Dynamics and stability of directional jumps in the desert locust

Locusts are known for their ability to jump large distances to avoid predation. The jump also serves to launch the adult locust into the air in order to initiate flight. Various aspects of this important behavior have been studied extensively, from muscle physiology and biomechanics, to the energy storage systems involved in powering the jump, and more. Less well understood are the mechanisms participating in control of the jump trajectory. Here we utilise video monitoring and careful analysis of experimental directional jumps by adult desert locusts, together with dynamic computer simulation, in order to understand how the locusts control the direction and elevation of the jump, the residual angular velocities resulting from the jump and the timing of flapping-flight initiation. Our study confirms and expands early findings regarding the instrumental role of the initial body position and orientation. Both real-jump video analysis and simulations based on our expanded dynamical model demonstrate that the initial body coordinates of position (relative to the hind-legs ground-contact points) are dominant in predicting the jumps’ azimuth and elevation angles. We also report a strong linear correlation between the jumps’ pitch-angular-velocity and flight initiation timing, such that head downwards rotations lead to earlier wing opening. In addition to offering important insights into the bio-mechanical principles of locust jumping and flight initiation, the findings from this study will be used in designing future prototypes of a bio-inspired miniature jumping robot that will be employed in animal behaviour studies and environmental monitoring applications.

Programmed cell death specifically eliminates one part of a locust pleuroaxillary muscle after the imaginal moult

In the hemimetabolous insect Locusta migratoria, fundamental restructuring occurs at the transition from flightless nymph to flight-capable adult. This transition involves all components of the flight circuit, which is present but not used for flight in nymphs. The meso- and metathoracic pleuroaxillary muscles, M85 and M114 respectively, constitute one component of this circuit. In the adult locust, these are flight-steering muscles, but their function in the nymph is as yet unknown. Our study reveals that adult and nymphal metathoracic pleuroaxillary muscles M114 differ profoundly. The nymphal muscle contains the distinct part M114c in addition to parts M114a and M114b characteristic of the adult. The contractions of M114c are slow and long-lasting, corresponding to its long sarcomeres and slow form of ATPase, and contrast with the adult muscle parts M114a and M114b in all of these features. We demonstrate a hormone-dependent degeneration of M114c after the adult moult. This degeneration can be blocked by actinomycin D and cycloheximide. It may thus be termed genetically programmed cell death, triggered after the adult moult and, as demonstrated here, functioning via the ATP-dependent ubiquitin pathway. Given the defined onset of degeneration after the adult moult, it is possible that M114c may fulfil a specific function in nymphs, during or shortly after moulting.

Plasticity of Synaptic Connections in Sensory-Motor Pathways of the Adult Locust Flight System

Wolf, Harald and Ansgar Büschges. Plasticity of synaptic connections in sensory-motor pathways of the adult locust flight system. J. Neurophysiol. 78: 1276–1284, 1997. We investigated possible roles of retrograde signals and competitive interactions in the lesion-induced reorganization of synaptic contacts in the locust CNS. Neuronal plasticity is elicited in the adult flight system by removal of afferents from the tegula, a mechanoreceptor organ at the base of the wing. We severed one hindwing organ and studied the resulting rearrangement of synaptic contacts between flight interneurons and afferent neurons from the remaining three tegulae (2 forewing, 1 hindwing). This was done by electric stimulation of afferents and intracellular recording from interneurons (and occasionally motoneurons). Two to three weeks after unilateral tegula lesion, connections between tegula afferents and flight interneurons were altered in the following way. 1) Axons from the forewing tegula on the operated side had established new synaptic contacts with metathoracic elevator interneurons. In addition, the amplitude of compound excitatory postsynaptic potentials elicited by electric stimulation was increased, indicating that a larger number of afferents connected to any given interneuron. 2) On the side contralateral to the lesion, connectivity between axons from the forewing tegula and elevator interneurons was decreased. 3) The efficacy of the (remaining) hindwing afferents appeared to be increased with regard to both synaptic transmission to interneurons and impact on flight motor pattern. 4) Flight motoneurons, which are normally restricted to the ipsilateral hemiganglion, sprouted across the ganglion midline after unilateral tegula removal and apparently established new synaptic contacts with tegula afferents on that side. The changes on the operated side are interpreted as occupation of synaptic space vacated on the interneurons by the severed hindwing afferents. On the contralateral side, the changes in synaptic contact must be elicited by retrograde signals from bilaterally arborizing flight interneurons, because tegula projections remain strictly ipsilateral. The pattern of changes suggests competitive interactions between forewing and hindwing afferents. The present investigation thus presents evidence that the CNS of the mature locust is capable of extensive synaptic rearrangement in response to injury and indicates for the first time the action of retrograde signals from interneurons.

Recovery of the flight system following ablation of the tegulae in immature adult locusts

The capacity of the flight system to recover from ablation of the tegulae was studied in immature adult Locusta migratoria and compared with recovery in mature adults. We ablated the hindwing tegulae or all tegulae in adult locusts either 1 day after the imaginal moult (immature locusts) or 2 weeks after the imaginal moult (mature locusts). We monitored recovery throughout the recovery period by using a stroboscope to measure the wingbeat frequency of tethered locusts. In addition, we measured other parameters of the flight motor pattern using electromyographic electrodes implanted into recovered locusts. Both methods of monitoring recovery yielded the same results. There was no reduction, during adult maturation, in the capacity of the locust flight system to recover from the loss of these proprioceptors. Plasticity of the locust flight system was therefore maintained in the mature adult locust. This suggests that the flight system is not fixed and simply implemented when the locust reaches adulthood, but that the circuitry can be remodelled throughout the animal's life to produce behaviour adapted to the needs and constraints of the individual.