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perception
a process in which the brain builds a model of the world, using both previous experience and signals from the sensory organs
awareness of objects and events -> meaning
visual perception
an experience-based interpretation of retinal images
primary visual pathway
retina --> thalamus (Lateral Geniculate Nucleus, LGN) --> cortex (vision, visual perception, object recognition, etc.)
ALSO: retina --> superior colliculus (eye movements)
primary visual cortex, AKA...
V1
striate cortex
Brodmann's area
the image in the eye
reduced and reversed
upside down and right to left
orderly, point to point
retinal image
output: retinal ganglion cells (sampling of image)
dense in center (small receptive fields)
poor in periphery (large receptive fields)
cortical magnification
expanded representation of the foveal (center) stimulus
a small area in central visual field is represented in a larger area of V1 than a large area in the peripheral vision
visual acuity depends on...
retinal location
fovea
highest density of receptors (cones)
lowest convergence (1:1)
highest cortical magnification
highest acuity
periphery
lower density of receptors (rods, cones)
high convergence (many:1)
low cortical magnification
acuity is lower
individual variance
people with higher foveal magnification have higher acuity
functional consequence of more tissue --> higher computational power --> better cortical processing --> better function
retinotopy
points of the visual world that are represented next to one another on the retina are represented next to one another in the visual cortex
orderly representation of the visual world in the cortex
receptive fields of adjacent neurons are systematically shifted in space
oversampling
each point in visual space is covered by thousands of receptive fields
microelectrode recording
Records the activity of a single cell
retinotopy
oversampling
metabolic markers
metabolic radio-isotope marker (C14, 2D glucose)
stimulation
active neurons consume more energy and accumulate more marker
pattern is revealed after processing tissue
drawback: postmortem, one per animal
optical imaging/fMRI
in a living animal, mapping representation of several features possible
measures changes of levels of the oxygenated hemoglobin in brain regions
after onset of grating motion:
- visual cortical neurons fire within milliseconds
- vascular response about 2 seconds later (BOLD negative, cerebral blood flow volume positive)
does not directly measure activity of neurons, but BOLD, due to neurons' activity
fMRI: stimuli
to study representation of the visual space on the cortex: alternate S1/S2, calculate the difference of responses
complementary stimuli in space activate complementary regions of visual cortex
fMRI: results
systematic representation of the visual space on cortex
neighboring points in visual field are represented at neighboring cortical locations
cortical magnification
consequence of retinotopy
stimuli at specific retinal locations activate specific locations in the brain
lines of activity along visual pathway
scotomas
scotoma
regions of absent vision
damage to brain region is reflected as scotoma
- of a certain portion of the visual field
- of a specific shape
possibility to attribute damages along the visual pathway to specific scotoma
receptive field
region on the retina in which visual stimuli influence the neuron's firing rate
lateral geniculate nucleus (LGN)
A structure in the thalamus
stronger surround inhibition
modulation from other brain structures
visual cortex neurons
neurons in the visual cortex extract specific features of the object (orientation) which are then used by the brain in the process of perception
orientation selectivity
emerges in visual cortex
neurons respond to certain orientations but not others
brought about by cooperative action of multiple mechanisms
parallel to perception
End-stopping or hypercomplex receptive fields
receptive fields in the visual cortex as detectors of: edges and strips, oriented bars of specific length
simple cell response
responds to oriented bars at specific location
complex cell response
responds to oriented bars within a certain area
Hubel and Wiesel
studied feature detection in visual cortex and discovered simple, complex, and hypercomplex cells
columns in visual cortex
modular organization
receptive fields of vertically located neurons have similar
- location in visual field
- preferred orientation
- dominant eye
intrinsic signals
distribution of the levels of oxygenated hemoglobin (BOLD) over the cortex --> a correlate of activated regions
orientation map
from single-condition maps, at each location, calculate preferred orientation (strongest response)
ocular dominance map
pool single-condition maps for all orientations for the right and left eye, calculate preferred eye
orientation columns
different orientations are systematically represented in the visual cortex
neurons responding preferentially to same orientations
ocular dominance columns
neurons responding preferentially to the same eye
hypercolumn
complete set of orientations for both eyes
blobs
groups of neurons within V1 that are sensitive to color
interblobs
cells between blobs that are sensitive to orientation and not wavelength (motion and spatial structure)
How does representation of the visual world change from the retina to the cortex?
retina: image
visual cortex:
- representation
- feature maps (retinotopy, oculodominance columns, orientation columns)
selective rearing
experiment in which kittens were reared in an environment of vertical stripes to determine the effect on orientation selectivity of cortical neurons.
