Search PubMed⌕ Search

Biomedical subjects

M Moulins

Publications and source records attributed to M Moulins.

At least 37 records · Page 2Linked to original sources

A disynaptic sensorimotor pathway in the lobster stomatogastric system.

1. In the lobster Homarus, muscle gm 1 that causes protraction of the medial tooth of the gastric mill system is innervated via a dorsal branch of the anterior gastric nerve by motoneurons (GM) arising in the stomatogastric ganglion (STG) (Fig. 1). 2. A ventral branch of the anterior gastric nerve (VAGN) contains a single unit that is mechanosensitive, responds to gentle pressure on the stomach wall in the vicinity of gm 1, and evokes reflex activation of GM motoneurons (Fig. 2). 3. This mechanoreceptor neuron (called anterior gastric receptor, AGR) has been identified morphologically (Fig. 3) and electrophysiologically (Figs. 4 and 5). The bipolar cell body is located in the dorsal ventricular nerve immediately posterior to the STG. It sends out long peripheral processes in the left and right VAGNs to ramify bilaterally in the epidermis of the stomach wall underlying muscle gm 1. The axon of the AGR runs anteriorly through the STG and projects to the left and right commissural ganglia (CoGs) via the stomatogastric (STN) and inferior esophageal nerves. 4. AGR activation of GM motoneurons disappears after cutting the STN, indicating that the reflex is mediated by an axonal pathway involving rostral ganglia (Fig. 6). 5. Electrophysiological (Fig. 7) and morphological (Fig. 8) methods were used to identify an interneuron (commissural gastric neuron, CG) located in each CoG and intercalated between AGR and GM. Axons of the two CGs project to the STG via the superior esophageal nerves and the STN. 6. Simultaneous intracellular recordings from the three cell types demonstrate that AGR excites CG, which in turn excites GM; in each case excitatory postsynaptic potentials follow presynaptic impulses one for one and at constant latency (Fig. 9). Raising the threshold for spiking with saline containing high divalent cation concentrations further indicates that both excitatory connections are monosynaptic and confirms that AGR does not directly excite GM motoneurons (Fig. 10). 7. The input/output properties of AGR in this disynaptic excitatory pathway (Fig. 11) are discussed as also are the functional implications of such a long-loop pathway for sensorimotor integration.

Afferent Pathways↗

Nonlinear interneuronal properties underlie integrative flexibility in a lobster disynaptic sensorimotor pathway.

1. In the lobster Homarus, a single mechanoreceptor neuron (anterior gastric receptor, AGR) associated with muscle gm 1 of the gastric medial tooth has access to motoneurons (GM) innervating this muscle via an excitatory synaptic pathway involving two bilateral interneurons (commissural gastric, CG) (see 31). 2. Studies on in vitro preparations of the stomatogastric nervous system show that despite its apparent simplicity, this disynaptic pathway can express considerable flexibility in information processing, as evident by a wide variety of GM output responses to sensory input from AGR (Fig. 1). 3. This input/output flexibility does not rely on multiple synaptic pathways operating in parallel with the interneuron CG, since it is demonstrated that AGR has access to GM only via CG (Fig. 2). 4. Short AGR impulse trains at different spike frequencies can give rise to similarly brief excitation of GM, or prolonged motoneuron responses. Moreover, graded increases in AGR discharge frequency can lead to a sudden increase in the intensity of GM responsiveness that otherwise grades linearly with receptor firing. Such step changes in gain (both in duration and magnitude) are due to synaptic triggering of regenerative "plateau" depolarizations in CG (Figs. 3 and 4). 5. Sustained tonic discharge in AGR can induce cyclic bursting activity in previously nonrhythmic GM neurons. Furthermore, the frequency of motoneuron bursts increases with the frequency of AGR tonic firing. Such changes in pattern are ascribed to synaptic triggering and modification of regenerative "oscillatory" depolarizations in CG (Fig. 5). 6. Higher levels of AGR firing can result either in strong activation of GM motoneurons or in complete inactivation of GM. This switch in sign of the motor response is dependent on base-line levels of activity in the receptor and is due to the capability of CG to fire action potentials only within a window of membrane potential (Figs. 6-8). The functional outcome of this cellular property of CG is that positive feedback from AGR to GM can be switched to negative feedback via the same excitatory synaptic pathway (Fig. 9). 7. We conclude that flexibility in sensorimotor integration can be an inbuilt feature even of hard-wired neuronal pathways; in the present case, changes in input/output relationships reside with intrinsic properties of an intercalated interneuron (Fig. 10).

