大鼠初级听皮层神经元对声音空间信息的编码

尽管大脑听皮层神经元对声音空间信息的编码已有不少的研究报道,但其编码机制并不十分清楚,相关研究在大鼠的初级听皮层也未见详细的研究报道．用神经电生理学方法在大鼠初级听皮层考察了151个听神经元的听空间反应域,分析了神经元对来自不同空间方位声刺激反应的放电数和平均首次发放潜伏期的关系．结果表明,多数(52.32%)神经元对来自对侧听空间的声刺激反应较强,表现为对侧偏好型特征,其他神经元分别归类为同侧偏好型(18.54%)、中间偏好型(18.54%)、全向型(3.31%)和复杂型(7.28%)．多数神经元偏好的听空间区域的几何中心位于记录部位对侧听空间的中部和上部．绝大多数初级听皮层神经元对来自偏好听空间的声刺激反应的放电数较多、反应潜伏期较短,对来自非偏好听空间的声刺激反应的放电数较少、反应潜伏期较长,放电数与平均首次发放潜伏期呈显著负相关．在对声音空间信息的编码中,大脑初级听皮层可能综合放电数和潜伏期的信息以实现对声源方位的编码．     

幼年蝙蝠下丘神经元的听空间特性

实验分别在出生后４周龄的幼年和成年鲁氏菊头蝠（Ｒｈｉｎｏｌｏｐｈｕｓｒｏｕｘｉ）上进行。使用移动声刺激装置,高频喇叭可在动物头部前方水平方向１８０度、垂直方向６０度的范围内移动。玻璃微电极记录单个神经元的听反应。实验考察了幼年和成年动物下丘神经元的听空间特性,共观察了３０１个神经元,其中幼年动物１４８个,成年动物１５３个。结果表明,４周龄的幼年动物下丘听神经元已表现出方向敏感性,即每个听神经元均有一个特定的最佳反应中心和反应域。但神经元听反应中心在听空间的分布相当弥散,大多数位于对侧水平方向２０—８０度、垂直方向上下１５度范围内。而成年动物听神经元反应中心的分布则相当集中,局限地分布于对侧水平方向２８－５０度,垂直方向０—１０度范围内,两者构成明显差异。     

Distributed coding of sound locations in the auditory cortex

Although the auditory cortex plays an important role in sound localization, that role is not well understood. In this paper, we examine the nature of spatial representation within the auditory cortex, focusing on three questions. First, are sound-source locations encoded by individual sharply tuned neurons or by activity distributed across larger neuronal populations? Second, do temporal features of neural responses carry information about sound-source location? Third, are any fields of the auditory cortex specialized for spatial processing? We present a brief review of recent work relevant to these questions along with the results of our investigations of spatial sensitivity in cat auditory cortex. Together, they strongly suggest that space is represented in a distributed manner, that response timing (notably first-spike latency) is a critical information-bearing feature of cortical responses, and that neurons in various cortical fields differ in both their degree of spatial sensitivity and their manner of spatial coding. The posterior auditory field (PAF), in particular, is well suited for the distributed coding of space and encodes sound-source locations partly by modulations of response latency. Studies of neurons recorded simultaneously from PAF and/or A1 reveal that spatial information can be decoded from the relative spike times of pairs of neurons - particularly when responses are compared between the two fields - thus partially compensating for the absence of an absolute reference to stimulus onset.     

Location coding by opponent neural populations in the auditory cortex

Although the auditory cortex plays a necessary role in sound localization, physiological investigations in the cortex reveal inhomogeneous sampling of auditory space that is difficult to reconcile with localization behavior under the assumption of local spatial coding. Most neurons respond maximally to sounds located far to the left or right side, with few neurons tuned to the frontal midline. Paradoxically, psychophysical studies show optimal spatial acuity across the frontal midline. In this paper, we revisit the problem of inhomogeneous spatial sampling in three fields of cat auditory cortex. In each field, we confirm that neural responses tend to be greatest for lateral positions, but show the greatest modulation for near-midline source locations. Moreover, identification of source locations based on cortical responses shows sharp discrimination of left from right but relatively inaccurate discrimination of locations within each half of space. Motivated by these findings, we explore an opponent-process theory in which sound-source locations are represented by differences in the activity of two broadly tuned channels formed by contra- and ipsilaterally preferring neurons. Finally, we demonstrate a simple model, based on spike-count differences across cortical populations, that provides bias-free, level-invariant localization—and thus also a solution to the “binding problem” of associating spatial information with other nonspatial attributes of sounds.     

