Finding out About Filling-In: a Guide to Perceptual Completion for Visual Science and the Philosophy of Perception

Total Page:16

File Type:pdf, Size:1020Kb

Finding out About Filling-In: a Guide to Perceptual Completion for Visual Science and the Philosophy of Perception BEHAVIORAL AND BRAIN SCIENCES (1998) 21, 723–802 Printed in the United States of America Finding out about filling-in: A guide to perceptual completion for visual science and the philosophy of perception Luiz Pessoa Department of Computer and Systems Engineering, Center of Technology, Federal University of Rio de Janeiro, llha do Fundao, Rio de Janeiro, RJ 21945-970, Brazil [email protected] www.cos.ufrj.br/~pessoa Evan Thompson1 Department of Philosophy and Centre for Vision Research, York University, North York, Ontario, Canada M3J 1P3 [email protected] www.yorku.ca/research/vision/evant.htm Alva Noë Department of Philosophy, University of California, Santa Cruz, Santa Cruz, CA 95064 [email protected] www2.ucsc.edu/people/anoe/ Abstract: In visual science the term filling-in is used in different ways, which often leads to confusion. This target article presents a tax- onomy of perceptual completion phenomena to organize and clarify theoretical and empirical discussion. Examples of boundary com- pletion (illusory contours) and featural completion (color, brightness, motion, texture, and depth) are examined, and single-cell studies relevant to filling-in are reviewed and assessed. Filling-in issues must be understood in relation to theoretical issues about neural–per- ceptual isomorphism and linking propositions. Six main conclusions are drawn: (1) visual filling-in comprises a multitude of different perceptual completion phenomena; (2) certain forms of visual completion seem to involve spatially propagating neural activity (neural filling-in) and so, contrary to Dennett’s (1991; 1992) recent discussion of filling-in, cannot be described as results of the brain’s “ignor- ing an absence” or “jumping to a conclusion”; (3) in certain cases perceptual completion seems to have measurable effects that depend on neural signals representing a presence rather than ignoring an absence; (4) neural filling-in does not imply either “analytic isomor- phism” or “Cartesian materialism,” and thus the notion of the bridge locus – a particular neural stage that forms the immediate substrate of perceptual experience – is problematic and should be abandoned; (5) to reject the representational conception of vision in favor of an “enactive” or “animate” conception reduces the importance of filling-in as a theoretical category in the explanation of vision; and (6) the evaluation of perceptual content should not be determined by “subpersonal” considerations about internal processing, but rather by considerations about the task of vision at the level of the animal or person interacting with the world. Keywords: blind spot; bridge locus; brightness; consciousness; filling-in; Gestalt theory; illusory contours; isomorphism; linking propo- sitions; perceptual completion; vision 1. Introduction that something is present in a particular region of visual space when it is actually absent from that region, but pres- Figure 1 illustrates the so-called neon color illusion: a red ent in the surrounding area. The idea is easiest to under- diamond is seen where there is only a lattice of red line seg- stand in the case of the blind spot. We have a blind spot in ments. The term neon color spreading (van Tuijl 1975) is of- each eye corresponding to the region where the optic nerve ten used to describe this phenomenon: the figure seems to leaves the retina and there are no photoreceptors (Fig. 2). result from the color having spread between the line seg- In everyday perception we are never aware of the blind ments into the background. The color is said to “fill in” the spot. The blind spots of the two eyes do not overlap, and background, thereby forming the figure. Nevertheless, one stimulus that falls on the blind spot of one retina will fall does not see any spreading or filling-in process; one sees outside the blind spot of the other retina. Even under only the figure. What line of reasoning, then, lies behind the monocular viewing the blind spot is not easily revealed. description of such figures in the terminology of “filling- Close one eye and fixate on a point on a uniformly colored in”? piece of paper: there is no experience of any gap or discon- Visual scientists use the terms filling-in and perceptual tinuity in the visual field. Now follow the instructions for completion to refer to situations in which subjects report the blind spot demonstration in Figure 3. The left dot dis- © 1998 Cambridge University Press 0140-525X/9X $12.50 723 Pessoa et al.: Perceptual completion appears and one perceives a uniform expanse of brightness and color. This is an example of perceptual completion or visual filling-in: the color and brightness surrounding the area corresponding to the blind spot are said to “fill in” that area so that a uniform expanse is perceived. The existence of such