Chapter 11: Executive Functions

• Executive Functions include the ability to plan actions to reach a goal, to use information
flexibly, to think abstractly, and to make inferences.
o Because of the multifaceted nature of these executive functions, more than one
function usually contributes to performance of many of the complex tasks discussed
in this chapter.
o Executive functions rely heavily on the frontal lobe, but executive deficits can occur
after posterior damage or damage to white-matter tracts connecting the frontal
regions to other brain areas (interacting network of brain structures) as well.
• Controlled versus Automatic Process (two-component system):
o Contention scheduling = automatic processing: Stimuli or situations become linked to
actions, routines, or processing schemes, and then groups of these routines become
linked to one another. In this manner, a single stimulus may result in a relatively
automatic string of actions, referred to as a schema.
o Supervisory attentional system = direct attention and guide action trough decision
processes. Active when no preexisting processing schemes are available, as in novel
situations or when typical responses must be overridden or inhibited
1. Frontal lobe damage disables the supervisory attentional system and thereby
leaves actions to be governed totally by contention scheduling
è people with frontal lobe damage fail to respond appropriately to
novel situations, show environmental dependency syndrome or
exhibit perseveration
o Alternative Modell, hierarchical
i. Lowest level of control: automatic processing of sensory information;
posterior regions
ii. Next level of control: processing of sensory information in term of guiding
the behaviour towards a goal; frontal lobe (executive functions)
iii. Highest level of control: self-reflection and metacognition; prefrontal cortex
• Goal-Centered Processing:
o Artificial intelligence approach: Reaching a goal requires the creation of a hierarchy
of simpler and more solvable subgoals. Then for each subgoal, the steps required to
achieve those subgoals must be determined and specified. It has been argued that
frontal lobe damage disrupts the ability to create such a hierarchical goal list
• Multifactor Models: executive function may consist of separable facets or factors
o Unity and diversity Model: executive function consists of one common factor (e.g.
maintain task goals) required for all executive function, as well as more specialized
functions (task switching, updating information) that are only required in specific
situations
1. Factors are linked to different brain regions in the frontal cortex

