Contributions of the striatum to learning, motivation, and performance: an associative account
Open accessAbstract
It has long been recognized that the striatum is composed of distinct functional sub-units that are part of multiple cortico-striatal-thalamic circuits. Contemporary research has focused on the contribution of striatal sub-regions to three main phenomena: learning of associations between stimuli, actions and rewards; selection between competing response alternatives; and motivational modulation of motor behavior. Recent proposals have argued for a functional division of the striatum along these lines, attributing, for example, learning to one region and performance to another. Here, we consider empirical data from human and animal studies, as well as theoretical notions from both the psychological and computational literatures, and conclude that striatal sub-regions instead differ most clearly in terms of the associations being encoded in each region.
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Written from the paper’s full text.
TL;DR
This review argues that striatal subregions are best distinguished by the type of associative content they encode, not by a simple learning-versus-performance or motivation-versus-performance division. It proposes that ventral striatum supports Pavlovian associations, dorsomedial striatum supports goal-directed instrumental associations, and dorsolateral striatum supports habitual stimulus–response associations.
Key findings
- The paper concludes that ventral and dorsal striatum are both implicated in learning and performance, so functional divisions along learning vs. performance lines are unlikely; instead, striatal regions support dissociable associative learning strategies.
- Ventral striatum is linked to Pavlovian associations, including conditioned skeletomotor responses and modulatory influences on instrumental performance such as general and specific Pavlovian-instrumental transfer.
- Within dorsal striatum, dorsomedial striatum is associated with goal-directed instrumental actions, whereas dorsolateral striatum is associated with habitual stimulus–response associations.
- The review notes unresolved issues, including whether Pavlovian signals in ventral striatum are model-free, model-based, or both, and inconsistent findings on ventral striatal responses in aversive contexts.
Methods
- Narrative review synthesizing empirical data from human and animal studies plus theoretical notions from psychological and computational literatures.
- Main analysis contrasts accounts that delineate striatal subregions by learning, performance, or motivation with an associative account emphasizing the content and nature of associative encoding.
- No original quantitative results or new empirical data are reported.
Limitations
- The paper states that very few human studies have empirically assessed the mapping of model-based and model-free reinforcement-learning signals to dorsomedial and dorsolateral striatum.
- The paper states that the neuroimaging literature is profoundly inconsistent regarding ventral striatal activity in aversive contexts, with some studies reporting increased and others decreased activity.
- The paper states that much less is understood about how dissociable striatal modules interact with each other and with cortex; it also notes its own discussion is geared toward appetitive learning, partly reflecting a general literature emphasis on reward processing in the striatum.
- Evident from the text: this is a narrative review without systematic search or meta-analysis, so its conclusions depend on the selected cited literature.
Model-written summary. Check it against the paper before relying on it.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
0 of 135 cited sources checked · 0 supported · 0 flagged
- Premisecitation
The ventral striatum is connected with limbic structures, including the amygdala, hippocampus, and medial orbitofrontal and anterior cingulate cortices.
“whereas the ventral striatum is connected with limbic structures, including the amygdala, hippocampus, and medial orbitofrontal and anterior cingulate cortices [1,2].”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [1] Lynd-Balta E Haber SN The organization of midbrain projections to the striatum in the primate: sensorimotor-related striatum versus ventral striatum. Neuroscience 1994 59 625 640 7516506 source
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- cites [2] Zahm DS Ventral striatopallidothalamic projection: IV. Relative involvements of neurochemically distinct subterritories in the ventral pallidum and adjacent parts of the rostroventral forebrain. J. Comp. Neurol 1996 364 340 362 8788254 source
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- Premisecitation
The ventral striatum is implicated in Pavlovian learning.
“Considerable evidence has amassed to implicate the ventral striatum (VS) in Pavlovian learning”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [16] Day JJ Associative learning mediates dynamic shifts in dopamine signaling in the nucleus accumbens. Nat. Neurosci 2007 10 1020 1028 17603481 source
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- cites [17] Day JJ Nucleus accumbens neurons encode Pavlovian approach behaviors: evidence from an autoshaping paradigm. Eur. J. Neurosci 2006 23 1341 1351 16553795 source
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- Premisecitation
In humans, activity in the DMS is correlated with computations of action-outcome contingency, a hallmark of goal-directed learning.
“in humans, activity in the DMS has been found to be correlated with computations of action-outcome contingency, a hallmark of goal-directed learning”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [21] Liljeholm M Neural correlates of instrumental contingency learning: differential effects of action-reward conjunction and disjunction. J. Neurosci 2011 31 2474 2480 21325514 PMC3269757 source
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- cites [22] Tanaka SC Calculating consequences: brain systems that encode the causal effects of actions. J. Neurosci 2008 28 6750 6755 18579749 PMC3071565 source
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- cites [23] Tricomi E A specific role for posterior dorsolateral striatum in human habit learning. Eur. J. Neurosci 2009 29 2225 2232 19490086 PMC2758609 source
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- Premisecitation
In humans, activity in a region of right posterior DLS tracks the behavioral development of habits.
“whereas activity in a region of right posterior DLS was found to track the behavioral development of habits”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [21] Liljeholm M Neural correlates of instrumental contingency learning: differential effects of action-reward conjunction and disjunction. J. Neurosci 2011 31 2474 2480 21325514 PMC3269757 source
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- cites [22] Tanaka SC Calculating consequences: brain systems that encode the causal effects of actions. J. Neurosci 2008 28 6750 6755 18579749 PMC3071565 source
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- cites [23] Tricomi E A specific role for posterior dorsolateral striatum in human habit learning. Eur. J. Neurosci 2009 29 2225 2232 19490086 PMC2758609 source
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- Premisecitation
The dorsomedial region of the striatum (caudate) is connected with frontal and parietal association cortices.
“In contrast, a dorsomedial region (i.e., caudate) is connected with frontal and parietal association cortices”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [1] Lynd-Balta E Haber SN The organization of midbrain projections to the striatum in the primate: sensorimotor-related striatum versus ventral striatum. Neuroscience 1994 59 625 640 7516506 source
Not checked in the free run.
- cites [2] Zahm DS Ventral striatopallidothalamic projection: IV. Relative involvements of neurochemically distinct subterritories in the ventral pallidum and adjacent parts of the rostroventral forebrain. J. Comp. Neurol 1996 364 340 362 8788254 source
Not checked in the free run.
- Premisecitation
The dorsolateral region of the striatum (putamen) is primarily connected to sensory and motor cortices.
“demonstrating that the dorsolateral region of the striatum (i.e., putamen) is primarily connected to sensory and motor cortices.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [1] Lynd-Balta E Haber SN The organization of midbrain projections to the striatum in the primate: sensorimotor-related striatum versus ventral striatum. Neuroscience 1994 59 625 640 7516506 source
Not checked in the free run.
- cites [2] Zahm DS Ventral striatopallidothalamic projection: IV. Relative involvements of neurochemically distinct subterritories in the ventral pallidum and adjacent parts of the rostroventral forebrain. J. Comp. Neurol 1996 364 340 362 8788254 source
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- Premisecitation
Tasks involving instrumental actions recruit both ventral and dorsal striatum.
