Direct measurement of intra-cochlear pressure waves

The cochlear travelling wave is fundamental to the ability of the mammalian auditory system to resolve frequency. The seashell-shaped outer bone of the cochlea (the auditory inner ear) contains a spiral of cochlear fluid and the sensory tissue known as the cochlear partition. Sound travels down the ear canal to the eardrum, causing its flexible tympanic membrane to vibrate. This vibration is transmitted to the cochlea via the ossides. Motion of the stapes (the stirrup ossicle) sets the cochlear fluid in motion, which in turn sets the cochlear partition near the states in motion. The motion of the cochlear partition ripples down the cochlear spiral as a travelling wave, stimulating the cochlea's sensory hair cells. The wave peaks near the base (the stapes end) of the cochlea for high frequency tones and near the apex for low frequencies. The fundamental elements of the cochlear travelling wave are fluid pressure and motion and partition forces and motion. However, the wave's direct experimental study has to date relied almost solely on measurements of the partition motion. Here I report finely spaced measurements of intracochlear pressure close to the partition, which reveal the fluid component of the cochlear wave. The penetration depth of the wave is very limited, approximately 15 microm. Over a range of frequencies at least an octave wide, the depth is independent of frequency.     

Stereocilin-deficient mice reveal the origin of cochlear waveform distortions

Although the cochlea is an amplifier and a remarkably sensitive and finely tuned detector of sounds, it also produces conspicuous mechanical and electrical waveform distortions. These distortions reflect nonlinear mechanical interactions within the cochlea. By allowing one tone to suppress another (masking effect), they contribute to speech intelligibility. Tones can also combine to produce sounds with frequencies not present in the acoustic stimulus. These sounds compose the otoacoustic emissions that are extensively used to screen hearing in newborns. Because both cochlear amplification and distortion originate from the outer hair cells-one of the two types of sensory receptor cells-it has been speculated that they stem from a common mechanism. Here we show that the nonlinearity underlying cochlear waveform distortions relies on the presence of stereocilin, a protein defective in a recessive form of human deafness. Stereocilin was detected in association with horizontal top connectors, lateral links that join adjacent stereocilia within the outer hair cell's hair bundle. These links were absent in stereocilin-null mutant mice, which became progressively deaf. At the onset of hearing, however, their cochlear sensitivity and frequency tuning were almost normal, although masking was much reduced and both acoustic and electrical waveform distortions were completely lacking. From this unique functional situation, we conclude that the main source of cochlear waveform distortions is a deflection-dependent hair bundle stiffness resulting from constraints imposed by the horizontal top connectors, and not from the intrinsic nonlinear behaviour of the mechanoelectrical transducer channel.     

Mechanoelectrical transduction of adult outer hair cells studied in a gerbil hemicochlea

Sensory receptor cells of the mammalian cochlea are morphologically and functionally dichotomized. Inner hair cells transmit auditory information to the brain, whereas outer hair cells (OHC) amplify the mechanical signal, which is then transduced by inner hair cells. Amplification by OHCs is probably mediated by their somatic motility in a mechanical feedback process. OHC motility in vivo is thought to be driven by the cell's receptor potential. The first steps towards the generation of the receptor potential are the deflection of the stereociliary bundle, and the subsequent flow of transducer current through the mechanosensitive transducer channels located at their tips. Quantitative relations between transducer currents and basilar membrane displacements are lacking, as well as their variation along the cochlear length. To address this, we simultaneously recorded OHC transducer currents (or receptor potentials) and basilar membrane motion in an excised and bisected cochlea, the hemicochlea. This preparation permits recordings from adult OHCs at various cochlear locations while the basilar membrane is mechanically stimulated. Furthermore, the stereocilia are deflected by the same means of stimulation as in vivo. Here we show that asymmetrical transducer currents and receptor potentials are significantly larger than previously thought, they possess a highly restricted dynamic range and strongly depend on cochlear location.     

