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Dirk Bucher

Publications and source records attributed to Dirk Bucher.

10 recordsLinked to original sources

Neuronal morphology and neuropil structure in the stomatogastric ganglion of the lobster, Homarus americanus.

The stomatogastric nervous system (STNS) has long been used as a model system for the study of central pattern generation, neuromodulation, and network dynamics. Anatomical studies of the crustacean stomatogastric ganglion (STG) in different species have mostly been restricted to subsets of neurons and/or general structural features. For the first time, we describe the morphology of all STG neurons belonging to the two circuits that produce the well-described pyloric and gastric rhythms in the lobster, Homarus americanus. Somata sit on the dorsal and lateral surface of the STG and send a single primary neurite into the core of the neuropil, which is mostly made up of larger lower order branches. The perimeter of the neuropil consists mostly of finer higher order branches. Immunohistochemical labeling for synaptic proteins is associated with the small diameter branches. Somata positions are not constant but show preferred locations across individuals. The number of copies is constant for all neuron types except the PY and GM neurons (PY neuron number ranges from 3 to 7, and GM neuron number ranges from 6 to 9). Branch structure is largely nondichotomous, and branches can deviate substantially from cylindrical shape. Diameter changes at branch points can be as large as 20-fold. Clearly, the morphology of a specific neuron type can be quite variable from animal to animal.

Animals↗

Understanding circuit dynamics using the stomatogastric nervous system of lobsters and crabs.

Studies of the stomatogastric nervous systems of lobsters and crabs have led to numerous insights into the cellular and circuit mechanisms that generate rhythmic motor patterns. The small number of easily identifiable neurons allowed the establishment of connectivity diagrams among the neurons of the stomatogastric ganglion. We now know that (a) neuromodulatory substances reconfigure circuit dynamics by altering synaptic strength and voltage-dependent conductances and (b) individual neurons can switch among different functional circuits. Computational and experimental studies of single-neuron and network homeostatic regulation have provided insight into compensatory mechanisms that can underlie stable network performance. Many of the observations first made using the stomatogastric nervous system can be generalized to other invertebrate and vertebrate circuits.

Animals↗

Central pattern generating neurons simultaneously express fast and slow rhythmic activities in the stomatogastric ganglion.

Neuronal firing patterns can contain different temporal information. It has long been known that the fast pyloric and the slower gastric motor patterns in the stomatogastric ganglion of decapod crustaceans interact. However, the bidirectional influences between the pyloric rhythm and the gastric mill rhythm have not been quantified in detail from preparations that spontaneously express both patterns in vitro. We found regular and stable spontaneous gastric and pyloric activity in 71% of preparations of the isolated stomatogastric nervous system of the lobster, Homarus americanus. The gastric [cycle period: 10.96 +/- 2.67 (SD) s] and pyloric (cycle period: 1.35 +/- 0.18 s) patterns showed bidirectional interactions and coordination. Gastric neuron firing showed preferred phases within the reference frame of the pyloric cycle. The relative timing and burst parameters of the pyloric neurons systematically changed within the reference frame of the gastric cycle. The gastric rhythm showed a tendency to run at cycle periods that were integer multiples of the pyloric periods, but coupling and coordination between the two rhythms were variable. We used power spectra to quantify the gastric and pyloric contributions to the firing pattern of each individual neuron. This provided us with a way to analyze the firing pattern of each gastric and pyloric neuron type individually without reference to either gastric or pyloric phase. Possible functional consequences of these network interactions for motor output are discussed.

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Invertebrate central pattern generation moves along.

Central pattern generators (CPGs) are circuits that generate organized and repetitive motor patterns, such as those underlying feeding, locomotion and respiration. We summarize recent work on invertebrate CPGs which has provided new insights into how rhythmic motor patterns are produced and how they are controlled by higher-order command and modulatory interneurons.

Animals↗

Animal-to-animal variability in motor pattern production in adults and during growth.

Which features of network output are well preserved during growth of the nervous system and across different preparations of the same size? To address this issue, we characterized the pyloric rhythms generated by the stomatogastric nervous systems of 99 adult and 12 juvenile lobsters (Homarus americanus). Anatomical studies of single pyloric network neurons and of the whole stomatogastric ganglion (STG) showed that the STG and its neurons grow considerably from juvenile to adult. Despite these changes in size, intracellularly recorded membrane potential waveforms of pyloric network neurons and the phase relationships in the pyloric rhythm were very similar between juvenile and adult preparations. Across adult preparations, the cycle period and number of spikes per burst were not tightly maintained, but the mean phase relationships were independent of the period of the rhythm and relatively tightly maintained across preparations. We interpret this as evidence for homeostatic regulation of network activity.

Animals↗

Constant amplitude of postsynaptic responses for single presynaptic action potentials but not bursting input during growth of an identified neuromuscular junction in the lobster, Homarus americanus.