perceptual completion phenom- ena at the subject level is uncontroversial. However, the term filling-in is often used in a controversial sense that goes beyond what subjects report. In this controversial sense, the term filling-in suggests that certain kinds of sub- ject-level perceptual completion phenomena are accom- plished by the brain providing information to make up for an absence – by the brain actively filling in the missing in- formation. However, whether there is neural filling-in is a matter of great debate in visual science. We will argue later Figure 1. Neon color illusion. When the segments in the dia- that there is considerable evidence for neural filling-in, but mond are red and the matrix is black (left), the entire diamond ap- that great care needs to be taken in thinking about the re- pears reddish even though only the segments are physically col- lation between neural filling-in and subject-level percep- ored red (right). tual completion. To appreciate the debates about filling-in it is necessary 1962; 1968). In other words, many neurons respond more first to review certain facts about vision and second to dis- strongly to boundaries than to regions or surfaces. Second, cuss certain conceptual and methodological points. Many psychophysical experiments (e.g., with stabilized images) of the objects we perceive have roughly uniform regions of have shown the importance of boundaries for proper sur- surface color and lightness. Now consider two facts. First, face perception (Krauskopf 1963; Yarbus 1967). In the clas- cells in the visual cortex in general do not respond to uni- sic study by Krauskopf (1963), an inner green disc was sur- form regions, but rather to discontinuities (Hubel & Wiesel rounded by a red annulus (Fig. 4). When the red-green boundary was stabilized on the retina (so that it always maintained a fixed position on the eye), subjects reported Luiz Pessoa is Assistant Professor of Computer and Systems Engineering at the Federal University of Rio de Janeiro, Brazil. He received a Ph.D. in Cognitive and Neural Systems from Boston University and B.Sc. and M.Sc. degrees from the Federal University of Rio de Janeiro. He has published articles on computational modeling of vision, visual psychophysics, and neural network models, among others. Evan Thompson is an Associate Professor in the Department of Philosophy and a member of the Centre for Vision Research at York University. He received his Ph.D. in philosophy from the University of Toronto. He is the author of Colour Vision: A Study in Cognitive Science and the Philosophy of Perception (Routledge Press, 1995), and co-author, with Francisco J. Varela and Eleanor Rosch, of The Embodied Mind: Cognitive Science and Human Experience (MIT Press, 1991). He has published a number of articles in philosophy and cognitive science, including a previous Behavioral and Brain Sciences target article on color vision. Alva Noë is Assistant Professor of Philosophy at the University of California, Santa Cruz. He received a Ph.D. in philosophy from Harvard Figure 2. The blind spot. (A) Vertebrate lens and retina. The University and a B.Phil. from Oxford blind spot corresponds to the area where the optic nerve leaves University, and has been a Research the retina and there are no photoreceptors (Sheperd 1994; used Associate of the Center for Cognitive by permission of Oxford University Press, Inc.). (B) Distribution Studies at Tufts University. He has of photoreceptors over the extent of the retina of the right eye. published articles on topics in the philosophy of Note the complete absence of photoreceptors in the blind spot re- perception, philosophy of mind, and other areas. gion (Sekuler & Blake 1985, p. 56; used by permission of the McGraw-Hill Companies). 724 BEHAVIORAL AND BRAIN SCIENCES (1998) 21:6 Pessoa et al.: Perceptual completion the case of brightness perception there is good evidence for neural filling-in that involves spatially propagating activity. However, the filling-in controversy is not only empirical, because it involves fundamental conceptual and method- ologic issues. To appreciate these issues it is necessary to in- troduce the concepts of the bridge locus and neural–per- ceptual isomorphism. The best way to introduce these concepts is by way of an example. Figure 5 presents another case often discussed in connection with filling-in, the Craik- O’Brien-Cornsweet effect (Cornsweet 1970; Craik 1940; O’Brien 1958). Two largely uniform regions of different brightness are seen, although most of the corresponding stimulus regions have exactly the same luminance. In fact, the two regions differ only in the luminance distribution at Figure 3. Blind spot demonstration. Close your right eye while fixating at the cross. Adjust the distance of the page in front of you the “cusp” edge separating the two regions. Why, then, do until the left dot disappears (start at a distance of approximately 8 we see a brightness step? inches). A similar procedure should be used with the left eye Here is one route leading to an answer to this question closed.