The Opponent Channel Population Code of Sound Location Is an Efficient Representation of Natural Binaural Sounds

In mammalian auditory cortex, sound source position is represented by a population of broadly tuned neurons whose firing is modulated by sounds located at all positions surrounding the animal. Peaks of their tuning curves are concentrated at lateral position, while their slopes are steepest at the interaural midline, allowing for the maximum localization accuracy in that area. These experimental observations contradict initial assumptions that the auditory space is represented as a topographic cortical map. It has been suggested that a “panoramic” code has evolved to match specific demands of the sound localization task. This work provides evidence suggesting that properties of spatial auditory neurons identified experimentally follow from a general design principle- learning a sparse, efficient representation of natural stimuli. Natural binaural sounds were recorded and served as input to a hierarchical sparse-coding model. In the first layer, left and right ear sounds were separately encoded by a population of complex-valued basis functions which separated phase and amplitude. Both parameters are known to carry information relevant for spatial hearing. Monaural input converged in the second layer, which learned a joint representation of amplitude and interaural phase difference. Spatial selectivity of each second-layer unit was measured by exposing the model to natural sound sources recorded at different positions. Obtained tuning curves match well tuning characteristics of neurons in the mammalian auditory cortex. This study connects neuronal coding of the auditory space with natural stimulus statistics and generates new experimental predictions. Moreover, results presented here suggest that cortical regions with seemingly different functions may implement the same computational strategy-efficient coding.     

Response variability in the mammalian auditory cortex: an objection to feature detection?

Research strategy in the auditory system has tended to parallel that in the visual system, where neurons have been shown to respond selectively to specific stimulus parameters. Auditory neurons have been shown to be sensitive to changes in acoustic parameters, but only rarely have neurons been reported that respond exclusively to only one biologically significant sound. Even at higher levels of the auditory system very few cells have been found that could be described as "vocalization detectors." In addition, variability in responses to artificial sounds have been reported for auditory cortical neurons similar to the response variability that has been reported in the visual system. Recent evidence indicates that the responses of auditory cortical neurons to species-specific vocalizations can also be labile, varying in both strength and selectivity. This is especially true of the secondary auditory cortex. This variability, coupled with the lack of extreme specificity in the secondary auditory cortex, suggests that secondary cortical neurons are not well suited for the role of "vocalization detectors."     

Learning-induced plasticity in animal and human auditory cortex

Recent data on learning-related changes in animal and human auditory cortex indicate functions beyond mere stimulus representation and simple recognition memory for stimuli. Rather, auditory cortex seems to process and represent stimuli in a task-dependent fashion. This implies plasticity in neural processing, which can be observed at the level of single neuron firing and the level of spatiotemporal activity patterns in cortical areas. Auditory cortex is a structure in which behaviorally relevant aspects of stimulus processing are highly developed because of the fugitive nature of auditory stimuli.     

When and where of auditory spatial processing in cortex: a novel approach using electrotomography

The modulation of brain activity as a function of auditory location was investigated using electro-encephalography in combination with standardized low-resolution brain electromagnetic tomography. Auditory stimuli were presented at various positions under anechoic conditions in free-field space, thus providing the complete set of natural spatial cues. Variation of electrical activity in cortical areas depending on sound location was analyzed by contrasts between sound locations at the time of the N1 and P2 responses of the auditory evoked potential. A clear-cut double dissociation with respect to the cortical locations and the points in time was found, indicating spatial processing (1) in the primary auditory cortex and posterodorsal auditory cortical pathway at the time of the N1, and (2) in the anteroventral pathway regions about 100 ms later at the time of the P2. Thus, it seems as if both auditory pathways are involved in spatial analysis but at different points in time. It is possible that the late processing in the anteroventral auditory network reflected the sharing of this region by analysis of object-feature information and spectral localization cues or even the integration of spatial and non-spatial sound features.     

