Detailed Model of Intersegmental Coordination in the Timing Network of the Leech Heartbeat Central Pattern Generator

2004 ◽  
Vol 91 (2) ◽  
pp. 958-977 ◽  
Author(s):  
Sami H. Jezzini ◽  
Andrew A. V. Hill ◽  
Pavlo Kuzyk ◽  
Ronald L. Calabrese
Keyword(s):  
Central Pattern Generator ◽  
Pattern Generator ◽  
Living System ◽  
Cycle Period ◽  
Intersegmental Coordination ◽  
Experimental Conditions ◽  
Detailed Model ◽  
Distributed Structure ◽  
Synaptic Interactions ◽  
Oscillatory Neuronal Networks

To address the general problem of intersegmental coordination of oscillatory neuronal networks, we have studied the leech heartbeat central pattern generator. The core of this pattern generator is a timing network that consists of two segmental oscillators, each of which comprises two identified, reciprocally inhibitory oscillator interneurons. Intersegmental coordination between the segmental oscillators is mediated by synaptic interactions between the oscillator interneurons and identified coordinating interneurons. The small number of neurons (8) and the distributed structure of the timing network have made the experimental analysis of the segmental oscillators as discrete, independent units possible. On the basis of this experimental work, we have made conductance-based models to explore how intersegmental phase and cycle period are determined. We show that although a previous simple model, which ignored many details of the living system, replicated some essential features of the living system, the incorporation of specific cellular and network properties is necessary to capture the behavior of the system seen under different experimental conditions. For example, spike frequency adaptation in the coordinating interneurons and details of asymmetries in intersegmental connectivity are necessary for replicating driving experiments in which one segmental oscillator was injected with periodic current pulses to entrain the activity of the entire network. Nevertheless, the basic mechanisms of phase and period control demonstrated here appear to be very general and could be used by other networks that produce coordinated segmental motor outflow.

Gating of Afferent Input by a Central Pattern Generator

1999 ◽  
Vol 81 (2) ◽  
pp. 950-953 ◽  
Author(s):  
Ralph A. DiCaprio
Keyword(s):  
Central Pattern Generator ◽  
Sensory Input ◽  
Motor Pattern ◽  
Pattern Generator ◽  
Afferent Input ◽  
Inhibitory Input ◽  
Cycle Period ◽  
Intracellular Recordings ◽  
Afferent Fibers ◽  
Sufficient Magnitude

Gating of afferent input by a central pattern generator. Intracellular recordings from the sole proprioceptor (the oval organ) in the crab ventilatory system show that the nonspiking afferent fibers from this organ receive a cyclic hyperpolarizing inhibition in phase with the ventilatory motor pattern. Although depolarizing and hyperpolarizing current pulses injected into a single afferent will reset the ventilatory motor pattern, the inhibitory input is of sufficient magnitude to block afferent input to the ventilatory central pattern generator (CPG) for ∼50% of the cycle period. It is proposed that this inhibitory input serves to gate sensory input to the ventilatory CPG to provide an unambiguous input to the ventilatory CPG.

scholarly journals The neuromuscular transform in a single segment of a segmented heart tube

2020 ◽  
Vol 124 (3) ◽  
pp. 914-929
Author(s):  
Angela Wenning ◽  
Young Rim Chang ◽  
Brian J. Norris ◽  
Ronald L. Calabrese
Keyword(s):  
Relaxation Time ◽  
Central Pattern Generator ◽  
Motor Neurons ◽  
Pattern Generator ◽  
Spike Frequency ◽  
Burst Duration ◽  
Cycle Period ◽  
Heart Tube ◽  
Single Segment ◽  
Over Time

Moving blood through the segmented heart tubes of leeches requires sequential constrictions driven by motor neurons controlled by a central pattern generator. In a single heart segment, we varied stimuli to explore the neuromuscular transform. Decreasing the cycle period, e.g., to increase volume pumped over time, without altering motor burst duration and intraburst spike frequency shortens relaxation time and decreases amplitude. The likely strategy to preserve constriction amplitude is to shorten burst duration while increasing spike frequency.

