A parallel cholinergic brainstem pathway for enhancing locomotor drive

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Abstract
This study identifies a population of cells in lamprey brainstem that are activated by cholinergic input from the mesencephalic locomotor region and in turn excite the reticulospinal neurons that drive spinal motor neurons, representing a previously unrecognized feedforward circuit. The brainstem locomotor system is believed to be organized serially from the mesencephalic locomotor region (MLR) to reticulospinal neurons, which in turn project to locomotor neurons in the spinal cord. We identified brainstem muscarinoceptive neurons in lampreys (Petromyzon marinus) that received parallel inputs from the MLR and projected back to reticulospinal cells to amplify and extend the duration of locomotor output. These cells responded to muscarine with extended periods of excitation, received direct muscarinic excitation from the MLR and projected glutamatergic excitation to reticulospinal neurons. Targeted blockade of muscarine receptors over these neurons profoundly reduced MLR-induced excitation of reticulospinal neurons and markedly slowed MLR-evoked locomotion. The presence of these neurons forces us to rethink the organization of supraspinal locomotor control, to include a sustained feedforward loop that boosts locomotor output.