Relearning sound localization with new ears

Because the inner ear is not organized spatially, sound localization relies on the neural processing of implicit acoustic cues. To determine a sound's position, the brain must learn and calibrate these cues, using accurate spatial feedback from other sensorimotor systems. Experimental evidence for such a system has been demonstrated in barn owls, but not in humans. Here, we demonstrate the existence of ongoing spatial calibration in the adult human auditory system. The spectral elevation cues of human subjects were disrupted by modifying their outer ears (pinnae) with molds. Although localization of sound elevation was dramatically degraded immediately after the modification, accurate performance was steadily reacquired. Interestingly, learning the new spectral cues did not interfere with the neural representation of the original cues, as subjects could localize sounds with both normal and modified pinnae.     

Spectral cues explain illusory elevation effects with stereo sounds in cats

Mammals localize sound sources in azimuth based on two binaural cues, interaural differences in the time of arrival and level of the sounds at the ears. In contrast, the cue for elevation is based on patterns of the broadband power spectra at each ear that result from the direction-dependent acoustic filtering properties of the head and pinnae. Although the exact form of this "spectral shape" cue is unknown, most attention has been directed toward a prominent direction-dependent energy minimum, or "notch," because its location in frequency, for both humans and cats, moves predictably from low to high as a source is moved from low to high elevations. However, there is little direct evidence that these spectral notches are important elevational cues in animals other than humans. Here we demonstrate a striking illusion in the localization of sounds in elevation by cats using stimulus configurations that elicit summing localization and the precedence effect that can be explained by spectral shape cues.     

Mismatch negativity on the cone of confusion

Localization of sounds by the auditory system is based on the analysis of three sources of information: interaural level differences (ILD, caused by an attenuation of the sound as it travels to the more distant ear), interaural time differences (ITD, caused by the additional amount of time it takes for the sound to arrive at the more distant ear), and spectral cues (caused by direction-specific spectral filter properties of the pinnae). Although in a number of psychophysiological studies cortical processes of ITD and ILD analysis were investigated, there is hitherto no evidence on the cortical processing of spectral cues for sound localization. The objective of the present experiment was to test whether it is possible to observe electrophysiological correlates of sound localization based on spectral cues. In an auditory oddball experiment, 80 ms of broadband noise from varying free field locations were presented to inattentive participants. Mismatch negativities (MMNs) were observed for pairs of standards and location deviants located symmetrically with respect to the interaural axis. As interaural time and level differences are identical for such pairs of sounds, the observed MMNs most likely reflect cognitive processes of sound localization utilizing the spectral filter properties of the pinnae. MMN latencies suggest that sound localization based on spectral cues is slower than ITD- or ILD-based localization.     

Evidence for a vestigial pinna‐orienting system in humans

Although some people can voluntarily move their ears, overt reflexive control of the pinnae has been lost during the course of primate evolution. Humans and apes do not move their ears to express emotion, they do not defensively retract them when startled, and they do not point them at novel, salient, or task‐relevant stimuli. Nevertheless, it is the thesis of this review that neural circuits for pinna orienting have survived in a purely vestigial state for over 25 million years. There are three lines of evidence: (1) Shifting the eyes hard to one side is accompanied by electromyographic (EMG) activity in certain ear muscles and by a barely visible (2–3 mm) curling of the dorsal edge of the pinna. (2) The capture of attention by a novel, unexpected sound emanating from behind and to one side has been found to trigger a weak EMG response in the muscle behind the corresponding ear. (3) Reflexive EMG bursts recorded during a selective attention task suggested that subjects were unconsciously attempting to orient their ears toward the relevant sounds. In addition to pinna orienting, the possibility that pinna startle might have survived in a vestigial state is also considered. It is suggested that the postauricular reflex to sudden, intense sounds constitutes a vestigial startle response, but that the reflex arc is dominated by a pathway that bypasses the main organizing center for startle.     

Binaural weighting of pinna cues in human sound localization

Human sound localization relies on binaural difference cues for sound-source azimuth and pinna-related spectral shape cues for sound elevation. Although the interaural timing and level difference cues are weighted to produce a percept of sound azimuth, much less is known about binaural mechanisms underlying elevation perception. This problem is particularly interesting for the frontal hemifield, where binaural inputs are of comparable strength. In this paper, localization experiments are described in which hearing for each ear was either normal, or spectrally disrupted by a mold fitted to the external ear. Head-fixed saccadic eye movements were used as a rapid and accurate indicator of perceived sound direction in azimuth and elevation. In the control condition (both ears free) azimuth and elevation components of saccadic responses were well described by a linear regression line for the entire measured range. For unilateral mold conditions, the azimuth response components did not differ from controls. The influence of the mold on elevation responses was largest on the ipsilateral side, and declined systematically with azimuth towards the side of the free ear. Near the midsagittal plane the elevation responses were clearly affected by the mold, suggesting a systematic binaural interaction in the neural computation of perceived elevation that straddles the midline. A quantitative comparison of responses from the unilateral mold, the bilateral mold and control condition provided evidence that the fusion process can be described by binaural weighted averaging. Two different conceptual schemes are discussed that could underlie the observed responses. Electronic Publication     

