Motor Cognition
Overview
•
•
•
Many people believe processes used to plan and
enact a movement can be used in problem solving
and reasoning
Moreover, the processes involved in perceiving
action are also involved in movement
This lecture will introduce key ideas involved in
motor cognition and memory for action needed
• in planning, and producing our own actions;
anticipating, perceiving, and interpreting the
actions of others
• In remembering actions
Motor Cognition
Terminology
•
•
Movement is a voluntary displacement of a body
part in space
Action – a series of movements performed to
achieve a goal
Motor Cognition
Perception-action cycle
•
Refers to the transformation of perceived patterns
(often visually) into coordinated patterns of
movement
• Examples. Returning a tennis ball, picking up a
mug of coffee, walking on uneven terrain
• According to this perspective much of human
behavior involves the interconnection of
perception and movement
• The mediating link between perception and
action is that a shared representation; that is,
the coding of perception and action is shared in
the brain
Motor Cognition
Motor processing in the brain
•
Neuroanatomy
• Area M1, the primary cortex. Neurons in this
region control fine motor movements, and send
fibers out to muscles themselves
• Premotor area (PM) sets up program for motor
sequences, and send input into M1
• Supplementary motor area (SMA) sets up and
executes motor plans
Motor Cognition
Motor processing in the brain
•
Neuroanatomy
• Think of these 3 areas as a hierarchy. M1 is at
the bottom of the hierarchy, and it enables
specific movements; PM higher up and it
enables sets of movements; at top SMA
represents overarching plans of action
Primary motor cortex (M1)
Posterior parietal cortex
Supplementary
motor cortex
(SMA)
Premotor cortex
Motor Cognition
Some evidence for roles of three areas
•
•
•
•
Mushiake (1991)
Recorded single-cell activity in monkeys in M1, PM,
and SMA
2 conditions of interest were IT (internally triggered)
and VT (visually triggered)
In both conditions monkeys were required to touch
3 pads on a panel; in the IT condition monkeys
needed to remember sequence and then touch the
panels in the remembered order; in the VT
condition monkeys touched panels as they were
illuminated
Motor Cognition
Some evidence for roles of three areas
•
•
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M1 cells were active in the premovement and
movement periods consistent with the hypothesis
that M1 is involved in movement
More SMA cells were active in the IT than the VT
condition, consistent with the idea that SMA is
important during planning since planning is more
important in IT than VT condition
More neurons were active in the VT than the IT
condition during the premovement and movement
periods
Motor Cognition
Some evidence for roles of three areas
• Conclusions
• Movement planning and production involves a
number of neuropsychological processes with
different functions. These processes occur in
different brain regions, and often occur
simultaneously
• Planning and then producing a response
involves different neural processes than
responding to environmental cues
Motor Cognition
Summary
•
•
We have reviewed the role of just 3 brain regions
involved in motor cognition
Other regions are also involved with their
involvement depending on the precise nature of the
task; in fact it’s been said that it takes the whole
brain to make a cup of coffee
Motor Cognition
Shared representation
•
•
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A considerable amount of data suggests that we
can efficiently represent the actions made by other
people (e.g., through viewing)
It has been hypothesized this occurs (and brainactivation studies support this) because the
representation of perceived action and produced
action is shared
These representations enable us to interpret the
actions of others, respond appropriately, and
efficiently learn how to physically produce actions
that were viewed
Motor Cognition
•
Mirror Neurons
• Mirror neurons—refers to neurons that fire
when organism (monkey) performs a specific
grasping movement, and when that same
grasping movement is observed being
performed by experimenter or monkey
Motor Cognition
•
Mirror Neurons
• Human evidence
• A variety of techniques have been used to
provide evidence for motor neurons in humans
• A transcranial magnetic stimulation study
showed increased excitability in the motor
system during observation of actions performed
by another person, but only for the brain regions
involved in the muscles used by the other
person
Motor Cognition
Apraxia
• Definition
• An impaired ability to generate skilled actions
that cannot be attributed to basic sensory or
motor disturbance
• Generally conceptualized to reflect a disruption
of a distributed praxis network
• Research has often focused on transitive
actions -- actions that involve manipulation of a
tool or object
• Examples– use of a hammer, spatula,
• Apraxia frequently observed after neurological
damage
• praxis network was thought to be left lateralized
Motor Cognition
Diagram of praxis network
Component Model Approach
Roy and Square, 1994; Roy, 1996
Sensory/Perceptual
System
Visual/Gestural
Information
Auditory/Verbal
Information
Visual Tool/Object
Information
Pantomime
Conceptual System
Production System
Knowledge of
Action
Knowledge of Tool/Object
Function
Response Selection
Image Generation
Delayed Imitation
Working Memory
Response
Organization/Control
Concurrent Imitation
Component Model Approach
Roy and Square, 1994; Roy, 1996
Sensory/Perceptual
System
Visual/Gestural
Information
Auditory/Verbal
Information
Main Responsibilities:
Analyzing visual gestural information
Identifying key features of tools and objects for use
Visual Tool/Object
Information
Component Model Approach
Roy and Square, 1994; Roy, 1996
Sensory/Perceptual
System
Conceptual System
Visual/Gestural
Information
Auditory/Verbal
Information
Knowledge of
Action
Visual Tool/Object
Information
Knowledge of Tool/Object
Function
Main Responsibilities:
Understanding tools, objects, and gestures
different types of conceptual knowledge are dissociable
Motor Cognition
•
Conceptual knowledge
• It appears that different types of conceptual
knowledge associated with action are
dissociable from each other as demonstrated in
the next slides
Motor Cognition
•
Conceptual knowledge
• Function knowledge vs manipulation knowledge
• Buxbaum and Saffran (2002) investigated
function and manipulation knowledge in apraxic
and non-apraxic patients with LHD (aside, px
were apraxic to gestural tests including tests of
pantomime)
• Function knowledge – which two items are most
similar in function (e.g., stapler, cellophane
tape, pen)
• Manipulation knowledge – which two items are
most similar in manner of manipulation (e.g.,
piano, typewriter, stove)
Motor Cognition
•
Conceptual knowledge
• Results apraxic patients were more impaired in
manipulation test, but than function test
• Kellenbach et al. (2003) used PET to
investigate brain activation associated with
function and manipulation judgments
• Results showed that when participants made
conceptual judgments about function, there was
activation of a left network consisting of the
ventral premotor cortex and the posterior middle
temporal gyrus
• When participants made manipulation judgment
an additional region, and additional region, the
intraparietal sulcus, was activated
Motor Cognition
•
Conceptual knowledge
The intraparietal sulcus plays an important role in
skilled object use (Heilman, 1993)
Motor Cognition
•
Conceptual knowledge
Visual-gestural knowledge
Beauchamp (2002) in neuroimaging study
showed that bilateral regions of the middle
temporal cortex were activated when tool
motion (i.e., gestural motion) was viewed in
comparison to human motion (i.e., person
jogging on the spot)
Motor Cognition
•
Conceptual knowledge
Beauchamp (2003) in neuroimaging study
showed that middle temporal gyrus responded
more strongly to tool motion videos and pointlight displays of tool motion
Motor Cognition
•
Point-light displays (aside)
• Animals and humans move in ways that are
distinctive and different from the way in which
nonhumans move. These patterns of movement
are called biological motion
• Johansson (1973) developed the ‘point-light
display’ technique to investigate movement.
