A cockroach that jumps

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

Motor patterns during kicking movements in the locust

Locusts (Schistocerca gregaria) use a distinctive motor pattern to extend the tibia of a hind leg rapidly in a kick. The necessary force is generated by an almost isometric contraction of the extensor tibiae muscle restrained by the co-contraction of the flexor tibiae (co-contraction phase) and aided by the mechanics of the femoro-tibial joint. The stored energy is delivered suddenly when the flexor muscle is inhibited. This paper analyses the activity of motor neurons to the major hind leg muscles during kicking, and relates it to tibial movements and the resultant forces.During the co-contraction phase flexor tibiae motor neurons are driven by apparently common sources of synaptic inputs to depolarized plateaus at which they spike. The two excitatory extensor motor neurons are also depolarized by similar patterns of synaptic inputs, but with the slow producing more spikes at higher frequencies than the fast. Trochanteral depressors spike at high frequency, the single levator tarsi at low frequency, and common inhibitors 2 and 3 spike sporadically. Trochanteral levators, depressor tarsi, and a retractor unguis motor neuron are hyperpolarized.Before the tibia extends all flexor motor neurons are hyperpolarized simultaneously, two common inhibitors, and the levator trochanter and depressor tarsi motor neurons are depolarized. Later, but still before the tibial movement starts, the extensor tibiae and levator tarsi motor neurons are hyperpolarized. After the movement has started, the extensor motor neurons are hyperpolarized further and the depressor trochanteris motor neurons are also hyperpolarized, indicating a contribution of both central and sensory feedback pathways.Variations in the duration of the co-contraction of almost twenty-fold, and in the number of spikes in the fast extensor tibiae motor neuron from 2–50 produce a spectrum of tibial extensions ranging from slow and weak, to rapid and powerful. Flexibility in the networks producing the motor pattern therefore results in a range of movements suited to the fluctuating requirements of the animal.     

Neural circuits for jumping in the locust

The locust jump consists of three distinct phases: Cocking: a rapid flexion of both hindleg tibia and locking of both tibia in full flexion. Co-contraction: simultaneous contractions in hindleg flexor and extensor muscles lasting about 0.5 s resulting in the storage of energy for the jump in elastic elements of the legs and muscles. Triggering: a sudden inhibition of flexor activity to allow the shortening of the contracted extensors and the release of the energy stored during the co-contraction phase. The neural circuitry controlling these three phases is now reasonably well understood. Some of its major features are: (1) pairs of large identifiable interneurons in the thoracic ganglia for evoking the cocking response (C-neurons) and for triggering the jump (M-neurons), (2) a central excitatory pathway from extensor to flexor tibiae motoneurons to ensure simultaneous activation of extensor and flexor motoneurons during the initial part of the co-contraction phase, (3) a positive feedback pathway from cuticular receptors to extensor motoneurons for maintaining extensor activity during the co-contraction phase, (4) proprioceptive feedback to the trigger interneurons for increasing their excitability during the co-contraction phase and thereby allowing a variety of external stimuli to activate the trigger neurons and evoke a jump, (5) presynaptic inhibition of visual pathways to the trigger neurons to ensure that the trigger neurons are not activated by the simultaneous occurrence of visual and auditory stimuli in the absence of proprioceptive input, and (6) a pair of multifunctional visual movement detecting neurons which can initiate cocking or trigger the jump depending on the animal's state.     

The mechanics of elevation control in locust jumping

How do animals control the trajectory of ballistic motions like jumping? Targeted jumps by a locust, which are powered by a rapid extension of the tibiae of both hind legs, require control of the take-off angle and speed. To determine how the locust controls these parameters, we used high speed images of jumps and mechanical analysis to reach three conclusions: (1) the extensor tibiae muscle applies equal and opposite torques to the femur and tibia, which ensures that tibial extension accelerates the centre of mass of the body along a straight line; (2) this line is parallel to a line drawn from the distal end of the tibia through the proximal end of the femur; (3) the slope of this line (the angle of elevation) is not affected if the two hind legs extend asynchronously. The mechanics thus uncouple the control of elevation and speed, allowing simplified and independent control mechanisms. Jump elevation is controlled mechanically by the initial positions of the hind legs and jump speed is determined by the energy stored within their elastic processes, which allows us to then propose which proprioceptors are involved in controlling these quantities.     

