Responses of neurons in the cat's superior colliculus to acoustic stimuli. I. Monaural and binaural response properties.
Using extracellular electrodes we studied acoustic responses in the superior colliculus (SC) of the barbiturate-anesthetized cat. Pure tonal stimuli were delivered through sealed and calibrated earphones and were presented either monaurally or binaurally with interaural intensity differences (IIDs) and interaural time differences (ITDs). Acoustically sensitive cells were found in the intermediate and deep layers of the SC throughout its rostrocaudal and mediolateral extent. Most cells (80%) discharged only at stimulus onset; the rest had more complex firing patterns. For 88% of our sample the mean first-spike latency measured at 20 dB above threshold ranged between 6 and 16 ms. The sharpness and threshold intensity of the frequency tuning curves varied widely. In the SC, the average characteristic frequency and threshold intensity were higher than in other auditory brain stem nuclei. Neurons whose characteristic frequency was low were never sharply tuned. The probability of response decreased when the repetition rate at which the stimuli were delivered increased. The mean stimulus interval at which spike count reached 50% of maximum was 360 ms. Most (83%) of the cells discharged only to monaural stimulation of the contralateral ear, 7% responded to tones applied to either ear and only 1% to only ipsilateral input. The remaining cells responded only to stimulation of both ears. With binaural stimuli, most neurons (80%) could be shown to receive input from both ears. Seventy percent of the binaural cells showed predominant binaural inhibition (BI), 25% binaural facilitation (BF), and 5% a more complex mixture. Because the majority of SC neurons had high characteristic frequencies, we examined their responses to IIDs. The spike count vs. IID functions of BI cells were monotonic and sigmoidal, those of BF cells were nonmonotonic and bell-shaped. The slopes and horizontal positions of the curves varied among neurons. IIDs favoring the contralateral ear were the most effective. For a given cell, increasing the mean binaural level extended the range of IIDs that evoked maximal discharge. A small number of cells was sensitive to physiologically significant interaural time differences of low-frequency tones or the envelopes of amplitude-modulated, high-frequency tones.