Thalamic activation modulates the responses of neurons in rat primary auditory cortex: an in vivo intracellular recording study

Auditory cortical plasticity can be induced through various approaches. The medial geniculate body (MGB) of the auditory thalamus gates the ascending auditory inputs to the cortex. The thalamocortical system has been proposed to play a critical role in the responses of the auditory cortex (AC). In the present study, we investigated the cellular mechanism of the cortical activity, adopting an in vivo intracellular recording technique, recording from the primary auditory cortex (AI) while presenting an acoustic stimulus to the rat and electrically stimulating its MGB. We found that low-frequency stimuli enhanced the amplitudes of sound-evoked excitatory postsynaptic potentials (EPSPs) in AI neurons, whereas high-frequency stimuli depressed these auditory responses. The degree of this modulation depended on the intensities of the train stimuli as well as the intervals between the electrical stimulations and their paired sound stimulations. These findings may have implications regarding the basic mechanisms of MGB activation of auditory cortical plasticity and cortical signal processing.     

弱背景噪声对大鼠听皮层神经元频率调谐的影响

在自然环境中,人和动物常在一定的背景噪声下感知信号声刺激,然而,关于低强度的弱背景噪声如何影响听皮层神经元对声刺激频率的编码尚不清楚.本研究以大鼠听皮层神经元的频率反应域为研究对象,测定了阈下背景噪声对79个神经元频率反应域的影响.结果表明,弱背景噪声对大鼠初级听皮层神经元的听反应既有抑制性影响、又有易化性影响.一般来说,抑制性影响使神经元的频率调谐范围和最佳频率反应域缩小,易化性影响使神经元的频率调谐范围和最佳频率反应域增大.对于少数神经元,弱背景噪声并未显著改变其频率调谐范围,但却改变了其最佳频率反应域范围.弱背景噪声对63.64%神经元的特征频率和55.84%神经元的最低阈值无显著影响.神经元频率调谐曲线的尖部比中部更容易受到弱背景噪声的影响.该研究结果有助于我们进一步理解复杂声环境下大脑听皮层对听觉信息的编码机制.     

Corticofugal regulation of auditory sensitivity in the bat inferior colliculus

Under free-field stimulation conditions, corticofugal regulation of auditory sensitivity of neurons in the central nucleus of the inferior colliculus of the big brown bat, Eptesicus fuscus, was studied by blocking activities of auditory cortical neurons with Lidocaine or by electrical stimulation in auditory cortical neuron recording sites. The corticocollicular pathway regulated the number of impulses, the auditory spatial response areas and the frequency-tuning curves of inferior colliculus neurons through facilitation or inhibition. Corticofugal regulation was most effective at low sound intensity and was dependent upon the time interval between acoustic and electrical stimuli. At optimal interstimulus intervals, inferior colliculus neurons had the smallest number of impulses and the longest response latency during corticofugal inhibition. The opposite effects were observed during corticofugal facilitation. Corticofugal inhibitory latency was longer than corticofugal facilitatory latency. Iontophoretic application of γ-aminobutyric acid and bicuculline to inferior colliculus recording sites produced effects similar to what were observed during corticofugal inhibition and facilitation. We suggest that corticofugal regulation of central auditory sensitivity can provide an animal with a mechanism to regulate acoustic signal processing in the ascending auditory pathway. Accepted: 15 July 1998     

Distribution of auditory motion-direction sensitive neurons in the barn owl's midbrain

Barn owls have neurons sensitive to acoustic motion-direction in the midbrain. We report here that acoustic motion-direction sensitive neurons with receptive-field centres in frontal auditory space are not randomly distributed. In the inferior colliculus and optic tectum in the left (right) brain, the responses of about two-thirds of the motion-direction sensitive neurons were sensitive to clockwise (counter-clockwise) motion. The midbrain contains maps of auditory space that represent about 15 degrees of ipsilateral and all of contralateral space. Since a similar bias in motion-direction sensitivity was observed for neurons with receptive-field centres in ipsilateral as well as for neurons with receptive fields centres in contralateral auditory space, the brain side at which a motion-direction sensitive neuron was recorded was a more important predictor for the preferred direction of a cell than the spatial direction of the centre of the receptive field. Within one dorso-ventral electrode pass motion-direction sensitivity typically stayed constant suggesting a clustered or even a columnar-like organization. We hypothesize from these distributions that the right brain is important for orientating movements to the left hemisphere and vice versa.     

