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297 Cards in this Set

  • Front
  • Back
What is memory for?
• Learning facts and keeping track
Information about
the world
… about objects
… about situations
… about language
Information about
… who we are
… what we have
-can't make new memories
-remembers old stuff
anterograde memory
-can learn now
-can't remember old stuff
retrograde memory
what was the key point of karl lashley's work
worked with rats. trained them to do maze. removed small portion of brains. rats did fine. didn't matter where in brain.
*no localization of memories*
what was the key point of penfields work
*storage of memories not localized in the brain
*stored across the cortex
(Penfield thought temporal region was important bc of woman who heard music when this part of her brain was stimulated....wrong)
medial temporal lobe (includes portions of hippocampus and other regions...) is important for
memory, amnesia
HM suffered intratable __ perhaps due to a bike accident at age 7.
in 1953, dr. scovile localized HM's epilepsy to __ and suggested surgical resection
Scoville removed parts of HM's MTL on both sides of his brain. HM lost__.
approximately two-thirds of his hippocampus, parahippocampal gyrus (and
all his entorhinal cortex) was destroyed, and amygdala.
His hippocampus appears entirely nonfunctional because the remaining 2
cm of hippocampal tissue is atrophic and because the entire _________
(forming the major sensory input to the hippocampus) was destroyed.
After the surgery he suffered from severe __amnesia: although
his short-term memory was intact, he could not commit new events to longterm
HM also suffered moderate ______amnesia, and could
not remember most events in the 3-4 day period before surgery, and some
events up to 11 years before.
However, his ability to form ___was still intact; thus he could, as an example, learn
new ____, despite not being able to remember learning them.
long-term procedural memories
motor skills
The selective nature of HM’s
disorder. Almost entirely restricted to _____.
He has _____damage restricted to the _____.
Such surgeries have never again been done in humans
HM’s perceptual, motor and
cognitive functions are _____.
HM performs ________on tests of visual acuity
HM can ____and _____objects
HM had ___than average IQ before operation and IQ
___slightly afterward
HM's language capacities largely ___.
HM's sense of humor ______
• Spatial capacities not depending on memory are
• ________ for most types of new learning
Almost no capacity
Overall: very pronounced memory problems beyond
short-term (working) memory
• In HM Childhood memories, including memory for English
language ______
• Some_______ for the period just before the
• Immediate or short-term memory ____
• Memory deficit immediately apparent after _________.
retrograde loss
short-term period elapses
Exceptions to new learning
problems: __________
• Intact mirror drawing learning, despite
not being able to recall taking the test
• This was a revelation in the field of
memory research (aka Lashley
• This is acquisition of a skill
mirror drawing
Exceptions to new learning
problems: ___________
• The serial reaction time task measures
learning in terms of Reaction time (RT) to
respond to a sequence of flashes
• When flashes are really random, no RT
speeding with practice
• When flashes obey “hidden” sequences
then the subject becomes sensitive to this
and shows speeding of RT
• This represents the acquisition of a skill
sequence learning
Exceptions to new learning
problems: __________
• Weather Prediction Paradigm
• On each trial, subject is shown 1 to 3 cards
• Must indicate if cards predict rain or sunshine
• Outcome is shown
• Outcome is probabilistically related to cards
• This makes it too difficult to learn using “explicit
• You learn it procedurally, as a skill, see later
Intact procedural but impaired
declarative memory
• Amnesics learn to
predict the weather
almost as well as
• And much better than
Parkinson’s patients
(with basal ganglia
• But amnesics
remember almost no
facts about the
testing episode
Weather Prediction Facts about episode
procedural learning
Intact procedural but impaired

• Amnesics learn to
predict the weather
almost as well as
• And much better than
Parkinson’s patients
(with basal ganglia
• But amnesics
remember almost no
facts about the
testing episode
declarative memory
Against Lashley’s view that cognition (and memory) is well
distributed throughout the brain, research on HM showed that it
can be _____, and that all sorts of dissociations amongst
memory systems are possible
the brain's ability to change as reult of experience
memory for info currently held "in mind" has limited capacity
memory for info that is stored but need not be consciously accessible; has an essentially unlimited capacity
longer term memory
silently mouthing words while performing some other task (typically memeory task)
articulatory suppression
a system for the temporary storage and manipulation of info
working memory
memories that can be consciously accessed
declarative memory
memories that cannot be consciously accessed (eg procedural memory)
non-declarative memory
explicit memory=
declartive memory
implicit memory
=nondeclarative memory
memory for skills such as riding a bike
proceduaral memory
conceptually based knowledge about he world, including knowledge of people, places the meaning of objects and words
semantic memory
memory of specific events in one's own life
episodic memory
memory for events that have occured after brain damage
anterograde memory
memory for events that occured before brain damage
retrograde memory
amnesia arising from long term alcoholism
korakoff's syndrome
the process by which moment to moment changes in brain activity are translated into permaneent structural changes in the brain
an increase in the long term responsiveness of a postsynaptic neuron in response to stimulation of a persynaptic neuron
long term potentiation (LTP)
the observation that memories from early in life tend to be preserved in amnesia
ribot's law
a memory test in which participants must decide whether a stimulus was/was not shown on a particular occasion
recognition memory
participants must produce previously seen stimuli without a full prompt being given (compare recognition memory)