Listening through different ears alters spatial response fields in ferret primary auditory cortex

The localization of sounds in space is based on spatial cues that arise from the acoustical properties of the head and external ears. Individual differences in localization cue values result from variability in the shape and dimensions of these structures. We have mapped spatial response fields of high-frequency neurons in ferret primary auditory cortex using virtual sound sources based either on the animal's own ears or on the ears of other subjects. For 73% of units, the response fields measured using the animals' own ears differed significantly in shape and/or position from those obtained using spatial cues from another ferret. The observed changes correlated with individual differences in the acoustics. These data are consistent with previous reports showing that humans localize less accurately when listening to virtual sounds from other individuals. Together these findings support the notion that neural mechanisms underlying auditory space perception are calibrated by experience to the properties of the individual.     

Blind subjects process auditory spectral cues more efficiently than sighted individuals

The goal of the present study was to investigate how monaural sound localization on the horizontal plane in blind humans is affected by manipulating spectral cues. As reported in a previous study (Lessard et al. 1998), blind subjects are able to calibrate their auditory space despite their congenital lack of vision. Moreover, the performance level of half of the blind subjects was superior to that of sighted subjects under monaural listening conditions. Here, we first tested ten blind subjects and five controls in free-field (1) binaural and (2) monaural sound localization tasks. Results showed that, contrary to controls and half the blind subjects, five of the blind listeners were able to localize the sounds with one ear blocked. The blind subjects who showed good monaural localization performances were then re-tested in three additional monaural tasks, but we manipulated their ability to use spectral cues to carry out their discrimination. These subjects thus localized these same sounds: (3) with acoustical paste on the pinna, (4) with high-pass sounds and unobstructed pinna and (5) with low-pass sounds and unobstructed pinna. A significant increase in localization errors was observed when their ability to use spectral cues was altered. We conclude that one of the reasons why some blind subjects show supra-normal performances might be that they more effectively utilize auditory spectral cues.     

A radioisotopic method to measure delayed type hypersensitivity in the mouse. I. Studies in sensitized and normal mice.

Delayed type hypersensitivity (DTH) lesions have been difficult to evaluate objectively in the mouse because they are usually assessed in terms of an increase in footpad swelling or ear thickness. We have developed a radioisotopic method which not only reduces the observer's error but also gives an objective measurement of the cellular activity in the lesions. In brief, 10 mul of the test antigen is injected intradermally into the left pinna and either nothing or the same volume of a control solution into the right. 10 h later, a 2-muCi pulse of 5-iodo-2'deoxyuridine-125I is given intravenously, the ears cut off at the hairline 16 h later and the radioactivity counted in a gamma spectrometer. The following was obtained as evidence that the increased radioactivity of the left pinna over the right was a measure of the extent of a DTH response: (1) the ear reaction was delayed in mice without serum antibodies becoming maximum at 24 h; (2) there was a mononuclear cell infiltration in the left pinna and autoradiographs revealed radioactive label bound to these cells; (3) athymic mice could not develop a 24-hour ear reaction, and (4) antigens known not to activate T cells did not elicit the ear response. Cell transfer studies will be described in a subsequent paper. Different sensitization regimes were required with different antigens in order to obtain the highest levels of DTH as tested by the ear response, and the maximal ear reaction occurred at different days. The ear reaction showed the specificity expected of a DTH response.     

Allocation of attention in dichotic listening: differential effects on the detection and localization of signals.

In 2 dichotic listening experiments, 96 normal right-handed adults attended selectively to the left and right ear and divided their attention equally between both ears. Participants listened for specified targets and reported the ear of entry. The material consisted of pairs of consonant-vowel syllables in Experiment 1 and pairs of rhyming consonant-vowel-consonant words in Experiment 2. Both experiments yielded a right-ear advantage for detection and for localization. Attention instructions had no effect on detection. However, focusing attention on 1 ear increased the number of targets attributed to that ear while decreasing the number of targets attributed to the opposite ear. The dissociation between detection and localization indicates that volitional shifts of attention influence late (response selection) processes rather than early (stimulus identification) processes. Selective-listening effects can be accounted for by a 2-stage model in which a fixed input asymmetry is modulated by a biased selection of responses.     

