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Malcolm Burrows

Publications and source records attributed to Malcolm Burrows.

12 recordsLinked to original sources
PubMed2006

Neural control and coordination of jumping in froghopper insects.

The thrust for jumping in froghopper insects is produced by a rapid, synchronous depression of both hind legs generated by huge, multipartite trochanteral depressor muscles in the thorax and smaller levator muscles in the coxae. A three-phase motor pattern activates these muscles in jumping. First, a levation phase lasts a few hundred milliseconds, in which a burst of spikes in the trochanteral levator motor neurons moves the hind legs into their fully cocked position and thus engages a mechanical lock between a coxa and a femur. Second, a cocked phase lasts a few seconds, in which a trochanteral depressor motor neuron spikes continuously at a frequency gradually rising to 50 Hz, although the hind legs remain stationary. Levator motor spikes are sporadic. Third, the jump movement lasts <1 ms, in which the spikes in the depressors stop abruptly and the legs rapidly depress. This pattern may vary in the speed of the initial levation and in the duration of the cocked phase. Recordings from the depressor muscles on both sides showed remarkable synchrony of their motor spikes. In one 4.9-long cocked phase all 174 spikes were synchronous and in another 27 s period of continuous spiking all but one of 1,176 spikes were synchronous. When a single hind leg moves rapidly, these depressor spikes are nevertheless independent of those of the other leg. These features of the motor pattern and the coupling between motor neurons to the two hind legs ensure powerful movements to propel rapid jumping.

Action Potentials↗
PubMed2006

Jumping performance of froghopper insects.

The kinematics of jumping in froghopper insects were analysed from high speed sequences of images captured at rates up to 8000 s(-1). In a jump, the attitude of the body is set by the front and middle legs, and the propulsion is delivered by rapid and synchronous movements of the hind legs that are 1.5 times longer than the other legs, but are only about half the length of the body and represent just 2% of the body mass. The wings are not moved and the front and middle legs may be raised off the ground before take-off. The hind legs are first cocked by a slow levation of the trochantera about the coxae so that the femora are pressed against the ventral, indented wall of the thorax, with the femoro-tibial joints tucked between the middle legs and body. Only the tips of the hind tarsi are in contact with the ground. In this position, the hind legs stay motionless for 1-2 s. Both trochantera are then synchronously and rapidly depressed about the coxae at rotational velocities of 75 500 deg. s(-1) and the tibiae extended, to launch a jump that in Philaenus reaches a height of 700 mm, or 115 body lengths. In the best jumps by Philaenus, take-off occurs within 0.875 ms of the start of movements of the hind legs at a peak velocity of 4.7 m s(-1) and involves an acceleration of 5400 m s(-2), equivalent to 550 times gravity. This jumping performance requires an energy output of 136 microJ, a power output of 155 mW and exerts a force of 66 mN.

Animals↗
PubMed2006

Morphology and action of the hind leg joints controlling jumping in froghopper insects.

The morphology and movements of key joints of the hind legs that generate the rapid jumping of froghoppers were analysed. The movements of an individual hind leg during a jump occur in three phases. First, the trochanter is slowly levated about the coxa so that the femur moves anteriorly and engages with a lateral protrusion on the coxa. Second, both hind legs are held in this fully levated (cocked) position without moving for a few seconds. Third, both hind legs depress and extend completely in less than 1 ms. The critical, power-generating movement underlying a jump is the rapid and simultaneous depression of the trochantera about the coxae. The lever arm of the hind trochanteral depressor muscle is smallest at the cocked position, but does not appear to go over the centre of the pivot. It then increases to a maximum after some 80 degrees of depression movement. By contrast, the lever arm of the trochanteral levator tendon is similar over the range of joint movements and is exceeded by that of the depressor only after 40 degrees of depression. Three prominent arrays of hairs on the trochantin, coxa and trochanter are appropriately positioned to act as proprioceptors signalling key movements in jumping. In the fully levated position, a protrusion on the dorsal, proximal surface of a hind femur engages with a protrusion from the ventral and lateral part of a coxa. These structures are not present on the front and middle legs. Both protrusions are covered with a dense array of small projections (microtrichia) that both increase the surface area and may interlock with each other. To depress rapidly in a jump these protrusions must disengage. If the hind leg of a dead froghopper is forcibly levated, it will lock in its cocked position, from which it can depress rapidly by movement of the coxo-trochanteral joint and disengagement of the femoral and coxal protrusions. A prominent click sound occurs at the start of a jump that results either from the initial movements of the coxo-trochanteral joint, or from the disengagement of the microtrichia on the coxa and femur. Larval Philaenus, which do not jump, lack a femoral protrusion and have no microtrichia in equivalent positions on either the coxa or femur.