context free memory in which the recognized item just feels familar
context dependent memory that involves remembering specific info from the study episode
info that is processed semantically is more likely to be remembered than info that is processed perceptually
levels of processing account
retrieval of a memory causes active inhibition of similar competing memories
retrieval induced forgetting
forgetting arising bc of a deliberate intention to forget
directed forgetting
the act of remembering consturcted in terms of making interferences about the past based on what is currently known and accessible
constructive memory
the process by which retrieved memorie are attributed to their original context
source monitoring
a memory that is either partly or wholly inaccurate but is accpted as areal memory by the person doing the remembering
false memory
a memory that is false and sometimes self-contradictory without an intention of a lie
______relates to vocab facts naming faces
takes ____ usually
known to depend son the ____: Amnesics very impaired
-this relates to higher level of of object recognition
semantic memory
multiple exposures
____ is memory for events
-usually 1 shot kind of memory
-known to depend on ___
-can be thought of as a ___ tying together items in your ____ memory
relates to emotion eg. ____ memories
one shot kind of memory
amnesia has a problem with ______
how moment to moment changes in brain activity are translated into permanent structural changes in the brain
-this relates to how new memories are formed but also can be used to account for ______
if consolidation takes days, wks, even years to achieve brain damage will affect events prior to the injury too
-this could aslo explain effects seen with _________
retrograde memory loss
electro convulsive shock (ECT)
make holes in memory (recent)
shock brain
used to treat serious depression
whatever memories are made of takes time
recall of events shows a
temporal gradient
memories from earlier in life are easier to recall than those from later in life
ribot's law
ribot's law can be explained by...
-older memories mayb ebecome more like stories and less like episodes
-each time an old event is rehearsed a new memory is made, so more resilient
cellular mechanisms of consolidation
Long term potentiation
can the synapses btw neurons in the hippocampus store information?
-input nerve cell was stimulated electronically
-found that high frequency tetanus produced an increase in synaptic strength in hippocampus lasting hours
-manifest in increased EPSP
2 types of consolidation
-fast synaptic consolidation... anywhere in nervous system
-slower system consolidation...
esp for hippocampus and declarative memory
adding new memory to cortex immediately would produce...
-in this model _____ learns quickly then integrates this info gradually elsewhere without disrupting existing memory
catastrophic interference
How memories may be made:
pathways into and out of the MTL
for information in
parietal (P) and temporal (TE) areas,
which are influenced by the frontal
cortex (FC), to develop into stable longterm
- neural activity must occur at the
time of learning along projections
from these areas to the MTL
- first to parahipp, perirhinal cortex
and entorhinal cortex
- then through several states of
hippocampus and back to Parietal
and Temporal areas
Role of MTL in encoding new
information confirmed with fMRI
Scan people while
they learn new
material (e.g.
complex color
...this study showed...
Activation at time
of scanning
predicts what is
Skills and habits=
procedural memory
The Weather Prediction paradigm
engages a different kind of memory
the Basal Ganglia (see lectures
on movement)
• This is because facts and episodes
aren’t relevant to learning to predict the
weather from cards
• Instead, it is brain-as-statisticalprocessor;
picking up associations
between stimuli and responses
• This is___________
nondeclarative, procedural,
implicit learning - acquiring a skill or
an fMRI
baseline baseline
Procedural learning
• “Boosting” ___________
led to more
response strategy
• “Boosting”
_________led to
even more place

This experiement was ______
double diss
Two types of declarative memory
Facts and episodes
MTL as
two explicit tests of memory
1. tests of recognition
2. tests of recall
in ____ memory diff tasks tax diff systems
explicit memory
_____ is easier than recall
subjects shown a list of words and then at test:
1. was given word previously preented on the list? (single probe recognition)
2. which of the two words shown togetherwas previously presented? (forced choice recogition)
tests of recognition
subjects shown a list of words and they recall them in
1. any order (free ___)
2. order given (serial ___)
3. given a prompt (cued ___)
tests of recall
are subregions of MTL specialized for recognition vs. familiarity?
-contentious issue
-fmri data suggest yes
-others have argued recollection is just a more vivid form of familiarity
fmri study of familiarity vs recollection
-subjects shown stream of words
-make animacy (living or nonliving) or size judgement depending on color of the word
-after scan subjects make recognition judgement depending on color of the word
-after scan , subjects make recognition judgement(1. def new...6. def old) and source memory (red or green)
-results: mean proportion of studied ('old') vs unstudied ('new') items endorsed at each confidence level
rhinal cortex
1. posterior hippocampus
2. posterior parahippocampal cortex show strong effects
familiarity and recollection are ...
two diff processes
this study suggests a a _____ btw regions for familiarity (______) and regions for recollection (______)
why do we forget?
forgetting may be important for efficient memory
it is desirable to forget where you parked your car on a particular (previous) day when you parked it in a diff place every day
-new info needs to be consolidated
-permanent memories take months to lay down
-watch tv...calms brain down...drinking alchol...better recall when you drink (moderation...)
levels of processing account for forgetting
-info that is prcocessed ____ is more likely to be remembered than info processed ____.
how would you remember better?
house-> big
house-> mouse
1st way
better recall when you process more deeply and give meaning to word
forgetting: a prob with encoding or retrieval?
