Variations on a segmental theme: muscle receptor organs and extensor neuromusculature in the squat lobster Munida quadrispina (Anomura, Galatheidae)

Extensor neuromusculature and the muscle receptor organs (MROs) associated with them have been conserved during the evolution of malacostracan crustaceans, despite species-specific differences between homologous segments in divergent taxa. Investigations of these differences could provide insight into how sensory and neuromuscular elements are modified to accommodate changing behavioural patterns. The most obvious differences between squat lobsters (galatheid anomurans) and macruran decapods, such as crayfish, are the greater dorso-ventral flattening of the galatheid abdomen and its flexed resting posture. To investigate whether the evolution of this altered posture affected extensor neuromusculature and MRO morphology and physiology, we used Methylene Blue staining, cobalt backfilling and extracellular recording techniques to describe these elements in the caudal thoracic and six abdominal segments of the squat lobster Munida quadrispina and compared our results with published descriptions of homologous elements in macrurans. In M. quadrispina, there is segmental variation both in the orientation of the MROs along the abdomen and in their physiological responses to stretch: apparent sensitivity is higher in caudal than rostral MROs. Homologues of three of the four accessory neurones found in crayfish occur, but AN#1 has a major dendrite not present in crayfish. Intersegmental differences in size and morphology of extensor motoneurones occur in M. quadrispina, as have been reported in crayfish, but are dissimilar in the two: abdominal ganglion 5 extensor motoneurones are the largest in M. quadrispina and the smallest in crayfish; this difference correlates with the difference in relative size of axial muscles along the abdomen reported previously for these species. M. quadrispina also differs from macrurans in having a single tonic, and no phasic, MRO on each side of the last abdominal segment. Together, these observations suggest that galatheids have evolved modified or additional neurobehavioural control(s) for the abdomen and tailfan.

FUNCTIONAL SPECIALIZATION OF THE SCOLOPARIA OF THE FEMORAL CHORDOTONAL ORGAN IN STICK INSECTS

The femoral chordotonal organ (fCO), one of the largest proprioceptive sense organs in the leg of the stick insect, is important for the control of the femur-tibia joint during standing and walking. It consists of a ventral scoloparium with about 80 sensory cells and a dorsal scoloparium with about 420 sensory cells. The present study examines the function of these scoloparia in the femur-tibia control loop. Both scoloparia were stimulated independently and the responses in the extensor tibiae motoneurones were recorded extra- and intracellularly. The ventral scoloparium, which is the smaller of the two, functions as the transducer of the femur-tibia control loop. Its sensory cells can generate the known resistance reflexes. The dorsal scoloparium serves no function in the femur-tibia control loop and its stimulation elicited no or only minor reactions in the extensor motoneurones. A comparison with other insect leg proprioceptors shows that a morphological subdivision of these organs often indicates a functional specialization.

Nonspiking Pathways in a Joint-control Loop of the Stick Insect Carausius Morosus

In the stick insect Carausius morosus (Phasmida) intracellular recordings were made from local nonspiking interneurones involved in the reflex activation of the extensor motoneurones of the femur-tibia joint during ramp-like stimulation of the transducer of this joint, the femoral chordotonal organ (ChO). The nonspiking interneurones in the femur-tibia control loop were characterized by their inputs from the ChO, their output properties onto the extensor motoneurones and their morphology. Eight different morphological and physiological types of nonspiking interneurones are described that are involved in the femur-tibia control loop. The results show that velocity signals from the ChO are the most important movement parameter processed by the nonspiking interneurones. Altering the membrane potential of these interneurones had marked effects on the reflex activation in the extensor motoneurones as the interneurones were able to increase or decrease the response of the participating motoneurones. The processing of information by the nonspiking pathways showed another remarkable aspect: nonspiking interneurones were found to process sensory information from the ChO onto extensor motoneurones in a way that seems not always to support the generation of the visible resistance reflexes in the extensor tibiae motoneurones in response to imposed flexion and extension movements of the joint. The present investigation demonstrated interneuronal pathways in the joint-control loop that show ‘assisting’ characteristics.

Intracellular Recordings from Interneurones and Motoneurones During Bilateral Kicks in the Locust: Implications for Mechanisms Controlling the Jump

Intracellular recordings were made from the neuropile processes of thoracic neurones of Locusta migratoria during bilateral kicks of the hindlegs. Electromyographic (EMG) recordings showed that the pattern of flexor and extensor tibiae muscle activity during kicks in this extensively dissected preparation was similar to that seen during a jump. Intracellular recordings from hindleg flexor and extensor motoneurones and from 13 identified interneurones revealed additional features of the motor programme for jumping and kicking and of the mechanism which triggers these events. There was a discrete burst of activity in the fast extensor tibiae (FETi) motoneurone at the end of the co-contraction phase, generated by a system that appeared to be separate from that triggering the kick. The excitatory connection from FETi to flexors was not responsible for initiating flexor activity and was of little functional importance in maintaining this activity during the co-contraction phase. The initial flexor excitation came from another, unidentified, central source. A pair of identified interneurones, the M-neurones, discharged with a high frequency burst just prior to the kick. Since these neurones inhibit hindleg flexor tibiae motoneurones, this observation provides further support for their proposed role as the neurones responsible for triggering kicks and jumps. Our data do not support the proposal that the activation of the M-neurones depends on them receiving progressively increasing proprioceptive input during the co-contraction phase. Throughout co-contraction, the M-neurones were hyperpolarized. Their activation was rapid and strong enough to cause them to discharge at rates as high as 400 spikes s−1. We suggest that the pulse-like activation of the M-neurones is produced centrally by a higher order system of interneurones. Another pair of previously identified interneurones, the C-neurones, were not necessary for the generation of the co-contraction phase of the motor programme. Their pattern of activity and their known connections indicated that they provide additional excitation to the flexors and extensors towards the end of co-contraction. Many other interneurones discharged either during co-contraction or when a kick was triggered. We conclude that the system generating the motor programme for a kick (jump) is more complex than proposed in previous studies.

Further Studies of Crayfish Escape Behaviour: II. Giant Axon-Mediated Neural Activity in the Appendages

Stereotyped responses were evoked in a number of motoneurones in the appendages of semi-intact crayfish when the command neurones for escape behaviour were activated. The medial giant neurones mediated short latency responses in pereiopod common inhibitor, promotor and extensor motoneurones, several abdominal first root neurones and one uropod exopodite promotor motoneurone. The lateral giant neurones mediated short latency responses in the pereiopod common inhibitor neurones, the same abdominal first root neurones and one uropod protopodite promotor motoneurone. These responses can be correlated with stereotyped movements of the appendages which occur in the normal escape behaviour of crayfish. Note: