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The human reward system and its associated brain regions

As for satisfying our needs - like food, sleep, sex, love and approval (to name but a few) - we need no extra motivation. Which doesn't mean that we may follow our pleasures in any form or by any means, for if we did so we might have to face serious consequences. In order to be able to find out what is appropriate in any given context, and what is not, we need a complex and reliable neural network with stable conductions of stimuli from the midbrain all the way up to the cortex.

Our social "warning system" has to shoulder a lot of tasks: the anticipation of consequences, the recognition, processing and evaluation of outside information (like facial expressions or context), as well as the detection of internal and external conflicts and errors... And even if this warning system failed us for once and we had to face the unpleasant outcomes we could still learn by the consequences of our decisions (like punishment or ostracism) and thereby avoid to make the same mistakes over and over again. An efficient reward system with a reliable cortical feedback enables us to adapt to our social environment and prevents us from running into trouble. Thus by way of stimulus-response learning we automatically form durable patterns of stimulus and response that (should) make our lives a lot easier. This learning process is called operant conditioning.


The intertwining of the following tasks makes for an efficient reward system:

1. The perception/ recognition of environmental information (like emotions etc. - for instance by way of empathy)

2. The processing and evaluation of environmental information and our own responses to that information

3. The error and conflict detection (internal and external discrepancies)

4. The anticipation of consequences

5. Decision making in coordination with current information, previous knowledge (memory), own values, subjective valence of stimuli, motivation and many more

6. The (situational) adaptation/ adjustment of behavior (response modulation, inhibition, amplification and so on...)

7. Operant conditioning (the long-term adaptation) and "reversal learning"


The following dysfunctions and disorders are characterised by disturbances of this information flow for social "fine tuning":

* Tics, obsessions and compulsions (as in Obsessive-Compulsive Disorder, Tourette Syndrome, Addictions, Pedophilia...)

* Misinterpretations or distortions of environmental information (as in Depression, Anxiety Disorders, PTSD, Dissociation, Schizophrenia...)

* Impaired inhibition/ affect regulation (as in Borderline PD, Pedophilia...)

* Social disorders


The following brain regions chiefly contribute to the human reward system:

OFC, MPFC, VMPFC, pACC, LPFC, Insular lobe, Amygdala, Hippocampus, ACC, Basal ganglia (Striatum, Caudate nucleus, Globus Pallidus, Nucleus Accumbens), Thalamus, Fornix and Substantia Nigra.

Brain Region Functions within the reward system
OFC

1.) Posterior: emotion recognition and empathy S; sympathy/ preference (unknown faces) S; strong connections to the amygdala and anterior parts of the temporal lobes as well as to the sensory association cortex S

2.) Interpretation S, processing of environmental emotional and motivational context information S, e.g. facial expressions S;

Sensory integration; selection, comparison, and evaluation of stimuli,

Appraisal of the emotional and motivational values of environmental information (while integrating the subject's prior experience) S; access to emotional memory S; episodic memory S

3.) Prediction (mOFC, S) and detection of errors

5.) Important for decision making S

6.) Adjustment: reaction to punishers - like ostracism - and reinforcers - like food and praise - (evaluation S, monitoring, learning, storage), and the resulting behavior; context-dependent regulation of amygdala-dependent autonomic and behavioral arousal governed by external cues S; fear extinction (mOFC) S

7.) Operant Conditioning:
Behavior modification (emotion control, impulse control, social adjustment) especially in view of the consequences like reward and punishment S, change of behavior ("reversal learning" S), learning by mistakes and unexpected outcomes S

Disorders: General impairments involving stimulus-reward reversal learning, response inhibition, and ability to judge the appropriateness of one's own behavior in the social context S; Anterolateral: negative affect, obsessions S; Borderline PD S, Posttraumatic SD, Addiction S, S, Depression S, Obsessive-compulsive disorder S, (right) S, S (dopaminergic hyperfunction), Anxiety disorders S, Chron. Schizophrenia S, S, (Heterosexual) Pedophilia (hypoactivity) S

Medial PFC

2.) Cognitive appraisal of affective stimuli even if they are complex and ambiguous) S relating to people (instead of objects); strong connections to the amygdala;

Social cognition and reward, social valence S and classification S; Processing of positive (not negative) visual stimuli S

"Contextualization" of stimuli; dysregulation of contextualization processes might play a key role in the generation of PTSD symptoms S

5.) Moral judgments (right/ wrong) S, Theory of Mind ToM S, S

Disorders: Reality distortion, Schizophrenia, Schizoaffective disorders S, Dissociation (left hemisphere) S, Anxiety Disorders S, PTSD S, Cocaine Addiction S, Anorexia Nervosa S, Autism S

Ventromedial PFC

1.) Social cognition S (Reasoning about the minds and motivation of other people, compassion to the suffering opponent S, empathy/ friendship S, self-referential thinking while mentalizing about similar others S, S, self-relatedness S