Deletions of Rhythmic Motoneuron Activity During Fictive Locomotion and Scratch Provide Clues to the Organization of the Mammalian Central Pattern Generator

2005 ◽  
Vol 94 (2) ◽  
pp. 1120-1132 ◽  
Author(s):  
Myriam Lafreniere-Roula ◽  
David A. McCrea
Keyword(s):  
Central Pattern Generator ◽  
Rhythmic Activity ◽  
Pattern Generator ◽  
Cycle Period ◽  
Fictive Locomotion ◽  
Integer Multiple ◽  
Extensor Motoneuron ◽  
Complete Failure ◽  
Motoneuron Activity ◽  
Decerebrate Cats

We examined the features of spontaneous deletions of bursts of motoneuron activity that can occur within otherwise rhythmic alternating flexor and extensor activity during fictive locomotion and scratch in adult decerebrate cats. Deletions of activity were observed both in hindlimb flexor and extensor motoneuron pools during brain stem–stimulation-evoked fictive locomotion but only in extensors during fictive scratch. Paired intracellular motoneuron recordings showed that deletions reduced the depolarization of homonymous motoneurons in qualitatively similar ways. Differences occurred in the extent to which activity in synergist motoneuron pools operating at other joints within the limb was reduced during deletions. The timing of the rhythmic activity that followed a deletion was often at an integer multiple of the preexisting locomotor or scratch cycle period. This maintenance of cycle period was also seen during deletions in which there was a complete failure of motoneuron depolarization. The activity of antagonist motoneurons was usually sustained during deletions with some rhythmic modulation at intervals of the preexisting cycle period. We discuss an organization of the central pattern generator for locomotion and scratch that functions as a single rhythm generator with separate and multiple pattern formation modules for controlling the hyper- and depolarization of subsets of motoneurons within the limb.

scholarly journals Using a Model to Assess the Role of the Spatiotemporal Pattern of Inhibitory Input and Intrasegmental Electrical Coupling in the Intersegmental and Side-to-Side Coordination of Motor Neurons by the Leech Heartbeat Central Pattern Generator

2008 ◽  
Vol 100 (3) ◽  
pp. 1354-1371 ◽  
Author(s):  
Paul S. García ◽  
Terrence M. Wright ◽  
Ian R. Cunningham ◽  
Ronald L. Calabrese
Keyword(s):  
Motor Neuron ◽  
Central Pattern Generator ◽  
Motor Neurons ◽  
Electrical Coupling ◽  
Pattern Generator ◽  
Living System ◽  
Phase Relations ◽  
Spatiotemporal Pattern ◽  
Intrinsic Properties ◽  
Synaptic Inputs

Previously we presented a quantitative description of the spatiotemporal pattern of inhibitory synaptic input from the heartbeat central pattern generator (CPG) to segmental motor neurons that drive heartbeat in the medicinal leech and the resultant coordination of CPG interneurons and motor neurons. To begin elucidating the mechanisms of coordination, we explore intersegmental and side-to-side coordination in an ensemble model of all heart motor neurons and their known synaptic inputs and electrical coupling. Model motor neuron intrinsic properties were kept simple, enabling us to determine the extent to which input and electrical coupling acting together can account for observed coordination in the living system in the absence of a substantive contribution from the motor neurons themselves. The living system produces an asymmetric motor pattern: motor neurons on one side fire nearly in synchrony (synchronous), whereas on the other they fire in a rear-to-front progression (peristaltic). The model reproduces the general trends of intersegmental and side-to-side phase relations among motor neurons, but the match with the living system is not quantitatively accurate. Thus realistic (experimentally determined) inputs do not produce similarly realistic output in our model, suggesting that motor neuron intrinsic properties may contribute to their coordination. By varying parameters that determine electrical coupling, conduction delays, intraburst synaptic plasticity, and motor neuron excitability, we show that the most important determinant of intersegmental and side-to-side phase relations in the model was the spatiotemporal pattern of synaptic inputs, although phasing was influenced significantly by electrical coupling.

Period Differences Between Segmental Oscillators Produce Intersegmental Phase Differences in the Leech Heartbeat Timing Network

2002 ◽  
Vol 87 (3) ◽  
pp. 1603-1615 ◽  
Author(s):  
Mark A. Masino ◽  
Ronald L. Calabrese
Keyword(s):  
Central Pattern Generator ◽  
Phase Relationship ◽  
Pattern Generator ◽  
The Other ◽  
Cycle Period ◽  
Pharmacological Manipulation ◽  
The Third ◽  
The Individual ◽  
Phase Relationships ◽  
Phase Differences

