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This study investigated the impact of bilateral bone conduction (BC) stimulation and sensorineural hearing loss on spatial release from masking, binaural intelligibility level difference, and lateralization. The study involved two groups of adults with mild to moderate sensorineural hearing loss: one group of 21 participants with symmetric hearing loss and another group of nine participants with asymmetric hearing loss. All tests were conducted through BC and air conduction (AC) headsets with non-individualized virtual positions of the sound sources and linear amplification based on individual hearing thresholds. The findings revealed a bilateral benefit for both groups of hearing-impaired individuals, with symmetric hearing loss yielding better results than asymmetric hearing loss. AC stimulation provided approximately twice the benefit in terms of dB compared to BC stimulation. A large part of this benefit originated from a favorable signal-to-noise ratio due to noise reduction from the head shadow. However, binaural processing was present in both hearing-impaired groups with bilateral BC stimulation. The ability to lateralize sounds based on interaural time delays was significantly impaired in participants with both types of hearing loss when stimulation was by BC. Despite these challenges, the study underscores the benefits of bilateral fitting of BC hearing aids, even in individuals with mild to moderate sensorineural hearing loss, whether symmetric or asymmetric., Competing Interests: Declarations. Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

How we move our bodies affects how we perceive sound. For instance, head movements help us to better localize the source of a sound and to compensate for asymmetric hearing loss. However, many auditory experiments are designed to restrict head and body movements. To study the role of movement in hearing, we developed a behavioral task called sound-seeking that rewarded freely moving mice for tracking down an ongoing sound source. Over the course of learning, mice more efficiently navigated to the sound. Next, we asked how sound-seeking was affected by hearing loss induced by surgical removal of the malleus from the middle ear. After bilateral hearing loss sound-seeking performance drastically declined and did not recover. In striking contrast, after unilateral hearing loss mice were only transiently impaired and then recovered their sound-seek ability over about a week. Throughout recovery, unilateral mice increasingly relied on a movement strategy of sequentially checking potential locations for the sound source. In contrast, the startle reflex (an innate auditory behavior) was preserved after unilateral hearing loss and abolished by bilateral hearing loss without recovery over time. In sum, mice compensate with body movement for permanent unilateral damage to the peripheral auditory system. Looking forward, this paradigm provides an opportunity to examine how movement enhances perception and enables resilient adaptation to sensory disorders., (© 2024. The Author(s).)

In real-world listening situations, individuals typically utilize head and eye movements to receive and enhance sensory information while exploring acoustic scenes. However, the specific patterns of such movements have not yet been fully characterized. Here, we studied how movement behavior is influenced by scene complexity, varied in terms of reverberation and the number of concurrent talkers. Thirteen normal-hearing participants engaged in a speech comprehension and localization task, requiring them to indicate the spatial location of a spoken story in the presence of other stories in virtual audio-visual scenes. We observed delayed initial head movements when more simultaneous talkers were present in the scene. Both reverberation and a higher number of talkers extended the search period, increased the number of fixated source locations, and resulted in more gaze jumps. The period preceding the participants' responses was prolonged when more concurrent talkers were present, and listeners continued to move their eyes in the proximity of the target talker. In scenes with more reverberation, the final head position when making the decision was farther away from the target. These findings demonstrate that the complexity of the acoustic scene influences listener behavior during speech comprehension and localization in audio-visual scenes., (© 2024. The Author(s).)

Locating sound sources such as prey or predators is critical for survival in many vertebrates. Terrestrial vertebrates locate sources by measuring the time delay and intensity difference of sound pressure at each ear 1-5 . Underwater, however, the physics of sound makes interaural cues very small, suggesting that directional hearing in fish should be nearly impossible 6 . Yet, directional hearing has been confirmed behaviourally, although the mechanisms have remained unknown for decades. Several hypotheses have been proposed to explain this remarkable ability, including the possibility that fish evolved an extreme sensitivity to minute interaural differences or that fish might compare sound pressure with particle motion signals 7,8 . However, experimental challenges have long hindered a definitive explanation. Here we empirically test these models in the transparent teleost Danionella cerebrum, one of the smallest vertebrates 9,10 . By selectively controlling pressure and particle motion, we dissect the sensory algorithm underlying directional acoustic startles. We find that both cues are indispensable for this behaviour and that their relative phase controls its direction. Using micro-computed tomography and optical vibrometry, we further show that D. cerebrum has the sensory structures to implement this mechanism. D. cerebrum shares these structures with more than 15% of living vertebrate species, suggesting a widespread mechanism for inferring sound direction., (© 2024. The Author(s).)