2.) Affective evaluation (complex display of multiple human faces, (ambiguous) reward related information) S, S, Appraisal of the emotional and motivational values of environmental information S; * Anterior: visualization of emotional events in the far future (q.v.: caudate nucleus), and assignment of emotional values to mental representations of future events that pertain to long-term goals S

3.) Error detection: activation during mood-incongruent recall of autobiographical memories S

5.) Decision making under risk S

6.) Adjustment: Fear; fear extinction S, motor inhibition S

Disorders: Borderline PD S, Depression S, S, S, PTSD S; Psychopathology S, Anxiety Disorders S

pACC

2.) Evaluation and/ or representation of reward value S, Valence of all stimuli, whether persons or objects S

Disorders: Bipolar Disorder S

Lateral PFC

2.) Context-dependent coding of stimuli, representation of contextual information (contexts: task, motivation, reward); integration of the cognitive and the motivational context for adaptive goal-directed behavior S

3.) Conflict detection: Anticipatory preparation and online adjustment in response to conflicts S

6.) Adjustment: Action monitoring S, Response inhibition S

Disorder: Pedophilia (hypoactivity) S

Insular lobe

1.) Body perception, subjective emotional experiences, conscious emotions, observation of noxious or disgusting stimuli S, empathy S

2.) Inequity coding S

5.) Decision making under risk S

6.) Motor adjustment following reward or punishment S;

'Pessimistic' bias: internal adaptation and preparation processes in order to act according to potential or certain unpleasant events S;

Response inhibition S; pain (anterior); suppression of pain (posterior) S

Disorders: Addiction, PTSD S, S (right, anterior), Depression S, S; Huntington's Disease S, Obsessive-Compulsive Disorder S, S among others

Amygdala

1.) Perception: Emotion, danger, disgust S, empathy S

2.) Processing of emotional and social information S, emotion recognition in others, emotional assessment of (amongst others negative S) facial expressions (are they threatening or not?) S

5.) Motivation S

6.) Adjustment: Modulation of memory-related processes (both enhancement and suppression of impulses) in other brain regions, such as the hippocampus S, S; recognition of negative items S

7.) Operant conditioning S (closely connected to the pOFC S); Fear S (conditioning) S

Disorders: Depression S, S, Borderline PD S (hyperactivity), PTSD S, Epilepsy S, Schizophrenia S, S, Autism and Paranoid Schizophrenia (right hemisphere) S, Autism (left hemisphere) S, Dissociative disorders S, Tourette Syndrome S, Obsessive-Compulsive Disorder S, Bipolar Disorder S, Cocaine Addiction S, Alcohol Addiction S, S (serotonergic system), Anxiety Disorders S

Hippocampus

4.) Anticipation: codes the degree of uncertainty of potential reward situations or rather the the strength of the association between the cue and the associated outcome; prerequisite for appropriate behavior S

5.) Previous knowledge: Encoding, retention and retrieval of spatial and episodic experiences S; memory S, above all episodic and autobiographic S, but not procedural, and only in part semantic (controversial, S, S, S); learning, recognition S

6.) Adjustment: Behavioral regulation and inhibition (e.g. of food intake for body weight regulation) S

7.) Conditioning: Spatial (CA3 subregion, S) and contextual learning and memory S, continuous encoding of ongoing experience, categorization S, pattern separation and pattern completion S

Disorders: Amnesia, PTSD S and other stress-related disorders S; Alzheimer's Disease S, Depression S, S, Anxiety Disorders S, Bipolar Disorder S, S, Dissociative disorders S, Obesity S, Tourette Syndrome S, Cushing's syndrome S, Multiple Sclerosis S; Huntington's Disease (posterior) S

Anterior Cingulate Cortex

1.) Perception: Ventral=affective S, attention on the internal emotional and motivational status S, empathy (with pain) S

2.) Processing: Information transfer: to the auditory association cortices, brain stem vocalization, and autonomic structures, in pathways that may mediate emotional communication and autonomic activation in emotional arousal S

3.) 'Error detection': Likelihood and magnitude of the consequences S, S, S; the conflict signal detected by the ACC is transmitted to other brain regions, such as the DLPFC, to increase the level of cognitive control S; Risk prediction, conflict monitoring and cognitive control S above all conflict detection between simultaneously active, competing representations S; conflict -> acts as teaching signal for avoidance learning S; dorsal: attention, working memory, error detection, conflict monitoring S

4.) Anticipation of rewards S

5.) (Reinforcement-related) decision making S, e.g. regarding how much effort to invest for rewards S, action evaluation, decision making regarding reward and punishment S, Posterior=evaluative

Problem solving, concentration upon a task, minimization of distractions S; dorsal: attention, working memory S, dorsal: response selection, anticipation of incoming information S, will bias behavioral decision making toward cognitively efficient tasks and strategies S

* Rostral: projection of positive future events, optimism S, detecting response conflict caused by irrelevant stimuli S

