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Department of Psychological and Brain Sciences
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Peter Ulric Tse

Associate Professor
Harvard University, 1992-1998, PhD Cognitive Psychology, with Patrick Cavanagh and Ken Nakayama

Max Planck Institute for Biological Cybernetics, 1999-2001, post-doc, fMRI, with Nikos Logothetis
Dartmouth College 1980-84 BA Physics/Math

Office Phone: (603) 646-4014, Moore 355
Fax: (603) 646-141; Lab phone:(603) 64664764



We use fMRI, DTI, and psychophysics in my lab to investigate the cognitive and neural bases of visual perception, attention and consciousness. Our research focuses on the visual perception of 3D form and motion, and how these two types of information interact with each other and with attention. Because the 2D visual image is inherently ambiguous, the visual system must construct 3D percepts on the basis of assumptions about the image-to-world mapping. One of our team's goals is to understand the assumptions that underlie the construction of visual percepts, and to understand the neuronal circuits that could realize such constructive processes. In addition to vision, attention and consciousness my lab has become increasingly interested in studying the neural bases of human creativity, free will and symbolic processing as well.
   see 3D scan

Interested in earning a PhD?

Are you interested in the biggest of the big questions and willing to work very hard? I hope to take one or two new graduate students into the lab in September 2012. I am looking for hard-working, smart, humorful, kind, creative individuals who are passionate about understanding human vision and uncovering neural correlates of visual consciousness, or understanding the neural bases of human creativity, symbolic processing or free will. Technical skills, like programming, and research experience are a big plus though not required. You can apply by going to The application deadline is December 15th 2011.

Some interesting demos of the visual effects we study                                            

1. Transformational apparent motion  example1  example 2  example 3  example 4   see refs. A, 12, 27

2. 3D transformational apparent motion  ref. 12 2nd-order transformational apparent motion  ref. 27

3. Wakes You can right click save this executable and play it on your computer: Wakes with 3 balls refs. 3 and 7

4. What is the 3D shape of this? And of this?

5. The subjective expansion of time    Clock example     refs. 18 and 22

6. How do we recover shape from contours?  1  2  3    ref. 11

7. Examples of volume and amodal completion:  1  2  3  4  5  6  7   refs. 1, 2, 4-6

8. Amodal completion is incomplete    refs. 1, 2, 4-6

9. Motion-induced color change  demo 1  demo2  Equiluminant version 1  Equiluminant version 2

10. Attention-induced brightness changes 1 2 3 4   ref. 20

11. Illusory Rebound Motion example2   refs. 21, 25, 29

12. Gradient onset and offset induced motion   example2   ref. 24

13. Feature mixing rather than filling-in. Fixate very carefully. The color of the top field will merge into a uniform color approximately like that of the bottom field. Example1. Example2. refs. 26, 42, and 47 read more.

14. The thinner ellipse seems to rotate faster, though the two ellipses in fact have the same angular velocity (Quicktime movie) ref. 28, 32, 37 read more.

15. The Bar-Cross-Ellipse Illusion. Is it a deforming cross, bars alternating in depth, a rigidly rotating ellipse behind four white square occluders, or a rigid cross viewed through a rotating elliptical window, complete with illusory contours? ref. 31.

16. The infinite regress illusion. Also on youtube. The object appears to move away from the fixation point, even though it is just moving up and down. ref. 33. An interesting variant - save to disk, then play in loop mode in quicktime. Art Shapiro demonstrates that the infinite regress illusion may account for the break of the curveball

17. Examples of the dancing bar illusion. If they do not play when you click them, right click to save as, then play the download in loop mode Dancing Bars Dancing Squares Dancing Gabors Expanding V Rubber Tree ref. 38.

18. Examples of motion fading. The cross takes longer to motion fade, though made of the same dots The cross appears to rotate faster than the square rotating at the same angular velocity The solid line cross takes longer to fade than the square ref. 35. The second demo above is closely related to the findings of papers 37 and 28.