刺激大鼠杏仁外侧核对皮层AⅠ区ON-OFF神经元反应及调谐曲线的影响

在30只氨基甲酸乙酯麻醉的SD大鼠上记录神经元单位放电,观察短纯音诱发的皮层AI区神经元ON－OFF反应的特性及电刺激杏仁外侧核（lateral amygdaloid nucleus,LA)对ON－OFF反应以及调谐曲线的影响。实验证实,AI区神经元ON－OFF反应的模式与纯音刺激的强度、频率及作用时程有关;刺激LA可以抑制ON－OFF反应的放电频数,使反应的阈值升高,或使反应放电构型发生变化;此外,刺激LA能使ON－OFF神经元的调谐曲线变窄,Q10数值增大。研究结果不仅表明ON－OFF神经元能对纯音刺激的时程、强度和频率等多种信息进行编码,而且还证明杏仁外侧核可以在皮层水平参与听觉信息的调制,削弱或衰减某些听觉信息,导致整个调谐曲线上移变窄,从而提高AI区ON－OFF神经元的频率选择性能,有利于检测外界嘈杂环境中特定的听觉信息。     

The auditory cortex

The division of the auditory cortex into various fields, functional aspects of these fields, and neuronal coding in the primary auditory cortical field (AI) are reviewed with stress on features that may be common to mammals. On the basis of 14 topographies and clustered distributions of neuronal response characteristics in the primary auditory cortical field, a hypothesis is developed of how a certain complex acoustic pattern may be encoded in an equivalent spatial activity pattern in AI, generated by time-coordinated firing of groups of neurons. The auditory cortex, demonstrated specifically for AI, appears to perform sound analysis by synthesis, i.e. by combining spatially distributed coincident or time-coordinated neuronal responses. The dynamics of sounds and the plasticity of cortical responses are considered as a topic for research. Accepted: 25 July 1997     

A Population Rate Code of Auditory Space in the Human Cortex

Background

Previous work on the human auditory cortex has revealed areas specialized in spatial processing but how the neurons in these areas represent the location of a sound source remains unknown.

Methodology/Principal Findings

Here, we performed a magnetoencephalography (MEG) experiment with the aim of revealing the neural code of auditory space implemented by the human cortex. In a stimulus-specific adaptation paradigm, realistic spatial sound stimuli were presented in pairs of adaptor and probe locations. We found that the attenuation of the N1m response depended strongly on the spatial arrangement of the two sound sources. These location-specific effects showed that sounds originating from locations within the same hemifield activated the same neuronal population regardless of the spatial separation between the sound sources. In contrast, sounds originating from opposite hemifields activated separate groups of neurons.

Conclusions/Significance

These results are highly consistent with a rate code of spatial location formed by two opponent populations, one tuned to locations in the left and the other to those in the right. This indicates that the neuronal code of sound source location implemented by the human auditory cortex is similar to that previously found in other primates.     

A common cortical substrate activated by horizontal and vertical sound movement in the human brain

Perception of movement in acoustic space depends on comparison of the sound waveforms reaching the two ears (binaural cues) as well as spectrotemporal analysis of the waveform at each ear (monaural cues). The relative importance of these two cues is different for perception of vertical or horizontal motion, with spectrotemporal analysis likely to be more important for perceiving vertical shifts. In humans, functional imaging studies have shown that sound movement in the horizontal plane activates brain areas distinct from the primary auditory cortex, in parietal and frontal lobes and in the planum temporale. However, no previous work has examined activations for vertical sound movement. It is therefore difficult to generalize previous imaging studies, based on horizontal movement only, to multidimensional auditory space perception. Using externalized virtual-space sounds in a functional magnetic resonance imaging (fMRI) paradigm to investigate this, we compared vertical and horizontal shifts in sound location. A common bilateral network of brain areas was activated in response to both horizontal and vertical sound movement. This included the planum temporale, superior parietal cortex, and premotor cortex. Sounds perceived laterally in virtual space were associated with contralateral activation of the auditory cortex. These results demonstrate that sound movement in vertical and horizontal dimensions engages a common processing network in the human cerebral cortex and show that multidimensional spatial properties of sounds are processed at this level.     

