1. Slide 1 Vision is Remarkable! Extremely complicated and costly process As a reward over the course of evolution, vision provided New ways of communication Ability to predict the trajectory of objects and events in time and space New froms of mental imagery and abstraction World of visual art Eyes like a camera can adjust to differences in illumination and focus itself on objects of interest Additional functions such as the ability to track moving objects and self-cleaning system Retina like film (but much more than that) Photoreceptors: Converts light energy into neural activity Output does not faithfully reproduce the intensity of the light falling on it Detects differences in intensity of light falling on different parts of it Image processing on the retina

2. Slide 2 Properties of Light Light  Electromagnetic radiation is all around us! radio, wireless phones, x-ray machines, sun  Light is the electromagnetic radiation that is visible to our eyes! (400-700 nm)  Energy content is proportional to frequency Hot colors: Orange, red : lower energy Cool colors: blue, violet: higher energy Colors are themselves “colored” by the brain

3. Slide 3 Properties of Light Optics  Light rays travel in straight lines until they interact with atoms and molecules in the medium Reflection =Bouncing of light rays off a surface =Most of what we see is reflected light Absorption =Transfer of light energy to a particle or surface =Basis for color perception - reflected light off a surface is absorbed in the retina Refraction =Bending of light rays from one medium to another - toward a line that is perpendicular to the surface =Due to the speed (of light travel) difference - the greater the difference, the greater the angle of refraction =Basis for image forming on the retina

4. Slide 4 The Structure of the Eye Gross Anatomy of the Eye  Pupil: Opening where light enters the eye  Iris: Control the amount of light coming into eye (aperture) The sphincter muscle lies around the very edge of the pupil. In bright light, the sphincter contracts, causing the pupil to constrict. The dilator muscle runs radially through the iris, like spokes on a wheel. This muscle dilates the eye in dim lighting. Of which pigmentation determines the eye color  Sclera: White of the eye provides tough wall of eyeball Extraocular muscles (3pairs) are embedded and control the movement of eyeball

5. Slide 5 The Structure of the Eye Gross Anatomy of the Eye  Eye’s orbit: bony socket of skull  Conjunctiva: membrane connecting sclera with eyelids  Cornea: Glassy transparent external surface of the eye (lens-like regractive power) Innervated by unmyelinated nerve endings : very sensitive to pressure (touch)  Optic nerve: Bundle of axons from the retina

6. Slide 6 The Structure of the Eye Ophthalmoscopic Appearance of the Eye  Blood vessels on the surface of Retina  Optic disk : A pale circular region Gate for entering blood vessels and Exiting optic nerve fibers Blind spot =No photoreceptors present =Brain is deceiving you!  Macula (spot): Central vision Relative absence of blood vessels - improves the quality of central vision  Fovea (pit): A thinner center of macula The center of retina - serves as a anatomical reference point =Nasal vs temporal =Superior vs inferior

7. Slide 7 The Structure of the Eye Ophthalmoscopic Appearance of the Eye  Blood vessels on the surface of Retina  Optic disk : A pale circular region Gate for entering blood vessels and Exiting optic nerve fibers Blind spot =No photoreceptors present =Brain is deceiving you!  Macula (spot): Central vision Relative absence of blood vessels - improves the quality of central vision  Fovea (pit): A thinner center of macula The center of retina - serves as a anatomical reference point =Nasal vs temporal =Superior vs inferior

8. Slide 8 The Structure of the Eye Cross-Sectional Anatomy of the Eye  Aqueous Humor fluid filling space between cornea and lens supply nourishment  Ciliary muscles Ligaments (zonule fibers) that suspend lens are attached Connect to sclera  Lens: Change shape to adjust focus Aqueous humor in anterior chamber Jelly-like vitreous humor in posterior chamber = its pressure serves to keep the spherical shape of eyeball

9. Slide 9 Image Formation by the Eye Eye collects light, focuses on retina, forms images Refraction of light by the cornea  Parallel lights from far distance must be bent by refraction  Air to aqueous humor change makes refraction on the surface of cornea  Focal distance depends on the curvature of cornea  Refractive power - reverse of Focal distance (diopter) Cornea has 42 diopters Depends on the slowing of light (Blurry vision underwater)