Funktionsmorphologie eines arthropodengelenks mit extrem gro ßem aktionswinkel das femoro-tibial-gelenk von Gerris najas (DeG.) (inseeta,heteroptera)

Summary The femoro-tibial joint of the middle leg of Gerris najas is a single-axis hinge with an effective angle of 180°. Morphology and kinematics of this joint are described. Short sclerites are inserted between the tibia and the tendon-like apodemes of its flexor and extensor muscles. Flexible at both ends, the sclerites extend the angle of leverage by 120° in the case of the extensor tibiae and by 60° in the case of the flexor tibiae. The effective lever length was determined quantitatively for the entire 180° (see Fig. 6).     

Muscular co-contraction during walking and landing from a jump: comparison between genders and influence of activity level.

BACKGROUND: Women have higher rates of knee ligament injury than men. Co-contraction of knee muscles is proposed to be an important mechanism to protect the joint from injuries. HYPOTHESIS: Females have lower co-contraction levels when compared to males. STUDY DESIGN: Exploratory, cross-sectional design. METHODS: Thirty-six men and women equally divided into four groups according to gender and activity level (sedentary and athletic) were compared in relation to vastus lateralis and biceps femoris co-contraction before heel strike during level walking and before floor contact during landing from a jump. Muscular co-contraction was assessed by surface electromyography. Correlations between co-contraction and ligament laxity, extensor and flexor work, and flexion/extension torque ratio were also analyzed. RESULTS: No differences between genders were found in the studied situations (p0.381). During walking, co-contraction was greater in sedentary women compared to athletic women (p=0.002). A moderate inverse correlation was found between co-contraction during walking and women extensor (r=-0.613; p=0.007) and flexor (r=-0.575; p=0.012) work. During landing from a jump, no variables correlated to co-contraction in any of the groups tested (r0.477; p0.061). CONCLUSION: Co-contraction levels were not different between genders. Results suggest that women compensate strength deficits by means of increasing activation levels, possibly to generate adequate joint stiffness to meet stabilization demands. However, this is not evident in a more stressful activity like landing from a jump. CLINICAL RELEVANCE: This study contributes to a better understanding of the factors related to joint protection in females, who are at a greater risk of ligament injuries.     

Excitatory interactions between antagonistic motor neurones underlying locust kicking and jumping during maturation after the adult moult

There is a change in the synaptic connections between motor neurones that underlie locust kicking and jumping during maturation following the adult moult. The fast extensor tibiae (FETi) motor neurone makes monosynaptic excitatory connections with flexor tibiae motor neurones that have previously been implicated in maintaining flexor activity during the co-contraction phase of jumping, in which energy generated by the muscles of a hind leg is stored. The amplitude of the FETi spike decreases when repetitively activated, and this decrement is larger in locusts immediately following the adult moult than in mature locusts. The decrement in␣the FETi spike is correlated with a greater decrease in the amplitude of the flexor excitatory postsynaptic potential (EPSP) in newly moulted locusts and in turn with the failure of these locusts to kick or jump. The results presented here indicate that the developmental change in the connections between the motor neurones contributes to the change in behaviour following the moult. Accepted: 28 April 1997     

The jumping mechanism of cicada Cercopis vulnerata (Auchenorrhyncha, Cercopidae): skeleton-muscle organisation, frictional surfaces, and inverse-kinematic model of leg movements

In Auchenorrhyncha, jumping is achieved by metathoracic muscles which are inserted into the trochanter of the hind leg. The synchronisation of movements of the hind legs is a difficult problem, as the leg extension that produces the jump occurs in less than 1 ms. Even slight asynchrony could potentially result in failure of a jump. Both the synchronisation of the movements of a pair of jumping legs, and their stabilisation during a jump, seem to be important problems for small jumping insects. The present study was performed in order to clarify some questions of the functional morphology of the leafhopper jumping mechanism. It is based on skeleton-muscle reconstruction, high-speed video recordings, transmission (TEM) and scanning electron microscopic (SEM) investigations of the cuticle, together with 3D inverse-kinematic modelling of angles and working zones of hind leg joints of cicada Cercopis vulnerata (Cercopidae). The complete extension of the hind leg takes less than 1 ms, which suggests that the jump is powered not only by the muscle system, but also by an elastic spring. Histological staining and fluorescence microscopy showed resilin-bearing structures, responsible for elastic energy storage, in the pleural area of the metathorax. Synchronisation of hind leg movements may be aided by microtrichia fields that are located on the medial surface of each hind coxa. In Auchenorrhyncha, hind coxae are rounded in their anterior and lateral parts, whereas medial parts are planar, and contact each other over a rather large area. The inverse-kinematic model of propulsive leg movements was used to draw the surface outlined by the medial surface of the coxa, during the jump movement. This is a cone surface, faced with its bulged-in side, medially. Surfaces outlined by the movements of both right and left coxae overlap in their anterior and posterior positions. In both extreme positions, coxae are presumably connected to each other by coupled microtrichia fields. Thus, in extreme positions, both coxae can be moved synchronously.     

