A new taxonomy for perceptual filling-in

Perceptual filling-in occurs when structures of the visual system interpolate information across regions of visual space where that information is physically absent. It is a ubiquitous and heterogeneous phenomenon, which takes place in different forms almost every time we view the world around us, such as when objects are occluded by other objects or when they fall behind the blind spot. Yet, to date, there is no clear framework for relating these various forms of perceptual filling-in. Similarly, whether these and other forms of filling-in share common mechanisms is not yet known. Here we present a new taxonomy to categorize the different forms of perceptual filling-in. We then examine experimental evidence for the processes involved in each type of perceptual filling-in. Finally, we use established theories of general surface perception to show how contextualizing filling-in using this framework broadens our understanding of the possible shared mechanisms underlying perceptual filling-in. In particular, we consider the importance of the presence of boundaries in determining the phenomenal experience of perceptual filling-in.

[1]  J Sergent,et al.  An investigation into perceptual completion in blind areas of the visual field. , 1988, Brain : a journal of neurology.

[2]  F. Kingdom,et al.  Border effects on brightness: a review of findings, models and issues. , 1988, Spatial vision.

[3]  Nava Rubin,et al.  Perceptual Completion across the Vertical Meridian and the Role of Early Visual Cortex , 2002, Neuron.

[4]  S. Shimojo,et al.  Neon Flank and Illusory Contour: Interaction between the Two Processes Leads to Color Filling-in , 1992, Perception.

[5]  S. Ullman,et al.  Neuroscience: Rewiring the adult brain , 2005, Nature.

[6]  H. F. van Tuijl A new visual illusion: neonlike color spreading and complementary color induction between subjective contours. , 1975, Acta psychologica.

[7]  Kai Hamburger,et al.  Perceptual filling-in from the edge of the blind spot , 2006, Vision Research.

[8]  Geraint Rees,et al.  Neural Correlates of Motion-induced Blindness in the Human Brain , 2010, Journal of Cognitive Neuroscience.

[9]  C. Gilbert,et al.  Topographic reorganization in the striate cortex of the adult cat and monkey is cortically mediated , 1995, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[10]  A. Dale,et al.  The Representation of Illusory and Real Contours in Human Cortical Visual Areas Revealed by Functional Magnetic Resonance Imaging , 1999, The Journal of Neuroscience.

[11]  M. Sur,et al.  Orientation Maps of Subjective Contours in Visual Cortex , 1996, Science.

[12]  R Gattass,et al.  Dynamic surrounds of receptive fields in primate striate cortex: a physiological basis for perceptual completion? , 1992, Proceedings of the National Academy of Sciences of the United States of America.

[13]  P. Weerd Perceptual filling-in: More than the eye can see. , 2006 .

[14]  R. von der Heydt,et al.  Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[15]  D. Levi,et al.  Two-dot alignment across the physiological blind spot , 1996, Vision Research.

[16]  C. Tyler,et al.  Induced twinkle aftereffect as a probe of dynamic visual processing mechanisms , 1995, Vision Research.

[17]  Niall C. Strang,et al.  Induced internal noise in perceptual artificial scotomas created by surrounding dynamic noise , 2007, Vision Research.

[18]  Daniel D. Dilks,et al.  Reorganization of visual processing in macular degeneration: Replication and clues about the role of foveal loss , 2008, Vision Research.

[19]  Janine D. Mendola,et al.  fMRI Measures of Perceptual Filling-in in the Human Visual Cortex , 2006 .

[20]  Y. Sakaguchi Contrast dependency in perceptual filling-in , 2006, Vision Research.

[21]  H. Esteky,et al.  Orientation-Selective Adaptation during Motion-Induced Blindness , 2004, Perception.

[22]  John J. Foxe,et al.  The Spatiotemporal Dynamics of Illusory Contour Processing: Combined High-Density Electrical Mapping, Source Analysis, and Functional Magnetic Resonance Imaging , 2002, The Journal of Neuroscience.

[23]  H. Komatsu The neural mechanisms of perceptual filling-in , 2006, Nature Reviews Neuroscience.

[24]  Su-Ling Yeh,et al.  A common mechanism for perceptual filling-in and motion-induced blindness , 2006, Vision Research.