- feature detectors are formed in development
- perception depends on neurons
- perception is determined by experience (what we see is what we have learned)
- horizontal reared cats only see horizontal orientation
critical period
representational maps are formed during development
experience is necessary for normal development of the brain (including structure)
experience is necessary for normal perception
plasticity
perception can be trained (within genetic and developmental limits)
magnocellular system
~10%
large cells
large receptive fields
phasic responses
fast axons, 30-40 m/s
movement, contrast, depth
"colorblind"
(Y projection to V1)
dorsal stream, WHERE/HOW
parvocellular system
~80%
smaller cells
small receptive fields
tonic responses
slower axons, 15-23 m/s
texture, color, shape, depth
"motionblind"
(X projection to V1)
ventral stream, WHAT
heterogeneous cells
~10% remaining cells
gamma cells
small somata
large dendritic fields
large receptive fields
tonic/phasic
slow axons, 2-18 m/s
analysis of movement and direction of movement
projections: subcortical structures
function: unconscious reaction, visual reflects
contralateral eye
the eye on the opposite side of the body from the LGN
ipsilateral eye
The eye on the same side of the head as the structure to which the eye sends inputs.
LGN layers
- right/left eye
- parvo (3-6) and magno (1-2) cellular
- retinotopy
layers of visual cortex
1-6
fibers
small pyramids
stellate cells
large pyramids
white matter
inhibitory interneurons throughout
systematic projection of parvo and magno systems from LGN to cortex
L 2/3 --> other cortical areas
L 5 --> callosal, colliculus suerior, pulivinar, pons
l 6 --> LGN (thalamus)
discrimination task
which part of the brain is responsible for the processing?
ablation method
Ungerleider and Mishkin: WHAT and WHERE pathways
WHERE pathway
parietal lobe
landmark discrimination
where/how
dorsal pathway
magnocelllular
WHAT pathway
temporal lobe
object discrimination
what
ventral pathway
parvocellular
one function of V1
sorting information to channels and streams for further selective processing
which part of the brain is responsible for a specific function?
- animal research: ablation method
- observation of patients with localized brain damage (neuropsychology)
- psychophysical experiments: dissociated tasks
double dissociation
mechanisms are different and independent
how to find function with neuropsychology
patients with localized brain damage
tasks: matching orientation vs posting card through slot
- ventral pathway damaged, selective impairment of matching orientation (WHAT) but not of action (WHERE/HOW)
- dorsal pathway damaged, impairment of WHERE HOW
different aspects of visual processing are performed by different cortical areas
psychophysical experiments: dissociated tasks
adjust to match orientation of stimulus with shifted frame vs grasping
visual illusion does not affect grasping
different aspects of visual processing take place in different areas
agnosia
the inability to recognize familiar objects
color agnosia (A18)
color anomia (A18-37)
akinetopsia (mediotemporal bilateral)
object agnosia (A 18,20,21 left and c callosum)
prospagnosia (A 20, 21 bilateral)
representation of faces
neurons process sensory information by extracting more and more complex features and combinations
specific representations of biologically most relevant features
grandmother neuron (or at least highly specialized neuron detector)
distributed representation
weighted activity of a few neuronal groups
high capacity, high reliability
but template coding is faster
representation of objects
extraction of features (translating images into feature codes)
assembling these codes into a percept
- distributed
- partially overlapped
- combination of feature codes
binding problem
which features belong to one object?
possible solutions:
- grouping by timing
- high frequency oscillations
- different synchronized ensembles represent the respective percept
gamma oscillations
faster and more accurate recognition (of objects) correlated with the increased synchrony of EEG across visual and auditory areas
relevant for perception and multimodal integration
features
most common (space, orientation, color)
biologically more relevant (elements of faces)
specificity code
the idea that an object could be represented by the firing of a specialized neuron that responds only to that object
limited capacity, low reliability
what the retina tells the brain
distribution of contrast differences (in lumination and color)
what strategy does the brain use while assembling feature codes?
structuralism vs Gestalt psychology
Gestalt psychology
the whole is different than theh sum of the parts
interpretation is more than a sum of parts (link to top-down processing)
honor physics and avoid accidents
heuristic, mental shortcut
help to make fast decisions about most probably arrangements of objects which produced that image
structuralism
sensations add up to create perception
figure-ground segregation
The perceptual separation of an object from its background.
