterpillar hunting beetle Calosoma affine and the Australian field cricket Teleogryllus commodus. Steady-state oxygen consumption (VO2ss) increased linearly with speed. Cmin was similar for crickets and cockroaches (8.0 and 8.5 ml O2 g-1km-1, respectively), but was substantially lower for beetles (4.6 ml O2 g-1km-1). The predicted value of Cmin for all three insects was within the 95% confidence intervals of the Cmin versus body mass function. However, the 95% confidence intervals extend approximately 2.5-fold above and 40% below the regression line, making the variation at any given body mass nearly sixfold. Normalizing for the rate of muscle force production by determining the metabolic cost per stride failed to account for the interspecific variation in the cost of locomotion observed in the three insects. Ground contact costs (i.e. VO2ss multiplied by leg contact time during a stride) in insects were similar to those measured in mammals (1.5-3.1 J kg-1) and were independent of speed, but did not explain the interspecific variation in the cost of locomotion. Muscles of the caterpillar hunting beetle may have a greater mechanical advantage than muscles of the Australian field cricket and American cockroach. Variation in musculo-skeletal arrangement, apart from variation in body mass, could translate into significant differences in the minimum cost of terrestrial locomotion.

Animals↗