[25]  Peter De Weerd,et al.  Mechanisms of Surface Completion: Perceptual Filling-In of Texture , 2003 .

[26]  V. Ramachandran,et al.  On the perception of illusory contours , 1994, Vision Research.

[27]  V. Ramachandran,et al.  Transparency: Relation to Depth, Subjective Contours, Luminance, and Neon Color Spreading , 1990, Perception.

[28]  Jess R. Kerlin,et al.  Cortical representation of space around the blind spot. , 2005, Journal of neurophysiology.

[29]  D. Hebb,et al.  Visual perception approached by the method of stabilized images. , 1960, Canadian journal of psychology.

[30]  B. Gulyás,et al.  Neuronal correlates of real and illusory contour perception: functional anatomy with PET , 1999, The European journal of neuroscience.

[31]  Ronald A. Rensink,et al.  Early completion of occluded objects , 1998, Vision Research.

[32]  Dov Sagi,et al.  Opposite Neural Signatures of Motion-Induced Blindness in Human Dorsal and Ventral Visual Cortex , 2008, The Journal of Neuroscience.

[33]  Jon Driver,et al.  Parallel detection of Kanizsa subjective figures in the human visual system , 1994, Nature.

[34]  E. Smith,et al.  Effects of Selective Attention on Perceptual Filling-in , 2006, Journal of Cognitive Neuroscience.

[35]  O. D. Creutzfeldt,et al.  Neuronal responses to borders with and without luminance gradients in cat visual cortex and dorsal lateral geniculate nucleus , 2004, Experimental Brain Research.

[36]  T. Maddess,et al.  The spatiotemporal properties of the Craik–O’Brien–Cornsweet effect are consistent with ‘filling-in’ , 1998, Vision Research.

[37]  N. Rubin,et al.  fMRI Activation in Response to Illusory Contours and Salient Regions in the Human Lateral Occipital Complex , 2003, Neuron.

[38]  K Nakayama,et al.  Stereoscopic Depth: Its Relation to Image Segmentation, Grouping, and the Recognition of Occluded Objects , 1989, Perception.

[39]  Alejandro Lleras,et al.  What You See Is What You Get , 2006, Psychological science.

[40]  Hong Zhou,et al.  Searching For The Neural Mechanism Of Color Filling-In , 2003 .

[41]  Takeo Watanabe,et al.  The primary visual cortex fills in color , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[42]  J. Kaas,et al.  Rapid reorganization of cortical maps in adult cats following restricted deafferentation in retina , 1992, Vision Research.

[43]  Erich W Graf,et al.  Modulating motion-induced blindness with depth ordering and surface completion , 2002, Vision Research.

[44]  Jeffrey S. Johnson,et al.  The recognition of partially visible natural objects in the presence and absence of their occluders , 2005, Vision Research.

[45]  Shimon Ullman,et al.  Filling-in of retinal scotomas , 2003, Vision Research.

[46]  R van Lier,et al.  Integrating Global and Local Aspects of Visual Occlusion , 1994, Perception.

[47]  F. Sengpiel,et al.  Reorganization of Visual Cortical Maps after Focal Ischemic Lesions , 2003, Journal of cerebral blood flow and metabolism : official journal of the International Society of Cerebral Blood Flow and Metabolism.

[48]  Dov Sagi,et al.  Motion-induced blindness in normal observers , 2001, Nature.

[49]  J. O'Regan,et al.  Solving the "real" mysteries of visual perception: the world as an outside memory. , 1992, Canadian journal of psychology.

[50]  B. Wandell,et al.  V1 projection zone signals in human macular degeneration depend on task, not stimulus. , 2008, Cerebral cortex.

[51]  U. Eysel,et al.  Increased receptive field size in the surround of chronic lesions in the adult cat visual cortex. , 1999, Cerebral cortex.

[52]  A. Nieder,et al.  Seeing more than meets the eye: processing of illusory contours in animals , 2002, Journal of Comparative Physiology A.

[53]  T. Hendler,et al.  Object-completion effects in the human lateral occipital complex. , 2002, Cerebral cortex.