Animals↗
PubMed2004

Projection patterns of posterior dorsal unpaired median neurons of the locust subesophageal ganglion.

Six neurons in a group of dorsal unpaired median (DUM) neurons with cell bodies in the posterior part-maxillary and labial neuromeres-of the subesophageal ganglion of locusts have two axons each that descend into both the left and the right halves of the ganglia of the ventral nerve cord. None of the neurons has peripheral axons, so they are interneurons. Electrophysiology shows that the axons of at least four neurons project to the terminal abdominal ganglion to which they conduct spikes at a velocity of 0.5-0.6 m. second(-1). In the somata, the spikes have a smaller amplitude and briefer duration at half height than the spikes of thoracic, efferent DUM neurons. Each neuron has bilaterally symmetrical branches within the subesophageal ganglion and in the thoracic ganglia. On the basis of the specific patterns of branches, and the neuropiles, tracts, and commissures in which they occur, three types of neurons (DUM SD 1-3) can be recognized. DUM SD 1 and 3 project to ventral regions of neuropile in the thoracic ganglia in which the efferent DUM neurons of these ganglia have no branches. DUM SD 2 projects to dorsal neuropiles. The projection patterns of these putatively octopaminergic neurons suggest that they could be the source of the octopaminergic modulation of networks underlying sensory processing and motor pattern generation within these ganglia. Within this group of posterior DUM neurons, two additional cells were stained that have axons ascending to the brain.

Animals↗
PubMed2004

Substantial changes in central nervous system neurotransmitters and neuromodulators accompany phase change in the locust.

Desert locusts (Schistocerca gregaria) can undergo a profound transformation between solitarious and gregarious forms, which involves widespread changes in behaviour, physiology and morphology. This phase change is triggered by the presence or absence of other locusts and occurs over a timescale ranging from hours, for some behaviours to change, to generations, for full morphological transformation. The neuro-hormonal mechanisms that drive and accompany phase change in either direction remain unknown. We have used high-performance liquid chromatography (HPLC) to compare amounts of 13 different potential neurotransmitters and/or neuromodulators in the central nervous systems of final instar locust nymphs undergoing phase transition and between long-term solitarious and gregarious adults. Long-term gregarious and solitarious locust nymphs differed in 11 of the 13 substances analysed: eight increased in both the brain and thoracic nerve cord (including glutamate, GABA, dopamine and serotonin), whereas three decreased (acetylcholine, tyramine and citrulline). Adult locusts of both extreme phases were similarly different. Isolating larval gregarious locusts led to rapid changes in seven chemicals equal to or even exceeding the differences seen between long-term solitarious and gregarious animals. Crowding larval solitarious locusts led to rapid changes in six chemicals towards gregarious values within the first 4 h (by which time gregarious behaviours are already being expressed), before returning to nearer long-term solitarious values 24 h later. Serotonin in the thoracic ganglia, however, did not follow this trend, but showed a ninefold increase after a 4 h period of crowding. After crowding solitarious nymphs for a whole larval stadium, the amounts of all chemicals, except octopamine, were similar to those of long-term gregarious locusts. Our data show that changes in levels of neuroactive substances are widespread in the central nervous system and reflect the time course of behavioural and physiological phase change.

Acetylcholine↗
PubMed2003

Localisation of Even-skipped in the mature CNS of the locust, Schistocerca gregaria.

Even-skipped (Eve) is a transcriptional repressor found within the nuclei of a subset of neurons in the embryonic central nervous system of insects. It is necessary for neuronal pathfinding and promotes adhesion of developing motor axons to the intersegmental nerve. Here we show by immunohistochemical staining that Eve is also expressed in the nuclei of a small set of neurons in the thoracic nervous system of larval and adult locusts, suggesting a postembryonic role in transcriptional control. Thirty-two to 39 Eve-positive neurons in four distinct subsets were found in the mesothoracic ganglion of fourth instar larvae and in adults: an anterior, bilaterally paired group of 2-4 neurons; a posterior, bilaterally paired group of 12-15 neurons; 1-2 neurons in a midline position, at the centre of the ganglion; and 1-2 neurons near the midline in a posterior position. These subsets did not change in position or number of neurons within them from fourth instars through to adults, 2 and 35 days post-final moult. The metathoracic ganglion had approximately 48 Eve-positive neurons in four subsets: an anterior, bilaterally paired group of 3-4 neurons; a posterior, bilaterally paired group of 8-10 neurons; a diffuse group of 19 neurons at the midline and a group of 4 neurons in a midline, posterior position. These results suggest that Eve demonstrates both phasic and persistent expression throughout the insect's lifetime and that changes in expression are segment-specific. We suggest Eve helps maintain neural phenotype in the adult CNS, possibly by regulating the expression of cell adhesion molecules.

