CORTEX FOR DECISION MAKING Sandrine Duverne Paris, Dec 02, 2019 - - PowerPoint PPT Presentation

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CORTEX FOR DECISION MAKING Sandrine Duverne Paris, Dec 02, 2019 - - PowerPoint PPT Presentation

Master of Integrative Biology Neuroscience Specialization UE 5NB04 INFORMATION PROCESSING IN THE PREFRONTAL CORTEX FOR DECISION MAKING Sandrine Duverne Paris, Dec 02, 2019 Decision making A SERIES OF INTERWEAVED CHOICES IN EVER-CHANGING


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Master of Integrative Biology Neuroscience Specialization UE 5NB04

INFORMATION PROCESSING IN THE PREFRONTAL CORTEX FOR DECISION MAKING

Sandrine Duverne

Paris, Dec 02, 2019

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Decision making

A SERIES OF INTERWEAVED CHOICES IN EVER-CHANGING ENVIRONMENT

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Perspective of cognitive sciences

Decision making refers to a set of cognitive processes that results in the selection of a belief or a course of action among several alternative possibilities.

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Characteristics of decision making

Decisions are :  either automatic/uncontrolled or voluntary/controlled.  either irrational/intuitive/emotional or rational/thoughtful/effortful  Based on both knowledge (subjective values, preferences, beliefs) and information in the immediate environment (context, instructions, rewards).  dynamic and temporally intermixed.

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How the brain makes decision

THE PREFRONTAL CORTEX ORCHESTRATES DECISION MAKING

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The prefrontal cortex

A very large part of the brain that plays a key role in high- level cognitive processes involved in goal-driven behaviors

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Cross-species evolution of the PFC

  • PFC grey matter in humans is up to 1.9-fold greater than in

macaques and 1.2-fold greater than in chimpanzees

  • Subcortical white matter underlying the PFC in humans is

2.4-fold greater than in macaques and 1.7-fold greater than in chimpanzees (Donahue et al., 2018)

Human cognitive uniqueness should therefore focus less on the frontal lobes in isolation and more on distributed neural networks.

(Barton & Venditti, 2013)

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Two approches of PFC functions in decision making

MODULAR APPROACH localization

Goal-directed choices can be subdivided into a set of discrete component processes with serial and localized neural implementation.

CIRCUIT-BASED APPROACH distribution

Goal-directed choices emerge from repeated computations that are distributed across many brain regions that perform similar computations.

Hunt & Hayden, Nature Rev Neurosci, 2017

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Tools and techniques

Temporal resolution EEG, MEG Spatial resolution fMRI, PET scans Connectivity modelling static functional connectivity dynamic functional connectivity

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The modular approach

  • f PFC functions

MODULES OF INFORMATION PROCESSING

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Localization of cognitive functions in the brain

An old hypothesis recycled to study the organization of cognitive functions in the brain.

Fodor, 1983 Gall & Surzheim , 1810

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Modularity of the mind

Modules are :  Domain specific: dedicated to a specific type of process or computation  Encapsulated: each module refer to

  • ther modules

 Associated with a specific neural structure  Their output converge and are centralized in a central system

The mind is composed of composed of neural structures (modules) with distinct evolutionarily-developed functions.

Fodor, The Modularity of Mind, 1983

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Decision making in the modular approach

A series of component processes :  evaluation of options  comparison of option values  selection of an appropriate action plan  monitoring of the outcome of the choice

Each component process is assumed to correspond to discrete neural computations implemented in distinct neural modules .

Hunt & Hayden, Nature Rev Neurosci, 2017

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Three main functional axes

Lateral PFC Control system Ventral PFC Valuation system Dorso-Medial PFC Motivational system

Each PFC axis

  • perates along

distinct dimensions of information processing

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COGNITIVE CONTROL IN LATERAL PFC

Hierarchical processing of information

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Cognitive Control

Attention

  • Selective

attention

  • Inhibitory

attention

  • Shifting

attention Memory

  • Working

memory

  • Episodic

memory

  • Prospective

memory Problem solving

  • Action selection
  • Conflict

resolution

  • Error detection
  • Planning

Set of processes that allow information processing and behavior to vary adaptively from moment to moment depending on current goals and environmental constrains

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Hierarchical organization of PFC functions

The frontal and posterior parts of the brain are two networks that are hierarchically organized:

  • Frontal system = motor memory
  • Dorsal system = perceptual memory

PFC functions are hierarchically organized in an anterior-posterior axis to process abstract to more concrete information.

