Location Signaling by Cortical Neurons

  • John C. Middlebrooks
  • Li Xu
  • Shigeto Furukawa
  • Brian J. Mickey
Part of the Springer Handbook of Auditory Research book series (SHAR, volume 15)


Localization is a fundamental task of hearing. Identification of the locations of sound sources permits animals to locate prey or to avoid predators. Spatial hearing improves signal detection and aids in the segregation of multiple sources, thus improving identification of sounds. The importance of sound localization for the auditory system is reflected in the organization of the auditory brainstem, which contains several discrete nuclei that appear to be specialized for analysis of particular acoustical cues for sound source location.


Receptive Field Sound Source Auditory Cortex Sound Level Primary Auditory Cortex 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Ahissar M, Ahissar E, Bergman H, Vaadia E (1992) Encoding of sound-source location and movement-activity of single neurons and interactions between adjacent neurons in the monkey auditory cortex. J Neurophysiol 67: 203–215.PubMedGoogle Scholar
  2. Andersen P, Knight L, Merzenich MM (1980) The thalamocortical and corticothalmic connections of AI, AII, and the anterior auditory field (AFF) in the cat: Evidence for two largely segregated systems of connections. J Comp Neurol 194: 663–701.PubMedCrossRefGoogle Scholar
  3. Barlow HB (1972) Single units and sensation: A neuron doctrine for perceptual psychology? Perception 1: 371–394.PubMedCrossRefGoogle Scholar
  4. Barone P, Clarey JC, Irons WA, Imig TJ (1996) Cortical synthesis of azimuth-sensitive single-unit responses with nonmonotonic level tuning: A thalamocortical comparison in the cat. J Neurophysiol 75: 1206–1220.PubMedGoogle Scholar
  5. Beitel RE, Kaas JH (1993) Effects of bilateral and unilateral ablation of auditory cortex in cats on the uniconditioned head orienting response to acoustic stimuli. J Neurophysiol 70: 351–369.PubMedGoogle Scholar
  6. Benson DA, Hienz RD, Goldstein Jr MH (1981) Single-unit activity in the auditory cortex of monkeys actively localizing sound sources: Spatial tuning and behavioral dependency. Brain Res 219: 249–267.PubMedCrossRefGoogle Scholar
  7. Blauert J (1969–1970) Sound localization in the median plane. Acustica 22:205–213.Google Scholar
  8. Blauert J (1983) Spatial Hearing. Cambridge, MA: MIT Press.Google Scholar
  9. Brown CH, Schessler T, Moody D, Stebbins W (1982) Vertical and horizontal sound localization in primates. J Acoust Soc Am 72: 1804–1811.PubMedCrossRefGoogle Scholar
  10. Brugge JF, Reale RA, Hind JE, Chan JCK, Musicant AD, Poon PWF (1994) Simulation of free-field sound sources and its application to studies of cortical mechanisms of sound localization in the cat. Hear Res 73: 67–84.PubMedCrossRefGoogle Scholar
  11. Brugge JF, Reale RA, Hind JE (1996) The structure of spatial receptive fields of neurons in primary auditory cortex of the cat. J Neurosci 16: 4420–4437.PubMedGoogle Scholar
  12. Bushara KO, Weeks RA, Ishii K, Catalan M, Tian B, Rauschecker JP, Hallett M (1999) Modality-specific frontal and parietal areas for auditory and visual spatial localization in humans. Nat Neurosci 2: 759–766.PubMedCrossRefGoogle Scholar
  13. Clarey JC, Irvine DRF (1990) The anterior ectosylvian auditory field in the cat: I. An electrophysiological study of its relationship to surrounding auditory cortical fields. J Comp Neurol 301: 289–303.PubMedCrossRefGoogle Scholar
  14. Clarey JC, Barone P, Imig T (1992) Physiology of Thalamus and Cortex. In: Popper AN, Fay RR (eds) The Mammalian Auditory Pathway: Neurophysiology. New York: Springer-Verlag, pp. 232–334.CrossRefGoogle Scholar
  15. Clarey JC, Barone P, Imig TJ (1994) Functional organization of sound direction and sound pressure level in primary auditory cortex of the cat. J Neurophysiol 72: 2383–2405.PubMedGoogle Scholar
  16. Efron B, Tibshirani R (1991) Statistical data analysis in the computer age. Science 253: 390–395.PubMedCrossRefGoogle Scholar