Two-tone distortion in the basilar membrane of the cochlea

When humans listen to pairs of thnes they hear additional tones, or distortion products, that are not present in the stimulus. Two-tone distortion products are also known as combination tones, because their pitches match combinations of the primary frequencies (f1 and f2, f2 greater than f1), such as f2-f1, (n + 1)f1-nf2 and (n + 1)f2-nf1 (n = 1, 2, 3...). Physiological correlates of the perceived distortion products exist in responses of auditory-nerve fibres and inner hair cells and in otoacoustic emissions (sounds generated by the cochlea, recordable at the ear canal). Because the middle ear responds linearly to sound and neural responses to distortion products can be abolished by damage to hair cells at cochlear sites preferentially tuned to the frequencies of the primary tones, it was hypothesized that distortion products are generated at these sites and propagate mechanically along the basilar membrane to the location tuned to the distortion-product frequency. But until now, efforts to confirm this hypothesis have failed. Here we report the use of a new laser-velocimetry technique to demonstrate two-tone distortion in basilar-membrane motion at low and moderate stimulus intensities.     

用小波变换实现电子耳蜗CIS语音信号的处理

为克服以往滤波器组参数调整复杂问题和实现电子耳蜗语音处理的快速数字化计算,提出了将小波变换应用于电子耳蜗的语音处理。介绍了电子耳蜗的机理及连续交替取样（ Ｃ Ｉ Ｓ）语音信号处理方案,并对耳蜗的频率分析特性与小波变换的时间—尺度分析特点进行了比较; 讨论了用小波变换实现 Ｃ Ｉ Ｓ语音信号处理方案的方法及仿真结果,给出尺度的选择依据,并与滤波器组的分析方法进行了比较。结果表明,用小波变换实现电子耳蜗的 Ｃ Ｉ Ｓ语音信号处理方案是可行的     

Graded and nonlinear mechanical properties of sensory hairs in the mammalian hearing organ

It is generally agreed that frequency selectivity of the mammalian hearing organ is mainly due to a graded elasticity of the basilar membrane. Recent measurements of basilar membrane motion hair cell receptor potentials and neural tuning curves show that frequency selectivity can be extremely sharp. It has been suggested that in non-mammalian species there are additional tuning mechanisms in the sensory hair cells themselves, either by virtue of their electrical membrane properties or through a gradation in length of their sensory hairs. Indeed, sensory hair mechanical tuning has been demonstrated in the lizard. We have investigated the mechanical properties of sensory hair bundles in the guinea pig organ of Corti, and report here that hair-bundle stiffness increases longitudinally towards the high-frequency end of the cochlea, decreases radially towards the outer rows of cells, and is greater for excitatory than for inhibitory deflection. On the basis of these findings, we suggest that sensory hairs confer frequency-specific, nonlinear mechanical properties on the hearing organ.     

人眼明度视觉空间频率传递特性的研究

为描述人眼视觉系统对空间复杂颜色刺激的信息处理机制,研究人眼视觉系统空间频率信息传递的多通道特性.设计心理物理实验,测量人眼视觉系统在感知均匀色空间CIELAB明度通道上的对比度敏感度数据.实验选取明度轴上5个明度值作为颜色样本,制作明度对比度随正弦波频率调制的图像刺激序列,挑选正常视觉观察者,采集5个明度样本的12个空间频率的对比度敏感度数据.对实验数据进行相关性分析发现:单个明度样本的不同空间频率对应的观察值之间是非线性相关的;随着明度样本数量增多,观察值之间的线性相关性增强.因此,人眼视觉系统对明度颜色通道的信息处理机制可用线性模型描述;明度颜色通道的空间频率传递特性可用3条空间频率调谐曲线表征;3条空间频率调谐曲线作用的频段不同,峰值不同,表现了人眼视觉系统具有不同的空间频率感受野.      

