Neural networks engaged in milliseconds and seconds time processing: Evidence from transcranial magnetic stimulation and patients with cortical or subcortical dysfunction

Philosophical Transactions of the Royal Society B: Biological Sciences

Volumn 364, Issue 1525, 2009, Pages 1907-1918

  Koch, Giacomo ^a,b   Oliveri, Massimiliano ^a,c   Caltagirone, Carlo ^a,b  

^a IRCCS FONDAZIONE SANTA LUCIA   (Italy)

^b UNIVERSITY OF ROME TOR VERGATA   (Italy)

^c UNIVERSITY OF PALERMO   (Italy)

Author keywords

Parkinson's disease; Repetitive transcranial magnetic stimulation; Stroke; Time perception; Timing; Transcranial magnetic stimulation

Indexed keywords

ARTIFICIAL NEURAL NETWORK; COGNITION; ESTIMATION METHOD; NERVOUS SYSTEM DISORDER; STROKE;

EID: 67650254570     PISSN: 09628436     EISSN: 14712970     Source Type: Journal    
DOI: 10.1098/rstb.2009.0018     Document Type: Review

References (109)

1. Artieda, J., Pastor, M. A., Lacruz, F. & Obeso, J. A. 1992 Temporal discrimination is abnormal in Parkinson's disease. Brain 115, 199-210. (doi:10.1093/brain/115.1.199)
2. Basso, G., Nichelli, P., Frassinetti, F. & di Pellegrino, G. 1996 Time perception in a neglected space. Neuroreport 7, 2111-2114. (doi:10.1097/00001756-199609020-00009)
3. Battaglia-Mayer, A., Caminiti, R., Lacquaniti, F. & Zago, M. 2003 Multiple levels of representation of reaching in the parieto-frontal network. Cereb. Cortex 13, 1009-1022. (doi:10.1093/cercor/13.10.1009)
4. Becchio, C. & Bertone, C. 2006 Time and neglect: abnormal temporal dynamics in unilateral spatial neglect. Neuropsychologia 44, 2775-2782. (doi:10.1016/j.neuropsychologia. 2006.06.011)
5. Bueti, D. & Walsh, V. 2009 The pariet al. cortex and the representation of time, space, number and other magnitudes. Phil. Trans. R. Soc. B 364, 1831-1840. (doi:10. 1098/rstb.2009.0028)
6. Bueti, D., Bahrami, B. & Walsh, V. 2008a Sensory and association cortex in time perception. J. Cogn. Neurosci. 20, 1054-1062. (doi:10.1162/jocn. 2008.20060)
7. Bueti, D., van Dongen, E. V. & Walsh, V. 2008b The role of superior temporal cortex in auditory timing. PLoS ONE 3, e2481. (doi:10.1371/journal.pone.0002481)
8. Buhusi, C. V. & Meck, W. H. 2005 What makes us tick? Functional and neural mechanisms of interval timing. Nat. Rev. Neurosci. 6, 755-765. (doi:10.1038/nrnl764)
9. Centonze, D. et al. 2008 Adaptations of glutamatergic synapses in the striatum contribute to recovery from cerebellar damage. Eur. J. Neurosci. 27, 2188-2196. (doi: 10.1111/j.1460-9568.2008.06182.x)
10. Coull, J. T., Vidal, F., Nazarian, B. & Macar, F. 2004 Functional anatomy of the attentional modulation of time estimation. Science 303, 1506-1508. (doi:10.1126/science. 1091573)
11. Danckert, J. & Ferber, S. 2006 Revisiting unilateral neglect. Neuropsychologia 44, 987-1006. (doi:10.1016/j.neuropsychologia. 2005.09.004)
12. Danckert, J., Ferber, S., Pun, C., Broderick, C., Striemer, C., Rock, S. & Stewart, D. 2007 Neglected time: impaired temporal perception of multisecond intervals in unilateral neglect. J. Cogn. Neurosci. 19, 1706-1720. (doi:10.1162/jocn. 2007.19.10.1706)
13. di Pellegrino, G., Basso, G. & Frassinetti, F. 1998 Visual extinction as a spatio-temporal disorder of selective attention. Neuroreport 9, 835-839. (doi:10.1097/00001756-199803300-00013)
