here are replies tanice Which of the following is not included in the factors that contributes to why action is computationally difficult?
a. Degree of Freedom
b. Sensory to motor transformation problem
c. Lack of awareness and skills necessary
d. Homunculus problem
(S) Alsah123:
Include
Amelia Which of the following describes when executive functions are needed to optimize performance?
a. A situation is novel.
b. Several cognitive processes need to be coordinated.
c. A situation does not require an automatic response.
d. A situation is complex.
e. All of the above.
Unit 4: Cognitive Control of Action
and Behavior
Our goal is to form a basic understanding on the neural
basis of cognitive control. Perception → Action
Part 1. From perception to action – cortical and
subcortical neural circuits (4 th ed. Ch 10; bb)
Part 2. The executive brain: (a) frontal lobe
hypothesis, EF tasks, EF disorders; (b) EF
anatomy, PFC functional divisions (4 th ed. Ch
15; bb)
Part 3. Motivation, reward, and decision making (bb)
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Part 2a: Frontal lobe hypothesis, EF
tasks, and EF disorders
A. What is executive control?
B. The frontal lobe hypothesis – Early evidence
for the importance of (pre)frontal cortex in
executive control
C. Executive functions in practice
D. Disorders associated with PFC
E. Neuroethics: Cognitive neuroscience and law
Note: We use the terms action control, cognitive
control, and executive control interchangeably.
The executive brain – Part 2a
Learning Objectives:
●
Identify daily life situations requiring executive control; define
executive control (EF) operationally; distinguish automatic vs
controlled mode of processing (model free vs model based)
●
Describe the two forms of PFC neural firing implicated for EF
●
Describe the history of frontal lobe hypothesis; e.g., describe
the classical “frontal lobe syndromes”, cases, and evolutionary
development
●
Define the cognitive domains of EF and describe the various
behavioral tasks used to test them
●
Describe PFC disorders: lateral vs. ventromedial PFC damage
●
Briefly describe cognitive neuroscience and law
Refer to the Leung lecture video
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A. What is executive control?
1. The human brain must perform a broad array of
oversight functions in a wide range of situations.
Norman & Shallice (1986) identify 5 general
situations requiring executive functions:
– Situations that require the overcoming of a
strong habitual response or resisting temptation
– Situations where responses are not welllearned or contain novel sequences of actions
– Situations involving planning for the future
– Situations involving reasoning and problem
solving (error correction or trouble shooting)
– Situations that require decision making
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What is executive control (cont.)
Empirical evidence from electrophysiology studies (cont.)
2. There is no homunculus (No “CEO” in the brain!)
Evidence 1: Memory-tuned neurons in the PFC
●
Working memory (e.g., mental arithmetic, reasoning)
●
Prefrontal neurons show ___________ firing during
temporary maintenance of task-related information
●
●
●
●
No single part of the brain can be identified as the
sole region for executive control
Executive control systems involve many cortical
and subcortical systems
However, the prefrontal cortex has been implicated
to play a critical role executive control
Two important pieces of empirical evidence from
electrophysiology studies (next two slides)
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Fill in the blank:
A. Intermittent
B. Ramping up
C. Ramping down
D. Continuous
Goldman-Rakic (1992)
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Empirical evidence from electrophysiology studies (cont.)
What is executive control (cont.)
Evidence 2: Rule-selective neurons in the PFC
3. Automatic vs. Controlled Processing (e.g.,
Schneider & Shiffrin, 1977)
●
Automatic Mode of behavior – responses are
made reflexively or automatically (Model free)
●
Executive control processes are needed to
overcome the limits of automatic processing
(Model based)
Wallis and Miller, 2001
Which allows flexible and adaptive behavior?
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What is executive control (cont.)
