A narrow ear canal reduces sound velocity to create additional acoustic inputs in a micro-scale insect ear

Located in the forelegs, katydid ears are unique among arthropods in having outer, middle and inner components, analogous to the mammalian ear. Unlike mammals, sound is received externally via two tympanic membranes in each ear, and internally via a narrow ear canal (EC) derived from the respiratory tracheal system. Inside the EC sound travels slower than in free air, causing temporal and pressure differences between external and internal inputs. The delay was suspected to arise as a consequence of the narrowing EC geometry. If true, a reduction in sound velocity should persist independently of the gas composition in the EC (e.g. air, CO2). Integrating laser Doppler vibrometry, micro-CT scanning, and numerical analysis on precise 3D geometries of each experimental animal EC, we demonstrate that the narrowing radius of the EC is the main factor reducing sound velocity. Both experimental and numerical data also show that sound velocity is reduced further when excess CO2 fills the EC. Likewise, the EC bifurcates at the tympanal level (one branch for each tympanic membrane) creating two additional narrow internal sound paths and imposing different sound velocities for each tympanic membrane. Therefore, external and internal inputs total to four sound paths for each ear (only one for the human ear). New research paths, and implication of findings in avian directional hearing are discussed

Spatial Hearing with Simultaneous Sound Sources: A Psychophysical Investigation

This thesis provides an overview of work conducted to investigate human spatial hearing in situations involving multiple concurrent sound sources. Much is known about spatial hearing with single sound sources, including the acoustic cues to source location and the accuracy of localisation under different conditions. However, more recently interest has grown in the behaviour of listeners in more complex environments. Concurrent sound sources pose a particularly difficult problem for the auditory system, as their identities and locations must be extracted from a common set of sensory receptors and shared computational machinery. It is clear that humans have a rich perception of their auditory world, but just how concurrent sounds are processed, and how accurately, are issues that are poorly understood. This work attempts to fill a gap in our understanding by systematically examining spatial resolution with multiple sound sources. A series of psychophysical experiments was conducted on listeners with normal hearing to measure performance in spatial localisation and discrimination tasks involving more than one source. The general approach was to present sources that overlapped in both frequency and time in order to observe performance in the most challenging of situations. Furthermore, the role of two primary sets of location cues in concurrent source listening was probed by examining performance in different spatial dimensions. The binaural cues arise due to the separation of the two ears, and provide information about the lateral position of sound sources. The spectral cues result from location-dependent filtering by the head and pinnae, and allow vertical and front-rear auditory discrimination. Two sets of experiments are described that employed relatively simple broadband noise stimuli. In the first of these, two-point discrimination thresholds were measured using simultaneous noise bursts. It was found that the pair could be resolved only if a binaural difference was present; spectral cues did not appear to be sufficient. In the second set of experiments, the two stimuli were made distinguishable on the basis of their temporal envelopes, and the localisation of a designated target source was directly examined. Remarkably robust localisation was observed, despite the simultaneous masker, and both binaural and spectral cues appeared to be of use in this case. Small but persistent errors were observed, which in the lateral dimension represented a systematic shift away from the location of the masker. The errors can be explained by interference in the processing of the different location cues. Overall these experiments demonstrated that the spatial perception of concurrent sound sources is highly dependent on stimulus characteristics and configurations. This suggests that the underlying spatial representations are limited by the accuracy with which acoustic spatial cues can be extracted from a mixed signal. Three sets of experiments are then described that examined spatial performance with speech, a complex natural sound. The first measured how well speech is localised in isolation. This work demonstrated that speech contains high-frequency energy that is essential for accurate three-dimensional localisation. In the second set of experiments, spatial resolution for concurrent monosyllabic words was examined using similar approaches to those used for the concurrent noise experiments. It was found that resolution for concurrent speech stimuli was similar to resolution for concurrent noise stimuli. Importantly, listeners were limited in their ability to concurrently process the location-dependent spectral cues associated with two brief speech sources. In the final set of experiments, the role of spatial hearing was examined in a more relevant setting containing concurrent streams of sentence speech. It has long been known that binaural differences can aid segregation and enhance selective attention in such situations. The results presented here confirmed this finding and extended it to show that the spectral cues associated with different locations can also contribute. As a whole, this work provides an in-depth examination of spatial performance in concurrent source situations and delineates some of the limitations of this process. In general, spatial accuracy with concurrent sources is poorer than with single sound sources, as both binaural and spectral cues are subject to interference. Nonetheless, binaural cues are quite robust for representing concurrent source locations, and spectral cues can enhance spatial listening in many situations. The findings also highlight the intricate relationship that exists between spatial hearing, auditory object processing, and the allocation of attention in complex environments

