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Neural mechanisms potentially contributing to the intersegmental phase lag in lamprey II.: Hemisegmental oscillations produced by mutually coupled excitatory neurons
KTH, Superseded Departments, Numerical Analysis and Computer Science, NADA.ORCID iD: 0000-0002-0550-0739
KTH, Superseded Departments, Numerical Analysis and Computer Science, NADA.ORCID iD: 0000-0002-2358-7815
1999 (English)In: Biological cybernetics, ISSN 0340-1200, E-ISSN 1432-0770, Vol. 81, no 4, 299-315 p.Article in journal (Refereed) Published
Abstract [en]

Most previous models of the spinal central pattern generator (CPG) underlying locomotion in the lamprey have relied on reciprocal inhibition between the left and right side for oscillations to be produced. Here, we have explored the consequences of using self-oscillatory hemisegments. Within a single hemisegment, the oscillations are produced by a network of recurrently coupled excitatory neurons (E neurons) that by themselves are not oscillatory but when coupled together through N-methyl-D-aspartate (NMDA) and x-amino-3-hydroxy-5-methyl-4-isoxazolepropionicacid (AMPA)/kainate transmission can produce oscillations. The bursting mechanism relies on intracellular accumulation of calcium that activates Ca2+-dependent KC The intracellular calcium is modeled by two different intracellular calcium pools, one of which represents the calcium entry following the action potential, Ca-AP pool, and the other represents the calcium inflow through the NMDA channels, Ca-NMDA pool. The Ca2+-dependent K+ activated by these two calcium pools are referred to as K-CaAP and K-CaNMDA respectively, and their relative conductances are modulated and increase with the background activation of the network. When changing the background stimulation, the bursting activity in this network can be made to cover a frequency range of 0.5-5.5 Hz with reasonable burst proportions if the adaptation is modulated with the activity. When a chain of such hemisegments are coupled together, a phase lag along the chain can be produced. The local oscillations as well as the phase lag is dependent on the axonal conduction delay as well as the types of excitatory coupling that are assumed, i.e. AMPA/kainate and/or NMDA. When the caudal excitatory projections are extended further than the rostral ones, and assumed to be of approximately equal strength, this kind of network is capable of reproducing several experimental observations such as those occurring during strychnine blockade of the left-right reciprocal inhibition. Addition of reciprocally coupled inhibitory neurons in such a network gives rise to antiphasic activity between the left and right side, but not necessarily to any change of the frequency if the burst proportion of the hemisegmental bursts is well below 50%. Prolongation of the C neuron projection in the rostrocaudal direction restricts the phase lag produced by only the excitatory hemisegmental network by locking together the interburst intervals at different levels of the spinal cord.

Place, publisher, year, edition, pages
1999. Vol. 81, no 4, 299-315 p.
Keyword [en]
DEPENDENT POTASSIUM CHANNELS, SPINAL-CORD, FICTIVE LOCOMOTION, GLYCINERGIC INHIBITION, SYNAPTIC DRIVE, NETWORK, MODEL, INTERNEURONS, SIMULATIONS, MODULATION
National Category
Engineering and Technology
Identifiers
URN: urn:nbn:se:kth:diva-13368DOI: 10.1007/s004220050564ISI: 000083208800003OAI: oai:DiVA.org:kth-13368DiVA: diva2:324696
Note
QC 20100616Available from: 2010-06-16 Created: 2010-06-16 Last updated: 2010-06-16Bibliographically approved
In thesis
1. Modeling of bursting mechanisms and coordination in a spinal central pattern generator
Open this publication in new window or tab >>Modeling of bursting mechanisms and coordination in a spinal central pattern generator
1998 (English)Doctoral thesis, comprehensive summary (Other scientific)
Abstract [en]

Mechanisms underlying lotal bursting as well as coordinationbetween different levels of a spinal CPG generating locomotionhave been investigated using computer simulations. A"primitive" jawless vertebrate, the lamprey, is used a.s aprototype model. Most simulations have been conducted using abiophysical neu ron model built on the Hodgkin-Huxley formalismand equipped with Nu+, K+,Ca²+, Kca, LVACa²+ and NMDA activated channels. Inhibitory andexcitatory AMPA/kainate and NMDA synapses are modeled as timedependent conductances with appropriate reversal potentials.For tomparison, Morris-Letar oscillators as well as adaptingleaky integrator-like units are also used.

The basic identified building blocks of the CPG, generatingalternating left right burst activity, tonsist of ipsilaterallyprojecting excitatory neurons (E) and contralaterallyprojecting inhibitory neurons (C). The model neurons are connected in the same way ss has been established experimentally.Sinte several complementary mechanisms may play a role, thepotential of two different neural mechanisms have been exploredwhich can provide burst activity at the segmen tal level, andintersegmental coordination. When alternating left-rightactivity is produced through an escape-like mechanism the quietside is able to become ac tive despite ongoing inhibition fromthe contralateral side. Reciprocal inhibition is then a crucialburst terminating factor. Burst frequency is strongly affectedby the effective inhibition and the drive to escape fromongoing inhibition. Several factors influence this process. Kcacurrents control spike frequency on the active sideand also a post-burst hyperpolarization on the inactive side.Postin hibitory rebound properties, carried by e.g. low voltageactivatedCa²+ currents further can promote escape. Phasicipsilateral excitation and NMDA membrane properties stabilizethe rhythm, especially in the lower frequency range. Severalexperimental observations can be explained based on the effectthese different factors have on effective inhibition andtendency for escape.

Bursting can, however, also be produced by a networkdeprived of inhibition, showing that powerful burst terminatingmechanisms not requiring inhibition exist. In the model withbiophysically detailed neurons such a mechanism could beactivation ofKcacurrents due to accumulation ofCa²+ during the active phase. As shown innon-spiking, as well as biophysically detailed models, aconstant burst proportion over a wide frequency range can beachieved by modulation of the rel ative strength of adaptationin such networks. The left-right inhibition causes left-rightalternation but may not affect the frequency of bursting.

When both types of lotal oscillatory networks are extendedlongitudinally, a rostral to caudal phase delay is producedwhen caudal projections are extended further than the rostralenes. However, the excitatory versus inhibitory projec tionsmay have different roles in the two alternative models. Thisrelative phase delay expressed as % of cycle duration,increases in general with frequency. The simulations suggestthat the conditions at the ends of the simulated chain arecritical for the resulting phase lag. The capability ofbuffering against frequency variations and rapid adjustmentsfollowing perturbations is discussed and com pared with chainsof relaxation oscillators and phase-coupled oscillators.

Place, publisher, year, edition, pages
Stockholm: KTH, 1998. 82 p.
Series
Trita-NA, ISSN 0348-2952 ; 98:10
Keyword
adaptation, central pattern generator, computer simulation, inter segmental coordination, lamprey, locomotion, neural network, rhythmogenesis
National Category
Engineering and Technology
Identifiers
urn:nbn:se:kth:diva-2673 (URN)91-7170-255-5 (ISBN)
Public defence
1998-06-16, 00:00
Note
QC 20100616Available from: 2000-01-01 Created: 2000-01-01 Last updated: 2010-06-16Bibliographically approved

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