-_______refers to what happens at the time of the stimulus
-______ refers to the time of trying to recall or recognize stimulus
-difficult to distinguish you may access a memory right now but you might in the future
-it would be wrong to conclude that you hadn't encoded it correctly and it would be wrong to conclude that retrieval as such was the problem
-also whether or not people can successfully retrieve a memory also depends subtly on the way they are tested
-memories might fade away _____ (eg weakening of synaptic connection)
-or there might be ___ mechanisms such as strategic contrl via inhibition
active, controlled, inhibitory mechanisms are
automatic side effect of act of retrieval is that other items get inhibited
retrieval induced forgetting
memories are voluntarily inhibited eg. think/no think paradigm
retrieval induced forgetting
memories are volutarily inhibited
directed forgetting
Train all cue-target
pairs to 50%
performance at test
Learn which are
suppression cues
If green ->Think
If red ->Nothink
Try to recall all
target words,
including those for
pairs not include in
phase 3
The Think/no-think paradigm
Results: think/no think
recall for NoThink targets
worse than for Think targets
• These results are interesting but
• Kind of like Freudian repression …
• Could suggest that active control
mechanism literally inhibits the memory
representation of the target word
• Perhaps the frontal cortex “stamps” in
MTL or other representations …?
• This is an example of cognitive (or
executive) control - we’ll revisit in later
•False recall of related words
False memories
False recall: why?
• False recall/recognition may occur because the non-presented
item has features whose memory trace is “reactivated” when the
features are primed by other other concepts
• Activity in ________has been observed for false recall
patients provide memories and information that
is false and even self-contradictory, without intending to lie
______have different memory problems from amnesics
• They don’t just forget the story, they embellish
Confabulation is associated with damage to
orbital frontal and
ventromedial frontal cortex rather than MTL
Why confabulation?
(3 expl)
• Failure of retrieval?
• Failure of source monitoring?
• Failure of executive control?
• Failure of retrieval?
– Episodic memories (as opposed to semantic ones) may be
retrieved using a strategy
– Damage to OFC will affect the retrieval operation - so
inappropriate fragments emerge for the current context
• Failure of source monitoring?
– Source monitoring refers to attributing memories to their
original context
– Requires an evaluation of the context in which the memory
was made (was it heard or seen? Was it internally imagined
or an external event?)
– But usually associated with lateral PFC, not OFC
• Failure of executive control?
– See later lectures on Executive Control
– Concerns control processes during task performance
– But usually associated with lateral PFC, not OFC
Frontal cortex is important for memory....
retrieval, and interacts with MTL
Recognizing (_____) is different from remembering/knowing
-It may depend on different sectors of ____
Forgetting is an important aspect of memory
-It could _____ or _____
False memories could be generated through ______
-Could lead to ____representation being activated
semantic priming
MTL something that occurs after frontal damage
The_______plays an important role in encoding and retrieval
of memory
frontal cortex
What is working memory?
• Short-term memory or working memory is memory for
information currently held “in mind” - i.e. it’s information that is
______ rather than _____
• Remember: HM has intact memory for immediate items, but as
soon as time elapses or he does something else, it’s gone
• George Miller famously argued that working memory span is for
• It seems possible that this could be true for _____ chunks rather
than items
• As we’ll see, ___is very important for
working memory - in concert with other regions
7 +/- 2 items
7 +/- 2
lateral prefrontal cortex
What is working memory? … more
• Baddeley posited that working memory consists
of 3 components:
– Central executive does the encoding and
– Two slave systems
- Phonological buffer, refreshed via the
articulatory loop
- The visuo-spatial scratch-pad for
nonlinguistic information
• In reality for working memory there are probably...
multiple multiple
“slave” systems; and the relation between these
and the “Central Executive” is still unclear
Certainly, working memory (or
short term memory) is an
important way station between
_______ and _______, as we have seen
sensory information
longterm memory
Lateral prefrontal cortex
• An important division is the ________
• Above is the _____
• Below is the ______
• Anterior is______
• There are many more areas in PFC - e.g.
Brodmann Areas (defined cytoarchitectonically)
• There may or may not be homologous areas in
the monkey
inferior frontal sulcus
dorsolateral PFC (DLPFC)
ventrolateral PFC (VLPFC)
the anterior PFC (APFC)
___________ - goal driven, topdown
• The cue designates a location in
space, but working memory is
needed to keep this ‘online’ for a
few seconds
______ or top-down attention engages
a ___________(both
• Demonstrated by many fMRI studies
dorsal frontoparietal network
typically associated with
working memory
• Connected to ______via
white matter tract
Dorsolateral PFC
Superior parietal lobe
Role of the dorsolateral PFC:
Lesion study in monkeys
______ condition
• Monkey displaces one object of 3
to get to food well
• At test: one between one of the
prior nonchosen objects and the
prior chosen one; monkey must
choose previously non-chosen
object to get reward (must monitor
prior choice across time)
• Monkey displaces one object
of 3 to get to food well
• At test: chosen object is
presented along with new one;
monkey must select the chosen
one (doesn’t require monitoring,
just recognition)
