Neuroscience: Exploring the Brain, 3e Chapter 9: The Eye Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Introduction • Significance of vision – Relationship between human eye & camera – Retina • Photoreceptors: Converts light energy into neural activity • Detects differences in intensity of light – Lateral geniculate nucleus (LGN) • First synaptic relay in the primary visual pathway • Visual information ascends to cortex interpreted and remembered Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Properties of Light • Light – Electromagnetic radiation – Wavelength, frequency, amplitude Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Properties of Light • Light – Energy is proportional to frequency – e.g., gamma radiation and cool colors - high energy – e.g., radio waves and hot colors - low energy Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Properties of Light • Optics – Study of light rays and their interactions • Reflection • Bouncing of light rays off a surface • Absorption • Transfer of light energy to a particle or surface • Refraction • Bending of light rays from one medium to another Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins The Structure of the Eye • Gross Anatomy of the Eye – Pupil: Opening where light enters the eye – Sclera: White of the eye – Iris: Gives color to eyes – Cornea: Glassy transparent external surface of the eye – Optic nerve: Bundle of axons from the retina Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins The Structure of the Eye • Ophthalmoscopic Appearance of the Eye Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins The Structure of the Eye • Cross-Sectional Anatomy of the Eye Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Image Formation by the Eye • Refraction of light by the cornea – Eye collects light, focuses on retina, forms images Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Image Formation by the Eye • Accommodation by the Lens – Changing shape of lens allows extra focusing power Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Image Formation by the Eye • The Pupillary Light Reflex – Connections between retina and brain stem neurons that control muscle around pupil – Continuously adjusting to different ambient light levels – Consensual – Pupil similar to the aperture of a camera Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Image Formation by the Eye • The Visual Field – Amount of space viewed by the retina when the eye is fixated straight ahead Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Image Formation by the Eye • Visual Acuity – Ability to distinguish two nearby points – Visual Angle: Distances across the retina described in degrees Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Microscopic Anatomy of the Retina • Direct (vertical) pathway: – Ganglion cells – Bipolar cells – Photoreceptors Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Microscopic Anatomy of the Retina • Retinal processing also influenced lateral connections: – Horizontal cells • Receive input from photoreceptors and project to other photoreceptors and bipolar cells – Amacrine cells • Receive input from bipolar cells and project to ganglion cells, bipolar cells, and other amacrine cells Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Microscopic Anatomy of the Retina • The Laminar Organization – Inside-out – Light passes through ganglion and bipolar cells before reaching photoreceptors Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Microscopic Anatomy of the Retina • Photoreceptor Structure – Converts electromagnetic radiation to neural signals – Four main regions • Outer segment • Inner segment • Cell body • Synaptic terminal – Types of photoreceptors • Rods and cones Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Microscopic Anatomy of the Retina • Regional Differences in Retinal Structure – Varies from fovea to retinal periphery – Peripheral retina • Higher ratio of rods to cones • Higher ratio of photoreceptors to ganglion cells • More sensitive to light Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Microscopic Anatomy of the Retina • Regional Differences in Retinal Structure (Cont’d) – Cross-section of fovea: Pit in retina where outer layers are pushed aside • Maximizes visual acuity – Central fovea: All cones (no rods) • 1:1 ratio with ganglion cells • Area of highest visual acuity Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Phototransduction • Phototransduction in Rods – Light energy interacts with photopigment to produce a change in membrane potential – Analogous to activity at G-protein coupled neurotransmitter receptor - but causes a decrease in second messenger Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Phototransduction • Phototransduction in Rods – Dark current: Rod outer segments are depolarized in the dark because of steady influx of Na+ – Photoreceptors hyperpolarize in response to light Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Phototransduction • Phototransduction in Rods – Light activated biochemical cascade in a photoreceptor – The consequence of this biochemical cascade is signal amplification Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Phototransduction • Phototransduction in Cones – Similar to rod phototransduction – Different opsins • Red, green, blue • Color detection – Contributions of blue, green, and red cones to retinal signal – Spectral sensitivity – Young-Helmholtz trichromacy theory of color vision Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Phototransduction • Dark and Light Adaptation 20–25 minutes All-cone daytime vision All-rod nighttime vision – Dark adaptation—factors • Dilation of pupils • Regeneration of unbleached rhodopsin • Adjustment of functional circuitry Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Phototransduction • Dark and Light Adaptation – Calcium’s Role in Light Adaptation • Calcium concentration changes in photoreceptors • Indirectly regulates levels of cGMP channels Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Processing • Transformations in the Outer Plexiform Layer – Photoreceptors release less neurotransmitter when stimulated by light – Influence horizontal cells and bipolar cells Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Processing • Receptive Field: “On” and “Off” Bipolar Cells – Receptive field: Stimulation in a small part of the visual field changes a cell’s membrane potential – Antagonistic center-surround receptive fields Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Processing • On-center Bipolar Cell – Light on (less glutamate); Light off -> more glutamate ‘Inverting’ synapse (inhib) Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Output • Ganglion Cell Receptive Fields – On-Center and Off-Center ganglion cells – Responsive to differences in illumination Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Output • Types of Ganglion Cells – Appearance, connectivity, and electrophysiological properties • M-type (Magno) and P-type (Parvo)ganglion cells in monkey and human retina Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Output • Color-Opponent Ganglion Cells Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Output • Parallel Processing – Simultaneous input from two eyes • Information from compared in cortex • Depth and the distance of object – Information about light and dark: ON-center and OFF-center ganglion cells – Different receptive fields and response properties of retinal ganglion cells: M- and P- cells, and nonMnonP cells Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Concluding Remarks • Light emitted by or reflected off objects in space imaged onto the retina • Transduction – Light energy converted into membrane potentials – Phototransduction parallels olfactory transduction • Electrical-to-chemical-electrical signal • Mapping of visual space onto retina cells not uniform Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins End of Presentation Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins Retinal Processing • Research in ganglion cell output by – Keffer Hartline, Stephen Kuffler, and Horace Barlow – Only ganglion cells produce action potentials • Research in how ganglion cell properties are generated by synaptic interactions in the retina – John Dowling and Frank Werblin – Other retinal neurons produce graded changes in membrane potential Copyright © 2007 Wolters Kluwer Health | Lippincott Williams & Wilkins