不同注意形式调节听感觉门控的神经机制

doi:10.3724/SP.J.1042.2020.01232

摘要/Abstract

摘要：

前脉冲抑制(prepulse inhibition, PPI)是听感觉门控的测量模型, 反映了听觉系统的早期信息选择功能。尽管PPI的主要神经环路位于脑干, 研究发现PPI可以被注意自上而下调节。然而, 已有研究并未区分不同注意(特征注意和空间注意)对PPI的特异性调节, 并且神经机制方面的研究集中于听皮层区域, 仍缺乏对皮层下机制的探讨。在以往研究的成果基础之上, 借助听觉信息加工的双通路模型, 采用行为测量、脑电和脑成像技术, 揭示特征和空间两种注意调节PPI的神经活动在听觉系统中的层次性神经表达。包括1)建立特征注意和空间注意调节PPI的统一行为模型, 考察两种注意调节PPI的时间动态性异同; 2)两种注意调节PPI的脑干分离机制, 即前脉冲刺激包络和精细结构成分加工在注意调节PPI中的作用差异; 3)两种注意调节PPI的关键脑区和脑网络差异。

关键词: 注意, 听感觉门控, 前脉冲抑制, 双通路模型

Abstract:

Prepulse inhibition (PPI) is considered as a measurement of sensory gating, reflecting the function of early auditory sensory information selection. Although the main neural correlates of PPI lie in the brainstem, previous researches have revealed that PPI could be top-down modulated by attention. However, different forms of attention, such as feature-based attention and spatial attention, have not been investigated together in the specific modulation of PPI. Moreover, the neural mechanism of attentional modulation of PPI in humans has been focused on the cortical level, whereas the subcortical mechanism has remained unclear. Based on the dual-pathway model for auditory processing, the present project aims to use behavioral testing, electroencephalogram recording and functional magnetic resonnance imaging, to investigate the various levels of neural representations of attentional modulation of PPI in the auditory system. The paper will 1) establish the behavioral paradigm of feature-based attentional and spatially attentional modulation of PPI, and examine the temporal dynamic processing differences between feature-based attentional modulation and spatially attentional modulation of PPI; 2) use brainstem frequency following responses (FFRs) to examine the subcortical neural mechanism of two types of attentional modulation of PPI, focusing on the envelope and fine structure processing differences of prepulse stimuli in the two types of attentional PPI modulation; 3) investigate the key brain areas and neural networks involved in feature-based attentional modulation and spatially attentional modulation of PPI.

Key words: attention, auditory sensory gating, prepulse inhibition, dual-pathway model

中图分类号:

B845

雷铭, 李朋波. (2020). 不同注意形式调节听感觉门控的神经机制. 心理科学进展 , 28(8), 1232-1245.

LEI Ming, LI Pengbo. (2020). Neural mechanism underlying the attentional modulation of auditory sensory gating. Advances in Psychological Science, 28(8), 1232-1245.

图/表 3

图1 惊反射和前脉冲抑制示意图。惊刺激单独呈现时, 惊反射幅度较大(上图); 惊刺激呈现之前的短时间内出现的前脉冲刺激会抑制随后出现的惊刺激引起的惊反射(下图), 被称为前脉冲抑制(PPI)

图2 知觉空间分离示意图。A声音(音乐符号)和B声音(宽波符号)之间物理关系(图a), 知觉空间分离(图b)和知觉空间重合(图c)