“whereas tasks involving instrumental actions have been shown to recruit both ventral and dorsal striatum [27,29,30].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [27] O'Doherty J Dissociable roles of ventral and dorsal striatum in instrumental conditioning. Science 2004 304 452 454 15087550 source
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- cites [29] Cooper JC Human dorsal striatum encodes prediction errors during observational learning of instrumental actions. J. Cogn. Neurosci 2012 24 106 118 21812568 PMC3576883 source
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- cites [30] Glascher J Determining a role for ventromedial prefrontal cortex in encoding action-based value signals during reward-related decision making. Cereb. Cortex 2009 19 483 495 18550593 PMC2626172 source
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- Premisecitation
Different sub-regions of the dorsal striatum are involved in habitual and goal-directed instrumental conditioning, respectively.
“In contrast, different sub-regions of the dorsal striatum appear to be involved in habitual and goal-directed instrumental conditioning, respectively.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [18] Yin HH Lesions of dorsolateral striatum preserve outcome expectancy but disrupt habit formation in instrumental learning. Eur. J. Neurosci 2004 19 181 189 14750976 source
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- cites [19] Yin HH Blockade of NMDA receptors in the dorsomedial striatum prevents action-outcome learning in instrumental conditioning. Eur. J. Neurosci 2005 22 505 512 16045503 source
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- cites [20] Yin HH The role of the dorsomedial striatum in instrumental conditioning. Eur. J. Neurosci 2005 22 513 523 16045504 source
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- Premisecitation
In rodents, lesions to the medial part of the dorsal striatum (DMS) impair goal-directed learning.
“whereas lesions to the medial part of the dorsal striatum (DMS) impair goal-directed learning [18–20].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [18] Yin HH Lesions of dorsolateral striatum preserve outcome expectancy but disrupt habit formation in instrumental learning. Eur. J. Neurosci 2004 19 181 189 14750976 source
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- cites [19] Yin HH Blockade of NMDA receptors in the dorsomedial striatum prevents action-outcome learning in instrumental conditioning. Eur. J. Neurosci 2005 22 505 512 16045503 source
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- cites [20] Yin HH The role of the dorsomedial striatum in instrumental conditioning. Eur. J. Neurosci 2005 22 513 523 16045504 source
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- Premisecitation
Human fMRI studies have found that VS activity correlates with reward prediction errors during tasks that feature exclusively Pavlovian reward associations.
“human fMRI studies have found that VS activity correlates with RPEs during tasks that feature exclusively Pavlovian reward associations [27,28]”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [27] O'Doherty J Dissociable roles of ventral and dorsal striatum in instrumental conditioning. Science 2004 304 452 454 15087550 source
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- cites [28] O'Doherty JP Temporal difference models and reward-related learning in the human brain. Neuron 2003 38 329 337 12718865 source
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- Premisecitation
In rodents, lesions of the lateral dorsal striatum (DLS) disrupt acquisition of habits.
“In rodents, lesions of the lateral dorsal striatum (DLS) disrupt acquisition of habits”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [18] Yin HH Lesions of dorsolateral striatum preserve outcome expectancy but disrupt habit formation in instrumental learning. Eur. J. Neurosci 2004 19 181 189 14750976 source
Not checked in the free run.
- cites [19] Yin HH Blockade of NMDA receptors in the dorsomedial striatum prevents action-outcome learning in instrumental conditioning. Eur. J. Neurosci 2005 22 505 512 16045503 source
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- cites [20] Yin HH The role of the dorsomedial striatum in instrumental conditioning. Eur. J. Neurosci 2005 22 513 523 16045504 source
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- Premisecitation
The medial part of the nucleus accumbens called the core mediates the retrieval and expression of CS-US associations.
“a medial part of the nucleus accumbens (Nacc) called the core, distinct from a more lateral part called the shell (Figure 1), has been shown to mediate the retrieval and expression of CS-US associations [33,34].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [33] Blaiss CA Janak PH The nucleus accumbens core and shell are critical for the expression, but not the consolidation, of Pavlovian conditioned approach. Behav. Brain Res 2009 200 22 32 19159648 PMC4667776 source
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- cites [34] Parkinson JA Dissociation in effects of lesions of the nucleus accumbens core and shell on appetitive pavlovian approach behavior and the potentiation of conditioned reinforcement and locomotor activity by D-amphetamine. J. Neurosci 1999 19 2401 2411 10066290 PMC6782569 source
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- Premisecitation
The opportunity to earn very large rewards can have a deleterious influence on performance, a phenomenon known as choking.
“the opportunity to earn very large rewards has also been shown to have a deleterious influence, a phenomenon known in the psychological literature as choking.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [39] Chib VS Neural mechanisms underlying paradoxical performance for monetary incentives are driven by loss aversion. Neuron 2012 74 582 594 22578508 PMC3437564 source
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- cites [40] Mobbs D Choking on the money: reward-based performance decrements are associated with midbrain activity. Psychol. Sci 2009 20 955 962 19594859 PMC2931754 source
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- Premisecitation
Pavlovian-instrumental transfer effects are largely dependent on the VS.
“These effects also appear to be largely dependent on the VS [43–45].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [43] Berridge KC Dissecting components of reward: ‘liking’, ‘wanting’, and learning. Curr. Opin. Pharmacol 2009 9 65 73 19162544 PMC2756052 source
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- cites [44] Smith KS Disentangling pleasure from incentive salience and learning signals in brain reward circuitry. Proc. Natl. Acad. Sci. U.S.A 2011 108 E255 E264 21670308 PMC3131314 source
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- cites [45] Wyvell CL Berridge KC Incentive sensitization by previous amphetamine exposure: increased cue-triggered ‘wanting’ for sucrose reward. J. Neurosci 2001 21 7831 7840 11567074 PMC6762900 source
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- Premisecitation
Recent neuroimaging studies have implicated the VS in both detrimental and facilitating effects of incentives on performance.
“Recent neuroimaging studies have implicated the VS in these detrimental, as well as in the facilitating, effects of incentives on performance [39,40].”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [39] Chib VS Neural mechanisms underlying paradoxical performance for monetary incentives are driven by loss aversion. Neuron 2012 74 582 594 22578508 PMC3437564 source
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- cites [40] Mobbs D Choking on the money: reward-based performance decrements are associated with midbrain activity. Psychol. Sci 2009 20 955 962 19594859 PMC2931754 source
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- Premisecitation
Lesions and transient inactivation of the VS significantly impair previously acquired conditioned responses to food-paired CSs.
“Lesions as well as transient inactivation of the VS significantly impair previously acquired conditioned responses (CRs) to food-paired CSs”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [33] Blaiss CA Janak PH The nucleus accumbens core and shell are critical for the expression, but not the consolidation, of Pavlovian conditioned approach. Behav. Brain Res 2009 200 22 32 19159648 PMC4667776 source
Not checked in the free run.
- cites [34] Parkinson JA Dissociation in effects of lesions of the nucleus accumbens core and shell on appetitive pavlovian approach behavior and the potentiation of conditioned reinforcement and locomotor activity by D-amphetamine. J. Neurosci 1999 19 2401 2411 10066290 PMC6782569 source
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- Premisecitation
Neurophysiological studies in non-human primates and rodents have found dissociable contributions of the DMS and DLS to early versus late stages of training.
“neuro-physiological studies in non-human primates [9], as well as in rodents [10], have also found dissociable contributions of the DMS and DLS to early versus late stages of training.”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [9] Miyachi S Differential activation of monkey striatal neurons in the early and late stages of procedural learning. Exp. Brain Res 2002 146 122 126 12192586 source
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- cites [10] Yin HH Dynamic reorganization of striatal circuits during the acquisition and consolidation of a skill. Nat. Neurosci 2009 12 333 341 19198605 PMC2774785 source
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- Premisecitation
Cues that signal reward delivery independently of instrumental action can invigorate instrumental performance, a phenomenon termed Pavlovian-instrumental transfer.
“Cues that signal reward delivery independently of whether or not an instrumental action is performed can nevertheless invigorate instrumental performance, a phenomenon termed Pavlovian-instrumental transfer (PIT) [41,42].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [41] Corbit LH Balleine BW The general and outcome-specific forms of Pavlovian-instrumental transfer are differentially mediated by the nucleus accumbens core and shell. J. Neurosci 2011 31 11786 11794 21849539 PMC3208020 source
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- cites [42] Corbit LH General and outcome-specific forms of Pavlovian-instrumental transfer: the effect of shifts in motivational state and inactivation of the ventral tegmental area. Eur. J. Neurosci 2007 26 3141 3149 18005062 source
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- Premisecitation
There is considerable evidence implicating the ventral striatum in generating skeletomotor reflexes elicited by Pavlovian cues.
“There is considerable evidence to implicate the ventral striatum in generating skeletomotor reflexes elicited by Pavlovian cues [33,34].”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [33] Blaiss CA Janak PH The nucleus accumbens core and shell are critical for the expression, but not the consolidation, of Pavlovian conditioned approach. Behav. Brain Res 2009 200 22 32 19159648 PMC4667776 source
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- cites [34] Parkinson JA Dissociation in effects of lesions of the nucleus accumbens core and shell on appetitive pavlovian approach behavior and the potentiation of conditioned reinforcement and locomotor activity by D-amphetamine. J. Neurosci 1999 19 2401 2411 10066290 PMC6782569 source
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- Premisecitation
One human study found evidence in support of the postulated computational dissociation between model-based and model-free RL signals in the DMS and DLS.
“One such study found evidence in support of the postulated computational dissociation [31]”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [31] Wunderlich K Mapping value based planning and extensively trained choice in the human brain. Nat. Neurosci 2012 15 786 791 22406551 PMC3378641 source
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- Premisecitation
Neuroimaging research has implicated the DMS, more strongly than the DLS, in inhibiting responding on No Go trials.
“Neuroimaging research has implicated the DMS, more strongly than the DLS, in inhibiting responding on No Go trials [36,37].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [36] Chevrier AD Dissociation of response inhibition and performance monitoring in the stop signal task using event-related fMRI. Hum. Brain Mapp 2007 28 1347 1358 17274022 PMC6871417 source
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- cites [37] Wager TD Common and unique components of response inhibition revealed by fMRI. Neuroimage 2005 27 323 340 16019232 source
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- Premisecitation
The DLS and DMS differ in their contribution to the inhibition of competing, but incorrect responses.
“The DLS and DMS also appear to differ in their contribution to the inhibition of competing, but incorrect responses”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [36] Chevrier AD Dissociation of response inhibition and performance monitoring in the stop signal task using event-related fMRI. Hum. Brain Mapp 2007 28 1347 1358 17274022 PMC6871417 source
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- cites [37] Wager TD Common and unique components of response inhibition revealed by fMRI. Neuroimage 2005 27 323 340 16019232 source
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- Premisecitation
Blockage of NMDA receptors in the DMS during action-outcome learning abolishes sensitivity to outcome devaluation in subsequent tests.
“blockage of NMDA receptors in the DMS during action-outcome learning abolishes sensitivity to outcome devaluation in subsequent tests [19].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [19] Yin HH Blockade of NMDA receptors in the dorsomedial striatum prevents action-outcome learning in instrumental conditioning. Eur. J. Neurosci 2005 22 505 512 16045503 source
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- Premisecitation
Numerous studies have found selective involvement of the DMS in tasks requiring cognitive control and working memory.
“numerous studies have found selective involvement of the DMS in various tasks that require cognitive control and working memory [6,35,38]”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [6] Mestres-Misse A An anterior-posterior gradient of cognitive control within the dorsomedial striatum. Neuroimage 2012 62 41 47 22617829 source
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- cites [35] Poldrack RA The neural correlates of motor skill automaticity. J. Neurosci 2005 25 5356 5364 15930384 PMC6725010 source
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- cites [38] Levy R Differential activation of the caudate nucleus in primates performing spatial and nonspatial working memory tasks. J. Neurosci 1997 17 3870 3882 9133405 PMC6573717 source
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- Interpretationreasoning
The DMS is involved in the encoding of goal-directed instrumental actions.
“whereas the DMS is involved in the encoding of goal-directed instrumental actions”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationreasoning
Depending on whether such responses are compatible or incompatible with the instrumental actions needed to obtain the reward, a behavioral effect of either facilitation or impairment might occur.
“Depending on whether such responses are compatible or incompatible with the instrumental actions needed to obtain the reward, a behavioral effect of either facilitation or impairment might occur.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Premisecitation
Liking responses are altered by blockage or stimulation of Nacc opioid receptors.
“Instead, such responses are altered by blockage or stimulation of Nacc opioid receptors [44,49]”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [44] Smith KS Disentangling pleasure from incentive salience and learning signals in brain reward circuitry. Proc. Natl. Acad. Sci. U.S.A 2011 108 E255 E264 21670308 PMC3131314 source
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- cites [49] Wassum KM Distinct opioid circuits determine the palatability and the desirability of rewarding events. Proc. Natl. Acad. Sci. U.S.A 2009 106 12512 12517 19597155 PMC2718390 source
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- Premisecitation
The ventral striatum is involved in the hedonic evaluation of stimuli, termed liking.
“Another important function recently attributed to the ventral striatum is the hedonic evaluation of stimuli, termed ‘liking’, which is commonly assessed using measures of affective facial reactions [48].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [48] Berridge KC Robinson TE What is the role of dopamine in reward: hedonic impact, reward learning, or incentive salience? Brain Res. Brain Res. Rev 1998 28 309 369 9858756 source
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- Premisecitation
Conditioned responses themselves exhibit sensitivity to outcome devaluation procedures.
“CRs themselves exhibit sensitivity to outcome devaluation procedures”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [55] Gottfried JA Encoding predictive reward value in human amygdala and orbitofrontal cortex. Science 2003 301 1104 1107 12934011 source
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- cites [56] Lex B Hauber W The role of nucleus accumbens dopamine in outcome encoding in instrumental and Pavlovian conditioning. Neurobiol. Learn. Mem 2010 93 283 290 19931626 source
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- cites [57] Singh T Nucleus accumbens core and shell are necessary for reinforcer devaluation effects on pavlovian conditioned responding. Front. Integr. Neurosci 2010 4 126 21088698 PMC2981418 source
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- Interpretationreasoning
The DLS is involved in the encoding of habitual stimulus-response associations.
“and the DLS in the encoding of habitual stimulus-response associations.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationcitation
Findings implicating the ventral striatum in incentive-based performance can arguably be accounted for in terms of the role of this structure in the expression of Pavlovian conditioned responses.
“Findings implicating the ventral striatum in incentive-based performance [12,13,39,40] can arguably also be accounted for in terms of the role of this structure in the expression of Pavlovian conditioned responses.”