Sound localization is essential to perceive the surrounding world and to interact with objects. This ability can be learned across time, and multisensory and motor cues play a crucial role in the learning process. A recent study demonstrated that when training localization skills, reaching to the sound source to determine its position reduced localization errors faster and to a greater extent as compared to just naming sources' positions, despite the fact that in both tasks, participants received the same feedback about the correct position of sound sources in case of wrong response. However, it remains to establish which features have made reaching to sound more effective as compared to naming. In the present study, we introduced a further condition in which the hand is the effector providing the response, but without it reaching toward the space occupied by the target source: the pointing condition. We tested three groups of participants (naming, pointing, and reaching groups) each while performing a sound localization task in normal and altered listening situations (i.e. mild-moderate unilateral hearing loss) simulated through auditory virtual reality technology. The experiment comprised four blocks: during the first and the last block, participants were tested in normal listening condition, while during the second and the third in altered listening condition. We measured their performance, their subjective judgments (e.g. effort), and their head-related behavior (through kinematic tracking). First, people's performance decreased when exposed to asymmetrical mild-moderate hearing impairment, more specifically on the ipsilateral side and for the pointing group. Second, we documented that all groups decreased their localization errors across altered listening blocks, but the extent of this reduction was higher for reaching and pointing as compared to the naming group. Crucially, the reaching group leads to a greater error reduction for the side where the listening alteration was applied. Furthermore, we documented that, across blocks, reaching and pointing groups increased the implementation of head motor behavior during the task (i.e., they increased approaching head movements toward the space of the sound) more than naming. Third, while performance in the unaltered blocks (first and last) was comparable, only the reaching group continued to exhibit a head behavior similar to those developed during the altered blocks (second and third), corroborating the previous observed relationship between the reaching to sounds task and head movements. In conclusion, this study further demonstrated the effectiveness of reaching to sounds as compared to pointing and naming in the learning processes. This effect could be related both to the process of implementing goal-directed motor actions and to the role of reaching actions in fostering the implementation of head-related motor strategies., (© 2024. The Author(s).)

The effects of music on bodily movement and feelings, such as when people are dancing or engaged in physical activity, are well-documented-people may move in response to the sound cues, feel powerful, less tired. How sounds and bodily movements relate to create such effects? Here we deconstruct the problem and investigate how different auditory features affect people's body-representation and feelings even when paired with the same movement. In three experiments, participants executed a simple arm raise synchronised with changing pitch in simple tones (Experiment 1), rich musical sounds (Experiment 2) and within different frequency ranges (Experiment 3), while we recorded indirect and direct measures on their movement, body-representations and feelings. Changes in pitch influenced people's general emotional state as well as the various bodily dimensions investigated-movement, proprioceptive awareness and feelings about one's body and movement. Adding harmonic content amplified the differences between ascending and descending sounds, while shifting the absolute frequency range had a general effect on movement amplitude, bodily feelings and emotional state. These results provide new insights in the role of auditory and musical features in dance and exercise, and have implications for the design of sound-based applications supporting movement expression, physical activity, or rehabilitation., (© 2022. The Author(s).)

Tinnitus therapies have been combined with the use of varieties of sound/noise. For masking external sounds, location of the masker in space is important; however, effects of the spatial location of the masker on tinnitus are less understood. We aimed to test whether a masking sound location would affect the perception level of simulated tinnitus. The 4 kHz simulated tinnitus was induced in the right ear of healthy volunteers through an open-type earphone. White noise was presented to the right ear using a single-sided headphone or a speaker positioned on the right side at a distance of 1.8 m for masking the simulated tinnitus. In other sessions, monaurally recorded noise localized within the head (inside-head noise) or binaurally recorded noise localized outside the head (outside-head noise) was separately presented from a dual-sided headphone. The noise presented from a distant speaker and the outside-head noise masked the simulated tinnitus in 71.1% and 77.1% of measurements at a lower intensity compared to the noise beside the ear and the inside-head noise, respectively. In conclusion, spatial information regarding the masking noise may play a role in reducing the perception level of simulated tinnitus. Binaurally recorded sounds may be beneficial for an acoustic therapy of tinnitus., (© 2022. The Author(s).)

Peripersonal space (PPS) is important for humans to perform body-environment interactions. However, many previous studies only focused on the specific direction of the PPS, such as the front space, despite suggesting that there were PPSs in all directions. We aimed to measure and compare the peri-trunk PPS in four directions (front, rear, left, and right). To measure the PPS, we used a tactile and an audio stimulus because auditory information is available at any time in all directions. We used the approaching and receding task-irrelevant sounds in the experiment. Observers were asked to respond as quickly as possible when a tactile stimulus was applied to a vibrator on their chest. We found that peri-trunk PPS representations exist with an approaching sound, irrespective of the direction.