[54]  J. Driver,et al.  Segmentation, attention and phenomenal visual objects , 2001, Cognition.

[55]  Yoshimichi Ejima,et al.  Magnetic responses of human visual cortex to illusory contours , 2002, Neuroscience Letters.

[56]  S Grossberg,et al.  Neural dynamics of brightness perception: Features, boundaries, diffusion, and resonance , 1984, Perception & Psychophysics.

[57]  R Shapley,et al.  Illusory contours activate specific regions in human visual cortex: evidence from functional magnetic resonance imaging. , 1995, Proceedings of the National Academy of Sciences of the United States of America.

[58]  H. V. Tuijl,et al.  A new visual illusion: Neonlike color spreading and complementary color induction between subjective contours , 1975 .

[59]  E. Leeuwenberg,et al.  Coding theory of visual pattern completion. , 1981, Journal of experimental psychology. Human perception and performance.

[60]  S. Dakin,et al.  Natural image statistics mediate brightness ‘filling in’ , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[61]  M. Livingstone,et al.  Perceptual and Physiological Evidence for a Role for Early Visual Areas in Motion-induced Blindness Introduction , 2022 .

[62]  R. von der Heydt,et al.  Color Filling-in under Steady Fixation: Behavioral Demonstration in Monkeys and Humans , 1999, Perception.

[63]  J. Hegdé,et al.  Preferential responses to occluded objects in the human visual cortex. , 2008, Journal of vision.

[64]  T. Watanabe,et al.  The relation between color spreading and illusory contours , 1990, Perception & psychophysics.

[65]  A. Dale,et al.  The representation of the ipsilateral visual field in human cerebral cortex. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[66]  Peter De Weerd,et al.  Responses of cells in monkey visual cortex during perceptual filling-in of an artificial scotoma , 1995, Nature.

[67]  Leila Montaser-Kouhsari,et al.  Orientation-selective adaptation to illusory contours in human visual cortex. , 2010, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[68]  E. Mingolla,et al.  Neon Color Spreading: A Review , 1997, Perception.

[69]  R. von der Heydt,et al.  Mechanisms of contour perception in monkey visual cortex. II. Contours bridging gaps , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[70]  D. Burr Implications of the Craik-O'Brien illusion for brightness perception , 1987, Vision Research.

[71]  Geraint Rees,et al.  Attentional influences on the dynamics of motion-induced blindness. , 2009, Journal of vision.

[72]  R S Weil,et al.  Neural correlates of perceptual filling-in of an artificial scotoma in humans , 2007, Proceedings of the National Academy of Sciences.

[73]  Christof Koch,et al.  Motion-induced blindness does not affect the formation of negative afterimages , 2004, Consciousness and Cognition.

[74]  Eric Halgren,et al.  Cortical activation to illusory shapes as measured with magnetoencephalography , 2003, NeuroImage.

[75]  John J. Foxe,et al.  Setting Boundaries: Brain Dynamics of Modal and Amodal Illusory Shape Completion in Humans , 2004, The Journal of Neuroscience.

[76]  Lothar Spillmann,et al.  Texture fading correlates with stimulus salience , 2001, Vision Research.

[77]  R. L. Gregory,et al.  Perceptual filling in of artificially induced scotomas in human vision , 1991, Nature.

[78]  V. Ramachandran Filling in the blind spot , 1992, Nature.

[79]  G Westheimer,et al.  A quantitative measure for short-term cortical plasticity in human vision , 1994, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[80]  J. Bakin,et al.  Visual Responses in Monkey Areas V1 and V2 to Three-Dimensional Surface Configurations , 2000, The Journal of Neuroscience.

[81]  Leslie G. Ungerleider,et al.  Perceptual filling-in: a parametric study , 1998, Vision Research.

[82]  Stephen Grossberg,et al.  Filling-in the Forms: Surface and Boundary Interactions in Visual Cortex , 1998 .

[83]  R. Fleming,et al.  The Interpolation of Object and Surface Structure , 2002, Cognitive Psychology.

[84]  Lothar Spillmann,et al.  Dynamic noise backgrounds facilitate target fading , 1992, Vision Research.