14. Droit-Volet, S. & Meck, W. H. 2007 How emotions colour our perception of time. Trends Cogn. Sci. 11, 504-513. (doi:10.1016/j.tics.2007. 09.008)
15. Eagleman, D. M. 2008 Human time perception and its illusions. Curr. Opin. Neurobiol. 18, 131-136. (doi:10. 1016/j.conb.2008.06.002)
16. Elvevåg, B., Brown, G. D., McCormack, T., Vousden, J. I. & Goldberg, T. E. 2004 Identification of tone duration, line length, and letter position: an experimental approach to timing and working memory deficits in schizophrenia. J. Abnorm. Psychol. 113, 509-521. (doi:10.1037/0021-843X.113.4.509)
17. Exner, C., Weniger, G. & Irle, E. 2004 Cerebellar lesions in the PICA but not SCA territory impair cognition. Neurology 63, 2132-2135.
18. Fernandez Del Olmo, M., Cheeran, B., Koch, G. & Rothwell, J. C. 2007 Role of the cerebellum in externally paced rhythmic finger movements. J. Neurophysiol. 98, 145-152. (doi: 10.1152/jn. 01088.2006)
19. Ferrandez, A. M., Hugueville, L., Lehéricy, S., Poline, J. B., Marsault, C. & Pouthas, V. 2003 Basal ganglia and supplementary motor area subtend duration perception: an fMRI study. Neuroimage 19, 1532-1544. (doi:10.1016/S1053-8119 (03) 00159-9)
20. Foltynie, T., Brayne, C. & Barker, R. A. 2002 The heterogeneity of idiopathic Parkinson's disease. J. Neurol. 249, 138-145. (doi:10.1007/PL00007856)
21. Genovesio, A., Tsujimoto, S. & Wise, S. P. 2006 Neuronal activity related to elapsed time in prefrontal cortex. J. Neurophysiol. 95, 3281-3285. (doi:10.1152/jn. 01011. 2005)
22. Gibbon, J., Malapani, C., Dale, C. L. & Gallistel, C. 1997 Toward a neurobiology of temporal cognition: advances and challenges. Curr. Opin. Neurobiol. 7, 170-184. (doi:10.1016/S0959-4388 (97) 80005-0)
23. Goldman-Rakic, P. S. 1995 Cellular basis of working memory. Neuron 14, 477-485. (doi:10.1016/0896-6273 (95) 90304-6)
24. Guehl, D., Burbaud, P., Lorenzi, C., Ramos, C., Bioulac, B., Semal, C. &Demany, L. 2008 Auditory temporal processing in Parkinson's disease. Neuropsychologia 46, 2326-2335. (doi:10.1016/j.neuropsychologia. 2008.03.007)
25. Harrington, D. L., Haaland, K. Y. & Hermanowicz, N. 1998a Temporal processing in the basal ganglia. Neuropsychology 12, 3-12. (doi:10.1037/0894-4105.12.1.3)
26. Harrington, D. L., Haaland, K. Y. & Knight, R. T. 1998b Cortical networks underlying mechanisms of time perception. J. Neurosci. 18, 1085-1095.
27. Harrington, D. L., Lee, R. R., Boyd, L. A., Rapcsak, S. Z. & Knight, R. T. 2004 Does the representation of time depend on the cerebellum? Effect of cerebellar stroke. Brain 127, 561-574. (doi:10.1093/brain/awh065)
28. Hellström, A. & Rammsayer, T. H. 2004 Effects of timeorder, interstimulus interval, and feedback in duration discrimination of noise bursts in the 50- and 1000-ms ranges. Acta Psychol. (Amst.) 116, 1-20. (doi:10.1016/j.actpsy.2003.11.003)
29. Hinton, S. C. & Meck, W. H. 2004 Frontal-striatal circuitry activated by human peak-interval timing in the supraseconds range. Brain Res. Cogn. Brain Res. 21, 171-182. (doi:10.1016/j.cogbrainres.2004.08.005)
30. Holmes, G. 1939 The cerebellum of man. Brain 62, 1-30. (doi:10.1093/brain/62.1.1)
31. Hoshi, E., Tremblay, L., Féger, J., Carras, P. L. & Strick, P. L. 2005 The cerebellum communicates with the basal ganglia. Nat. Neurosci. 8, 1491-1493. (doi:10.1038/nnl544)