Recommended publications
  • Guess Who? Answer
    Guess Who? Answer soon became interested in ophthalmology and was appointed Hansen Grut's assistant in 1879. Bjerrum's scientific concern was the relationship between visual perception of form and the resolving power in localized areas of the retina. He demonstrated this in his thesis entitled ' UndersØgeleser over Formsans og Lyssands i forskellige Øjensyngdomme (Investigations on the form sense and light sense in various eye diseases). This title is deliberately given in Danish to indicate that through his entire lifetime it was mandatory for him to write his publications in Danish. An antipathy against German, in those days the language of science, may have been gained in a childhood so filled with tension regarding nationalism. The scientific achievement that made the name Bjerrum universally known was conceived during his work on the relationship between visual acuity and the perception of the bright stimuli in various retinal zones. In accordance with his own modest attitude, this discovery was published in 1889 in a small paper which in translation was called 'An addendum to the usual examination of the visual field of glaucoma'. At that time Bjerrum was studying the visual field by means of small white objects. The idea Jannik Peterson Bjerrum of this investigation was to record the performance of every single functional unit of the retina. As a Danish ophthalmologist. Born 1851, died 1920 minimum such units in Bjerrum's opinion would subtend a visual angle of one minute of arc (in the Jannik Petersen Bjerrum was born 26th December 1851 macular region). However, even a small test object in Skarbak, a village in the most southern part of would subtend a visual angle exceeding two degrees Jutland in the border district between the Danish and accordingly cover a multitude of functional units.
    [Show full text]
  • Imaging Retinotopic Maps in the Human Brain ⇑ Brian A
    Vision Research 51 (2011) 718–737 Contents lists available at ScienceDirect Vision Research journal homepage: www.elsevier.com/locate/visres Review Imaging retinotopic maps in the human brain ⇑ Brian A. Wandell , Jonathan Winawer Psychology Department, Stanford University, Stanford, CA 94305, United States article info abstract Article history: A quarter-century ago visual neuroscientists had little information about the number and organization of Received 5 April 2010 retinotopic maps in human visual cortex. The advent of functional magnetic resonance imaging (MRI), a Received in revised form 2 August 2010 non-invasive, spatially-resolved technique for measuring brain activity, provided a wealth of data about Available online 6 August 2010 human retinotopic maps. Just as there are differences amongst non-human primate maps, the human maps have their own unique properties. Many human maps can be measured reliably in individual sub- Keywords: jects during experimental sessions lasting less than an hour. The efficiency of the measurements and the Visual field maps relatively large amplitude of functional MRI signals in visual cortex make it possible to develop quanti- Retinotopy tative models of functional responses within specific maps in individual subjects. During this last quarter- Human visual cortex Functional specialization century, there has also been significant progress in measuring properties of the human brain at a range of Optic radiation length and time scales, including white matter pathways, macroscopic properties of gray and white mat- Visual field map clusters ter, and cellular and molecular tissue properties. We hope the next 25 years will see a great deal of work that aims to integrate these data by modeling the network of visual signals.