Quote in the paperSubstantial claim0 of 4 sources checked- cites [12] Pessiglione M How the brain translates money into force: a neuroimaging study of subliminal motivation. Science 2007 316 904 906 17431137 PMC2631941 source
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- cites [13] Schmidt L Neural mechanisms underlying motivation of mental versus physical effort. PLoS Biol 2012 10 e1001266 22363208 PMC3283550 source
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- cites [39] Chib VS Neural mechanisms underlying paradoxical performance for monetary incentives are driven by loss aversion. Neuron 2012 74 582 594 22578508 PMC3437564 source
Not checked in the free run.
- cites [40] Mobbs D Choking on the money: reward-based performance decrements are associated with midbrain activity. Psychol. Sci 2009 20 955 962 19594859 PMC2931754 source
Not checked in the free run.
- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationreasoning
Selective activation of the VS or DMS during early stages of training reflects the respective dominance of Pavlovian and goal-directed instrumental processes, rather than learning per se.
“From this perspective, selective activation of the VS or DMS during early stages of training reflects the respective dominance of Pavlovian and goal-directed instrumental processes, rather than learning per se.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationcitation
Striatal regions support dissociable associative learning strategies that may respectively dominate at various stages of training, depending on the task.
“a more parsimonious interpretation is that striatal regions support dissociable associative learning strategies that may respectively dominate at various stages of training, depending on the task [3,50,51].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [3] Balleine B Glimcher PW Multiple forms of value learning and the function of dopamine. Neuroeconomics: Decision Making and the Brain 2008 Academic Press
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- cites [50] Balleine BW O'Doherty JP Human and rodent homologies in action control: corticostriatal determinants of goal-directed and habitual action. Neuropsychopharmacology 2010 35 48 69 19776734 PMC3055420 source
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- cites [51] Yin HH Reward-guided learning beyond dopamine in the nucleus accumbens: the integrative functions of cortico-basal ganglia networks. Eur. J. Neurosci 2008 28 1437 1448 18793321 PMC2756656 source
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- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationcitation
Performance of highly skilled motor behavior or instrumental responses that necessitate approach towards aversive stimuli might be impaired by incompatible reflexes elicited by Pavlovian cues.
“Conversely, performance of highly skilled motor behavior or of instrumental responses that necessitate approach towards aversive stimuli might be impaired by incompatible reflexes elicited by Pavlovian cues [39].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [39] Chib VS Neural mechanisms underlying paradoxical performance for monetary incentives are driven by loss aversion. Neuron 2012 74 582 594 22578508 PMC3437564 source
Not checked in the free run.
- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Premisecitation
There are strong projections between the VS and the medial orbitofrontal cortex, an area well known for its involvement in utility estimation.
“It is also consistent with the strong projections between the VS and the medial orbitofrontal cortex (mOFC), an area well known for its involvement in utility estimation [53,54].”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [53] Hare TA Dissociating the role of the orbitofrontal cortex and the striatum in the computation of goal values and prediction errors. J. Neurosci 2008 28 5623 5630 18509023 PMC6670807 source
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- cites [54] Plassmann H Appetitive and aversive goal values are encoded in the medial orbitofrontal cortex at the time of decision making. J. Neurosci 2010 30 10799 10808 20702709 PMC6634706 source
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- Interpretationcitation
The separation of motivational and hedonic processes is preserved throughout the Nacc-VP circuit.
“suggesting that the separation of motivational and hedonic processes is preserved throughout the Nacc-VP circuit [44].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [44] Smith KS Disentangling pleasure from incentive salience and learning signals in brain reward circuitry. Proc. Natl. Acad. Sci. U.S.A 2011 108 E255 E264 21670308 PMC3131314 source
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- Interpretationcitation
Pavlovian retrieval of affective aspects of unconditioned stimuli contributes to the elicitation of hedonic, emotional, conditioned responses indicative of liking.
“Pavlovian retrieval of affective aspects of unconditioned stimuli contributes to the elicitation of hedonic, emotional, conditioned responses indicative of ‘liking’ [48].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [48] Berridge KC Robinson TE What is the role of dopamine in reward: hedonic impact, reward learning, or incentive salience? Brain Res. Brain Res. Rev 1998 28 309 369 9858756 source
Not checked in the free run.
- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationcitation
Pavlovian retrieval of sensory-specific features of unconditioned stimuli might evoke stimulus representations previously linked to particular instrumental responses through stimulus-response learning, and consequently elicit habitual performance of those responses at the point of Pavlovian retrieval.
“Pavlovian retrieval of sensory-specific features of unconditioned stimuli might evoke stimulus representations that have been previously linked to particular instrumental responses through stimulus-response learning and that, consequently, elicit habitual performance of those responses at the point of Pavlovian retrieval [52].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [52] Balleine BW Ostlund SB Still at the choice-point: action selection and initiation in instrumental conditioning. Ann. N. Y. Acad. Sci 2007 1104 147 171 17360797 source
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- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationreasoning
Pavlovian processes may play a role in the estimation of outcome utility, central to accounts of goal-directed instrumental performance.
“in this capacity, Pavlovian processes may also play a role in the estimation of outcome utility, central to accounts of goal-directed instrumental performance.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationcitation
Dissociable neurobiological substrates in the VS mediate motivational and hedonic processes.
“suggesting that dissociable neurobiological substrates in the VS mediate motivational and hedonic processes.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [44] Smith KS Disentangling pleasure from incentive salience and learning signals in brain reward circuitry. Proc. Natl. Acad. Sci. U.S.A 2011 108 E255 E264 21670308 PMC3131314 source
Not checked in the free run.
- cites [49] Wassum KM Distinct opioid circuits determine the palatability and the desirability of rewarding events. Proc. Natl. Acad. Sci. U.S.A 2009 106 12512 12517 19597155 PMC2718390 source
Not checked in the free run.
- Premisecitation
Behavioral expressions of liking are unaffected by amphetamine injection into the Nacc.
“behavioral expressions of liking are unaffected by amphetamine injection into the Nacc [43,44].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [43] Berridge KC Dissecting components of reward: ‘liking’, ‘wanting’, and learning. Curr. Opin. Pharmacol 2009 9 65 73 19162544 PMC2756052 source
Not checked in the free run.
- cites [44] Smith KS Disentangling pleasure from incentive salience and learning signals in brain reward circuitry. Proc. Natl. Acad. Sci. U.S.A 2011 108 E255 E264 21670308 PMC3131314 source
Not checked in the free run.
- Interpretationreasoning
It is unlikely that striatal regions differ functionally in terms of their respective contributions to learning versus performance.
“It is unlikely therefore, that these regions differ functionally in terms of their respective contributions to learning vs performance [10,11].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [10] Yin HH Dynamic reorganization of striatal circuits during the acquisition and consolidation of a skill. Nat. Neurosci 2009 12 333 341 19198605 PMC2774785 source
Not checked in the free run.
- cites [11] Atallah HE Separate neural substrates for skill learning and performance in the ventral and dorsal striatum. Nat. Neurosci 2007 10 126 131 17187065 source
Not checked in the free run.
- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Interpretationreasoning
Performance of an instrumental action involving approach towards a food location may be facilitated by Pavlovian cues that elicit compatible conditioned reflexes.
