1. The Problems With Time Neuroenigmas Lecture January 19, 2007

2. Neural processing is not instantaneous

4. 40 ms Neural processing is not instantaneous

5. 40 ms 100 ms Neural processing is not instantaneous

6. When is this a problem?

8. 100 mph fast ball moves 15 feet in 100 ms

9. How might we compensate for neural delays?

10. Sensory compensation Motor compensation

11. http://www.interchange.ubc.ca/vsearch/research/objectupdating/obj_index.html

12. Theories to explain flash-lag effect Motion extrapolation Motion integration and post diction Latency differences for moving and stationary items Attentional capture by flashed item Citation for review

14. Flag errors in soccer games: the flash-lag effect brought to real life Perception, 2002, volume 31, pages 1205 ^ 1210 Marcus Vinicius C Baldo, Ronald D Ranvaud, Edgard Morya DOI:10.1068/p3422

19. Psychophysics +

20. -1.0 -0.5 0 0.5 1.0 Psychophysics + 0 0.2 0.4 0.6 0.8 1 Isolated Flash Subject JD Position (deg) Proportion reported right -1.0 -0.5 0 0.5 1.0

21. Psychophysics + 0 0.2 0.4 0.6 0.8 1 Isolated Flash Subject JD Position (deg) Proportion reported right

22. Psychophysics +

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34. +

35. -1.0 -0.5 0 0.5 1.0 Psychophysics + 0 0.2 0.4 0.6 0.8 1 Flash in motion Subject JD Position (deg) Proportion reported right -1.0 -0.5 0 0.5 1.0

36. Psychophysics + 0 0.2 0.4 0.6 0.8 1 Flash in motion Subject JD Position (deg) Proportion reported right -1.0 -0.5 0 0.5 1.0.39 o

37. Physiology

38. Tootell, R. B., M. S. Silverman, et al. Science (1982) Retinotopy in cat

40. Prediction: Under conditions that give rise to the perceived shift, receptive fields should shift in a direction opposite to motion direction.

41. RF Physiology

42. RF Physiology

43. RF Physiology flash positions

44. RF Physiology flash positions

45. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

46. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

47. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

48. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

49. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

50. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

51. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

52. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

53. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

54. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

55. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

56. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

57. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

58. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

59. Physiology 0100 400 0 10 20 30 40 50 Time (ms) firing rate

60. Physiology 1.2 2.02.83.64.45.2 6.0.4 0 5 10 15 20 25 30 35 40 45 0100 400 0 10 20 30 40 50 Time (ms) firing rate Position (deg) right left baseline

61. Physiology 0 5 10 15 20 25 30 35 40 45 0100200500 0 10 20 30 40 50 Time (ms) firing rate 1.2 2.02.83.64.45.2 6.0.4 firing rate Position (deg) right left baseline

62. Physiology 0 5 10 15 20 25 30 35 40 45 0100400 0 10 20 30 40 50 Time (ms) firing rate 1.2 2.02.83.64.45.2 6.0.4 firing rate Position (deg) right left baseline

63. Physiology 0 50 0 5 10 15 20 25 30 35 40 45 1.2 2.02.83.64.45.2 6.0.4 firing rate Position (deg) right left baseline

64. Physiology 0 50 0150 0 50 0 5 10 15 20 25 30 35 40 45 1.2 2.02.83.64.45.2 6.0.4 firing rate Position (deg) right left baseline

65. Physiology 0 50 0150 0 50 0 5 10 15 20 25 30 35 40 45 1.2 2.02.83.64.45.2 6.0.4 1 o firing rate Position (deg) right left baseline

66. flash positions RF Physiology

67. 0 5 10 15 20 25 30 35 40 45 1.2 2.02.83.64.45.2 6.0.4 firing rate Position (deg)