Fuster, Trends NeuroSci, 1997

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The notion of task set

Task sets :  a configuration of cognitive processes that is actively maintained for subsequent task performance  Dorsolateral PFC functions are organized to selectively represent, update and implement a specific task  Competition between neural representations of task sets during task switching  Facilitate task performance by establishing context-dependent rules

Sakai, Ann Rev Neurosci, 2003 Collins & Frank, PLoS Comput Biol, 2016

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Cascade of cognitive control processing

Abstraction

subsets of actions are linked with contecxtual cues to form task sets

  • Sensory control : select actions based on stimulus-response

associations

  • Contextual control : select task set based on contextual cues

Temporal processing

ensemble of task sets are maintained and temporally extended

  • Episodic control : select ensembles of task set based on temporal

episode (frequencies of cues)

  • Branching control : monitor alternative task sets = not currently

performed but can be quickly reactivated

Koechlin & Summerfield, Trends Cog Sci, 2007

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Take home message

Cognitive control in lateral PFC

  • Hierarchical organization
  • Contextual control: abstraction of task

set based on contextual cues

  • Temporal control: maintenance of task

sets during episodes with repeated cues

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VALUATION SYSTEM IN VENTRAL PFC

Reward-based choices

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Valuation system

Reward

  • Part of the

dopaminergic system with direct projection from the ventral striatum Subjective values

  • Expected values

and reward- probabilities associated with stimuli

  • Common currency

across stimuli Social cognition

  • Emotion
  • Self-estim and self-

representation

  • Theory of mind

Set of processes that allow information processing and behavior to vary adaptively from moment to moment depending on available rewards in the environment.

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Ventromedial PFC & value-based choices

Daw et al., Nature 2006 4-arm bandit choice task

Markov Decision Process to maximize expected gain by choosing between competing choices associated with stockastic rewards

Reward-related activations

vmPFC activations linearly increase with rewads and choices

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Doya, HFSPJ 2007 Dopaminergic loops Reinforcement learning and dopaminergic loops

Dopaminergic system and reinforcement learning

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MODEL-FREE DECISION IN STRIATUM

  • Estimation of cache values with

temporal-difference learning

  • Actions are selected on the basis of

stored value representations

  • Long-term values are estimated directly

from experience MODEL-BASED DECISION IN PFC

  • Estimation of the state and reward

transitions

  • A knowledge of action-outcome

relationships is used to anticipate long- run consequences of candidate actions

  • Long-term reward probabilities are

estimated by iteratively searching through a decision tree

PFC vs Striatum

Daw et al, Nature Neurosci 2005

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Model-free and model-based choices

Glascher et al., Neuron 2010

Decision tree paradigm sequential two-choice Markov decision State and reward prediction error in striatum vs lateral PFC State prediction correct signal in striatum and vmPFC State-based > standard RL

Hampton et al., J Neurosci 2006

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Take home message

Valuation system in ventral PFC

  • Reward dependent
  • Value-based model of the environment
  • Signature of choice value
  • abstract model of task structure to guide

behavioral choices: combines model-free striatal signals and model-based lateral PFC signals

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MOTIVATIONAL CONTROL IN DORSO-MEDIAL PFC

A key interface between reward and control systems

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Cognitive control

  • Error monitoring
  • Conflict detection
  • Motivational

control

  • Task switching
  • Regulation of

cognitive control Reward-based decisions

  • Volatility of rewards
  • Unchosen task

values

  • Foraging
  • Choice difficulty
  • Preference signals

Effort

  • Sustained effort
  • Regulate alertness
  • Speed-accuracy

tradre-off

Dorso-medial PFC functions

Mediating role of the dorsal anterior cingulate cortex (dACC) as building the links between rewards and actions

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Dorsal ACC & Cognitive control

Error monitoring

van Schie et al, 2004

Conflict detection

Carter & van Veen, 2008

Hierarchical motivation

Kouneiher et al, 2009 Holroyd & Coles, 2002

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Dorsal ACC & value-based choices

Volatility of reward

ACC learns the value of information in an uncertain world

Behrens et al., Nature Neurosci 2007

Foraging

ACC encodes the average value of the foraging environment and cost of foraging

Kolling et al., Science, 2012

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Dorsal ACC as selector or detector of alternative choices

ACC activations scale with choice difficulty Shenhav et al., Nature Neurosci 2014 ACC activations signal the need to switch and depart from the default choice Donoso et al., Science 2014