4. Main cognitive domains of executive functions
B. Frontal lobe hypothesis
(i) Working memory – maintenance & manipulation
• The prefrontal cortex (PFC) has long been
associated with controlling behavior and higher
cognitive functioning
(ii) Inhibition – suppression of an automatic behavior
that is not appropriate in the current context
(iii) Task Switching (initiation & shifting) – changing
from one behavior to another
(iv) Simulation – Generating a potential future state,
necessary for planning and reasoning
(v) Reward Evaluation – Track the consequences of
behavior
Miller and Cohen, 2001
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The case about W.R.
Watch the
Youtube video
W.R. had suffered a seizure during his last year in law
school, after staying up all night and drinking large
amounts of coffee so that he could study for a midterm
exam. Since then, his life suddenly seemed to change
course. [10 years past by.] His family thought he was
experiencing an early midlife crisis. But, W.R.’s chief
complaint was that “he had lost his ego”. A CT scan
revealed that W.R. had an astrocytoma – a large tumor
invaded his lateral prefrontal cortex in the left
hemisphere. W.R.’s prognosis was poor. Though his
brother shed tears upon hearing the news, W.R.
remained relatively passive and detached. W.R. Had
lost the ability to engage in goal-oriented behavior.
(Knight & Grabowecky, 1995)
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1. Classical frontal lobe “syndromes”
Patients with (pre)frontal damage show a constellation
of cognitive, emotional, and behavioral deficits
(e.g., Phineas Gage)
●
Are often superficially normal
can identify objects and sounds and perform motor
tasks with dexterity
➢
can comprehend and carry on simple conversations
➢
●
Yet do poorly when faced with real-world challenges
some are easily distracted and inflexible, apathetic, lack
insight into the consequences of their actions
➢
some are impulsive, risk taking and socially
inappropriate, etc.
➢
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2. History- Comparative Anatomy
3. Evolutionary development
Leonardo Bianchi (1900s) – found that bilateral
prefrontal cortex lesion in monkeys/dogs
resulted in:
●
reduced working memory,
●
inability to perform complex tasks
●
inability to formulate steps needed to reach a
goal
●
altered emotional attachments
●
changes in social skills
●
increased apathy and distractibility
a) Korbinian Brodmann (Early 20th century)
Ranked mammals by the size of their prefrontal
cortex
Found that, proportionally, humans have the
largest PFC
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b) A more recent view: Brain size (relative or
absolute) does not explain differences in cognitive
capabilities between mammals. (e.g.,
Semenderferi et al. 1997, 2002)
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C. Executive Functions in Practice
Examples of common experimental tasks for
testing executive functions
Problem solving: Tower of London task (Shallice,
1982), FAS test
●
Fluid intelligence (Duncan et al.)
●
Working memory tasks: e.g., self-ordering
●
Inhibition: e.g., Stroop task, Go/No-go,
Antisaccade
●
Task switching: WCST, Rules for different
stimulus-response associations
Watch demos
●
Semendeferi et al. 2002
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Executive functions in practice (cont.)
Executive functions in practice (cont.)
Task-setting & Problem Solving tasks – related to
lay notions of (fluid) intelligence
●
●
Working memory tasks
Tower of London task: PFC activated in functional
imaging during task (healthy participants), and damage to
PFC results in poor performance (Shallice, 1982)
In the verbal domain: Cognitive Estimates (Shallice &
Evans, 1978) e.g. “How many camels are in Holland?”
Cognitive fluency (FAS test): generate as many words
beginning with “F” (or “A” or “S”) in one minutes
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Delayed recognition tasks
From Bo & Seldler
A self- ordered pointing task based
on Petrides and Milner (1982).
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Executive functions in practice (cont.)
Executive functions in practice (cont.)
Inhibition tasks
Task switching tasks
The Wisconsin Card Sort
task (WCST)
The test requires shift in
strategy following an
unexpected rule change via
trial and error
Stroop: more complex, involves
components other than inhibition
(e.g. neuroimaging and lesion
studies suggest involvement of
anterior cingulate cortex
Individuals with Lateral PFC
damage show perseveration
deficits
Simpler tasks (see diagrams on
the right): involves the
ventrolateral PFC and FEF
Based on Milner (1963).
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Task switching (cont.)