32. Ivry, R. B. & Schlerf, J. E. 2008 Dedicated and intrinsic models of time perception. Trends Cogn. Sci. 12, 273-280. (doi:10.1016/j.tics.2008.04.002)
33. Ivry, R. B. & Spencer, R. M. 2004 The neural representation of time. Curr. Opin. Neurobiol. 14, 225-232. (doi:10.1016/j.conb.2004.03. 013)
34. Ivry, R. B., Keele, S. W. & Diener, H. C. 1988 Dissociation of the lateral and medial cerebellum in movement timing and movement execution. Exp. Brain Res. 73, 167-180. (doi:10.1007/BF00279670)
35. Ivry, R. B., Spencer, R. M., Zelaznik, H. N. & Diedrichsen, J. 2002 The cerebellum and event timing. Ann. NY Acad. Sci. 978, 302-317. (doi:10.1111/j.1749-6632.2002.tb0 7576.x)
36. Jahanshahi, M., Jones, C. R., Dirnberger, G. & Frith, C. D. 2006 The substantia nigra pars compacta and temporal processing. J. Neurosci. 26, 12266-12273. (doi:10.1523/JNEUROSCI.2540-06.2006)
37. Jones, C. R., Rosenkranz, K., Rothwell, J. C. & Jahanshahi, M. 2004 The right dorsolateral prefrontal cortex is essential in time reproduction: an investigation with repetitive transcranial magnetic stimulation. Exp. Brain Res. 158, 366-372. (doi:10.1007/s00221-004-1912-3)
38. Jones, C. R., Malone, T. J., Dirnberger, G., Edwards, M. & Jahanshahi, M. 2008 Basal ganglia, dopamine and temporal processing: performance on three timing tasks on and off medication in Parkinson's disease. Brain Cogn. 68, 30-41. (doi:10.1016/j.bandc.2008.02.121)
39. Jueptner, M., Rijntjes, M., Weiller, C., Faiss, J. H., Timmann, D., Mueller, S. P. & Diener, H. C. 1995 Localization of a cerebellar timing process using PET. Neurology 45, 1540-1545.
40. Kelly, R. M. & Strick, P. L. 2003 Cerebellar loops with motor cortex and prefrontal cortex of a nonhuman primate. J. Neurosci. 23, 8432-8444.
41. Koch, G., Oliveri, M., Carlesimo, G. A. & Caltagirone, C. 2002 Selective deficit of time perception in a patient with right prefrontal cortex lesion. Neurology 59, 1658-1659.
42. Koch, G., Oliveri, M., Torriero, S. & Caltagirone, C. 2003 Underestimation of time perception after repetitive transcranial magnetic stimulation. Neurology 60, 1844-1846.
43. Koch, G., Brusa, L., Caltagirone, C., Oliveri, M., Peppe, A., Tiraboschi, P. & Stanzione, P. 2004a Subthalamic deep brain stimulation improves time perception in Parkinson's disease. Neuroreport 15, 1071-1073. (doi:10.1097/00001756-200404290-00028)
44. Koch, G., Oliveri, M., Brusa, L., Stanzione, P., Torriero, S. & Caltagirone, C. 2004b High-frequency rTMS improves time perception in Parkinson disease. Neurology 63, 2405-2406.
45. Koch, G., Brusa, L., Oliveri, M., Stanzione, P. & Caltagirone, C. 2005a Memory for time intervals is impaired in left hemi-Parkinson patients. Neuropsychohgia 43, 1163-1167. (doi:10.1016/j. neuropsychologia.2004.11.017)
46. Koch, G., Oliveri, M., Torriero, S., Carlesimo, G. A., Turriziani, P. & Caltagirone, C. 2005b rTMS evidence of different delay and decision processes in a frontopariet al. neuronal network activated during spatial working memory. Neuroimage 24, 34-39. (doi:10.1016/j.neuroimage. 2004.09.042)
47. Koch, G., Oliveri, M., Torriero, S., Salerno, S., Lo Gerfo, E. & Caltagirone, C. 2007a Repetitive TMS of cerebellum interferes with millisecond time processing. Exp. Brain Res. 179, 291-299. (doi:10.1007/s00221-006-0791-1)
