Respiratory responses to shivering produced by external and central cooling in the pigeon

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Abstract
Respiratory responses of pigeons to spinal cord cooling (5–6°C) in neutral environment (T a=28°C), to ambient cooling (T a=5°C), and to simultaneous spinal cord and ambient cooling were measured. Spinal cord cooling produced shivering and a 242% increase in heat production (M); expiratory flow rate (VE) increased 216%, a result of increases in both respiratory frequency (160%) and tidal volume (140%). Increases produced by ambient cooling compared to thermoneutral controls were slightly, but not significantly, less than those during spinal cord cooling: M=203%, VE=199%, respiratory frequency (f R)=146%, tidal volume (V T)=138%. Spinal cord cooling at low ambient temperature produced greater increases in shivering, heat production and respiration compared to thermoneutral controls than either type of cooling alone: M=337%, VE,f R=198%,V T=178%. The oxygen extraction from the ventilatory gas remained relatively constant among the different groups.f R,V T and VE were all significantly linearly related to M over the wide range studied. These relationships were independent of whether cooling was central or external. Respiratory changes induced by the onset and end of spinal cord cooling were immediate and closely correlated with the magnitude of shivering. It is unlikely that changes in arterial and venous blood gases during shivering effected the major portion of the respiratory response. Thus, it is suggested that a control mechanism of the respiratory center via afferents from the shivering muscles is important in increasing respiration during central or external cooling.