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POWERPOINT PRESENTATION FOR BIOPSYCHOLOGY, 9TH EDITION BY JOHN P.J. PINEL P R E PA R E D B Y J E F F R E Y W. G R I M M WESTERN WASHINGTON UNIVERSITY COPYRIGHT © 2014 PEARSON EDUCATION, INC. ALL RIGHTS RESERVED. This multimedia product and its contents are protected under copyright law. The following are prohibited by law: • any public performance or display, including transmission of any image over a network; • preparation of any derivative work, including the extraction, in whole or in part, of any images; • any rental, lease, or lending of the program. Chapter 6 The Visual System How We See Copyright © 2014 Pearson Education, Inc. All rights reserved. Learning Objectives LO1: Describe the human eyes and how their properties influence vision. LO2: Describe the structure of the human retina, and explain how the structure of the retina influences vision. LO3: The retina-geniculate-striate system is retinotopic, with signals from each retina arriving in both ipsilateral and contralateral cortices. Explain with diagrams. LO4: Explain the Mach Band demonstration and the important point that it makes about vision. LO5: Describe Hubel and Wiesel’s method, and summarize their major findings. LO6: Explain color constancy and its important implication. LO7: Describe and discuss the dorsal and ventral streams and their relation to prosopagnosia and akinetopsia. Copyright © 2014 Pearson Education, Inc. All rights reserved. What Do We See? Somehow a distorted and upside-down 2D retinal image is transformed into the 3-D world we perceive. Two types of research are needed to study vision. Research probing the components of the visual system Research assessing what we see Copyright © 2014 Pearson Education, Inc. All rights reserved. Light Enters the Eye and Reaches the Retina No species can see in the dark, but some are capable of seeing when there is little light. Light can be thought of as: Particles of energy (photons) Waves of electromagnetic radiation Humans see light between 380–760 nanometers. Wavelength: perception of color Intensity: perception of brightness Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.2 The electromagnetic spectrum and the colors associated with wavelengths visible to humans. Copyright © 2014 Pearson Education, Inc. All rights reserved. The Pupil and the Lens Light enters the eye through the pupil, whose size changes in response to changes in illumination. Sensitivity: the ability to see when light is dim Acuity: the ability to see details Copyright © 2014 Pearson Education, Inc. All rights reserved. The Pupil and the Lens (Con’t) Lens: focuses light on the retina Ciliary muscles alter the shape of the lens as needed. Accommodation: the process of adjusting the lens to bring images into focus Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.4 The human eye, a product of 600 million years of evolution (Lamb, Collin, & Pugh, 2007). Copyright © 2014 Pearson Education, Inc. All rights reserved. Eye Position and Binocular Disparity Convergence: eyes must turn slightly inward when objects are close Binocular disparity: the difference between the images on the two retinas Both are greater when objects are close— together, they provide the brain with a 3-D image and distance information. Copyright © 2014 Pearson Education, Inc. All rights reserved. The Retina and Translation of Light into Neural Signals The retina is, in a sense, inside-out. Light passes through several cell layers before reaching its receptors. Vertical pathway: receptors > bipolar cells > retinal ganglion cells Lateral Communication Horizontal cells Amacrine cells Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.5 The cellular structure of the mammalian retina. Copyright © 2014 Pearson Education, Inc. All rights reserved. The Retina and Translation of Light into Neural Signals (Con’t) Blind spot: no receptors where information exits the eye The visual system uses information from cells around the blind spot for “completion,” filling in the blind spot. Fovea: high-acuity area at center of retina Thinning of the ganglion cell layer reduces distortion due to cells between the pupil and the retina. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.6 A section of the retina. The fovea is the indentation at the center of the retina; it is specialized for high-acuity vision. Copyright © 2014 Pearson Education, Inc. All rights reserved. Cone and Rod Vision Duplexity theory of vision: cones and rod mediate different kinds of vision. Cones: photopic (daytime) vision High-acuity color information in good lighting Rods: scotopic (nighttime) vision High-sensitivity, allowing for low-acuity vision in dim light, but lacks detail and color information There is more convergence in the rod system, increasing sensitivity while decreasing acuity. Only cones are found at the fovea. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.8 A schematic representation of the convergence of cones and rods on retinal ganglion cells. There is a low degree of convergence in cone-fed pathways and a high degree of convergence in rod-fed pathways. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.9 The distribution of cones and rods over the human retina. The figure illustrates the number of cones and rods per square millimeter as a function of distance from the center of the fovea. (Based on Lindsay & Norman, 1977.) Copyright © 2014 Pearson Education, Inc. All rights reserved. Spectral Sensitivity Lights of the same intensity but different wavelengths may not all look equally bright. A spectral sensitivity curve shows the relationship between wavelength and brightness. There are different spectral sensitivity curves for photopic (cone) vision and scotopic (rod) vision. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.10 Human photopic (cone) and scotopic (rod) spectral sensitivity curves. The peak of each curve has been arbitrarily set at 100%. Copyright © 2014 Pearson Education, Inc. All rights reserved. Eye Movement We continually scan the world with small and quick eye movements: saccades. These bits of information are then integrated. Stabilize retinal image; see nothing. The visual system responds to change. Copyright © 2014 Pearson Education, Inc. All rights reserved. Visual Transduction: The Conversion of Light to Neural Signals Transduction: conversion of one form of energy to another Visual transduction: conversion of light to neural signals by visual receptors Pigments absorb light. Absorption spectrum describes spectral sensitivity. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.11 The adsorption spectrum of rhodopsin compared with the human scotopic spectral sensitivity curve. Copyright © 2014 Pearson Education, Inc. All rights reserved. Visual Transduction: The Conversion of Light to Neural Signals (Con’t) Rhodopsin is the pigment found in rods. A G-protein-linked receptor that responds to light rather than to neurotransmitters. In the Dark Na+ channels remain partially open (partial depolarization), releasing glutamate. When Light Strikes Na+ channels close. Rods hyperpolarize, inhibiting glutamate release. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.12 The inhibitory response of rods to light. When light bleaches rhodopsin molecules, the rods’ sodium channels close; as a result, the rods become hyperpolarized and release less glutamate. Copyright © 2014 Pearson Education, Inc. All rights reserved. From Retina to Primary Visual Cortex The retinal-geniculate-striate pathways include about 90 percent of axons of retinal ganglion cells. The left hemiretina of each eye (right visual field) connects to the right lateral geniculate nucleus (LGN); the right hemiretina (left visual field) connects to the left LGN. Most LGN neurons that project to primary visual cortex (V1, striate cortex) terminate in the lower part of cortical layer IV. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.13 The retina-geniculatestriate system: the neural projections from the retinas through the lateral geniculate nuclei to the left and right primary visual cortex (striate cortex). The colors indicate the flow of information from various parts of the receptive fields of each eye to various parts of the visual system. (Based on Netter, 1962.) Copyright © 2014 Pearson Education, Inc. All rights reserved. Retinotopic Organization Information received at adjacent portions of the retina remains adjacent in the striate cortex (retinotopic). More cortex is devoted to areas of high acuity—like the disproportionate representation of sensitive body parts in somatosensory cortex. About 25 percent of primary visual cortex is dedicated to input from the fovea. Copyright © 2014 Pearson Education, Inc. All rights reserved. The M and P Channels Magnocellular Layers (M Layers) Big cell bodies; bottom two layers of LGN Particularly responsive to movement Input primarily from rods Parvocellular Layers (P Layers) Small cell bodies; top four layers of LGN Color, detail, and still or slow objects Input primarily from cones Copyright © 2014 Pearson Education, Inc. All rights reserved. The M and P Channels (Con’t) The channels project to slightly different areas in lower layer IV in striate cortex; M neurons are just above the P neurons. The channels project to different parts of visual cortex beyond V1. Copyright © 2014 Pearson Education, Inc. All rights reserved. Seeing Edges Contrast Enhancement Mach bands: nonexistent stripes the visual system creates for contrast enhancement Makes edges easier to see A consequence of lateral inhibition Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.14 The illusory bands visible in this figure are often called Mach bands, although Mach used a different figure to generate them in his studies (see Eagleman, 2001). Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.15 How lateral inhibition produces contrast enhancement. (Based on Ratliff, 1972.) Copyright © 2014 Pearson Education, Inc. All rights reserved. Receptive Fields of Visual Neurons The Area of the Visual Field within which It Is Possible for a Visual Stimulus to Influence the Firing of a Given Neuron Hubel and Wiesel looked at receptive fields in the retinal ganglion, LGN, and lower layer IV of striate cortex of a cat. Copyright © 2014 Pearson Education, Inc. All rights reserved. Receptive Fields: Neurons of the Retina-Geniculate-Striate System Similarities seen at all three levels: Receptive fields of foveal areas are smaller than those in the periphery. Neurons’ receptive fields are circular in shape. Neurons are monocular. Many neurons at each level had receptive fields with excitatory and inhibitory area. Copyright © 2014 Pearson Education, Inc. All rights reserved. Receptive Fields: Neurons of the Retina-Geniculate-Striate System (Con’t) Many cells have receptive fields with a center-surround organization: excitatory and inhibitory regions separated by a circular boundary. Some cells are on-center and some are offcenter. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.16 The receptive fields of an oncenter cell and an off-center cell. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.17 The responses of an oncenter cell to contrast. Copyright © 2014 Pearson Education, Inc. All rights reserved. Receptive Fields: Simple and Complex Cortical Cells In lower layer IV of the striate cortex, neurons with circular receptive fields (as in retinal ganglion cells and LGN) are rare. Most neurons in V1 are either: Simple—receptive fields are rectangular with “on” and “off” regions—or Complex—also rectangular, with larger receptive fields, and respond best to a particular stimulus anywhere in their receptive fields Copyright © 2014 Pearson Education, Inc. All rights reserved. Receptive Fields: Simple and Complex Cortical Cells (Con’t) SIMPLE Rectangular “On” and “off” regions, like cells in layer IV Orientation and location sensitive All are monocular. COMPLEX Rectangular Larger receptive fields Do not have static “on” and “off” regions Not location sensitive Motion sensitive Many are binocular. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.18 Examples of visual fields of simple cortical cells. Copyright © 2014 Pearson Education, Inc. All rights reserved. Columnar Organization of Primary Visual Cortex Cells with simpler receptive fields send information on to cells with more complex receptive fields. Functional vertical columns exist such that all cells in a column have the same receptive field and ocular dominance. Ocular dominance columns: as you move horizontally, the dominance of the columns changes. Retinotopic organization is maintained. Copyright © 2014 Pearson Education, Inc. All rights reserved. Contextual Influences in Visual Processing Plasticity appears to be a fundamental property of visual cortex function. E.g., receptive field properties depend on the scene in which the stimuli to its field are embedded. Copyright © 2014 Pearson Education, Inc. All rights reserved. Seeing Color: Component and Opponent Processing Component Theory (Trichromatic Theory) Proposed by Young, refined by Helmholtz Three types of receptors, each with a different spectral sensitivity Copyright © 2014 Pearson Education, Inc. All rights reserved. Seeing Color: Component and Opponent Processing (Con’t) Opponent-process theory was proposed by Hering. Two different classes of cells encoding color, and another class encoding brightness Each encodes two complementary color perceptions. This theory accounts for color afterimages and colors that cannot appear together (reddish green or bluish yellow). Copyright © 2014 Pearson Education, Inc. All rights reserved. Seeing Color: Component and Opponent Processing (Con’t) Both theories are correct: coding of color by cones seems to operate on a purely component basis; opponent processing of color is seen at all subsequent levels. Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.19 The absorption spectra of the three classes of cones. Copyright © 2014 Pearson Education, Inc. All rights reserved. Color Constancy and the Retinex Theory Color constancy: color perception is not altered by varying reflected wavelengths. Retinex theory (Land): color is determined by the proportion of light of different wavelengths that a surface reflects. Relative wavelengths are constant, so perception is constant. Dual-opponent color cells are sensitive to color contrast. Found in cortical “blobs” Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.20 The method of Land’s (1977) color-vision experiments. Subjects viewed Mondrians illuminated by various proportions of three different wavelengths: a short wavelength, a middle wavelength, and a long wavelength. Copyright © 2014 Pearson Education, Inc. All rights reserved. Cortical Mechanisms of Vision and Conscious Awareness Flow of Visual Information Thalamic relay neurons, to 1˚ visual cortex (striate), to 2˚ visual cortex (prestriate), to Visual association cortex As visual information flows through hierarchy, receptive fields: Become larger Respond to more complex and specific stimuli Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.21 The visual areas of the human cerebral cortex. Copyright © 2014 Pearson Education, Inc. All rights reserved. Damage to Primary Visual Cortex Scotomas Areas of blindness in contralateral visual field due to damage to primary visual cortex Detected by perimetry test Completion Patients may be unaware of scotoma; missing details are supplied by “completion.” Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.23 The completion of a migraine-induced scotoma as described by Karl Lashley (1941). Copyright © 2014 Pearson Education, Inc. All rights reserved. Damage to Primary Visual Cortex (Con’t) Blindsight Response to visual stimuli outside conscious awareness of “seeing” Possible explanations of blindsight Islands of functional cells within scotoma Direct connections between subcortical structures and secondary visual cortex; not available to conscious awareness Copyright © 2014 Pearson Education, Inc. All rights reserved. Functional Areas of Secondary and Association Visual Cortex Neurons in each area respond to different visual cues, such as color, movement, or shape. Lesions of each area results in specific deficits. Anatomically distinct: about 12 functionally distinct areas have been identified so far. Retinotopically Organized Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.24 Some of the visual areas that have been identified in the human brain. Copyright © 2014 Pearson Education, Inc. All rights reserved. Dorsal and Ventral Streams Dorsal stream: pathway from primary visual cortex to dorsal prestriate cortex to posterior parietal cortex The “where” pathway (location and movement), or Pathway for the control of behavior (e.g., reaching) Ventral stream: pathway from primary visual cortex to ventral prestriate cortex to inferotemporal cortex The “what” pathway (color and shape), or Pathway for the conscious perception of objects Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.26 The “where” versus “what” and the “control of behavior” versus “conscious perception” theories make different predictions. Copyright © 2014 Pearson Education, Inc. All rights reserved. Prosopagnosia Inability to Distinguish among Faces Most prosopagnosics’ recognition deficits are not limited to faces. Prosopagnosia is associated with damage to the ventral stream between the occipital and temporal lobes. Prosopagnosics may be able to recognize faces in the absence of conscious awareness. Prosopagnosics have different skin conductance responses to familiar faces compared to unfamiliar faces, even though they reported not recognizing any of the faces. Copyright © 2014 Pearson Education, Inc. All rights reserved. Akinetopsia Deficiency in the Ability to See Movement Progress in a Normal, Smooth Fashion Can Be Induced by a High Dose of Certain Antidepressants Associated with Damage to the Middle Temporal (MT) Area of the Cortex Copyright © 2014 Pearson Education, Inc. All rights reserved. FIGURE 6.27 The location of the fusiform face area and the MT area: Damage to the fusiform face area is associated with prosopagnosia, damage to the MT area is associated with akinetopsia. Copyright © 2014 Pearson Education, Inc. All rights reserved.