16.899A: Physiology (contd) Lavanya Sharan January 24th, 2011 Before we start, a few caveats • A lot is not known about how the human visual system works. • We (Alyosha + Lavanya) don’t know a lot about physiology. • But, before you worry, a few lines from Marr… Slide source: Nancy Kanwisher & Jim DiCarlo We care about ‘big picture’ • In this class, we are interested in the underlying software/algorithm/computations • Not in specifics of the `particular hardware’ • Want back pocket models for various components of the human visual system – Very few of these exist. • Our closest cousins: computational neuroscientists/cognitive scientists/psychophysicists Overview of `particular hardware’ • • • • • • • Retina, LGN What are visual areas? Tools for studying human visual system Area V1 Beyond area V1 What and where pathways Summary of what (and how little) we know Primary Visual Pathway 1. Retina 2. Thalamus – Lateral Geniculate Nucleus (LGN) • divided into magno and parvo layers 3. Primary visual cortex (V1) 4. Extrastriate visual areas • Each visual hemifield projects to the opposite hemisphere Slide source: Jody Culham 7 Slide source: Nancy Kanwisher & Jim DiCarlo Primary Visual Pathway 1. Retina 2. Thalamus – Lateral Geniculate Nucleus (LGN) • divided into magno and parvo layers 3. Primary visual cortex (V1) 4. Extrastriate visual areas • Each visual hemifield projects to the opposite hemisphere Slide source: Jody Culham What is a Visual Area? 1. Function – an area has a unique pattern of responses to different stimuli 2. Architecture – different brain areas show differences between cortical properties (e.g., thickness of different layers, sensitivity to various dyes) 3. Connectivity – Different areas have different patterns of connections with other areas 4. Topography – many sensory areas show topography (retinotopy, somatotopy, tonotopy) – boundaries between topographic maps can indicate boundaries between areas (e.g., separate maps of visual space in visual areas V1 and V2 Slide source: Jody Culham Why are there so many visual areas? MAGNO • quick and dirty PARVO • slow and detailed Slide source: Jody Culham Source: Felleman & Van Essen, 1991 Source: Mapping the MInd cover image More brain, more visual areas Slide source: Jody Culham Why not a really big visual area? • As areas become larger, longer interconnections are required • Limits on cortical thickness and connections may constrain max area size Slide source: Jody Culham Parallel processing is more efficient Teach neural network to identify “what” and “where” One neural network with 18 nodes (~neurons) devoted to both tasks versus One neural networks with two streams of 9 nodes each (total = 18) After 300 training trials, the two stream model outperformed the single-system model Slide source: Jody Culham Rueckl, Cave & Kosslyn, 1989 Different Tasks Require Different Information • different regions may need to use different coding systems dorsal stream: viewer-centred ventral stream: object-centred Slide source: Jody Culham Wiring Constraints Source: Van Essen, 1997 David Van Essen proposes that as the brain develops, areas that are richly interconnected will be pulled together to form a gyrus (and those that are weakly interconnected form sulci). Slide source: Jody Culham Optimized Connections Multidimensional Scaling • strength of connections can be used to infer spatial layout • expected layout of visual areas matches anatomy amazingly well Parietal Occipital Temporal Malcolm Young Slide source: Jody Culham Tools for mapping human areas • Neuropsychological Lesions • Temporary Disruption • transcranial magnetic stimulation (TMS) • Electrical and magnetic signals • electroencephalography (EEG) • magnetoencephalography (MEG) • Brain Imaging • positron emission tomography (PET) • functional magnetic resonance imaging (fMRI) Slide source: Jody Culham Slide source: Nancy Kanwisher & Jim DiCarlo Overview of `particular hardware’ • • • • • • • Retina, LGN What are visual areas? Tools for studying human visual system Area V1 Beyond area V1 What and where pathways Summary of what (and how little) we know Cortical Receptive Fields Single-cell recording from visual cortex David Hubel & Thorston Wiesel © Stephen E. Palmer, 2002 Cortical Receptive