68. Physiology 0 5 10 15 20 25 30 35 40 45 1.2 2.02.83.64.45.2 6.0.4 firing rate Position (deg)

69. Physiology 0 5 10 15 20 25 30 35 40 45 * * ** * * * 1.2 2.02.8.4 Inward Outward firing rate Position (deg)

70. Physiology 10121416 0 5 10 15 20 25 30 35 40 45 * * * * * ** * * * * 1.2 2.02.8.4 Inward Outward firing rate Position (deg)

71. Physiology 10121416 0 5 10 15 20 25 30 35 40 45 * * * * * ** * * * * Shift Index = 1.2 o 1.2 2.02.8.4 Inward Outward firing rate Position (deg)

72. Example cell Physiology 0.4.81.21.62.02.42.8 0 50 100 150 200 firing rate position (deg) 0.4.81.21.62.02.42.8 0 10 20 30 40 50 firing rate position (deg) 0.4.81.21.62.02.42.8 0 10 20 30 40 50 60 firing rate position (deg) 0.4.81.21.62.02.42.8 0 10 20 30 40 50 60 70 80 firing rate position (deg) * ** * * * * * * * * * ** * * * * * * * * * * * * * S.I. = 1.2 o S.I. = 0.4 o S.I. = 0.8 o

73. Physiology -2.002.0 0 5 10 15 20 Mean S.I. =.31 o p <.0001 n = 59 number of cells Position (deg)

74. -2.002.0 0 5 10 15 20 Physiology Full Motion sequence Mean S.I. =.31 o p <.0001 n = 59 number of cells Position (deg)

75. Physiology Full Motion sequence Shift across population =.31 o p <.0001

76. Physiology Full Motion sequence Shift across population =.31 o p <.0001 Flash Terminal sequence

77. Physiology Full Motion sequence Shift across population =.31 o p <.0001 Flash Terminal sequence Shift across population =.34 o p <.0001

78. Physiology Full Motion sequence Shift across population =.31 o p <.0001 Flash Terminal sequence Shift across population =.34 o p <.0001 Single Color sequence

79. Physiology Full Motion sequence Shift across population =.31 o p <.0001 Flash Terminal sequence Shift across population =.34 o p <.0001 Single Color sequence Shift across population =.12 o p =.14

80. Physiology Non-Color Selective Cells Similar results to full population

81. Summary

82. Spatial representation of the the flashed element is distorted

83. Summary Spatial representation of the the flashed element is distorted Apparent in the first responses

84. Summary Spatial representation of the the flashed element is distorted Apparent in the first responses Similar in magnitude to mislocalization in humans

85. Summary Spatial representation of the the flashed element is distorted Apparent in the first responses Similar in magnitude to mislocalization in humans Motion alone is not sufficient to induce shift

86. Summary Spatial representation of the the flashed element is distorted Apparent in the first responses Similar in magnitude to mislocalization in humans Motion alone is not sufficient to induce shift Dissociation between perception and V4 physiology in the flashed terminal condition

87. Summary Spatial representation of the the flashed element is distorted Apparent in the first responses Similar in magnitude to mislocalization in humans Motion alone is not sufficient to induce shift Dissociation between perception and V4 physiology in the flashed terminal condition V4 distortion does not require motion following the flash

88. Probability Position Full motion condition Flash terminal condition

89. Bhavin R. Sheth, Romi Nijhawan, & Shinsuke Shimojo. (2000). Changing objects lead briefly flashed ones. Nature Neuroscience 3, 489-495.

90. David Alais & David Burr. (2003). The flash-lag effect occurs in audition and crossmodally. Current Biology 13, 59-63.

91. R.H. Cai, K. Jacobson, R. Baloh, M. Schlag-Rey & J. Schlag. (2000) Vestibular signals can distort the perceived spatial relationship of retinal stimuli. Experimental Brain Research 135, 275-278.

92. Alan Johnston & Shinya Nishida (2001). “Time perception: brain time or event time. Current Biology 11, R427-R430. K. Moutoussis and S. Zeki, Functional segregation and temporal hierarchy of the visual perceptive systems. Proc R Soc Lond [Biol] Biol Sci 264 (1997), pp. 1407– 1414. Color Motion Asynchrony Illusion