Action Potentials↗

Control by an identified modulatory neuron of the sequential expression of plateau properties of, and synaptic inputs to, a neuron in a central pattern generator.

Recordings from the lateral gastric (LG) neuron, which forms part of the gastric mill central pattern generator in the red lobster, Palinurus vulgaris, indicate that regenerative membrane properties (plateau properties) and synaptic inputs interact sequentially rather than simultaneously to determine its discharge pattern. LG thus presents a composite discharge, consisting of 2 separate segments of firing and one silent period. The first firing segment depends on regenerative membrane properties; this is the endogenous component, or segment, of LG's discharge. The second firing segment is the result of extrinsic synaptic input, forming the synaptic component of LG's discharge. The relative importance of these 2 components can vary, and thus LG's discharge ranges from one in which LG fires only as a result of its endogenous component to one in which its endogenous component is entirely absent and only the synaptic component underlies action potentials. Activity in an identified modulatory neuron suppresses the endogenous segment and enhances the synaptic segment of LG's discharge. This long-lasting effect in turn changes phase relationships within the gastric mill network and provides mechanisms for producing flexibility in the gastric pattern generator and for ensuring that a specific motor output is generated by a flexible neural network.

Action Potentials↗

Suppressive control of a rhythmic central pattern generator by an identified modulatory neuron in crustacea.

The activity of the 14 neuron network which organizes the pyloric motor rhythm in the stomatogastric ganglion of the lobster, Homarus gammarus, is controlled by neuromodulatory inputs which have been described as having mainly 'permissive' effects. By contrast, here we identify a neuron, the pyloric suppressor (PS) neuron which exerts a 'suppressive' effect on the pyloric activity. We show that PS neuron discharge can terminate in a long-lasting manner, spontaneous pyloric rhythmic activity. Its effect results from a direct suppression of the endogenous ability of the pyloric pacemaker neurons to produce rhythmic bursts of action potentials. Thus the output of the pyloric neuronal network appears to be finely tuned by neuromodulatory influences having opposite effects.

Action Potentials↗

Suppression of oscillatory activity in crustacean pyloric neurons: implication of GABAergic inputs.

Generation of rhythmic pyloric motor output in the crustacean stomatogastric ganglion results from synaptic connections and cellular properties of a 14-cell network of pyloric neurons. These cellular properties are under the influences of modulatory inputs, which act, for the most part, in an activating mode, i.e., they enhance the bursting properties of the pyloric neurons and/or their ability to express their regenerative properties. Here we attempt to demonstrate that the pyloric motor output is also under the control of suppressive afferent inputs that are able to stop the pyloric rhythm in a long-lasting manner. Immunohistochemistry, using GABA antibodies, indicates that GABAergic-like fibers are present in both the stomatogastric ganglion and its afferent nerve. Bath-applied GABA suppresses spontaneous pyloric rhythmic activity. This is due to an inability of the pyloric pacemakers to express their bursting properties. The suppressive effect of GABA is blocked by picrotoxin and mimicked by muscimol. Isolating the pyloric neurons from all descending spiking influences with tetrodotoxin demonstrates that exogenously applied GABA acts directly on the pyloric neurons. To confirm the existence of a physiological suppressive system for the pyloric motor pattern, we show that the stimulation of an afferent nerve, known to contain GABA-like fibers, also causes the cessation of rhythmic activity and the inability of the pyloric neurons to express their bursting properties.

Animals↗

Oscillatory neural networks.

