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乌龟耳蜗中听觉神经纤维和毛细胞的频率选择性。

The frequency selectivity of auditory nerve fibres and hair cells in the cochlea of the turtle.

作者信息

Crawford A C, Fettiplace R

出版信息

J Physiol. 1980 Sep;306:79-125. doi: 10.1113/jphysiol.1980.sp013387.

DOI:10.1113/jphysiol.1980.sp013387
PMID:7463380
原文链接:https://pmc.ncbi.nlm.nih.gov/articles/PMC1282996/
Abstract
  1. The electrical responses of single auditory nerve fibres or cochlear hair cells were recorded in the isolated half-head of the turtle Pseudemys scripta elegans. Responses to sound stimuli presented to the tympanum could be recorded for at least 4 hr after isolation.2. Impulses were recorded extracellularly from single auditory nerve fibres. For tones of suprathreshold intensity the impulses occurred with a preferred phase relation (i.e. they were phase-locked) to the cycles of the sound stimulus. Nerve fibres had sharp tuning curves (Q(10 db) = 0.5-7.5) with single characteristic frequencies (c.f.) ranging from about 30 to 700 Hz. Best threshold sensitivities of fibres at their c.f. were in the region of 30-40 db sound pressure level with respect to 20 muPa.3. Intracellular recordings were made from hair cells in the basilar papilla. Following injection of a fluorescent dye into a cell through the recording electrode, the dye was localized in a single hair cell in a transverse section of the cochlea.4. Hair cells had resting potentials of about -50 mV, and, to low frequency tones, gave periodic responses graded with the intensity and frequency of the stimulus. Recordings were obtained from cells with characteristic frequencies between 70 and 670 Hz.5. The voltage response to a pure tone at low sound pressure was sinusoidal for all frequencies of stimulation; at higher sound pressures a number of non-linearities were apparent in the response wave form. One of these was a steady depolarizing component, which, relative to the periodic component of the response, was most prominent at high frequencies.6. The amplitude of the response evoked in a hair cell by a low intensity tone was linearly related to the sound pressure; for loud sounds, the response eventually reached a saturating amplitude, which in some cells was as great as 30-45 mV peak-to-peak.7. The linear sensitivity of a hair cell is defined as the r.m.s. voltage for a linear response of the cell at its c.f. divided by the sound pressure at the tympanum. In the most sensitive cells this value was 30-90 mV/Pa.8. If the frequency selectivity of a hair cell was expressed in terms of the sound pressure needed to produce a constant amplitude of response, the sharpness of this frequency selectivity was found to be virtually independent of the response criterion for responses between 1 and 10 mV; in the cells which gave the largest responses, the frequency selectivity expressed in this way was comparable to that of the nerve fibres. Cells with smaller maximum responses often had broader tuning curves.9. Responses of hair cells to short low intensity tone bursts at the c.f. built up approximately exponentially during the tone, and decayed away exponentially when the tone was terminated. The terminal oscillations were at the c.f. of the cell, and independent of the frequency of stimulation.10. From the time constant of the build up and decay of the linear response to a tone burst at the c.f. the sharpness of tuning of the cell was estimated and found to agree with that obtained from the responses of the cell to continuous tones. The most highly tuned cells had quality factors (Q(3 db)) in the range 5-10.11. The c.f. of a hair cell was correlated with its position along the basilar membrane. Low frequency hair cells were located towards the apical or lagenar end and high frequency cells were found towards the basal or saccular end. On the assumption of an exponential distribution of c.f. with distance, each octave occupied about 94 mum along the membrane.12. A hair cell's response to a click was a decaying oscillation at the characteristic frequency of the cell. From the initial polarity of the responses to condensation and rarefaction clicks it was concluded that the hair cell depolarized as a result of movements of the basilar membrane towards the scala vestibuli, and hyperpolarized for motion towards the scala tympani.13. In the absence of deliberate sound stimulation, the hair cell voltage fluctuated continuously about its mean level. The principal frequency components in the noise were concentrated around the c.f. of the cell. The voltage noise in the hair cells showed no significant cross-correlation with sound pressure fluctuations at the tympanum.
https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b8eb/1282996/871baf664ad8/jphysiol00716-0129-a.jpg
https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b8eb/1282996/ea66b6742561/jphysiol00716-0128-a.jpg
https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b8eb/1282996/871baf664ad8/jphysiol00716-0129-a.jpg
https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b8eb/1282996/ea66b6742561/jphysiol00716-0128-a.jpg
https://cdn.ncbi.nlm.nih.gov/pmc/blobs/b8eb/1282996/871baf664ad8/jphysiol00716-0129-a.jpg
摘要
  1. 在隔离的秀丽锦龟半脑标本中记录了单根听神经纤维或耳蜗毛细胞的电反应。在隔离后,对施加于鼓膜的声音刺激的反应至少可记录4小时。