Echolocating bats adapt their emissions to succeed in noisy environments. In the present study we investigated if echolocating humans can detect a sound-reflecting surface in the presence of noise and if intensity of echolocation emissions (i.e. clicks) changes in a systematic pattern. We tested people who were blind and had experience in echolocation, as well as blind and sighted people who had no experience in echolocation prior to the study. We used an echo-detection paradigm where participants listened to binaural recordings of echolocation sounds (i.e. they did not make their own click emissions), and where intensity of emissions and echoes changed adaptively based on participant performance (intensity of echoes was yoked to intensity of emissions). We found that emission intensity had to systematically increase to compensate for weaker echoes relative to background noise. In fact, emission intensity increased so that spectral power of echoes exceeded spectral power of noise by 12 dB in 4-kHz and 5-kHz frequency bands. The effects were the same across all participant groups, suggesting that this effect occurs independently of long-time experience with echolocation. Our findings demonstrate for the first time that people can echolocate in the presence of noise and suggest that one potential strategy to deal with noise is to increase emission intensity to maintain signal-to-noise ratio of certain spectral components of the echoes.

Auditory roughness elicits aversion, and higher activation in cerebral areas involved in threat processing, but its link with defensive behavior is unknown. Defensive behaviors are triggered by intrusions into the space immediately surrounding the body, called peripersonal space (PPS). Integrating multisensory information in PPS is crucial to assure the protection of the body. Here, we assessed the behavioral effects of roughness on auditory-tactile integration, which reflects the monitoring of this multisensory region of space. Healthy human participants had to detect as fast as possible a tactile stimulation delivered on their hand while an irrelevant sound was approaching them from the rear hemifield. The sound was either a simple harmonic sound or a rough sound, processed through binaural rendering so that the virtual sound source was looming towards participants. The rough sound speeded tactile reaction times at a farther distance from the body than the non-rough sound. This indicates that PPS, as estimated here via auditory-tactile integration, is sensitive to auditory roughness. Auditory roughness modifies the behavioral relevance of simple auditory events in relation to the body. Even without emotional or social contextual information, auditory roughness constitutes an innate threat cue that elicits defensive responses.

Hearing aid and cochlear implant (CI) users often struggle to locate and segregate sounds. The dominant sound-localisation cues are time and intensity differences across the ears. A recent study showed that CI users locate sounds substantially better when these cues are provided through haptic stimulation on each wrist. However, the sensitivity of the wrists to these cues and the robustness of this sensitivity to aging is unknown. The current study showed that time difference sensitivity is much poorer across the wrists than across the ears and declines with age. In contrast, high sensitivity to across-wrist intensity differences was found that was robust to aging. This high sensitivity was observed across a range of stimulation intensities for both amplitude modulated and unmodulated sinusoids and matched across-ear intensity difference sensitivity for normal-hearing individuals. Furthermore, the usable dynamic range for haptic stimulation on the wrists was found to be around four times larger than for CIs. These findings suggest that high-precision haptic sound-localisation can be achieved, which could aid many hearing-impaired listeners. Furthermore, the finding that high-fidelity across-wrist intensity information can be transferred could be exploited in human-machine interfaces to enhance virtual reality and improve remote control of military, medical, or research robots.

Before the onset of hearing, cochlea-generated patterns of spontaneous spike activity drive the maturation of central auditory circuits. In the glycinergic sound localization pathway from the medial nucleus of the trapezoid body (MNTB) to the lateral superior olive (LSO) this spontaneous activity guides the strengthening and silencing of synapses which underlies tonotopic map refinement. However, the mechanisms by which patterned activity regulates synaptic refinement in the MNTB-LSO pathway are still poorly understood. To address this question, we recorded from LSO neurons in slices from prehearing mice while stimulating MNTB afferents with stimulation patterns that mimicked those present in vivo. We found that these semi-natural stimulation patterns reliably elicited a novel form of long-term potentiation (LTP) of MNTB-LSO synapses. Stimulation patterns that lacked the characteristic high-frequency (200 Hz) component of prehearing spike activity failed to elicit potentiation. LTP was calcium dependent, required the activation of both g-protein coupled GABA B and metabotropic glutamate receptors and involved an increase in postsynaptic glycine receptor-mediated currents. Our results provide a possible mechanism linking spontaneous spike bursts to tonotopic map refinement and further highlight the importance of the co-release of GABA and glutamate from immature glycinergic MNTB terminals.

Users of hearing-assistive devices often struggle to locate and segregate sounds, which can make listening in schools, cafes, and busy workplaces extremely challenging. A recent study in unilaterally implanted CI users showed that sound-localisation was improved when the audio received by behind-the-ear devices was converted to haptic stimulation on each wrist. We built on this work, using a new signal-processing approach to improve localisation accuracy and increase generalisability to a wide range of stimuli. We aimed to: (1) improve haptic sound-localisation accuracy using a varied stimulus set and (2) assess whether accuracy improved with prolonged training. Thirty-two adults with normal touch perception were randomly assigned to an experimental or control group. The experimental group completed a 5-h training regime and the control group were not trained. Without training, haptic sound-localisation was substantially better than in previous work on haptic sound-localisation. It was also markedly better than sound-localisation by either unilaterally or bilaterally implanted CI users. After training, accuracy improved, becoming better than for sound-localisation by bilateral hearing-aid users. These findings suggest that a wrist-worn haptic device could be effective for improving spatial hearing for a range of hearing-impaired listeners.