Goal-Directed Behaviors
• Initiation of Behavior:
o Patients with executive dysfunction show psychological inertia (= tendency of a body
at rest to stay at rest or a body in motion to stay in motion unless acted upon by an
outside force); difficulties starting an action or a behavior, but once engaged in it,
they have great difficulty stopping it
 o reduction in spontaneous speech
1. disturbed cost-benefit analysis of actions (after damage to anterior cingulate
cortex); difficulty in initiating behavior is because the goal or potential
outcome does not seem worth the effort
• Creation and Maintenance of o Goal or Task Set
o Patients with frontal lobe damage wander off task; engaging in irrelevant activity
o Dorsolateral prefrontal regions aid in creating and maintaining an attentional set (=
process that designates which information is task-relevant)
1. prefrontal areas helps to stay on task, especially when irrelevant information
is particularly distracting
2. other regions than lateral prefrontal cortex might be involved in maintaining
an attentional set as well, e.g. anterior cingulate, the frontal operculum and
anterior insula
o frontopolar cortex involved in maintaining more than one task set at time
1. damage to frontopolar cortex results in impaired management of multiple
goals
o more activity in dorsolateral prefrontal cortex (DLPFC) and n portions of the anterior
cingulate cortex when people choose what task they will perform versus when
instructed; frontopolar regions might be involved in selecting potential task sets as
well
• Sequencing and Planning
o Anterior regions of the brain are important for sequencing movements and thoughts
o People with frontal lobe damage have difficulty in sequencing and planning (assessed
with the self-ordered pointing task à DLPFC plays an important role)
1. DLPFC may play a specific role because planning requires that information in
working memory is continually re-ordered and updated
o ventral regions of lateral prefrontal cortex (i.e., Broca’s area) appear to play an
important role in aspects of language that involve sequencing, such as syntax
o Broca’s area responds to sequential processing demands of linguistic and musical
information
o Patients with frontal lobe damage are less likely to report that they use strategies,
and when they do use a strategy, it tends to be ill-defined or invoked inconsistently;
impaired in choosing which sequence or strategy best allows a goal to be attained
(assessed with the Tower of London Task à DLPFC and rostrolateral prefrontal
cortex play a role)
1. rostrolateral prefrontal cortex is involved in higher-order processing of
sequences to reach a goal
• Shifting Set and Modifying Strategies
o The classic neuropsychological test used to examine task-switching is the Wisconsin
Card Sorting Test (WCST); test is sensitive to frontal lobe damage.
o People with executive dysfunction perseverate, which is the action of continuing to
engage in the same behaviour; are impaired in modifying their strategies
o Typical task-switching-task consists of repeat trials (he person performs the same
task as on the prior trial) and switch trials (the person performs a different task
than on the prior trial)
1. response times are longer for switch trials than for repeat trials, known as
the switch cost (reflects the need to inhibit or overcome the prior task set,
sometimes referred to as task-set inertia, and the need to configure the
system for the current/new task set)
 2. Increasing the cue–stimulus interval (indicating which task has to be
performed in the next trial) reduces the switch cost, most likely because
the person has additional time to configure the new task set before that
task set must be used on the stimulus. Typically, however, even with long
delays, it is impossible to completely erase or eliminate the switch cost,
presumably reflecting task-set inertia.
o Patients with left frontal lobe damage have a specific deficit in task-switching,
especially when there are no strong or obvious cues as to which task should be
performed when à no specific brain region involved in task switching, more general
mechanism
1. BUT regions may vary depending of the type of switch and the degree of
similarity or dissimilarity between the task sets
• Self-Monitoring and Evaluation
o First, metacognitive awareness, that is, an overall evaluation of one’s performance, is
disrupted in patients with left or right frontal lesions.
o In addition, frontal damage, especially right frontal damage, impairs the ability to
detect errors during a sustained attention task and to modify ongoing behaviors to
take a corrective action.
o We have a particular set of brain mechanisms that helps to monitor our performance
and detect error: The error-related negativity (ERN) occurs approximately 100 ms
after an error has been made.
1. the larger the error, the larger the amplitude of the ERN
2. arises from rostral regions of the anterior cingulate, located on the medial
portion of the frontal lobe
3. ERN is just providing a rather undifferentiated signal that something is amiss
o Awareness of an error seems to be indexed by another component, the error
positivity (Pe), which frequently follows the ERN by about 200–300 ms
o Alternative views:
1. medial prefrontal cortex does not detect errors but monitors for conflict
(cingulate sends then a signal to dorsolateral prefrontal cortex to ramp up
the top-down control to reduce conflict, and hence errors, in subsequent
behavior)
2. medial prefrontal cortex determines whether exerting control is worth the
effort and cost in a given situation
3. cingulate is focused on determining whether the action leads to a useful or
good outcome; is sensitive to negative outcomes or the loss of rewards à
history of the outcomes of an individual’s recent behaviors can be used to
guide subsequent actions and responses so as to avoid future “errors”
o Influenced by peoples’ characteristics: high level of worry or anxiety tend to increase
ERN versus substance abuse or ADHS decrease ERN and Pe
o Anterior cingulate is not the sole “error detector” of the brain; Patients with
cingulate damage are able to report errors. Instead they typically show post-error
slowing, which is the phenomenon that people respond more slowly on the next trial
after an error.
o Patients with damage to lateral prefrontal cortex, can compromise monitoring and
evaluation, e.g. self-corrective actions.
• Inhibition:
o Disrupted inhibition leads to perseveration, one of the hallmarks of executive
dysfunction
 o Patients with frontal lobe damage perform poorly on tasks that require inhibitory
control, esp. when well linked association must be overridden
o Assessed with Go/No-Go tasks: In this task, the person responds by pushing a button
when certain visual stimuli appear (Go trials) and withholds response to other stimuli
(No-Go trials)
o Another task to assess inhibition control is the stop-signal task. In contrast to Go/No-
Go tasks, a person must cancel an ongoing response
o Both tasks activate a wide range of brain regions spanning (dorso)lateral prefrontal
cortex, the anterior cingulate, SMA, pre-SMA, insula, and parietal regions
o Response inhibition may be a more specific example of a more general function, that
of interference resolution, which is the ability to resolve conflict between competing
information or distracting information that might interfere with performing a task.

