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J Neurophysiol (October 1, 2008). doi:10.1152/jn.90729.2008
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Submitted on July 2, 2008
Revised on September 18, 2008
Accepted on September 23, 2008

Asymmetric operation of the locomotor central pattern generator in the neonatal mouse spinal cord

Toshiaki Endo1 and Ole Kiehn2*

1 Karolinska Institutet
2 Karolinska Institute

* To whom correspondence should be addressed. E-mail: o.kiehn{at}ki.se.

The rhythmic voltage oscillations in motor neurons (MNs) during locomotor movements reflect the operation of the pre-MN central pattern generator (CPG) network. Recordings from MNs can, therefore, be used as a method to deduct the organization of CPGs. Here, we use continuous conductance measurements and decomposition methods to quantitatively assess the weighting and phase tuning of synaptic inputs to different flexor and extensor MNs during locomotor-like activity in the isolated neonatal mice lumbar spinal cord preparation. Whole-cell recordings were obtained from 22 flexor and 18 extensor MNs in rostral and caudal lumbar segments. In all flexor and the large majority of extensor MNs the extracted excitatory and inhibitory synaptic conductances alternate but with a predominance of inhibitory condutances, most pronounced in extensors. These conductance changes are consistent with a "push-pull" operation of locomotor CPG. The extracted excitatory and inhibitory synaptic conductances varied between 2 and 56% of the mean total conductance. Analysis of the phase tuning of the extracted synaptic conductances in flexor and extensor MNs in the rostral lumbar cord showed that the flexor phase-related synaptic conductance changes have sharper locomotor phase tuning than the extensor phase-related conductances, suggesting a modular organization of pre-motor CPG networks consisting of reciprocally coupled, but differently composed, flexor and extensor CPG networks. There was a clear difference between phase tuning in rostral and caudal MNs suggesting a distinct operation of CPG networks in different lumbar segments. The highly asymmetric features were preserved throughout all ranges of locomotor frequencies investigated and with different combinations of locomotor-inducing drugs. The asymmetric nature of CPG operation and phase tuning of the conductance profiles provide important clues to the organization of the rodent locomotor CPG and are compatible with a multi layered and distributed structure of the network.




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