  17. Eggermont JJ (1998) Azimuth coding in primary auditory cortex of the cat. II. Relative latency and interspike interval representation. J Neurophysiol 80: 2151–2161.PubMedGoogle Scholar
  18. Eggermont JJ, Mossop JE (1998) Azimuth coding in primary auditory cortex of the cat. I. Spike synchrony versus spike count representations. J Neurophysiol 80: 2133–2150.PubMedGoogle Scholar
  19. Eisenman LM (1974) Neural encoding of sound location: An electrophysiological study in auditory cortex (AI) of the cat using free field stimuli. Brain Res 75: 203–214.PubMedCrossRefGoogle Scholar
  20. Evans EF (1967) Cortical representation. In: de Reuck AUS, Knight J (eds) Symposium on Hearing Mechanisms in Vertebrates. London: CIBA, Churchill, pp. 272–295.Google Scholar
  21. Furukawa S, Xu L, Middlebrooks JC (2000) Coding of sound-source location by ensembles of cortical neurons. J Neurosci 20: 1216–1228.PubMedGoogle Scholar
  22. Georgopoulos AP, Schwartz AB, Kettner RE (1986) Neuronal population coding of movement direction. Science 233: 1416–1419.PubMedCrossRefGoogle Scholar
  23. Greene TC (1929) The ability to localize sound: a study of binaural hearing in patients with tumor of the brain. Arch Surg 18: 1825–1841.CrossRefGoogle Scholar
  24. Grunewald A, Linden JF, Andersen RA (1999) Responses to auditory stimuli in macaque lateral intraparietal area. I. Effects of training. J Neurophysiol 82: 330–342.PubMedGoogle Scholar
  25. Hebrank J, Wright D (1974) Spectral cues used in the localization of sound sources on the median plane. J Acoust Soc Am 56: 1829–1834.PubMedCrossRefGoogle Scholar
  26. Heffner HE, Heffner RS (1990) Effect of bilateral auditory cortex lesions on sound localization in Japanese macaques. J Neurophysiol 64: 915–931.PubMedGoogle Scholar
  27. Heffner RS, Heffner HE (1988) Sound localization acuity in the cat: Effect of azimuth, signal duration, and test procedure. Hear Res 36: 221–232.PubMedCrossRefGoogle Scholar
  28. Huang AY, May BJ (1996) Sound orientation behavior in cats. II. Mid-frequency spectral cues for sound localization. J Acoust Soc Am 100: 1070–1080.PubMedCrossRefGoogle Scholar
  29. Imig TJ, Irons WA, Samson FR (1990) Single-unit selectivity to azimuthal direction and sound pressure level of noise bursts in cat high-frequency primary auditory cortex. J Neurophysiol 63: 1448–1466.PubMedGoogle Scholar
  30. Jenison RL (1998) Models of direction estimation with spherical-function approximated cortical receptive fields. In: Brugge JF, Poon PWF (eds) Central Auditory Processing and Neural Modeling. New York: Plenum, pp. 161–174.CrossRefGoogle Scholar
  31. Jenison RL, Reale RA, Hind JE, Brugge JF (1998) Modeling of auditory spatial receptive fields with spherical approximation functions. J Neurophysiol 80: 2645–2656.PubMedGoogle Scholar
  32. Jenkins WM, Masterton RB (1982) Sound localization: Effects of unilateral lesions in central auditory system. J Neurophysiol 47: 987–1016.PubMedGoogle Scholar
  33. Jenkins WM, Merzenich MM (1984) Role of cat primary auditory cortex for sound-localization behavior. J Neurophysiol 52: 819–847.PubMedGoogle Scholar
  34. Kavanagh GL, Kelly JB (1987) Contribution of auditory cortex to sound localization by the ferret (Mustela putorius). J Neurophysiol 57: 1746–1766.PubMedGoogle Scholar
  35. Klingon GH, Bontecou DC (1966) Localization in auditory space. Neurology 16: 879–886.CrossRefGoogle Scholar
  36. Korte M, Rauschecker JP (1993) Auditory spatial tuning of cortical neurons is sharpened in cats with early blindness. J Neurophysiol 70: 1717–1721.PubMedGoogle Scholar
  37. Linden JF, Grunewald A, Andersen RA (1999) Responses to auditory stimuli in macaque lateral intraparietal area. II. Behavioral modulation. J Neurophysiol 82: 343–358.PubMedGoogle Scholar
  38. Litovsky RY, Colburn HS, Yost WA, Guzman SJ (1999) The precedence effect. J Acoust Soc Am 106: 1633–1654.PubMedCrossRefGoogle Scholar
  39. Makous JC, Middlebrooks JC (1990) Two-dimensional sound localization by human listeners. J Acoust Soc Am 87: 2188–2200.PubMedCrossRefGoogle Scholar