Adaptation to systematic visual distortions is well-documented but there is little evidence of similar adaptation to radical changes in audition. We use a pseudophone to transpose the sound streams arriving at the left and right ears, evaluating the perceptual effects it provokes and the possibility of learning to locate sounds in the reversed condition. Blindfolded participants remain seated at the center of a semicircular arrangement of 7 speakers and are asked to orient their head towards a sound source. We postulate that a key factor underlying adaptation is the self-generated activity that allows participants to learn new sensorimotor schemes. We investigate passive listening conditions (very short duration stimulus not permitting active exploration) and dynamic conditions (continuous stimulus allowing participants time to freely move their heads or remain still). We analyze head movement kinematics, localization errors, and qualitative reports. Results show movement-induced perceptual disruptions in the dynamic condition with static sound sources displaying apparent movement. This effect is reduced after a short training period and participants learn to find sounds in a left-right reversed field for all but the extreme lateral positions where motor patterns are more restricted. Strategies become less exploratory and more direct with training. Results support the hypothesis that self-generated movements underlie adaptation to radical sensorimotor distortions.

Cochlear implants (CIs) have enabled hundreds of thousands of profoundly hearing-impaired people to perceive sounds by electrically stimulating the auditory nerve. However, CI users are often very poor at locating sounds, which leads to impaired sound segregation and threat detection. We provided missing spatial hearing cues through haptic stimulation to augment the electrical CI signal. We found that this "electro-haptic" stimulation dramatically improved sound localisation. Furthermore, participants were able to effectively integrate spatial information transmitted through these two senses, performing better with combined audio and haptic stimulation than with either alone. Our haptic signal was presented to the wrists and could readily be delivered by a low-cost wearable device. This approach could provide a non-invasive means of improving outcomes for the vast majority of CI users who have only one implant, without the expense and risk of a second implantation.

We analyse the effects of exploration feedback on reaching measures of perceived auditory peripersonal space (APS) boundary and the auditory distance perception (ADP) of sound sources located within it. We conducted an experiment in which the participants had to estimate if a sound source was (or not) reachable and to estimate its distance (40 to 150 cm in 5-cm steps) by reaching to a small loudspeaker. The stimulus consisted of a train of three bursts of Gaussian broadband noise. Participants were randomly assigned to two groups: Experimental (EG) and Control (CG). There were three phases in the following order: Pretest-Test-Posttest. For all phases, the listeners performed the same task except for the EG-Test phase where the participants reach in order to touch the sound source. We applied models to characterise the participants' responses and provide evidence that feedback significantly reduces the response bias of both the perceived boundary of the APS and the ADP of sound sources located within reach. In the CG, the repetition of the task did not affect APS and ADP accuracy, but it improved the performance consistency: the reachable uncertainty zone in APS was reduced and there was a tendency to decrease variability in ADP.

Although sound position is initially head-centred (egocentric coordinates), our brain can also represent sounds relative to one another (allocentric coordinates). Whether reference frames for spatial hearing are independent or interact remained largely unexplored. Here we developed a new allocentric spatial-hearing training and tested whether it can improve egocentric sound-localisation performance in normal-hearing adults listening with one ear plugged. Two groups of participants (N = 15 each) performed an egocentric sound-localisation task (point to a syllable), in monaural listening, before and after 4-days of multisensory training on triplets of white-noise bursts paired with occasional visual feedback. Critically, one group performed an allocentric task (auditory bisection task), whereas the other processed the same stimuli to perform an egocentric task (pointing to a designated sound of the triplet). Unlike most previous works, we tested also a no training group (N = 15). Egocentric sound-localisation abilities in the horizontal plane improved for all groups in the space ipsilateral to the ear-plug. This unexpected finding highlights the importance of including a no training group when studying sound localisation re-learning. Yet, performance changes were qualitatively different in trained compared to untrained participants, providing initial evidence that allocentric and multisensory procedures may prove useful when aiming to promote sound localisation re-learning.

How the human auditory system learns to map complex pinna-induced spectral-shape cues onto veridical estimates of sound-source elevation in the median plane is still unclear. Earlier studies demonstrated considerable sound-localisation plasticity after applying pinna moulds, and to altered vision. Several factors may contribute to auditory spatial learning, like visual or motor feedback, or updated priors. We here induced perceptual learning for sounds with degraded spectral content, having weak, but consistent, elevation-dependent cues, as demonstrated by low-gain stimulus-response relations. During training, we provided visual feedback for only six targets in the midsagittal plane, to which listeners gradually improved their response accuracy. Interestingly, listeners' performance also improved without visual feedback, albeit less strongly. Post-training results showed generalised improved response behaviour, also to non-trained locations and acoustic spectra, presented throughout the two-dimensional frontal hemifield. We argue that the auditory system learns to reweigh contributions from low-informative spectral bands to update its prior elevation estimates, and explain our results with a neuro-computational model.