    [Show full text]
  • Interaction Among Ocularity, Retinotopy and On-Center/Off­ Center Pathways During Development
    INTERACTION AMONG OCULARITY, RETINOTOPY AND ON-CENTER/OFF­ CENTER PATHWAYS DURING DEVELOPMENT Shigeru Tanaka Fundamental Research Laboratories, NEC Corporation, 34 Miyukigaoka, Tsukuba, Ibaraki 305, Japan ABSTRACT The development of projections from the retinas to the cortex is mathematically analyzed according to the previously proposed thermodynamic formulation of the self-organization of neural networks. Three types of submodality included in the visual afferent pathways are assumed in two models: model (A), in which the ocularity and retinotopy are considered separately, and model (B), in which on-center/off-center pathways are considered in addition to ocularity and retinotopy. Model (A) shows striped ocular dominance spatial patterns and, in ocular dominance histograms, reveals a dip in the binocular bin. Model (B) displays spatially modulated irregular patterns and shows single-peak behavior in the histograms. When we compare the simulated results with the observed results, it is evident that the ocular dominance spatial patterns and histograms for models (A) and (B) agree very closely with those seen in monkeys and cats. 1 INTRODUCTION A recent experimental study has revealed that spatial patterns of ocular dominance columns (ODes) observed by autoradiography and profiles of the ocular dominance histogram (ODH) obtained by electrophysiological experiments differ greatly between monkeys and cats. ODes for cats in the tangential section appear as beaded patterns with an irregularly fluctuating bandwidth (Anderson, Olavarria and Van Sluyters 1988); ODes for monkeys are likely to be straight parallel stripes (Hubel, Wiesel and LeVay, 1977). The typical ODH for cats has a single peak in the middle of the ocular dominance corresponding to balanced response in ocularity (Wiesel and Hubel, 1974).
    [Show full text]
  • Twelfth International Visual Field Symposium
    PERIMETRY UPDATE 1996/1997 PERIMETRY UPDATE 199611997 Proceedings of the Xllth International Perimetric Society Meeting Wijrzburg, Germany, June 4-8, 1996 edited by Michael Wall and Anders Heijl KUGLER PUBLICATIONS Amsterdam / New York ISBN 90-6299-139-4 Distributors For the LJ S A and Canada: DEMOS 386 Park Avenue South, Suite 201 New York, NY 10016 Telefax (+212) 683-0072 For all other countries Kugler Publications P.O. Box 11188 1001 GD Amsterdam, The Netherlands 0 Copyright 1997 Kugler Publications All rights reserved No part of this book may be translated or reproduced in any form by print, photoprint, microfilm, or any other means without prior written permission of the publisher. Table of Contents v TABLE OF CONTENTS Preface xi New methods of perimetry Contrast sensitivity perimetry in experimental glaucoma: investigations with degenerate gratings R.S. Harwerth and E.L. Smith, III 3 The role of spatial and temporal factors in frequency-doubling perimetry C.A. Johnson and S. Demirel 13 Motion detection perimetry: properties and results M. Wall, C.F. Brito and K. Kutzko 21 Stimulus orientation can affect motion sensitivity in glaucoma M.C. Westcott, F.W. Fitzke and R.A. Hitchings 35 Short-wavelength automated perimetry and motion automated perimetry in glaucoma P.A. Sample, Ch.F. Bosworth, I. Irak and R.N. Weinreb 43 Blue-on-yellow perimetry in patients with ocular hypertension H. Maeda, Y. Tanaka and T. Sugiura 45 Mass screening for visual field defects with snowfield campimetry: results of a field study using local TV broadcasting A.C. Gisolf, J. Kirsch, H.K.
    [Show full text]
  • Retinotopy Versus Face Selectivity in Macaque Visual Cortex
    Retinotopy versus Face Selectivity in Macaque Visual Cortex Reza Rajimehr1,2, Natalia Y. Bilenko1, Wim Vanduffel1,3, 1 and Roger B. H. Tootell Downloaded from http://mitprc.silverchair.com/jocn/article-pdf/26/12/2691/1782578/jocn_a_00672.pdf by MIT Libraries user on 17 May 2021 Abstract ■ Retinotopic organization is a ubiquitous property of lower- Distinct subregions within face-selective patches showed tier visual cortical areas in human and nonhuman primates. In either (1) a coarse retinotopic map of eccentricity and polar macaque visual cortex, the retinotopic maps extend to higher- angle, (2) a retinotopic bias to a specific location of visual order areas in the ventral visual pathway, including area TEO field, or (3) nonretinotopic selectivity. In general, regions in the inferior temporal (IT) cortex. Distinct regions within IT along the lateral convexity of IT cortex showed more overlap cortex are also selective to specific object categories such as between retinotopic maps and face selectivity, compared with faces. Here we tested the topographic relationship between regions within the STS. Thus, face patches in macaques can be retinotopic maps and face-selective patches in macaque visual subdivided into smaller patches with distinguishable retino- cortex using high-resolution fMRI and retinotopic face stimuli. topic properties. ■ INTRODUCTION has been recently challenged by the demonstration of small Visual cortical areas are typically defined on the basis of receptive fields in anterior IT cortex (DiCarlo & Maunsell, differences in histology (cytoarchitecture and myelo- 2003; Op De Beeck & Vogels, 2000). architecture), anatomical connections, functional proper- How do the retinotopic maps in IT cortex relate to ties, and retinotopic organization (Felleman & Van Essen, the functionally defined category-selective areas? Neuro- 1991).
