具有延时作用的基底膜主动耦合模型
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摘要
本文通过研究耳蜗中基底膜运动对不同频率和强度分辨能力,讨论其中外毛细胞自身运动具有的时延效应。为了探索这个问题,我们假设外毛细胞受到听力系统上层传出神经的反馈调控,通过建立具有外毛细胞双向耦合作用的基底膜振动模型,考查加入外毛细胞时延效应后对模型仿真效果的影响,例如基底膜的频率选择性,非线性等特点。
首先,在大量以往的对听力系统特别是耳蜗内毛细胞运动机制和效果的研究的基础上,根据中枢听觉系统的下行传导通路,也就是上橄榄复核的神经元对耳蜗毛细胞支配作用的认识,我们提出了支配外毛细胞的传出神经在外毛细胞电致运动(电刺激诱发细胞体长度改变)的过程中通过改变这一电致运动的速度,而起到听觉中枢对整个基底膜运动的反馈调控作用。其次,在具有外毛细胞双向耦合作用的基底膜振动模型上,加入外毛细胞电致运动的延时参数来进行建模仿真。通过仿真结果与实验数据进行对比,结果证明修正后的模型符合外毛细胞自身作用机制的生理特征,从一个新的角度诠释了外毛细胞运动对基底膜振动作用的贡献。
In this paper, the delay action of outer hair cells in its electromotility is discussed by investigating the frequency selectivity and intensity discrimination of basilar membrane vibration. To explor this question, we introduce a hypothesis that there is descending feedback of auditory efferent onto outer hair cells by proposing a novel cochlear amplifier model with delay action based on the cochlea's microanatomy, as well as outer hair cell (OHC) motility, to account for the cochlea's characteristic behavior such as the frequency selectivity and nolinear of basilar membrane.
Firstly, based on the research of auditory system especially the outer hair cells motility, according the descending efferent, as the dominant role of superior olive complex on cochlear hair cells, we propese the feedback control of descending efferent on basilar membrane by changing the speed in the electromotility of OHC. Then, the delay action-based mathematical cochlear model produced responses that are comparable to physiological measurements. The results illustrate a novel understanding of the OHC contribution on basilar membrane's characteristic behavior.
首先,在大量以往的对听力系统特别是耳蜗内毛细胞运动机制和效果的研究的基础上,根据中枢听觉系统的下行传导通路,也就是上橄榄复核的神经元对耳蜗毛细胞支配作用的认识,我们提出了支配外毛细胞的传出神经在外毛细胞电致运动(电刺激诱发细胞体长度改变)的过程中通过改变这一电致运动的速度,而起到听觉中枢对整个基底膜运动的反馈调控作用。其次,在具有外毛细胞双向耦合作用的基底膜振动模型上,加入外毛细胞电致运动的延时参数来进行建模仿真。通过仿真结果与实验数据进行对比,结果证明修正后的模型符合外毛细胞自身作用机制的生理特征,从一个新的角度诠释了外毛细胞运动对基底膜振动作用的贡献。
In this paper, the delay action of outer hair cells in its electromotility is discussed by investigating the frequency selectivity and intensity discrimination of basilar membrane vibration. To explor this question, we introduce a hypothesis that there is descending feedback of auditory efferent onto outer hair cells by proposing a novel cochlear amplifier model with delay action based on the cochlea's microanatomy, as well as outer hair cell (OHC) motility, to account for the cochlea's characteristic behavior such as the frequency selectivity and nolinear of basilar membrane.
Firstly, based on the research of auditory system especially the outer hair cells motility, according the descending efferent, as the dominant role of superior olive complex on cochlear hair cells, we propese the feedback control of descending efferent on basilar membrane by changing the speed in the electromotility of OHC. Then, the delay action-based mathematical cochlear model produced responses that are comparable to physiological measurements. The results illustrate a novel understanding of the OHC contribution on basilar membrane's characteristic behavior.
引文
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[4]Bekesy von G.:1961,'Concerning the Pleasures of Observing, and the Mechanics of the Inner Ear',in A. B. Soner (ed.), Les Prix Nobel en 1961, P. A. Norstedt, Stockholm, pp.184-208.
[5]Bekesy, von G.:1974,'Some Biophysical Experiments from Fifty Years Ago', Reprint from Annual Review of Physiology 36,1-16.