Watch demo
online
Watch demo
online
More simple studies developed that
isolate different aspects of a switch
(and used more in healthy
participants rather than patients)
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D. Disorders associated with the PFC
Executive control functions are not localized to
particular PFC regions. However, it can be useful
to discuss different dysfunctions in associated with
different regions.
Relate to the EF components (in the earlier slides)
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Disorders associated with lateral PFC damage (cont.)
1. Lateral PFC damage
Impairment in “initiation” can be seen following
damage to the lateral PFC
a) Apathy – diminished motivation or affect
b) Abulia – lack of ability to act or make decision
c) Frontal dysexecutive syndrome – unable to
initiate and change actions, motor movements,
and mental plan
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d) Disinhibition
●
Inability to stay focus in the face of distraction
(Bob Knight and colleagues)
●
May have normal working memory capacity, but
make many more errors
●
Unable to control impulses (e.g., schizophrenia,
ADHD)
●
Deficits in these tasks conditions: working
memory updating, go/no-go, go/stop
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Disorders associated with ventromedial PFC damage (cont.)
2. Ventromedial PFC damage
c) Frontal Disinhibition Syndrome associated with
bilateral ventral and medial frontal damage
●
Normal performance on response selection and
working memory tasks.
●
Lack of insight
Impairment in “inhibition” can be seen following
damage to the ventromedial PFC
a) Socially inappropriate behavior and/or
dependency on information from the immediate
sensory environment
b) Environmental dependency syndrome
• Imitation and utilization (Lhermitte, 1980s)
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E. Neuroethics: Cognitive
Neuroscience and Law
• Do “psychopaths” show
unusual patterns of brain
activity?
• Can neuroscience data be
used in court?
– Nita Farahany: The cat is
already out of the bag!
– Buckholtz: Group vs
individual difference
– Kiehl: fMRI is just one
variable
d) Acquired Sociopathy (Damasio) – the more
severe form of frontal disinhibition syndrome
●
Blunted emotional affect, poor decisions in social
situations, difficulty interacting with others
●
Unlike congenital sociopathy, patients can state
appropriate rules of behavior, can distinguish
good from bad, and can feel remorse for their
actions.
●
Somatic Marker Hypothesis
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Watch the ted
talk
Summary
1. Executive control processes play a supervisory role
in cognitive functions, allowing behavior to be modified
flexibly in a given environmental context.
2. Typical executive functions include working memory,
inhibition, task switching, simulation of behavior
consequences, and reward evaluation.
3. Specific behavioral tests have been developed
experimentally to test different domains of executive
control, but many involve multiple component processes.
4. The prefrontal cortex has been implicated to play a
critical role in executive control because of its strategic
anatomical position.
Courtesy
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5. In general, the lateral PFC facilitates cognitive
control processes such as initiation, selection,
inhibition, simulation, and abstraction of novel
rules for behavior, whereas the ventromedial PFC
facilitates behavioral control in situations with wellestablished behavioral rules, as in social situations
or in interaction with objects in the environment.