48. Koch, G., Fernandez Del Olmo, M., Cheeran, B., Ruge, D., Schippling, S., Caltagirone, C. & Rothwell, J. C. 20076 Focal stimulation of the posterior pariet al. cortex increases the excitability of the ipsilateral motor cortex. J. Neurosci. 27, 6815-6822. (doi:10.1523/JNEUROSCI. 0598-07.2007)
49. Koch, G. et al. 2008 Impaired reproduction of second but not millisecond time intervals in Parkinson's disease. Neuropsychohgia 46, 1305-1313. (doi:10.1016/j.neuro psychologia.2007.12.005)
50. Koekkoek, S. K., Hulscher, H. C., Dortland, B. R., Hensbroek, R. A., Elgersma, Y., Ruigrok, T. J. & De Zeeuw, C. I. 2003 Cerebellar LTD and learningdependent timing of conditioned eyelid responses. Science 301, 1736-1739. (doi:10.1126/science.l088383)
51. Kotani, S., Kawahara, S. & Kirino, Y. 2003 Purkinje cell activity during learning a new timing in classical eyeblink conditioning. Brain Res. 994, 193-202. (doi:10.1016/j.brainres.2003.09.036)
52. Lee, K. H., Egleston, P. N., Brown, W. H., Gregory, A. N., Barker, A. T. & Woodruff, P. W. 2007 The role of the cerebellum in subsecond time perception: evidence from repetitive transcranial magnetic stimulation. J. Cogn. Neurosci. 19, 147-157. (doi:10.1162/jocn. 2007.19.1.147)
53. Lewis, P. A. & Miall, R. C. 2003a Distinct systems for automatic and cognitively controlled time measurement: evidence from neuroimaging. Curr. Opin. Neurobiol. 13, 250-255. (doi:10.1016/S0959-4388 (03) 00036-9)
54. Lewis, P. A. & Miall, R. C. 2003b Brain activation patterns during measurement of sub- and supra-second intervals. Neuropsychologia 41, 1583-1592. (doi:10.1016/S0028-3932 (03) 00118-0)
55. Lewis, P. A. & Miall, R. C. 2006 Remembering the time: a continuous clock. Trends Cogn. Sci. 10, 401-406. (doi:10.1016/j.tics.2006.07. 006)
56. Lewis, S. G., Foltynie, T., Blackwell, A. D., Robbins, T. W., Owen, A. M. & Barker, R. A. 2005 Heterogeneity of Parkinson's disease in the early clinical stages using a data driven approach. J. Neurol. Neurosurg. Psychiatry 76, 343-348. (doi:10.1136/jnnp. 2003.033530)
57. Macar, R., Coull, J. & Vidal, F. 2006 The supplementary motor area in motor and perceptual time processing: fMRI studies. Cogn. Process. 7, 89-94. (doi: 10.1007/s 10339-005-0025-7)
58. Malapani, C., Dubois, B., Rancurel, G. & Gibbon, J. 1998a Cerebellar dysfunctions of temporal processing in the seconds range in humans. Neuroreport 9, 3907-3912. (doi:10.1097/00001756-199812010-00026)
59. Malapani, C., Rakitin, B., Levy, R., Meck, W. H., Deweer, B., Dubois, B. & Gibbon, J. 1998b Coupled temporal memories in Parkinson's disease: a dopamine-related dysfunction. J. Cogn. Neurosci. 10, 316-331. (doi:10.1162/089892998562762)
60. Malapani, C., Deweer, B. & Gibbon, J. 2002 Separating storage from retrieval dysfunction of temporal memory in Parkinson's disease. J. Cogn. Neurosci. 14, 311-322. (doi:10.1162/089892902317236920)
61. Mangels, J. A., Ivry, R. B. & Shimizu, N. 1998 Dissociable contributions of the prefrontal and neocerebellar cortex to time perception. Brain Res. Cogn. Brain Res. 7, 15-39. (doi:10.1016/S0926-6410 (98) 00005-6)
62. Maricq, A. V. & Church, R. M. 1983 The differential effects of haloperidol and methamphetamine on time estimation in the rat. Psychopharmacology 79, 10-15. (doi:10.1007/BF00433008)
