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Vision: From Eye to Brain (Chap 3, Part B)

Lecture 7

Jonathan Pillow Sensation & Perception (PSY 345 / NEU 325) Princeton University, Spring 2015

1 more “channels”: spatial channels spatial frequency: the number of cycles of a per unit of visual angle (usually specified in degrees) • think of it as: # of bars per unit length

low frequency intermediate high frequency

2 Why sine ? • Provide useful decomposition of images

Technical term: Fourier decomposition

3 Fourier decomposition • mathematical decomposition of an image (or sound) into sine waves. reconstruction: “image” 1

2 sine waves

3 sine waves

4 sine waves

4 “Fourier Decomposition” theory of V1 claim: role of V1 is to do “Fourier decomposition”, i.e., break images down into a sum of sine waves

• Summation of two spatial sine waves • any pattern can be broken down into a sum of sine waves

5 Fourier decomposition • mathematical decomposition of an image (or sound) into sine waves.

Original image High Low Frequencies

6 original

low medium high

7 Retinal Ganglion Cells: tuned to spatial frequency

Response of a ganglion cell to sine gratings of different frequencies

8 The contrast sensitivity function

Human contrast sensitivity illustration of this sensitivity

9 Image Illustrating Spatial Frequency Channels

10 Image Illustrating Spatial Frequency Channels

11 If it is hard to tell who this famous person is, try squinting or defocusing

“Lincoln illusion” Harmon & Jules 1973

12 “Gala Contemplating the Mediterranean Sea, which at 30 meters becomes the portrait of Abraham Lincoln (Homage to Rothko)”

- Salvador Dali (1976)

13 “Gala Contemplating the Mediterranean Sea, which at 30 meters becomes the portrait of Abraham Lincoln (Homage to Rothko)”

- Salvador Dali (1976)

14 lateral geniculate nucleus (LGN): one on each side of the brain • this is where axons of retinal ganglion cells synapse

Organization: • represents contralateral visual field • segregated into eye- specific layers • segregated into M and P layers

Ipsilateral: Referring to the same side of the body Contralateral: Referring to the opposite side of the body

15 Primary Visual Cortex

• Striate cortex: known as primary visual cortex, or V1 • “Primary visual cortex” = first place in cortex where visual information is processed

(Previous two stages: retina and LGN are pre-cortical)

16 Receptive Fields: monocular vs. binocular

- LGN cells: responds to one eye or the other, never both

LGN

V1 - V1 cells: can respond to input from both eyes • By the time information gets to V1, inputs from both eyes have been combined (but V1 neurons still tend to have a preferred eye - they spike more to input from one eye)

17 Topography: mapping of objects in space onto the visual cortex

• contralateral representation - each visual field (L/R) represented in opposite hemisphere

• cortical magnification - unequal representation of fovea vs. periphery in cortex - a misnomer, because “magnification” already present in retina

(that is, the amount of space in cortex for each part of the visual field is given by the number of fibers coming in from LGN)

18 Acuity in V1

Visual acuity declines in an orderly fashion with eccentricity—distance from the fovea (in deg)

19 V1 receptive fields: elongated regions of space

Major change in representation: • Circular receptive fields (retina & LGN) replaced by elongated “stripe” receptive fields in cortex • Has ~ 200 million cells! • (vs. 1 million Retinal Ganglion Cells)

20 Orientation tuning: • neurons in V1 respond more to bars of certain orientations • response rate falls off with difference from preferred orientation “preferred orientation” “preferred

21 Receptive Fields in V1

Many cortical cells respond especially well to: • Moving lines • Bars • Edges • Gratings • Direction of motion

Ocular dominance: • Cells in V1 tend to have a “preferred eye” (i.e., respond better to inputs from one eye than the other)

22 Simple vs. Complex Cells

23 Receptive Fields in V1

Cells in V1 respond best to bars of light rather than to spots of light • “simple” cells: prefer bars of light, or prefer bars of dark • “complex” cells: respond to both bars of light and dark

[Hubel & Weisel movie]