objects may overlap and their borders may be hidden
knowing the rules and objects help
color helps
neuronal responses as early as V1 can distinguish the figure from the background
viewpoint invariance
objects may be viewed from different perspectives and have different size but are still perceived the same
accidental viewpoint
a viewing position that produces some regularity in the visual image that is not present in the world
image may be ambiguous, allowing more than one interpretation
contrast borders (what retina tells brain) do not necessarily correspond to object borders
defining object borders
results of cortical processing (figure/background segregation) trump results of retinal processing (contrast borders)
how to study an illusion
- quantitative study of an illusion
- clear design of psychophysical experiments
- masking methods
quantitative study of illusions
forced choice paradigm - was shape fat or thin
with masking
variables:
- angle
- distance between elements
- presentation time
measure:
- probability of correct response
- detection threshold
1. illusory contours detected almost as good as real
limit of spatial
2. variable distance between elements - limit of spatial integration
3. variable presentation time
4. backward masking
masking
for perception of illusory contours, ~100 ms presentation is sufficient
interrupts afterimage
backwards masking
perception of illusory contours requires additional ~140-200 ms of processing time
interrupts afterimage and processing
grouping
knowing the occluding object helps
seeing the obstacle helps to recover a hidden pattern
mid-vision
- bring together what belongs together
- split asunder what does not belong together
- use what you know
- avoid accidents and ambiguity of interpretation
perceptual grouping, figure ground segregation, gestalt
Gestalt rules
grouping or organization is not in the stimulus
Law of Proximity
Law of Similarity
Law of Good Continuation
Law of Closure
Also: parallelism, symmetry, meaningfulness, common fate (motion)
formation of perceptual unit characterized by
- superadditivity (whole more than sum of parts)
- transposability (whole preserved despite large changes to parts)
not stimuli alone, but active mechanisms of their processing are involved in perception
figure
in front of the ground
owns the border
is more memorable
- lower area more likely perceived as figure, no difference right/left
- meaningful part more likely perceived as figure
brightness perception
dependent on context and experience
high-level vision
recognition: categorization or matching stimulus representation with representation in memory
recognition and brain activity
recognition
global before local
knowledge of what to expect enhances recognition of the context-appropriate details
global superiority effect
large-scale objects first, then details
the gist of a scene perceived first
contextual influence on recognition
scene, stimulus, mask, question
stimuli which are appropriate to the context are recognized better than those which do not belong to the context
recognition and brain activity
activation of face fusiform area when Harrison Ford shown
conservative conclusion: brain activity correlates with recognition
provocative conclusion: sufficiently high brain activity is necessary for correct recognition
delayed matching to sample experiment
is the test stimulus from same category as sample?
neurons in different brain regions accomplish different tasks
- inferotemporal neurons identification (perception)
- prefrontal neurons preserving information during the delay
is it possible to tell what a person is seeing by measuring brain activity?
voxel experiment - corresponding brain activity with images
72-92% of cases image was correctly identified
sensory stimuli produce unique patterns of brain activity, leading to specific perceptions
prediction: best fit from a pre-defined set
templates
fast
easy to categorize
help to segregate objects
but...
...for viewpoint invariance you need a lot of them
...one neuron one object code is not efficient
image description models
advanced template approach
- templates are created during experience
- more templates for more familiar objects
- viewpoint invariance does not always work, but for most familiar objects it does
experiment: learning letter like figures - no viewpoint invariance
recognition by components theory (RBC)
volumetric features
geons (geometric ions):
- non accidental properties, NAPs
- easily recognizable from most viewpoints (viewpoint invariant)
- simple formalization of description
36 geons - basic shapes
but...
...poor performance with line drawings
...letters
...classes of objects
perception is not always viewpoint invariant
experiments with letter like figures/greebles: poor recognition from new viewpoints (more errors with more deviation from learned viewpoint)
inverted faces, figure ground segregation of familiar contours
support for image description models
multiple committees
categorization:
- entry level category
- subordinate and superordinate categories
- different brain areas (several processes in parallel)
with training, entry level shifts down (previous subordinate becomes entry)
representations in ventral visual object pathway
- small number of category specific regions in addition to a more general purpose region that responds to any kind of objects
- genetically defined regions: face recognition
--- even if damaged early in life, no substitute
perceptual committees
a general scheme of hierarchical processing
feature demons --> cognitive demons --> decision demon
lateral inhibition in neurons and perception
space: antagonistic receptive fields
color: opponent mechanisms
orientation: cross-orientation inhibition
perception: one at a time interpretation of a retinal image
winner take all mechanism