Dorsolateral PFC important for
_____not ________.
• affect monitoring but not
MDL - mid-dorsolateral PFC
• have no
effect, and there is no effect on NC
(normal control) animals
PA - periarcuate lesions (more
ventral and posterior)
• ______ is
important for
monitoring/maintenance but not for
dorsolateral PFC
______is probably important for
In ________ Subject sorts cards according to number, color or
object rule
• After some time, experimenter says “wrong” and
subject must stop sorting with that rule and find the
new one
• Lesions to_______ produce many persevative errors after the change of rule - the
subject keeps sorting according to the previously
correct (but currently incorrect) rule
• Part of the problem probably relates to poor _____- but things are more complicated as we’ll see in lectures on Executive Function
Wisconsin Card Sort Test
dorsolateral PFC
working memory
• Monkey remembers where food is across
a delay period
• Neuron in DLPFC fires strongly during
delay period
Single unit recording in monkeys:
delayed recall
Neural responses do
occur at stimulus
presentation and at
response and at reset,
but they are strongly
sustained throughout
the delay period
Single unit recording: Oculomotor
delayed response
How does working memory work?
Prefrontal cortex may be temporary
repository for “active” material
• May communicate with posterior
sensory areas to retrieve, rehearse
and maintain information
• Consistent with extensive
connectivity with posterior sensory
fMRI study with the N-back task
• The authors compare 0-back with
1-back with 2-back with 3-back
• On each trial there are several
fMRI scans to get a measure of
the stimulus, response and delay
• The design allows one to examine
the effect of time and of working
memory load
• Brain regions that “care” about
working memory maintenance
should show sustained activation
across time; those that “care”
about visual and response
aspects should change with time
Dorsolateral PFC maintains activity
• The authors perform two
– 1. Which regions show an
increase of fMRI signal with
increasing load?
– 2. Which regions show
increased signal for scans 2
and 3 vs. 1 and 4?
Visual and other cortical areas
showed an effect of _____- i.e.
they were sensitive to the
• DLFPC and SMG (parietal)
showed an effect of ____- I.e.
they are relate to holding memory
information in mind across time
showed an effect of load - I.e.
they are relate to holding memory information in mind across time
DLFPC and SMG (parietal)
A similar study: DLPFC activity
________with _______memory
in terms of awareness, ownership, plan as to what you want to do
action (vs. movement)
-tons of ways to pick up your keys
-potentially infinite number of solutions
-you can calculate and action from scratch each time
degrees of freedom problem
-specification or pattern of muscles
-higher level script for motor commands (single muscle program)
-eg. handwriting does not change when you use an effector
motor programs
handwriting does not change when you use _________...use other hand, still looks like your handwriting
actions are directed towards
-can't act in a vacuum (need objects)
-need visuomotor or sensory motor integration
-incorporate ____ and ____ visual system pathways
what pathway
where pathway
incoporate where your body parts are
bringing together of sensory and motor info for action
sensory motor transformation
frontal lobe=
primary motor
-antieror to primary somatosensory cortex and posterior to premotor cortex
primary motor cortex
-there is a homunculus in the ________ for control of body parts, just as there is in _________
motor cortex
somatosensory cortex
In the primary motor cortex
-areas such as the ___ have larger representation bc of fine control
In the primary motor cortex
___ controls right side of the body and vice versa
Left M1
damage to one side of the brain affecting M1 thus produces ____ (paralyzed on 2 side of the body)
motor cortex contains _________ for specififying voluntary movements of body parts.
projects down the ______
motor programs
corticospinal tract
motor programs are collections of lots of neurons which encode *specific movements* via
patterns of muscle activation
eyes are unusual- are controlled by _________ in laeteral superior frontal gyrus (avove superior frontal sulcus) and not by M1
frontal eye fields
Axons in M1 project down the white matter tract bundles into
1. pons (jaw and face)
2. cervical level of the spinal cord (hand)
3. lumbar level (legs, feet, trunk)
Brain machine interface in
neural signals in ____ may be sufficient to control simple prosthetic devices
-arms plugged into primary motor cortex (implant into primary motor cortex)
-moves arms by thinking
Anterior to M1 is the premotor cortex which has two major divisions
1. lateral permotor cortex
2. medial premotor (supplementary motor area= SMA)
role of lateral premotor cortex
Lateral premotor: acts with objects in
the environment (reaching for pen):
more externalised
– Lateral premotor cortex has _____than the motor
programs of M1 - not just muscle
representation but set of movements
(e.g. grasp)
motor plans at a higher level
role of SMA:
deals with spontaneous welllearned
actions, more internalised
TMS shows _________ in function
for SMA vs M1
• TMS delivered over SMA and M1 in 3
– “simple” button press
– “scale” buttons presses
– “complex” - playing a prelearned
musical piece
• TMS over SMA disrupted complex only
• TMS over M1 disrupted simple and
• Conclusion: SMA has critical role in
organizing forthcoming movements in
complex motor sequences that are
rehearsed from memory (i.,e. internal
Prefrontal cortex (PFC) lies ______to premotor cortex
PFC regions are involved in _____ cognition rather than action
Premotor regions prepare actions (to ________ triggered
internally or externally
Prefrontal cortex mediates their selection and
working memory
Prefrontal cortex and motor
Frontal lobe damage does
not impair movement or
execution of actions per se
• Instead, ...