[85]  M Dojat,et al.  Moving illusory contours activate primary visual cortex: an fMRI study. , 2000, Cerebral cortex.

[86]  Vivian O'Brien,et al.  Contour Perception, Illusion and Reality* , 1958 .

[87]  F. Campbell,et al.  A comparison of threshold and suprathreshold appearance of gratings with components in the low and high spatial frequency range , 1978, The Journal of physiology.

[88]  Stephen A. Engel,et al.  Interocular rivalry revealed in the human cortical blind-spot representation , 2001, Nature.

[89]  N. Kanwisher,et al.  Reorganization of Visual Processing in Macular Degeneration , 2005, The Journal of Neuroscience.

[90]  Ikuya Murakami,et al.  Neural responses in the primary visual cortex of the monkey during perceptual filling-in at the blind spot , 2002, Neuroscience Research.

[91]  T. S. Lee,et al.  Dynamics of subjective contour formation in the early visual cortex. , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[92]  K. Craik The nature of psychology , 1966 .

[93]  R. W. DITCHBURN,et al.  Vision with a Stabilized Retinal Image , 1952, Nature.

[94]  Su-Ling Yeh,et al.  Linking motion-induced blindness to perceptual filling-in , 2004, Vision Research.

[95]  Peter De Weerd,et al.  Perceptual filling-in: More than the eye can see. , 2006, Progress in brain research.

[96]  Earl L. Smith,et al.  Recovery of binocular responses by cortical neurons after early monocular lesions , 2001, Nature Neuroscience.

[97]  Yoichi Sugita,et al.  Grouping of image fragments in primary visual cortex , 1999, Nature.

[98]  Heiko Neumann,et al.  Visual filling-in for computing perceptual surface properties , 2001, Biological Cybernetics.

[99]  Taosheng Liu,et al.  Temporally Unfolding Neural Representation of Pictorial Occlusion , 2006, Psychological science.

[100]  H. Gerrits,et al.  The filling-in process in patients with retinal scotomata. , 1969, Vision research.

[101]  Julie M. Harris,et al.  Filling-in the details on perceptual fading , 2001, Vision Research.

[102]  Interocular Transfer in the Disappearance of Stabilized Images , 1959 .

[103]  S. Dakin,et al.  Illusory Stimuli Can Be Used to Identify Retinal Blind Spots , 2007, PloS one.

[104]  S Grossberg,et al.  3-D vision and figure-ground separation by visual cortex , 2010, Perception & psychophysics.

[105]  Xoana G. Troncoso,et al.  Microsaccades Counteract Visual Fading during Fixation , 2005, Neuron.

[106]  Lars Muckli,et al.  Separate cortical stages in amodal completion revealed by functional magnetic resonance adaptation , 2007, BMC Neuroscience.

[107]  Geraint Rees,et al.  Monocular signals in human lateral geniculate nucleus reflect the Craik-Cornsweet-O'Brien effect. , 2009, Journal of vision.

[108]  T. Wiesel,et al.  Receptive field dynamics in adult primary visual cortex , 1992, Nature.

[109]  M. Concetta Morrone,et al.  Neuronal Mechanisms for Illusory Brightness Perception in Humans , 2005, Neuron.

[110]  F. Boselie,et al.  A critical discussion of Kellman and Shipley's (1991) theory of occlusion phenomena , 1992, Psychological research.

[111]  Taosheng Liu,et al.  Retinotopic mapping of the visual cortex using functional magnetic resonance imaging in a patient with central scotomas from atrophic macular degeneration. , 2004, Ophthalmology.

[112]  Daniel D. Dilks,et al.  Human Adult Cortical Reorganization and Consequent Visual Distortion , 2007, The Journal of Neuroscience.

[113]  P. de Weerd,et al.  Effects of Selective Attention on Perceptual Filling-in , 2006 .

[114]  M. Seghier,et al.  Functional Neuroimaging Findings on the Human Perception of Illusory Contours , 2022 .

[115]  Geraint Rees,et al.  Neural correlates of perceptual completion of an artificial scotoma in human visual cortex measured using functional MRI , 2008, NeuroImage.