In Focus: Can You Inhibit a Memory?

• In the Think/No-Think task, individuals are taught cue-target pairs that are random associates
(such as the word pair “roach–ideal”) during a training phase. They are trained on the pairs
so that when given the cue, they can produce or identify the target with a high degree of
accuracy, ensuring that the memory exists. In the next phase, the experimental phase,
participants are presented with just the cue. For some cues, there is a designation (e.g., a
green box around the cue) indicating that the person should try to remember the target
associated with that cue. These are known as Think trials. For other cues, the designation
(e.g., a red box around the cue) indicates that the person should try to inhibit the associated
item from coming to consciousness. These are known as No-Think trials. In the final phase,
the test phase, performance is measured by giving the participants the cue and asking them
to produce the item that went with it.
• In a study performed by one of the co-authors of this book, we were able to provide
evidence that inhibition of memory can indeed occur. […] We used the Think/No-Think task
in our study, but unlike prior studies, we used face–picture pairs. […] Instead of asking
someone if they are re-remembering or inhibiting a memory, we could use activity in ventral
visual processing regions as a proxy for what they were doing! When people were thinking
about an item, we should observe an increase in activity compared to a baseline (in this case,
just a simple fixation cross on the screen); when they were not thinking about the item, we
should observe a decrease in activity below baseline in these regions. In fact, this is what we
found. […] At the same time, we could examine the degree to which there is activity in the
hippocampus, which plays an important role in memory retrieval. Therefore, if people are
really inhibiting the memory on No-Think trials, we should also observe decreases in activity
in the hippocampus in this condition compared to the fixation baseline. Likewise, if memories
are being retrieved on Think trials, then activity in the hippocampus should be greater
compared to the fixation baseline. This expectation was also met. Thus, we demonstrated
that a memory can indeed be inhibited.
• Regions responsible for inhibition spanning the right middle and inferior frontal gyrus;
prefrontal cortex can down-regulate activity in the hippocampus to inhibit the overall
process of memory retrieval (= No-Think trials)