Animals↗
PubMed2003

Proprioceptors monitoring forces in a locust hind leg during kicking form negative feedback loops with flexor tibiae motor neurons.

In preparation for jumping and kicking, a locust slowly generates large forces in the femoral muscles of its hind legs and stores them in elastic distortions of the tendons and femoral cuticle. At the femoro-tibial joints, the semi-lunar processes are bent, the cuticle of the dorsal distal femur is crumpled, and the femur is expanded in a mediolateral direction. We have analysed whether these distortions are monitored by sense organs and whether the information they provide is used to limit the forces generated and thus prevent structural damage to the joint. The two sensory neurons comprising the lump receptor lie in a groove in the ventral part of the distal femur. The sensory neurons spike if force is applied to the flexor tendon when the joint is fully flexed, but not when it is extended. They also spike as the tendon of the flexor muscle slides into the ventral femoral groove when the tibia is fully flexed during the co-contraction phase of kicking. Their spike frequency correlates with the extent of bending of a semi-lunar process that provides a quantifiable measure of the joint distortions. If the tibia is not fully flexed, however, then muscle contractions still cause distortions of the joint but these are not signalled by sensory spikes from the lump receptor. The lump receptor, therefore, does not respond primarily to the joint distortions but to the movements or force in the flexor tendon. Contractions of the flexor tibiae muscle caused by spikes in individual flexor motor neurons can evoke spikes in sensory neurons from the lump receptor when the joint is fully flexed. In turn, the sensory neurons cause a hyperpolarisation in particular flexor motor neurons in a polysynaptic negative feedback loop. The lump receptor could, therefore, regulate the output of the flexor motor neurons and, thus, limit the amount of force generated during co-contraction. It may also contribute to the inhibition of the flexors at the end of co-contraction that allows rapid kicking movements to occur.

Animals↗
PubMed2003

Jumping and kicking in bush crickets.

Bush crickets have long, thin hind legs but jump and kick rapidly. The mechanisms underlying these fast movements were analysed by correlating the activity of femoral muscles in a hind leg with the movements of the legs and body captured in high-speed images. A female Pholidoptera griseoaptera weighing 600 mg can jump a horizontal distance of 300 mm from a takeoff angle of 34 degrees and at a velocity of 2.1 m s(-1), gaining 1350 microJ of kinetic energy. The body is accelerated at up to 114 m s(-2), and the tibiae of the hind legs extend fully within 30 ms at maximal rotational velocities of 13500 deg. s(-1). Such performance requires a minimal power output of 40 mW. Ruddering movements of the hind legs may contribute to the stability of the body once the insect is airborne. During kicking, a hind tibia is extended completely within 10 ms with rotational velocities three times higher at 41800 deg. s(-1). Before a kick, high-speed images show no distortions of the hind femoro-tibial joints or of the small semi-lunar groove in the distal femur. Both kicks and jumps can be generated without full flexion of the hind tibiae. Some kicks involve a brief, 40-90 ms, period of co-contraction between the extensor and flexor tibiae muscles, but others can be generated by contraction of the extensor without a preceding co-contraction with the flexor. In the latter kicks, the initial flexion of the tibia is generated by a burst of flexor spikes, which then stop before spikes occur in the fast extensor tibiae (FETi) motor neuron. The rapid extension of the tibia can follow directly upon these spikes or can be delayed by as long as 40 ms. The velocity of tibial movement is positively correlated with the number of FETi spikes. The hind legs are 1.5 times longer than the body and more than four times longer than the front legs. The mechanical advantage of the hind leg flexor muscle over the extensor is greater at flexed joint angles and is enhanced by a pad of tissue on its tendon that slides over a protuberance in the ventral wall of the distal femur. The balance of forces in the extensor and flexor muscles, coupled with their changing lever ratio at different joint positions, appears to determine the point of tibial release and to enable rapid movements without an obligatory co-contraction of the two muscles.

Animals↗
PubMed2003

Mechanosensory-induced behavioural gregarization in the desert locust Schistocerca gregaria.