Corticofugal modulation of initial neural processing of sound information from the ipsilateral ear in the mouse

Background

Cortical neurons implement a high frequency-specific modulation of subcortical nuclei that includes the cochlear nucleus. Anatomical studies show that corticofugal fibers terminating in the auditory thalamus and midbrain are mostly ipsilateral. Differently, corticofugal fibers terminating in the cochlear nucleus are bilateral, which fits to the needs of binaural hearing that improves hearing quality. This leads to our hypothesis that corticofugal modulation of initial neural processing of sound information from the contralateral and ipsilateral ears could be equivalent or coordinated at the first sound processing level.

Methodology/Principal Findings

With the focal electrical stimulation of the auditory cortex and single unit recording, this study examined corticofugal modulation of the ipsilateral cochlear nucleus. The same methods and procedures as described in our previous study of corticofugal modulation of contralateral cochlear nucleus were employed simply for comparison. We found that focal electrical stimulation of cortical neurons induced substantial changes in the response magnitude, response latency and receptive field of ipsilateral cochlear nucleus neurons. Cortical stimulation facilitated auditory response and shortened the response latency of physiologically matched neurons whereas it inhibited auditory response and lengthened the response latency of unmatched neurons. Finally, cortical stimulation shifted the best frequencies of cochlear neurons towards those of stimulated cortical neurons.

Conclusion

Our data suggest that cortical neurons enable a high frequency-specific remodelling of sound information processing in the ipsilateral cochlear nucleus in the same manner as that in the contralateral cochlear nucleus.     

Coding of communication calls in the subcortical and cortical structures of the auditory system

The processing of species-specific communication signals in the auditory system represents an important aspect of animal behavior and is crucial for its social interactions, reproduction, and survival. In this article the neuronal mechanisms underlying the processing of communication signals in the higher centers of the auditory system--inferior colliculus (IC), medial geniculate body (MGB) and auditory cortex (AC)--are reviewed, with particular attention to the guinea pig. The selectivity of neuronal responses for individual calls in these auditory centers in the guinea pig is usually low--most neurons respond to calls as well as to artificial sounds; the coding of complex sounds in the central auditory nuclei is apparently based on the representation of temporal and spectral features of acoustical stimuli in neural networks. Neuronal response patterns in the IC reliably match the sound envelope for calls characterized by one or more short impulses, but do not exactly fit the envelope for long calls. Also, the main spectral peaks are represented by neuronal firing rates in the IC. In comparison to the IC, response patterns in the MGB and AC demonstrate a less precise representation of the sound envelope, especially in the case of longer calls. The spectral representation is worse in the case of low-frequency calls, but not in the case of broad-band calls. The emotional content of the call may influence neuronal responses in the auditory pathway, which can be demonstrated by stimulation with time-reversed calls or by measurements performed under different levels of anesthesia. The investigation of the principles of the neural coding of species-specific vocalizations offers some keys for understanding the neural mechanisms underlying human speech perception.     

Object-oriented echo perception and cortical representation in echolocating bats

Echolocating bats can identify three-dimensional objects exclusively through the analysis of acoustic echoes of their ultrasonic emissions. However, objects of the same structure can differ in size, and the auditory system must achieve a size-invariant, normalized object representation for reliable object recognition. This study describes both the behavioral classification and the cortical neural representation of echoes of complex virtual objects that vary in object size. In a phantom-target playback experiment, it is shown that the bat Phyllostomus discolor spontaneously classifies most scaled versions of objects according to trained standards. This psychophysical performance is reflected in the electrophysiological responses of a population of cortical units that showed an object-size invariant response (14/109 units, 13%). These units respond preferentially to echoes from objects in which echo duration (encoding object depth) and echo amplitude (encoding object surface area) co-varies in a meaningful manner. These results indicate that at the level of the bat's auditory cortex, an object-oriented rather than a stimulus-parameter-oriented representation of echoes is achieved.