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Executive processes Properties of executive processes
Executive processes coordinate mental activity so that a particular goal is achieved They do this by modulating the operation of other processes (modulate = to guide or modify); however, executive processes do not actually carry out these activities
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Executive processes Different types of executive processes
Several different types of executive processes have been identified. There is currently controversy about whether there are several types of executive processes or a single type (we are assuming several) 1. selective attention. (sometimes called executive attention (sustained attention) – this type of attention is believed to act on the contents of working memory and directs future mental activity in order to achieve a goal (e.g., driver focuses on traffic signs in order to take correct exit) 2. switching attention – switching attention from one activity to another (e.g., having a conversation and driving) Inhibiting information that has been perceived (e.g., ignoring music on radio when traffic is heavy and one is driving) Scheduling a sequence of activities – planning the order in which a activities will be carried out in order to achieve goal(s) (e.g., order in which you will pick up groceries, dry cleaning, medications) Performance monitoring – review performance to ensure it is acceptable (e.g., checking speed limit while driving)
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Executive processes It is generally agreed that the frontal lobes play an important role when executive functions are carried out The frontal executive hypothesis proposes that executive processes are primarily mediated by the prefrontal cortex (PFC) PFC – anterior to motor cortex (and for some authors the premotor cortex) PFC includes dorsolateral PFC, anterior cingulate, Broca’s area, and the medial and orbital regions of the frontal lobes
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Executive processes Frontal lobes:
Prefrontal cortex: massive network that links the motor, perceptual, and limbic (emotional network) regions Limbic system include amygdala, cingulate gyrus, orbitofrontal cortex and parts of basal ganglia
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Executive functions
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Executive functions Frontal lobe:
Major subdivisions of prefrontal cortex: lateral prefrontal cortex, ventromedial prefrontal cortex, and anterior cingulate
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Executive processes Frontal lobes:
About third of cerebral cortex in humans; frontal lobes are much larger in humans and are much larger than in other mammals, especially its anterior aspect Frontal lobes separated from parietal lobes by central sulcus and from temporal lobes by lateral sulcus
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Kolb & Whishaw frontal lobe figure
Lateral area 6 premotor cortex Medial area 6 supplementary motor cortex Area 8 frontal eye field
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Executive processes Dorsolateral prefrontal cortex (areas 9 and 46)
Medial frontal cortex (areas 25 and 32) Inferior (ventral) prefrontal cortex (areas 11, 12, 13, and 14); also called orbitofrontal cortex
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Executive processes Anatomical properties of PFC that are crucial for executive processing Perceptual, motor, cortical, and subcortical brain structures project to PFC This makes it possible to combine information from diverse sources, thereby enabling complex behavior PFC has multiple projections to multiple brain structures This permits it to modulate (exert top-down guidance) on other neural processes
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Executive processes Working memory and lateral prefrontal cortex
A previous lecture discuss working memory Baddeley’s model of working memory was presented
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Baddeley’s working memory model
Visuo-spatial sketchpad Phonological loop Central Executive
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Executive processes Content-based model of working memory
This is known as a content-based model of working memory because it assumes that different systems represent different content Phonological stores linguistic info and visusospatial represents visual and spatial information Model based on behavioral data reviewed in prior lecture
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Executive processes Process-based models of working memory
It has been hypothesized that different regions of the PFC are recruited to carry out different tasks In other words different processes are associated with different brain regions
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Executive processes Different types of working memory
In some tasks a participant is presented a stimulus, then is required to internally maintain a representation of that stimulus, until a probe is presented (maintenance condition) In an n-back task the participant is required to keep in mind a stream of stimuli and respond only if the stimulus was presented n-back Requires maintenance + manipulation
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Executive functions Illustration of n-back task
Note, the need to maintain information in memory and manipulate it
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Executive processes D’Esposito et al. (1998) Meta-analysis
D’Esposito and colleagues performed a meta-analysis to investigate content-based and process-based accounts of working memory meta-analysis – quantitative review of findings from several studies
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Executive processes D’Esposito et al. (1998)
One might expect based on the content account of working memory that the phonological loop might be left lateralized and the spatial sketchpad should be right lateralized as it is for perception To investigate this possibility D’Esposito categorized studies based on their content (spatial, nonspatial)
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Executive processes D’Esposito et al. (1998)