• Small light sources attached wrists, knees,
ankles, shoulders, and heads of actors who
performed various movements (e.g., walking,
running, dancing)
Motor Cognition
•
Conceptual knowledge
Park and Roy (in prep) showed that patients with
LHD, but not RHD were strongly impaired on
function and manipulation tests but that patients
with LHD and RHD were impaired on tests of
visual-gestural knowledge
Motor Cognition
•
•
Conclusions
These studies suggest that different types of
conceptual knowledge associated with action are
dissociable from each other
• Three types of knowledge have been studied in
depth. These are knowledge of:
• Function
• Manipulation
• Visual-gestural knowledge of tool motion
Motor Cognition
•
•
Imitation in this model can be accomplished in two
different ways
• 1. directly from perception to action; and
• 2. indirectly through long-term memory
Evidence to support this comes from studies which
have shown:
• The general observation that meaningless
actions can be imitated accurately in cognitively
unimpaired individuals
• Findings of dissociation between imitation and
pantomime (e.g., Goldenberg & Hagmann,
1997; Ochipa et al., 1994); stronger
lateralization to pantomime than to imitation
• (interpret on basis of model)
Motor Cognition
•
What is acquired when we view purposeful action
• It would appear that we derive the goal of the
action
• (e.g., see a person reaching hand across table,
grasping a mug of coffee, and moving is toward
lips would be described as …drinking a cup of
coffee. In other words viewed actions tend to be
described in terms of the goal of the action
Motor Cognition
Imitation
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•
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Imitation -- ability to understand the intent of a
viewed action and then to reproduce it
This needs to be distinguished from mimicry, which
is reproduction of a behavior without understanding
(e.g., a parrot mimics human speech)
Meltzoff and Moore (1977) showed that newborn
infants can imitate viewed action (sticking out
tongue; opening mouth etc.)
By 6 months of age infants can imitate actions on
objects (e.g., shaking a rattle)
Motor Cognition
Imitation
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•
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As infants grow older deferred imitation abilities
increase (i.e., memory for imitated action)
data show that infants as young as 18 months
appear to represent intentions of actions not just
the action itself
E.g., children who watched an actor try to pull apart
a dumbbell but failed, were more likely to try and
pull apart the dumbbell than if they watched a
mechanical device attempt to pull apart a dumbbell
Motor Cognition
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Components of imitation
Decety et al. (1997) in neuroimaging studies
compared brain activation of subjects as they
viewed meaningless actions either intending to
recognize or to imitate the viewed action
Additional brain regions activated when intending
imitate meaningless actions: supplementary motor
area (SMA), the middle frontal gyrus, the premotor
cortex, the anterior cingulate, and the superior and
inferior parietal cortices in both hemispheres.
Motor Cognition
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•
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Conclusion: intentions (top-down) processes of
participant influenced observation of action.
Regions activated during observation also are the
ones involved in action generation.
Observing an action with the intention to perform
that action involves regions similar to those used to
generate the action
when intending to recognize an action activated
regions were the memory encoding structures (the
parahippocampal gyrus)
Motor Cognition
•
When actions are viewed — separating intention
from means – a neuroimaging perspective
• Several people have proposed that actions are
often understood in terms of the intentions
(goals) they achieve and the means used
(movements) to achieve these goals (e.g.,
Heider)
• Chaminade (2002) investigated using
neuroimaging the neural regions associated
with goals and means
Motor Cognition
•
When actions are viewed — separating intention
from means – a neuroimaging perspective
• Participants saw an actor make Lego
constructions: participants viewed the goal
alone (hand moving away from block in
specified location); the means alone (the hand
grasping and moving the block); or the entire
action. All participants imitated action just
observed
Motor Cognition
•
When actions are viewed — separating intention
from means – a neuroimaging perspective
• Findings– when participants imitated action or
means, the medial prefrontal cortex was
activated; this region appears to play a critical
role in inferring other people’s intentions
• When participants imitated goal the left
premotor cortex activated
• Conclusion
• In normally functioning adults imitating a
gesture activates neural regions associated with
the intentions underlying the action
•
Motor Cognition
Mental simulation
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Since imagery and perception activate
similar brain regions it seems
reasonable to hypothesize that one
way to reason is to simulate (or
imagine) the consequence of
performing a planned action
Motor Cognition
•
Simulation theories of action understanding
• It has been proposed that actions of others are
understood by putting yourself in their place
(either by observation or imagination)
• This permits you to derive their intentions and
generate an action plan (but how can you do
this since intentions and actions are internal
and unobservable?)