Spatial Hearing with Simultaneous Sound Sources: A Psychophysical Investigation

This thesis provides an overview of work conducted to investigate human spatial hearing in situations involving multiple concurrent sound sources. Much is known about spatial hearing with single sound sources, including the acoustic cues to source location and the accuracy of localisation under different conditions. However, more recently interest has grown in the behaviour of listeners in more complex environments. Concurrent sound sources pose a particularly difficult problem for the auditory system, as their identities and locations must be extracted from a common set of sensory receptors and shared computational machinery. It is clear that humans have a rich perception of their auditory world, but just how concurrent sounds are processed, and how accurately, are issues that are poorly understood. This work attempts to fill a gap in our understanding by systematically examining spatial resolution with multiple sound sources. A series of psychophysical experiments was conducted on listeners with normal hearing to measure performance in spatial localisation and discrimination tasks involving more than one source. The general approach was to present sources that overlapped in both frequency and time in order to observe performance in the most challenging of situations. Furthermore, the role of two primary sets of location cues in concurrent source listening was probed by examining performance in different spatial dimensions. The binaural cues arise due to the separation of the two ears, and provide information about the lateral position of sound sources. The spectral cues result from location-dependent filtering by the head and pinnae, and allow vertical and front-rear auditory discrimination. Two sets of experiments are described that employed relatively simple broadband noise stimuli. In the first of these, two-point discrimination thresholds were measured using simultaneous noise bursts. It was found that the pair could be resolved only if a binaural difference was present; spectral cues did not appear to be sufficient. In the second set of experiments, the two stimuli were made distinguishable on the basis of their temporal envelopes, and the localisation of a designated target source was directly examined. Remarkably robust localisation was observed, despite the simultaneous masker, and both binaural and spectral cues appeared to be of use in this case. Small but persistent errors were observed, which in the lateral dimension represented a systematic shift away from the location of the masker. The errors can be explained by interference in the processing of the different location cues. Overall these experiments demonstrated that the spatial perception of concurrent sound sources is highly dependent on stimulus characteristics and configurations. This suggests that the underlying spatial representations are limited by the accuracy with which acoustic spatial cues can be extracted from a mixed signal. Three sets of experiments are then described that examined spatial performance with speech, a complex natural sound. The first measured how well speech is localised in isolation. This work demonstrated that speech contains high-frequency energy that is essential for accurate three-dimensional localisation. In the second set of experiments, spatial resolution for concurrent monosyllabic words was examined using similar approaches to those used for the concurrent noise experiments. It was found that resolution for concurrent speech stimuli was similar to resolution for concurrent noise stimuli. Importantly, listeners were limited in their ability to concurrently process the location-dependent spectral cues associated with two brief speech sources. In the final set of experiments, the role of spatial hearing was examined in a more relevant setting containing concurrent streams of sentence speech. It has long been known that binaural differences can aid segregation and enhance selective attention in such situations. The results presented here confirmed this finding and extended it to show that the spectral cues associated with different locations can also contribute. As a whole, this work provides an in-depth examination of spatial performance in concurrent source situations and delineates some of the limitations of this process. In general, spatial accuracy with concurrent sources is poorer than with single sound sources, as both binaural and spectral cues are subject to interference. Nonetheless, binaural cues are quite robust for representing concurrent source locations, and spectral cues can enhance spatial listening in many situations. The findings also highlight the intricate relationship that exists between spatial hearing, auditory object processing, and the allocation of attention in complex environments

Behavioural and neural correlates of binaural hearing

The work in this thesis involves two separate projects. The first project involves the behavioural measurement of auditory thresholds in the ferret (Mustela Putorius). A new behavioural paradigm using a sound localisation task was developed which produces reliable psychophysical detection thresholds in animals. Initial attempts to use the task failed and after further investigation improvements were made. These changes produced a task that successfully produced reliably low thresholds. Different methods of testing, and the number of experimental trials required, here then explored systemically. The refined data collection method was then used to investigate frequency resolution in the ferret. These data demonstrated that the method was suitable for measuring perceptual frequency selectivity. It revealed that the auditory filters of ferrets are broader than several other species. In some cases this was also broader than neural estimates would suggest. The second project involved the measurement of neural data in the Guinea Pig (Cavia porecellus). More specifically the project aimed to test the ability of the primary auditory cortex (AI) to integrate high frequency spatial cues. Two experiments were required to elucidate these data. The first experiment demonstrated a relationship between frequency and space, though these data proved noisy. A second experiment was conducted, focussing on improving the quality of the data this allowed for a more quantitative approach to be applied. The results highlighted that though AI neurons are responsive over a broad frequency range, inhibitory binaural interactions integrate spatial information over a smaller range. Binaural interactions were only strong when sounds in either ear were closely matched in frequency. In contrast, excitatory binaural interactions did not generally depend on the interaural frequency difference. These findings place important constraints on the across frequency integration of binaural level cues