Chapter 12 – Cognitive Control

Part 2

B&C3: Cognitive Neuropsychology

Hanneke den Ouden
103 Overview

§ Introduction
§ lessons from history on thinking about control
§ central issues
§ anatomy of the frontal lobe
§ deficits of cognitive control

§ Aspects of cognitive control

§ maintain your goal
§ define on your goal: value-based decision making
§ make a plan to get to your goal
§ stay on track: information selection / filtering
§ monitor your progress towards your goal
104 Overview

§ Introduction
§ lessons from history on thinking about control
§ central issues
§ anatomy of the frontal lobe
§ deficits of cognitive control

§ Aspects of cognitive control

§ maintain your goal
§ define on your goal: value-based decision making
§ make a plan to get to your goal
§ stay on track: information selection / filtering
§ monitor your progress towards your goal
Achieving your goal:
Goal planning
106 Plan of action

§ 3 components
§ develop subgoals
§ anticipate consequences
§ identify requirements to achieve goals
107 Plan of action

§ 3 components
§ develop subgoals
§ anticipate consequences
§ identify requirements to achieve goals

Goals are
hierarchical!
108 Hierarchical organization of prefrontal cortex: 3 axes
Anterior-posterior PFC
§ posterior: more concrete
§ anterior: more abstract
§ uniquely human lateral frontal pole

Ventral-dorsal PFC
§ ventral: maintenance
§ dorsal: manipulation

Lateral-medial PFC
§ lateral: environment / context
medial personal / self oriented
Neubert et al Neuron 2014
 What is the evidence for this
posterior-anterior hierarchy of abstraction?
110 Testing the hierarchy of abstraction
Task: learn to follow increasingly abstract rules

Badre & D’Esposito 2007

111 Testing the hierarchy of abstraction
Task: learn to follow increasingly abstract rules
A. Response task: If red, press left

Stimuli Rule

Badre & D’Esposito 2007

112 Testing the hierarchy of abstraction
Task: learn to follow increasingly abstract rules
A. Response task: If red, press left
B. Feature task: if red, then look out for
the triangle. If present, press ‘left’

Badre & D’Esposito 2007

113 Testing the hierarchy of abstraction
Task: learn to follow increasingly abstract rules
A. Response task: If red, press left
B. Feature task: if red, then look out for
the triangle. If present, press ‘left’
C. Dimension task: If red, then check the
shape of the 2 stimuli. If they match,
then press ‘left’. If blue, then check the
size of the 2 stimuli… etc

Badre & D’Esposito 2007

114 Testing the hierarchy of abstraction
Task: learn to follow increasingly abstract rules
A. Response task: If red, press left
B. Feature task: if red, then look out for
the triangle. If present, press ‘left’
C. Dimension task: If red, then check the
shape of the 2 stimuli. If they match,
then press ‘left’. If blue, then check the
size of the 2 stimuli… etc
D. Dimension task II: On in the odd task
block numbers, do task C with ‘red-
shape’ and ‘blue-size’, on even blocks
the reverse

Badre & D’Esposito 2007

115 Testing the hierarchy of abstraction
Increasingly abstract rules…. ….Increasingly anterior representation
A. Response task: If red, press left
B. Feature task: if red, then look out for
the triangle. If present, press ‘left’
C. Dimension task: If red, then check the
shape of the 2 stimuli. If they match,
then press ‘left’. If blue, then check the
size of the 2 stimuli… etc
D. Dimension task II: On in the odd task
block numbers, do task C with ‘red-
shape’ and ‘blue-size’, on even blocks
the reverse

Badre & D’Esposito 2007

116 Hierarchy of abstraction
Increasingly abstract rules…. ….Increasingly anterior representation
A. Response task: If red, press left
B. Feature task: if red, then look out for
the triangle. If present, press ‘left’
C. Dimension task: If red, then check the
shape of the 2 stimuli. If they match, ns
then press ‘left’. If blue, then check the sio
size of the 2 stimuli… etc Le
D. Dimension task II: On in the odd task
block numbers, do task C with ‘red-
shape’ and ‘blue-size’, on even blocks
the reverse