Motor Cognition
Mental simulation
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It has been shown that practicing with mental imagery can
help participants in their performance of the task
It has been shown that mental imagery practice has positive
effects on complex motor skill learning (e.g., putting a golf
ball)
Yue & Cole (1992) showed compared finger strength of two
groups– group 1 performed repeated isometric exercises;
group 2 received motor imagery instruction and imagined
making finger movements without actually making them
Both groups had increased finger strength– group 1 by 30%
and group 2 by 22%
Conclusion– possible to increase strength without actually
repeated muscle activation
Motor Cognition
Mental simulation
• It has also been shown that viewing an action can
facilitate enactment of that action
• Priming refers to the facilitation of processing by
previous performance of a task
• Motor priming refers to the facilitative effect that
watching a movement or action has on making a
similar motor response. Motor priming has been
observed in a variety of experimental situations
Motor Cognition
Mental simulation
• fMRI studies have shown that the neural difference
between motor imagery and motor performance is
not a matter of “what”, but “how much”
• i.e., similar brain regions are activated, but the level
of activation is significantly lower; in one study
imagery activation was 30% of that found in actual
execution (Roth, 1996)
Motor Cognition
Mental simulation
• Ruby & Decety (2001) Nature Neuroscience
investigated the question of agency
• Background—if viewing an action activates regions
involved in performing an action, how do people
distinguish actions they perform vs those they
observe (i.e., attribution of action to self or another
agent)
• Previous studies have shown that the first-person
perspective (imagining oneself) is associated with
activation of inferior parietal, premotor, and SMA
on left side
Motor Cognition
Mental simulation
• This study asked what areas are activated when we imagine
not ourselves performing an action, but another person
performing that action
• Method
• Participants were scanned while they simulated everyday
actions (e.g., winding a watch) with right hand (all Ps were
right handed)
• Ps instructed to imagine themselves perform the action or to
imagine another person performing the action
• Perspectives initiated by presenting a photo or from a spoken
sentence describing the action
Motor Cognition
Conclusions
• First person perspective versus imaging another
person acting was associated with activation of
common neural resources
• Consistent with notion that a common code is used to
perceive, imagine, and produce actions
• However, specific regions are activated when
imagining oneself performing an action versus another
person. These regions may be used to determine
agency
Motor Cognition
Mental simulation
• Results
• Both self perspective and other perspective activated
common regions – supplementary motor area (SMA),
premotor cortex, precuneus (an area located in parietal
lobe), and occipital-temporal lobe
• However, when compared to the first-person perspective,
the third-person perspective selectively activated the
frontopolar cortex, the precuneus, and the right inferior
portion of the parietal lobe
• See figure in next slide
Ruby & Decety (2001)
Motor Cognition & Memory
•
Memory for action
• Subject-performed task (SPT) paradigm
requires participant(P) to perform actions
according to verbal instructions given by
experimenter (e.g., roll the ball, fold the paper,
lift the pen) at study
• At test P’s memory for these actions is tested
• Control condition– P hears instructions but does
not perform actions
• Result—memory for enacted action phrases is
superior to that for events encoded without
enactment
Presentation relies on Nilsson (2000) In Craik
and Tulving Oxford Handbook of memory
Motor Cognition & Memory
•
Memory for action--theories
• Non-strategic encoding theory of Cohen
• This theory proposed that enacted actions are
encoded nonstrategically unlike verbal and
other types of events
Motor Cognition & Memory
•
Memory for action--theories
• Multimodal theory of Backman and Nilsson (1984, 1985)
• Enactment during encoding automatically leads to
multimodal processing, which produces a rich encoding
of information (multimodal because there is auditory,
visual, and haptic input) in SPT condition
• subsequently proposed that physical (perceptual)
properties were encoded nonstrategically, whereas verbal
components were encoded strategically.
• SPTs contained verbal and physical properties whereas
VTs contained verbal component only (Backman et al.
1986)
Motor Cognition & Memory
•
Memory for action--theories
• (Backman et al. 1989) proposed that the
physical component of the dual code is
encoded incidentally and retrieved implicitly,
whereas verbal component is encoded
intentionally and retrieved explicitly
Motor Cognition & Memory
•
Memory for action--theories
• Engelkamp and Zimmer (1984, 1985 etc.)
proposed that encoding SPTs is governed by
separate motor, visual, and verbal programs
that produce separate modality-specific
representations
• Motor encoding is more efficient than the other
types of encoding and this results in the
enactment effect
Motor Cognition & Memory
•
Memory for action--theories
• Motor coding improves item-specific encoding,
whereas visual and verbal processing result in
relational encoding between the items
• Two types of relational encoding: 1. integration
of actions within a list; 2. integration of noun
and verb in a command
• Hypothesized that type 1 integration is
independent of enactment, and type 2
integration is hindered by enactment
Motor Cognition & Memory
•
•
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Memory for action—data in support of Cohen
1. no levels of processing effect
2. no primacy effect
3. no generation effect
4. no rate of processing effect
5. no difference in memory performance for
children of different ages, for mentally retarded and
controls or for elderly.