“performance of an instrumental action that involves approach towards a food location may be facilitated by the presence of Pavlovian cues that elicit compatible conditioned reflexes (i.e., directed at the same location).”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Premisecitation
Under certain training conditions, Pavlovian-instrumental transfer effects exhibit selectivity for an action that earns the same reward as signaled by the Pavlovian cue, suggesting outcome-specific representations.
“under certain training conditions, PIT effects exhibit a clear selectivity, such that instrumental responding is enhanced specifically for an action that earns the same reward as that signaled by the Pavlovian cue, suggesting the involvement of outcome-specific representations (i.e., specific PIT).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [41] Corbit LH Balleine BW The general and outcome-specific forms of Pavlovian-instrumental transfer are differentially mediated by the nucleus accumbens core and shell. J. Neurosci 2011 31 11786 11794 21849539 PMC3208020 source
Not checked in the free run.
- Interpretationreasoning
Encoding novel events may serve as a basis for exploration.
“An important aspect of encoding novel events is that they may serve as a basis for exploration.”
Quote in the paperNot a substantial claim - Premisecitation
The neuroimaging literature is profoundly inconsistent on the role of the striatum in aversive learning, with some studies reporting increased VS activity in aversive contexts and others reporting decreased activity.
“the neuroimaging literature is profoundly inconsistent on this point, with some studies reporting increased VS activity in aversive contexts (Figure 2c) [62–64] and others reporting decreasing activity in this area during the prediction, learning, and receipt of aversive outcomes [65,66].”
Quote in the paperSubstantial claim0 of 5 sources checked- cites [62] Delgado MR Neural systems underlying aversive conditioning in humans with primary and secondary reinforcers. Front. Neurosci 2011 5 71 21637321 PMC3101377 source
Not checked in the free run.
- cites [63] Jensen J Direct activation of the ventral striatum in anticipation of aversive stimuli. Neuron 2003 40 1251 1257 14687557 source
Not checked in the free run.
- cites [64] Wrase J Different neural systems adjust motor behavior in response to reward and punishment. Neuroimage 2007 36 1253 1262 17521924 source
Not checked in the free run.
- cites [65] Cooper JC Available alternative incentives modulate anticipatory nucleus accumbens activation. Soc. Cogn. Affect. Neurosci 2009 4 409 416 19843618 PMC2799954 source
Not checked in the free run.
- cites [66] Tom SM The neural basis of loss aversion in decision-making under risk. Science 2007 315 515 518 17255512 source
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- Premisecitation
DA neurons across the mesolimbic, mesocortical and nigrostriatal pathways respond phasically to novel environmental stimuli regardless of their particular valence (appetitive, aversive, or neutral).
“DA neurons across the mesolimbic, mesocortical and nigrostriatal pathways have been shown to respond phasically to novel environmental stimuli [70], regardless of their particular valence (i.e., appetitive, aversive, or neutral).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [70] Horvitz JC Mesolimbocortical and nigrostriatal dopamine responses to salient non-reward events. Neuroscience 2000 96 651 656 10727783 source
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- Premisecitation
Some data indicate that over-trained responses can be independent of dopamine more generally.
“and of DA more generally [77]”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [77] Choi WY Extended habit training reduces dopamine mediation of appetitive response expression. J. Neurosci 2005 25 6729 6733 16033882 PMC6725359 source
Not checked in the free run.
- Interpretationreasoning
The dorsomedial striatum implements far more complex functions than those outlined by RL theory.
“suggesting that this structure implements far more complex functions than those outlined by RL theory”
Quote in the paperSubstantial claim- Rests on an assumption: Deductive reasoning and problem solving have no obvious connection to reward learning.
- Premisecitation
The Nacc core and shell have both been shown to be necessary for the effects of outcome-specific devaluation on preparatory conditioned responses.
“the Nacc core and shell have both been shown to be necessary for the effects of outcome-specific devaluation on preparatory CRs [56,57].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [56] Lex B Hauber W The role of nucleus accumbens dopamine in outcome encoding in instrumental and Pavlovian conditioning. Neurobiol. Learn. Mem 2010 93 283 290 19931626 source
Not checked in the free run.
- cites [57] Singh T Nucleus accumbens core and shell are necessary for reinforcer devaluation effects on pavlovian conditioned responding. Front. Integr. Neurosci 2010 4 126 21088698 PMC2981418 source
Not checked in the free run.
- Premisecitation
Neuroimaging studies show that with extremely extended training (several weeks), slowly evolving BOLD signals in the primary motor cortex begin to discriminate between practiced and novel sequences.
“Additional support for this view comes from neuroimaging studies showing that, with extremely extended training (i.e., several weeks), slowly evolving BOLD signals in the primary motor cortex (M1) begin to discriminate between practiced and novel sequences [79].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [79] Ungerleider LG Imaging brain plasticity during motor skill learning. Neurobiol. Learn. Mem 2002 78 553 564 12559834 source
Not checked in the free run.
- Interpretationcitation
The same stimulus can be perceived and neurally encoded as both pleasant and aversive depending on the subject's internal context.
“Thus, the same stimulus was perceived, and neurally encoded, as both pleasant and aversive depending on the subject's internal context.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
- Premisecitation
Some data indicate that over-trained responses can be independent of the dorsolateral striatum specifically.
“some data indicate that over-trained responses can be independent of the DLS specifically [76]”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [76] Turner RS Bolam JP Sequential motor behavior and the basal ganglia. The basal ganglia VIII (Advances in Behavioral Biology) 2005 563 574 Springer
Not checked in the free run.
- Premisecitation
Some behavioral evidence from rodents suggests that novelty may serve as an instrumental reinforcer, such that rats will press a lever that produces an apparently neutral light stimulus more than a lever that does not yield any outcome.
“Indeed, some behavioral evidence from rodents suggests that novelty may serve as an instrumental reinforcer, such that rats will press a lever that produces an apparently neutral light stimulus more than a lever that does not yield any outcome [74].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [74] Reed P Intrinsic reinforcing properties of putatively neutral stimuli in an instrumental two-lever discrimination. Anim. Learn. Behav 1996 24 38 45 source
Not checked in the free run.
- Premisecitation
There is overwhelming evidence for a role of the dorsolateral striatum in performance of well-learned motor programs.
“there is overwhelming evidence for a role of the DLS in performance of well-learned motor programs [5,7–10,18]”
Quote in the paperSubstantial claim0 of 6 sources checked- cites [5] Jankowski J Distinct striatal regions for planning and executing novel and automated movement sequences. Neuroimage 2009 44 1369 1379 19059350 source
Not checked in the free run.
- cites [7] Grafton ST Functional mapping of sequence learning in normal humans. Cogn. Neurosci 1995 7 497 510 23961907 source
Not checked in the free run.
- cites [8] Jueptner M Anatomy of motor learning. I. Frontal cortex and attention to action. J. Neurophysiol 1997 77 1313 1324 9084599 source
Not checked in the free run.
- cites [9] Miyachi S Differential activation of monkey striatal neurons in the early and late stages of procedural learning. Exp. Brain Res 2002 146 122 126 12192586 source
Not checked in the free run.
- cites [10] Yin HH Dynamic reorganization of striatal circuits during the acquisition and consolidation of a skill. Nat. Neurosci 2009 12 333 341 19198605 PMC2774785 source
Not checked in the free run.