Higher-Order Thinking
• Abstract and Conceptual Thinking:
o Executive dysfunction leads to the inability to process material in an abstract rather
than a concrete manner
o Frontal areas involved in the understanding the meaning of metaphoric sentences
and (verbal or visuospatial) analogy reasoning (e.g. “PLANET:SUN +
ELECTRON:NUCLEUS”)
• Rules and Inference
o Primate frontal cortex has neurons that can encode and categorize information in a
rule-like manner.
o Abstraction and rule-like understanding (can’t figure out the criterion by which the
cards should be sorted in the Wisconsin Card Sorting Test) is compromised in at least
some patients with frontal lobe damage.
o Different brain activity in frontal areas in rule implementation vs. rule discovery vs.
rule deduction
è Ventrolateral Prefrontal Cortex plays a role in retrieving stored knowledge
that allows the retrieval of rules
è DLPFC is more involved in selecting or influencing how rules should be used
to guide responding than in actually selecting the rules (alternative
viewpoints: plays a direct role in abstracting rules OR holds rules in working
memory)
è Left lateral Cortex plays a critical role in rule generation but not for rule
recognition
o “Ah-ha” moments during problem solving = insight (correct solution suddenly
popping into mind)
è Burst of Synchronized gamma activity (information from different brain
regions can be bound together to emerge into consciousness) from right
temporal areas and increased alpha activity (increased filtering out of
sensory information; keep sensory information from disrupting your mental
effort) prior to insight
o Absence of frontal cortex “interference” and control would free us from the
restriction provided by the rule-based behaviour, allowing and enhancing creativity
(in certain situations); thinking of novel and unusual uses for objects or making
remote associations
• Response to Novelty
o P3a occurs when a novel or unexpected stimulus captures attention […] We can be
relatively certain that frontal regions of the brain contribute to the generation of this
potential because (1) the P3a decreases in amplitude after lesions to prefrontal
cortex; (2) the amplitude of the P3a is correlated with the volume of gray matter in
the frontal lobes in neurologically intact men; and (3) high-density electrode arrays
suggest a frontal source
o People with executive dysfunction have trouble being cognitively flexible, e.g.
patients with frontal lobe damage cannot generate alternative plans of action.
o Orbitofrontal damage impedes the ability to exhibit normal reversal learning, in
which an individual reverses a previous response.
o Monkeys with damage to frontopolar cortex are less distracted than control
monkeys when faced with distractions or free rewards; their ability to reorient
potential task goals toward novel situations or opportunities is compromised
• Judgment and Decision Making
 o Patients with frontal lobe damage have difficulty with estimates (doesn’t rely on
stored memory), e.g. length of the spinal cord in women, prices of objects
o When people decide to stick with the ongoing strategy, activation is observed in
orbitofrontal regions (contributes to determining the subjective value of a stimulus).
However, when they decide to switch strategies, activation is observed bilaterally in
frontopolar cortex (responsible for selecting overall task goals; important for
changing strategies)
o Dorsolateral (prefrontal) regions may play an important role in overcoming yielding
to the temptation of a current reward

Organization of the Brain for Executive Function

• Nested-hierarchy Model
o One model posits a nested hierarchy of control processes within prefrontal cortex.
According to this model, there is a gradient from posterior to anterior, with more
posterior regions selecting sensory information, then regions that select with regard
to the context in which that sensory information occurs, then regions that select
information with regard to the current episode or event, and finally regions that
select with regard to the context of prior episodes or events.
• Another model argues that selection of material occurs in posterior regions of prefrontal
cortex on the basis of more concrete dimensions, and that the representations used for
selection become more abstract as one moves in an anterior direction. The more choices
(decision tree branches) are to make, the more anterior regions are needed.

• Another model views control as implemented in a cascade involving lateral and medial
prefrontal regions. Posterior regions of dorsolateral prefrontal cortex send a top-down signal
to bias activity toward those regions of posterior cortex that will be needed to process task-
relevant information, essentially imposing an attentional set. […] Posterior regions of the
dorsal anterior cingulate help to bias processing toward the use of certain information to
select a response and more anterior regions evaluate the appropriateness of the response.
Other models posit that when anterior cingulate activity is increased, it provides a signal back
to lateral prefrontal cortex to increase top-down control.
A Central Role for Working Memory in Executive Function

• How might difficulties in working memory account for some of the deficits in executive
functioning? If one cannot maintain information in working memory, one may not be able to
keep a goal in mind; thus, this deficit interferes with a person’s ability to direct behavior
toward a goal or to formulate a strategy for attaining the goal. In addition, difficulty in
keeping information on-line may disrupt a person’s understanding of temporal relations
between items and events à sequencing would be quite difficult. Moreover, if one is not
able to retain multiple pieces of information simultaneously in working memory, there will
be difficulty in creating or following rules, in making inferences, and/or in understanding the
relations between items. […] If one is unable to clear out what is held in working memory,
perseveration will result.
• Likewise, executive aspects of attentional control are required to determine what
information is selected to be maintained or manipulated in working memory, and which of
the contents currently in working memory are most relevant for current task demands.