Badre & D’Esposito 2007

Why is representing the hierarchy of goals important in real life?
118 Hierarchy of goals in real life
§ Task: Help a family to live on a smaller budget
119 Hierarchy of goals in real life
§ Task: Help a family to live on a smaller budget

no PFC lesion

evaluate expenditures
(need to eat, place to sleep)

remove non-crucial See if crucial expenditures

expenditures can be reduced
120 Hierarchy of goals in real life
§ Task: Help a family to live on a smaller budget

no PFC lesion PFC lesion patient

evaluate expenditures find biggest expenditures

(need to eat, place to sleep) (rent)

remove non-crucial See if crucial expenditures

expenditures can be reduced
121 Hierarchy of goals in real life
§ Task: Help a family to live on a smaller budget

no PFC lesion PFC lesion patient

evaluate expenditures find biggest expenditures

(need to eat, place to sleep) (rent)

remove non-crucial See if crucial expenditures move out

expenditures can be reduced (and buy a tent)
Implicit goals Stay alive and
happy

Stay healthy

Live on a smaller
Have a home
budget

Move out and live in a tent

123 Hierarchy of goals in real life
§ Task: Help a family to live on a smaller budget

Healthy control PFC lesion patient

evaluate expenditures find biggest expenditures

(need to eat, place to sleep) (rent)

remove non-crucial See if crucial expenditures move out

expenditures can be reduced (and buy a tent)

ve rate
e
pers
Hierarchical goal planning and definition of
appropriate sub-goals is disrupted
in PFC lesion patients
125 Control over complex actions require that we

§ maintain our current goal (reduce expenditure)

§ be able to shift from one subgoal to another (e.g. saving money, but
also having a place to live)

But also

§ focus on the information that is relevant to achieving that goal and

ignore irrelevant information
126 Overview

§ Introduction
§ lessons from history on thinking about control
§ central issues
§ anatomy of the frontal lobe
§ deficits of cognitive control

§ Aspects of cognitive control

§ maintain your goal
§ define on your goal: value-based decision making
§ make a plan to get to your goal
§ stay on track: information selection / filtering
§ monitor your progress towards your goal
127 Dynamic Filtering - Shimamura

§ Prefrontal cortex as dynamic filter

§ Not just maintenance

§ Goals modify the salience of information

§ actively select information
128 Dynamic Filtering - Shimamura

§ Prefrontal cortex as dynamic filter

§ Not just maintenance

§ Goals modify the salience of information

§ actively select information

§ Problems: interference (e.g. Stroop)

§ PFC patients have problems
§ PFC is important for filtering (not just WM)
129 Think of a verb associated with this object

§ Menti code: 88 37 21 9
131 Filtering Demands - Thompson-Schill et al

§ Semantic generation
§ High filtering: numerous associates
§ Requires selection

low filtering high filtering

132 Filtering Demands - Thompson-Schill et al

§ Semantic generation
§ High filtering: numerous associates
§ Requires selection

high > low filtering:

(left) inferior frontal cortex
PFC is important for filtering (select/ignore) information as/when it
becomes important

So what happens when you are not very good at filtering?

When is this a benefit?
PFC is important for filtering (select/ignore) information as/when it
becomes important

Hypothesis:

Impaired selection process in PFC lesions leads to openness to

creative/ atypical solutions
135 Openness to atypical solutions in PFC lesions
Match stick task
136 Openness to atypical solutions in PFC lesions
Match stick task
137 Openness to atypical solutions in PFC lesions
Match stick task
138 Openness to atypical solutions in PFC lesions
Match stick task
139 Openness to atypical solutions in PFC lesions
Match stick task
140 Openness to atypical solutions in PFC lesions
Match stick task

Double
Dissociation!

one hipothesys
another hypothesis
Impaired selection process in PFC lesions leads to
openness to creative/ atypical solutions

… an immature PFC in childhood & adolescence

is perhaps an advantage when you are learning
about the world!
So how does the PFC select information?

Input or output filtering?

Inhibition or facilitation of information?
143 Testing sensory input filtering in frontal lesions

§ Unattended auditory clicks

§ evoked potential (EP) measured in auditory
cortex of lesion patients

Knight et al (1995)
144 Testing sensory input filtering in frontal lesions

§ Unattended auditory clicks

§ Measure evoked potential (EP) in auditory
cortex

Lesion patients:
§ Parietal: no difference

Knight et al (1995)
145 Testing sensory input filtering in frontal lesions

§ Unattended auditory clicks

§ Measure evoked potential (EP) in auditory
cortex

Lesion patients:
§ Parietal: no difference

§ Temporal lobe lesion: loss of information

Knight et al (1995)
146 Testing sensory input filtering in frontal lesions

§ Unattended auditory clicks

§ Measure evoked potential (EP) in auditory
cortex

Lesion patients:
§ Parietal: no difference

§ Temporal lobe lesion: loss of information

§ Frontal lobe lesion: loss of inhibition of

auditory cortex!