Despite the fact that a large number of neuronal oscillators have been described, there are only a few good examples that illustrate how they operate at the cellular level. For most, there is some isolated information about different aspects of the oscillator network, but too little to explain the whole mechanism. Two quite remarkable features do seem to be emerging from ongoing studies, however. One is that there are very few generalizable features common to neural oscillators. Many utilize reciprocal inhibitory circuits and endogenous burst-generating currents to some extent. All that have been well worked out utilize a combination of both cellular and network properties, but little else in the way of common mechanism is noteworthy. Perhaps the most interesting aspect of recent work is the ability of a particular oscillator to produce a large repertoire of different outputs. This is separate and in addition to changes occurring via phasic sensory feedback. It is in fact a radical functional "rewiring" of the network in response to neuromodulators. The CPG circuits represent only the most basic form of a given pattern. Finally, concerning the role of sensory feedback in generating oscillatory patterns, the concept of the CPG as a group of neurons able to produce oscillatory patterns without any sensory feedback is, in our opinion, still valid. There is no doubt that some oscillators may be quite weak when isolated, but they can still produce bursts with firing sequences similar to those seen in vivo. The fact that sensory feedback can both control and enhance the oscillations has never been in doubt. Similarly, entrainment of the pattern by sensory feedback does not mean that the receptor is part of the generator, only that it has access to it (as do command and coordinating fibers). The real question remains: Can a group of cells produce an oscillatory pattern without phasic sensory input? We must answer this affirmatively even for the insect-flight motor CPG, while emphasizing the fact that for this system sensory feedback plays a larger role than in most other CPGs. Most neural oscillators will probably fall on some continuum between those like insect flight, which need and use a large amount of phasic feedback, and those that can oscillate in a near-normal manner without it.

Animals↗

All-or-none control of the bursting properties of the pacemaker neurons of the lobster pyloric pattern generator.

In Crustacea the central pattern generator for the pyloric motor rhythm (filtration to the midgut) is known to be located within the stomatogastric ganglion (STG); its cycling activity is known to be organized by three endogenous burster neurons acting as pacemakers and driving 11 follower neurons. In Homarus, recordings from the isolated stomatogastric nervous system (Fig. 1) indicate that (1) the pyloric output can be generated only when the STG is afferented (i.e., connected to the more rostral oesophageal and commissural ganglia) (Fig. 2) and (2) the deafferentation of the STG results in a complete loss of the bursting properties of the pacemaker neurons (Fig. 4). Manipulation of the STG inputs responsible for unmasking the properties of the pacemakers strongly suggests that (1) they are not phasic inputs (Fig. 5) and (2) they are long-term acting inputs (Fig. 6). These results provide evidence for a neural all-or-none control of the bursting properties of the pacemaker neurons of a motor pattern generator.

Afferent Pathways↗

Firing between two spike thresholds: implications for oscillating lobster interneurons.

An identified interneuron in the lobster commissural ganglia fires spikes only between membrane potential values of -60 and -30 millivolts. The membrane potential of this neuron can also oscillate, and interaction between these two properties has important implications in determining the firing pattern of the neuron itself and the modalities of driving of a distant postsynaptic neuron.

Animals↗

Modulatory effects of a single neurons on the activity of the pyloric pattern generator in Crustacea.

The firing of a single neuron (named anterior pyloric modulator: APM) of the esophageal ganglion considerably modifies the pyloric rhythm of the rock lobster. These modifications, characterized by a long delay to onset and a long duration, include increased frequency and amplitude of oscillations of the motor neurons, changes in the efficacy of certain synapses within the network, and voltage-dependent modifications of membrane properties of some motor neurons. APM thus seems to be a true modulatory neuron. The APM-provoked changes resemble changes seen in the whole animal, making this a suitable system for an analysis of modulation on several levels.

Animals↗

Control of rhythmic behaviour by a hierarchy of linked oscillators in crustacea.

In Homarus, the central pattern generators for the rhythmic motor activities of the gastric teeth and the pyloric chamber are located in the stomatogastric ganglion. It is shown that independent gastric and pyloric oscillators are also contained in higher nervous centres (the commissural ganglia) and provide a phasic rhythmic input to the stomatogastric pattern generators. This demonstrates that rhythmic behaviour can be organized by a hierarchy of linked oscillators each capable of producing the rhythm.

Animals↗

A corollary discharge of total foregut motor activity is monitored by a single interneurone in the lobster Homarus gammarus.

1. In Homarus, an identified interneurone (the L cell), which possesses the largest cell body in the commissural ganglion and projects to the brain, exhibits a complex firing pattern (Fig. 2 a). 2. It is shown that the L cell discharges with each of the 4 pattern generators of the stomatogastric nervous system which organize the rhythmic motor activity of the foregut (Fig. 2 b-e). 3. Manipulation of the membrane potential of the L cell does not induce any change in the 4 rhythms (Fig. 3), and it is concluded that the L cell is driven by the 4 pattern generators. 4. The functional meaning of this complex corollary discharge of the total foregut motor activity is discussed.

Animals↗