Vision in vertebrates How do we see ? Aims To describe how sensory systems fulfil their need to transduce energy compare inputs lead to appropriate behaviour using the vertebrate visual system Main parts of visual system eye lens, retina brain - primates lateral geniculate nucleus visual cortex brain - other vertebrates superior colliculus / optic tectum Warning Not all vertebrates work the same way differences in anatomy differences in retinal & CNS function even between closely related species, e.g. frog and toad always look and see what species the work was done on! Visual systems frog - optic tectum man Lens light is a wave 3 * 108 m/s focus light onto the retina maps places in outside world to places on retina in 1:1 fashion Retina structure photoreceptors - at back of retina layers of cells output from ganglion cells - at front of retina Blindspot Structure of retina retina at back of eye ganglion cells photoreceptors Structure of retina Electron - micrograph Structure of retina Diagram light Photoreceptors Rods and Cones Transduction Can we perceive photons? Colour vision Rods and Cones rods 100 * more sensitive most cones at fovea rod density highest around fovea Therefore, turn eye to see in different places Photoreceptor rhodopsin in membrane discs inside outer segment membrane voltage determined by cell membrane light Transduction in the dark, channels are open Transduction Transduction movie Transduction Sensitivity increased by gain in enzymes gain in channel gain at synapse vesicle ribbon increases number of vesicles released this reduces quantal noise at synapse Physiological recording suction pipette records inward current in outer segment bright dim Can we perceive photons? Macaque rods able to detect individual photons people can see light flashes when 1 in 100 rods will get a photon with 0.2s Colour vision most common form is red-green deficiency Colour vision Diurnal animals & birds have colour vision Humans and OldWorld monkeys are tri-chromatic most monkeys dichromatic May have evolved to detect when fruit is ripe Colour vision Diurnal animals/birds have colour vision Humans and Old-World monkeys are trichromatic most monkeys dichromatic May have evolved to detect when fruit is ripe other mechanisms of color vision exist oil droplets in amphibians, turtles gene homology Colour vision gene duplication on X chromosome Summary so far At retina, world is spatially mapped light level is encoded by current color is used (but not in all animals) very sensitive Retina is Layered Diagram light light Physiology Dowling - Necturus (mudpuppy) light Physiology receptor is inhibited by light Sign conserving /reversing synapses horizontal cells mediate lateral inhibition light Physiology ganglion cells signal to brain difference in light between adjacent receptors amacrine cells signal on or off Not light level On-Off responses ganglion cells usually respond to changes in light results from lateral inhibition Lateral inhibition Hermann Grid common to vision, touch, hearing... Summary so far At retina, world is spatially mapped light level is encoded by current color is used (but not in all animals) very sensitive At ganglion cells on/off & surround /center not a 1:1 relation between light level and signal this enhances dynamic range Blindspot axons of ganglion cells run over surface and turn to give optic nerve Ganglion cells project to LGN Lateral geniculate nucleus Mapping of cells visual field mapped spatially different ganglion cells project to different layers LGN sensitive to lines ganglion cells respond to spots LGN to lines different line orientations for each LGN cell LGN projects to Visual cortex visual = striate cortex = V1 Orientation selectivity in cortex LGN orientation is maintained, with a pinwheel pattern Ocular dominance LGN kept data from the eyes separate in visual cortex, data converges. Some cells have dominant input from R, some from L cells in same column have same dominance V1 Cortex orientation and ocular dominance work together grey is contralateral eye Depth perception Use both eyes to calculate how far away objects are Hypothesis 1: rangefinder Hypothesis 2: measure overlap of images Disparity... 1) rotate eyes 2) compare signals from different parts of the retina 2 wins out Depth perception Muller - Lyer Ponzo which lines are longer ? Hering Blindsight loss of visual cortex may show evidence for blindsight patient cannot “see” but can follow targets with their eyes patient can discriminate words projection to superior colliculus may be responsible Deconstruction of signal Visual system does not froward a direct “photographic copy” Reconstruction ? Kanizsa illusion temporal lobe Summary transduction well understood; high gain system retina well understood; lateral inhibitory mechanism LGN and V1 cortex fairly well understood; lateral & temporal inhibition; binocular vision further processing still to be elucidated