63. Matell, M. S. & Meck, W. H. 2004 Cortico-striatal circuits and interval timing: coincidence detection of oscillatory processes. Brain Res. Cogn. Brain Res. 21, 139-170. (doi:10.1016/j.cogbrainres.2004.06.012)
64. Matell, M. S., Meck, W. H. & Nicolelis, M. A. 2003 Interval timing and the encoding of signal duration by ensembles of cortical and striatal neurons. Behav. Neurosci. 117, 760-773. (doi:10.1037/0735-7044. 117.4.760)
65. Matell, M. S., King, G. R. & Meck, W. H. 2004 Differential modulation of clock speed by the administration of intermittent versus continuous cocaine. Behav. Neurosci. 118, 150-156. (doi:10.1037/0735-7044.118.1.150)
66. Mattay, V. S., Tessitore, A., Callicott, J. H., Bertolino, A., Goldberg, T. E., Chase, T. N., Hyde, T. M. & Weinberger, D. R. 2002 Dopaminergic modulation of cortical function in patients with Parkinson's disease. Ann. Neurol. 51, 156-164. (doi:10.1002/ana.l0078)
67. Mauk, M. D. & Buonomano, D. V. 2004 The neural basis of temporal processing. Annu. Rev. Neurosci. 27, 307-340. (doi:10.1146/ annurev. neuro.27.070203.144247)
68. Meck, W. H. 1996 Neuropharmacology of timing and time perception. Brain Res. Cogn. Brain Res. 3, 227-242. (doi:10.1016/0926-6410 (96) 00009-2)
69. Meck, W. H. & Benson, A. M. 2002 Dissecting the brain's internal clock: how frontal-striatal circuitry keeps time and shifts attention. Brain Cogn. 48, 195-211. (doi:10. 1006/brcg.2001.1313)
70. Merchant, H., Luciana, M., Hooper, C., Majestic, S. & Tuite, P. 2008 Interval timing and Parkinson's disease: heterogeneity in temporal performance. Exp. Brain Res. 184, 233-248. (doi:10.1007/s00221-007-1097-7)
71. Michon, J. A. 1985 The compleat time experiencer. In Time, mind, and behavior (eds J. A. Michon & J. L. Jackson), pp. 21-52. Berlin, Germany: Springer.
72. Mimura, M., Kinsbourne, M. & O'Connor, M. 2000 Time estimation by patients with frontal lesions and by Korsakoff amnesics. J. Int. Neuropsychol. Soc. 6, 517-528. (doi:10.1017/S1355617700655017)
73. Mottaghy, F. M., Gangitano, M., Sparing, R., Krause, B. J. & Pascual-Leone, A. 2002 Segregation of areas related to visual working memory in the prefrontal cortex revealed by rTMS. Cereb. Cortex 12, 369-375. (doi:10.1093/cercor/12.4.369)
74. Mottaghy, F. M., Gangitano, M., Krause, B. J. & Pascual- Leone, A. 2003 Chronometry of pariet al. and prefrontal activations in verbal working memory revealed by transcranial magnetic stimulation. Neuroimage 18, 565-575. (doi:10.1016/S1053-8119 (03) 00010-7)
75. Nenadic, I., Gaser, C., Volz, H. P., Rammsayer, T., Hager, F. & Sauer, H. 2003 Processing of temporal information and the basal ganglia: new evidence from fMRI. Exp. Brain Res. 148, 238-246.
76. Nichelli, P., Always, D. & Grafman, J. 1996 Perceptual timing in cerebellar degeneration. Neuropsychologia 34, 863-871. (doi:10.1016/0028-3932 (96) 00001-2)
77. Nobre, A. C. & O'Reilly, J. 2004 Time is of the essence. Trends Cogn. Sci. 8, 387-389. (doi:10.1016/j.tics.2004. 07.005)
78. O'Boyle, D. J., Freeman, J. S. & Cody, F. W. 1996 The accuracy and precision of timing of self-paced, repetitive movements in subjects with Parkinson's disease. Brain 119, 51-70. (doi:10.1093/brain/119.1. 51)