24 Column: a vertical arrangement of neurons

• ocular dominance • orientation column: column: for particular for a particular location in location in cortex, neurons cortex, neurons have same have same preferred eye preferred orientation

25 Hypercolumn - contains all possible columns

• Hypercolumn: 1-mm block of V1 containing “all the machinery necessary to look after everything the visual cortex is responsible for, in a certain small part of the visual world” (Hubel, 1982

• Each hypercolumn contains a full set of columns - has cells responding to every possible orientation, and inputs from left right eyes

26 web demos receptive fields http://sites.sinauer.com/wolfe4e/wa03.04.html columns http://sites.sinauer.com/wolfe4e/wa03.05.html

27 Adaptation

28 Adaptation: the Psychologist’s Electrode

“tilt after-effect”

29 Adaptation: the Psychologist’s Electrode

“tilt after-effect”

• perceptual illusion of tilt, provided by adapting to a pattern of a given orientation

• supports idea that the human contains individual neurons selective for different orientations

30 Adaptation: the Psychologist’s Electrode

Adaptation: the diminishing response of a sense organ to a sustained stimulus

• An important method for deactivating groups of neurons without surgery

• Allows selective temporary “knock out” of group of neurons by activating them strongly

31 Effects of adaptation on population response and perception

Before Adaptation 0 degree stimulus

unadapted population resp to 0 deg

Stimulus presented =

32 Effects of adaptation on population response and perception

Then adapt to 20º Before Adaptation

unadapted population resp to 0 deg

Stimulus presented =

33 Selective adaptation alters neural responses and perception

After Adaptation perceptual effect of adaptation is repulsion away from the adapter

Stimulus presented =

34 Selective Adaptation: The Psychologist’s Electrode

Selective adaptation for spatial frequency: Evidence that human visual system contains neurons selective for spatial frequency

35 Adaptation that is specific to spatial frequency (SF)

1. adapt 2. test

3. percept

36 Adaptation that is specific to spatial frequency (SF)

1. adapt 2. test

3. percept

37 Adaptation that is specific to spatial frequency (SF)

1. adapt 2. test

3. percept

38 Adaptation that is specific to spatial frequency AND orientation

1. adapt 2. test

3. No adaptive percept

39 Adaptation that is specific to spatial frequency AND orientation

1. adapt 2. test

3. No adaptive percept

40 Adaptation that is specific to spatial frequency AND orientation

1. adapt 2. test

3. No adaptive percept

41 Selective Adaptation: The Psychologist’s Electrode

Orthodox viewpoint: • If you can observe a particular type of adaptive after-effect, there is a certain neuron in the brain that is selective (or tuned) for that property

THUS (for example):

There are no neurons tuned for spatial frequency across all orientations, because adaptation is orientation specific.

42 Selective Adaptation: The Psychologist’s Electrode

adapting spatial freq

contrast sensitivity after threshold increases near width of “channels” adaptation to a sine the adapted frequency that contribute to wave with a frequency contrast sensitivity of 7 cycles/degree.

43 Selective Adaptation: The Psychologist’s Electrode

adapting spatial freq

Therefore: • adaptation reveals separate channels devoted to orientation and spatial frequencies • width of adaptive effect reveals the width of the channel

44 The Development of Spatial Vision • how can you study the vision of infants who can’t yet speak?

1. preferential-looking paradigm - infants prefer to look at more complex stimuli

45 The Development of Spatial Vision • how can you study the vision of infants who can’t yet speak?

2. visually evoked potentials (VEP) - measure brain’s electrical activity directly

46 The Development of Spatial Vision

young children: not very sensitive to high spatial frequencies

• Visual system is still developing ! Cones and rods are still developing and taking final shape ! Retinal ganglion cells are still migrating and growing connections with the fovea ! The fovea itself has not fully developed until about 4 years of age

47 Summary • early visual pathway: retina -> LGN -> V1 • “contralateral” representations in visual pathway • (vs. sensitivity) • spatial frequency channels • • spatial frequency sensitivity & tuning • V1 receptive fields, orientation tuning • Hubel & Weisel experiments • simple vs. complex cells • cortical magnification • cortical columns, hypercolumns • adaptation

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