Considerable experimental and theoretical effort has been exerted to understand how constant intersegmental phase relationships are produced between oscillators in segmentally organized pattern generating networks. The phase relationship between the segmental oscillators in the isolated timing network of the leech heartbeat central pattern generator is quite regular within individual preparations. However, it varies considerably among different preparations. Our goal is to determine how the phase relationships in this network are established. Here we assess whether inherent period differences, as suggested by the excitability-gradient hypothesis, play a role in establishing the phase relationships between the two coupled segmental oscillators of the leech heartbeat timing network. To do this we developed methods for reversibly uncoupling the segmental oscillators (sucrose knife) and pharmacological manipulation of the individual oscillators (split bath). Differences in inherent cycle periods between the third and fourth segmental oscillators (G3 and G4) were present in most (20 of 26) preparations. These period differences correlated with the phase differences observed between the segmental oscillators in the recoupled timing network, such that the oscillator with the faster cycle period, regardless of the segment in which it was located, led in phase in proportion to its period difference with the other oscillator. The phase differences between the original (coupled) and recoupled states of individual preparations were similar. Thus application and removal of the sucrose knife did not alter the period difference between the segmental oscillators in the timing network. Pharmacological manipulation of the uncoupled oscillators to alter the period difference between the oscillators led to similar correlated phase differences in the recoupled timing network. Across all experiments the uncoupled segmental oscillator with the faster cycle period established the cycle period of the timing network when recoupled. In conclusion, our findings indicate that an excitability-gradient plays a role in establishing the phase relationship between the segmental oscillators of the leech heartbeat central pattern generator since inherent period differences present between the oscillators are correlated to the phase relationships of the coupled/recoupled timing network.

Modeling Alternation to Synchrony with Inhibitory Coupling: A Neuromorphic VLSI Approach

2000 ◽  
Vol 12 (10) ◽  
pp. 2259-2278 ◽  
Author(s):  
Gennady S. Cymbalyuk ◽  
Girish N. Patel ◽  
Ronald L. Calabrese ◽  
Stephen P. DeWeerth ◽  
Avis H. Cohen
Keyword(s):  
Central Pattern Generator ◽  
Large Scale ◽  
Pattern Generator ◽  
Integrated System ◽  
Simple Explanation ◽  
Synaptic Coupling ◽  
Silicon Neurons ◽  
Inhibitory Coupling ◽  
Synchronous Oscillations ◽  
Oscillatory Neuronal Networks

We developed an analog very large-scale integrated system of two mutually inhibitory silicon neurons that display several different stable oscillations. For example, oscillations can be synchronous with weak inhibitory coupling and alternating with relatively strong inhibitory coupling. All oscillations observed experimentally were predicted by bifurcation analysis of a corresponding mathematical model. The synchronous oscillations do not require special synaptic properties and are apparently robust enough to survive the variability and constraints inherent in this physical system. In biological experiments with oscillatory neuronal networks, blockade of inhibitory synaptic coupling can sometimes lead to synchronous oscillations. An example of this phenomenon is the transition from alternating to synchronous bursting in the swimming central pattern generator of lamprey when synaptic inhibition is blocked by strychnine. Our results suggest a simple explanation for the observed oscillatory transitions in the lamprey central pattern generator network: that inhibitory connectivity alone is sufficient to produce the observed transition.

Forcing of coupled nonlinear oscillators: studies of intersegmental coordination in the lamprey locomotor central pattern generator

1990 ◽  
Vol 64 (3) ◽  
pp. 862-871 ◽  
Author(s):  
T. L. Williams ◽  
K. A. Sigvardt ◽  
N. Kopell ◽  
G. B. Ermentrout ◽  
M. P. Remler
Keyword(s):  
Central Pattern Generator ◽  
Coupled Oscillators ◽  
Nonlinear Oscillators ◽  
Pattern Generator ◽  
Fictive Locomotion ◽  
Frequency Range ◽  
Intersegmental Coordination ◽  
Coupled Nonlinear Oscillators ◽  
Phase Lags ◽  
A Chain

1. This paper reports the results of an investigation of the basic mechanisms underlying intersegmental coordination in lamprey locomotion, by the use of a combined mathematical and biological approach. 2. Mathematically, the lamprey central pattern generator (CPG) is described as a chain of coupled nonlinear oscillators; experimentally, entrainment of fictive locomotion by imposed movement has been investigated. Interpretation of the results in the context of the theory has allowed conclusions to be drawn about the nature of ascending and descending coupling in the lamprey spinal CPG. 3. Theory predicts and data show that 1) the greater the number of oscillators in the chain, the smaller is the entrainment frequency range and 2) it is possible to entrain both above and below the rest frequency at one end but only above or below at the other end. 4. In the context of the experimental results, the theory indicates the following: 1) ascending coupling sets the intersegmental phase lags, whereas descending coupling changes the frequency of the coupled oscillators; 2) there are differences in the ascending and descending coupling other than strength; and it also suggests that 3) coupling slows down the oscillators.