Results partially support the hypothesis that spatial tasks tend to be associated with activation of the right prefrontal cortex, whereas nonspatial tasks are associated with activation of the left prefrontal cortex, but not However, there was bilateral activation in many studies See top panel
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D’Esposito et al. (1998) Top panel shows active foci for spatial and nonspatial tasks (content) Bottom panel shows active foci for maintenance and maintenance + tasks
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Executive processes D’Esposito et al. (1998)
to investigate the process-based account of working memory, tasks were categorized as requiring maintenance or maintenance plus (maintenance + manipulation) Results showed that tasks requiring maintenance plus had more dorsolateral activation See bottom panel
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D’Esposito et al. (1998) Top panel shows active foci for spatial and nonspatial tasks (content) Bottom panel shows active foci for maintenance and maintenance + tasks
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Executive functions Stroop test
This test assesses the ability to maintain a goal and ignore/suppress habitual (prepotent) responses
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Executive functions Stroop effect demonstration
in this next slide I want you to name the ink colours of the words as rapidly as possible
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Stroop (slide 1) red yellow blue red green red yellow green
blue green red yellow red green red yellow
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Executive functions Stroop effect demonstration
in this next slide I want you to name the ink colours of the words as rapidly as possible
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Stroop (slide 2) red yellow blue red green red yellow green
blue green red yellow red green red yellow
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Executive functions Stroop effect demonstration
in this next slide I want you to name the ink colours of the colour patches as rapidly as possible
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Stroop (slide 3)
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Executive functions Stroop effect
The increase in time it takes to name of color when the word name does not match the color versus naming color patches is called the color-word interference effect (slide 1 time/slide 3 slide) Note: different versions of the Stroop assess interference in slightly different ways
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Executive functions Age Dot time (DT) M (SD) Color word time (CWT)
Interference CWT/DT Errors Color word 18-39 11.0 (2.5) 22.1 (7.2) 2.0 (0.6) 0.8 (1.0) 75-74 13.3 (3.6) 32.6 (9.6) 2.6 (0.9) 0.6 (1.2) Dot time = name dot color; color word time = name word color DT = Name color of dot; CWT = Name color of word designating color Troyer, Leach, Strauss (2006), 13, 20-35
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Executive functions Stroop effect
Age effects – age significantly increases the magnitude of the interference effect Gender differences are not always present in the interference score The higher the IQ score the lower the interference effect
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Executive functions Stroop effect
Standard interpretation of Stroop – participant must selectively attend to the name of the ink color and ignore the word name (of a color)
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Executive functions Wisconsin Card Sort
Used to assess for frontal lobe damage 4 stimulus cards are arranged in front of a participant; cards vary on 3 dimensions: shape, color, number Participants are given a deck of cards and must match each card with 1 of the stimulus cards, but are not told on which dimension they are matching Participants are told “right” or “wrong”
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Executive functions
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Executive functions Participants are given a deck of cards and must match each card to 1 of the 4 stimulus cards Participants guess at first, but since they are given feedback, they learn the correct attribute After sorting about 10 cards correctly, the examiner changes the attribute without warning Normal participants soon figure out correct attribute for sorting
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Executive functions Frontal lobe patients and normal participants do not differ in learning first critical trial, but they differ in the ability to switch attributes Normal participants switch after a few trials of negative feedback; frontal lobe patients are less able to switch
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Executive functions Executive attention is needed whenever multiple representations in working memory or multiple processes are competing for control of behavior and thought In Stroop task there is competition, but this sort of competition is a feature of a broad range of tasks E.g., it has been shown that naming the color of a picture of a banana is slowed when it is not yellow (e.g., red) in general when there is an incompatibility between an automatic response and a correct response you get Stroop-like effects
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Executive functions Stimulus response compatibility exists when the response required is compatible with the way people would naturally respond to that stimulus E.g., high pitch respond – “up”; low pitch respond – “down” E.g., stimulus presented on left or right side of display requires a response on same side as stimulus was presented
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Executive functions Stimulus response compatibility is strong and is observed even when position of object is irrelevant to response (Simon, 1990) E.g., suppose task is to make a right-handed response when a circle is presented and a left-handed response to a triangle; reaction time is faster when the circle or triangle is presented on the side of the response Interpretation – when there is an automatic connection between a stimulus and a response, little executive attention is required; when 2 sources of information are incompatible, attention must be paid in order to focus on the relevant information and inhibit/ignore the irrelevant information