图3 本研究研究内容示意图

参考文献 82

[1]	杜忆, 李量. (2011). 对听感觉运动门控自上而下调节的动物模型和神经机制. 心理科学进展, 19(7), 944-958.
[2]	雷铭, 田晴, 王传跃, 李量. (2017). 精神疾病前脉冲抑制研究. 中华行为医学与脑科学杂志, 26(2), 188-192.
[3]	Aiken, S. J., & Picton, T. W. (2008). Envelope and spectral frequency-following responses to vowel sounds. Hearing Research, 245(1-2), 35-47. doi: 10.1016/j.heares.2008.08.004 URL pmid: 18765275
[4]	Akhoun, I., Gallégo, S., Moulin, A., Ménard, M., Veuillet, E., Bergervachon, C., ... Thaivan, H. (2008). The temporal relationship between speech auditory brainstem responses and the acoustic pattern of the phoneme /ba/ in normal-hearing adults. Clinical Neurophysiology, 119(4), 922-933. doi: 10.1016/j.clinph.2007.12.010 URL pmid: 18291717
[5]	Ananthakrishnan, S., Krishnan, A., & Bartlett, E. (2016). Human frequency following response: Neural representation of envelope and temporal fine structure in listeners with normal hearing and sensorineural hearing Loss. Ear and Hearing, 37(2), e91-e103. doi: 10.1097/AUD.0000000000000247 URL pmid: 26583482
[6]	Arnott, S. R., Binns, M. A., Grady, C. L., & Alain, C. (2004). Assessing the auditory dual-pathway model in humans. NeuroImage, 22(1), 401-408. URL pmid: 15110033
[7]	Blumenthal, T. D., Reynolds, J. Z., & Spence, T. E. (2015). Support for the interruption and protection hypotheses of prepulse inhibition of startle: Evidence from a modified attention network test. Psychophysiology, 52(3), 397-406. doi: 10.1111/psyp.12334 URL pmid: 25234706
[8]	Braff, D., Stone, C., Callaway, E., Geyer, M., Glick, I., & Bali, L. (1978). Prestimulus effects on human startle reflex in normals and schizophrenics. Psychophysiology, 15(4), 339-343. doi: 10.1111/j.1469-8986.1978.tb01390.x URL pmid: 693742
[9]	Campbell, L. E., Hughes, M., Budd, T. W., Cooper, G., Fulham, W. R., Karayanidis, F., ... Schall, U. (2007). Primary and secondary neural networks of auditory prepulse inhibition: A functional magnetic resonance imaging study of sensorimotor gating of the human acoustic startle response. European Journal of Neuroscience, 26(8), 2327-2333. doi: 10.1111/j.1460-9568.2007.05858.x URL pmid: 17908169
[10]	Chandrasekaran, B., & Kraus, N. (2010). The scalp-recorded brainstem response to speech: Neural origins and plasticity. Psychophysiology, 47(2), 236-246. doi: 10.1111/j.1469-8986.2009.00928.x URL pmid: 19824950
[11]	Chen, J., He, Y., Zhu, Z., Zhou, T., Peng, Y., Zhang, X., & Fang, F. (2014). Attention-dependent early cortical suppression contributes to crowding. The Journal of Neuroscience, 34(32), 10465-10474. doi: 10.1523/JNEUROSCI.1140-14.2014 URL pmid: 25100582
[12]	Chimento, T. C., & Schreiner, C. E. (1990). Selectively eliminating cochlear microphonic contamination from the frequency-following response. Electroencephalography and Clinical Neurophysiology, 75(1-2), 88-96.
[13]	Coffey, E. B., Musacchia, G., & Zatorre, R. J. (2017). Cortical correlates of the auditory frequency-following and onset responses: EEG and fMRI evidence. Journal of Neuroscience, 37(4), 830-838. doi: 10.1523/JNEUROSCI.1265-16.2016 URL pmid: 28123019
[14]	Crowley, K. E., & Colrain, I. M. (2004). A review of the evidence for P2 being an independent component process: Age, sleep and modality. Clinical Neurophysiology, 115(4), 732-744. doi: 10.1016/j.clinph.2003.11.021 URL pmid: 15003751
[15]	Csomor, P. A., Preller, K. H., Geyer, M. A., Studerus, E., Huber, T., & Vollenweider, F. X. (2014). Influence of aripiprazole, risperidone, and amisulpride on sensory and sensorimotor gating in healthy “low and high gating” humans and relation to psychometry. Neuropsychopharmacology, 39(10), 2485-2496. doi: 10.1038/npp.2014.102 URL pmid: 24801767
[16]	Davis, M. (2006). Neural systems involved in fear and anxiety measured with fear-potentiated startle. American Psychologist, 61(8), 741-752. doi: 10.1037/0003-066X.61.8.741 URL pmid: 17115806