    [Show full text]
  • VISUAL FIELD Pathway Extends from the „Front‟ to the „Back‟ of the RETINA Brain
    NOTE: To change the image on this slide, select the picture and delete it. Then click the Pictures icon in the placeholde r to insert your own image. Visual Pathway Disorders Amr Hassan, MD, FEBN Associate professor of Neurology - Cairo University Optic nerve • Anatomy of visual pathway • How to examine • Visual pathway disorders • Quiz 2 Optic nerve • Anatomy of visual pathway • How to examine • Visual pathway disorders • Quiz 3 Optic nerve The Visual Pathway VISUAL FIELD Pathway extends from the „front‟ to the „back‟ of the RETINA brain. ON OC OT LGN OPTIC RADIATIONS ON = Optic Nerve OC = Optic Chiasm OT = Optic Tract LGN = Lateral Geniculate Nucleus of Thalamus VISUAL CORTEX 5 The Visual Pathway Eyes & Retina Light >> lens >> retina (inverted and reversed image). Eyes & Retina Eyes & Retina • Macula: oval region approximately 3-5 mm that surrounds the fovea, also has high visual acuity. • Fovea: central fixation point of each eye// region of the retina with highest visual acuity. Eyes & Retina • Optic disc: region where axons leaving the retina gather to form the Optic nerve. Eyes & Retina • Blind spot located 15° lateral and inferior to central fixation point of each eye. Object to be seen Peripheral Retina Central Retina (fovea in the macula lutea) 12 Photoreceptors © Stephen E. Palmer, 2002 Photoreceptors Cones • Cone-shaped • Less sensitive • Operate in high light • Color vision • Less numerous • Highly represented in the fovea >> have high spatial & temporal resolution >> they detect colors. © Stephen E. Palmer, 2002 Photoreceptors Rods • Rod-shaped • Highly sensitive • Operate at night • Gray-scale vision • More numerous than cons- 20:1, have poor spatial & temporal resolution of visual stimuli, do not detect colors >> vision in low level lighting conditions © Stephen E.
    [Show full text]
  • Functional Field of Vision
    Functional Field of Vision Lea Hyvärinen Lea-Test Ltd, Apollonkatu 6 A 4, FIN-00100 Helsinki The size and the quality of visual field are important basic functions to be assessed as a part of functional vision assessment. Evaluation of functional field losses can be organized as follows: 1. Major field losses: A. Field losses due to pathway damage - half field loss, right, left remaining motion perception in ‘blind’ field half - quadrant losses B. Field losses due to disorders of the eyes - ROP - Congenital Glaucoma - Coloboma - Retinitis Pigmentosa and related disorders C. Central Scotoma 2. Minor field losses 3. Distortion of the image 4. Perceptual losses without measurable field loss 5. Restriction of functional visual field due to motor problems It may be wise to clarify some definitions: Anterior visual pathways contain eyes and the pathway up to the lateral geniculate nucleus (LGN), and posterior visual pathways contain the nerve fibres from the LGN to the primary visual cortex. Anterior visual impairment is due to damage to anterior pathways. Posterior visual impairment is due to damage to posterior visual pathways but may also be due to abnormal function in higher visual functions in the associative visual cortices without pathway damage. Information on visual field is given as drawings that depict the loss of function as it is projected onto the physical space around us. However, because central parts of visual field occupy much more cortical area in the primary visual cortex, they are magnified (= cortical magnification) in relation to the peripheral parts of the visual field (as illustrated in Figure 1).