79. Oliveri, M., Rossini, P. M., Filippi, M. M., Traversa, R., Cicinelli, P., Palmieri, M. G., Pasqualetti, P. & Caltagirone, C. 2000 Time-dependent activation of parieto-frontal networks for directing attention to tactile space. A study with paired transcranial magnetic stimulation pulses in right-brain-damaged patients with extinction. Brain 123, 1939-1947. (doi:10.1093/brain/123.9.1939)
80. Oliveri, M., Turriziani, P., Carlesimo, G. A., Koch, G., Tomaiuolo, F., Panella, M. & Caltagirone, C. 2001 Parieto-frontal interactions in visual-object and visualspatial working memory: evidence from transcranial magnetic stimulation. Cereb. Cortex 11, 606-618. (doi:10.1093/cercor/11.7.606)
81. Oliveri, M., Koch, G., Torriero, S. & Caltagirone, C. 2005 Increased facilitation of the primary motor cortex following 1 Hz repetitive transcranial magnetic stimulation of the contralateral cerebellum in normal humans. Neurosci. Lett. 376, 188-193. (doi:10.1016/j.neulet.2004. 11.053)
82. Oliveri, M., Torriero, S., Koch, G., Salerno, S., Petrosini, L. & Caltagirone, C. 2007 The role of transcranial magnetic stimulation in the study of cerebellar cognitive function. Cerebellum 6, 95-101. (doi:10.1080/14734220701213421)
83. O'Reilly, J. X., Mesulam, M. M. & Nobre, A. C. 2008 The cerebellum predicts the timing of perceptual events. J. Neurosci. 28, 2252-2260. (doi:10.1523/JNEUROSCI. 2742-07.2008)
84. Passingham, D. & Sakai, K. 2004 The prefrontal cortex and working memory: physiology and brain imaging. Curr. Opin. Neurobiol. 14, 163-168. (doi:10.1016/j.conb.2004. 03.003)
85. Pastor, M. A., Artieda, J., Jahanshahi, M. & Obeso, J. A. 1992 Time estimation and reproduction is abnormal in Parkinson's disease. Brain 115, 211-225. (doi:10.1093/brain/115.1.211)
86. Paulsen, J. S., Zimbelman, J. L., Hinton, S. C., Langbehn, D. R., Leveroni, C. L., Benjamin, M. L., Reynolds, N. C. & Rao, S. M. 2004 fMRI biomarker of early neuronal dysfunction in presymptomatic Huntington's disease. AJNR Am. J. Neuroradiol. 25, 1715-1721.
87. Perbal, S., Deweer, B., Pillon, B., Vidailhet, M., Dubois, B. & Pouthas, V. 2005 Effects of internal clock and memory disorders on duration reproductions and duration productions in patients with Parkinson's disease. Brain Cogn. 58, 35-48. (doi:10.1016/j.bandc.2005.02.003)
88. Pouthas, V., Garnero, L., Ferrandez, A. M. & Renault, B. 2000 ERPs and PET analysis of time perception: spatial and temporal brain mapping during visual discrimination tasks. Hum. Brain Mapp. 10, 49-60. (doi:10.1002/(SICI) 1097-0193 (200006) 10:2<49::AID-HBM10>3.0. CO;2-8)
89. Rakitin, B. C., Scarmeas, N., Li, T., Malapani, C. & Stern, Y. 2006 Single-dose levodopa administration and aging independently disrupt time production. J. Cogn. Neurosci. 18, 376-387. (doi:10.1162/jocn. 2006.18.3.376)
90. Rammsayer, T. & Classen, W. 1997 Impaired temporal discrimination in Parkinson's disease: temporal processing of brief durations as an indicator of degeneration of dopaminergic neurons in the basal ganglia. Int. J. Neurosci. 91, 45-55.
91. Rammsayer, T. H. 1993 On dopaminergic modulation of temporal information processing. Biol. Psychol. 36, 209-222. (doi:10.1016/0301-0511 (93) 90018-4)
92. Rammsayer, T. H. 1997 Are there dissociable roles of the mesostriatal and mesolimbocortical dopamine systems on temporal information processing in humans? Neuropsychobiology 35, 36-45. (doi:10.1159/000119328)