[17]	Dawson, M. E., Hazlett, E. A., Filion, D. L., Nuechterlein, K. H., & Schell, A. M. (1993). Attention and schizophrenia: Impaired modulation of the startle reflex. Journal of Abnormal Psychology, 102(4), 633-641. doi: 10.1037//0021-843x.102.4.633 URL pmid: 8282934
[18]	Dawson, M. E., Schell, A. M., Hazlett, E. A., Nuechterlein, K. H., & Filion, D. L. (2000). On the clinical and cognitive meaning of impaired sensorimotor gating in schizophrenia. Psychiatry Research, 96(3), 187-197. doi: 10.1016/s0165-1781(00)00208-0 URL pmid: 11084215
[19]	de Haan, E. H. F., & Cowey, A. (2011). On the usefulness of “what” and “where” pathways in vision. Trends in Cognitive Sciences, 15(10), 460-466. doi: 10.1016/j.tics.2011.08.005 URL pmid: 21906989
[20]	Ding, N., Melloni, L., Zhang, H., Tian, X., & Poeppel, D. (2016). Cortical tracking of hierarchical linguistic structures in connected speech. Nature Neuroscience, 19(1), 158-164. doi: 10.1038/nn.4186 URL pmid: 26642090
[21]	Ding, N., Patel, A.D., Chen, L., Butler, H., Luo, C., & Poeppel, D. (2017). Temporal modulations in speech and music. Neuroscience and Biobehavioral Reviews, 81, 181-197. doi: 10.1016/j.neubiorev.2017.02.011 URL pmid: 28212857
[22]	Ding, Y., Xu, N., Gao, Y., Wu, Z., & Li, L. (2019). The role of the deeper layers of the superior colliculus in attentional modulations of prepulse inhibition. Behavioural Brain Research, 364, 106-113. doi: 10.1016/j.bbr.2019.01.052 URL pmid: 30707906
[23]	Du, Y., Kong, L., Wang, Q., Wu, X., & Li, L. (2011a). Auditory frequency-following response: A neurophysiological measure for studying the "cocktail-party problem". Neuroscience and Biobehavioral Reviews, 35(10), 2046-2057. doi: 10.1016/j.neubiorev.2011.05.008 URL pmid: 21645541
[24]	Du, Y., Li, J. Y., Wu, X. H., & Li, L. (2009). Precedence-effect-induced enhancement of prepulse inhibition in socially reared but not isolation-reared rats. Cognitive Affective & Behavioral Neuroscience, 9, 44-58.
[25]	Du, Y., Wu, X., & Li, L. (2010). Emotional learning enhances stimulus-specific top-down modulation of sensorimotor gating in socially reared rats but not isolation-reared rats. Behavioural Brain Research, 206(2), 192-201. doi: 10.1016/j.bbr.2009.09.012 URL pmid: 19761801
[26]	Du, Y., Wu, X., & Li, L. (2011b). Differentially organized top-down modulation of prepulse inhibition of startle. The Journal of Neuroscience, 31(38), 13644-13653. doi: 10.1523/JNEUROSCI.1292-11.2011 URL pmid: 21940455
[27]	Filion, D. L., Dawson, M. E., & Schell, A. M. (1998). The psychological significance of human startle eyeblink modification: A review. Biological Psychology, 47(1), 1-43. doi: 10.1016/s0301-0511(97)00020-3 URL pmid: 9505132
[28]	Freyman, R. L., Balakrishnan, U., & Helfer, K. S. (2004). Effect of number of masking talkers and auditory priming on informational masking in speech recognition. The Journal of the Acoustical Society of America, 115(5), 2246-2256. doi: 10.1121/1.1689343 URL
[29]	Freyman, R. L., Balakrishnan, U., & Helfer, K. S. (2008). Spatial release from masking with noise-vocoded speech. The Journal of the Acoustical Society of America, 124(3), 1627-1637. doi: 10.1121/1.2951964 URL pmid: 19045654
[30]	Fritz, J. B., Elhilali, M., David, S. V., & Shamma, S. A. (2007). Auditory attention—focusing the searchlight on sound. Current Opinion in Neurobiology, 17(4), 437-455. doi: 10.1016/j.conb.2007.07.011 URL pmid: 17714933
[31]	Galbraith, G. C. (1994). Two-channel brain-stem frequency- following responses to pure tone and missing fundamental stimuli. Electroencephalography and Clinical Neurophysiology, 92(4), 321-330. URL pmid: 7517854
[32]	Galbraith, G. C., Olfman, D. M., & Huffman, T. M. (2003). Selective attention affects human brain stem frequency- following response. Neuroreport, 14(5), 735-738. doi: 10.1097/00001756-200304150-00015 URL pmid: 12692473