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Take home message

Motivational control in Dorso- medial PFC

  • Interface between cognitive control and

valuation system

  • Signature of the need to increase cognitive

control

  • Signature of difficulty to depart from default

choice

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SUMMARY OF THE MODULAR APPROACH

Strenghts and limits

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  • Simplification of system complexity
  • Ecologically flexible: hierarchically inter-

connected components support the emergence of adaptive behavior and cognition

  • Ecologically robust: Modular systems can

evolve under changing or competitive selection criteria

The modular approach

STRENGHTS RESOLUTION LIMITS

  • Topological scales: each module contains sub-

modules, which contain sub-sub-modules (the homunculus fallacy of infinite regress)

  • Spatial scale: how to link small-scale and

large-scale networks

  • Temporal scale: how to account for evolution
  • f functions over time (or across distinct

pathologies)

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The circuit-based approach of PFC functions

BRAIN DYNAMICS FOR DECISION MAKING

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Distribution of cognitive functions in the brain

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Small to large-scale circuits

Highlights of large-scale brain circuits:  Functional connectivity: parallel and antagonistic pathways complicate circuit  neuronal dynamics shape the activity

  • f circuits

 neuromodulation reconfigures circuit properties  circuits interact to generate behavior

Neural circuits interconnect to

  • ne another to carry

information and form large scale brain networks.

Bargmann & Marder, Nature Methods, 2013

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Large-scale functionnal circuits

  • Distributed: Association cortex

comprises multiple distributed networks of distinct connectivity profiles that are embedded within largely parallel, interdigitated circuits.

  • Hierarchical: The hierarchy across

networks reveals how a distributed network might serve as a bridge between sensory and motor networks

Yeo et al., J Neurophysiol 2011

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Decision making in the circuit-based approach

Three overarching principles :  Decisions (and values) are formed in a distributed fashion across many brain regions.  The distributed networks are highly recurrent in nature.  The networks are organized into functional and temporal hierarchies.

Choices are formed in a distributed fashion across many brain regions that act in concert and perform similar computations.

Hunt & Hayden, Nature Rev Neurosci, 2017

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COGNITIVE CONTROL BEYOND LATERAL PFC

Sensorimotor transformations in the fronto- parietal network

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Two frontoparietal circuits in attention

Dual attention system

  • Dorsal fronto-parietal system

Goal-directed (top-down) attention for selection of stimuli and responses

  • Ventral fronto-parietal system

Stimulus-driven (bottom-up) attention for detection of relevant stimuli 'circuit breaker' of the dorsal system

Corbetta & Schulman, Nature Reviews Neurosci 2002

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Distinct posterior parietal cortex circuits in attention and memory

  • In perceptual attention: dorsal regions (pIPS and

SPL) are related to visuospatial attention and eye movements

  • In memory retrieval: the latIPS tracks the sense of

familiarity or perceived memory strength

Dynamic competition between dorsal and ventral parietal regions contributes to reorienting retrieval in episodic memory.

Sestieri et al., Nature Rev Neurosci 2017

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The Intra-parietal sulcus as a ‘mnemonic accumulator’

  • drift-diffusion model in perceptual decision : decisions result

from the continuous accumulation of noisy information until one

  • f two response thresholds is reached

 Evidence accumulation and motor intention are coupled: network-level interactions organize environmental sampling into rhythmic cycles (Fiebelkorn et al., Neuron 2018)

  • Accumulating memory-decision evidence: episodic retrieval is

the process of accumulating evidence about the relatedness between probes and items in the memory set  Evidence accumulation and motor intention are NOT coupled: IPS tracks perceived oldness, regardless of whether the response is correct or incorrect

Sestieri et al., Nature Rev Neurosci 2017

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Take home message

Cognitive control beyond lateral PFC

Two frontoparietal circuits of cognitive control:

  • Perceptual decisions based on sensory-

evidence accumulation

  • Episodic retrieval based on memory-

evidence accumulation

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VALUATION SYSTEM BEYOND VENTRAL PFC

Linking past and future memories in the default mode circuit

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The default mode network

Large-scale network with a set of hubs densively connected with surounding regions Task-negative functional activations

  • Default mode network in passive tasks: resting state, stimulus

independent thoughts, spontaneous activity dynamic, monitoring external environment broadly for unexpected events

  • Control network in active goal-directed tasks negatively regulates

the DMN

Raichle et al., PNAS 2001 ; Grecius e al., PNAS 2003

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DM network in goal-directed task

Interacting subsystems

  • Medial temporal lobe provides information from

prior experiences

  • Medial prefrontal subsystem facilitates the flexible

use of this information for self-relevant mental simulations.

