MIT OpenCourseWare http://ocw.mit.edu 9.01 Introduction to NeuroscienceFall 2007 For information about citing these materials or our Terms of Use, visit: http://ocw.mit.edu/terms.Chapter 9 and 10 Review – Vision http://www.brown.edu/Courses/BN01/images/review/vision1.pdf http://www.brown.edu/Courses/BN01/images/review/vision2.pdf http://www.brown.edu/Courses/BN01/images/review/vision3.pdf General Properties of light (wavelength, color) Refraction • bending of light that occurs when light rays travel from one medium to another • from air to water, light rays bend toward a line that is perpendicular to the water surface The Big Picture Visual stimuli Æ Eye (photoreceptors, bipolar cells, ganglion cells) Æ optic nerve (chiasm, tract) Æ LGN (thalamus) Æ primary visual cortex Æ dorsal and ventral visual areas Eye • Anatomy: iris, aqueous humor, ciliary muscles, vitreous humor, sclera, optic nerve, retina, fovea (highest acuity, only photoreceptors, no blood vessels), cornea, lens, extraocular muscles, optic disk (blind spot, where optic nerve exists retina, where blood vessels originate), macula, pupil • Focusing an image on the retina requires: o Refraction by the cornea o Accommodation by the lens (important for focusing objects that are close)  Ciliary muscles contract (swell) Æ ligaments loosen Æ fatter lens, more refraction  Too fat a lens Æ myopia (nearsightedness), correct with concave eyeglass; too flat a lens Æ hyperopia (farsightedness), correct with convex eyeglass • Retinal organization o Photoreceptors – hyperpolarize to light; depolarize (release glutamate) when dark  Two types: rods and cones • fovea has cones, periphery has more rods • rods specialized for low light and are used at night • cones specialized for color, used during the day • higher photoreceptor-to-ganglion ratio in the periphery = less acuity  Transduction: photoreceptors have steady influx of sodium ions because cGMP keeps channels open; light activates opsin (or photopigment; called rhodopsin for rods); opsin activates G-protein which activates phosphodiesterase; phosphodiesterase breaks down cGMP, closing channels; cell hyperpolarizes o Bipolar cells (retinal processing)  Antagonistic center-surround receptive fields (important for contrast)  OFF bipolar (responds to glutamate by depolarizing; gated cation channels)  ON bipolar (responds to glutamate by hyperpolarizing; G-protein-coupled receptors) o Ganglion cells (retinal output)  Also center-surround; receive input from corresponding type of bipolar cell  Mainly responsive to differences in illumination  Types: P-type (small, 90%), M-type (large, 5%), nonM-nonP (5%)  Color opponency • Other topics: horizontal and amacrine cells; light and dark adaptation; pupillary light reflexOptic Nerve, Chiasm, Tract • At chiasm, axons from nasal retinas decussate; result: left visual field information carried by right optic tract; right visual field information carried by left optic tract • What parts of your vision is lost when: o Left optic nerve is cut? Input from left o Chiasm is cut down the middle? Peripheral vision o Left optic tract is cut? Right visual field • Targets of projection: LGN (thalamus) to striate cortex, hypothalamus, superior colliculus. Lateral Geniculate Nucleus • Six layers labeled 1 through 6 (most ventral layer is 1) • Retinal information separated by eye and ganglion cell type: o Ipsilateral axons synapse on LGN layers 2, 3, 5; contralateral axons on 1, 4, 6 o Magnocellular LGN layers (1 and 2) receive input from M-type ganglion cells; parvocellular LGN layers (3-6) receive input from P-type; and koniocellular layers (lie just ventral to each numbered layer) receive input from non-M-non-P • LGN neuron receptive fields almost identical to those of ganglion cells that innervate them o Magnocellular LGN neurons: large center-surround fields, insensitive to differences in wavelength, respond to stimulation with transient burst of action potentials o Parvocellular LGN neurons: small center-surround; many exhibit color opponency; sustained increase in frequency of action potentials o Koniocellular LGN neurons: center-surround; light/dark or color opponency Striate Cortex (V1, Area 17, primary visual cortex) • Six layers (I through VI; IV divided into A, B, and C; IVC into α and β) • Input o Most axons from LGN terminate in IVC – cell type and eye separation maintained in IVC  Magnocellular LGN neurons project to IVCα; parvocellular to IVCβ  Right, left eye inputs separated via ocular dominance columns (autoradiography) o Koniocellular LGN axons project to II and III blob regions • Processing o Radial connections (perpendicularly across all layers), horizontal connections (in one layer) o Blobs (seen with cytochrome oxidase) run along II, III, V, VI o IVC neurons project to II, III, IVB Æ some information integrated, processed; II and III receive binocular input • Output o II, III, IVB Æ cortical areas o V Æ superior colliculus, pons o VI Æ back to LGN • Receptive fields o Binocularity o Orientation selectivity o Direction selectivity o Simple and complex receptive fields o Blob receptive fields • Other: cortical modulesBeyond Striate Cortex • Dorsal stream (“where” pathway) – motion processing o Area MT (V5) – respond to movement in range of directions o Area MST – additional movement sensitivity (linear, radial, circular motion) • Ventral stream (“what” pathway) – object shape, color processing o Area V4 – shape and color perception (achromatopsia – loss of color vision) o Area IT – faces (prosopagnosia – difficulty recognizing faces) Coding• Retinotopy (2D surface of retina is mapped onto 2D surface of LGN, striate) o Mapping of visual field often distorted (greater representation of fovea) o Discrete point of light can activate many cells in the retina, more in target structures, because of overlapping receptive fields; activity in cortex is broad distribution with peak at specific retinotopic location o Not literal map; no pictures in the brain • Parallel processing o Left and right eyes o ON and OFF bipolar and ganglion cells o M- and P-type ganglion cells o Three channels in V1  Analysis of MOTION: magnocellular pathway = M-type ganglion cells Æ magnocellular layers of LGN Æ IVCα of cortex Æ IVB;