[116]  L. Pessoa,et al.  Finding out about filling-in: a guide to perceptual completion for visual science and the philosophy of perception. , 1998, The Behavioral and brain sciences.

[117]  S. Grossberg,et al.  The watercolor illusion and neon color spreading: a unified analysis of new cases and neural mechanisms. , 2005, Journal of the Optical Society of America. A, Optics, image science, and vision.

[118]  K. Gegenfurtner,et al.  Temporal properties of the chromatic and achromatic Craik–O’Brien–Cornsweet effect , 2007, Vision Research.

[119]  Michael D Crossland,et al.  Spatial alignment over retinal scotomas. , 2009, Investigative ophthalmology & visual science.

[120]  J. Mattingley,et al.  Preattentive Filling-in of Visual Surfaces in Parietal Extinction , 1997, Science.

[121]  Hidehiko Komatsu,et al.  Neural responses in the macaque v1 to bar stimuli with various lengths presented on the blind spot. , 2005, Journal of neurophysiology.

[122]  Georg Schweigart,et al.  Activity‐dependent receptive field changes in the surround of adult cat visual cortex lesions , 2002, The European journal of neuroscience.

[123]  P. Kellman,et al.  A theory of visual interpolation in object perception , 1991, Cognitive Psychology.

[124]  C. Hung,et al.  Building surfaces from borders in Areas 17 and 18 of the cat , 2001, Vision Research.

[125]  L. Pessoa,et al.  Filling-in: From perceptual completion to cortical reorganization. , 2003 .

[126]  A. Metha,et al.  Artificial scotoma-induced perceptual distortions are orientation dependent and short lived , 2004, Visual Neuroscience.

[127]  C. Hung,et al.  Cortical processing of a brightness illusion. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[128]  G. Butterworth,et al.  Michotte's experimental phenomenology of perception , 1994 .

[129]  Rafael Malach,et al.  Rapid completion effects in human high-order visual areas , 2004, NeuroImage.

[130]  Alice Mado Proverbio,et al.  Electrophysiological indexes of illusory contours perception in humans , 2002, Neuropsychologia.

[131]  C. Gilbert,et al.  Axonal sprouting accompanies functional reorganization in adult cat striate cortex , 1994, Nature.

[132]  H. A. Pham,et al.  Perceptual deficits after lesions of inferotemporal cortex in macaques. , 2000, Cerebral cortex.

[133]  G. Sáry,et al.  Illusory shape representation in the monkey inferior temporal cortex , 2007, The European journal of neuroscience.

[134]  I. Murakami,et al.  Motion aftereffect after monocular adaptation to filled-in motion at the blind spot , 1995, Vision Research.

[135]  Stuart Anstis,et al.  Filling-in afterimage colors between the lines , 2009, Current Biology.

[136]  R. Haber,et al.  Visual Perception , 2018, Encyclopedia of Database Systems.

[137]  Peter Ulric Tse,et al.  Volume Completion , 1999, Cognitive Psychology.

[138]  F. Kruggel,et al.  Hemodynamic and Electroencephalographic Responses to Illusory Figures: Recording of the Evoked Potentials during Functional MRI , 2001, NeuroImage.

[139]  N. Logothetis,et al.  Lack of long-term cortical reorganization after macaque retinal lesions , 2005, Nature.

[140]  J. Widdicombe,et al.  Physical characteristics of the chest and lungs and the work of breathing in different mammalian species , 1961, The Journal of physiology.

[141]  S. Pollmann,et al.  Retinotopic Activation in Response to Subjective Contours in Primary Visual Cortex , 2008, Frontiers in human neuroscience.

[142]  ANDREW SMITH,et al.  Tilt aftereffects with subjective contours , 1975, Nature.

[143]  L. Spillmann,et al.  The Neon Color Effect in the Ehrenstein Illusion , 1981, Perception.

[144]  Guy Orban,et al.  Illusory, motion, and luminance-defined contours interact in the human visual system , 1994, Vision Research.

[145]  K Nakayama,et al.  Experiencing and perceiving visual surfaces. , 1992, Science.

[146]  S. Sharma,et al.  fMRI Measures of Perceptual Filling-in in the Human Visual Cortex , 2006, Journal of Cognitive Neuroscience.