Psychophysical Tests of the Hypothesis of a Bottom-Up Saliency Map in Primary Visual Cortex

A unique vertical bar among horizontal bars is salient and pops out perceptually. Physiological data have suggested that mechanisms in the primary visual cortex (V1) contribute to the high saliency of such a unique basic feature, but indicated little regarding whether V1 plays an essential or peripheral role in input-driven or bottom-up saliency. Meanwhile, a biologically based V1 model has suggested that V1 mechanisms can also explain bottom-up saliencies beyond the pop-out of basic features, such as the low saliency of a unique conjunction feature such as a red vertical bar among red horizontal and green vertical bars, under the hypothesis that the bottom-up saliency at any location is signaled by the activity of the most active cell responding to it regardless of the cell's preferred features such as color and orientation. The model can account for phenomena such as the difficulties in conjunction feature search, asymmetries in visual search, and how background irregularities affect ease of search. In this paper, we report nontrivial predictions from the V1 saliency hypothesis, and their psychophysical tests and confirmations. The prediction that most clearly distinguishes the V1 saliency hypothesis from other models is that task-irrelevant features could interfere in visual search or segmentation tasks which rely significantly on bottom-up saliency. For instance, irrelevant colors can interfere in an orientation-based task, and the presence of horizontal and vertical bars can impair performance in a task based on oblique bars. Furthermore, properties of the intracortical interactions and neural selectivities in V1 predict specific emergent phenomena associated with visual grouping. Our findings support the idea that a bottom-up saliency map can be at a lower visual area than traditionally expected, with implications for top-down selection mechanisms.

[1]  B. Bergum,et al.  Attention and performance IX , 1982 .

[2]  Stefan Treue,et al.  Feature-based attention influences motion processing gain in macaque visual cortex , 1999, Nature.

[3]  Z Li,et al.  Contextual influences in V1 as a basis for pop out and asymmetry in visual search. , 1999, Proceedings of the National Academy of Sciences of the United States of America.

[4]  D. V. van Essen,et al.  Neuronal responses to static texture patterns in area V1 of the alert macaque monkey. , 1992, Journal of neurophysiology.

[5]  D. Foster,et al.  Asymmetries in oriented-line detection indicate two orthogonal filters in early vision , 1991, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[6]  C. Koch,et al.  A saliency-based search mechanism for overt and covert shifts of visual attention , 2000, Vision Research.

[7]  H. Nothdurft Salience of Feature Contrast , 2005 .

[8]  C. Gilbert,et al.  Improvement in visual sensitivity by changes in local context: Parallel studies in human observers and in V1 of alert monkeys , 1995, Neuron.

[9]  M. Goldberg,et al.  Neuronal Activity in the Lateral Intraparietal Area and Spatial Attention , 2003, Science.

[10]  Zhaoping Li,et al.  Towards a theory of striate cortex , 1994 .

[11]  Zhaoping Li,et al.  Toward a Theory of the Striate Cortex , 1994, Neural Computation.

[12]  D. Mumford,et al.  Neural activity in early visual cortex reflects behavioral experience and higher-order perceptual saliency , 2002, Nature Neuroscience.

[13]  M. Wong-Riley,et al.  Activity correlates of cytochrome oxidase-defined compartments in granular and supragranular layers of primary visual cortex of the macaque monkey , 1995, Visual Neuroscience.

[14]  Jay Hegdé,et al.  How Selective Are V1 Cells for Pop-Out Stimuli? , 2003, The Journal of Neuroscience.

[15]  John K. Tsotsos,et al.  Neurobiology of Attention , 2005 .

[16]  R. Desimone,et al.  Neural mechanisms of selective visual attention. , 1995, Annual review of neuroscience.

[17]  D. V. van Essen,et al.  Response profiles to texture border patterns in area V1 , 2000, Visual Neuroscience.

[18]  R. Snowden,et al.  Texture segregation and visual search: a comparison of the effects of random variations along irrelevant dimensions. , 1998, Journal of experimental psychology. Human perception and performance.

[19]  D. Meyer,et al.  A Neural System for Error Detection and Compensation , 1993 .

[20]  H. Jones,et al.  Visual cortical mechanisms detecting focal orientation discontinuities , 1995, Nature.

[21]  N. P. Bichot,et al.  Perceptual and motor processing stages identified in the activity of macaque frontal eye field neurons during visual search. , 1996, Journal of neurophysiology.

[22]  S Ullman,et al.  Shifts in selective visual attention: towards the underlying neural circuitry. , 1985, Human neurobiology.

[23]  D. Hubel,et al.  Receptive fields and functional architecture of monkey striate cortex , 1968, The Journal of physiology.

[24]  S. Shipp The brain circuitry of attention , 2004, Trends in Cognitive Sciences.

[25]  B. S. Rubenstein,et al.  Spatial variability as a limiting factor in texture-discrimination tasks: implications for performance asymmetries. , 1990, Journal of the Optical Society of America. A, Optics and image science.

[26]  D. Hubel,et al.  Anatomy and physiology of a color system in the primate visual cortex , 1984, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[27]  Susan L. Franzel,et al.  Guided search: an alternative to the feature integration model for visual search. , 1989, Journal of experimental psychology. Human perception and performance.