19. Examples of the drifting edge illusion. Look at a point not directly on the drifting grating. Notice how the barberpole appears to move when it in fact is completely stationary. Only the grating is moving, yet the edge appears to move up and down. If the movies do not play when you click them, right click to save as, then play the download in loop mode. If the gratings flicker, try making the size of the window the movie is playing in smaller. Drifting edge illusion avi file Drifting edge illusion version 2 avi file Drifting edge illusion mov file DEI vertical DEI horizontal No DEI Single edge DEI horizontal barberpole with gap without gap ref. 41.

20. First consider Stuart Anstis' original dots version. Motion appears to slow down under a global perceptual organization. In the next example, note that Ls on left seem to move more than Ls on right. When the global pattern is seen, there appears to be less rotational motion. This slowdown occurs even in the absence of rotational motion. ref. 43 below. See non-rotational case here.

21. Smooth pursuit motion suppression. See how motion slows down during smooth pursuit. ref. 52 below.

22. Attention-biased after-image rivalry. What you attend to influences what afterimage you experience. ref. 58 below.

23. Voluntary attention modulates motion-induced mislocalization. View movies in loop mode. Fixate the fixation spot in the center. The colored disks are in fact vertically aligned, but if you attend carefully to the white layer, the colored spot pair will appear slanted one way, and if you attend to the black layer, the colored spot pair will appear slanted the other way. Single version avi file. Single version mov file Double version avi file Double version mov file This version rules out cyclotorsion as the cause, avi file This version rules out cyclotorsion as the cause, mov file Translation version, avi file Translation version, mov file ref. 54 below.


Selected Publications

Journal articles

60. Tse, P. U., Sarnoff, R., Miller, J., Busby, S., Porter, K. B. and Wheatley, T. P. (2011). The temporal dynamics of an expectation of biological motion. Submitted.

59. Frank, S. M., Reavis, E. A., Tse, P. U. and Greenlee, M. W. (2011). From inefficient visual search to pop-out in one week: Neural mechanisms of feature conjunction learning. Submitted.

58. Tse, P. U., Reavis, E. A., Kohler, P. J, Caplovitz, G. P. and Wheatley, T. P. (2011). Attention alters perceived features by defining the domain of preconscious operations. Submitted. Nature commentary. See demo #22 above.

57. Tse, P. U. (2011). A criterial neuronal code underlies downward mental causation and free will. Submitted.

56. Kohler, P. J, Fogelson, S. V., Reavis, E. A., Meng, M., Guntupalli, S., Hanke, M., Halchenko, Y. O., Connolly, A. C. Haxby, J. V. and Tse, P. U. (2011). Pattern classification precedes regional-average hemodynamic response in early visual cortex. Submitted.

55. Schlegel, A. S., Rudelson, J. R., and Tse, P. U. (2011). White matter structure changes as adults learn a second language. Submitted.

54. Porter, K. B., Caplovitz. G. P., Kohler, P. J., Ackerman, C. M. and Tse, P. U. (2011). Rotational and translational motion interact independently with form. Attention, Perception and Psychophysics. In press.

53. Tse, P. U., Whitney, D., Anstis, S. and Cavanagh, P. (2011). Voluntary attention modulates motion-induced mislocalization. J Vis. 11(3):12 See demo #23 above.

52. Tse, P. U., Kohler, P. J, and Sheinberg, D. L. (2010). The attenuation of visual motion during smooth pursuit. Submitted. See demo #21 above.

51. Caplovitz, G. P. and Tse, P. U. (2010).   Extrastriate cortical activity reflects segmentation of motion into independent sources. Neuropsychologia. In press.  Supplementary movie. Download and play in loop mode.

50. Kohler, P. J, Caplovitz, G. P., Hsieh, P.-J., Sun, J. and Tse, P. U. (2010).  Motion fading is driven by perceived, not actual angular velocity. Vision Research, doi:10.1016/j.visres.2010.03.023. See demo #18 above.