Desert locusts show an extreme form of phenotypic plasticity, changing between a cryptic solitarious phase and a swarming gregarious phase that differ in many aspects of behaviour, physiology and appearance. Solitarious locusts show rapid behavioural phase change in response to tactile stimulation directed to the hind femora. Repeatedly touching as little as one quarter of the anterior (outer) surface area of a hind femur produced full behavioural gregarization within 4 h. Solitarious locusts have approximately 30% more mechanosensory trichoid sensilla on the hind femora than do gregarious locusts but have similar or fewer numbers of sensilla elsewhere on the legs. Tactile stimulation of a hind femur in solitarious locusts that had been restrained so that they could not move their legs failed to induce any behavioural gregarization. Patterned electrical stimulation of metathoracic nerve 5, which innervates the hind leg, however, produced full gregarization in restrained locusts. Our data show for the first time that the gregarizing signal combines both exteroceptive and proprioceptive components, which travel in both nerves 5B1 and 5B2, and provides us with a powerful experimental method with which to elicit and study neuronal plasticity in this system. Acetic acid odour, a strong chemosensory stimulus that activates the same local processing pathways as exteroceptive stimuli, failed to elicit behavioural gregarization, suggesting an early segregation in the central nervous system of the mechanosensory signals that leads to gregarization.

Acetic Acid↗
PubMed2002

Jumping and kicking in the false stick insect Prosarthria teretrirostris: kinematics and motor control.

The false stick insect Prosarthria teretrirostris looks and behaves like a real stick insect but can jump and kick rapidly and powerfully like a locust, to which it is more closely related. It has an elongated body with slender hind legs that are some 2.5 times longer than the front and middle legs. A male with a body 67 mm long and weighing 0.28 g can jump 90 cm with a take-off angle of 40 degrees and velocity of 2.5 ms(-1), requiring an energy expenditure of 850 microJ. The body is accelerated at 165 ms(-2) for only 30 ms. The larger and heavier females (mean body length 104 mm and weighing 1.5 g) can jump on average a distance of 49 cm. During jumping, the tibiae of the hind legs are extended in 30 ms with maximum rotational velocities of 11.5 degrees per ms, but during kicking, when there is no body weight to support, extension is complete in 7 ms with rotational velocities as high as 48 degrees per ms. The short time available to accelerate the body indicates that the movements are not powered by direct muscle contractions and that there must be storage of elastic energy in advance. The motor patterns responsible for generating the necessary forces in the hind legs for jumping and kicking are similar and consist of three phases; an initial flexion of the tibia is followed by a co-contraction of the small flexor and large extensor tibiae muscles lasting several hundred milliseconds while the tibia remains fully flexed. Finally, the flexor motor neurons stop spiking so that the tibia is able to extend rapidly. The small semi-lunar processes at the femoro-tibial joints are not distorted, so that they cannot act as energy stores. Some 7% of the energy is stored transiently by bending the thin tibiae during the initial acceleration phase of a jump and releasing it just before take-off. The jumping and kicking mechanisms of Prosarthria teretrirostris have features in common with those used by locusts but also have their own characteristics. The evolution of jumping in Orthoptera is discussed in this context.

Animals↗
PubMed2002

Jumping in a winged stick insect.

The Thailand winged stick insect (Sipyloidea sp.) flees rapidly from a disturbance by jumping forwards when stimulated on the abdomen and backwards when stimulated on the head. The mechanisms underlying these fast movements were analysed by measuring movements of the body and legs from images captured at 250 Hz. A forward jump of both adults and nymphs involves movements of the abdomen and the middle and hind pairs of legs. The abdomen is raised and swung forwards by flexion at the joint with the metathorax and at the joint between the meso- and metathorax. At the same time, the tibiae of the hind and middle legs are extended and their femora depressed. The femoro-tibial joints of the legs are not fully flexed before a jump, and no structures in these joints appear to store muscular energy. The whole jumping sequence takes approximately 100 ms and results in take-off angles of 10-35 degrees at velocities of 0.6-0.8 m s(-1) and with an acceleration of 10 m s(-2). The abdominal angular velocity was 2000 degrees s(-1) and the tip of the abdomen moved at linear velocities of some 1 m s(-1), while the maximum rate of tibial extension was 4000 degrees s(-1). Rapid backward movements result either in the collapse of the body onto the ground, with a displacement away from the stimulus of approximately half a body length, or in the propulsion of the insect off its perch. Neither movement involves curling of the abdomen. From a horizontal posture, the forward jumps result in a displacement of a few body lengths. More lift can be generated in adults by elevating the hind wings as the abdomen is swung forwards and depressing them as the legs lose contact with the ground. In this way, jumps can lead directly to flapping flight. Take-off into flight can, however, be achieved without the abdominal movements seen during jumping. From a vertical posture, a forward jump propels the insect upwards and backwards before it falls to the ground horizontally displaced from its perch. Backward movements result in the insect falling with little horizontal displacement from its perch.

Animals↗