The respiratory control system, broadly speaking, comprises a controller, sensors, and a plan. This hierarchical structure, in which there is central processing of afferent input, is important for coordinating respiratory movements with behaviors such as eating, speaking, and moving. The controller is a neuronal network within the central nervous system (CNS), which is responsible for generating and modulating individual breaths and the overall breathing pattern. Often referred to as the respiratory central pattern generator (rCPG), the controller comprises reciprocally connected neuronal populations in the medulla and pons. Neural output from the rCPG drives the activity of various motor neuron pools. Motor neurons in the spinal cord (e.g., phrenic and intercostal) innervate the respiratory pump muscles, while brain stem motorneurons innervate upper airway muscles. The so-called “plant” is animportant component of breathing control and includes the CO2stores, which are made up of lung stores and circulating blood volume including hemoglobin. Closed loop feedback to the controller is supplied by chemoreceptors and mechanoreceptors. The consistent cycling of the ventilatory pattern is generated spontaneously from the spatial and functional architecture of the rCPG. Intrinsic membrane properties of rhythmically active neurons within the rCPG are capable of producing automatic periodicity.4 In addition, reciprocal (excitatory and inhibitory) synaptic connections between neuronal populations in the medulla and pons are believed to be critical for the automatic generation of the respiratory rhythm.
The neural respiratory cycle comprises three phasesInspiration (I) involves ramp-like increases in inspiratory motor neuron firing, which drive phrenic nerve activity throughout this phase. The first phase of expiration (E1) is often called post-inspiration because inspiratory motor neurons are still active. Persistent inspira-
tory motor activity during E1, which declines throughout this phase, acts to slow the exit of air from the lungs. Finally, during the second phase of expiration (E2), expiratory muscles are typically electrically silent. During this phase of passive relaxation, gas is expelled as the
lungs and chest wall return to their equilibrium state (i.e., functional residual capacity). However, under conditions where respiratory drive is increased, expiratory muscles including the internal intercostal and abdominal muscles become active during E2. This notion is an
example of how the central controller, influenced by sensory feedback, modulates and alters the integrated motor response of the system.