49. Hsieh, P.-J. and Tse, P. U. (2010).  BOLD Signal in Both Ipsilateral and Contralateral Retinotopic Cortex Modulates with Perceptual Fading. Supplementary figures are here. PLoS One. 5(3), e9638.

48. Tse, P. U., Baumgartner, F. J., and Greenlee, M. W. (2010).  Event-related functional MRI of cortical activity evoked by microsaccades, small visually-guided saccades, and eyeblinks in human visual cortex. Figures are here. Supplementary figures are here. Neuroimage. 49, 805-816.

47. Hsieh, P.-J. and Tse, P. U. (2010). 'Brain-reading' of perceived colors reveals a feature mixing mechanism underlying perceptual filling-in in cortical area V1. Human Brain Mapping. epub Jan. 2010. See demo #13 above.

46. Hsieh, P.-J. and Tse, P. U. (2009).  Motion fading and the motion after-effect share a common process of neural adaptation. Attention, Perception & Psychophysics. 71(4),724-733.

45. Hsieh, P.-J. and Tse, P. U. (2009).  Microsaccade rate varies with subjective visibility during motion-induced blindness. PLOS One. 4, 4, e5163.

44. Tse, P. U., Caplovitz, G. P., and Hsieh, P.-J. (2009).  Corrigendum to 'Microsaccade directions do not predict directionality of illusory brightness changes of overlapping transparent surfaces' [Vision Research 46 (2006) 3823-3830]. Vision Research 49, 790.e1-e7. This corrects and replaces reference 30 below.

43. Kohler, P. K., Caplovitz, G. P., and and Tse, P. U. (2009).  The whole moves less than the spin of its parts. Attention, Perception & Psychophysics. 71(4), 675-679. Demo of Anstis' stimuli. Experimental stimuli. Note that Ls on left seem to move more than Ls on right. The same can be seen in Stuart Anstis' original dot version. When the global pattern is seen, there appears to be less rotational motion. See demo #20 above.

42. Hsieh, P.-J. and Tse, P. U. (2009).  Feature mixing rather than feature replacement during perceptual filling-in. Vision Research 49(4):439-50. Demo #13 above. See also ref.26 below.

41. Caplovitz, G. P., Paymer, N., and Tse, P. U. (2008).  The Drifting Edge Illusion: A stationary edge abutting an oriented drifting grating appears to move because of the 'other aperture problem.' Vision Research. 48(22):2403-14. Demo #19 above

40. Chiao, J. Y., Adams, R. B., Tse, P. U., Lowenthal, W. T., Richeson, J. A. and Ambady, N. (2008).  Knowing who's boss: fMRI and ERP investigations of social dominance perception. Group Processes and Intergroup Relations, 11(2) 201-214; special issue on Social Neuroscience.

39. Caplovitz, G. P., Barroso, D. J., Hsieh, P.-J. and Tse, P. U. (2007).  fMRI reveals that non-local processing in ventral retinotopic cortex underlies perceptual grouping by temporal synchronyDemo of stimuli. Download and play in loop mode. Human Brain Mapping, 29(6):651-61.

38. Tse, P. U. and Hsieh, P.-J. (2007). Component and intrinsic motion integrate in 'dancing bar' illusion. Biological Cybernetics, 96(1):1-8. Epub 2007 Jan 26. Demo #17 above

37. Caplovitz, G. P. and Tse, P. U. (2007).  Rotating Dotted Ellipses: Motion perception driven by grouped figural rather than local dot motion signals. Vision Research. Jun 2007;47(15):1979-1991

36. Troncoso, X. G., Tse, P. U., Macknik, S. L., Caplovitz,, G. P., Hsieh, P.-J. Schlegel, A. A., Martinez-Conde, S. (2007).  BOLD activation varies parametrically with corner angle in all human retinotopic areas. Perception, 36, 808-820.