Symptoms of frontal
the actions become
poorly organized - do not
necessarily reflect the goals
of the subject (“frontal
• Symptoms of frontal
– Perseveration
– Utilization behavior
Goal driven action: the supervisory
attention system (SAS)
• Key distinction: actions performed
________ vs those that require some
online control
Driving a car can be very automatic:
changing gear, stopping, going, turning are
presumably driven by ______ -
without the need for much awareness
• But parking in a narrow space requires
an interruption of automatic behavior, and
setting up of a novel sequence -> _________
motor schemas
The SAS model describes the
relationship between automatic schemas,
the control system and motor output in
terms of ________
contention scheduling
Goal driven action=
the supervisory
attention system (SAS)
The Supervisory Attentional System likely
has its home
in the frontal cortex
The frontal cortex exerts “executive
control” over: (4)
– Premotor cortex (motor plans),
– primary motor cortex (motor
– posterior cortex (e.g. parietal and
temporal representations)
– Subcortical regions (e.g. basal ganglia)
________ is:
(Miller and Cohen, 2001)
The optimization, by prefrontal cortex, of cognitive, sensory or motor
processing in posterior cortical and subcortical modules
Executive Control
The SAS model accounts for ______ after brain damage
action impairments
Damage to frontal cortex produces symptoms of
_______ behavior by
damaging the putative Supervisory Attentional
• Partly could relate to working memory problems,
but …
• Automatic schemas or habits or routines run wild
without ....
Perseveration and Utilization

...appropriate contention scheduling and
SAS monitoring
• Subjects sits in front of a oscilloscope timer with a rotating dot. EEG electrodes on scalp.
• Subject presses a button (or moves wrist) within a certain time frame.
• No limits placed on # times subject could perform the action within this period.
• During the experiment, the subject noted the position of the dot on the oscilloscope timer when
"he/she was first aware of the wish or urge to act"
• Pressing the button records position of dot on the oscillator, electronically.
• By comparing the marked time of the button's pushing and the subject's conscious decision to
act, researchers were able to calculate the total time of the trial from the subject's initial volition
through to the resultant action.
• On average, approximately ____ elapsed between the first appearance of conscious will to
press the button and the act of pressing it.
• Brain activity involved in the initiation of the action, primarily centered in the SMA, occurred, on
average, around ____ before the trial ended with the pushing of the button.
• So, researchers recorded mounting brain activity related to the resultant action as many as
___before subject reported first awareness of conscious will to act.
• So, apparently conscious decisions to act were preceded by an unconscious buildup of electrical
charge within the brain - ______________

the Bereitschaftspotential or readiness potential.
– ______is at the apex
in terms of conceptual
cognitive plans
– It organizes, selects,
controls lower-level motor
plans and programs, e.g.
in premotor cortex
– ______generates plan
for motor programs
– Activation of arm and
other effectors occurs via
primary motor cortex
- higher plans and
working memory
internally generated
generated actions
• Lateral premotor
motor programs and
movement execution
memory for object names and
where, how and visuomotor integration
Parietal cortex
e.g. V5 - motion area
Occipital regions,
fine timing
Why is it impossible to tickle
A sense of ownership over one!s actions
could be maintained by predicting the
sensory outcomes of our actions
This can happen via a ______
• A representation of the motor command
("efference copy!) is used to predict the
sensory consequences of action
• This could explain why tickling yourself
doesn!t work - you sensory system gets a
copy of the motor command before it
forward model
Our actions vs others’ actions
• Mimicry vs imitation
• Humans may reproduce (and understand?) the
actions of others by imitation, i.e. by representing
the goal state.
• An important finding in neuroscience relevant to
our understanding of the goal state and action is
the discovery of _____
• An important finding was that
neurons within region F5 of the
monkey brain ___ as well as the _________
show increased
activity for goal-directed action
• Both when monkey performs a
goal-directed action or when it
views a human or another monkey
performing a goal-directed action
• It is not certain what is the
corresponding region in the human
brain, but it may include ventral
premotor cortex as well Broca!s
region (or Brodmann area 44)
mirror neurons, or the mirror
(ventral lateral premotor cortex)
parietal lobe
An important finding was that
neurons within region F5 of the
monkey brain (ventral lateral
premotor cortex) as well as the
parietal lobe show increased
activity for _________
goal-directed action
Both when monkey performs a
goal-directed action or when it
views a human or another monkey
performing a___________
goal-directed action
It is not certain what is the
corresponding region in the human
brain, but it may include
premotor cortex as well Broca!s
region (or Brodmann area 44)
• A mirror neuron in F5
responds strongly when the
human movement is a _______ - i.e. to grasp the
• The neuron also responds
strongly when the monkey
makes the _____
Monkey observing human action
• A mirror neuron in F5
responds strongly when the
human movement is a goaldriven
action - i.e. to grasp the
• When there is no object the
neuron hardly cares.