10. Slide 10 Image Formation by the Eye Accommodation by the Lens  Changing shape of lens allows for extra focusing power (~12 diopters)  Important for focusing images of objects within 9m ranges - requires greater refraction for diverging (not parallel) rays  Accommodations by ciliary muscle contraction tension of zonule fubers Decrease lens becomes rounder increased curvature lose this function with age (presbyopia)

11. Slide 11 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 : bilateral reflex  Pupil similar to the aperture of a camera increase the depth of focus by the constriction of pupil same as increasing the f-stop

12. Slide 12 Image Formation by the Eye The Visual Field  Amount of space viewed by the retina when the eye is fixated straight ahead Visual Acuity  Ability to distinguish two nearby points : depends on several factors including the spacing of photoreceptors in the retina and the precision of eye’s refraction  Visual Angle: Distances across the retina described in degrees

13. Slide 13 Microscopic Anatomy of the Retina Photoreceptors:  Cells that convert light energy into neural activity Direct (vertical) pathway:  Horizontal cells, Amacrine cells : modify the responses of bipolar cells and ganglion cells via lateral connections Ganglion cells  Output from the retina Photoreceptorsbipolar cells ganglion cells

14. Slide 14 Microscopic Anatomy of the Retina The Laminar Organization of the Retina  Cells organized in layers  Inside-out : Upper cells are relatively transparent Pigmented epithelium is critical to maintain photoreceptors and photopigments

15. Slide 15 Microscopic Anatomy of the Retina Photoreceptor Structure  Electromagnetic radiation to neural signals  Four main regions Outer segment =Stack of membraneous disks that contain photopigments =Lights are absorbed by photopigments and lead to changes in membrane potential Inner segment Cell body Synaptic terminal  Types of photoreceptors Rods =have more disks and higher photopigments concentration - 1000 times more sensitive to light than cones =Scotopic retina Cones detect colors =Photopic retina

16. Slide 16 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 - lower acuity More sensitive to light (rods are specialized for low light)  Rods outnumber cones in the human retina (20 to 1)

18. Slide 18 Microscopic Anatomy of the Retina Regional Differences in Retinal Structure  Cross-section of fovea: Pit in retina due to lateral displacement of the cells above the photoreceptors Maximizes visual acuity by allowing light to strike photoreceptors directly (no scattering)  Central fovea: All cones (no rods) 1:1 ratio with ganglion cells Area of high visual acuity

19. Slide 19 Phototransduction Phototransduction in Rods  Depolarization in the dark: “Dark current” -30 mV Due to steady influx of Na+ through cGMP gated sodium channel Constant production of cGMP by guanylyl cyclase  Hyperpolarization in the light Light reduces cGMP to close Na+ channel - hyperpolarize transducin

20. Slide 20 Phototransduction Phototransduction in Rods  Depolarization in the dark: “Dark current” to -30 mV Due to steady influx of Na+ through cGMP gated sodium channel Constant production of cGMP by guanylyl cyclase  Hyperpolarization in the light Light reduces cGMP to close Na+ channel - hyperpolarize PDE

21. Slide 21  Rhodopsin Photopigment that absorb electromagnetic radiation Receptor protein that is embedded in the membrane of the stacked disks in the rod outer segments Receptor protein with a prebound chemical agonist [Opsin (GPCR) + retinal (vitamin A derivative)] Bleaching : change in conformation of retinal by light absorption leading to the activation of opsin (by dissociation) Phototransduction 11-cis-retinal all-trans-retinal

22. Slide 22  Light - retinal - opsin - transducin - PDE - cGMP - cGMP gated Na + channel  Signal amplification : very low number of photons can be detected Phototransduction