6. Damage to the lateral PFC is associated with
dysexecutive syndromes, whereas damage to the
ventromedial frontal cortex is associated with
disinhibition syndromes such as acquired
sociopathy
Psy355 Human Brain Function
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You should be able to answer the following:
What behavioral situations may require control processes
What evidences shown at the neuronal level that are relevant for EF
●
What are the main cognitive domains of executive functions
●
Briefly describe some early evidence for the frontal lobe EF hypothesis
●
What cognitive deficits do patients with frontal lobe damage most
commonly experience
●
According to the evolutionary studies of PFC, does brain size explain
cognitive capabilities between mammals? Why and Why not
●
Give a couple examples of cognitive tasks used for testing each EF
●
Describe disorders associated with the PFC
●
Compare and contrast effects of lateral PFC damage vs ventromedial
PFC damage
●
●
Practice what you have learned by doing the chapter 15 quiz
Unit 4: Cognitive Control of Action
and Behavior
Our goal is to form a basic understanding on the neural
basis of cognitive control. Perception → Action
Part 1. From perception to action – cortical and
subcortical neural circuits (4th ed. Ch 10; bb)
Part 2. The executive brain: (a) frontal lobe
hypothesis, EF tasks, EF disorders; (b) EF
anatomy, PFC functional divisions (4th ed. Ch
15; bb)
Part 3. Motivation, reward, and decision making (bb)
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The executive brain – Part 2b
Learning Objectives:
Recognize and identify the key brain regions that are parts
of the functional anatomy of executive control
Describe the unique connections to PFC and identify the
different subdivisions of PFC
Recognize the functional organization hypotheses of
executive control by comparing different brain areas and
their functions
Compare and contrast the roles of PFC areas, posterior
parietal, and subcortical circuits in executive functions
Relate brain functions to the cognitive components of EF
Refer to the lecture video and
the textbook chapter
●
●
●
●
●
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Part 2b. Functional Organization of
Executive Functions
A. Functional Anatomy of
Executive Control
A. Functional Anatomy of Executive Control
B. “Multiple Demand Network”
C. Functional Organization Hypotheses
1. Orbital/Ventromedial vs. Lateral PFC
2. Left vs. Right Lateral PFC
3. Posterior vs. Anterior Lateral PFC
4. Anterior Cingulate vs. Lateral PFC
1. Brain regions involved in
cognitive control:
o
Lateral (Dorsolateral &
Ventrolateral) PFC
o
Ventromedial PFC & OFC
o
Anterior Cingulate Cortex
o
Posterior Parietal Cortex
o
Basal Ganglia
Figure 15.1 Adapted
from Fuster (1989)
Figure 15.2 DLPFC, VLPFC,
anterior PFC,ACC, pre-SMA, OFC
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Connections to the PFC (cont.)
2. Connections to the PFC
(i) Projections to the PFC (direct inputs):
The prefrontal cortex
has numerous,
diverse, and
bidirectional
connections with
other many brain
regions.
o
o
o
o
o
Thalamus: the largest input to the PFC (primarily from
mediodorsal nucleus)
Secondary sensory/motor or association cortices
– i.e., not directly from primary motor/sensory
EXCEPTION: The OFC receives inputs from primary
olfactory, gustatory, and somatosensory cortices.
Posterior parietal cortex
Limbic regions: hippocampus, amygdala, ventral
tegmental area (VTA)
Adapted from Miller & Cohen (2001)
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Connections to the PFC (cont.)
(ii) The PFC sends projections back (direct
outputs) to many of the same regions from
which it receives inputs. Besides,
o
o
It is the only cortical region directly connected to
the hypothalamus
It sends direct outputs to the basal ganglia,
though inputs from the basal ganglia are
indirect
3. PFC’s close allies
(i) Posterior Parietal cortex (PPC)
plays a critical role in the allocation This material was produced for the sole use of s
of attention
o
co-activated with the lateral PFC
during executive control tasks
o
closely interconnected with the
PFC
o
Early evidence – Mountcastle’s
work in awake behaving monkeys
PPC neurons respond to the behavioral
responses associated with the stimuli rather the
stimuli per se
Attention vs. intention – probably both
o
o
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PFC’s close allies (cont.)
PFC’s close allies: Posterior Parietal cortex (PPC) (cont.)
(ii) Basal Ganglia
Current hypothesis:
o
Parietal: creating and updating a set of
stimulus-response associations
o
Prefrontal: formulating an appropriate plan of
behavior in a given context
o
o
o
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PFC’s close allies: Basal Ganglia (cont.)
