Encoding of target range and its representation in the auditory cortex of the mustached bat

The time course of acoustic events is a critical element for the recognition of biologically meaningful sounds. Echolocating bats analyze the time intervals between their emitted biosonar pulses and the echoes returning from objects to assess target distance (range). In this study, we have explored the auditory cortex of the mustached bat, Pteronotus parnellii rubiginosus, using pairs of acoustic stimuli mimicking the multiharmonic biosonar signals (pulses) used by this species and their echoes. A discrete field of auditory cortex dorsorostral to the tonotopically organized primary field contains neurons which are insensitive to pure tone, frequency-modulated (FM), or noise stimuli presented singly. Rather, they respond strongly to pairs of stimuli, specifically, the fundamental FM component of the pulse paired with an FM component of one of the higher harmonics of the echo. We call these neurons FM1-FMn facilitation neurons. There are three separate longitudinal clusters in this cortical area containing FM1-FM2, FM1-FM3, and FM1-FM4 neurons, respectively. Moreover, FM1-FMn neurons are specifically sensitive to the time delay between the two FM components, i.e., the time delay of the echo from the pulse. Thus, they can decode target range. Two types of delay-sensitive neurons were found. Tracking neurons, whose response to echo delay varied according to repetition rate and stimulus duration, were found rarely. Delay-tuned neurons, which were tuned to specific time delays (best delays) of the echo from the pulse, were much more evident. Both types of neurons are organized into columns with similar best delays, and the best delay of delay-tuned neurons was found to increase systematically along the cortical surface in the rostrocaudal direction. This area, therefore, contains a neural representation of target range along this best delay axis. Such an axis exists in each of the clusters of FM1-FM2, FM1-FM3, and FM1-FM4 neurons. This is a new type of cortical organization which is not tonotopic but which represents an important acoustic cue related to the time course of acoustic events.