Title: The Cerebral Cortex and Memory
1The Cerebral Cortex and Memory STUDY QUESTIONS
What is the functional organization of the
cerebral cortex? Is the organization of the
cortex fixed during the course of development?
Is the organization of the cortex fixed in
adulthood?
2Cortical localization general principles a.
anterior -- motor processing. b. posterior fields
perceptual processing c. both the anterior and
posterior cortex involve processing
hierarchies. i. In the anterior cortex, the
primary motor the muscles of the body are mapped
out in a topographic organization, with adjacent
areas of cortex representing muscle groups in
adjacent areas of the body. ii. In the
posterior cortex there are distinct primary areas
for each sensory modality.
3- the posterior cortex
- distinct primary areas for each sensory modality.
- Receptive field
- a small circumscribed spatial region of the
sensory field within which cells respond to
stimulation. Cells often respond preferentially
to specific trigger features of the stimulus. - (2) The receptive fields and other trigger
features are organized in a topographic map of
the sensory field and of other relevant sensory - dimensions.
4- Primary visual cortex
- i. cells with these response properties are
organized topographically along two dimensions. - Ocular dominance
- preferences for activation by the ipsilateral
(same side of the head) or contralateral
(opposite side) eye. - (2) Orientation selectivity
- preferences for an optimal orientation of the
contrasting edge
5- Representation of these two dimensions
- Between layers, arranged in columns,
- through the depths of the cortical layers,
neurons have very similar properties - Within layers,
- Ipsilateral and contralateral ocular dominance
columns alternate, and - orientation columns are arranged in a systematic
sequence. - The combination of a full set of ocular dominance
and orientation columns that represent the same
small receptive field area is known as a
"hypercolumn." - Sets of such modules are organized systematically
to provide a full representation of the
contralateral visual field for each hemisphere.
6The tuning and modification of cortical
processing networks by experience (the classic
studies by David Hubel and Torsten Wiesel) i.
first observed in the primary visual cortex
associated with development in young animals. ii.
showed that response properties of primary visual
cortex neurons are plastic, that is, modifiable
by experience, during a "critical period" of
the first 4 weeks of life.
7- David Hubel and Torsten Wiesel
- Plasticity in ocular dominance
- closure of one eye ? a shift in ocular dominance
of all cells toward a preference for the active
eye. - Plasticity in orientation selectivity
- restricting exposure to stimuli with only certain
orientations of visual contrast ? a shift of all
cells toward selectivity for the trained
orientation. - Such manipulations are not nearly so effective
after this early critical period ends, - a finding that led most investigators to conclude
that cortical organization becomes fixed in
adulthood. - Mechanisms
- the reorganization of sensory maps, as well as
the normal initial organizing of sensory maps,
arises from a competition of activity among
different inputs to each cell.
8- Reorganization of the visual cortex in adulthood,
part 1 - plasticity of the adult cortex revealed by
creating small and selective damage to a part of
the normal inputs to the cortex (Gilbert) - Experimental manipulation
- a laser light ? very small lesion of the retina
- Results
- initially produces a correspondingly small area
of silent primary visual cortex. - Short period of time (minutes or hours) after
lesion - cells at the border of the deafferented area
become responsive to stimulation of intact visual
field areas. - Cortical cells with receptive fields located near
the boundary of the lesion expand in size - after a 2-month recovery period,
- the topography of the cortex is reorganized such
that the formerly disconnected zone becomes
responsive to neighboring parts of the visual
field
9- Reorganization of the visual cortex in adulthood,
part 2 - Cortical changes under normal sensory and
behavioural experience (Gilbert) - Experimental manipulation
- "artificial scotoma"
- Results
- After a 10-minute period of conditioning the
cell, its receptive field expanded severalfold in
length - c. The results suggested
- an ongoing process of modulation of receptive
field size, adapting in different ways to
different scenes. - normal sensory experience, not just peripheral
lesions, the changes take place on a brief time
scale. - e. Implications
- the short-term plasticity must involve in some
way a change in the synaptic weight of existing
connections, altering the patterns of activation
of intrinsic circuits.
10cortex
Retina or screen
Before lesion
B
A
B
A
After lesion
B
A
B
B
11- Rearrangements of the somatotopic representation
(Merzenich group ) on monkeys through a variety
of traumatic and more natural interventions in
somatosensory input. - digit removal ?
- the cortical representation of neighbouring
digits invades the cortical zone whose afferents
have been removed - Surgical joining of the digits ?
- the establishment of a continuous somatic
representation of formerly discontinuous zones
for each digit. - c. Tactile discrimination training (for several
weeks) ? - larger cortical representations of the
stimulated digits, and larger receptive fields in
the expanded areas. - cortical physiological activity acquired during
training is well correlated with the behavioural
performance in a discrimination task.
12How do higher areas of cortex respond to learning
- inferotemporal cortex (IT)? a. Two Visual
pathways and IT b. IT is the highest-order
cortical visual processing area, whose function
is the identification of objects by their visual
qualities-and this area is thought to be the site
of long-term storage of memory about visual
objects.
13evidence
- a. effects of damage to this area
- i. in humans, results in visual agnosia, a
selective deficit in visual object recognition - b. observed to be activated in various PET or
fMRI studies of neurologically intact individuals
performing tasks requiring visual object
recognition
14- c. Normal sensory response properties of IT cells
- i. 1st type
- responsive to whole objects positioned almost
anywhere' within the visual fields. - (2) respond similarly to a particular stimulus
regardless of its - size,
- contrast from the background
- form,
- location in the visual field
- motion.
15- The selectivity of IT cells is sometimes highly
specific. - the first explorations of IT described a cell
that responded best to the silhouette of a
monkey's hand, - Other cells responded to the shape of a banana or
a toilet brush (used to clean monkey cages) - (3) most widely studied are IT cells that respond
best to faces - (a) The responses of these cells are relatively
invariant to size, color, contrast, and position. - (b) some neurons respond to
- (i) particular features of faces
- (ii) particular face orientation
- (iii) face identity
162nd type of IT neurons Change their firing
patterns in accordance with their recent past
history (short-term or working memory). (1)
delayed match to sample task an animal is
presented with a sample cue, followed by a memory
delay during which that sample has to be
remembered. Then one or more choice stimuli are
presented and the animal is required to respond
depending on whether the choice cue is the same
as the sample (a match) or not (a nonmatch).
17Joaquin Fuster and his colleagues performed the
first studies (a) In one version of their task
the monkey was presented with a color cue and was
required to retain it for up to 20 seconds prior
to the choice. They identified cells that
fired differentially to specific colors of the
sample and choice. (b) Some of these cells
maintained high levels of activity during the
memory delay, and this activity was specific
to the sample cue. (c) Other studies have
revealed that many inferotemporal cells show
suppressed responses to repeated stimuli,
and suggested that this phenomenon may provide
signals for short-term memory.
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