  2. 从单根听神经纤维进行细胞外冲动记录。对于阈上强度的纯音,冲动与声音刺激的周期具有优选的相位关系(即它们是锁相的)。神经纤维具有尖锐的调谐曲线(Q(10分贝)=0.5 - 7.5),单个特征频率(c.f.)范围约为30至700赫兹。纤维在其特征频率处的最佳阈值灵敏度相对于20微帕的声压级在30 - 40分贝的范围内。

  3. 从基底乳头的毛细胞进行细胞内记录。通过记录电极将荧光染料注入细胞后,染料定位在耳蜗横切面上的单个毛细胞中。

  4. 毛细胞的静息电位约为 - 50毫伏,对于低频纯音,会产生与刺激强度和频率分级相关的周期性反应。从特征频率在70至670赫兹之间的细胞获得了记录。

  5. 在低声压下,对所有刺激频率的纯音的电压反应都是正弦形的;在较高声压下,反应波形中出现了一些非线性现象。其中之一是稳定的去极化成分,相对于反应的周期性成分,在高频时最为突出。

  6. 低强度纯音在毛细胞中诱发的反应幅度与声压呈线性关系;对于强声音,反应最终达到饱和幅度,在一些细胞中,峰 - 峰值高达30 - 45毫伏。

  7. 毛细胞的线性灵敏度定义为细胞在其特征频率处线性反应的均方根电压除以鼓膜处的声压。在最敏感的细胞中,该值为30 - 90毫伏/帕。

  8. 如果用产生恒定反应幅度所需的声压来表示毛细胞的频率选择性,发现这种频率选择性的锐度实际上与1至10毫伏反应之间的反应标准无关;在产生最大反应的细胞中,以这种方式表示的频率选择性与神经纤维的相当。最大反应较小的细胞通常具有较宽的调谐曲线。

  9. 毛细胞对特征频率处短的低强度音爆的反应在音爆期间近似指数增长,并在音爆终止时指数衰减。终末振荡是细胞的特征频率,且与刺激频率无关。

  10. 根据对特征频率处音爆的线性反应的增长和衰减时间常数,估计了细胞调谐的锐度,发现与从细胞对连续纯音的反应中获得的结果一致。调谐最高的细胞的品质因数(Q(3分贝))在5 - 10的范围内。

  11. 毛细胞的特征频率与其沿基底膜的位置相关。低频毛细胞位于靠近顶端或壶腹端,高频细胞则靠近基部或球囊端。假设特征频率随距离呈指数分布,每个倍频程沿膜占据约94微米。

  12. 毛细胞对咔嗒声的反应是在细胞特征频率处的衰减振荡。根据对压缩和稀疏咔嗒声反应的初始极性得出结论,毛细胞因基底膜向前庭阶移动而发生去极化,因向前庭阶移动而发生超极化。

  13. 在没有故意声音刺激的情况下,毛细胞电压围绕其平均水平持续波动。噪声中的主要频率成分集中在细胞的特征频率附近。毛细胞中的电压噪声与鼓膜处的声压波动没有明显的互相关。

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