Outer hair cells in the mammalian cochlea and noise-induced hearing loss

Hair cells in the mammalian cochlea transduce mechanical stimuli into electrical signals leading to excitation of auditory nerve fibres. Because of their important role in hearing, these cells are a possible site for the loss of cochlear sensitivity that follows acoustic overstimulation. We have recorded from inner and outer hair cells (IHC, OHC) in the guinea pig cochlea during and after exposure to intense tones. Our results show functional changes in the hair cells that may explain the origin of noise-induced hearing loss. Both populations of hair cells show a reduction in amplitude and an increase in the symmetry of their acoustically evoked receptor potentials. In addition, the OHCs also suffer a sustained depolarization of the membrane potential. Significantly, the membrane and receptor potentials of the OHCs recover in parallel with cochlear sensitivity as measured by the IHC receptor potential amplitude and the auditory nerve threshold. Current theories of acoustic transduction suggest that the mechanical input to IHCs may be regulated by the OHCs. Consequently, the modified function of OHCs after acoustic overstimulation may determine the extent of the hearing loss following loud sound.     

Force generation by mammalian hair bundles supports a role in cochlear amplification

It is generally accepted that the acute sensitivity and frequency discrimination of mammalian hearing requires active mechanical amplification of the sound stimulus within the cochlea. The prevailing hypothesis is that this amplification stems from somatic electromotility of the outer hair cells attributable to the motor protein prestin. Thus outer hair cells contract and elongate in synchrony with the sound-evoked receptor potential. But problems arise with this mechanism at high frequencies, where the periodic component of the receptor potential will be attenuated by the membrane time constant. On the basis of work in non-mammalian vertebrates, force generation by the hair bundles has been proposed as an alternative means of boosting the mechanical stimulus. Here we show that hair bundles of mammalian outer hair cells can also produce force on a submillisecond timescale linked to adaptation of the mechanotransducer channels. Because the bundle motor may ultimately be limited by the deactivation rate of the channels, it could theoretically operate at high frequencies. Our results show the existence of another force generator in outer hair cells that may participate in cochlear amplification.     

Orthogonal linear polarized lasers (Ⅱ)--Study on the physical phenomena

The physical phenomena and corresponding theoretical analysis of orthogonal polarized laser are reviewed. Four lasers (or systems) with orthogonal polarized beams are involved. For the birefringence dual frequency laser, its physical phenomena discussed include the alternation between strong mode competition and medium mode competition in cavity tuning; the range of frequency difference of strong mode competition (about 0-40 MHz); four polarization statuses (o-light oscillating but e-light extinguishing, both o-light and elight oscillating, e-light oscillating but o-light extinguishing, both o-light and e-light extinguishing) in cavity tuning; the tuning curves of frequency difference; the influence of optical activity of quartz crystal on polarization direction; and the aberrance of frequency splitting.For the Birefringence-Zeeman dual frequency laser, we focus on its intensity tuning and frequency difference tuning. For the feedback system of orthogonally polarized laser, we discuss the mutual suppression between two orthogonal frequencies, intensity exchange between two orthogonal frequencies and double of intensity fringe frequency. For orthogonally polarized LD-pumped Nd: YAG microchip laser, its property of the dependence of intensity sensitivity on frequency difference is described.     

Orthogonal linear polarized lasers (II)--Study on the physical phenomena

The physical phenomena and corresponding theoretical analysis of orthogonal polarized laser are reviewed. Four lasers (or systems) with orthogonal polarized beams are involved. For the birefringence dual frequency laser, its physical phenomena discussed include the alternation between strong mode competition and medium mode competition in cavity tuning; the range of frequency difference of strong mode competition (about 0?40 MHz); four polarization statuses (o-light oscillating but e-light extinguishing, both o-light and e-light oscillating, e-light oscillating but o-light extinguishing, both o-light and e-light extinguishing) in cavity tuning; the tuning curves of frequency difference; the influence of optical activity of quartz crystal on polarization direction; and the aberrance of frequency splitting. For the Birefringence-Zeeman dual frequency laser, we focus on its intensity tuning and frequency difference tuning. For the feedback system of orthogonally polarized laser, we discuss the mutual suppression between two orthogonal frequencies, intensity exchange between two orthogonal frequencies and double of intensity fringe frequency. For orthogonally polarized LD-pumped Nd: YAG microchip laser, its property of the dependence of intensity sensitivity on frequency difference is described.     