Proprioceptive feedback in locust kicking and jumping during maturation

Campaniform sensilla monitor the forces generated by the leg muscles during the co-contraction phase of locust (Schistocerca gregaria) kicking and jumping and re-excite the fast extensor (FETi) and flexor tibiae motor neurones, which innervate the leg muscles. Sensory signals from a campaniform sensillum on the proximal tibia were compared in newly moulted locusts, which do not kick and jump, and mature locusts which readily kick and jump. The activity pattern of FETi during co-contraction was mimicked by stimulating the extensor tibiae muscle. Less force was generated and the spike frequency of the sensory neurone from the sensillum was significantly lower in newly moulted compared to mature locusts. Depolarisation of both FETi and flexor motor neurones as a result of sensory feedback was consequently less in newly moulted than in mature locusts. The difference in the depolarisation was greater than the decrease in the afferent spike frequency suggesting that the central connections of the afferents are modulated. The depolarisation could generate spikes in FETi and maintain flexor spikes in mature but not in newly moulted locusts. This indicates that feedback from the anterior campaniform sensillum comprises a significant component of the drive to both FETi and flexor activity during co-contraction in mature animals and that the changes in this feedback contribute to the developmental change in behaviour.Abbreviations aCS anterior campaniform sensillum - ETi extensor tibiae - FETi fast extensor tibiae motor neurone - FlTi flexor tibiae - pCS posterior campaniform sensillum     

Comparative morphology of the tibial flexor and extensor tendons in insects

Abstract. The relative size, orientation, and degrees of sclerotization of the tibial flexor and extensor tendons are compared in nineteen orders of insects. The sclerotized, independently movable tibial flexor sclerite, known previously only from Alticinae and Carabidae (Coleoptera), is found in some other Coleoptera, Megaloptera, Neuroptera, Hymenoptera and Heteroptera. The Heteroptera also have another small sclerite at the base of the tibial extensor tendon. The tibial flexor sclerite is presumed to provide additional strength and leverage to the flexion of the tibia in certain insect groups; it may also provide protection for the ventral side of the femoro-tibial joint of the leg.     

A Three-Dimensional Musculoskeletal Model of the Human Knee Joint. Part 2: Analysis of Ligament Function

A three-dimensional model of the knee is used to study ligament function during anterior-posterior (a-p) draw, axial rotation, and isometric contractions of the extensor and flexor muscles. The geometry of the model bones is based on cadaver data. The contacting surfaces of the femur and tibia are modeled as deformable; those of the femur and patella are assumed to be rigid. Twelve elastic elements are used to describe the geometry and mechanical properties of the cruciate ligaments, the collateral ligaments, and the posterior capsule. The model is actuated by thirteen musculotendinous units, each unit represented as a three-element muscle in series with tendon. The calculations show that the forces applied during a-p draw are substantially different from those applied by the muscles during activity. Principles of knee-ligament function based on the results of in vitro experiments may therefore be overstated. Knee-ligament forces during straight a-p draw are determined solely by the changing geometry of the ligaments relative to the bones: ACL force decreases with increasing flexion during anterior draw because the angle between the ACL and the tibial plateau decreases as knee flexion increases; PCL force increases with increasing flexion during posterior draw because the angle between the PCL and the tibial plateau increases. The pattern of ligament loading during activity is governed by the geometry of the muscles spanning the knee: the resultant force in the ACL during isometric knee extension is determined mainly by the changing orientation of the patellar tendon relative to the tibia in the sagittal plane; the resultant force in the PCL during isometric knee flexion is dominated by the angle at which the hamstrings meet the tibia in the sagittal plane.     