[33]	Glaser, E. M., Suter, C. M., Dasheiff, R., & Goldberg, A. (1976). The human frequency-following response: Its behavior during continuous tone and tone burst stimulation. Electroencephalography and Clinical Neurophysiology, 40(1), 25-32. doi: 10.1016/0013-4694(76)90176-0 URL pmid: 55345
[34]	Goodale, M. A., & Milner, A. D. (1992). Separate Visual Pathways for Perception and Action. Trends in Neurosciences, 15(1), 20-25. doi: 10.1016/0166-2236(92)90344-8 URL pmid: 1374953
[35]	Graham, F. (1975). The more or less startling effects of weak prestimulation. Psychophysiology, 12(3), 238-248. doi: 10.1111/j.1469-8986.1975.tb01284.x URL pmid: 1153628
[36]	Hairston, W. D., Letowski, T. R., & Mcdowell, K. (2013). Task-related suppression of the brainstem frequency following response. PloS One, 8(2), e55215. doi: 10.1371/journal.pone.0055215 URL pmid: 23441150
[37]	Hazlett, E. A., Buchsbaum, M. S., Tang, C. Y., Fleischman, M. B., Wei, T. C., Byne, W., & Haznedar, M. M. (2001). Thalamic activation during an attention-to-prepulse startle modification paradigm: A functional MRI study. Biological Psychiatry, 50(4), 281-291. doi: 10.1016/s0006-3223(01)01094-0 URL pmid: 11522263
[38]	Hazlett, E. A., Levine, J., Buchsbaum, M. S., Silverman, J. M., New, A., Sevin, E. M., ... Siever, L. J. (2003). Deficient attentional modulation of the startle response in patients with schizotypal personality disorder. American Journal of Psychiatry, 160(9), 1621-1626. doi: 10.1176/appi.ajp.160.9.1621 URL pmid: 12944337
[39]	Hazlett, E. A., Romero, M. J., Haznedar, M. M., New, A. S., Goldstein, K. E., Newmark, R. E., ... Buchsbaum, M. S. (2007). Deficient attentional modulation of startle eyeblink is associated with symptom severity in the schizophrenia spectrum. Schizophrenia Research, 93(1-3), 288-295. doi: 10.1016/j.schres.2007.03.012 URL pmid: 17478083
[40]	Javitt, D. C., & Sweet, R. A. (2015). Auditory dysfunction in schizophrenia: Integrating clinical and basic features. Nature Reviews Neuroscience, 16(9), 535-550. doi: 10.1038/nrn4002 URL pmid: 26289573
[41]	Jones, L. A., Hills, P. J., Dick, K. M., Jones, S. P., & Bright, P. (2016). Cognitive mechanisms associated with auditory sensory gating. Brain and Cognition, 102, 33-45. doi: 10.1016/j.bandc.2015.12.005 URL pmid: 26716891
[42]	Kraus, N., Anderson, S., & White-Schwoch, T. (2017). The Frequency-following response: A window into human communication. Boston, MA: Springer International Publishing.
[43]	Krishnan, A., Xu, Y., Gandour, J. T., & Cariani, P. A. (2004). Human frequency-following response: Representation of pitch contours in Chinese tones. Hearing Research, 189(1-2), 1-12. doi: 10.1016/S0378-5955(03)00402-7 URL pmid: 14987747
[44]	Kumari, V., Antonova, E., & Geyer, M. A. (2008). Prepulse inhibition and "psychosis-proneness" in healthy individuals: An fMRI study. European Psychiatry, 23(4), 274-280. URL pmid: 18222069
[45]	Landis, C., & Hunt, W. A. (Eds.). (1939). The startle pattern. New York: Farrar and Rinehart.
[46]	Lehmann, A., & Schönwiesner, M. (2014). Selective attention modulates human auditory brainstem responses: Relative contributions of frequency and spatial cues. PloS One, 9(1), e85442. doi: 10.1371/journal.pone.0085442 URL pmid: 24454869
[47]	Lei, M., Luo, L., Qu, T., Jia, H., & Li, L. (2014). Perceived location specificity in perceptual separation-induced but not fear conditioning-induced enhancement of prepulse inhibition in rats. Behavioural Brain Research, 269, 87-94. doi: 10.1016/j.bbr.2014.04.030 URL pmid: 24780867
[48]	Lei, M., Zhang, C., & Li, L. (2018). Neural correlates of perceptual separation-induced enhancement of prepulse inhibition of startle in humans. Scientific Reports, 8(1), 1-10. doi: 10.1038/s41598-017-17765-5 URL pmid: 29311619