23. Slide 23 Phototransduction Phototransduction in Cones  Similar to rod phototransduction Rods are hyperpolarized constantly - saturated Daytime vision depends on cones, whose photopigments require more energy to become bleached  Different opsins : major difference Red, green, blue cones  Color detection Contributions of blue, green, and red cones to retinal signal Young-Helmholtz trichromacy theory of color vision = Brain assigns colors based on a comparison of the readout of the three color types =Color blindness - significant spectral abnormality (beyond normal variation), mostly due to genetic errors = Abnormal red-green vision is the most common abnormality that is more frequently found in men = Peak sensitivity of rods is to a wavelength of 500 nm (blue- green)

24. Slide 24 Phototransduction Dark and Light Adaptation  Dark adaptation— Increasing the sensitivity to light (10 6 fold) Dilation of pupils - 2-8 mm diameter; 16 fold Regeneration of unbleached rhodopsin Adjustment of functional circuitry - signals from more rods are available to each ganglion cells All-cone daytime visionAll-rod nighttime vision 20–25 minutes

25. Slide 25 Phototransduction Dark and Light Adaptation  Calcium’s Role in Light Adaptation Blinding sensation reflects saturation of both rods and cones (hyperpolarization) Cones gradually adapt their membrane potential to - 35 mV cGMP-gated Na+ channel also allow Ca++ that inhibit guanylyl cyclase (balancing GC in the dark) Under the light, low Ca++ concentration in the hyperpolarized cones gradually activates GC, recovering cGMP - open the Na+ channel again Ca++ also affect photopigments and PDE to reset their responses to light What we see is the relative difference in light level, not the absolute level!

26. Slide 26 Retinal Processing Only ganglion cells fire APs!  All other cells respond with graded changes in membrane potential : difficult to detect! Transformations in the Outer Plexiform Layer  Photoreceptors form synapses with bipolar cells and horizontal cells  Output of photoreceptors is generated by dark rather than light  Dark is the preferred stimulus When a shadow passes across a photoreceptor, it responds by depolarizing and releasing neurotransmitter glutamate

27. Slide 27 Retinal Processing Bipolar Cell Receptive Fields  Two classes of bipolar cells OFF bipolar cells : depolarized through glutamate gated Na+ channel - active when photoreceptors are depolarized (dark) ON bipolar cells : GPCR respond to glutamate to hyperpolarize - active (depolarized) when there’s less glutamate (light)  Receptive Field Area of retina that changes the cell’s mem- brane potential upon Light stimulation Consists of center and surround Antagonistic center- surround receptive fields ; complex interaction of horizontal cells, photoreceptors, and bipolar cells

28. Slide 28 Light on surround On center becomes off

29. Slide 29 Retinal Output Ganglion Cell Receptive Fields  On-Center and Off-Center cells (corresponding connections with bipolar cells)  Center-surround cancellation  Responsive to differences in illumination within their receptive fields

30. Slide 30 Retinal Output Ganglion Cell Receptive Fields  The center-surround organization of the receptive fields leads to a neural response that emphasizes the contrast at light-dark edges  Illusion of the perception of light level can be explained by the level of inhibition by surround

31. Slide 31 Retinal Output Types of Ganglion Cells  Two types of ganglion cells in monkey and human retina M-type (Magno, 5%), P-type (Parvo, 90%), nonM-nonP (5%)

32. Slide 32 Retinal Output Color-Opponent Ganglion Cells  Some P cells and nonM-nonP cells  Response to one wavelength in the receptive field center is canceled by another wavelength in the receptive field surround  Red versus green and blue versus yellow White light will equally activate both center and surround to cancel each other Ganglion cells output  A stream of information concerning three different comparisons : Light vs dark, red vs green, blue vs yellow

33. Slide 33 Retinal Output Parallel Processing - independent but simultaneous information processing  Simultaneous input from two eyes Information from two streams is compared in the central visual system to give the perception of ‘Depth’  Information about light and dark ON-center and OFF-center ganglion cells provide independent streams of information  Different receptive fields and response properties of retinal ganglion cells M cells : sensitive to subtle contrasts over large receptive field and contribute to low-res vision P- cells : small receptive field contribute to high-res vision (detail) nonM-nonP cells : color opponency