35. Hsieh, P.-J. and Tse, P. U. (2007). Grouping inhibits motion fading by giving rise to virtual trackable features. Journal of Experimental Psychology: Human Perception and Performance, 33(1):57-63 . Demo #18 above

34. Tse, P. U. and Caplovitz, G. P. (2006). Contour discontinuities subserve two types of form analysis that underlie motion processing. In: 'Progress in Brain Research: Visual Perception Part I, Fundamentals of Vision: Low and Mid-Level processes in perception.' Elsevier. 154:271-92.

33. Tse, P. U. and Hsieh, P.-J. (2006). The infinite regress illusion reveals faulty integration of local and global motion signals. Vision Research, 46(22):3881-5. Demo #16 above

32. Caplovitz, G. P. and Tse, P. U. (2006). V3A processes contour curvature as a trackable feature for the perception of rotational motion. Cerebral Cortex. 17(5):1179-89. Color versions of Figures. Fig.1 Fig.2 Fig.3 Fig.4 Fig.5 SuppFig.1 SuppFig.2

31. Caplovitz, G. P. and Tse, P. U. (2006). The Bar-Cross-Ellipse Illusion: alternating percepts of rigid and non-rigid motion based on contour ownership and trackable feature assignment. Perception. 35(7):993-7.. Demo #15 above.

30. Tse, P. U., Caplovitz, G. P., and Hsieh, P.-J. (2006). Microsaccade directions do not predict directionality of illusory brightness changes of overlapping transparent surfaces. Vision Research, 46(22):3823-30 . This is replaced by the corrigendum above in reference 44.

29. Hsieh, P.-J., Caplovitz, G. P., and Tse, P. U. (2006). Bistable Illusory Rebound Motion: Event-related functional magnetic resonance imaging of perceptual states and switches. Neuroimage, 15;32(2):728-39. Demo #11 above

28. Caplovitz, G. P., Hsieh, P.-J., and Tse, P. U. (2006). Mechanisms underlying the perceived angular velocity of a rigidly rotating object. Vision Research, 46(18):2877-93.

27. Tse, P. U. (2006). Neural correlates of transformational apparent motion. Neuroimage, 31(2): 766-73.

26. Hsieh, P.-J. and Tse, P. U. (2006). Illusory color mixing upon perceptual fading and filling-in does not result in 'forbidden colors.' Vision Research, 46(14):2251-8. Demo #13 above

25. Hsieh, P.-J. and Tse, P. U. (2006). Stimulus factors affecting Illusory Rebound Motion. Vision Research, 46(12):1924-33.

24. Hsieh, P.-J., Caplovitz, G. P., and Tse, P. U. (2006). Illusory motion induced by the offset of stationary luminance-defined gradients. Vision Research, 46(6-7):970-8.

23. Tse, P. U., Martinez-Conde, S., Schlegel, A., and Macknik, S. (2005). Visibility and visual masking of simple targets are confined to areas in the occipital cortex beyond human V1/V2. Proceedings of the National Academy of Sciences, 102, 47, 17178-17183. Scroll down to see demo of standing wave of invisibility.

22. Eagleman, D. M., Tse, P. U., Janssen, P., Nobre, A. C., Buonomano, D., and Holcombe, A. O. (2005). Time and the brain: how subjective time relates to neural time. Journal of Neuroscience, 25(45):10369-71.

21. Hsieh, P.-J., Caplovitz, G. P., and Tse, P. U. (2005). Illusory Rebound Motion and the motion continuity heuristic. Vision Research, 45(23):2972-85.

20. Tse, P. U. (2005). Voluntary attention modulates the brightness of overlapping transparent surfaces. Vision Research, 45(9):1095-8. Demo #10 above.

