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Endogenous excitatory drive to the respiratory system in rapid eye movement sleep in cats

John Orem, Andrew T. Lovering, Witali Dunin-Barkowski and Edward H. Vidruk*

1. A putative endogenous excitatory drive to the respiratory system in rapid eye movement (REM) sleep may explain many characteristics of breathing in that state, e.g. its irregularity and variable ventilatory responses to chemical stimuli. This drive is hypothetical, and determinations of its existence and character are complicated by control of the respiratory system by the oscillator and its feedback mechanisms. In the present study, endogenous drive was studied during apnoea caused by mechanical hyperventilation. We reasoned that if there was a REM-dependent drive to the respiratory system, then respiratory activity should emerge out of the background apnoea as a manifestation of the drive.

2. Diaphragmatic muscle or medullary respiratory neuronal activity was studied in five intact, unanaesthetized adult cats who were either mechanically hyperventilated or breathed spontaneously in more than 100 REM sleep periods.

3. Diaphragmatic activity emerged out of a background apnoea caused by mechanical hyperventilation an average of 34 s after the onset of REM sleep. Emergent activity occurred in 60 % of 10 s epochs in REM sleep and the amount of activity per unit time averaged approximately 40 % of eupnoeic activity. The activity occurred in episodes and was poorly related to pontogeniculo-occipital waves. At low CO₂ levels, this activity was non-rhythmic. At higher CO₂ levels (less than 0·5 % below eupnoeic end-tidal percentage CO₂ levels in non-REM (NREM) sleep), activity became rhythmic.

4. Medullary respiratory neurons were recorded in one of the five animals. Nineteen of twenty-seven medullary respiratory neurons were excited in REM sleep during apnoea. Excited neurons included inspiratory, expiratory and phase-spanning neurons. Excitation began about 43 s after the onset of REM sleep. Activity increased from an average of 6 impulses s^-1 in NREM sleep to 15·5 impulses s^-1 in REM sleep. Neuronal activity was non-rhythmic at low CO₂ levels and became rhythmic when levels were less than 0·5 % below eupnoeic end-tidal levels in NREM sleep. The level of CO₂ at which rhythmic neuronal activity developed corresponded to eupnoeic end-tidal CO₂ levels in REM sleep.

5. These results demonstrate an endogenous excitatory drive to the respiratory system in REM sleep and account for rapid and irregular breathing and the lower set-point to CO₂ in that state.

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