• The neuron does care about
goal-driven action even when
the final destination is obscured
by a screen - as long as monkey
saw the object there beforehand
• Neuron does not care about
the reach if there is no object
goaldriven action
movement too
What vs. where/how
• The _______ has been characterized as “Where”,
but it can also be characterized as “How” if we emphasize
output (action) rather than input (perception)
• Remember: damage to visual stream causes apperceptive
agnosia - this is a problem with ________
• Damage to dorsal stream (parietal lobe) produces _______ but also severe problems with acting towards objects
(especially under visual guidance) - optic ataxia
• This is a probem with __________
dorsal visual pathway
object recognition
attentional neglect
sensory-motor integration
Typically, left parietal
damage affects _______
and vice-versa
The deficit is not purely
motoric (“bad” hand functions
ok in “good part” side of
space ) and not purely visual
(“good” hand functions poorly
in “bad side” of space).
Instead, such patients have
profound problems
________ sensory input
(visual) with the motor system
right hand
• 3D structural description of the object
(tool) is computed based on grouping of
parts (edges, form, depth, etc.)
• The representation is probably in the
typical (normalized) view and it probably
located in ______
• The tool will have access to semantic
representations (e.g. what to do with it)
and have a name - probably a function of
the medial temporal lobe
• But tools are different from other objects
in that they have specific gestures
associated with them - likely stored in _________ - perhaps because most people are right handed
infero-temporal cortex - ventral
visual stream
left inferior parietal lobe
Left inferior parietal cortex and ideomotor apraxia
• Subjects had to generate either an action or a name in response to an
object, OR they had to generate an action or a name in response to an
• Generating an action to an object (pantomiming) activated ________
• Patients who cannot produce an appropriate action on command or to
a visual object (or picture of an object) have ______
left inferior parietal cortex
ideomotor apraxia
Basal ganglia anatomy
5 key structures:
• Caudate
• Putamen
• Globus Pallidus
• Subthalamic Nucleus
• Substanta Nigra
Basal ganglia circuitry
• There are excitatory and
inhibitory connections
• Cortex send
projection to striatum
• Striatum sends _________
projections to Gpe and Gpi
• Gpe and Gpi send _____
connections on to thalamus
• Thalamus sends ________
connection on to cortex
Movements are selected by means
of the
direct pathway
Striatum inhibits Gpi, thus removing
inhibition on thalamus, and
thalamocortical drive
Movements can be inhibited by the
indirect pathway
Striatum inhibits Gpe, thus removing
inhibition on Gpi, thus increasing
inhibition on thalamus, and
thalamocortical drive
• Single gene, genetic disorder
• Autosomal dominant
• 5 to 10 per 100 000
• Leads to massive striatal cell
loss (and cortical loss too)
• Choreic movements
- classic HD symptom
- involuntary ‘writhing’
Choreic movements
______relates to loss of striatal
projection neurons to pallidum
_______ lead to
loss of projection neurons to
globus pallidus (especially
• This affects the _______
• Means the “braking” of
movements is weak - so motor
commands are disinhibited -
i.e. chorea
Loss of striatal cells
indirect pathway
• Affects 0.15% of the population
• Rigidity, dystonia, bradykinesia
• Aetiology uncertain - multiple factors
• Neuropathology now well
understood - partly because of MPTP
• These people developed ..after
ingesting the drug
• Leads, like ..., to loss of dopamine
cells from substantia nigra pars
compacta - which project to striatum
Parkinsons D
_____ caused by loss of dopamine
neurons projecting to striatum
• Loss of enervation of striatum
leads to abnormal ability for
subjects to initiate movements
via cortical-striatal circuits
• Direct pathway drive is too
• Gpi is not inhibited enough
• Gpi output to thalamus too
• Thalamocortical output weak
Direct pathway is too weak
Why the basal ganglia?
• Most movements are executed
via M1
• So why are the basal ganglia
• Answer: They allow complex
interchanges between cognitive,
motor, emotional, and
motivational information
• All this kind of processing is
brought to bear to determine the
best response
They let through the response that is most “active” in
terms of best fitting goals, emotion, and motivation (think dam)
Basal ganglia act as a response
selection device
PFC reaches largest relative
size in ______
• Comprises 1/3 of cortex
Increase in PFC must relate
to (and parallel)
abstract representation,
planning and thinking
We do these things so much
better than the great apes
and we have a much ____
• PFC damage affects these
functions, but not much else
Major Sulci of PFC
• Precentral
• Superior frontal
• Inferior frontal
• Sylvian
Major Gyri of PFC
• Superior frontal gyrus
• Middle frontal gyrus
• Inferior frontal gyrus
• The pattern of direct connectivity, via white matter
association and projection pathways is complex
• Two examples we’ve already seen concern:
- prefrontal to striatum connections
- prefrontal to parietal connections
• PFC has other extensive connections with other brain
regions, and there is an extensive pattern of
connectivity within PFC, but that is beyond this course.
