Sound Localization of World and Head-Centered Space in Ferrets

The location of sounds can be described in multiple coordinate systems that are defined relative to ourselves, or the world around us. Evidence from neural recordings in animals point towards the existence of both head-centered and world-centered representations of sound location in the brain; however, it is unclear whether such neural representations have perceptual correlates in the sound localization abilities of non-human listeners. Here, we establish novel behavioral tests to determine the coordinate systems in which ferrets can localize sounds. We found that ferrets could learn to discriminate between sound locations that were fixed in either world-centered or head-centered space, across wide variations in sound location in the alternative coordinate system. Using probe sounds to assess broader generalization of spatial hearing, we demonstrated that in both head and world-centered tasks, animals used continuous maps of auditory space to guide behavior. Single trial responses of individual animals were sufficiently informative that we could then model sound localization using speaker position in specific coordinate systems and accurately predict ferrets’ actions in held-out data. Our results indicate that an animal model in which neurons are known to be tuned to sound location in egocentric and allocentric reference frames can also localize sounds in multiple head and world-centered spaces

Dissociable neural correlates of multisensory coherence and selective attention

Previous work has demonstrated that performance in an auditory selective attention task can be enhanced or impaired, depending on whether a task-irrelevant visual stimulus is temporally coherent with a target auditory stream or with a competing distractor. However, it remains unclear how audiovisual (AV) temporal coherence and auditory selective attention interact at the neurophysiological level. Here, we measured neural activity using electroencephalography (EEG) while human participants (men and women) performed an auditory selective attention task, detecting deviants in a target audio stream. The amplitude envelope of the two competing auditory streams changed independently, while the radius of a visual disc was manipulated to control the audiovisual coherence. Analysis of the neural responses to the sound envelope demonstrated that auditory responses were enhanced independently of the attentional condition: both target and masker stream responses were enhanced when temporally coherent with the visual stimulus. In contrast, attention enhanced the event-related response (ERP) evoked by the transient deviants, independently of AV coherence. Finally, in an exploratory analysis, we identified a spatiotemporal component of ERP, in which temporal coherence enhanced the deviant-evoked responses only in the unattended stream. These results provide evidence for dissociable neural signatures of bottom-up (coherence) and top-down (attention) effects in AV object formation.Significance StatementTemporal coherence between auditory stimuli and task-irrelevant visual stimuli can enhance behavioral performance in auditory selective attention tasks. However, how audiovisual temporal coherence and attention interact at the neural level has not been established. Here, we measured EEG during a behavioral task designed to independently manipulate AV coherence and auditory selective attention. While some auditory features (sound envelope) could be coherent with visual stimuli, other features (timbre) were independent of visual stimuli. We find that audiovisual integration can be observed independently of attention for sound envelopes temporally coherent with visual stimuli, while the neural responses to unexpected timbre changes are most strongly modulated by attention. Our results provide evidence for dissociable neural mechanisms of bottom-up (coherence) and top-down (attention) effects on AV object formation

Behaviourally modulated hippocampal theta oscillations in the ferret persist during both locomotion and immobility

Theta oscillations are a hallmark of hippocampal activity across mammals and play a critical role in many hippocampal models of memory and spatial navigation. To reconcile the cross-species differences observed in the presence and properties of theta, we recorded hippocampal local field potentials in rats and ferrets during auditory and visual localisation tasks designed to vary locomotion and sensory attention. Here, we show that theta oscillations occur during locomotion in both ferrets and rats, however during periods of immobility, theta oscillations persist in the ferret, contrasting starkly with the switch to large irregular activity (LIA) in the rat. Theta during immobility in the ferret is identified as analogous to Type 2 theta that has been observed in rodents due to its sensitivity to atropine, and is modulated by behavioural state with the strongest theta observed during reward epochs. These results demonstrate that even under similar behavioural conditions, differences exist between species in the relationship between theta and behavioural state

Simultaneous Assessment of Speech Identification and Spatial Discrimination: A Potential Testing Approach for Bilateral Cochlear Implant Users?