[49]	Li, L., Du, Y., Li, N., Wu, X., & Wu, Y. (2009). Top-down modulation of prepulse inhibition of the startle reflex in humans and rats. Neuroscience & Biobehavioral Reviews, 33(8), 1157-1167. doi: 10.1016/j.neubiorev.2009.02.001 URL pmid: 19747594
[50]	Li, L., Qi, J. G., He, Y., Alain, C., & Schneider, B. A. (2005). Attribute capture in the precedence effect for long-duration noise sounds. Hearing Research, 202(1-2), 235-247. doi: 10.1016/j.heares.2004.10.007 URL pmid: 15811715
[51]	Luo, L., Wang, Q., & Li, L. (2017). Neural representations of concurrent sounds with overlapping spectra in rat inferior colliculus: Comparisons between temporal-fine structure and envelope. Hearing Research, 353, 87-96. doi: 10.1016/j.heares.2017.06.005 URL pmid: 28655419
[52]	Marsh, J. T., Worden, F. G., & Smith, J. C. (1970). Auditory frequency-following response: Neural or artifact? Science, 169(3951), 1222-1223. doi: 10.1126/science.169.3951.1222 URL pmid: 5450700
[53]	Meng, Q., Ding, Y., Chen, L., & Li, L. (2020). The medial agranular cortex mediates attentional enhancement of prepulse inhibition of the startle reflex. Behavioural Brain Research, 383, 112511. doi: 10.1016/j.bbr.2020.112511 URL pmid: 31987934
[54]	Musacchia, G., Sams, M., Skoe, E., & Kraus, N. (2007). Musicians have enhanced subcortical auditory and audiovisual processing of speech and music. Proceedings of the National Academy of Sciences of the United States of America, 104(40), 15894-15898. doi: 10.1073/pnas.0701498104 URL pmid: 17898180
[55]	Näätänen, R., & Picton, T. (1987). The N1 wave of the human electric and magnetic response to sound: A review and an analysis of the component structure. Psychophysiology, 24(4), 375-425. doi: 10.1111/j.1469-8986.1987.tb00311.x URL pmid: 3615753
[56]	Ping, J., Li, N., Galbraith, G. C., Wu, X., & Li, L. (2008). Auditory frequency-following responses in rat ipsilateral inferior colliculus. Neuroreport, 19(14), 1377-1380. URL pmid: 18766015
[57]	Poje, A. B., & Filion, D. L. (2017). Erratum to: The effects of multiphasic prepulses on automatic and attention- modulated prepulse inhibition. Cognitive Processing, 18, 271. doi: 10.1007/s10339-017-0823-8 URL pmid: 28616664
[58]	Romanski, L. M., Tian, B., Fritz, J., Mishkin, M., Goldman-Rakic, P. S., & Rauschecker, J. P. (1999). Dual streams of auditory afferents target multiple domains in the primate prefrontal cortex. Nature Neuroscience, 2(12), 1131-1136. doi: 10.1038/16056 URL
[59]	Röskam, S., & Koch, M. (2006). Enhanced prepulse inhibition of startle using salient prepulses in rats. International Journal of Psychophysiology, 60(1), 10-14. doi: 10.1016/j.ijpsycho.2005.04.004 URL
[60]	Rajji, T. K., & Mulsant, B. H. (2008). Nature and course of cognitive function in late-life schizophrenia: A systematic review. Schizophrenia Research, 102(1-3), 122-140. URL pmid: 18468868
[61]	Russo, N., Nicol, T., Musacchia, G., & Kraus, N. (2004). Brainstem responses to speech syllables. Clinical Neurophysiology, 115(9), 2021-2033. doi: 10.1016/j.clinph.2004.04.003 URL pmid: 15294204
[62]	Skoe, E., & Kraus, N. (2012). A little goes a long way: How the adult brain is shaped by musical training in childhood. Journal of Neuroscience, 32(34), 11507-11510. URL pmid: 22915097
[63]	Smith, C. W., & Cornblatt, B. (2005). Attention deficits in the development of schizophrenia: Recent evidence from genetic high-risk and prodromal studies. Current Psychosis & Therapeutics Reports, 3(4), 152-156.
[64]	Smith, Z. M., Delgutte, B., & Oxenham, A. J. (2002). Chimaeric sounds reveal dichotomies in auditory perception. Nature, 416(6876), 87-90. doi: 10.1038/416087a URL pmid: 11882898
[65]	Song, J., Skoe, E., Wong, P., & Kraus, N. (2008). Plasticity in the adult human auditory brainstem following short-term linguistic training. Journal of Cognitive Neuroscience, 20(10), 1892-1902. URL pmid: 18370594