These results all support notion that enactment (in
contrast to verbal encoding) does not require
strategic processing at encoding as hypothesized
by Cohen
Motor Cognition & Memory
Memory for action—data in support of
multimodal coding theory
These data are supported by studies that
have used a divided attention as Ps encode
SPTs at study (e.g., bounce the ball). At test
memory for perceptual (color of object) and
conceptual aspects (recall SPT) was tested
(from Backman, Nilsson, Herlitz, Nyberg, &
Stigsdotter, 1991)
Results shown in next slide
Motor Cognition & Memory
25
20
15
full
divided
10
5
0
verbal
color
Motor Cognition & Memory
Memory for action—data in support of
multimodal coding theory
This next experiment very similar to previous
one:
Ps encode SPTs at study (e.g., bounce the
ball) under full and divided attention.
At test memory for perceptual (weight of
object) and conceptual aspects (recall SPT)
was tested (from Backman, Nilsson, Herlitz,
Nyberg, & Stigsdotter, 1991)
Results shown in next slide
Motor Cognition & Memory
25
20
15
full
divided
10
5
0
verbal
weight
Motor Cognition & Memory
Conclusions
•recall of perceptual components is less
strongly affected by DA than recall of the
verbal instruction
•Supports hypothesis that verbal
component requires strategic processing
and physical (perceptual) component is
more automatic or non-strategic
•Potential problem with experiment is that
color recall is a form of cued recall,
whereas verbal recall is a form of free
recall. (This problem was addressed in
Backman et. al. 1993).
What’s a tool?
“a manipulable object that is used to transform an actor’s motor
output into a predictable mechanical action for the purpose of
attaining a specific goal (Frey, 2007)”
• Simple vs. complex tools
- Simple tools amplify the movement of the upper limbs (e.g.,
using a stick to extend reach)
- Complex tools provide a mechanical advantage and convert
hand movements into qualitatively different actions (e.g., using
scissors to cut paper)
What do I need to know to use this tool?
colour of the tool
function of the tool
manner of grasping
the tool
identity of the
recipient
colour of the recipient
how the tool physically
is used
learned motor skill
Memory Systems
Declarative
Memory
knowing “what”
Procedural
Memory
knowing “how”
Memory for Tools
Declarative
Memory
Procedural
Memory
?
Skilled Tool
Use
Tool Attributes
Motor Skill
Acquisition
?
Functional
Associative
Perceptual
Tool Grasping
Grasp-to-Use
Grasp-to-Move
Overview of Roy & Park
(2010)
• Investigated memory systems involved in the acquisition of different
types of complex tool knowledge in a single study
• Examined extent to which an amnesic individual could acquire
knowledge and skills related to novel complex tools
Why study amnesia?
- Individuals with amnesia are impaired in acquiring new declarative
knowledge, but have intact procedural learning
Ideal population to study dissociation between declarative and
procedural aspects of tool knowledge!
Method
Participants
• D.A.
- 58 year old man with 17 years of education
- Diagnosed with retrograde and anterograde amnesia after
contracting herpes encephalitis in 1993
Neuroanatomical Profile
Cognitive Functioning
Damaged /
Impaired
Medial temporal lobe
structures (bilaterally), right
anterior temporal lobe
Delayed memory
Spared /
Unimpaired
Dorsal frontal, superior and
inferior parietal, and posterior
cingulate regions
Immediate memory, visual
naming, fluency, digit span, and
executive functioning
• 6 healthy age and education-matched controls (3 males, 3 females)
Method
Materials
• 15 novel unimanual complex tools constructed using
K’NEX
• Tools were designed to act on a recipient (e.g., plastic wheel) to
perform a specific function (e.g., move wheel down a path)
• Tool function, manner of grasping, or manner of use cannot be
inferred based on physical appearance
Example of Novel Complex Tool
Procedure
S1
S2
3 days
S3
3 weeks
• Each session (S1, S2, S3) had 3 phases:
1) Pre-test
2) Training
3) Post-test
Procedure
1) Pre-test
- Recall test (e.g. tool function, tool colour)
- Recognition test
- Grasp-to-command
- Use-to-command
2) Training Phase
- 2 blocks (10 target tools x 2)
- Video demonstration followed by practice
- Limit of 90 seconds to complete one errorless trial
- Experimenter provided feedback
3) Post-test (same format as Pre-test)
Procedure
• Task order remained the same across sessions except....