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Executive functions Cohen and colleagues have developed a neural network model of Stroop task It proposes that in addition to initial visual perceptual processing of color (occipital lobes) and visual words (temporal lobes), two additional attentional processes are required
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Executive functions Stroop effect demonstration
in this next slide I want you to name the ink colours of the words as rapidly as possible
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Executive functions 1. attention controller
This process keeps track of the task goal – this is necessary because during incompatible trials in the color word condition, the name of the color font and the name of the word, which designates a color are two competing responses In the color-word condition it does this by activating more strongly the processes associated with the task goal (in this case the color of the word)
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Executive functions 2. conflict monitor
This process monitors the amount of conflict between potential responses. When there is more conflict as there would be in the word-colour condition, the conflict monitor increases executive attention
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Executive functions Stroop effect
Other points – neuroimaging and lesion studies are consistent with hypothesis that frontal lobes are associated with Stroop Data also suggest that performance is mediated by a more broadly based system
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Executive functions Neuroimaging evidence
Jonides and colleagues (2002) performed a meta-analysis of Stroop studies and related studies Results showed that the anterior cingulate and dorsolateral PFC were activated; this is consistent with theory because it is known from other research that the anterior cingulate is activated mediates conflict and the dorsolateral PFC is involved in executive attention
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Executive processes Executive processes & effects on longer-term memory Individuals with PFC damage may be impaired in their ability to organize temporally events in memory Milner (1995) performed a recency experiment in which participants were required to discriminate which of two events was presented more recently
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Executive processes Memory for temporal order Milner (1995)
Participants were presented pairs of stimuli (e.g., 2 pictures of objects) Every so often a probe card is presented with ? Task: to choose picture with more recently presented object
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Executive processes Milner (1995) Memory for temporal order
Experimental condition– both objects had been presented previously Control condition – one picture presented previously one picture new (Recognition test)
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Executive processes Memory for temporal order Milner (1995)
3 groups of participants were tested Controls Unilateral damage to dorsolateral PFC px Unilateral damage to temporal px Px underwent surgery for relief from focal epilepsy
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Milner recency discrimination experiment
Top panel shows stimuli used in exp’t Bottom panel shows that PFC participants were impaired relative to other 2 groups on recency discrimination test but not on recognition test
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Executive processes Memory for temporal order Milner (1995)
Other findings – this experiment was also performed with word stimuli Results showed that the effect was lateralized – Patients with LHD were more impaired on recency discrimination when words were used, whereas px with RHD were more impaired with pictures
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Executive functions Memory for temporal order
Milner et al. (1991) performed an experiment in which frontal and parietal patients were shown pairs of items Occasionally the pairs had a question mark between them Participants task was to decide which item had been presented more recently When only 1 item had been actually presented the test was an item recognition test When both items had been presented, the test requires order (and item) information
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Executive functions Memory for temporal order
Patients with frontal lobe damage were impaired in their memory for order but not item information Patients with parietal lobe damage were impaired in their memory for item but not order information
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Executive processes Memory for temporal order Self-ordered pointing
Keeping track of previous experienced events (memory for temporal order) has been examined using a self-ordered pointing task In this task participant is presented n cards with n objects depicted on it The same objects are present on each card but their order is scrambled from card to card
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Executive processes Memory for temporal order Self-ordered pointing
Task of participant is to point to the a different object on each card (one that hasn’t yet been pointed to) Results Frontal lobe patients made more errors than controls; discrepency between 2 groups increased with n, the number of objects and cards
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Executive processes Source memory
Source memory refers to when we learned a fact or the context in which a fact was learned –e.g., who told you fact or in what context you viewed a face Source memory appears to require frontal lobe function
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Executive processes Source memory Janowsky, Shimamura, & Squire (1999)
In this experiment participants were taught new facts (e.g., “The name of the dog on the cracker box is Bingo”)
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Executive processes Source memory Janowsky, Shimamura, & Squire (1999)
6-8 days later participants were asked to answer questions about these newly learned facts and other facts that might have been acquired outside of the experiment; if they recalled the fact they were asked questions about when they learned the fact (during the previous session or reading, school etc.)