[6]R. G. Kessel and R. H. Kardon. Tissues and Organs:A Text-Atlas of Scanning Electron Microscopy. W. H. Freeman, New York,1979
[7]Barany, E., A contribution to the physiology of bone conduction. Acta Otol.,1938. Suppl. 26
[8]Bekesy, von G., Experiments in Hearing.1960, New York:John Wiley& Sons.
[9]Brownell WE, Bader CR, Bertrand D, et al. Evoked mechanical responses of isolated cochlear outer hair cells. Science,1985,227:194-196
[10]P. Dallos. Overview:Cochlear neurobiology. In P. Dallos, A. N. Popper, and R. R. Fay, editors, The Cochlea, Springer Handbook of Auditory Research. Springer-Verlag, New York,1996.
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[12]Galambos R, Davis H. Action potentials from single auditory fibers. Science,1948.108: 513.
[13]D. Purves. Neuroscience. Sinauer Associates.
[14]Wever E G. Theory of hearing. Am. J. Psychol.1951,64:626—628
[15]Rasmussen, G.L., The olivary peduncle and other fiber projections of the superior olivary comples. J Comp Neurol,1946.84:p.141-219
[16]王坚,姜涛,曾凡钢.听觉科学概论.2005.中国科学技术出版社.P：301
[17]Villa, A.E., et al., Corticofugal modulation of the information processing in the auditory thalamus of the cat. Exp Brain Res,1991.86(3):p.506—517
[18]Desmedt, J. E. and K. Mechelse, Suppression of acoustic input by thalamic stimulation. Proc Soc Exp Biol (NY),1958.99:p.772-775
[19]Sun, X. D., et al., Corticofugal influence on the responses of bat inferior collicular neurons to sound stimulation. Brain Res,1989.495(1):p,1-8
[20]Jen, PH., Q.C. Chen, and X. D. Sun, Corticofugal regulation of auditory sensitivity in the bat inferior colliculus. J Comp Physiol [A],1998.183(6):p.131-134
[21]Fex, J., Efferent inhibition in the cochlea related to hair-cell dc activity:study of postsynaptic activity of the crossed olivocochlear fibres in the cat. J Acoust Soc Am, 1967.41(3):p.666-675
[22]Rajan, R., Electrical stimulation of the inferior colliculus at low rates protects the cochlea from auditory desensitization. Brain Res,1990.506(2):p.192-204
[23]Xiao, Z. and N. Suga, Modulation of cochlear hair cells by the auditory cortex in the mustached bat. Nat Neurosci,2002.5(1):p.57-63
[24]Hallin, R. G. and H. E. Torebjork, Electrically induced A and C fiber responses in intact human skin nerves. Exp Brain Res,1973.16(3):p.309-320
[25]Fitzgerald, M. and C. J. Woolf, Effects of cutaneous nerve and intraspinal conditioning of C-fibre afferent terminal excitability in decerebrate spinal rats. J Physiol,1981.318:p. 25-39
[26]Gifford, M. L. and J.J.Guinan, Jr., Effects of electrical stimulation of medial olivochlear neurons on ipsilateral and contralateral cochlear responses. Hear Res,1987.29(2-3):p. 179-194
[27]Wiederhold, M.L. Variations in the effects of electric stimulation of the crossed olivocochlear bundle of cat single auditory nerve fiber responses to tone bursts. J Acoust Soc Am,1970.48:p.966-977
[28]Wiederhold, M.L. and N.Y. Kiang, Effects of electric stimulation of the crossed olivocochlear bundle on single auditory-nerve fibers in the cat. J Acoust Soc Am,1970. 48(4):p.950-965
[29]Yan, J. and G. Ehret, Corticofugal modulation of midbrain sound processing in the house mouse. Eur J Neurosci,2002.16(1):p.119-128.
[30]Brown, M. C., A.L. Nuttall, and R.I. Masta, Intracellular recordings from cochlear inner hair cells:Effects of stimulation of the crossed olivocochlear efferents.. Science,1983. 222:p.69-72
[31]Murugasu, E. and I.J. Russell, The effect of efferent stimulation on basilar membrane displacement in the basal turn of the guinea pig cochlea. J Neurosci,1996.16(1):p. 325-332
[32]Russell, I.J. and E. Murugasu, Medial efferent inhibition suppresses basilar membrane responses to near characteristic frequency tones of moderate to high intensities. J Acoust Soc Am,1997.102(6):p.1734-1738
[33]Dolan, D.F., M.H. Gou, and A.L. Nuttall, Frequency-dependent enhancement of basilar membrane velocity during olivocochlear bundle stimulation. J Acoust Soc Am,1997. 102(3):p.3587-3596
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