fMRI and lesion studies: Basal
ganglia are important for forming
behaviorally relevant
associations between stimuli and
responses and the future
implementation of those
associations
o
Reward-based learning:
differential involvement of basal
ganglia and prefrontal systems
during a task requiring object
switch, rule switch, no switch, or
switch both (Cools et al. 2004)
o
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Receive extensive inputs
from the frontal lobes
Send extensive
projections to the
thalamus and cortical
areas
Important for motor (and
cognitive) control – e.g.,
Parkinson’s disease
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B. “Multiple Demand Network”
(Duncan, 2010)
Figure 15.16 Adapted
from Ducan (2010)
Lateral PFC, parietal, and ACC are involved in
non-automatic tasks
(i) More active during conditions requiring
executive function or solving new problems
(ii) PFC damage impairs both but doesn’t impact
previously learned knowledge
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(iii) ‘Multiple Demand Network’ – illustration
Fluid Intelligence
(e.g. Raven’s Test)
C. Functional Organization Hypotheses
Crystallized Intelligence (e.g. WAIS)
• Information: “What is the capital of
France?“
• Comprehension: “What is the thing
to do if you find an injured person
laying on the sidewalk?“
• Similarities: “How are a snake and
an alligator alike?“
• Vocabulary: “What is the meaning
of the word ‘articulate’?“
• Arithmetic: “John bought three
books for five dollars each and paid
ten percent sales tax. How much did
he pay all together?”
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1. Orbital/ventro-medial
vs. Lateral PFC
(Emotion control vs.
Cognitive Control)
2. Left vs. Right Lateral
PFC
3. Posterior vs. anterior
lateral PFC
4. Anterior cingulate vs.
lateral PFC (monitoring
vs. control)
Figure 15.2
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B. Functional Organization Hypotheses
1. Orbitofrontal/Ventromedial frontal cortex
vs. Lateral PFC (“hot” vs. “cold” control) Figure
i) Switching rewards vs.
Switching dimensions in
marmosets (Dias et al. 1996):
• Similar to WCST, with
compound stimuli made of
lines and shapes
• OFC lesions: poor reversal
learning
• Lateral PFC lesions: poor
shifting between dimensions
15.13
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1. Orbitofrontal/Ventromedial vs Lateral PFC (cont.)
ii) The Somatic Marker Hypothesis (Damásio 1996):
1. Emotional value of events (e.g. a risky situation)
are store as “somatic markers”
2. Somatic markers are assumed to be “stored” in
orbitofrontal and ventromedial frontal cortex
3. Retrieving that event reinstates the somatic
marker, which guides subsequent behavior
4. Somatic markers may be unconscious or
conscious
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1. Orbitofrontal/Ventromedial vs Lateral PFC (cont.)
1. Orbitofrontal/Ventromedial vs Lateral PFC (cont.)
A task of the somatic marker hypothesis
Figure 15.14
The gambling task has been criticized
o
deficit might relate to reversal learning since
the bad decks initially came with a larger reward
(e.g., Maia and McClelland, 2004)
o
the same patients are unimpaired if this first trial
was omitted (Fellows and Farah, 2003)
o
Others found a link between failure on reversal
learning and poor regulation of social behavior
(Hornak et al. 2004)
o
Delay discounting (and WM) seems to better
differentiates the OFC and lateral PFC functions
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Iowa Gambling task
(Bechara et al., 1994)
Patients with OFC damage
fail to show anticipatory SCR
to the risky cards, but show
SCR in other circumstances
(e.g. after loss)
Double dissociation with
working memory deficits and
‘cold’ tests of EF (lateral PFC
damage)
C. Functional Organization Hypotheses
1. Orbital/ventro-medial
vs. Lateral PFC
(Emotion control vs.
Cognitive Control)
2. Left vs. Right Lateral
PFC
3. Posterior vs. anterior
lateral PFC
4. Anterior cingulate vs.
lateral PFC (monitoring
vs. control)
LEFT
• Task setting
Figure 15.2
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C. Functional Organization Hypotheses
1. Orbital/ventro-medial
vs. Lateral PFC
(Emotion control vs.