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Executive processes Source memory Janowsky, Shimamura, & Squire (1999)
Results Controls and frontal lobe patients did not differ in recall of facts Frontal lobe participants were impaired in recall of source of facts
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Executive processes Source memory Glisky et al. (1995)
Investigated role of memory and executive function on item and source memory Neuropsychological tests assessed memory (temporal) and executive (frontal) function Performance on these two types of tests was weakly correlated
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Executive processes Source memory Glisky et al. (1995)
Study: participants heard sentences describing events (e.g., The boy went to the store to buy apples and oranges) Half were read aloud in man’s voice and half in a woman’s voice
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Executive processes Source memory Glisky et al. (1995) Memory test
Item memory: recognition test in which pairs of studied and unstudied sentences were presented together Source memory: participant heard sentence read aloud in a man or woman’s voice and decided which voice matched the studied sentence
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Executive processes Source memory Glisky et al. (1995) Results
Controls and frontal lobe patients did not differ in recall of facts Frontal lobe participants were impaired in recall of source of facts
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Glisky (1995) Source memory relatively more impaired than item memory by low frontal function Item memory relatively more impaired than source memory by low temporal function
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Executive processes Executive processes & effects on longer-term memory Individuals with PFC damage may be impaired in their ability to organize temporally events in memory Milner (1995) performed a recency experiment in which participants were required to discriminate which of two events was presented more recently
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Executive functions Sequencing
Many activities of everyday living require people to plan and then carry out a sequence of activities, which must satisfy certain requirements in order to achieve a goal Studies have shown that patients with PFC damage are impaired on certain sequencing activities, but not all sequencing activities
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Executive functions Sequencing – how is temporal order coded?
An important component of sequencing is coding temporal order of events Evidence suggests that there are several different ways in which temporal order might be coded and that the coding of temporal order often involves separate processes from the coding of item information
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Executive functions Sequencing – how is temporal order coded?
Sternberg, (1966, 1967) compared performance on an item recognition (was letter presented?) and an order task (what is the next letter?) Item recognition – memory set (BGRD); probe (g); response (“yes”) Order task -- memory set (BGRD); probe (g); response (“r”)
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Sternberg, (1966,1967)
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Executive functions Sequencing – how is temporal order coded?
Sternberg, (1966, 1967) note the large slope differences between the two tasks Results suggest that different processes are involved item and order tasks; one difficulty with experiment is that one task requires recognition while the other requires recall; however, many studies are consistent with the idea that item and order information are stored and processed differently
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Executive functions Sequencing – how is temporal order coded?
For example, there is evidence to suggest that memory for order information (information about associations) depends critically upon the hippocampus, but item information can be carried out independently of the hippocampus There is also evidence indicating that sequential information can be stored and processed in a variety of ways
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Executive functions Sequencing – how is temporal order coded?
There are many ways to code temporal order and there is evidence for each of these ways depending upon the task and other factors 1. Associative – e.g., X R C B L ; code as X precedes R; R precedes C; C precedes L etc. 2. order tags – e.g., X R C B L; code as X is first; R is second; C is third, etc. 3. familiarity – e.g., a form of representation in which strength or familiarity of item is continuously represented, so more recent items are stronger than earlier items, allowing participants to make a judgement on that basis (e.g., B is stronger then C)
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Executive functions Sequencing related items
In many everyday situations, sequences of actions are related to each other, and in many cases have been performed several times in the past E.g., eating out typically involves being greeted, taken to a table, ordering a meal, eating, getting a bill, paying, and leaving (Schank & Abelson, 1977) It is also possible to generate a novel script (e.g., opening a beauty salon)
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Executive functions Sequencing related items
Sirigu et al. (1985) performed an experiment in which patients with PFC damage, damage to the posterior cortex, and normal controls were tested on familiar and novel scripts Participants were asked to generate familiar (going to a restaurant) and novel (opening a beauty salon) actions and then they were asked to order the actions into correct sequences
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Executive functions Sequencing related items
No significance difference in the number or type of actions generated Individuals with PFC damage made more errors than other 2 groups when asked to order generated actions in correct sequence and the pattern was amplified with novel scripts Similar findings were obtained when the different groups were given cards with actions for scripts written on them
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Executive functions Brain imaging studies
Buckner & Wheeler (2001) reviewed findings from neuroimaging of remembering Several studies have shown consistently with a variety of stimuli (words, faces, pictures) that during memory recall the dorsolateral prefrontal cortex and the anterior frontal-polar cortex are activated