With increasing numbers of children and adults receiving bilateral cochlear implants, there is an urgent need for assessment tools that enable testing of binaural hearing abilities. Current test batteries are either limited in scope or are of an impractical duration for routine testing. Here, we report a behavioral test that enables combined testing of speech identification and spatial discrimination in noise. In this task, multitalker babble was presented from all speakers, and pairs of speech tokens were sequentially presented from two adjacent speakers. Listeners were required to identify both words from a closed set of four possibilities and to determine whether the second token was presented to the left or right of the first. In Experiment 1, normal-hearing adult listeners were tested at 15° intervals throughout the frontal hemifield. Listeners showed highest spatial discrimination performance in and around the frontal midline, with a decline at more eccentric locations. In contrast, speech identification abilities were least accurate near the midline and showed an improvement in performance at more lateral locations. In Experiment 2, normal-hearing listeners were assessed using a restricted range of speaker locations designed to match those found in clinical testing environments. Here, speakers were separated by 15° around the midline and 30° at more lateral locations. This resulted in a similar pattern of behavioral results as in Experiment 1. We conclude, this test offers the potential to assess both spatial discrimination and the ability to use spatial information for unmasking in clinical populations

Experienced hearing aid users' perspectives of assessment and communication within audiology: a qualitative study using digital methods.

OBJECTIVE: To explore experienced hearing aid users' perspectives of audiological assessments and the patient-audiologist communication dynamic during clinical interactions. DESIGN: A qualitative study was implemented incorporating both an online focus group and online semi-structured interviews. Sessions were audio-recorded and transcribed verbatim. Iterative-inductive thematic analysis was carried out to identify themes related to assessment and communication within audiology practice. STUDY SAMPLES: Seven experienced hearing aid users took part in an online focus group and 14 participated in online semi-structured interviews (age range: 22 - 86 years; 9 males, 11 females). RESULTS: Themes related to assessment included the unaided and aided testing procedure and relating tests to real world hearing difficulties. Themes related to communication included the importance of deaf aware communication strategies, explanation of test results and patient centred care in audiology. CONCLUSION: To ensure hearing aid services meet the needs of the service users, we should explore user perspectives and proactively adapt service delivery. This approach should be ongoing, in response to advances in hearing aid technology. Within audiology, experienced hearing aid users' value (1) comprehensive, relatable hearing assessment, (2) deaf aware patient-audiologist communication, (3) accessible services and (4) a personalised approach to recommend suitable technology and address patient specific aspects of hearing loss

Reversible inactivation of ferret auditory cortex impairs spatial and non-spatial hearing

A key question in auditory neuroscience is to what extent are brain regions functionally specialized for processing specific sound features such as location and identity. In auditory cortex, correlations between neural activity and sounds support both the specialization of distinct cortical subfields, and encoding of multiple sound features within individual cortical areas. However, few studies have tested the contribution of auditory cortex to hearing in multiple contexts. Here we determined the role of ferret primary auditory cortex in both spatial and non-spatial hearing by reversibly inactivating the middle ectosylvian gyrus during behavior using cooling (n=2 females) or optogenetics (n=1 female). Optogenetic experiments utilized the mDLx promoter to express Channelrhodopsin2 in GABAergic interneurons and we confirmed both viral expression (n=2 females) and light-driven suppression of spiking activity in auditory cortex, recorded using Neuropixels under anesthesia (n=465 units from 2 additional untrained female ferrets). Cortical inactivation via cooling or optogenetics impaired vowel discrimination in co-located noise. Ferrets implanted with cooling loops were tested in additional conditions that revealed no deficits for identifying vowels in clean conditions, or when the temporally coincident vowel and noise were spatially separated by 180 degrees. These animals did however show impaired sound localization when inactivating the same auditory cortical region implicated in vowel discrimination in noise. Our results demonstrate that, as a brain region showing mixed selectivity for spatial and non-spatial features of sound, primary auditory cortex contributes to multiple forms of hearing.SIGNIFICANCE STATEMENT:Neurons in primary auditory cortex are often sensitive to the location and identity of sounds. Here we inactivated auditory cortex during spatial and non- spatial listening tasks using cooling, or optogenetics. Auditory cortical inactivation impaired multiple behaviors, demonstrating a role in both the analysis of sound location and identity and confirming a functional contribution of mixed selectivity observed in neural activity. Parallel optogenetic experiments in two additional untrained ferrets linked behavior to physiology by demonstrating that expression of Channelrhodopsin 2 permitted rapid light-driven suppression of auditory cortical activity recorded under anesthesia

The Non-Lemniscal Auditory Cortex in Ferrets: Convergence of Corticotectal Inputs in the Superior Colliculus