Knight et al (1995)
Frontal lobe minimises impact of irrelevant perceptual information

Prediction:
When we minimise interfering information,
this should help frontal lesion patients.
148 Inhibition of sensory input to suppress WM content interference

PFC lesion: Interference

149 Inhibition of sensory input to suppress WM content interference
150 Inhibition of sensory input to suppress WM content interference

PFC lesion:
Improved performance!
151 Inhibition of task-irrelevant sensory input helps goal-directed control

§ PFC lesion: patients show larger response to

irrelevant auditory stimuli and absence of
differentiation for attention

§ PFC lesion: monkeys perform fine when no

interfering visual stimululation is present during
delay
So how does the PFC select information?

Input or output filtering?

Inhibition or facilitation of information?
So how does the PFC select information?

Input or output filtering?

Inhibition or facilitation of information?
154 Output filtering: inhibition (or selection) of action

§ hold your horses, time to think

§ opposite of action selection? or part of it?
155 Inhibition of action
156 Testing inhibition of action: Stop signal task
Go Trial (75%)

à Go

Go RT
157 Testing inhibition of action: Stop signal task
Go Trial (75%)

à Go

Go RT

Stop Trial (25%)

STOP!

Succesful stop
à
Stop Signal Delay
158 Testing inhibition of action: Stop signal task
Go Trial (75%)

à Go

Go RT

Stop Trial (25%)

STOP!
Variable time
Succesful stop between onset
à of stimulus
and stop signal
Failed stop (SSD)
Stop Signal Delay
159 Inhibition action: Stop Signal Task (fMRI)

STOP!

à
Stop Signal Delay
160 Inhibition action: Stop Signal Task (fMRI)

§ Right inferior frontal gyrus lesion patients

STOP!

à
Stop Signal Delay
Signals of the brain
161 Inhibition action: Stop Signal Task (fMRI)

§ Right inferior frontal gyrus lesion patients are slower to abort

§ have a longer SSRT (= stopping process)
§ i.e. fail with shorter SSDs

STOP!

à
Stop Signal Delay
 After 4 lectures by Hanneke, you now all shout in concert…:

“Ok, so lesioning the left IFG reduces your ability to inhibit action,
but what is the IFG actually doing when it is intact?”
163 Frontal cortex for stopping action

Go
Succesful stop
Failed stop (Go)
164 Frontal cortex for stopping action

Should stop

Go
Succesful stop
Failed stop (Go)

Should go
165
Hmm, so IFG activity does not tell us why you failed

Should stop

Go
Succesful stop
Failed stop (Go)

Should go
166 Inhibition action: Stop Signal Task (fMRI)

§ Motor cortex
§ failed stop: early high activation
§ when IFG couldn’t stop you anymore

Go
Succesful stop
Failed stop (Go)
167 Stopping network

§ White matter tract tracing (DTI) from IFC reveals

‘stopping network’
168 Stopping network

§ White matter tract tracing (DTI) from IFC reveals

‘stopping network’

§ Proposed division of labour:

§ Motor cortex implements action
§ Right IFC generates command to stop
§ Subthalamic nucleus: implements command – tells the
whole (pre)motor cortex to stop

§ (next, an action can be selectively released)

Wessel & Aron 2018 Neuron

169 Subthalamic nucleus: implementation of global stop?
§ Parkinson’s Disease patients treated with DBS of STN

§ DBS inhibits STN à disinhibits cortex

170 Subthalamic nucleus: implementation of global stop?
§ Parkinson’s Disease patients treated with DBS of STN

§ DBS inhibits STN à disinhibits cortex

§ … motor disinhibition: improves movement

initiation

§ … cognitive disinhibition: loss of impulse control on

reward task!
171 Inhibition

§ ‘Organisation’ of inhibition implemented by (right) inferior frontal gyrus

§ Execution by subthalamic nucleus
§ which then globally suppresses the cortex

§ Inhibition of action important for controlling impulsive behaviour

§ Compromised in addiction (impulse control)
172 Overview

§ Introduction
§ lessons from history on thinking about control
§ central issues
§ anatomy of the frontal lobe
§ deficits of cognitive control