[28]  T. Sejnowski,et al.  Representation of Color Stimuli in Awake Macaque Primary Visual Cortex , 2003, Neuron.

[29]  R. Desimone,et al.  Interacting Roles of Attention and Visual Salience in V4 , 2003, Neuron.

[30]  Eds L. Itti,et al.  The primary visual cortex creates a bottom-up saliency map , 2005 .

[31]  H. Müller,et al.  Visual search for dimensionally redundant pop-out targets: Evidence for parallel-coactive processing of dimensions , 2001, Perception & psychophysics.

[32]  A. Sillito,et al.  Surround suppression in primate V1. , 2001, Journal of neurophysiology.

[33]  Zhaoping Li V1 mechanisms and some figure-ground and border effects. , 2003, Journal of physiology, Paris.

[34]  S. Kastner,et al.  Stimulus context modulates competition in human extrastriate cortex , 2005, Nature Neuroscience.

[35]  E. J. Tehovnik,et al.  Saccadic eye movements evoked by microstimulation of striate cortex , 2003, The European journal of neuroscience.

[36]  J. Allman,et al.  Stimulus specific responses from beyond the classical receptive field: neurophysiological mechanisms for local-global comparisons in visual neurons. , 1985, Annual review of neuroscience.

[37]  J. Mollon,et al.  Do masks terminate the icon? , 2006, Quarterly journal of experimental psychology.

[38]  J. Bergen,et al.  Texture segregation and orientation gradient , 1991, Vision Research.

[39]  T. Wiesel,et al.  Clustered intrinsic connections in cat visual cortex , 1983, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[40]  E DITORS,et al.  Who and what. , 1975, Pediatrics.

[41]  Michael H. Herzog,et al.  Local Interactions in Neural Networks Explain Global Effects in Gestalt Processing and Masking , 2003, Neural Computation.

[42]  J. Nelson,et al.  Intracortical facilitation among co-oriented, co-axially aligned simple cells in cat striate cortex , 2004, Experimental Brain Research.

[43]  J. Lund,et al.  Intrinsic laminar lattice connections in primate visual cortex , 1983, The Journal of comparative neurology.

[44]  K. Nakayama,et al.  Sustained and transient components of focal visual attention , 1989, Vision Research.

[45]  L. Zhaoping,et al.  A theory of a saliency map in primary visual cortex (V1) tested by psychophysics of colour–orientation interference in texture segmentation , 2006 .

[46]  Nathalie Guyader,et al.  Investigation of the relative contributions of 3-D and 2-D image cues in texture segmentation , 2005 .

[47]  Michael H. Herzog,et al.  Effects of grouping in contextual modulation , 2002, Nature.

[48]  H. Nothdurft Salience from feature contrast: additivity across dimensions , 2000, Vision Research.

[49]  B. Julesz Textons, the elements of texture perception, and their interactions , 1981, Nature.

[50]  A. Sillito,et al.  Spatial organization and magnitude of orientation contrast interactions in primate V1. , 2002, Journal of neurophysiology.

[51]  Li Zhaoping Visual segmentation without classification: A proposed function for primary visual cortex , 1998 .

[52]  A. Treisman,et al.  A feature-integration theory of attention , 1980, Cognitive Psychology.

[53]  Victor A. F. Lamme,et al.  Figure–ground activity in primary visual cortex (V1) of the monkey matches the speed of behavioral response , 2003, Neuroscience Letters.

[54]  John Duncan,et al.  A neural basis for visual search in inferior temporal cortex , 1993, Nature.

[55]  D. V. van Essen,et al.  Response modulation by texture surround in primate area V1: Correlates of “popout” under anesthesia , 1999, Visual Neuroscience.

[56]  John K. Tsotsos Analyzing vision at the complexity level , 1990, Behavioral and Brain Sciences.

[57]  M. Landy,et al.  Discrimination of orientation-defined texture edges , 1995, Vision Research.

[58]  Nathalie Guyader,et al.  Interference with Bottom-Up Feature Detection by Higher-Level Object Recognition , 2007, Current Biology.

[59]  Z Li,et al.  Visual segmentation by contextual influences via intra-cortical interactions in the primary visual cortex. , 1999, Network.

[60]  B Julesz,et al.  "Where" and "what" in vision. , 1985, Science.

[61]  Zhaoping Li A saliency map in primary visual cortex , 2002, Trends in Cognitive Sciences.

[62]  J. Jonides Voluntary versus automatic control over the mind's eye's movement , 1981 .

[63]  C. Gilbert,et al.  Synaptic physiology of horizontal connections in the cat's visual cortex , 1991, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[64]  Li Zhaoping,et al.  The Primary Visual Cortex Creates a Bottom-up Saliency Map , 2005 .

[65]  S. Yantis,et al.  Cortical mechanisms of space-based and object-based attentional control , 2003, Current Opinion in Neurobiology.

[66]  Z Li,et al.  Pre-attentive segmentation in the primary visual cortex. , 1998, Spatial vision.

[67]  J. Duncan,et al.  Visual search and stimulus similarity. , 1989, Psychological review.

[68]  M. Goldberg,et al.  The representation of visual salience in monkey parietal cortex , 1998, Nature.