Sensory representations are typically endowed with intrinsic noise, leading to variability and inaccuracies in perceptual responses. The Bayesian framework accounts for an optimal strategy to deal with sensory-motor uncertainty, by combining the noisy sensory input with prior information regarding the distribution of stimulus properties. The maximum-a-posteriori (MAP) estimate selects the perceptual response from the peak (mode) of the resulting posterior distribution that ensure optimal accuracy-precision trade-off when the underlying distributions are Gaussians (minimal mean-squared error, with minimum response variability). We tested this model on human eye- movement responses toward broadband sounds, masked by various levels of background noise, and for head movements to sounds with poor spectral content. We report that the response gain (accuracy) and variability (precision) of the elevation response components changed systematically with the signal-to-noise ratio of the target sound: gains were high for high SNRs and decreased for low SNRs. In contrast, the azimuth response components maintained high gains for all conditions, as predicted by maximum-likelihood estimation. However, we found that the elevation data did not follow the MAP prediction. Instead, results were better described by an alternative decision strategy, in which the response results from taking a random sample from the posterior in each trial. We discuss two potential implementations of a simple posterior sampling scheme in the auditory system that account for the results and argue that although the observed response strategies for azimuth and elevation are sub-optimal with respect to their variability, it allows the auditory system to actively explore the environment in the absence of adequate sensory evidence.

This study investigated the influence of body motion on an echolocation task. We asked a group of blindfolded novice sighted participants to walk along a corridor, made with plastic sound-reflecting panels. By self-generating mouth clicks, the participants attempted to understand some spatial properties of the corridor, i.e. a left turn, a right turn or a dead end. They were asked to explore the corridor and stop whenever they were confident about the corridor shape. Their body motion was captured by a camera system and coded. Most participants were able to accomplish the task with the percentage of correct guesses above the chance level. We found a mutual interaction between some kinematic variables that can lead to optimal echolocation skills. These variables are head motion, accounting for spatial exploration, the motion stop-point of the person and the amount of correct guesses about the spatial structure. The results confirmed that sighted people are able to use self-generated echoes to navigate in a complex environment. The inter-individual variability and the quality of echolocation tasks seems to depend on how and how much the space is explored.

Acute auditory processing in bats is typically associated with echolocation. A subset of bats, called gleaners, listens to prey-generated noise to hunt surface-dwelling prey. Gleaners depend less on echolocation to hunt and, therefore, accurate localization of prey-generated noise is necessary for foraging success. Here we studied azimuth sound localization behavior in the pallid bat, a gleaning bat in which spatial encoding has been studied extensively. We tested pallid bats on a relatively difficult open loop task (single sound, duration ≤ 200 ms). The bats were trained to face the midline when stimulus was presented, and this was confirmed with video analysis. Bats localized broadband noise (5-30 kHz) from 1 out of 11 speakers spaced evenly across the horizontal plane of the frontal sound field. Approach to the correct speaker was rewarded. Pallid bats show accurate localization near the midline with mean errors between 3-6°. Remarkably, the accuracy does not decline significantly at peripheral locations with bats averaging  <~7° error upto 72° off midline. Manipulation of stimulus bandwidth shows that higher frequencies (20-30 kHz) are necessary for accurate localization. Comparative studies of gleaning bats will reveal convergent adaptations across auditory systems for non-echolocation-based behaviors in bats.

Speech recognition starts with representations of basic acoustic perceptual features and ends by categorizing the sound based on long-term memory for word meaning. However, little is known about whether the reverse pattern of lexical influences on basic perception can occur. We tested for a lexical influence on auditory spatial perception by having subjects make spatial judgments of number stimuli. Four experiments used pointing or left/right 2-alternative forced choice tasks to examine perceptual judgments of sound location as a function of digit magnitude (1-9). The main finding was that for stimuli presented near the median plane there was a linear left-to-right bias for localizing smaller-to-larger numbers. At lateral locations there was a central-eccentric location bias in the pointing task, and either a bias restricted to the smaller numbers (left side) or no significant number bias (right side). Prior number location also biased subsequent number judgments towards the opposite side. Findings support a lexical influence on auditory spatial perception, with a linear mapping near midline and more complex relations at lateral locations. Results may reflect coding of dedicated spatial channels, with two representing lateral positions in each hemispace, and the midline area represented by either their overlap or a separate third channel.

