CENTRAL PATTERN GENERATORS

DEFINITION

Repetitive (cyclic) movements underlie crucial behaviors like locomotion

(walking, swimming, crawling, and flying), as well as essential vegetative

functions like respiration, heartbeat in some invertebrates, chewing, and gut

movements. Although both sensory feedback and neuromuscular dynamics play

an important role in shaping rhythmic motor output, the basic rhythmic activity

patterns are generated by central circuits called central pattern generators (CPGs).

These CPGs can, when properly activated, produce rhythmic network activity in

the absence of external timing cues that is without rhythmic sensory feedback or

rhythm activation by descending neurons.

Cellular and Network Mechanisms of Rhythm

Generation
Rhythmic motor patterns almost always consist of sequential or alternating

activation of antagonistic groups of muscles, and this sequence is seen in the

timing of bursts of action potentials between groups of neurons in the underlying

CPG. How this rhythmic activity is generated and how bursting and specific

relative timing between different neurons is achieved depend on the circuit

architecture, the properties of synaptic connections, and the intrinsic membrane

properties of CPG neurons.

Synaptic Properties

Even though a wide variety of synapse types may be found in any given CPG,

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synchronization of neurons that fire in the same phase of the pattern is often

achieved or aided by electrical coupling through gap junctions between these

neurons, whereas alternating activity is often based on reciprocal inhibitory

connections.

Many chemical synaptic interactions in CPGs are based on spike-mediated

mechanisms; that is, transmitter is released in response to presynaptic action

potentials. However, local interneurons often also rely on graded synaptic

transmission; that is, they release transmitter as a graded function of presynaptic

membrane potential. In general, it is not clear what the functional significance of

graded versus spike-mediated transmission is in the specific context of rhythm

generation. However, in a given circuit, the dynamics of these two types of

transmission and their interaction with intrinsic neuronal membrane generation of

motor patterns in the absence of sensory feedback. (a) In the locust, rhythmic motor

output to flight muscles can be generated in the difference thoracic nerve cord in an

immobilized animal with wind stimulation to the head. Alternating activity between

elevator and depressor motor neurons is generated even though no movement is

produced and no sensory feedback reaches the central nervous system.

(b) Transverse brain stem slices containing the central pattern generating neurons for

respiration in the pre-Bo¨tzingercomplex. .Population activity is recorded

extracellularly (lower trace), and integrated (upper trace) to show rhythmic activity

that corresponds to inspiration. XII, hypoglossal nucleus; X, dorsal motor nucleus of

vagus; NA, nucleus ambiguus; SP5, spinal trigeminal nucleus; IO, inferior olive.

Properties can be quite different, and CPG possibilities of both for dynamic

circuit operation. Many synapses show considerable short term plasticity such as

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depression, facilitation, or mixtures of both. In CPGs, synaptic dynamics can

strongly influence the frequency of rhythmic network activity and the relative

timing between different groups of neurons. This is because in rhythmic systems,

depression and facilitation cause the strength of a synaptic connection to be

dependent on rhythm frequency and on the duty cycle, the number of spikes, or

the spike frequency of the presynaptic neuron. In the heart beat CPG, the synaptic

strength of connections from a single presynaptic neuron onto several post-

synaptic neurons peaks at different times during a single presynaptic burst, as a

result of different synaptic dynamics at these connections. Because of synaptic

depression, some synaptic connections in the crustacean stomato gastric ganglion

become weaker and peak later the faster the rhythm is, and this can help maintain

stable relative timing between neurons at different rhythm frequencies.

Intrinsic Neuronal Properties

As in all neural networks, activity does not depend only on synaptic

connections but also on a potentially rich complement of voltage gated ion

channels that determine a neuron’s intrinsic membrane properties and

therefore how it behaves in isolation and in response to synaptic input. A

number of intrinsic properties are commonly found in CPG neurons among

them are endogenous bursting properties, post inhibitory rebound, plateau

properties, spike frequency adaptation, and restorative sag potentials.

Endogenous bursting means that a neuron can generate membrane oscillations in

the absence of synaptic inputs. This is an important mechanism of rhythm

generation in a number of systems.

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 In other neurons, bursting can be generated on rebound from

inhibitory synaptic input. This is an important mechanism for the production

of alternating activity, and the time course of this rebound is an important

determinant of relative timing between alternating rhythmic neurons.

The term plateau potential, or bistable membrane behavior, describes the

ability of neurons to generate sustained firing in response to brief and relatively

weak depolarizing input. If an input crosses a certain threshold, firing activity

outlasts the stimulus significantly and can either self terminate eventually or be

shut off by (again, relatively weak) hyperpolarizing input. Plateau properties can

therefore uncouple neuronal firing from the duration and strength of excitatory

input.

(OVERVIEW OF CENTRAL PATTERN GENERATORS)

Central pattern generators are neural circuits that produce the pattern of

neural activity that underlie rhythmic motor behaviors such as walking,

swimming and feeding. These patterns are generated centrally, without the need

for sensory feedback or other patterned input.