As lobsters grow from early juveniles to adults their body size increases more than 20-fold, raising the question of how function is maintained during these ongoing changes in size. To address this question we studied the pyloric 1 (p1) muscle of the stomach of the lobster, Homarus americanus. The p1 muscle receives multiterminal innervation from one motor neuron, the lateral pyloric neuron of the stomatogastric ganglion. Staining with antibodies raised against synaptotagmin showed that as the muscle fibers increased in length, the spacing between the terminal innervation increased proportionally, so the number of synaptic contact regions/muscle fiber did not change. Muscle fibers were electrically coupled in both juveniles and adults. The amplitude of single intracellularly recorded excitatory junctional potentials evoked by motor nerve stimulation was the same in both juveniles and adults. Nonetheless, the peak depolarizations reached in response to ongoing pyloric rhythm activity or in response to high-frequency trains of stimuli similar to those produced during the pyloric rhythm were approximately twofold larger in juveniles than in adults. This suggests that homeostatic regulation of synaptic connections may operate at the level of the amplitude of the single synaptic potential rather than on the summed depolarization evoked during strong rhythmic activity.

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Similar network activity from disparate circuit parameters.

It is often assumed that cellular and synaptic properties need to be regulated to specific values to allow a neuronal network to function properly. To determine how tightly neuronal properties and synaptic strengths need to be tuned to produce a given network output, we simulated more than 20 million versions of a three-cell model of the pyloric network of the crustacean stomatogastric ganglion using different combinations of synapse strengths and neuron properties. We found that virtually indistinguishable network activity can arise from widely disparate sets of underlying mechanisms, suggesting that there could be considerable animal-to-animal variability in many of the parameters that control network activity, and that many different combinations of synaptic strengths and intrinsic membrane properties can be consistent with appropriate network performance.

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Synaptic drive contributing to rhythmic activation of motoneurons in the deafferented stick insect walking system.

A general feature of motor patterns for locomotion is their cyclic and alternating organization. In walking, for example, rhythmic activity in leg motoneurons innervating antagonistic muscles of a joint is primarily antiphasic within each cycle. We investigate which role central pattern generating networks play in the generation of leg motoneuron activity in the absence of sensory feedback. We elicited activity in antagonistic flexor and extensor tibiae motoneurons in the deafferented mesothoracic ganglion of the stick insect by mechanically stimulating the head or abdomen, while recording intracellularly from their neuropilar processes. In most cases, tactile stimulation induced coactivation of tibial motoneurons. However, in approximately 25% of the trials, tibial motoneurons generated alternating cycles consisting of bursts of action potentials that were terminated by strong inhibitory synaptic inputs. Injection of depolarizing current increased the amplitude of the inhibitory phase of the oscillation, while hyperpolarizing current decreased it and revealed a tonic depolarization of the motor neurons during the bout of rhythmic motor activity. The same results were gathered from recording tibial leg motoneurons during 'twitching' motor activity in decerebrated animals. Our results indicate that alternating rhythmic motoneuron activity in the deafferented stick insect walking system results from phasic inhibitory drive provided by central pattern generating networks. This inhibitory input patterns the firing of the motoneurons that results from a tonic depolarizing drive. This tonic depolarizing drive was also observed in tibial motoneurons of the deafferented mesothoracic ganglion during walking movements of the intact ipsilateral front leg.

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Axonal dopamine receptors activate peripheral spike initiation in a stomatogastric motor neuron.

We studied the effects of dopamine on the stomatogastric ganglion (STG) of the lobster, Homarus americanus. The two pyloric dilator (PD) neurons are active in the pyloric rhythm, have somata in the STG, and send axons many centimeters to innervate muscles of the stomach. Dopamine application to the stomatogastric nervous system when the PD neurons were rhythmically active evoked additional action potentials during the PD neuron interburst intervals. These action potentials were peripherally generated at a region between the STG and the first bilateral branch, approximately 1 cm away from the STG, and traveled antidromically to the neuropil and orthodromically to the pyloric dilator muscles. Focal applications of dopamine to the nerves showed that spikes could be initiated in almost the entire peripheral axon of the PD neurons. Dopamine also evoked spikes in isolated peripheral axons. The concentration threshold for peripheral spike initiation was at or below 10-9 m dopamine. Thus, the peripheral axon can play an important role in shaping the output signaling to the muscles by the motor neuron.

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Interjoint coordination in the stick insect leg-control system: the role of positional signaling.

Interjoint coordination is essential for proper walking behavior in multi-jointed insect legs. We have shown previously that movement signals from the femur-tibia (FT) joint can shape motor activity of the adjacent coxa-trochanter (CT) joint in the stick insect, Carausius morosus. Here, we present data on the role of position signals from the FT-joint on activity generated in motoneurons (MNs) of the CT-joint. We show that the probability of occurrence of stance (with depression in the CT-joint) or swing movements (with levation in the CT-joint) at the start of walking sequences is influenced by the angle of the FT-joint in the resting animal. We tested the influence of FT-joint angle on pharmacologically induced rhythmic activity of CT-joint depressor (DprTr) and levator (LevTr) MNs. The burst duration, mean spike rate within bursts, and duty cycle for each MN pool were found to depend on FT position. For LevTr MNs, these parameters progressively increased as the FT-joint was moved from extension to flexion, and the opposite was true for DprTr MNs. The cycle period of CT-MN rhythmicity also depended on FT position. In addition, we sometimes observed that the motor output shifted completely to one MN pool at extreme positions, suggesting that the central rhythm-generating network for the CT-joint became locked in one phase. These results indicate that position signals from the FT-joint modulate rhythmic activity in CT-joint MNs partly by having access to central rhythm generating networks of the CT-joint.

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