1. Perceptual Systems

2. Readings, Class 1: Wolfe Ch 1,2 Kandell et al Chapters 26 Sources: Wolfe, J, Kluender, K, Levi, D. et al Sensation & Perception 2012 3 rd ed Sinauer – 15% discount and free shipping if ordering online from Sinauer Kandel, Schwartz & Jessel Principles of Neural Science McGraw-Hill 5 th ed Gazzaniga, Ivry, Mangun Cognitive Neuroscience Norton, 3 rd ed

3. Visual Perception: what do we want to explain? How do we get visual information from the world and use it to control behavior? Traditional sub-areas - visual sensitivity color vision spatial vision temporal vision binocular vision/ depth perception texture perception motion perception surfaces, segmentation object perception attention perceptual learning spatial orientation eye movements

4. The constructive nature of perception: a process of guessing the state of the world from sometimes incomplete sensory data. Constructive in the sense that it relies on memory representations of past experience

5. Major transformations of the light signal in the retina: 1.Temporal filtering – visual response slower than input signal. 2. Spatial filtering – local signals are combined across space to varying degrees. 3. Light adaptation – retina modifies responsiveness depending on average light level. 4.Color coding – trichromacy and color opponency

6. The Eye and Retina Pigment epithelium reduces scatter Important vegetative functions iris pupil Note – blind spot - cf damage to peripheral retina Visualize retinal blood vessels.

7. 18mm 0.3mm = 1 deg visual angle xa tan(a/2) = x/d a = 2 tan - 1 x/d Visual Angle d 1 diopter = 1/focal length in meters 55 diopters = 1/.018 Most of the optical power of the eye is accomplished by the cornea

8. Optical correction errors Presbyopia = stiffening of lens with age so it is no longer variable Blur circle

9. Figure 2.9 Photoreceptor density across the retina Note: color vision in peripheral retina Note: peripheral cones are fatter. Rods similar to foveal cones

10. Visual Acuity matches photoreceptor density Relative visual acuity Receptor density 1 foveal cone= 0.5 min arc

11. Two of the factors limiting visual acuity are - optics of the eye - size and spacing of photoreceptors -(in central fovea, a cone is about 0.5 min arc) -Grating versus vernier acuity: Snellen (letter chart versus threading a needle)

12. Sine wave gratings Acuity is the highest frequency pattern that is just visible – ie the narrowest stripes A similar measure is made by the Snellen letter chart: E Vernier acuity is an order of magnitude better than grating acuity. How can this be?

13. Figure 2.9 Photoreceptor density across the retina Question: Rods are small and dense. Why isn’t acuity better in the peripheral retina?

14. Transduction: light into electrical signals “dark light” Note sluggish response

16. Major transformations of the light signal in the retina: 1.Temporal filtering – visual response slower than input signal. photoreceptor response is slow – increases sensitivity 2. Spatial filtering – local signals are combined across space to varying degrees. Acuity for fine patterns determined by optics and photoreceptor layout. 3. Light adaptation – retina modifies responsiveness depending on average light level. 4.Color coding – trichromacy and color opponency

17. Probability of absorption of a photon depends on wavelength (but receptor doesn’t know what wavelength it absorbed)

18. Why blue flowers are brighter and red flowers are darker at dusk. Peak night day Note: peak sensitivity in day about the same wavelength as maximum output of sun.

19. Convergence: many rods converge onto a single rod bipolar cell, and several cones converge onto a diffuse bipolar cell. This allows the signal to be amplified. M= magnocellular, P= Parvocellular Midget system preserves acuity in the central fovea

20. Horizontal and amacrine cells form inhibitory surrounds of ganglion cells. Why ON and OFF cells?

21. Hecht, Schlaer, & Pirenne, 1942 A single quantum is sufficient to excite a rod photoreceptor. A few quanta within a small area is sufficient to give a sensation of light. Measure number of quanta for a just detectable sensation of light – about 100 quanta. Of those 100 quanta, about 90 are lost on the way to the retina form scatter in the eye. So 10 quanta incident on the retina lead to a sensation of light. Light has a Poisson distribution, so the probability that more than one photon falls on a single rod is very small. Therefore, a single photon must excite a rod, and 10 photons excite a retinal ganglion cell. This signal is transmitted to the brain with minimal loss and generates a sensation of light.