Auditory collusion and a coupled couple of outer hair cells.

The discrepancies between measured frequency responses of the basilar membrane in the inner ear and the frequency tuning found in psychophysical experiments led to Bekesy's idea of lateral inhibition in the auditory nervous system. We now know that basilar membrane tuning can account for neural tuning, and that sharpening of the passive travelling wave depends on the mechanical activity of outer hair cells (OHCs)3, but the mechanism by which OHCs enhance tuning remains unclear. OHCs generate voltage-dependent length changes at acoustic rates, which deform the cochlear partition. Here we use an electrical correlate of OHC mechanical activity, the motility-related gating current, to investigate mechano-electrical interactions among adjacent OHCs. We show that the motility caused by voltage stimulation of one cell in a group evokes gating currents in adjacent OHCs. The resulting polarization in adjacent cells is opposite to that within the stimulated cell, which may be indicative of lateral inhibition. Also such interactions promote distortion and suppression in the electrical and, consequently, the mechanical activity of OHCs. Lateral interactions may provide a basis for enhanced frequency selectivity in the basilar membrane of mammals.     

Prestin is the motor protein of cochlear outer hair cells

The outer and inner hair cells of the mammalian cochlea perform different functions. In response to changes in membrane potential, the cylindrical outer hair cell rapidly alters its length and stiffness. These mechanical changes, driven by putative molecular motors, are assumed to produce amplification of vibrations in the cochlea that are transduced by inner hair cells. Here we have identified an abundant complementary DNA from a gene, designated Prestin, which is specifically expressed in outer hair cells. Regions of the encoded protein show moderate sequence similarity to pendrin and related sulphate/anion transport proteins. Voltage-induced shape changes can be elicited in cultured human kidney cells that express prestin. The mechanical response of outer hair cells to voltage change is accompanied by a 'gating current', which is manifested as nonlinear capacitance. We also demonstrate this nonlinear capacitance in transfected kidney cells. We conclude that prestin is the motor protein of the cochlear outer hair cell.     

Prestin is required for electromotility of the outer hair cell and for the cochlear amplifier

Hearing sensitivity in mammals is enhanced by more than 40 dB (that is, 100-fold) by mechanical amplification thought to be generated by one class of cochlear sensory cells, the outer hair cells. In addition to the mechano-electrical transduction required for auditory sensation, mammalian outer hair cells also perform electromechanical transduction, whereby transmembrane voltage drives cellular length changes at audio frequencies in vitro. This electromotility is thought to arise through voltage-gated conformational changes in a membrane protein, and prestin has been proposed as this molecular motor. Here we show that targeted deletion of prestin in mice results in loss of outer hair cell electromotility in vitro and a 40-60 dB loss of cochlear sensitivity in vivo, without disruption of mechano-electrical transduction in outer hair cells. In heterozygotes, electromotility is halved and there is a twofold (about 6 dB) increase in cochlear thresholds. These results suggest that prestin is indeed the motor protein, that there is a simple and direct coupling between electromotility and cochlear amplification, and that there is no need to invoke additional active processes to explain cochlear sensitivity in the mammalian ear.     

鼠耳蝠下丘听神经元对超声刺激的反应特性

实验在５只鼠耳蜗上进行，共观察了２２７个对超声刺激发生反应的下丘听神经元。这些神经元听反应的最佳频率在１４．３－７６．２ｋＨｚ之间；潜伏期在３．０－１２．０毫秒之间，多数神经元（７９．３％）为５．０－７．０毫秒；最低阈值位于－２０．０－７０．０ｄＢ ＳＰＬ之间；多数神经元（８２．０５％）的调谐曲线为宽阔型，少数（１７．９５％）为狭窄型；反应最佳频率沿下丘背腹轴呈明显有序地排列。     

Structural alteration of hair cells in the contralateral ear resulting from extracochlear electrical stimulation