19. Tse, P. U. (2004). Unser Ziel muss eine Gestalt-Neurowissenschaft sein. Gestalt Theory, 287-292.

18. Tse, P. U., Rivest, J., Intriligator, J. and Cavanagh, P. (2004). Attention and the subjective expansion of time. Perception & Psychophysics, 66(7), 1171-1189. Demo #5 above

17. Tse, P. U., Sheinberg, D. L., and Logothetis, N. K. (2004). The distribution of microsaccade directions need not reveal the location of attention. Psychological Science, 15(10):708-10.

16. Tse, P. U. (2004). Mapping the distribution of spatial attention using change blindness as a probe. Cognitive Science, 28, 2, 241-258.

15. Tse, P. U. (2003). If vision is 'veridical hallucination', what keeps it veridical?. Commentary (p. 426-427) on Gestalt isomorphism and the primacy of subjective conscious experience: a Gestalt Bubble model" by Steven Lehar. Behavioral and Brain Sciences, 26(4):375-408.

14. Tse, P. U., Sheinberg, D. L., and Logothetis, N. K. (2003). Attentional enhancement opposite a peripheral flash revealed by change blindness, Psychological Science, 14, 2, 1-8.

13. Tse, P. U., Sheinberg, D. L., and Logothetis, N. K. (2002). Fixational eye movements are not affected by abrupt onsets that capture attention. Vision Research, 42, 1663-1669.

12. Tse, P. U. and Logothetis, N. K. (2002). The duration of 3-D form analysis in transformational apparent motion, Perception & Psychophysics, 64(2), 244-265.

11. Tse, P. U. (2002). A contour propagation account of surface filling-in and volume formation. Psychological Review, 109, 1, 91-115.

10. Kristjansson, A. and Tse, P. U. (2001). Curvature discontinuities are cues for rapid shape analysis. Perception & Psychophysics, 63, 390-403.

9. Tse, P. and Cavanagh, P. (2000). Chinese and Americans see opposite apparent motions in a Chinese character. Cognition, 74, B27-B32.

8. Tse, P. (2000). The sawtooth illusion. Perception. 29(7), 874-876.

7. Holcombe, A. O., Intriligator, J., & Tse, P. U. (2000). The spokes illusion originates at an early motion processing stage. Perception & Psychophysics, 62(8), 1619-1624. Demo #3 above

6. Albert, M. and Tse, P. (2000). The role of surface attachment in perceived volumetric shape. Perception, 29, 409-420.

5. Tse, P. U. (1999). Volume completion. Cognitive Psychology, 39. Demo #7 above , 37-68.

4. Tse, P. U. (1999). Complete mergeability and amodal completion. Acta Psychologica, 102, 165-201.

3. Holcombe, A. O., Macknik, S. L., Intriligator, J., Seiffert,A. E., and Tse, P. U. (1999). Wakes and spokes: New motion-induced brightness illusions. Perception, 28, 10, 1231-1242. Demo #3 above.

2. Tse, P. and Albert, M. (1998). Amodal Completion in the absence of image tangent discontinuities. Perception, 27, 455-464.

1. Tse, P. (1998). Illusory volumes from conformation. Perception, 27, 8, 977-994.


5. Tse, P. U. (2009). Attention causes the subjective expansion of time. In: 'Attention and Time', edited by Nobre and Coull. Oxford U. Press.

4. Greenlee, M. W. and Tse, P. U. (2006). Functional neuroanatomy of the human visual system: a review of functional MRI studies. In: 'Essentials in Ophthalmology,' Springer Verlag, Germany.

3. Tse, P. (2006). How the evolution of symbolic cognition transformed human morality. In: 'Moral Psychology, Volume 1: The Evolution of Morality.' MIT Press. Edited by Sinnott-Armstrong. In press 2007.

2. Tse, P. U. and Hughes, H. C. (2004). Visual Form Perception. In: 'The Encyclopedia of Neuroscience.' Adelman, G. and Smith, B. (Eds.). Elsevier.