A Three-Dimensional Musculoskeletal Model of the Human Knee Joint. Part 2: Analysis of Ligament Function

A three-dimensional model of the knee is used to study ligament function during anterior-posterior (a-p) draw, axial rotation, and isometric contractions of the extensor and flexor muscles. The geometry of the model bones is based on cadaver data. The contacting surfaces of the femur and tibia are modeled as deformable; those of the femur and patella are assumed to be rigid. Twelve elastic elements are used to describe the geometry and mechanical properties of the cruciate ligaments, the collateral ligaments, and the posterior capsule. The model is actuated by thirteen musculotendinous units, each unit represented as a three-element muscle in series with tendon. The calculations show that the forces applied during a-p draw are substantially different from those applied by the muscles during activity. Principles of knee-ligament function based on the results of in vitro experiments may therefore be overstated. Knee-ligament forces during straight a-p draw are determined solely by the changing geometry of the ligaments relative to the bones: ACL force decreases with increasing flexion during anterior draw because the angle between the ACL and the tibial plateau decreases as knee flexion increases; PCL force increases with increasing flexion during posterior draw because the angle between the PCL and the tibial plateau increases. The pattern of ligament loading during activity is governed by the geometry of the muscles spanning the knee: the resultant force in the ACL during isometric knee extension is determined mainly by the changing orientation of the patellar tendon relative to the tibia in the sagittal plane; the resultant force in the PCL during isometric knee flexion is dominated by the angle at which the hamstrings meet the tibia in the sagittal plane.     

Design and Demonstration of a Locust-Like Jumping Mechanism for Small-Scale Robots

A jumping mechanism can be an efficient mode of motion for small robots to overcome large obstacles on the ground and rough terrain.In this paper,we present a 7 g prototype of locust-inspired jumping mechanism that uses springs,wire,reduction gears,and a motor as the actuation components.The leg structure and muscles of a locust or grasshopper were mimicked using springs and wire,springs for passive extensor muscles,and a wire as a flexor muscle.A small motor was used to slowly charge the spring through a lever and gear system,and a cam with a special profile was used as a clicking mechanism for quick release of elastic energy stored in the springs to create a sudden kick for a quick jump.Performance analysis and experiments were conducted for comparison and performance estimation of the jumping mechanism prototype.Our prototype could produce standing jumps over obstacles that were about 14 times its own size (approximate to 71 cm) and a jumping distance of 20 times its own size (approximate to 100 cm).     

Effects of load inversion in cockroach walking

To examine how walking patterns are adapted to changes in load, we recorded leg movements and muscle activities when cockroaches (Periplaneta americana) walked upright and on an inverted surface. Animals were videotaped to measure the hindleg femoro-tibial joint angle while myograms were taken from the tibial extensor and flexor muscles. The joint is rapidly flexed during swing and extended in stance in upright and inverted walking. When inverted, however, swing is shorter in duration and the joint traverses a range of angles further in extension. In slow upright walking, slow flexor motoneurons fire during swing and the slow extensor in stance, although a period of co-contraction occurs early in stance. In inverted walking, patterns of muscle activities are altered. Fast flexor motoneurons fire both in the swing phase and early in stance to support the body by pulling the animal toward the substrate. Extensor firing occurs late in stance to propel the animal forward. These findings are discussed within the context of a model in which stance is divided into an early support and subsequent propulsion phase. We also discuss how these changes in use of the hindleg may represent adaptations to the reversal of the effects of gravity.     

Influence of eccentric strength of knee extensor muscles on biomechanical factors of a vertical drop jump

During a vertical drop jump (VDJ), the human neuromuscular system absorbs and reuses external loads applied to the lower extremity by coordinating the musculoskeletal system. This study aims to investigate the influence of the eccentric strength of the knee extensor muscles on the biomechanical factors of a VDJ. Participants were divided into two groups based on the eccentric strength of their knee extension muscles: low eccentric (L_ECC) and high eccentric (H_ECC) strength groups. The VDJ joint kinematics and kinetics of the lower extremity, the fascicle behavior of the vastus lateralis, and the muscle activation of the knee extensor muscles were simultaneously recorded during maximum-effort VDJ. Compared with the L_ECC group, the H_ECC group showed a higher jump, greater knee and ankle joint stiffness, and smaller fascicle length change. These findings suggest that the eccentric strength capacity of the knee extensor muscles accounts for the different biomechanical strategies (bouncing-type for H_ECC and absorbing-type for L_ECC) observed between the groups. Consequently, the eccentric strength of the knee extensor muscle may be an essential factor in determining the biomechanical strategy for VDJ and should be considered in the jumping performance enhancement training paradigm.     