93. Rammsayer, T. H. & Lima, S. D. 1991 Duration discrimination of filled and empty auditory intervals: cognitive and perceptual factors. Percept. Psychophys. 50, 565-574.
94. Rao, S. M., Harrington, D. L., Haaland, K. Y., Bobholz, J. A., Cox, R. W. & Binder, J. R. 1997 Distributed neural systems underlying the timing of movements. J. Neurosci. 17, 5528-5535.
95. Rao, S. M., Mayer, A. R. & Harrington, D. L. 2001 The evolution of brain activation during temporal processing. Nat. Neurosci. 4, 317-323. (doi:10.1038/85191)
96. Schmahmann, J. D. & Sherman, J. C. 1998 The cerebellar cognitive affective syndrome. Brain 121, 561-579. (doi:10. 1093/brain/121.4.561)
97. Schrag, A., Quinn, N. P. & Ben-Shlomo, Y 2006 Heterogeneity of Parkinson's disease. J. Neurol. Neurosurg. Psychiatry 77, 275-276.
98. Schubotz, R. I., Friederici, A. D. & von Cramon, D. Y 2000 Time perception and motor timing: a common cortical and subcortical basis revealed by fMRI. Neuroimage 11, 1-12. (doi:10.1006/nimg.1999.0514)
99. Selemon, L. D. & Goldman-Rakic, P. S. 1988 Common cortical and subcortical targets of the dorsolateral prefrontal and posterior pariet al. cortices in the rhesus monkey: evidence for a distributed neural network subserving spatially guided behavior. J. Neurosci. 8, 4049-4068.
100. Shea-Brown, E., Rinzel, J., Rakitin, B. C. & Malapani, C. 2006 A firing rate model of Parkinsonian deficits in interval timing. Brain Res. 1070, 189-201. (doi:10.1016/j.brainres.2005.10.070)
101. Smith, A., Taylor, E., Lidzba, K. & Rubia, K. 2003 A right hemispheric frontocerebellar network for time discrimination of several hundreds of milliseconds. Neuroimage 20, 344-350. (doi:10.1016/S1053-8119 (03) 00337-9)
102. Snyder, J. J. & Chatterjee, A. 2004 Spatial-temporal anisometries following right pariet al. damage. Neuropsychologia 42, 1703-1708. (doi:10.1016/j.neuropsychologia.2004. 04.003)
103. Spencer, R. M. & Ivry, R. B. 2005 Comparison of patients with Parkinson's disease or cerebellar lesions in the production of periodic movements involving event-based or emergent timing. Brain Cogn. 58, 84-93. (doi:10.1016/j.bandc.2004.09.010)
104. Spencer, R. M., Zelaznik, H. N., Diedrichsen, J. & Ivry, R. B. 2003 Disrupted timing of discontinuous but not continuous movements by cerebellar lesions. Science 300, 1437-1439. (doi:10.1126/science.l083661)
105. Vicario, C. M., Pecoraro, P., Turriziani, P., Koch, G., Caltagirone, C. & Oliveri, M. 2008 Relativistic compression and expansion of experiential time in the left and right space. PLoS ONE 3, el716. (doi: 10.1371/journal. pone.0001716)
106. Walsh, V. 2003 A theory of magnitude: common cortical metrics of time, space and quantity. Trends Cogn. Sci. 7, 483-488. (doi:10.1016/j.tics.2003.09.002)
107. Witt, K. et al. 2008 Neuropsychological and psychiatric changes after deep brain stimulation for Parkinson's disease: a randomised, multicentre study. Lancet Neurol. 7, 605-614. (doi:10.1016/S1474-4422 (08) 70114-5)
108. Wittmann, M. & Paulus, M. P. 2008 Decision making, impulsivity and time perception. Trends Cogn. Sci. 12, 7-12. (doi:10.1016/j.tics. 2007.10.004)
109. Wolpert, D. M. & Miall, R. C. 1996 Forward models for physiological motor control. Neural Netw. 9, 1265-1279. (doi:10.1016/S0893-6080 (96) 00035-4)