22. Center-surround organization of bipolar and ganglion cells Center-surround organization means that responses to uniform lights are reduced Light spot excites cell Dark spot excites cell Biggest response to a spot in center

23. Figure 3.6 Sine wave gratings illustrating low (a), medium (b), and high (c) spatial frequencies These grating stimuli are called “Gabor patches”. Spatial frequency is measured in Cycles per degree, and contrast is a measure of the difference in intensity between light and dark bars.

24. Figure 3.7 The contrast sensitivity function (red line): the window of visibility

26. Perceptual consequences of center surround antagonism Brightness is coded by the differences in illumination between adjoining regions This results from center-surround organization.

27. Perceptual consequences of center surround antagonism Brightness is coded by the differences in illumination between adjoining regions

28. Major transformations of the light signal in the retina: 1.Temporal filtering – reduced response to high temporal frequencies – Temporal integration – a strong 1 msec flash is equivalent to a weaker 50 msec flash. 2. Spatial filtering: - Anatomical organization of photoreceptors provides high acuity in fovea with rapid fall-off in the periphery. (photoreceptor density) -Convergence of photoreceptors onto ganglion cells also leads to acuity limitations in the peripheral retina. (1 cone per midget cell in fovea) - Center-surround antagonism reduces sensitivity to uniform fields. 3. Light adaptation 4.Color coding

29. Light adaptation: the problem Need to respond over a range of 10 10 – but ganglion cells can only signal 0-200 spikes/sec Receptor adaptation Response on different background intensities tvi curve ΔI/I = 1 Ganglion cells change sensitivity as well as photoreceptors. Perceptual consequence of light adaptation: hard to tell ambient light intensity

30. Loss of sensitivity at low temporal frequencies (slow rate of change of intensity) is a consequence of light adaptation (sensitivity changes with average light level) (afterimage fading)

31. Figure 2.17 Dark adaptation curve Sensitivity recovers when the retina is in the dark, rapidly for cones, slowly for rods. (afterimages)

32. Major transformations of the light signal in the retina: 1.Temporal filtering – reduced response to high temporal frequencies – Temporal integration – a strong 1 msec flash is equivalent to a weaker 50 msec flash. 2. Spatial filtering: - Anatomical organization of photoreceptors provides high acuity in fovea with rapid fall-off in the periphery. (photoreceptor density) -Convergence of photoreceptors onto ganglion cells also leads to acuity limitations in the peripheral retina. (1 cone per midget cell in fovea) - Center-surround antagonism reduces sensitivity to uniform fields. 3. Light adaptation – sensitivity regulation - adjustment of operating range to mean light level. (Light level 10 10 range, ganglion cells, 10 2 range.) 4.Color opponency. Organization of 3 cone photoreceptors into color opponent signals (Luminance, Red-Green, Yellow-Blue)

34. Retinotopic Organization and Cortical Magnification The brain uses more physical space for signals from the fovea than the periphery Adjacent points in the world Project to adjacent points in cortex

35. Signals from each eye are adjacent in LGN but remain segregated in different layers. Convergence occurs in V1. Two kinds of cells in retina project to different layers in LGN M=magno=big P=parvo=small K= konio

36. Magno and parvo cells have different spatial and temporal sensitivities. Function of the different M and P pathways is unclear. Note: attempts to Isolate a pathway psychophysically were unsuccessful

37. Figure 2.17 Dark adaptation curve

38. Cone Photoreceptors are densely packed in the central fovea Note: despite lower density of cones in peripheral retina, color vision is basically the same across the visual field.

39. Figure 2.11 Blue, green, and red represent the S-, M-, and L- cones, respectively, of a living human being in a patch of retina at 1 degree from the fovea Two of the factors limiting visual acuity are – optics of the eye -size and spacing of photoreceptors -(in central fovea, a cone is about 0.5 min arc)