D.A.’s S3 Post-test
- Recall test
- Recognition test
- Grasp-to-command
- Use-to-command
- Use-to-command Recipient cued (RC) trial
Changes made to bring D. A.’s performance off the floor
Hypotheses
1) D.A. would demonstrate unimpaired motor skill acquisition
associated with novel complex tools (i.e., becoming faster in
using the tools across training trials)
2) D.A. would be impaired in his ability to recall the
properties (functional and perceptual) of the novel tools
3) D.A. would be impaired on tasks that required him to
consciously demonstrate the appropriate grasp and trained
use of the novel complex tools.
4) There would be no effect of the 3-week delay on measures
of procedural memory in either D.A. and controls, but that
there would be an effect of the 3-week delay on measures
of declarative memory in the controls
Training
Completion Time (sec)
Completion Time (+/- SE)
by Training Trial
80
60
Controls
40
D.A.
20
0
T1
T2
S1
T3
T4
T5
T6
S2
S3
(3 days) (3 weeks)
• No differences between D.A. and controls in any training trial
• Completion time decreased by approximately 3.4 seconds per trial in
controls and 6.3 seconds per trial in D.A.
• No effect of the 3-week delay found in either D.A. or controls
Recall
80
100
Accuracy (%)
Accuracy (%)
100
Functional Associative Accuracy (+/SE) by Test Trial
60
40
20
0
S1 Pre S1 S2 Pre S2 S3 Pre S3
Post
Post
Post
(3 days)
(3 weeks)
Perceptual Accuracy
(+/-SE) by Test Trial
80
60
Controls
D.A.
40
20
0
S1
Pre
S1
S2
Post Pre
(3 days)
S2
S3
Post Pre
(3 weeks)
S3
Post
• For functional associative recall, D.A.’s performance was worse than controls
in all trials except in S3 Post
• For perceptual recall, D.A.’s performance was worse than controls in all trials
except in S3 Pre and S3 Post
• Performance for controls is significantly worse after the 3-week delay for
both categories
Grasp-to-command
Accuracy (%)
100
Grasp-to-command Accuracy (+/SE) by
Test Trial
80
60
Controls
40
DA
20
0
S1 S1 S2 S2 S3 S3 RC
Pre Post Pre Post Pre Post
(3 days) (3 weeks)
• D.A.’s grasp-to-command accuracy was worse than controls only in S3
Post
• Grasp-to-command accuracy in controls was worse after the 3-week
delay
100
Use-to-Command Completion
Time (+/- SE) by Test Trial
100
80
Accuracy (%)
Completion Time (sec)
Use-to-command
60
40
20
0
Use-to-Command Accuracy
(+/- SE) by Test Trial
80
Controls
60
D.A.
40
Lure Tools (D.A.
Only)
20
0
S1
Pre
S1
S2
S2
S3
Post Pre Post Pre
(3 days) (3 weeks)
S3
Post
RC
S1
Pre
S1
S2
S2
S3
Post Pre Post Pre
(3 days) (3 weeks)
S3
Post
RC
• D.A.’s completion time was worse than controls at S2 Post and S3 Pre, and
his accuracy was worse than controls at all trials
• D.A.’s performance on both measures in the RC trial was better with the
target tools than the lure tools
• Controls were slower and less accurate after the 3-week delay
Conclusion
Declarative
Memory
Tool Attributes
Memory
Functional
Associative
Perceptual
Procedural
Memory
Skilled Tool
Use
Memory
for Tool
Knowledg
e
Motor Skill
Acquisition
Tool
Knowledge
Tool Grasping
Grasp-to-Use
Grasp-to-Move