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Executive functions Brain imaging studies e.g., Buckner et al. (1995)
Participants studied a list of words (couple, string) At test, during imaging, participants were shown the beginning of the word and were intstructed to recall the word (e.g., cou---, stri---- ) Results on next slide illustrate the pattern of activation
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Buckner et al. (2001) Typical pattern of activation during recall of studied material – blue, dorsolateral (BA 44/6); green anterior frontal-polar (BA 10)
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Buckner et al. (2001) 2 brain regions appear to have distinct functional properties Selectivity of anterior suggests it may have a specific memory function; late onset consistent strategic retrieval/monitoring
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Executive functions Memory retrieval success
Evidence that MTL and PFC involved during memory retrieval, but are there additional regions? Habib and Lepage (1999) meta-analysis compared activation of old and new items Results showed that a network consisting of left parietal cortex, left anterior frontal lobe (near frontal-polar region) responded more to old items (i.e., retrieval success) This finding has been replicated in more recent studies with a variety of materials
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Buckner et al. (2001) Panel a. paradigm
Exp’t in which activation of old versus new items is compared
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Buckner et al. (2001) Panel b. fMRI imaging show differential activation of brain regions associated with retrieval of success; note left parietal cortex activation (green arrow)
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Buckner et al. (2001) Panel c. Event-related potential (ERP) shows a rapid positive waveform that develops over left parietal sites during remembered old items (red) as compared to new items (green)
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Executive functions Spontaneous confabulations
Individuals, who spontaneously confabulate have also been used to investigate role of the frontal lobes and executive function in memory Spontaneous confabulation – statements are actions that reflect unintentional but obvious distortions of memory “honest lying” Spontaeous confabulation is found sometimes but not always after px who survive aneurysms of the anterior communicating artery (ACoA)
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ACoA Ventral view of arteries in brain
Damage to ACoA may result in memory loss, personality change and amnesia Damage to ACoA often results in PFC damage
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Executive functions Spontaneous confabulations
Gilboa et al. (2006) investigated spontaneous confabulations Participants were ACoA individuals who confabulate, ACoA individuals who do not confabulate, and controls
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Executive functions Spontaneous confabulations
Temporal context judgment experiment In this experiment participants were required to make temporal context judgments about pictures of objects (Was this object presented earlier in this list; i.e., not a previous list?) Results Not all ACoA patients were impaired on the temporal context confusion experiment, but those with ventromedial prefrontal cortex damage were impaired
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Executive functions Spontaneous confabulations
Familiar narratives experiment (fairy tales and bible stories) In this experiment participants were asked to recall familiar narratives Results: spontaneous confabulators did not differ from ACoAs in terms of details recalled, but differed in terms of the number and type of errors (e.g., incorporation of details from other stories, idiosyncratic details)
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Semantic narrative performance
Top: mean details produced Bottom: proportion errors – T = total errors; D = distortions; E = external details (other stories); I = idiosyncratic errors
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Executive functions Spontaneous confabulations Further analyses
Only px with ventromedial prefrontal and orbitofrontal cortical damage were spontaneous confabulators
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Executive functions Spontaneous confabulations
Conclusion: spontaneous confabulation does not appear to be a result of confusing true memories in time (evidence: spontaneous confabulator errors such as semantic narrative external details) instead it appears to be a difficulty of strategic retrieval and difficulties of monitoring Strategic retrieval refers to a type of memory retrieval in which the target memory is not directly elicited by the retrieval cue (e.g., what did you do on your birthday?), but appears to require problem solving
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Executive functions Spontaneous confabulations
Evidence for strategic retrieval 1. Evidence from current study 2. Finding that spontaneous confabulation is observed even for remote memories acquired prior to brain damage (when encoding of memories was intact); suggests stored memory is relatively intact and problem is in retrieval
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Executive functions
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Executive functions Spontaneous confabulations Gilboa et al. proposed:
Strategic retrieval helps frame the memory problem and memory search Constrains memory search Once a memory is retrieved strategic retrieval monitors recovered memory for plausibility
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Executive functions (social, personality and emotional functions)
Overview Gage walked away from accident, could describe the accident the next day, and within a month was deemed able to resume work as a foreman It became clear that Gage was “no longer Gage” Prior to injury Gage was a sober, responsible, intelligent, home body, with no peculiar or bad habits; he was a responsible, valued employee After injury he was erratic, given to grossest profanity, impatient, unwilling to listen to advice, and unable to plan effectively Subsequent research has shown that damage to frontal lobes can lead to dramatic changes in personality while keeping perception, consciousness, and most cognitive functions intact
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