Echolocation is the ability to use sound-echoes to infer spatial information about the environment. People can echolocate for example by making mouth clicks. Previous research suggests that echolocation in blind people activates brain areas that process light in sighted people. Research has also shown that echolocation in blind people may replace vision for calibration of external space. In the current study we investigated if echolocation may also draw on 'visual' resources in the sighted brain. To this end, we paired a sensory interference paradigm with an echolocation task. We found that exposure to an uninformative visual stimulus (i.e. white light) while simultaneously echolocating significantly reduced participants' ability to accurately judge object size. In contrast, a tactile stimulus (i.e. vibration on the skin) did not lead to a significant change in performance (neither in sighted, nor blind echo expert participants). Furthermore, we found that the same visual stimulus did not affect performance in auditory control tasks that required detection of changes in sound intensity, sound frequency or sound location. The results suggest that processing of visual and echo-acoustic information draw on common neural resources.

A natural auditory scene often contains sound moving at varying velocities. Using a velocity contrast paradigm, we compared sensitivity to velocity changes between continuous and discontinuous trajectories. Subjects compared the velocities of two stimulus intervals that moved along a single trajectory, with and without a 1 second inter stimulus interval (ISI). We found thresholds were threefold larger for velocity increases in the instantaneous velocity change condition, as compared to instantaneous velocity decreases or thresholds for the delayed velocity transition condition. This result cannot be explained by the current static "snapshot" model of auditory motion perception and suggest a continuous process where the percept of velocity is influenced by previous history of stimulation.

Spatial hearing evolved independently in mammals and birds and is thought to adapt to altered developmental input in different ways. We found, however, that ferrets possess multiple forms of plasticity that are expressed according to which spatial cues are available, suggesting that the basis for adaptation may be similar across species. Our results also provide insight into the way sound source location is represented by populations of cortical neurons.

Across all sensory modalities, the effect of context-dependent neural adaptation can be observed at every level, from receptors to perception. Nonetheless, it has long been assumed that the processing of interaural time differences, which is the primary cue for sound localization, is nonadaptive, as its outputs are mapped directly onto a hard-wired representation of space. Here we present evidence derived from in vitro and in vivo experiments in gerbils indicating that the coincidence-detector neurons in the medial superior olive modulate their sensitivity to interaural time differences through a rapid, GABA(B) receptor-mediated feedback mechanism. We show that this mechanism provides a gain control in the form of output normalization, which influences the neuronal population code of auditory space. Furthermore, psychophysical tests showed that the paradigm used to evoke neuronal GABA(B) receptor-mediated adaptation causes the perceptual shift in sound localization in humans that was expected on the basis of our physiological results in gerbils.

The owl captures prey using sound localization. In the classical model, the owl infers sound direction from the position of greatest activity in a brain map of auditory space. However, this model fails to describe the actual behavior. Although owls accurately localize sources near the center of gaze, they systematically underestimate peripheral source directions. We found that this behavior is predicted by statistical inference, formulated as a Bayesian model that emphasizes central directions. We propose that there is a bias in the neural coding of auditory space, which, at the expense of inducing errors in the periphery, achieves high behavioral accuracy at the ethologically relevant range. We found that the owl's map of auditory space decoded by a population vector is consistent with the behavioral model. Thus, a probabilistic model describes both how the map of auditory space supports behavior and why this representation is optimal.

Activity in the primary auditory cortex (A1) is essential for normal sound localization behavior, but previous studies of the spatial sensitivity of neurons in A1 have found broad spatial tuning. We tested the hypothesis that spatial tuning sharpens when an animal engages in an auditory task. Cats performed a task that required evaluation of the locations of sounds and one that required active listening, but in which sound location was irrelevant. Some 26-44% of the units recorded in A1 showed substantially sharpened spatial tuning during the behavioral tasks as compared with idle conditions, with the greatest sharpening occurring during the location-relevant task. Spatial sharpening occurred on a scale of tens of seconds and could be replicated multiple times in ∼1.5-h test sessions. Sharpening resulted primarily from increased suppression of responses to sounds at least-preferred locations. That and an observed increase in latencies suggest an important role of inhibitory mechanisms.

Descending projections from sensory areas of the cerebral cortex are among the largest pathways in the brain, suggesting that they are important for subcortical processing. Although corticofugal inputs have been shown to modulate neuronal responses in the thalamus and midbrain, the behavioral importance of these changes remains unknown. In the auditory system, one of the major descending pathways is from cortical layer V pyramidal cells to the inferior colliculus in the midbrain. We examined the role of these neurons in experience-dependent recalibration of sound localization in adult ferrets by selectively killing the neurons using chromophore-targeted laser photolysis. When provided with appropriate training, animals normally relearn to localize sound accurately after altering the spatial cues available by reversibly occluding one ear. However, this ability was lost after eliminating corticocollicular neurons, whereas normal sound-localization accuracy was unaffected. The integrity of this descending pathway is therefore critical for learning-induced localization plasticity.