  40. May BJ, Huang AY (1996) Sound orientation behavior in cats. I. Localization of broadband noise. J Acoust Soc Am 100: 1059–1069.PubMedCrossRefGoogle Scholar
  41. Mazzoni P, Bracewell RM, Barash S, Andersen RA (1996) Spatially tuned auditory responses in area LIP of macaques performing delayed memory saccades to acoustic targets. J Neurophysiol 75: 1233–1241.PubMedGoogle Scholar
  42. Meredith MA, Clemo HR (1989) Auditory cortical projection from the anterior ectosylvian sulcus (field AES) to the superior colliculus in the cat: An anatomical and electrophysiological study. J Comp Neurol 289: 687–707.PubMedCrossRefGoogle Scholar
  43. Merzenich MM, Knight PL, Roth GL (1973) Cochleotopic organization of primary auditory cortex in the cat. Brain Res 63: 343–346.PubMedCrossRefGoogle Scholar
  44. Mickey BJ, Middlebooks JC (2001) Responses of auditory cortical neurons to pairs of sounds: correlates of fusion and localization. J Neurophysiol 86: 1333–1350.PubMedGoogle Scholar
  45. Middlebrooks JC (1992) Narrow-band sound localization related to external ear acoustics. J Acoust Soc Am 92: 2607–2624.PubMedCrossRefGoogle Scholar
  46. Middlebrooks JC (1998) Location coding by auditory cortical neurons. In: Poon PWF, Brugge JF (eds) Central auditory processing and neural modeling. New York: Plenium Press, pp. 139–148.CrossRefGoogle Scholar
  47. Middlebrooks JC, Green DM (1991) Sound localization by human listeners. Ann Rev Psychol 42: 135–159.CrossRefGoogle Scholar
  48. Middlebrooks JC, Knudsen EI (1984) A neural code for auditory space in the cat’s superior colliculus. J Neurosci 4: 2621–2634.PubMedGoogle Scholar
  49. Middlebrooks JC, Pettigrew JD (1981) Functional classes of neurons in primary auditory cortex of the cat distinguished by sensitivity to sound location. J Neurosci 1: 107–120.PubMedGoogle Scholar
  50. Middlebrooks JC, Xu L, Eddins AC, Green DM (1998) Codes for sound-source location in nontonotopic auditory cortex. J Neurophysiol 80: 863–881.PubMedGoogle Scholar
  51. Mills AW (1958) On the minimum audible angle. J Acoust Soc Am 30: 237–246.CrossRefGoogle Scholar
  52. Mishkin M, Ungerleider LG, Macko KA (1983) Object vision and spatial vision: two cortical pathways. TINS 414–417.Google Scholar
  53. Musicant AD, Chan JCK, Hind JE (1990) Direction-dependent spectral properties of cat external ear: New data and cross-species comparisons. J Acoust Soc Am 87: 757–781.PubMedCrossRefGoogle Scholar
  54. Palmer AR, King AJ (1982) The representation of auditory space in the mammalian superior colliculus. Nature 299: 248–249.PubMedCrossRefGoogle Scholar
  55. Pinek B, Duhamel JR, Cave C, Brouchon M (1989) Audio-Spatial deficits in humans: Differential effects associated with left versus right hemisphere parietal damage. Cortex 25: 175–186.PubMedGoogle Scholar
  56. Poirier P, Lassonde M, Villemure J-G, Geoffroy G, Lepore F (1994) Sound localization in hemispherectomized patients. Neuropsychologica 32: 541–553.CrossRefGoogle Scholar
  57. Poirier P, Jiang H, Lepore F, Guillemot JP (1997) Positional, directional and speed selectivities in the primary auditory cortex of the cat. Hear Res 113: 1–13.PubMedCrossRefGoogle Scholar
  58. Populin LC, Yin TC (1998) Behavioral studies of sound localization in cat. J Neurosci 18: 2147–2160.PubMedGoogle Scholar
  59. Rajan R, Aitkin LM, Irvine DRF, McKay J (1990a) Azimuthal sensitivity of neurons in primary auditory cortex of cats. I. Types of sensitivity and the effects of variations in stimulus parameters. J Neurophysiol 64: 872–887.Google Scholar
  60. Rajan R, Aitkin LM, Irvine DRF (1990b) Azimuthal sensitivity of neurons in primary auditory cortex of cats. II. Organization along frequency-band strips. J Neurophysiol 64: 888–902.Google Scholar
  61. Rauschecker JP (1998) Parallel processing in the auditory cortex of primates. Audiol Neurootol 3: 86–103.PubMedCrossRefGoogle Scholar
  62. Reale RA, Imig TJ (1980) Tonotopic organization in auditory cortex of the cat. J Comp Neurol 192: 265–291.PubMedCrossRefGoogle Scholar