- cites [18] Yin HH Lesions of dorsolateral striatum preserve outcome expectancy but disrupt habit formation in instrumental learning. Eur. J. Neurosci 2004 19 181 189 14750976 source
Not checked in the free run.
- Premisecitation
Tasks such as deductive reasoning and problem solving seem to recruit strongly the dorsomedial striatum.
“seem nonetheless to recruit strongly the DMS [80,81]”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [80] Rodriguez-Moreno D Hirsch J The dynamics of deductive reasoning: an fMRI investigation. Neuropsychologia 2009 47 949 961 18835284 source
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- cites [81] Stocco A Anderson JR Endogenous control and task representation: an fMRI study in algebraic problem-solving. J. Cogn. Neurosci 2008 20 1300 1314 18284348 source
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- Premisecitation
In rodents, the firing of ventral pallidal neurons in response to an intense salt solution tracks behavioral changes in hedonic processing between normal and salt-deprived states.
“the response patterns of VP neurons closely tracked such behavioral changes, showing a dramatic increase in response to the salt-solution in the deprived relative to the normal state [69].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [69] Tindell AJ Ventral pallidum firing codes hedonic reward: when a bad taste turns good. J. Neurophysiol 2006 96 2399 2409 16885520 source
Not checked in the free run.
- Premisecitation
In the ventral striatum specifically, responses to novel stimuli have been shown with fast-scan voltammetry and other techniques measuring extracellular DA concentrations, as well as with single unit recordings and fMRI.
“In the VS specifically, responses to novel stimuli have been shown with fast-scan voltammetry and other techniques measuring extra-cellular DA concentrations, as well as with single unit recordings and fMRI [71–73].”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [71] Ljungberg T Responses of monkey dopamine neurons during learning of behavioral reactions. J. Neurophysiol 1992 67 145 163 1552316 source
Not checked in the free run.
- cites [72] Rebec GV Real-time assessments of dopamine function during behavior: single-unit recording, iontophoresis, and fast-scan cyclic voltammetry in awake, unrestrained rats. Alcohol. Clin. Exp. Res 1998 22 32 40 9514283 source
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- cites [73] Wittmann BC Striatal activity underlies novelty-based choice in humans. Neuron 2008 58 967 973 18579085 PMC2535823 source
Not checked in the free run.
- Interpretationnone given
There is a dissociation within the dorsal striatum, between medial and lateral structures, in the implementation of goal-directed and habitual instrumental strategies.
“Moreover, there is a dissociation within the dorsal striatum – between medial and lateral structures – in the implementation of goal-directed and habitual instrumental strategies.”
Quote in the paperNot a substantial claim - Interpretationreasoning
Through its role in the learning and expression of Pavlovian conditioned responses, rather than through its role in motivation per se, the ventral striatum supports a range of modulatory influences on instrumental performance, including general invigoration, response selection, and potentially even goal-directed outcome evaluation.
“Finally, through its role in the learning and expression of Pavlovian conditioned responses, rather than, perhaps, through its role in motivation per se, the ventral striatum supports a range of modulatory influences on instrumental performance, including general invigoration (e.g., general PIT), response selection (specific PIT), and potentially even goal-directed outcome evaluation.”
Quote in the paperSubstantial claim - Interpretationreasoning
Striatal subregions appear to implement distinct forms of associative encoding, rather than being divided along lines of learning versus performance.
“We have noted that, rather than being divided along lines of learning versus performance, striatal subregions appear to implement distinct forms of associative encoding.”
Quote in the paperSubstantial claim
- May combine two claims. 19
- Classified as probably not a substantial claim. 11
May combine two claims.
PremisecitationAmphetamine injection into the Nacc enhances Pavlovian-instrumental transfer without affecting base rates of instrumental responding.
“amphetamine injection into the Nacc enhances PIT, without affecting base rates of instrumental responding [45].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [45] Wyvell CL Berridge KC Incentive sensitization by previous amphetamine exposure: increased cue-triggered ‘wanting’ for sucrose reward. J. Neurosci 2001 21 7831 7840 11567074 PMC6762900 source
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe actor/critic model cannot account for the known differences between goal-directed and habitual instrumental actions, and the differential functions of the DMS and DLS.
“A major limitation of the actor/critic model is that it cannot account for the known differences between goal-directed and habitual instrumental actions, and the differential functions of the DMS and DLS in supporting these mechanisms.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationA large body of research implicates the dorsal striatum in the implementation of already learned instrumental motor behaviors, with dissociations between the DLS and DMS.
“A large body of research has also implicated the dorsal striatum in the implementation of already learned instrumental motor behaviors, often with dissociations emerging between the DLS and DMS [7–10].”
Quote in the paperNot a substantial claim0 of 4 sources checked- cites [7] Grafton ST Functional mapping of sequence learning in normal humans. Cogn. Neurosci 1995 7 497 510 23961907
Not checked in the free run.
- cites [8] Jueptner M Anatomy of motor learning. I. Frontal cortex and attention to action. J. Neurophysiol 1997 77 1313 1324 9084599
Not checked in the free run.
- cites [9] Miyachi S Differential activation of monkey striatal neurons in the early and late stages of procedural learning. Exp. Brain Res 2002 146 122 126 12192586 source
Not checked in the free run.
- cites [10] Yin HH Dynamic reorganization of striatal circuits during the acquisition and consolidation of a skill. Nat. Neurosci 2009 12 333 341 19198605 PMC2774785 source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationAnother human study found evidence for a linear mix of model-based and model-free signals within the same overlapping areas.
“whereas another study, using a similar design, instead found evidence for a linear mix of model-based and model-free signals within the same overlapping areas [32].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [32] Daw ND Model-based influences on humans’ choices and striatal prediction errors. Neuron 2011 69 1204 1215 21435563 PMC3077926 source
Not checked in the free run.
May combine two claims.
PremisecitationCues indicating reward availability elicit increases in VS activity proportional to the amount of signaled reward, and these signals correlate with performance enhancement for larger compared to smaller rewards.
“Cues that indicate that a certain amount of reward is available given successful performance of an instrumental action, or even of a complex cognitive task, elicit increases in VS activity proportional to the amount of signaled reward and these signals correlate with the degree of performance enhancement found for larger compared to smaller rewards [12,13].”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [12] Pessiglione M How the brain translates money into force: a neuroimaging study of subliminal motivation. Science 2007 316 904 906 17431137 PMC2631941 source
Not checked in the free run.
- cites [13] Schmidt L Neural mechanisms underlying motivation of mental versus physical effort. PLoS Biol 2012 10 e1001266 22363208 PMC3283550 source
Not checked in the free run.
May combine two claims.
PremisecitationIn serial reaction time tasks, the DMS appears active during learning of novel sequences, whereas the DLS is active during performance of well-learned sequences.
“using a serial reaction time (SRT) task, in which participants respond to a sequence of consecutively presented stimuli, several neuroimaging studies have reported that, whereas the DMS appears to be active during learning of novel sequences, the DLS is active during performance of well-learned sequences [7,8]”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [7] Grafton ST Functional mapping of sequence learning in normal humans. Cogn. Neurosci 1995 7 497 510 23961907
Not checked in the free run.
- cites [8] Jueptner M Anatomy of motor learning. I. Frontal cortex and attention to action. J. Neurophysiol 1997 77 1313 1324 9084599
Not checked in the free run.