During the maintenance of visuospatial information, neural activity in the frontal eye field (FEF) persists and is thought to be an important neural mechanism for visual working memory. We used functional magnetic resonance imaging to examine whether human FEF activity persists when maintaining auditory space and whether it is selective for retinal versus extra-retinal space. Subjects performed an audiospatial working-memory task using sounds recorded from microphones placed in each subject's ear canals, which preserved the interaural time and level differences that are critical for sound localization. Putative FEF activity persisted when maintaining auditory-cued space, even for locations behind the head to which it is impossible to make saccades. Therefore, human FEF activity represents both retinal and extra-retinal space.

Studies of auditory cortex are often driven by the assumption, derived from our better understanding of visual cortex, that basic physical properties of sounds are represented there before being used by higher-level areas for determining sound-source identity and location. However, we only have a limited appreciation of what the cortex adds to the extensive subcortical processing of auditory information, which can account for many perceptual abilities. This is partly because of the approaches that have dominated the study of auditory cortical processing to date, and future progress will unquestionably profit from the adoption of methods that have provided valuable insights into the neural basis of visual perception. At the same time, we propose that there are unique operating principles employed by the auditory cortex that relate largely to the simultaneous and sequential processing of previously derived features and that therefore need to be studied and understood in their own right.

Although ears capable of detecting airborne sound have arisen repeatedly and independently in different species, most animals that are capable of hearing have a pair of ears. We review the advantages that arise from having two ears and discuss recent research on the similarities and differences in the binaural processing strategies adopted by birds and mammals. We also ask how these different adaptations for binaural and spatial hearing might inform and inspire the development of techniques for future auditory prosthetic devices.

Studies of cortical connections or neuronal function in different cerebral areas support the hypothesis that parallel cortical processing streams, similar to those identified in visual cortex, may exist in the auditory system. However, this model has not yet been behaviorally tested. We used reversible cooling deactivation to investigate whether the individual regions in cat nonprimary auditory cortex that are responsible for processing the pattern of an acoustic stimulus or localizing a sound in space could be doubly dissociated in the same animal. We found that bilateral deactivation of the posterior auditory field resulted in deficits in a sound-localization task, whereas bilateral deactivation of the anterior auditory field resulted in deficits in a pattern-discrimination task, but not vice versa. These findings support a model of cortical organization that proposes that identifying an acoustic stimulus ('what') and its spatial location ('where') are processed in separate streams in auditory cortex.

Neurons in the lateral superior olive (LSO) compute sound location based on differences in interaural intensity, coded in ascending signals from the two cochleas. Unilateral destruction of the neuronal feedback from the LSO to the cochlea, the lateral olivocochlear efferents, disrupted the normal interaural correlation in response amplitudes to sounds of equal intensity. Thus, lateral olivocochlear feedback maintains the binaural balance in neural excitability required for accurate localization of sounds in space.

The optic tectum of the barn owl contains a map of auditory space. We found that, in response to moving sounds, the locations of receptive fields that make up the map shifted toward the approaching sound. The magnitude of the receptive field shifts increased systematically with increasing stimulus velocity and, therefore, was appropriate to compensate for sensory and motor delays inherent to auditory orienting behavior. Thus, the auditory space map is not static, but shifts adaptively and dynamically in response to stimulus motion. We provide a computational model to account for these results. Because the model derives predictive responses from processes that are known to occur commonly in neural networks, we hypothesize that analogous predictive responses will be found to exist widely in the central nervous system. This hypothesis is consistent with perceptions of stimulus motion in humans for many sensory parameters.

Interaural time difference (ITD) is a critical cue to sound-source localization. Traditional models assume that sounds leading at one ear, and perceived on that side, are processed in the opposite midbrain. Using functional magnetic resonance imaging we demonstrate that as the ITDs of sounds increase, midbrain activity can switch sides, even though perceived location remains on the same side. The data require a new model for human ITD processing.

High-level circuits in the brain that control the direction of gaze are intimately linked with the control of visual spatial attention. Immediately before an animal directs its gaze towards a stimulus, both psychophysical sensitivity to that visual stimulus and the responsiveness of high-order neurons in the cerebral cortex that represent the stimulus increase dramatically. Equivalent effects on behavioural sensitivity and neuronal responsiveness to visual stimuli result from focal electrical microstimulation of gaze control centres in monkeys. Whether the gaze control system modulates neuronal responsiveness in sensory modalities other than vision is unknown. Here we show that electrical microstimulation applied to gaze control circuitry in the forebrain of barn owls regulates the gain of midbrain auditory responses in an attention-like manner. When the forebrain circuit was activated, midbrain responses to auditory stimuli at the location encoded by the forebrain site were enhanced and spatial selectivity was sharpened. The same stimulation suppressed responses to auditory stimuli represented at other locations in the midbrain map. Such space-specific, top-down regulation of auditory responses by gaze control circuitry in the barn owl suggests that the central nervous system uses a common strategy for dynamically regulating sensory gain that applies across modalities, brain areas and classes of vertebrate species. This approach provides a path for discovering mechanisms that underlie top-down gain control in the central nervous system.