§ Aspects of cognitive control

§ maintain your goal
§ define on your goal: value-based decision making
§ make a plan to get to your goal
§ stay on track: information selection / filtering
§ monitor your progress towards your goal
Monitoring goal achievement
174 Ensuring that goal-oriented behavior succeeds

§ Monitoring system in the medial frontal cortex (ACC)

§ Novel responses
§ Required response competes with
strong habitual response
§ Error correction
§ Difficult or dangerous situations
175 Monitoring in ACC
Clusters in ACC, but extends also beyond and therefore: medial frontal cortex
§ nice central location!
ACC Connectivity: In a good place to monitor!
What does the ACC do?
Many hypotheses….
178 Function of the ACC: hypotheses

1. Attentional hierarchy: coordinate & select different WM buffers

§ problem: homunculus

2. Error Detection: ERP signal in ACC when making an error

§ problem: also active for difficult tasks when you don’t make an error (e.g. Stroop)

3. Conflict Monitoring

4. Cost of Control
186 3. Conflict monitoring hypothesis

§ Response Conflict
§ Lateral cortex represents task goals
§ Medial frontal cortex monitors if that goal is achieved (not just errors)
§ Conflict: allocate resources…
§ especially when the automatic response is (likely) incorrect
187 Stroop task
§ Cue – long delay – stimulus
§ Measures: task selection vs response conflict
§ Difficulty: reading (easy) vs color naming (hard)
188 Stroop task
§ Cue – long delay – stimulus
§ Measures: task selection vs response conflict
§ Difficulty: reading (easy) vs color naming (hard)
§ Congruency: congruent vs incongruent
189 Stroop task: double dissociation ACC vs DLPFC
§ Instruction:

§ Stimulus:
190 Stroop task: double dissociation ACC vs DLPFC
§ Instruction: PFC as a function of task difficulty:
hard (name color) > easy (name word)

§ Stimulus:
191 Stroop task: double dissociation ACC vs DLPFC
§ Instruction: PFC as a function of task difficulty:
hard (name color) > easy (name word)

§ Stimulus: ACC as a function of response conflict

(incongruent > congruent).
192 Stroop task: double dissociation ACC vs DLPFC
§ Conflict registered in ACC leads to higher activation
on subsequent trial in DLPFC
193 Stroop task: double dissociation ACC vs DLPFC
§ Conflict registered in ACC leads to higher activation
on subsequent trial in DLPFC
§ Adaptation of task goal/setting
194 Conflict monitoring hypothesis

§ Medial frontal cortex detects conflict and raises alarm (pay attention, things are
harder than we thought!)

§ Lateral cortex represents & adapts task goals: implements ‘control’

Problem
§ ACC activity seems more to reflect likelihood of errors than degree of conflict
195 4. Cost –Benefit analysis of Control hypothesis (Shenhav e.a. 2013)

§ Stroop task:
§ very cognitively ‘costly’ to
perform the colour naming

§ So how much will I benefit if I

ramp up the control?

ued…
ntin
e co
to b
196 Summary overview
Lateral prefrontal cortex
§ working memory
§ filtering / inhibition
§ selective attention (activate posterior regions)

Ventromedial /orbitofrontal cortex

§ value coding
§ coming up: emotion, social cognition

Frontal pole
§ abstract rule learning
§ hierarchical action goals
§ reasoning

Medial frontal cortex / ACC

§ monitoring ongoing activity (modulate the degree of cognitive
control)
Cognitive control
Question & Answer session
198 Marshmallow test – who does what?
199 Marshmallow test – who does what?
§ https://www.menti.com/83568743

§ LPFC, ACC (= MPFC), VMPFC, frontopolar cortex…

§ Who does what? Match them together
§ is monitoring how hard this is: keep an eye on the prize!
Make sure to not start eating the marshmallow
§ makes the action plan: to get 2 marshmallows, i need to not
eat it now, sing a song, look away
§ computes how much you like eating marshmallows. And how
much you prefer it now over later. But also that you prefer 2
over 1
200 Lateral Prefrontal Cortex

§ Working Memory
§ Inhibition of Prepotent Responses
§ Selective Attention
201 Frontal Pole

§ Hierarchical Representation of Action

Goals
§ Abstract rule formation
§ Memory Retrieval
202 Medial Frontal Cortex

§ Error Detection / monitoring

§ Resolving Conflict
§ a.k.a. Anterior Cingulate Cortex
203 Ventro-Medial Prefrontal cortex