May combine two claims.
PremisecitationBoth lesions and transient inactivation of the DMS abolish the sensitivity of previously acquired actions to outcome devaluation and contingency degradation.
“both lesions and transient inactivation of the DMS abolish the sensitivity of previously acquired actions to outcome devaluation and contingency degradation – behavioral assays of goal-directed performance [20].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [20] Yin HH The role of the dorsomedial striatum in instrumental conditioning. Eur. J. Neurosci 2005 22 513 523 16045504 source
Not checked in the free run.
May combine two claims.
PremisecitationPavlovian-instrumental transfer effects can be general, emerging even when the instrumental action earns a different reward than that signaled by the cue and being attenuated by general motivational shifts from hunger to satiety.
“PIT effects emerge even when the instrumental action earns a different reward than that signaled by the cue and are attenuated by general motivational shifts from hunger to satiety [42], suggesting that the cue induces a general motivational state (i.e., general PIT).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [42] Corbit LH General and outcome-specific forms of Pavlovian-instrumental transfer: the effect of shifts in motivational state and inactivation of the ventral tegmental area. Eur. J. Neurosci 2007 26 3141 3149 18005062 source
Not checked in the free run.
Classified as probably not a substantial claim.
PremisecitationGoal-directed instrumental behavior can be accounted for by a model-based type of reinforcement learning.
“This shortcoming has been addressed by the proposal that goal-directed instrumental behavior can be accounted for by means of a ‘model-based’ type of RL”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [25] Daw ND Uncertainty-based competition between prefrontal and dorsolateral striatal systems for behavioral control. Nat. Neurosci 2005 8 1704 1711 16286932 source
Not checked in the free run.
May combine two claims.
InterpretationcitationDMS disruptions impair the expression of goal-directed behavior, suggesting that the DMS plays a critical role during performance.
“Thus, DMS disruptions impair the expression of goal-directed behavior, suggesting that this structure plays a critical role during performance.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [20] Yin HH The role of the dorsomedial striatum in instrumental conditioning. Eur. J. Neurosci 2005 22 513 523 16045504 source
Not checked in the free run.
May combine two claims.
PremisecitationFindings from rodent lesion and inactivation studies suggest that the Nacc shell and core may mediate specific and general Pavlovian-instrumental transfer, respectively.
“Findings from rodent lesion and inactivation studies suggest that the Nacc shell and core may mediate specific and general PIT, respectively [41].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [41] Corbit LH Balleine BW The general and outcome-specific forms of Pavlovian-instrumental transfer are differentially mediated by the nucleus accumbens core and shell. J. Neurosci 2011 31 11786 11794 21849539 PMC3208020
Not checked in the free run.
May combine two claims.
PremisecitationPavlovian conditioned responses, as well as BOLD signals in the VS, show clear sensitivity to outcome-specific devaluation.
“Pavlovian CRs, as well as BOLD signals in the VS, show clear sensitivity to outcome-specific devaluation [55–57].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [55] Gottfried JA Encoding predictive reward value in human amygdala and orbitofrontal cortex. Science 2003 301 1104 1107 12934011 source
Not checked in the free run.
- cites [56] Lex B Hauber W The role of nucleus accumbens dopamine in outcome encoding in instrumental and Pavlovian conditioning. Neurobiol. Learn. Mem 2010 93 283 290 19931626
Not checked in the free run.
- cites [57] Singh T Nucleus accumbens core and shell are necessary for reinforcer devaluation effects on pavlovian conditioned responding. Front. Integr. Neurosci 2010 4 126 21088698 PMC2981418
Not checked in the free run.
May combine two claims.
PremisecitationBoth dopaminergic and opioidergic manipulations of the Nacc modulate the firing of VP neurons in response to reward-proximal Pavlovian cues, but only opioid manipulations alter VP firing in response to unconditioned stimuli.
“although both dopaminergic and opioidergic manipulations of the Nacc modulate the firing of VP neurons in response to (reward proximal) Pavlovian cues, only opioid manipulations alter VP firing in response to unconditioned stimuli”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [44] Smith KS Disentangling pleasure from incentive salience and learning signals in brain reward circuitry. Proc. Natl. Acad. Sci. U.S.A 2011 108 E255 E264 21670308 PMC3131314 source
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe evidence reviewed implicates the ventral and dorsal striatum (both the DLS and DMS) in the learning as well as the performance of reward-related behaviors.
“The evidence reviewed here has implicated the ventral and dorsal striatum (both the DLS and DMS) in the learning as well as the performance of reward-related behaviors.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
May combine two claims.
PremisecitationHuman neuroimaging studies have demonstrated involvement of the medial VS in a form of Pavlovian-instrumental transfer that may depend on general motivational processes, and of the ventrolateral striatum in specific Pavlovian-instrumental transfer.
“More recently, the involvement of the medial VS in a form of PIT that may depend on general motivational processes [46], and of the ventrolateral striatum in specific PIT [47], has been demonstrated in human neuroimaging studies”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [46] Talmi D Human pavlovian-instrumental transfer. J. Neurosci 2008 28 360 368 18184778 PMC2636904
Not checked in the free run.
- cites [47] Bray S The neural mechanisms underlying the influence of Pavlovian cues on human decision making. J. Neurosci 2008 28 5861 5866 18509047 PMC6670800
Not checked in the free run.
May combine two claims.
Interpretationnone givenThe ventral striatum is involved in the encoding of Pavlovian associations, supporting generation of conditioned skeletomotor responses.
“the ventral striatum is involved in the encoding of Pavlovian associations, supporting generation of conditioned skeletomotor responses”
Quote in the paperNot a substantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationSome studies have reported that aversive stimuli inhibit DA activity of midbrain neurons, whereas others have found that they elicit phasic DA release in the VS.
“whereas some studies have reported that aversive stimuli inhibit the DA activity of midbrain neurons (e.g., [67]), others have found that they elicit phasic DA release in the VS (e.g., [68]).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [67] Mirenowicz J Schultz W Preferential activation of midbrain dopamine neurons by appetitive rather than aversive stimuli. Nature 1996 379 449 451 8559249 source
Not checked in the free run.
- cites [68] Budygin EA Aversive stimulus differentially triggers subsecond dopamine release in reward regions. Neuroscience 2012 201 331 337 22108611 PMC3258373 source
Not checked in the free run.
Probably not a substantial claim: may not pass S1 (asserted), S3 (load-bearing).
PremisecitationWith extended training, control is transferred to less plastic but considerably faster cortical-cortical projections.
“with extended training, control is transferred to these less plastic, but considerably faster, cortical-cortical projections [78]”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [78] Ashby FG Cortical and basal ganglia contributions to habit learning and automaticity. Trends Cogn. Sci 2010 14 208 215 20207189 PMC2862890 source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationExploratory sampling might be guided by uncertainty about the relationships between actions and rewards.
“For example, exploratory sampling might be guided by uncertainty about the relationships between actions and rewards [82].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [82] Badre D Rostrolateral prefrontal cortex and individual differences in uncertainty-driven exploration. Neuron 2012 73 595 607 22325209 PMC3285405
Not checked in the free run.
Probably not a substantial claim: may not pass S1 (asserted), S3 (load-bearing).
PremisecitationReinforcement-learning in the striatum provides a basis for successful Hebbian learning in sensory and premotor cortices.