1. Tse, P., Cavanagh, P., and Nakayama, K. (1998). The role of parsing in high-level motion processing. In: 'High-level motion processing - Computational, neurobiological and psychophysical perspectives.' Watanabe, T. (Ed.). MIT Press, pp. 249-266.

Science Books

Visual Perception Part 1, Volume 154: "Fundamentals of Vision: Low and Mid-Level Processes in Perception (Progress in Brain Research) by Susana Martinez-Conde, S. Macknik, Luis M. Martinez, Jose-Manuel Alonso and Peter U. Tse (Hardcover - Nov 13, 2006)

Visual Perception Part 2, Volume 155: "Visual Perception Part 2, Volume 155: Fundamentals of Awareness, Multi-Sensory Integration and High-Order Perception (Progress in Brain Research) by Susana Martinez-Conde, S. Macknik, Luis M. Martinez, Jose-Manuel Alonso and Peter U. Tse (Hardcover - Dec 28, 2006)


Other Books

Kansai Japanese: The language of Osaka, Kyoto, and Western Japan (1993).

Sing Japanese: The Fun Approach to Studying Japanese (1995).

Harvard Vision: Student Essays on Our Collective Future-Vol. IV (1996).

Other writings/chapters

I occasionally write essays about broader societal issues that I care about. Several of the chapters below are from the Harvard Vision book series that I worked on as a graduate student in the 1990s.

The Future of Science and Religion (1994).

The Reformation of Civil Ethics (1995).

Privacy Under Siege (1996).

The Future of Love and family in America (1997).

Thoughts on human morality and the evolution of our symbolic mind (2007).

Conference talks/posters

Other media

Interview with me on the Jim Lehrer newshour. transcript Watch the segment. Scroll down to Dec. 25, 2002.

From 2005-2009 National Geographic supported our team trying to camera trap a supposed bipedal ape in the region of Kerinci Seblat National Park, Sumatra. My initial interest in this was to determine whether there might still be any extant Homo Floresiensis. It was a long shot, but can you imagine if there had been? The reason we chose to look here is that locals spoke of a bipedal ape called 'orang pendek'. In the end we concluded that there was most likely not such an animal, at least in this region of Sumatra. National Geographic made one episode in part about our team's search called 'Is it Real?: Apeman' that you can find on netflix. Foreign team members were myself, Alex Schlegel (now at Dartmouth in grad school), Tim Mowrer and Murray Collins. Here is the team crossing the Tebo river to place camera traps in a place called Batang Ulas.

Lab Members


Current Graduate Students:

Peter Kohler with Eliza Peter's website

Alex Schlegel 1

Sergey Fogelson 1

Eric Reavis 1


Past Graduate Students:

Gideon P. Caplovitz (post-doc at Sabine Kastner's lab, Princeton, now faculty at Reno) picture 1 2

Po-Jang (Brown) Hsieh (post-doc at Nancy Kanwisher's lab at MGH Harvard, now faculty at Duke Singapore)


Past Researchers

Katie Porter 1 (Now at Alfonso Caramazza's lab at Harvard)

Mark Samco 1 2 3 

Diego J. Barroso 1

Nora Paymer


Current RAs

Scottie Alexander1

                                                                                                See the face in the neurons above?

Past RAs

Melissa Henley, Cynthia Santos, Christina Ackerman,Kendra Vierbickas, Omar Pardesi, Tamer B. Shabaneh, Jeremy Warburg


Other lab pics 1

Lab Retreat, Culebra, Puerto Rico Dec. 2004 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Some family pics kids2003 kids2006 Lilia+Henry2010 Mel+Henry Mel+Lilia Mel+Eliza Me+Eliza my brother Mike Mike+Henry family2006 My sister Barb family03 My father Kim Fung My mother Helga Our land

                                          Vision Sciences Laboratory


Last Updated: 8/23/06