Effects of movement speed and joint position on knee flexor torque in healthy and post-surgical subjects

Coactivation of knee flexors during knee extension assists in joint stability by exerting an opposing torque to the anterior tibial displacement induced by the quadriceps. This opposing torque is believed to be generated by eccentric muscle actions that stiffen the knee, thereby attenuating strain to joint ligaments, particularly the anterior cruciate ligament (ACL). However, as the lengths of knee muscles vary with changes in joint position, the magnitude of flexor/extensor muscle force coupling may likewise vary, possibly affecting the capacity for active knee stabilization. The purpose of this study was to assess the effect of changes in movement speed and joint position on eccentric/concentric muscle action relationships in the knees of uninjured (UNI) and post-ACL-surgery (INJ) subjects (n = 14). All subjects were tested for maximum eccentric and concentric torque of the contralateral knee flexors and extensor muscles at four isokinetic speeds (15 degrees-60 degrees x s(-1)) and four joint position intervals (20 degrees-60 degrees of knee flexion). Eccentric flexor torque was normalized to the percentage of concentric flexor torque generated at each joint position interval for each speed tested (flexor E-C ratio). In order to estimate the capacity of the knee flexors to resist active knee extension, the eccentric-flexor/concentric-extensor ratios were also computed for each joint position interval and speed (flexor/extensor E-C ratio). The results revealed that eccentric torque surpassed concentric torque by 3%-144% across movement speeds and joint position intervals. The magnitude of the flexor E-C ratio and flexor/extensor E-C increased significantly with speed in both groups of subjects (P < 0.05) and tended to rise with muscle length as the knee was extended; peak values were generated at the most extended joint position (20 degrees-30 degrees). Although torque development patterns were symmetrical between the contralateral limbs in both groups, between-group comparisons revealed significantly higher flexor/extensor E-C ratios for the INJ group compared to the UNI group (P < 0.05), particularly at the fastest speed tested (60 degrees x s(-1)). The results indicate that joint position and movement speed influence the eccentric/concentric relationships of knee flexors and extensors. The INJ subjects appeared to accommodate to surgery by developing the eccentric function of their ACL and normal knee flexors, particularly at higher speeds and at more extended knee joint positions. This may assist in the dynamic stabilization of the knee at positions where ACL grafts have been reported to be most vulnerable to strain.     

Are hamstrings activated to counteract shear forces during isometric knee extension efforts in healthy subjects?

The hamstring muscles have the potential to counteract anterior shear forces at the knee joint by co-contracting during knee extension efforts. Such a muscle recruitment pattern might protect the anterior cruciate ligament (ACL) by reducing its strain. In this study we investigated to what extent co-activation of the knee flexors during extension efforts is compatible with the hypothesis that this co-activation serves to counteract anterior tibial shear forces during isometric knee extension efforts in healthy subjects. To this aim, it is investigated whether co-activation varies with the required knee extension moment, with the knee joint angle, and with the position of the external flexing force relative to the knee joint. With unaltered moment and muscle activation, distal positioning of the flexing force on the tibia causes higher resultant (muscular plus external) forward shear forces at the knee as compared to proximal positioning. In ten subjects, knee flexor and extensor EMG was measured during a quasi-isometric positioning task for a range (5-50 degrees) of knee flexion angles. It was found that the co-activation of the knee flexors increased with the extension moment, but this increase was less than proportional (p<0.001). The extension moment increased 2.7 to 3.4 times, whereas the activation of Biceps Femoris and Semitendinosus increased only a factor 1.3 to 2.0 (joint angle dependent). Furthermore, a strong increase in co-activation was seen near full extension of the knee joint. The position of the external extension load on the tibia did not affect the level of co-contraction. It is argued that these results do not suggest a recruitment pattern that is directed at reduction of anterior shear forces in the knee joint during sub-maximal isometric knee extension efforts in healthy subjects.     

Femoral stiffness and jumping in grasshoppers and locusts

There is a close relationship between maximal jumping force and animal mass. Likewise, the relative stiffness of the hind femoral exoskeletal cuticle is very highly correlated with both animal mass and the force exerted during a jump. It is suggested that femoral stiffness has functional importance in the extensor energy storage system utilized in jumping by grasshoppers and locusts.