The effects of locomotor-respiratory coupling on the pattern of breathing in horses

1. To investigate the effect of locomotor activity on the pattern of breathing in quadrupeds, ventilatory response was studied in four healthy horses during horizontal and inclined (7%) treadmill exercise at different velocities (1.4-6.9 m s(-1)) and during chemical stimulation with a rebreathing method. Stride frequency (f(s)) and locomotor-respiratory coupling (LRC) were also simultaneously determined by means of video recordings synchronized with respiratory events. 2. Tidal volume (V(T)) was positively correlated with pulmonary ventilation (V(E)) but significantly different linear regression equations were found between the experimental conditions (P < 0.0001), since the chemical hyperventilation was mainly due to increases in V(T), whereas the major contribution to exercise hyperpnoea came from changes in respiratory frequency (f(R)). 3. The average f(R) at each exercise level was not significantly different from f(S), although there was not always a tight 1:1 LRC. At constant speeds, f(S) was independent of the treadmill slope and hence the greater V(E) during inclined exercise was due to increased V(T). 4. At any ventilatory level, the differences in breathing patterns between locomotion and rebreathing or locomotion at different slopes derived from different set points of the inspiratory off-switch mechanism. 5. The percentage of single breaths entrained with locomotor rhythm rose progressively and significantly with treadmill speed (P < 0.0001) up to a 1:1 LRC and was significantly affected by treadmill slope (P < 0.001). 6. A LRC of 1:1 was systematically observed at canter (10 out of 10 trials) and sometimes at trot (5 out of 14) and it entailed (i) a 4- to 5-fold reduction in both V(T) and f(R) variability, and (ii) a gait-specific phase locking of inspiratory onset during the locomotor cycle. 7. It is concluded that different patterns of breathing are employed during locomotion and rebreathing due to the interference between locomotor and respiratory functions, which may play a role in the optimization and control of exercise ventilation in horses.