Chronic electrical stimulation of the auditory nerve in patients with profound sensori-neural deafness is becoming increasingly routine. Therefore, it is important to understand more about the long-term consequences of this procedure. Hitherto, structural studies in animals after electrocochlear stimulation have concentrated on the stimulated cochlea. Here we have examined the effects of unilateral extracochlear electrical stimulation on the spiral organ of both the ipsilateral and contralateral ears of the mature guinea pig, and have found alterations in the structure of the outer hair cells and their efferent nerve terminals in the contralateral as well as the ipsilateral cochlea. This is the first evidence for a structural influence of efferent activity on the cochlea. Although the importance of the efferent system, consisting of the crossed and uncrossed olivo-cochlear bundles, is well established in providing central control of the sensory pathways, its exact role in hearing is incompletely understood. However, it is known that the outer hair cells and their efferent innervation are important in their contribution to inner hair cell responses and in modulating the micromechanics of the whole cochlea. These efferent functions now appear to be related to an important part of cochlear morphology, and are also relevant to our understanding of cochlear neurobiology, normal development and the management of hearing disability in both adult and child.     

Role of frequency band integration in sharpening frequency tunings of the inferior colliculus neurons in the big brown bat,<Emphasis Type="Italic">Eptesicus fuscus</Emphasis>

By means of a particular two-tone stimulationparadigm in combination of using a pair of electrodes forsimultaneously recording from two inferior colliculus (IC)neurons, the current in vivo study is undertaken to explorethe role of frequency band integration (FBI) in sharpening offrequency tuning in the big brown bat, Eptesicus fuscus.Three major results are found: (1) The paired neurons cor-related to FBI are located not only within the same frequencyfilter bandwidth (FFB), but also across different FFBs. Therelations of their frequency tuning curves (FTCs) are mainlyof two types: the flank-overlapped and overlaid patterns. (2)Although the sharpness of FTCs between paired neurons ismutual, the sharpening efficiency of neurons located withinthe same FFB is higher than that of neurons across FFBs,and the FTCs of neurons with the best frequencies (BF) of 20--30 kHz are most strongly sharpened. (3) The strength ofFBI is weak near the BF but gradually increased with fre-quencies away from the BF of sound stimuli. This suggeststhat the dynamical FBI of the IC neurons located within and across the FFBs might be involved in the formation of func-tional FFB structures.     

Electrokinetic shape changes of cochlear outer hair cells

Rapid mechanical changes have been associated with electrical activity in a variety of non-muscle excitable cells. Recently, mechanical changes have been reported in cochlear hair cells. Here we describe electrically evoked mechanical changes in isolated cochlear outer hair cells (OHCs) with characteristics which suggest that direct electrokinetic phenomena are implicated in the response. OHCs make up one of two mechanosensitive hair cell populations in the mammalian cochlea; their role may be to modulate the micromechanical properties of the hearing organ through mechanical feedback mechanisms. In the experiments described here, we applied sinusoidally modulated electrical potentials across isolated OHCs; this produced oscillatory elongation and shortening of the cells and oscillatory displacements of intracellular organelles. The movements were a function of the direction and strength of the electrical field, were inversely related to the ionic concentration of the medium, and occurred in the presence of metabolic uncouplers. The cylindrical shape of the OHCs and the presence of a system of membranes within the cytoplasm--laminated cisternae--may provide the anatomical substrate for electrokinetic phenomena such as electro-osmosis.     

豚鼠耳蜗基底膜的振动特性

为探讨豚鼠耳蜗基底膜的听觉机制,利用多普勒激光测振技术进行了声音在28只成年健康豚鼠耳中传递的测试实验.将豚鼠麻醉后切断头取出听泡,采用电钻暴露其耳蜗基底膜,利用多普勒激光测振仪在给定的声压级、频率的纯音激励条件下测量豚鼠外耳道耳蜗基底膜的振动速度,并测量了中耳镫骨的振动速度.结果表明,豚鼠外耳道基底膜具有频率选择特性,而且其基底膜的振动具有行波特性.