[66]	Song, J. H., Skoe, E., Banai, K., & Kraus, N. (2012). Training to improve hearing speech in noise: Biological mechanisms. Cerebral Cortex, 22(5), 1180-1190.
[67]	Song, K., Meng, M., Chen, L., Zhou, K., & Luo, H. (2014). Behavioral oscillations in attention: Rhythmic α pulses mediated through θ band. Journal of Neuroscience, 34(14), 4837-4840.
[68]	Swerdlow, N. R., Karban, B., Ploum, Y., Sharp, R., Geyer, M. A., & Eastvold, A. (2001). Tactile prepuff inhibition of startle in children with Tourette’s syndrome: In search of an “fMRI-friendly” startle paradigm. Biological Psychiatry, 50(8), 578-585.
[69]	Swerdlow, N. R., Weber, M., Qu, Y., Light, G. A., & Braff, D. L. (2008). Realistic expectations of prepulse inhibition in translational models for schizophrenia research. Psychopharmacology, 199(3), 331-388. doi: 10.1007/s00213-008-1072-4 URL pmid: 18568339
[70]	Wang, Q., & Li, L. (2017). Differences between auditory frequency-following responses and onset responses: Intracranial evidence from rat inferior colliculus. Hearing Research, 357, 25-32. URL pmid: 29156225
[71]	Wang, Y., & Luo, H. (2017). Behavioral oscillation in face priming: Prediction about face identity is updated at a theta-band rhythm. Progress in Brain Research, 36(1), 84-90.
[72]	Wilson, J. R., & Krishnan, A. (2005). Human frequency- following responses to binaural masking level. Journal of the American Academy of Audiology, 16(3), 184-192. URL pmid: 15844743
[73]	Woods, D. L., Knight, R. T., & Scabini, D. (1993). Anatomical substrates of auditory selective attention: Behavioral and electrophysiological effects of posterior association cortex lesions. Brain Research Cognitive Brain Research, 1(4), 227-232.
[74]	Wu, C., Zheng, Y., Li, J., Wu, H., She, S., Liu, S., ... Li, L. (2016). Brain substrates underlying auditory speech priming in healthy listeners and listeners with schizophrenia. Psychological Medicine, 1-11. doi: 10.1017/S0033291720002366 URL pmid: 32684185
[75]	Wu, M., Li, H., Gao, Y., Lei, M., Teng, X., Wu, X., & Li, L. (2012). Adding irrelevant information to the content prime reduces the prime-induced unmasking effect on speech recognition. Hearing Research, 283(1), 136-143.
[76]	Wu, X., Wang, C., Chen, J., Qu, H., Li, W., Wu, Y., ... Li, L. (2005). The effect of perceived spatial separation on informational masking of Chinese speech. Hearing Research, 199(1-2), 1-10. URL pmid: 15574295
[77]	Wu, Z. M., Ding, Y., Jia, H. X., & Li, L. (2016). Different effects of isolation-rearing and neonatal MK-801 treatment on attentional modulations of prepulse inhibition of startle in rats. Psychopharmacology, 233(17), 1-14.
[78]	Yang, N. B., Tian, Q., Fan, Y., Bo, Q. J., Zhang, L., Li, L., & Wang, C. Y. (2017). Deficits of perceived spatial separation induced prepulse inhibition in patients with schizophrenia: Relationships to symptoms and neurocognition. BMC Psychiatry, 17(1), 135-142. doi: 10.1186/s12888-017-1276-4 URL pmid: 28399842
[79]	Yeomans, J. S., Li, L., Scott, B. W., & Frankland, P. W. (2002). Tactile, acoustic and vestibular systems sum to elicit the startle reflex. Neuroscience and Biobehavioral Reviews, 26(1), 1-11. URL pmid: 11835980
[80]	Zhang, C., Lu, L., Wu, X., & Li, L. (2014). Attentional modulation of the early cortical representation of speech signals in informational or energetic masking. Brain and Language, 135, 85-95.
[81]	Zheng, Y., Wu, C., Li, J., Wu, H., She, S., Liu, S., ... Li, L. (2016). Brain substrates of perceived spatial separation between speech sources under simulated reverberant listening conditions in schizophrenia. Psychological Medicine, 46(3), 477-483. doi: 10.1017/S0033291715001828 URL pmid: 26423774
[82]	Zündorf, I. C., Lewald, J., & Karnath, H.-O. (2016). Testing the dual-pathway model for auditory processing in human cortex. NeuroImage, 124, 672-681. URL pmid: 26388552