Descending cortical inputs to the superior colliculus (SC) contribute to the unisensory response properties of the neurons found there and are critical for multisensory integration. However, little is known about the relative contribution of different auditory cortical areas to this projection or the distribution of their terminals in the SC. We characterized this projection in the ferret by injecting tracers in the SC and auditory cortex. Large pyramidal neurons were labeled in layer V of different parts of the ectosylvian gyrus after tracer injections in the SC. Those cells were most numerous in the anterior ectosylvian gyrus (AEG), and particularly in the anterior ventral field, which receives both auditory and visual inputs. Labeling was also found in the posterior ectosylvian gyrus (PEG), predominantly in the tonotopically organized posterior suprasylvian field. Profuse anterograde labeling was present in the SC following tracer injections at the site of acoustically responsive neurons in the AEG or PEG, with terminal fields being both more prominent and clustered for inputs originating from the AEG. Terminals from both cortical areas were located throughout the intermediate and deep layers, but were most concentrated in the posterior half of the SC, where peripheral stimulus locations are represented. No inputs were identified from primary auditory cortical areas, although some labeling was found in the surrounding sulci. Our findings suggest that higher level auditory cortical areas, including those involved in multisensory processing, may modulate SC function via their projections into its deeper layers

Housing and Environmental Enrichment of the Domestic Ferret: A Multi-Sector Survey

Ferrets (Mustela putorius furo) are kept and used in multiple sectors of society, but little is known about how they are housed and what environmental enrichment (EE) they may benefit from. We aimed to help guide caretakers about what housing and EE can be provided for ferrets. Through an online questionnaire of ferret caretakers, including pet, laboratory, zoological collection, rescue and working animal sectors internationally, we described ferret housing, opportunities for exploration, EE provision and caretaker opinions on ferrets' preferred EE types, and problematic EE. In total, 754 valid responses from 17 countries were analysed, with most (82.4%) coming from pet owners. Most ferrets were housed socially, with housing varying across sectors from single-level cages to free-range housing in a room or outdoor enclosure; pet owners mostly used multi-level cages. The most commonly reported EE included hammocks, tunnels and tactile interaction with caretakers. Respondents reported that ferrets particularly enjoyed digging substrates, tunnels, human interaction and exploration. The most frequently reported problems were that ingestion of unsuitable chew toys and rubber items could cause internal blockages, narrow tunnels could trap ferrets, and certain fabrics that could catch claws. This suggests a need for increased awareness of the risks of these EE types and for more commercially available safety-tested ferret EE. Scent trails were relatively rarely provided but were reported to be enjoyed and harmless, so we recommend that these should be provided more commonly. Our results suggest that there is scope to improve ferret housing and EE provision to benefit ferret welfare across all sectors

The role of temporal coherence and temporal predictability in the build-up of auditory grouping

The cochlea decomposes sounds into separate frequency channels, from which the auditory brain must reconstruct the auditory scene. To do this the auditory system must make decisions about which frequency information should be grouped together, and which should remain distinct. Two key cues for grouping are temporal coherence, resulting from coherent changes in power across frequency, and temporal predictability, resulting from regular or predictable changes over time. To test how these cues contribute to the construction of a sound scene we present listeners with a range of precursor sounds, which act to prime the auditory system by providing information about each sounds structure, followed by a fixed masker in which participants were required to detect the presence of an embedded tone. By manipulating temporal coherence and/or temporal predictability in the precursor we assess how prior sound exposure influences subsequent auditory grouping. In Experiment 1, we measure the contribution of temporal predictability by presenting temporally regular or jittered precursors, and temporal coherence by using either narrow or broadband sounds, demonstrating that both independently contribute to masking/unmasking. In Experiment 2, we measure the relative impact of temporal coherence and temporal predictability and ask whether the influence of each in the precursor signifies an enhancement or interference of unmasking. We observed that interfering precursors produced the largest changes to thresholds

Relative sound localisation abilities in human listeners

Spatial acuity varies with sound-source azimuth, signal-to-noise ratio, and the spectral characteristics of the sound source. Here, the spatial localisation abilities of listeners were assessed using a relative localisation task. This task tested localisation ability at fixed angular separations throughout space using a two-alternative forced-choice design across a variety of listening conditions. Subjects were required to determine whether a target sound originated to the left or right of a preceding reference in the presence of a multi-source noise background. Experiment 1 demonstrated that subjects' ability to determine the relative location of two sources declined with less favourable signal-to-noise ratios and at peripheral locations. Experiment 2 assessed performance with both broadband and spectrally restricted stimuli designed to limit localisation cues to predominantly interaural level differences or interaural timing differences (ITDs). Predictions generated from topographic, modified topographic, and two-channel models of sound localisation suggest that for low-pass stimuli, where ITD cues were dominant, the two-channel model provides an adequate description of the experimental data, whereas for broadband and high frequency bandpass stimuli none of the models was able to fully account for performance. Experiment 3 demonstrated that relative localisation performance was uninfluenced by shifts in gaze direction