Attention on the internal emotional and motivational status, Motivation, Theory of Mind ToM (bilateral) S; Anterior=executive

6.) Adjustment: Response inhibition S, self-regulation, emotion regulation S

7.) Conditioning: Fear (rACC, gating the efficiency of amygdala-dependent auditory fear conditioning learning) S, S; pain S, S; learning through consequences, avoidance learning S, behavioral adaptation after delivered punishments S

Disorders: Borderline PD S, S, PTSD S, S (rostral), Schizophrenia S, S, S, ADHD, Obsessive-Compulsive Disorder S, S, S, S, Autism S, Depression (BA 25, S, S, S, S), Anorexia Nervosa S, Bipolar Disorder S, S, Addiction (Cocaine, rostral) S, Addiction (Nicotine, Dopamine) S

Basal ganglia

1.) Perception: Attention (Dopamine) S

2.) Processing of different emotional and cognitive stimuli (reward system, reinforcement, addiction, habits) S (see below)

6.) Adjustment: Motion selection, initiation and control: vabove all inhibition of unwanted/ inappropriate movements but also coordination of volitional movements, postures S, modulation of pain perception S

Disorders: Obsessive-Compulsive Disorder S; ADHD S, S, Tourette Syndrome S, Schizophrenia S; Depression S; Sexual disorders S among others

Striatum

1.) Perception: Activation by rewards S (dopamine, ventral) S and by aversive, novel, unexpected or very intense stimuli

6.) Adjustment: Sexual arousal and behavior S

* Ventral: Preparation, initiation and execution of reward-related behavior as a result of successful integration of (relevant) emotional and cognitive information (strong connections to the OFC and the ACC) S even when rewards were cued but omitted S

* Dorsal: adjustments after delivered rewards S, categorization S

Disorders: Striatofrontal dysfunction: Depression S, S, Chronic Schizophrenia S, Obsessive-Compulsive Disorder S (serotonergic hypofunction), Borderline PD S, Pedophilia S, ADHD S, Tremor S, Parkinson's Disease (Dopamine), Huntington's Disease (GABA) S, S; Athetosis, Dyskinesia; Addiction (Dopamine) S, S; Anorexia Nervosa S

Caudate nucleus

2.) Love S, motherly love S, pain S

4.) Visualization of emotional events in the near future; concrete simulations of action plans to achieve rewarding situations in the near future S

5.) Memory S, working memory S

6.) Left: Detection of a change in the 'context' and the subsequent formulation of a new 'rule' S; the linking of action to outcome S

7.) Conditioning: Perceptual skill learning S; Reaction to positive and negative feedback during learning S

Disorders: Obsessive-Compulsive Disorder S, S; Major Depression (left hemisphere) S, Parkinson's Disease (Serotonin) S, Huntington's Disease S, S; Schizophrenia S; Autism S; Addiction (Cocaine) S

Globus pallidus

Part of the reward system S

6.) Adjustment: Motion facilitation and inhibition; sexual arousal and behavior S

Disorders: Tourette Syndrome (GABA) S, Parkinson's Disease (GPI) S, Huntington's Disease S, Anhedonia/ Depression S; Addiction (Cocaine; ventral) S, loss of craving S; Pedophilia S among others

Nucleus accumbens 1.) Perception: Sympathy/ preference (unknown faces) S

2.) Processing: Empathy/ friendship S

5.) Evaluation/ decision making: self-reference S

7.) Conditioning: emotional learning, operant conditioning, motivation S

Disorders: Addiction S, Depression S, S; Bipolar Disorder S

Thalamus

1.) Facial emotion identification S; attention (dopamine) S

6.) Adjustment: Speech processing (e.g. ambivalence) S, working memory S; motor adjustment following delivered reward/ punishment S; modulation of pain perception S; 'Pessimistic' bias: internal adaptation and preparation processes in order to act according to potential or certain unpleasant events (medial) S; sexual arousal and behavior S

Disorders: Addiction (Nicotine) S, Obsessive-Compulsive Disorder S, S, S, S; Parkinson's Disease S, Tourette Syndrome S, Huntington's Disease S; Schizophrenia S, S, S; Insomnia S; Multiple Sclerosis S; Autism S; Pedophilia S, Wernicke-Korsakoff syndrome (dorsomedial nucleus) S

Fornix

* Posterior fibers:
-> Septal nuclei (reward, reinforcement)
-> Nucleus accumbens (reward, conditioning, motivation)

Disorders: Amnesia, Wernicke-Korsakoff Syndrome S; Schizophrenia S, S, S (controversial S), Alzheimer's Disease S, Parkinson's Disease S

Substantia nigra

Dopamine: Movement planning and initiation ('starter function'), motivation, drive etc. S, S

Disorders: Parkinson's Disease, Huntington's Disease S, Stuttering S, Cocaine abuse (in the uterus) S, Alcoholism (MCP-1) S, Alzheimer's Disease (iron) S, Tourette Syndrome S among others

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