Connectivity of PFC to other
brain regions
DLPFC and VLPFC connect to
parietal lobe
• _____for
• ________ for top-down
Ventral fronto-parietal
Dorsal fronto-parietal
So what does the PFC do?
• We have already seen that it’s important for:
– Spatial attention (e.g. top-down attention)
– Working memory maintenance across delay
– Short-term (working) memory to facilitate encoding and
retrieval of material via MTL and cortical networks
• Higher-level representations of action plans and schemas to
enable movement and behavior
• Another, similar, way of thinking about
executive functions is in terms of automatic
vs controlled
• As we saw before, this is how the
Supervisory Attentional System is
supposed to work:
• Sensory information can automatically
trigger associated motor programs, leading
to movement, but when there’s a conflict, or
unusual situation, the SAS kicks in and
does contention scheduling
• We could regard the SAS as a set of
executive functions
Automatic/habitual vs controlled
1. Situations involving planning and decision making
2. Situations involving error correction and trouble
3. Situations where responses are not well-learned or
contain novel sequences
4. Situations judged dangerous or technically difficult
5. Situations requiring overcoming strong habitual
responses or resisting temptation
Executive functions in practice
• There are a few real world scenarios where control over
automatic tendencies is needed:
• The “Stockings of
• Subjects inspects the
• This is a 3-move
• Subjects decides (in
head) exactly which
moves to get yellow ball
into right hand pocket
Tasks of planning and decision making
• The Wisconsin Card Sort Test requires subject to
sort a new card on each trial according to 3
possible rules
– Number, Color or Shape
• Experimenter says, “correct”, “correct”, “correct”,
but suddenly, “wrong”
• Subject has to register the error, control the
tendency to sort on the previously correct rule,
remember what she previously did, remember the
currently relevant rules, choose a new one
• The experimenter could say “correct” or “wrong”
on the next trial
• Depending, the subject has to process this new
Tasks of error-correction and
trouble shooting
• Requires subject to develop a strategy to deal with a
novel situation
• E.g. the F-A-S test
• Subject asked to produce as many words as possible
beginning with a given letter, e.g. “F”
• Do not use proper names
• Do not repeat words
• Do not repeat related words such as “fast” and “fasting”
Novel situations
• Driving car vs parking car in tight spot
• Danger is typically associated with
arousal and emotion
• Emotional/arousing signals may
engage the ______ and feed into VLPFC
and DLPFC to generate response
• In this way, automatic or habitual
responding could be overriden
Dangerous or difficult situations
Overcoming habitual responses
• This can be tested in many ways:
- Stroop paradigm
- Eriksen Flanker task
- Go/NoGo or Stop signal
• Subject responds to central letter in
display, here it “S”
• Central letter can be flanked with
congruent letters, i.e Ss or
incongruent ones, i.e. Hs
• Reaction Time and error rate is
longer for incongruent compared to
congruent condition
• In incongruent condition, there is
_________ and control is
needed to ensure the correct
The Eriksen Flanker Test
response interference
memory maintenance/monitoring
Dorsomedial and
conflict monitoring
Orbital (ventromedial)
Ventrolateral and
Anterior frontal and
higher-level goals
Dorsolateral PFC and
monitoring/maintenance of memory
• DLPFC is BAs 46 and 9
• We’ve seen in prior lectures that
DLPFC is important for working
memory ______- i.e. what are the
rules of the task? What am I doing
now? Which object was the relevant
• Monkey displaces one object of 3
to get to food well
• At test: one between one of the
prior nonchosen objects and the
prior chosen one; monkey must
choose previously non-chosen
object to get reward (must monitor
prior choice across time)
Monitoring condition
• Monkey displaces one object
of 3 to get to food well
• At test: chosen object is
presented along with new one;
monkey must select the chosen
one (doesn’t require monitoring,
just recognition)
Recognition condition
Dorsolateral PFC important for
monitoring not
• _________ PFC lesions
affect monitoring but not recognition
• ______ have no effect,
and there is no effect on NC (normal
control) animals
• Thus ________ is important
for monitoring/maintenance but not
for recognition
• ___is probably important for
MDL - mid-dorsolateral
PA - periarcuate lesions (more
ventral and posterior)
dorsolateral PFC
• Monitoring is a variant
of working memory
• Although some have
argued that monitoring
and maintenance are
different functions (see
your book)
• Neural responses in
monkey DLFPC
strongly maintained
across the memory
Single unit recording: Oculomotor
delayed response
DLPFC activity increases with
memory load
___damage affects
Wisconsin Card Sort
• Human subjects with ______
lesions are impaired at shifting their
response to a new rule after
negative feedback
• Trouble maintaining task rules?
• Trouble monitoring self and
• ____frontal cortex
– BA 24/32 (aka Anterior
Cingulate aka ACC) and
– pre-Supplementary motor
area (aka, preSMA, aka medial
premotor, BA 6)
Dorsomedial frontal cortex and
“conflict monitoring”
The dorsomedial frontal cortex is
typically activated for ___
• In the Stroop task
incongruent trials are
contrasted with congruent
• RT is slower on incongruent
• fMRI signal is increased in
preSMA and ACC
voltage at surface of skull
• Raw time-course at dozens of
electrodes is voltage across time
(relative to a reference electrode
on back of neck)
• EEG signal is really noisy
(treat as 1 trial)
• ______is event-related
• You get it by averaging
• 10, less noisy
• 100 averaged trials starts
to have a typical pattern of
positive and negative peaks
Many ERP studies have shown an
“error” or “conflict” response over
dorsomedial PFC
• In Stroop and other conflict
tasks you average, e.g. 100
incongruent trials and e.g.