§ Value computation
§ a.k.a. orbitofrontal cortex

§ à see upcoming lectures

204 Control, anatomy, deficit (518/519)

§ Cognitive control is the collection of mental abilities that involve planning,

controlling, and regulating the flow of information processing.
§ Cognitive control gives us the flexibility required for goal-oriented behaviour
§ Prefrontal cortex includes four major components: lateral prefrontal cortex,
frontal pole, medial frontal cortex, and orbitofrontal cortex. These parts of the
brain have become very prominent in mammals, and especially in primates
§ Patients with frontal lobe lesions have difficulty executing a plan and may
exhibit stimulus-driven behaviour
§ Deficits in cognitive control are found in numerous psychiatric disorders as well
as when mental healthy is compromised by situational factors such as stress or
loneliness
205 Goal oriented behaviour (p526)

§ Working memory can be conceptualized as the information formed by the

combination of a task goal and the perceptual and long-term knowledge
relevant for achieving that goal. This form of dynamic memory emerges from
the interaction of prefrontal cortex and the rest of the brain.

§ Neurons in the PFC of monkeys show sustained activity throughout the delay
period in delayed-response tasks. These cells provide a neural correlate for
keeping a representation active after the triggering stimulus is longer visible

§ Various frameworks have been proposed to uncover function specialization

within the prefrontal cortex. Three gradients have been described to account for
PFD processing differences: ventral-dorsal, anterior-posterior, and lateral-
medial.
206 Decision Making (p 539)

§ A decision involves the selection of one option among several. It typically

involves an evaluation of the expected outcome (reward) associated with each
option.

§ The subjective value of an item is made up of multiple variables that include

payoff amount, context, probability, effort/cost, temporal discounting, novelty,
and preference.

§ Single-cell recordings in monkeys and fMRI studies in humans have implicated

fontal regions, including the orbitofrontal cortex, in value representation.
207 Decision Making - Dopamine neurons (p 539)
§ Reward prediction error (RPE) is the difference between the expected reward
and what is actually obtained. The RPE is used as a learning signal to update
value information as expectancies and the valence of rewards change. The
activity of some DA neurons provides a neuronal code of prediction errors.

§ DA neurons also appear to code other variables that may be important for goal-
oriented behaviour and decision making, such as signaling the salience of
information in the environment.
208 Goal Planning (p544)
§ Successful execution of an action plan involves three components: (1)
identifying the goal and developing subgoals, (2) anticipating consequences
when choosing among goals, and (3) determining what is required to achieve
the goals.

§ Goal-oriented behaviour requires the retrieval and selection of task-relevant

information. The PFC can be conceptualised as a dynamic filtering mechanism
through which the task-relevant information is activated and maintained in
working memory
209 Goal Planning II (p544)
§ Cognitive control is also essential when we need to maintain multiple goals at
the same time, especially when those goals are unrelated. the bike and the bridge

§ Through the selection of task-relevant information the prefrontal cortex helps to

make action selection more efficient. This benefit of using experience to guide
action selection may also come at a cost in terms of considering novel ways to
act, given a specific situation.
210 Mechanisms of goal-based selection (p 553)

§ Goal-oriented behavior involves the amplification of task relevant information

and the inhibition of task-irrelevant information. Amplification and inhibition
may entail separate processes given that aging selectively affects the ability to
inhibit task-irrelevant information.

§ Patients with prefrontal cortex damage lose inhibitory control. For example,
they cannot inhibit task-irrelevant information.
211 Mechanisms of goal-based selection II (p 553)

§ A network spanning prefrontal cortex and posterior cortex provides the neural
substrates for interactions between goal representations and perceptual
information.

§ The inhibition of action constitutes another form of cognitive control. The right
inferior frontal gyrus and the subthalamic nucleus are important for this form of
control.

§ Active video game playing has been hypothesised to improve some aspects of
cognitive function, such as task switching, perhaps because games require
coordinating multiple subgoals
212 Monitoring (p563)

§ The medial frontal cortex, including the anterior cingulate cortex (ACC) is
thought to be a critical part of a monitoring system, identifying situations in
which cognitive control is required.

§ The error-related negativity or feedback-related negativity (ERN or ERF) signals

are event related potentials that occur when unexpected feedback is produced.
This response is generated by the medial frontal cortex.

§ The medial frontal cortex is engaged when response conflict is high. Through its
interactions with lateral regions of the prefrontal cortex, a monitoring system
can regulate the level of cognitive control.
reserve slides