先天性外耳道畸形多层螺旋CT的分析与研究

目的通过分析先天性外耳道畸形的多层螺旋cT表现,为临床提供准确、有效的影像信息。方法回顾性分析56例（64耳）婴儿（45天一6个月）先天性外耳道畸形患者的cT表现。结果56例先天性外耳道畸形患者双侧畸形者8例,单侧畸形者48例;合并中耳或内耳畸形者62耳,仅外耳畸形者2耳;鼓部无或轻度发育不良12耳,鼓部重度发育不良19耳,鼓部未发育33耳。结论多层螺旋cT对先天性外耳道畸形具有重要的诊断价值,是定制临床治疗方案的重要依据。     

Effects of altering spectral cues in infancy on horizontal and vertical sound localization by adult ferrets

We investigated the behavioral consequences of removing the pinna and concha of the external ear bilaterally in infancy on the sound localization ability of adult ferrets. Altering spectral cues in this manner has previously been shown to disrupt the development of the neural representation of auditory space in the superior colliculus. Using broadband noise stimuli, we tested pinnae-removed ferrets and normal ferrets in three sound localization tasks. In each case, we found that both groups of animals performed significantly better when longer duration noise bursts were used. In a relative localization task, we measured the acuity with which the ferrets could discriminate between two speakers in the horizontal plane. The speakers were placed symmetrically either around the anterior midline or around a position 45 degrees lateral to the midline. In this task, the pinnae-removed ferrets achieved very similar scores to the normal ferrets. By contrast, in another relative localization task that measured localization ability in the midsagittal plane, pinnae-removed ferrets performed less well than normals. In an absolute localization task, 12 speakers were spaced at 30 degrees intervals in the horizontal plane at the level of the ferrets' ears. Overall, the pinnae-removed ferrets also performed poorly in this task compared with normal ferrets: they made significantly fewer correct responses, larger localization errors and more front-back errors. Both normal and pinnae-removed animals showed an improvement in performance with practice, although the pattern of improvement differed for each group. The largest improvements in localization accuracy were achieved by the pinnae-removed ferrets, particularly at the frontal positions, and their performance eventually approached that of the normal animals. Nevertheless, some intergroup differences were still present. In particular, the pinnae-removed ferrets continued to make significantly more front-back errors than the normals. These deficits can be attributed to differences in the spectral localization cues available to the animals. Acoustical measurements showed that, compared with normal animals, the head-related transfer functions in the horizontal plane were largely ambiguous around the interaural axis and also contained fewer location-dependent features in the midsagittal plane.     

双侧内耳统计形状模型

目的 探讨建立双侧内耳统计形状模型的方法和影响因素,并探讨使用平均双侧内耳模型作为带空间信息的标准模型。 方法 通过Otsu法以及手动分割37例双侧内耳模型,对模型进行配准并生成统计形状模型,观察测量标准模型空间信息。 结果 Otsu 法可以分割获取高质量内耳模型,模型配准后使用Statismo 0.81软件可以生成稳定的内耳统计形状模型,其导出的平均模型空间方向信息具有代表性。结论 双侧内耳统计形状模型可以导出平均模型作为带空间方向信息的标准模型。     

Instability of hemispheric asymmetry in dyslexic children.

The study tested the hypothesis of abnormal brain asymmetry in dyslexic children. Two dyslexic groups classified as "phonetic disorder" and "language disorder" and a control group participated in two experiments. In both experiments was employed a dichotic listening procedure consisting in recalling pairs of words presented simultaneously to two ears. In Experiment I the children were to recognize four words presented at a level of loudness typical for natural speech. In Experiment II only two words were used in each trial but they were presented at a low intensity level. The recognition scores for stimuli presented to the left and right ears were compared. In Experiment I all groups of children showed a typical right ear/left hemisphere superiority, i.e. their recognition scores were higher for the right than for the left ear. Dyslexics, however, performed significantly less well. In Experiment II the control children and those from "phonetic disorder" group again performed better when words were presented to their right ears. Unlike those two groups, the children from "language disorder group" showed the right hemisphere superiority, i.e. they performed better in the left ear presentation condition. The results do not support the hypothesis that dyslexic children have abnormal lateralization of verbal functions. They suggest that the pattern of hemispheric asymmetry in dyslexics is less stable and depends both on the kind of dyslexia and on task variation.     

