The transient potassium outward current has different roles in modulating the pyloric and gastric mill rhythms in the stomatogastric ganglion.

The crustacean stomatogastric nervous system is a classic model for understanding the effects of modulating ionic currents and synapses at both the cell and network levels. The stomatogastric ganglion in this system contains two distinct central pattern generators: a slow gastric mill network that generates flexible rhythmic outputs (8-20 s) and is often silent, and a fast pyloric network that generates more consistent rhythmic outputs (0.5-2 s) and is always active in vitro.

Role of Ih in differentiating the dynamics of the gastric and pyloric neurons in the stomatogastric ganglion of the lobster, Homarus americanus.

The hyperpolarization-activated inward cationic current (Ih) is known to regulate the rhythmicity, excitability, and synaptic transmission in heart cells and many types of neurons across a variety of species, including some pyloric and gastric mill neurons in the stomatogastric ganglion (STG) in Cancer borealis and Panulirus interruptus However, little is known about the role of Ih in regulating the gastric mill dynamics and its contribution to the dynamical bifurcation of the gastric mill and pyloric networks. We investigated the role of Ih in the rhythmic activity and cellular excitability of both the gastric mill neurons (medial gastric, gastric mill) and pyloric neurons (pyloric dilator, lateral pyloric) in Homarus americanus Through testing the burst period between 5 and 50 mM CsCl, and elimination of postinhibitory rebound and voltage sag, we found that 30 mM CsCl can sufficiently block Ih in both the pyloric and gastric mill neurons. Our results show that Ih maintains the excitability of both the pyloric and gastric mill neurons.

Invertebrate central pattern generator circuits.

There are now a reasonable number of invertebrate central pattern generator (CPG) circuits described in sufficient detail that a mechanistic explanation of how they work is possible. These small circuits represent the best-understood neural circuits with which to investigate how cell-to-cell synaptic connections and individual channel conductances combine to generate rhythmic and patterned output. In this review, some of the main lessons that have appeared from this analysis are discussed and concrete examples of circuits ranging from single phase to multiple phase patterns are described.

Neural mechanisms underlying the generation of the lobster gastric mill motor pattern.

The lobster gastric mill central pattern generator (CPG) is located in the stomatogastric ganglion and consists of 11 neurons whose circuitry is well known. Because all of the neurons are identifiable and accessible, it can serve as a prime experimental model for analyzing how microcircuits generate multiphase oscillatory spatiotemporal patterns. The neurons that comprise the gastric mill CPG consist of one interneuron, five burster neurons and six tonically firing neurons.

Robust microcircuit synchronization by inhibitory connections.

Microcircuits in different brain areas share similar architectural and biophysical properties with compact motor networks known as central pattern generators (CPGs). Consequently, CPGs have been suggested as valuable biological models for understanding of microcircuit dynamics and particularly, their synchronization. We use a well known compact motor network, the lobster pyloric CPG to study principles of intercircuit synchronization.

Probing the dynamics of identified neurons with a data-driven modeling approach.

In controlling animal behavior the nervous system has to perform within the operational limits set by the requirements of each specific behavior. The implications for the corresponding range of suitable network, single neuron, and ion channel properties have remained elusive. In this article we approach the question of how well-constrained properties of neuronal systems may be on the neuronal level.

Pacemaker and network mechanisms of rhythm generation: cooperation and competition.

The origin of rhythmic activity in brain circuits and CPG-like motor networks is still not fully understood. The main unsolved questions are (i) What are the respective roles of intrinsic bursting and network based dynamics in systems of coupled heterogeneous, intrinsically complex, even chaotic, neurons? (ii) What are the mechanisms underlying the coexistence of robustness and flexibility in the observed rhythmic spatio-temporal patterns? One common view is that particular bursting neurons provide the rhythmogenic component while the connections between different neurons are responsible for the regularisation and synchronisation of groups of neurons and for specific phase relationships in multi-phasic patterns. We have examined the spatio-temporal rhythmic patterns in computer-simulated motif networks of H-H neurons connected by slow inhibitory synapses with a non-symmetric pattern of coupling strengths.

Biomimetic robots and biology.

Using robots that operate in the real world as opposed to computer simulations of animal behavior is a form of modeling that may provide some biological insights. However, since engineering principles and materials differ significantly from those used in biology, one should be extremely cautious in interpreting robot biomimicry as providing an explanation of biological mechanisms.

Artificial synaptic modification reveals a dynamical invariant in the pyloric CPG.

The sequential firing of neurons in central pattern generators (CPGs) is generally thought to be a result of an interaction between intrinsic cellular and synaptic properties of the component neurons. Due to experimental limitations, it is usually difficult to address the role of each of these properties separately. We have done so by using the crustacean stomatogastric CPG and the dynamic clamp technique to measure how the network responds to the selective modification of an individual important synapse.

Models wagging the dog: are circuits constructed with disparate parameters?

In a recent article, Prinz, Bucher, and Marder (2004) addressed the fundamental question of whether neural systems are built with a fixed blueprint of tightly controlled parameters or in a way in which properties can vary largely from one individual to another, using a database modeling approach. Here, we examine the main conclusion that neural circuits indeed are built with largely varying parameters in the light of our own experimental and modeling observations. We critically discuss the experimental and theoretical evidence, including the general adequacy of database approaches for questions of this kind, and come to the conclusion that the last word for this fundamental question has not yet been spoken.