Mammals can hear sounds extending over a vast range of sound levels with remarkable accuracy. How auditory neurons code sound level over such a range is unclear; firing rates of individual neurons increase with sound level over only a very limited portion of the full range of hearing. We show that neurons in the auditory midbrain of the guinea pig adjust their responses to the mean, variance and more complex statistics of sound level distributions. We demonstrate that these adjustments improve the accuracy of the neural population code close to the region of most commonly occurring sound levels. This extends the range of sound levels that can be accurately encoded, fine-tuning hearing to the local acoustic environment.

Neurons are often considered to be the computational engines of the brain, with synapses acting solely as conveyers of information. But the diverse types of synaptic plasticity and the range of timescales over which they operate suggest that synapses have a more active role in information processing. Long-term changes in the transmission properties of synapses provide a physiological substrate for learning and memory, whereas short-term changes support a variety of computations. By expressing several forms of synaptic plasticity, a single neuron can convey an array of different signals to the neural circuit in which it operates.

The recognition and localization of sound signals is fundamental to acoustic communication. Complex neural mechanisms are thought to underlie the processing of species-specific sound patterns even in animals with simple auditory pathways. In female crickets, which orient towards the male's calling song, current models propose pattern recognition mechanisms based on the temporal structure of the song. Furthermore, it is thought that localization is achieved by comparing the output of the left and right recognition networks, which then directs the female to the pattern that most closely resembles the species-specific song. Here we show, using a highly sensitive method for measuring the movements of female crickets, that when walking and flying each sound pulse of the communication signal releases a rapid steering response. Thus auditory orientation emerges from reactive motor responses to individual sound pulses. Although the reactive motor responses are not based on the song structure, a pattern recognition process may modulate the gain of the responses on a longer timescale. These findings are relevant to concepts of insect auditory behaviour and to the development of biologically inspired robots performing cricket-like auditory orientation.

We do not yet know why whales occasionally strand after sonar has been deployed nearby, but such information is important for both naval undersea activities and the protection of marine mammals. Jepson et al. suggest that a peculiar gas-forming disease afflicting some stranded cetaceans could be a type of decompression sickness (DCS) resulting from exposure to mid-range sonar. However, neither decompression theory nor observation support the existence of a naturally occurring DCS in whales that is characterized by encapsulated, gas-filled cavities in the liver. Although gas-bubble formation may be aggravated by acoustic energy, more rigorous investigation is needed before sonar can be firmly linked to bubble formation in whales.

Auditory localization relies on the detection and interpretation of acoustic cues that change in value as the head and external ears grow. Here we show that the maturation of these structures is an important determinant for the development of spatial selectivity in the ferret auditory cortex. Spatial response fields (SRFs) of high-frequency cortical neurons recorded at postnatal days (P) 33-39 were broader, and transmitted less information about stimulus direction, than in older ferrets. They also exhibited slightly broader frequency tuning than neurons recorded in adult animals. However, when infant neurons were stimulated through virtual ears of adults, SRFs sharpened significantly and the amount of transmitted information increased. This improvement was predicted by a model that generates SRF shape from the localization cue values and the neurons' binaural spectrotemporal response properties. The maturation of spatial response characteristics in auditory cortex therefore seems to be limited by peripheral rather than by central factors.

Synapse elimination and strengthening are central mechanisms for the developmental organization of excitatory neuronal networks. Little is known, however, about whether these processes are also involved in establishing precise inhibitory circuits. We examined the development of functional connectivity before hearing onset in rats in the tonotopically organized, glycinergic pathway from the medial nucleus of the trapezoid body (MNTB) to the lateral superior olive (LSO), which is part of the mammalian sound localization system. We found that LSO neurons became functionally disconnected from approximately 75% of their initial inputs, resulting in a two-fold sharpening of functional topography. This was accompanied by a 12-fold increase in the synaptic conductance generated by maintained individual inputs. Functional elimination of MNTB-LSO synapses was restricted to the period when these glycinergic/GABAergic synapses are excitatory. These results provide new insights into the mechanisms by which precisely organized inhibitory circuits are established during development.

Auditory and visual target locations are encoded differently in the brain, but must be co-calibrated to maintain cross-sensory concordance. Mechanisms that adjust spatial calibration across modalities have been described (for example, prism adaptation in owls), though rudimentarily in humans. We quantified the adaptation of human sound localization in response to spatially compressed vision (0.5x lenses for 2-3 days). This induced a corresponding compression of auditory localization that was most pronounced for azimuth (minimal for elevation) and was restricted to the visual field of the lenses. Sound localization was also affected outside the field of visual-auditory interaction (shifted centrally, not compressed). These results suggest that spatially modified vision induces adaptive changes in adult human sound localization, including novel mechanisms that account for spatial compression. Findings are consistent with a model in which the central processing of sound location is encoded by recruitment rather than by a place code.