    [Show full text]
  • Neon Colour Spreading in Three-Dimensional Illusory Objects in Humans
    Neuroscience Letters 281 (2000) 119±122 www.elsevier.com/locate/neulet Neon colour spreading in three-dimensional illusory objects in humans Marja Liinasuoa,*, Ilpo Kojob, Jukka HaÈ kkinenc, Jyrki Rovamod aInstitute of Biomedicine, Department of Physiology, P.O. Box 9, 00014 University of Helsinki, Helsinki, Finland bThe Finnish Institute of Occupational Health, Brain Work Laboratory, Topeliuksenkatu 41 a A, 00250 Helsinki, Finland cDepartment of Psychology, General Psychology Division, P.O. Box 13, 00014 University of Helsinki, Helsinki, Finland dDepartment of Optometry and Vision Sciences, University of Wales, College of Cardiff, P.O. Box 905, Cardiff CF1 3XF, UK Received 11 October 1999; received in revised form 7 January 2000; accepted 10 January 2000 Abstract We studied whether neon spreading can be induced within three-dimensional illusory triangles. Kanizsa triangles were induced by black pacman disks consisting of red sectors with curved sides. Viewing our stimuli monocularly produced two-dimensional illusory contours and surfaces as well as neon spreading in each ®gure. Triangles appeared concave or convex under stereoscopical viewing. Neon colour spreading was induced within illusory ®gures bending in three-dimensional space, suggesting that neural contour completion and surface ®lling-in interact across depth. Surpris- ingly, neon spreading was induced above the intervening surface even when the inducers were below the surface. Neon colour and illusory con®guration were preserved behind the intervening surface only when it appeared transparent. q 2000 Elsevier Science Ireland Ltd. All rights reserved. Keywords: Vision; Perception; Illusion; Three-dimensionality; Colour; Neon spreading Illusory ®gures with one to three dimensions are created zontal and vertical crossing lines so that some line elements by the neural processes of visual area V2 [15] between some are replaced by different colour or luminance, the surround- physically existing objects, known as inducers.
    [Show full text]
  • 17-2021 CAMI Pilot Vision Brochure
    Visual Scanning with regular eye examinations and post surgically with phoria results. A pilot who has such a condition could progress considered for medical certification through special issuance with Some images used from The Federal Aviation Administration. monofocal lenses when they meet vision standards without to seeing double (tropia) should they be exposed to hypoxia or a satisfactory adaption period, complete evaluation by an eye Helicopter Flying Handbook. Oklahoma City, Ok: US Department The probability of spotting a potential collision threat complications. Multifocal lenses require a brief waiting certain medications. specialist, satisfactory visual acuity corrected to 20/20 or better by of Transportation; 2012; 13-1. Publication FAA-H-8083. Available increases with the time spent looking outside, but certain period. The visual effects of cataracts can be successfully lenses of no greater power than ±3.5 diopters spherical equivalent, at: https://www.faa.gov/regulations_policies/handbooks_manuals/ techniques may be used to increase the effectiveness of treated with a 90% improvement in visual function for most One prism diopter of hyperphoria, six prism diopters of and by passing an FAA medical flight test (MFT). aviation/helicopter_flying_handbook/. Accessed September 28, 2017. the scan time. Effective scanning is accomplished with a patients. Regardless of vision correction to 20/20, cataracts esophoria, and six prism diopters of exophoria represent series of short, regularly-spaced eye movements that bring pose a significant risk to flight safety. FAA phoria (deviation of the eye) standards that may not be A Word about Contact Lenses successive areas of the sky into the central visual field. Each exceeded.