100 congruent trials
• Large _____
negativity (on incongruent
trials with mistake) found
over dorsomedial electrodes
• This is a form of conflict
• Stimuli consist of face plus name
• Sometimes incongruent,
sometimes congruent
• Face activates ….
- fusiform face area!
• Word activates ….
- gotcha!
-word form area
fMRI experiment investigates
how conflict may generate control
How conflict-monitoring *may*
generate control
• These results are a
cartoon ….
• On a high-conflict trial
(incongruent), ACC is
• Supposedly recruits
DLPFC which sends ____ to FFA or to
Word Area to ______the
Hence ….
frontal cortex
topdown signals to
Problems with this study
• Not clear that how DLPFC is
connected to FFA and Word
• Not clear that ACC signals
conflict and then recruits
DLPFC, as opposed to other
way round
• Not clear exactly what is role of
-working memory for tasks
- top down signals (how?)
cortex corresponds to BA 11
• Phineas Gage is the famous case
in history
• Selective damage to orbital
frontal cortex left him with profound
problems with decision-making,
especially involving risk and social
• But remainder of
• Orbital (or ventromedial) frontal
• ______is intimately wired up with the limbic (emotion
processing) areas of the brain - see later lectures
• It may send an “emotional” input into the deliberative process
• Emotion processing can be
measured autonomically, e.g.
via the galvanic skin response
• Voltage at skin changes with
more sweating
• It is reported that patients with
OFC damage show less (or no)
GSR response to emotional
• Hence - their decision-making
may be impaired because they
don’t get the “body markers”
The “somatic marker hypothesis”
there are potentially infininite number of motor solutions for acting on object
degrees of freedom problem
there are potentially an infinite number of motor solutions for acting on an object
degrees of freedom problem
a stored routine that specifies certain mtor parameters of an action (eg. relative time of strokes)
motor program
a cluster of perceptual processes that relate to the skin and body and includes touch pain theraml sensation and limb postion
knowledge of the position of the limbs in psace
linking together of perceputual knowledge of objects in space and knowledge of the position of one's body to enable objects to be acted on
sensory motor transformation
the problem of explaining volitional acts without assuming a cognitive processt hat is itself volitional ("a man within a man")
homuculus problem
responsible for execution of voluntary movements of the body
primary motor cortex
damage to one side of the primary motor cortex results in a failure ot voluntarily move the other sid eof the body
responsible for voluntary movement of the eyes
frontal eye fields
the lateral area is important for the linking action with objects int he enviornment the medial area is known as the supplementary motor area and deals with well learned actions and action sequences
premotor cortex
deals iwth well-learned actions particularly action sequences that do not place strong demands on monitoring the enviornment
supplementary motor area (SMA)
repeatng an action that has already been performed and is no longer relevant
impulsively acting on irrelevant objects in the enviornment
utilization behavior
an organized set of stored info (eg familiar action routines)
the mechanism that selelcts on particular schema to be enacted form a host of competing schemas
contention scheduling
failure in tasks of routine activity that involve setting up and maintaining diff subgoals but with no basic deficits in object recognition or gesturing the use of isolated objects (also called action disorganization syndrome)
frontal apraxia
a representiation of the motor command (a so-called "efference copy") is used to predict the sensory consequences of a n action
forward model
the ability to reproduce the behavior of another through obervation
a neuron that responds to goal directed actions performed by oneself or by others
mirror neuron
an impairment of visually guided reaching
optic ataxia
a sensation that amputated or paralysed limb is still present
phantom limb
an object that affords certain actions for specific goals
an in ability to produce appropriate gestures on command given either an obect, a word, or a command
ideomotor apraxia
certain structural properties of objects imply certain usages
a disease associated iwth basal ganglia and characteried by a lacck o self initiated movement
parkinson's disease
a test of exectutive functions involving rule induction and rule use
wisconsin card sorting
response interference from naming the ink colour of written color name (eg word blue is printed in red ink and participants asked to say ink color ie red)
stroop test
a model of executive functions that assumes that controlled behavior arises through selection and implementation of schemas
supervisory system model
remembering to do things in the future (eg to deliver sthing or take meds)
prospective memory
the prcoess of relating info currently held in mind back to the task requirements
retaining focus on the task requirements over a period of time
sustained attention
a working memory task; the participant must decide whether the currently presented sitmulus is the same as the one persented immediately before (1back) or two items ibefore (2back) or three items before (3back)
N-back task
carrying out several tasks in sucession; requires both task switching and maintaing future goals while current goals are being dealt with
an electrical potential (error potential) that can be detected a the scalp when an error is made
error related negativity
a situation in which a perpotent incorrect response needs to be overcome to perform a task successfully (as in stroop color naming)
response conflict
discarding a pervious schema and establishing a new one
task switching