Fields Single-cell recording from visual cortex Time © Stephen E. Palmer, 2002 Cortical Receptive Fields Three classes of cells in V1 Simple cells Complex cells Hypercomplex cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Simple Cells: “Line Detectors” B. Dark Line Detector Firing Rate Horizontal Position © Stephen E. Palmer, 2002 Cortical Receptive Fields Simple Cells: “Edge Detectors” C. Dark-to-light Edge Detector Firing Rate D. Light-to-dark Edge Detector Firing Rate Horizontal Position Horizontal Position © Stephen E. Palmer, 2002 Cortical Receptive Fields Constructing a line detector Retina Receptive Fields LGN CenterSurround Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Complex Cells STIMULUS NEURAL RESPONSE 00o Time © Stephen E. Palmer, 2002 Cortical Receptive Fields Complex Cells STIMULUS NEURAL RESPONSE o 60 0 Time © Stephen E. Palmer, 2002 Cortical Receptive Fields Complex Cells STIMULUS NEURAL RESPONSE o 90 0 Time © Stephen E. Palmer, 2002 Cortical Receptive Fields Complex Cells STIMULUS NEURAL RESPONSE o 120 0 Time © Stephen E. Palmer, 2002 Cortical Receptive Fields Constructing a Complex Cell Retina Receptive Fields Cortical Area V1 Simple Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Hypercomplex Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Hypercomplex Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Hypercomplex Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Hypercomplex Cells “End-stopped” Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields “End-stopped” Simple Cells © Stephen E. Palmer, 2002 Cortical Receptive Fields Constructing a Hypercomplex Cell RETINA Receptive Fields CORTICAL AREA V1 Complex Cell End-stopped Cell © Stephen E. Palmer, 2002 Overview of `particular hardware’ • • • • • • • Retina, LGN What are visual areas? Tools for studying human visual system Area V1 Beyond area V1 What and where pathways Summary of what (and how little) we know Overview of visual areas Logothetis 1999; from http://psychology.uwo.ca/fMRI4Newbies/RetinotopicandEarlyVisualAreas.html Macaque & human visual areas are similar Tootell et al. 2003; from http://psychology.uwo.ca/fMRI4Newbies/RetinotopicandEarlyVisualAreas.html Slide source: Nancy Kanwisher & Jim DiCarlo Retinotopy (Tootell et al. 1982) Adjacent parts of visual field are mapped to adjacent parts of cortex. Not all visual areas have retinotopy, may be graded. Slide source: Nancy Kanwisher & Jim DiCarlo Slide source: Nancy Kanwisher, Jim DiCarlo, David Heeger Slide source: Nancy Kanwisher, Jim DiCarlo, David Heeger Two visual pathways The two visual processing streams for different visual percepts: “What” (ventral stream)- object recognition • main input from “slow and detailed” parvo system “Where” or “How” (dorsal stream) - spatial perception, motor planning • main input from “quick and dirty” magno system Slide source: Jody Culham Source: Mishkin & Ungerleider, 1982 Two visual pathways The two visual processing streams for different visual percepts: “What” (ventral stream)- object recognition • main input from “slow and detailed” parvo system “Where” or “How” (dorsal stream) - spatial perception, motor planning • main input from “quick and dirty” magno system Slide source: Jody Culham Source: Mishkin & Ungerleider, 1982 The “What” Pathway Other Visual Areas • contain more complex receptive fields Temporal Lobe • contains many specialized areas for recognizing various things body motion faces places bodies objects Slide source: Jody Culham The “Where” or “How” Pathway grasping and reaching attention head movements Parietal Lobe • contains many specialized areas for using vision to guide actions in space motion perception eye movements Slide source: Jody Culham Slide source: Nancy Kanwisher & Jim DiCarlo Summary • Low-level areas Filter banks, SIFT, HOG for color, orientation, spatial frequencies, motion… • High-level areas Desired output from computer vision systems e.g., segmentation, robust object/scene/texture recognition, motion understanding and planning… • Middle-level area Where the magic happens – No one (neuroscientists, psychologists, computer scientists, etc.) really understands this stage of processing. more, come find us for pointers to papers/books/readings and • For people to talk to.