  63. Recanzone GH (1998) Rapidly induced auditory plasticity:The ventriloquism aftereffect. Proc Nat Acad Sci USA 95: 869–875.PubMedCrossRefGoogle Scholar
  64. Recanzone GH, Guard DC, Phan ML, Su T-IK (2000) Correlation between the activity of single auditory cortical neurons and sound-localization behavior in the macaque monkey. J Neurophysiol 83: 2723–2739.PubMedGoogle Scholar
  65. Reinoso-Suarez F, Roda JM (1985) Topographical organization of the cortical afferent connections to the cortex of the anterior ectosylvian sulcus in the cat. Exp Brain Res 59: 313–324.PubMedCrossRefGoogle Scholar
  66. Rice JJ, May BJ, Spirou GA, Young ED (1992) Pinna-based spectral cues for sound localization in cat. Hear Res 58: 132–152.PubMedCrossRefGoogle Scholar
  67. Roda JM, Reinoso-Suarez R (1983) Topographical organization of the thalamic projections to the cortex of the anterior ectosylvian sulcus in the cat. Exp Brain Res 49: 131–139.PubMedCrossRefGoogle Scholar
  68. Romanski LM, Tian B, Fritz J, Mishkin M, Goldman-Rakic PS, Rauschecker JP (1999) Dual streams of auditory afferents target multiple domains in the primate prefrontal cortex. Nat Neurosci 2: 1131–1136.PubMedCrossRefGoogle Scholar
  69. Ruff RM, Hersh NA, Pribram KH (1981) Auditory spatial deficits in the personal and extrapersonal frames of reference due to cortical lesions. Neuropsychologica 19: 435–443.CrossRefGoogle Scholar
  70. Russo GS, Bruce CJ (1994) Frontal eye field activity preceding aurally guided saccades. J Neurophysiol 71: 1250–1253.PubMedGoogle Scholar
  71. Sanchez-Longo LP, Forster FM (1958) Clinical significance of impairment of sound localization. Neurology 8: 119–125.PubMedCrossRefGoogle Scholar
  72. Sovijarvi ARA, Hyvarinen J (1974) Auditory cortical neurons in the cat sensitive to the direction of sound source movement. Brain Res 73: 455–471.PubMedCrossRefGoogle Scholar
  73. Stricanne B, Andersen RA, Mazzoni P (1996) Eye-centered, head-centered, and intermediate coding of remembered sound locations in area LIP. J Neurophysiol 75: 1233–1241.Google Scholar
  74. Thompson GC, Cortez AM (1983) The inability of squirrel monkeys to localize sound after unilateral ablation of auditory cortex. Behav Brain Res 8: 211–216.PubMedCrossRefGoogle Scholar
  75. Thompson GC, Masterton RB (1978) Brainstem auditory pathway involved in reflexive head orientation to sound. J Neurophysiol 41: 1183–1202.PubMedGoogle Scholar
  76. Ungerleider LG, Mishkin M (1982) Two cortical visual systems. In: Ingle DJ, Goodale MA, Mansfields RJW (eds) Analysis of Visual Behaviour. Cambridge: MIT Press, pp. 549–586.Google Scholar
  77. Vaadia E, Benson DA, Hienz RD, Goldstein JR, Moise H (1986) Unit study of monkey frontal cortex: Active localization of auditory and visual stimuli. J Neurophysiol 56: 934–952.PubMedGoogle Scholar
  78. Wortis SB, Pfeiffer AZ (1948) Unilateral auditory-spatial agnosia. J Nery Ment Dis 108: 181–186.CrossRefGoogle Scholar
  79. Xu L, Middlebrooks JC (2000) Individual differences in external-ear transfer functions of cats. J Acoust Soc Am 107: 1451–1459.PubMedCrossRefGoogle Scholar
  80. Xu L, Furukawa S, Middlebrooks JC (1998) Sensitivity to sound-source elevation in non-tonotopic auditory cortex. J Neurophysiol 80: 882–894.PubMedGoogle Scholar
  81. Xu L, Furukawa S, Middlebrooks JC (1999) Auditory cortical responses in the cat to sounds that produce spatial illusions. Nature 399: 688–691.PubMedCrossRefGoogle Scholar
  82. Young ED, Rice.1.1, Tong SC (1996) Effects of pinna position on head-related transfer functions in the cat. J Acoust Soc Am 99: 3064–3076.Google Scholar
  83. Zattore RJ, Penhume VB (2001) Spatial localization after excision of human auditory cortex. J Neurosci 21: 6321–6328.Google Scholar
  84. Zattore RJ, Ptito A, Villemure J-G (1995) Preserved auditory spatial localization following cerebral hemispherectomy. Brain 118: 879–889.CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media New York 2002

Authors and Affiliations

  • John C. Middlebrooks
  • Li Xu
  • Shigeto Furukawa
  • Brian J. Mickey

There are no affiliations available

Personalised recommendations