scholarly journals Maintenance of motor pattern phase relationships in the ventilatory system of the crab

1997 ◽  
Vol 200 (6) ◽  
pp. 963-974
Author(s):  
R Dicaprio ◽  
G Jordan ◽  
T Hampton
Keyword(s):  
Membrane Potential ◽  
Central Pattern Generator ◽  
Motor Pattern ◽  
Pattern Generator ◽  
Rate Of Change ◽  
Cycle Period ◽  
Potential Oscillation ◽  
Cycle Frequency ◽  
Membrane Potential Oscillation ◽  
Phase Relationships

The central pattern generator responsible for the gill ventilation rhythm in the shore crab Carcinus maenas can produce a functional motor pattern over a large (eightfold) range of cycle frequencies. One way to continue to generate a functional motor pattern over such a large frequency range would be to maintain the relative timing (phase) of the motor pattern as cycle frequency changes. This hypothesis was tested by measuring the phase of eight events in the motor pattern from extracellular recordings at different rhythm frequencies. The motor pattern was found to maintain relatively constant phase relationships among the various motor bursts in this rhythm over a large (sevenfold) range of cycle frequencies, although two phase-maintaining subgroups could be distinguished. Underlying this phase maintenance is a corresponding change in the time delay between events in the motor pattern ranging from 470 to 1800 ms over a sevenfold (300­2100 ms) change in cycle period. Intracellular recordings from ventilatory neurons indicate that there is very little change in the membrane potential oscillation in the motor neurons with changes in cycle frequency. However, recordings from nonspiking interneurons in the ventilatory central pattern generator reveal that the rate of change of the membrane potential oscillation of these neurons varies in proportion to changes in cycle frequency. The strict biomechanical requirements for efficient pumping by the gill bailer, and the fact that work is performed in all phases of the motor pattern, may require that this motor pattern maintain phase at all rhythm frequencies.

scholarly journals Control of a central pattern generator by an identified modulatory interneurone in crustacea. II. Induction and modification of plateau properties in pyloric neurones

1983 ◽  
Vol 105 (1) ◽  
pp. 59-82
Author(s):  
P. S. Dickinson ◽  
F. Nagy
Keyword(s):  
Motor Pattern ◽  
Pattern Generator ◽  
Membrane Properties ◽  
Plateau Potentials ◽  
Stomatogastric Nervous System ◽  
Experimental Conditions ◽  
Pyloric Network ◽  
Synaptic Interactions ◽  
Voltage Dependent ◽  
A Current

In the isolated stomatogastric nervous system of the lobster Fasus lalandii, the strong modifications of the pyloric motor pattern induced by firing of the single anterior pyloric modulator neurone (APM) are due primarily to modulation by APM activity of the regenerative membrane properties which are responsible for the ‘burstiness’ of all the pyloric neurones and particularly of the non-pacemaker neurones (constrictor motoneurones). This modulation has been studied under experimental conditions where the main extrinsic influences usually received by the pyloric constrictor neurones (intra-network synaptic interactions, activity of pacemaker neurones, and phasic central inputs from two premotor centres) are minimal. Under these conditions a brief discharge of neurone APM induces long plateaus of firing in all of the pyloric neurones. The non-pacemaker neurones of the pyloric network are not simply passive follower neurones, but can produce regenerative depolarizations (plateau potentials) during which the neurones fire spikes. The ability of the pyloric constrictor neurones to produce plateau potentials (plateau properties) contributes greatly to the generation of the rhythmical pyloric motor pattern. When these neurones spontaneously express their plateau properties, firing of neurone APM amplifies these properties. When most of the central inputs usually received by the pyloric constrictor neurones are experimentally suppressed, these neurones can no longer produce plateau potentials. In such conditions, firing of the single modulatory neurone APM can reinduce plateau properties of the pyloric constrictor neurones. In addition, firing in APM neurone slows down the active repolarization phase which terminates the plateau potentials of pyloric constrictor neurones. This effect is long-lasting and voltage-dependent. Modulation by APM of the plateau properties of the pyloric neurones also changes the sensitivity of these neurones to synaptic inputs. This effect can explain the strong modifications that an APM discharge exerts on a current pyloric motor pattern. Moreover, it might render the motoneurones of the pyloric pattern generator more sensitive to inputs from a command oscillator, and contribute to switching on the pyloric motor pattern.

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