Sexual dimorphism in multiple aspects of 3D facial symmetry and asymmetry defined by spatially dense geometric morphometrics

Accurate measurement of facial sexual dimorphism is useful to understanding facial anatomy and specifically how faces influence, and have been influenced by, sexual selection. An important facial aspect is the display of bilateral symmetry, invoking the need to investigate aspects of symmetry and asymmetry separately when examining facial shape. Previous studies typically employed landmarks that provided only a sparse facial representation, where different landmark choices could lead to contrasting outcomes. Furthermore, sexual dimorphism is only tested as a difference of sample means, which is statistically the same as a difference in population location only. Within the framework of geometric morphometrics, we partition facial shape, represented in a spatially dense way, into patterns of symmetry and asymmetry, following a two-factor anova design. Subsequently, we investigate sexual dimorphism in symmetry and asymmetry patterns separately, and on multiple aspects, by examining (i) population location differences as well as differences in population variance-covariance; (ii) scale; and (iii) orientation. One important challenge in this approach is the proportionally high number of variables to observations necessitating the implementation of permutational and computationally feasible statistics. In a sample of gender-matched young adults (18-25 years) with self-reported European ancestry, we found greater variation in male faces than in women for all measurements. Statistically significant sexual dimorphism was found for the aspect of location in both symmetry and asymmetry (directional asymmetry), for the aspect of scale only in asymmetry (magnitude of fluctuating asymmetry) and, in contrast, for the aspect of orientation only in symmetry. Interesting interplays with hypotheses in evolutionary and developmental biology were observed, such as the selective nature of the force underpinning sexual dimorphism and the genetic independence of the structural patterns of fluctuating asymmetry. Additionally, insights into growth patterns of the soft tissue envelope of the face and underlying skull structure can also be obtained from the results.     

Acoustic factors govern developmental sharpening of spatial tuning in the auditory cortex

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.     

Right parietal cortex is involved in the perception of sound movement in humans

Changes in the delay (phase) and amplitude of sound at the ears are cues for the analysis of sound movement. The detection of these cues depends on the convergence of the inputs to each ear, a process that first occurs in the brainstem. The conscious perception of these cues is likely to involve higher centers. Using novel stimuli that produce different perceptions of movement in the presence of identical phase and amplitude modulation components, we have demonstrated human brain areas that are active specifically during the perception of sound movement. Both functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) demonstrated the involvement of the right parietal cortex in sound movement perception with these stimuli.     

Two-dimensional sound-localization behavior of early-blind humans

To investigate whether the visual system is crucial for adequate calibration of acoustic localization cues, sound-localization performance of early blind humans was compared with that of sighted controls. Because a potential benefit of vision is mainly expected for targets within the two-dimensional (2D) frontal hemifield, localization was tested within this target range, while using sounds of various durations and spectral content. Subjects were instructed to point, in separate experimental sessions, either with their left arm, or with their nose, in the direction of the perceived target position as accurately as possible. The experiments required the use of all available sound-localization cues such as interaural differences in phase and intensity, as well as the complex spectral shape cues provided by the pinnae. In addition, for long-duration stimuli, subjects could have had access to head motion-induced acoustic feedback. Moreover, the two pointing methods allowed us to assess different response strategies for the two groups. In an additional series, subjects were instructed to respond as quickly as possible. The results show that, in general, 2D sound-localization performance of blind subjects was indistinguishable from that of sighted subjects, both for broad-band noise and for pure tones. In the fast head-pointing task, the latency distributions of both groups were equal. These findings suggest that visual feedback is not required to calibrate the available localization cues – including the idiosyncratic and complex spectral shape cues for elevation. Instead, the localization abilities of blind people show that the putative supervising role of vision may be supported, or taken over, by other non-visual feedback systems. The results do not provide support for the hypothesis that blind people can hypercompensate for the loss of vision in the frontal hemifield by developing superior sound-localization abilities. Despite the general correspondence in localization behavior, some specific differences related to pointing strategies as well as to those between blind and sighted subjects were apparent. Most importantly, the reconstructed origin (bias) of arm pointing was located near the shoulder for the blind subjects, whereas it was shifted and located near the cyclopean eye for the sighted subjects. The results indicate that both early blind and sighted humans adequately transform the head-centered acoustic target coordinates into the required reference frame of either motor system, but that the adopted response strategy may be specific to the subject group and pointer method. Electronic Publication     

Effects of sex,age, and forced attention on dichotic listening in children: A longitudinal study

Seventy‐six children (38 boys, 38 girls) were tested with dichotic presentations of consonant‐vowel (CV) syllables under three different attentional instructions. Each child was tested twice: at age 8 and 1 year later (at age 9). The study investigated effects of laterally biased attention in dichotic performance in children. In the nonforced (NF) recall condition, children were free to report from both ears; in the forced‐right (FR) condition, they were instructed to attend to and report from only the right ear; in the forced‐left (FL) condition, they were instructed to attend to and report from only the left ear. The results showed a significant right‐ear advantage in both boys and girls and at both test occasions for the NF and FR conditions. During the FL condition, boys still reported more items from the right ear at age 8 but no at age 9, whereas girls reported an equal number of items from both ears at both test occasions. Thus, although hemisphere asymmetry did not change across age, the ability to laterally shift attention did and more so for boys than for girls.