Cognitive Control)
2. Left vs. Right Lateral
PFC
3. Posterior vs. anterior
lateral PFC
4. Anterior cingulate vs.
lateral PFC (monitoring
vs. control)
2. Left vs. Right Lateral PFC
Stuss et al. 1995
• E.g. Tower of
London, Open-ended
problems
• Task switching –
much slower to
switch
RIGHT
• Task monitoring
• E.g., Keeping ‘on
task’, Instructed of
the rule
• Task switching –
more likely to revert
to previous rule
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
3. Posterior vs. Anterior Lateral PFC
Koechlin & Summerfield (2007)
i. The executive
system is hierarchical
• Organized from
posterior to anterior
ii. ‘Branching’ maintaining multiple
goals over time
(‘multitasking’)
Figure 15.2
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
3. Posterior vs. Anterior Lateral PFC (cont.)
iii. Anterior PFC (BA10) shows increased
activation when holding a goal in mind whilst
simultaneously performing a sub-goal (Koechlin et
al. 1999)
• Damage to this region impairs multi-tasking
but not other tests of executive function (Roca
et al. 2011)
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
Figure 15.18
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
C. Functional Organization Hypotheses
1. Orbital/ventro-medial
vs. Lateral PFC
(Emotion control vs.
Cognitive Control)
2. Left vs. Right Lateral
PFC
3. Posterior vs. anterior
lateral PFC
4. Anterior cingulate vs.
lateral PFC (monitoring
vs. control)
Figure 15.2
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
4. Anterior Cingulate (ACC) vs. Lateral PFC
i.
Lateral prefrontal cortex
more associated with
task-switching, working
memory, and control
processes
4. Anterior Cingulate (ACC) (cont.)
iii. Conflict-monitoring model:
• Initially demonstrated using the Stroop task
• Response conflict detected → Control processes
are recruited to resolve conflicts
• Neuroimaging evidence: greater ACC activation
on incongruent trials, and greater ACC activity is
associated with greater lateral PFC activity on
the subsequent congruent trial
ii. ACC more associated
with engaging and
disengaging control
processes
Figure 15.20
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
4. Anterior Cingulate (ACC) (cont.)
Summary on functional organization of EF
iv. Error and response conflict detection
• Monkeys with ACC lesion don’t trouble shoot
after making an error (error+1 trial worse than
correct+1 trial)
• ERP: Error-related negativity
• fMRI: increased ACC activity on error trial
Figure 15.21
whereas lateral PFC
activity greatest on
error+1 trials
➢
ACC “detects” but
doesn’t correct
errors or conflicts
1.Orbital/ventro-medial vs. Lateral PFC
2.Left vs. Right Lateral PFC
3.Posterior vs. anterior lateral PFC
4.Anterior cingulate vs. lateral PFC
5.PFC’s close allies: posterior parietal and basal
ganglia
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
Summary
1. The PFC receives projections from many sensory and motor
association areas and sends projections back to most of the
same regions.
2. The OFC and ventromedial PFC have strong connections with
regions involved in processing emotions; damages to these areas
would affect conditions involving changes in emotional
value/state.
2. The lateral PFC has strong connections to sensory and motor
association areas; damages to these areas would affect
conditions involving changes in task-relevant stimulus features or
action planning.
3. There is evidence of a posterior-to-anterior organization of
executive functions with the anterior most part of the PFC
implicated in multi-tasking.
Review the text and
carefully evaluate the
evidence and theories to
understand these
different notions of
functional organization
Summary
5. Other regions such as the posterior parietal, anterior
cingulate cortex and basal ganglia are also involved in
supporting executive control processes.
• The anterior cingulate cortex monitors behavior and
signals the need for increase allocation of cognitive
control
• The posterior parietal cortex is implicated in allocating
attention and updating stimulus-response associations
• The basal ganglia initiate and gate at least some cognitive
control functions and motor acts
4. Left and right lateral PFC also show differential involvement in
cognitive control (e.g., task-setting vs. task-monitoring).
6. Others proposed a general framework of multiple demand
network, with a group of regions (PFC, ACC, and PPC) that
are commonly active during cognitive control of behavior.
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
This material was produced for the sole use of students at Stony Brook University currently
enrolled in PSY355 as per the TEACH Act Copyright © 2020 Hoi-Chung Leung. All rights reserved.
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