“it has been suggested that reinforcement-learning in the striatum provides a basis for successful Hebbian learning in sensory and premotor cortices”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [78] Ashby FG Cortical and basal ganglia contributions to habit learning and automaticity. Trends Cogn. Sci 2010 14 208 215 20207189 PMC2862890 source
Not checked in the free run.
Classified as probably not a substantial claim.
InterpretationcitationVentral striatal responses are strongly contextually dependent.
“One possible reason for these variable findings might be that ventral striatal responses are strongly contextually dependent.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).
InterpretationreasoningIn many cases, the basis for exploration of non-optimal response alternatives is likely more principled than that afforded by the soft-max rule.
“However, in many cases, the basis for exploration of non-optimal response alternatives, permitting discovery of actions that are more rewarding than those sampled thus far, is likely more principled than that afforded by the soft-max rule.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIn RL theory, one simple way to implement action selection in either a model-based or model-free learner is to use a soft-max distribution, in which a free parameter controls the degree to which choices are biased towards the highest valued action.
“In RL theory, one simple way to implement action selection in either a model-based or model-free learner is to use a soft-max distribution [24,25], in which a free parameter controls the degree to which choices are biased towards the highest valued action.”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [24] Sutton R Barto A Reinforcement Learning: An Introduction 1998 MIT Press
Not checked in the free run.
- cites [25] Daw ND Uncertainty-based competition between prefrontal and dorsolateral striatal systems for behavioral control. Nat. Neurosci 2005 8 1704 1711 16286932 source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationSeveral modified RL algorithms have been proposed that incorporate novel event signaling, either as a surrogate of reward or as a component of the estimated state value.
“Several modified RL algorithms have been proposed that incorporate novel event signaling, either as a surrogate of reward or as a component of the estimated state value [75].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [75] Kakade S Dayan P Dopamine: generalization and bonuses. Neural Netw 2002 15 549 559 12371511
Not checked in the free run.
May combine two claims.
InterpretationreasoningHumans, as well as other animals, are capable of learning about the specific features of Pavlovian outcomes, and the VS appears to play a role in such effects.
“it is clear that humans, as well as other animals, are capable of learning about the specific features of Pavlovian outcomes and that the VS appears to play a role in such effects.”
Quote in the paperSubstantial claim- Rests on an assumption: The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
May combine two claims.
InterpretationreasoningThe striatum as a whole is implicated in learning of both instrumental actions and Pavlovian conditioned responses, in the expression of such learned behaviors, and in controlling the motivation to respond.
“In this article, we have reviewed evidence implicating the striatum as a whole in a number of distinct processes underlying reinforcement-related motor behavior: in learning of both instrumental actions and Pavlovian conditioned responses, in the expression of such learned behaviors, and in controlling the motivation to respond.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationDrift diffusion models have been shown to successfully capture perceptual decision-making, value-based decision-making, and the firing rates of neurons in the lateral intraparietal area of the monkey brain.
“DDMs have been shown to successfully capture perceptual [84] and value-based [85] decision-making, as well as the firing rates of neurons in the lateral intraparietal area of the monkey brain [84].”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [84] Churchland AK Decision-making with multiple alternatives. Nat. Neurosci 2008 11 693 702 18488024 PMC2453226 source
Not checked in the free run.
- cites [85] Krajbich I Visual fixations and the computation and comparison of value in simple choice. Nat. Neurosci 2010 13 1292 1298 20835253
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe ventral striatum is more involved in Pavlovian conditioned responses, whereas the dorsal striatum is involved in instrumental action.
“Specifically, the ventral striatum is more involved in Pavlovian conditioned responses, whereas the dorsal striatum is involved in instrumental action.”
Quote in the paperNot a substantial claim
The paper relies on this without support.
The functional roles of striatal sub-regions can be inferred by comparing lesion, inactivation, and neuroimaging findings across rodents, non-human primates, and humans.
“Here, we consider empirical data from human and animal studies, as well as theoretical notions from both the psychological and computational literatures”
The paper relies on this without support.
Deductive reasoning and problem solving have no obvious connection to reward learning.
“which have no obvious connection to reward learning”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The dorsolateral striatum is more involved in stimulus-response learning and habitual control, the dorsomedial striatum is linked to action-outcome learning and goal-directed action, and ventral striatum to Pavlovian values.
“For example, the dorsolateral striatum is thought to be more involved in stimulus-response learning and habitual control, the dorsomedial striatum has been linked to action-outcome learning and goal-directed action, and ventral striatum to Pavlovian values (Glascher et al., 2010, Liljeholm and O’Doherty, 2012, Mannella et al., 2013, O’Doherty et al., 2004).”
From Active Inference, homeostatic regulation and adaptive behavioural control (2015) · cites this paper as Liljeholm and O’Doherty, 2012
The striatum is a key neural structure involved in reward-related processing, from recognizing and evaluating rewards to learning from them to predict the best potential reward in the future.
“The striatum is a key neural structure involved in reward-related processing, from recognizing and evaluating rewards to learning from them as to predict the best potential reward in the future for review see 1, 10, 11.”
From The social brain and reward: Social information processing in the human striatum (2014) · cites this paper as [11]
Reward-related signals in the striatum are crucial for alerting and motivating an organism to a reward opportunity, associating predictive cues with reward outcomes, and learning through trial and error in order to adapt behavior to attain rewards in the future.
“Thus, reward-related signals in the striatum appear to be crucial for alerting and motivating an organism to a reward opportunity, associating predictive cues with reward outcomes, and learning through trial and error in order to adapt behavior to attain rewards in the future1, 10, 11.”
From The social brain and reward: Social information processing in the human striatum (2014) · cites this paper as [11]
The citation lists haven’t been fetched yet
References 107 · Cited by 317
- Authors
- Liljeholm, M., & O’Doherty, J. P.
- Affiliations
- California Institute of Technology (US)
- Corresponding author
- Liljeholm, Mimi
- O’Doherty, John P.
- Venue
- Trends in Cognitive Sciences
- Publisher
- Elsevier BV
- Published
- 2012-09
- Volume
- 16
- Issue
- 9
- Pages
- 467-475
- ISSN
- 1364-6613 (ISSN-L)
- PMC
- PMC3449003
- OpenAlex
- W2008441351
- Semantic Scholar
- 7bfa5ff0c9c4c443185c7c6bb00c721a495a204d
- Cited by
- 317
- References
- 107
- Citations by year
- 2026: 11 · 2025: 17 · 2024: 22 · 2023: 21 · 2022: 17 · 2021: 33 · 2020: 25 · 2019: 23 · 2018: 25 · 2017: 34 · 2016: 19 · 2015: 30 · 2014: 23 · 2013: 14 · 2012: 2
- FWCI
- 1.75
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Trends in Cognitive Sciences (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- CaltechAUTHORS (California Institute of Technology) (submittedVersion)
- Funders
- National Institute on Drug Abuse (R01DA033077)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural and Behavioral Psychology Studies
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- striatum, associative learning, reward learning, motivation, response selection, functional specialization, computational models of learning, motor behavior
- MeSH terms
- Animals, Association Learning, Basal Ganglia, Brain Mapping, Corpus Striatum, Humans, Inhibition, Psychological, Motivation, Neural Pathways, Psychomotor Performance, Reward, Functional Neuroimaging
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