    [Show full text]
  • Explore Your Blind Spot Discover How the Mind Hides Its
    Explore your blind spot Discover how the mind hides its tracks by Tom Stafford Smashwords Edition (version 1.4, 2 July 2015) Copyright 2011 Tom Stafford This work is licensed under a Creative Commons Attribution-NonCommercial- ShareAlike 3.0 Unported License. Thank you for downloading this free eBook. You are welcome to share it with your friends. This book may be reproduced, copied and distributed for non-commercial purposes. You can even modify it, as long as the modified version is covered by the same licence. http://creativecommons.org/ Tom Stafford lives on the internet at http://idiolect.org.uk Follow him on twitter: @tomstafford Other ebooks by Tom: For argument’s sake: evidence that reason can change minds (2015) Control Your Dreams (2011) The Narrative Escape (2010) Your guide is Tom Stafford: This is a picture of the back of my eye, you can see the blood vessels and the optic disc, the light circle where they converge. It is this disc which produces the blind spots in our vision. You will need: pen, paper, eyes Your journey will take: minutes Category: perception Trig points: 3 The Treasure: Proud of your sharp sight? Perhaps you should think again. For each eye, there is a blind spot, an area near the middle of your vision for which you cannot see anything. Normally the way your vision works hides these blind spots from your awareness, but it isn't hard to show they're there. Visual blind spots are a good example of how our conscious experience is fundamentally based on our biological machinery.
    [Show full text]
  • Preference for Concentric Orientations in the Mouse Superior Colliculus
    ARTICLE Received 12 Jan 2015 | Accepted 25 Feb 2015 | Published 2 Apr 2015 DOI: 10.1038/ncomms7773 OPEN Preference for concentric orientations in the mouse superior colliculus Mehran Ahmadlou1 & J Alexander Heimel1 The superior colliculus is a layered structure important for body- and gaze-orienting responses. Its superficial layer is, next to the lateral geniculate nucleus, the second major target of retinal ganglion axons and is retinotopically organized. Here we show that in the mouse there is also a precise organization of orientation preference. In columns perpendicular to the tectal surface, neurons respond to the same visual location and prefer gratings of the same orientation. Calcium imaging and extracellular recording revealed that the preferred grating varies with retinotopic location, and is oriented parallel to the concentric circle around the centre of vision through the receptive field. This implies that not all orientations are equally represented across the visual field. This makes the superior colliculus different from visual cortex and unsuitable for translation-invariant object recognition and suggests that visual stimuli might have different behavioural consequences depending on their retinotopic location. 1 Netherlands Institute for Neuroscience, an institute of the Royal Academy of Arts and Sciences, Cortical Structure & Function group, Meibergdreef 47, 1105 BA Amsterdam, The Netherlands. Correspondence and requests for materials should be addressed to J.A.H. (email: [email protected]). NATURE COMMUNICATIONS | 6:6773 | DOI: 10.1038/ncomms7773 | www.nature.com/naturecommunications 1 & 2015 Macmillan Publishers Limited. All rights reserved. ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms7773 ells in the primary visual cortex (V1) respond optimally to single and multi-units recorded at different depths preferred the edges or lines of a particular orientation.
    [Show full text]
  • Attention and Blind-Spot Phenomenology
    Attention and Blind-Spot Phenomenology Liang Lou & Jing Chen Department of Psychology Grand Valley State University Allendale, Michigan 49401 U.S.A. [email protected] [email protected] Copyright (c) Liang Lou & Jing Chen 2003 PSYCHE, 9(02), January 2003 http://psyche.cs.monash.edu.au/v9/psyche-9-02-lou.html KEYWORDS: Attention, blind spot, filling-in, consciousness, visual perception, phenomenology. ABSTRACT: The reliability of visual filling-in at the blind spot and how it is influenced by the distribution of spatial attention in and around the blind spot were studied. Our data suggest that visual filling-in at the blind spot is 1) less reliable than it has been assumed, and 2) easier under diffused attention around the blind spot than under focal attention restricted in the blind spot. These findings put important constraints on understanding the filling-in in terms of its neural substantiation. Recent neurophysiological studies suggest that V1 neurons corresponding to the blind spot in retinotopic map extend their receptive fields far beyond the blind spot and are not silent during the filling-in (Komatsu, Kinoshita, and Murakami, 2000). For those neurons to subserve filling-in, it may be crucially important for top-down attention to match their receptive fields. 1. Introduction The visual blind spot is formed at the back of each eye in an area called optic disk, which is essentially a hole in the retina through which the axons of ganglion cells bundle to exit the eye to form the optic nerve. It is "blind" because no photoreceptors exist there for receiving information from the world.
    [Show full text]