  • DM network competes with executive network to

facilitate the retrieval and integration of relevant information, stored in their modality-specific cortical areas. Buckner, Ann N Y Acad Sci 2008

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Flexibility of functional connectivity according to task demand

Rapid adjustments in control may occur via flexible task-dependent shifts in frontoparietal coupling patterns.

  • External attention circuit: negative coupling with DN nodes and

positive coupling change with task demand  Functional connectivity varies with top-down signals of task-relevant

  • bjects and locations

 Sensorimotor processing facilitate moment-to-moment interactions with the environment

  • Internal attention circuit: shift away from negative coupling in DN

nodes when attention is directed toward one’s own thoughts and away from perceptual inputs  abstract “top-level management” of thought exerts a general constraint according to task-relevant material

Dixon et al., PNAS 2018

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Take home message

Valuation system beyond ventral PFC DM network as an internal attention system:

  • facilitate the retrieval and integration of

relevant information

  • flexible task-dependent shifts with

frontoparietal coupling

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MOTIVATIONAL CONTROL BEYOND DORSO-MEDIAL PFC

A key interface between the default mode and control systems

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The salience network

Pyramidal cells van Economo cells

Eisenberger, N., 2012

The salience network is involved in identifying biologically and cognitively relevant events to guide flexible behavior.

  • Anterior insula: typically associated with social and

affective tasks involving pain, empathy, disgust, and introspective processes, external and internal signals

  • dACC : typically associated with cognitive control and

valuation systems

  • Van Economo cells: wider axons that facilitate rapid relay of

signals from the AI and dACC to the motor system

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Dynamic interaction with other brain networks

Switching mechanisms in the ‘spotlight of attention’

  • SN recruits the central executive and task

control regions to maintain cognitive set and manipulate information in working memory

  • SN helps suppressing the default-mode

network to keep attention focused on task- relevant goals Menon, Brain Mapping: An Encyclopedic Reference 2015

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Brain circuits in creativity

Beaty et al., PNAS 2018

The creative connectome

High-creative network: set of frontal and parietal regions typically associated with default, salience, and executive brain systems. The creative brain is marked by a tendency to selectively and simultaneously engage these large-scale circuits to a greater degree than the less creative brain. Default and executive network coupling is mediated by salience network and supports creative idea production.

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Take home message

Motivational control beyond Dorso- medial PFC

Detection of salient stimuli to guide flexible behaviors throught two properties:

  • Switching mechanisms between the FPC

and DMN

  • Gate between the FPC and DMN for

creative idea generation

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SUMMARY OF THE CIRCUIT- BASED APPROACH

Stenghts and limits

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  • Signal circularity : how to dissociate

qualitatively distinct signals exchanged within a functional circuit

  • Signal directionality : how to dissociate the

sequences of signals embedded in a single functional circuit

The circuit-based approach

  • Ecologically robust and flexible: Computation

are distributed across several regions that support adaptive behavior

  • Recurrent signal: temporally extended

algorithms, in which noisy estimates of value are integrated sequentially over time

STRENGHTS LIMITS

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Conclusions

HOW TO LINK THE MODULAR AND CIRCUIT- BASED APPROACHES

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Methological and analytical strenghts

MODULAR APPROACH

  • Experimental toolbox: Behavioral and

functional data are sampled in a controlled environment

  • Theory-based analyses: Analyses of

behavioral and functional data aim at testing hypotheses

CIRCUIT-BASED APPROACH

  • Functionnal connectivity toolbox:

constrained by assumptions on anatomical connectivity

  • Data-driven (and theory-based)

analyses: Analyses of behavioral and functional data are either free of or constrained by hypothesis testing

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Challenges

MODULAR APPROACH

  • Need to account for variability in

behavioral responses: no one-to-one mapping of input-output in functional activations and behaviors

  • Need to account for changes in the

modules of a system: how to account for different developmental trajectories in life and for commonalities and differences across pathologies

CIRCUIT-BASED APPROACH

  • Need to understand the circuit

dynamics: how a system’s output results from the interactions of its components (causality)

  • Need for parsimony: enormous numbers
  • f assumptions about circuit parameters

that are likely to change in different modulatory states.

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Thanks for your attention!

SDUVERNE@CREACOG.COM