Oscillations and oscillatory behavior in small neural circuits.

In order to determine the dynamical properties of central pattern generators (CPGs), we have examined the lobster stomatogastric ganglion using the tools of nonlinear dynamics. The lobster pyloric and gastric mill central pattern generators can be analyzed at both the cellular and network levels because they are small, i.e.

Consistent dynamics suggests tight regulation of biophysical parameters in a small network of bursting neurons.

The neuronal firing patterns in the pyloric network of crustaceans are remarkably consistent among animals. Although this characteristic of the pyloric network is well-known, the biophysical mechanisms underlying the regulation of the systems output are receiving renewed attention. Computer simulations of the pyloric network recently demonstrated that consistent motor output can be achieved from neurons with disparate biophysical parameters among animals.

Mechanisms underlying type I mGluR-induced activation of lobster gastric mill neurons.

In addition to ionotropic effects, glutamate and acetylcholine have metabotropic modulatory effects on many neurons. Here we show that in the stomatogastric ganglion of the lobster, glutamate, one of the main ionotropic neurotransmitters, modulates the excitability of gastric mill neurons. The neurons in this well-studied system produce rhythmic output to a subset of lobster foregut muscles.

StdpC: a modern dynamic clamp.

With the advancement of computer technology many novel uses of dynamic clamp have become possible. We have added new features to our dynamic clamp software StdpC ("Spike timing-dependent plasticity Clamp") allowing such new applications while conserving the ease of use and installation of the popular earlier Dynclamp 2/4 package. Here, we introduce the new features of a waveform generator, freely programmable Hodgkin-Huxley conductances, learning synapses, graphic data displays, and a powerful scripting mechanism and discuss examples of experiments using these features.

The role of sensory network dynamics in generating a motor program.

Sensory input plays a major role in controlling motor responses during most behavioral tasks. The vestibular organs in the marine mollusk Clione, the statocysts, react to the external environment and continuously adjust the tail and wing motor neurons to keep the animal oriented vertically. However, we suggested previously that during hunting behavior, the intrinsic dynamics of the statocyst network produce a spatiotemporal pattern that may control the motor system independently of environmental cues.

A neural infrastructure for rhythmic motor patterns.

It is possible to work out the neural circuity of many invertebrate central pattern generators (CPGs) thereby providing a basis for linking cellular processes to actual behaviors. This review summarizes the infrastructure of the two CPGs in the lobster stomatogastric ganglion in terms of circuitry, ionic conductances and chemical modulation by amines and peptides. Analysis of the circuit using modeling techniques including the use of electronic neurons closes the chapter.

A network of electronic neural oscillators reproduces the dynamics of the periodically forced pyloric pacemaker group.

Low-dimensional oscillators are a valuable model for the neuronal activity of isolated neurons. When coupled, the self-sustained oscillations of individual free oscillators are replaced by a collective network dynamics. Here, dynamical features of such a network, consisting of three electronic implementations of the Hindmarsh-Rose mathematical model of bursting neurons, are compared to those of a biological neural motor system, specifically the pyloric CPG of the crustacean stomatogastric nervous system.

Dopamine modulation of spike dynamics in bursting neurons.

The pyloric network of the lobster stomatogastric ganglion is a prime example of an oscillatory neural circuit. In our previous study on the firing patterns of pyloric neurons we observed characteristic temporal structures termed 'interspike interval (ISI) signatures' which were found to depend on the synaptic connectivity of the network. Dopamine, a well-known modulator of the pyloric network, is known to affect inhibitory synapses so it might also tune the fine temporal structure of intraburst spikes, a phenomenon not previously investigated.

Winnerless competition between sensory neurons generates chaos: A possible mechanism for molluscan hunting behavior.

In the presence of prey, the marine mollusk Clione limacina exhibits search behavior, i.e., circular motions whose plane and radius change in a chaotic-like manner.

Synaptic modulation of the interspike interval signatures of bursting pyloric neurons.

The pyloric network of the lobster stomatogastric nervous system is one of the best described assemblies of oscillatory neurons producing bursts of action potentials. While the temporal patterns of bursts have been investigated in detail, those of spikes have received less attention. Here we analyze the intraburst firing patterns of pyloric neurons and the synaptic interactions shaping their dynamics in millisecond time scales not performed before.

Recovery of hidden information through synaptic dynamics.

The role of synaptic dynamics in processing neural information is investigated in a neural information channel with realistic model neurons having chaotic intrinsic dynamics. Our neuron models are realized in simple analogue circuits, and our synaptic connections are realized both in analogue circuits and through a dynamic clamp program. The information which is input to the first chaotic neuron in the channel emerges partially absent and partially 'hidden'.

Inhibitory synchronization of bursting in biological neurons: dependence on synaptic time constant.

Using the dynamic clamp technique, we investigated the effects of varying the time constant of mutual synaptic inhibition on the synchronization of bursting biological neurons. For this purpose, we constructed artificial half-center circuits by inserting simulated reciprocal inhibitory synapses between identified neurons of the pyloric circuit in the lobster stomatogastric ganglion. With natural synaptic interactions blocked (but modulatory inputs retained), these neurons generated independent, repetitive bursts of spikes with cycle period durations of approximately 1 s.