Bonsai trees in your head: how the pavlovian system sculpts goal-directed choices by pruning decision trees
Open accessAbstract
When planning a series of actions, it is usually infeasible to consider all potential future sequences; instead, one must prune the decision tree. Provably optimal pruning is, however, still computationally ruinous and the specific approximations humans employ remain unknown. We designed a new sequential reinforcement-based task and showed that human subjects adopted a simple pruning strategy: during mental evaluation of a sequence of choices, they curtailed any further evaluation of a sequence as soon as they encountered a large loss. This pruning strategy was Pavlovian: it was reflexively evoked by large losses and persisted even when overwhelmingly counterproductive. It was also evident above and beyond loss aversion. We found that the tendency towards Pavlovian pruning was selectively predicted by the degree to which subjects exhibited sub-clinical mood disturbance, in accordance with theories that ascribe Pavlovian behavioural inhibition, via serotonin, a role in mood disorders. We conclude that Pavlovian behavioural inhibition shapes highly flexible, goal-directed choices in a manner that may be important for theories of decision-making in mood disorders.
AI snapshot
Written from the paper’s full text.
TL;DR
Humans planning sequences prune decision trees by curtailing evaluation after large losses, a Pavlovian response that persists even when counterproductive. Computational modeling shows this specific pruning is distinct from general discounting and loss aversion, and correlates with subclinical depressive symptoms.
Key findings
- Subjects adopted a simple pruning strategy: they stopped evaluating a sequence when they encountered a large loss, even when this was costly. In group −140 pruning was never disadvantageous, but in groups −70 and −100 it was increasingly counterproductive, yet subjects continued to prune.
- The most parsimonious model was “Pruning & Learned,” which separated general pruning from specific pruning after large punishments. Specific pruning exceeded general pruning in 14/15, 12/16, and 14/15 subjects in the −70, −100, and −140 groups, respectively. This model correctly predicted 91% of choices; the full look-ahead model predicted 77% of choices and had a mean predictive probability of 0.71.
- Pruning was evident above and beyond loss aversion. Subjects were approximately 1.4 times more sensitive to punishments than to equal-sized rewards, but adding loss aversion to the best model did not improve parsimony.
- BDI score positively correlated with the specific pruning parameter, but not with general pruning. A negative correlation between BDI and reward sensitivity did not survive correction for multiple comparisons. NEO agreeableness positively correlated with the Learned Pavlovian influence and survived full correction for 60 comparisons; no significant correlation with NEO neuroticism was found.
Methods
- Forty-six volunteers (23 female, mean age 23.84 years) were recruited and screened; groups received different large losses: −70 (n=15), −100 (n=16), and −140 (n=15). Subjects learned a deterministic six-state transition matrix, then completed two blocks of 24 choice episodes with sequence lengths of 2–8; the first block was discarded and the second analyzed. On half the trials, subjects planned their last 2–4 moves together without intermittent feedback.
- Computational models included full look-ahead, discounting, pruning (general and specific), learned Pavlovian attraction/repulsion, loss aversion, and a habitual SARSA model. Models were fit with Bayesian model fitting and compared using integrated Bayesian Information Criterion at the group level.
- Psychometric correlates were assessed with a weighted hierarchical multivariate regression, orthogonalizing age, sex, IQ, Digit Span, NEO measures, STAI, and BDI.
Limitations
- The paper states that the sample was healthy volunteers with only very mild depressive symptoms, so the BDI correlation cannot be taken as direct evidence that mildly raised BDI scores measure vulnerability to depression; predictions about clinical depression and serotonin depletion remain to be tested. It also states that no significant correlation with NEO neuroticism was found.
- The paper notes that in group −140 there was always an optimal sequence avoiding the large loss, making that group’s behavioral statistic difficult to interpret; at depths of 6 or more, the number of events was small, limiting power; and some missing age/STAI data excluded subjects from psychometric analyses.
- Evident from the text: the task uses a fixed deterministic transition matrix and specific reinforcement structure, and the conclusions depend on the fitted model space; model comparison did not support separate group differences in pruning despite apparent behavioral differences.
Model-written summary. Check it against the paper before relying on it.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
0 of 29 cited sources checked · 0 supported · 0 flagged
- Findingown data
Bayesian model comparison found strong evidence for separating discounting into general and specific pruning parameters.
“Again, we applied Bayesian model comparison and found strong evidence for such a separation (Figure 4C).”
Quote in the paperSubstantial claim - Findingown data
BDI was positively correlated with the specific pruning parameter.
“We found that BDI was positively correlated with the specific pruning parameter ().”
Quote in the paperSubstantial claim- Rests on an assumption: The BDI validly measures sub-clinical depressive symptoms/mood disturbance.
- Findingown data
The correlation between BDI and specific pruning was specific in that there was no such correlation with general pruning.
“Furthermore, this effect was specific in that there was no such correlation with general pruning .”
Quote in the paperSubstantial claim- Rests on an assumption: The BDI validly measures sub-clinical depressive symptoms/mood disturbance.
- Findingown data
Subjects were significantly seduced by the effect of past rewards (or repulsed by punishments), in a way counterproductive to optimal control.
“Subjects were significantly seduced by the effect of these past rewards (or repulsed by punishments), again in a way that was counterproductive to optimal control.”
Quote in the paperSubstantial claim - Findingown data
Subjects were worse at choosing optimal sequences when the depth was greater.
“Subjects were worse at choosing optimal sequences when the depth was greater.”
Quote in the paperSubstantial claim
- Findingown data
Human subjects adopted a simple pruning strategy: during mental evaluation of a sequence of choices, they curtailed further evaluation of a sequence as soon as they encountered a large loss.
“We designed a new sequential reinforcement-based task and showed that human subjects adopted a simple pruning strategy: during mental evaluation of a sequence of choices, they curtailed any further evaluation of a sequence as soon as they encountered a large loss.”
Quote in the paperSubstantial claim - Findingown data
Subjects showed a substantial increase in their propensity to prune in the face of a large negative outcome.
“subjects also showed a substantial increase in their propensity to prune in the face of a large negative outcome (parameter in the model).”
Quote in the paperSubstantial claim- Rests on an assumption: The large losses in the task are aversive/punishing events capable of evoking Pavlovian behavioural inhibition.
- Findingown data
There was a positive correlation between NEO agreeableness and the weight of the 'Learned Pavlovian' influence which survived full correction for 60 comparisons.
“We found a positive correlation between NEO agreeableness and the weight of the ‘Learned Pavlovian’ influence which survived full correction for 60 comparisons .”
Quote in the paperNot a substantial claim - Findingown data
The tendency to excise subtrees lying below punishments persisted even when counterproductive in terms of outcomes.
“That is, the tendency to excise subtrees lying below punishments persisted even when counterproductive in terms of outcomes.”
Quote in the paperSubstantial claim - Findingown data
Subjects were no less likely to prune even when we rendered it increasingly disadvantageous (moving from group −140 to group −70).
“It was striking that subjects were no less likely to prune (Figure 2C–D) even when we rendered it increasingly disadvantageous (moving from group −140 to group −70),.”
Quote in the paperSubstantial claim - Findingown data
Loss models suggest subjects were more sensitive to punishments than rewards.
“The ‘Loss’ models do suggest that subjects were more sensitive to punishments than rewards (Figure 5C).”
Quote in the paperNot a substantial claim - Findingown data
The tendency towards Pavlovian pruning was selectively predicted by the degree to which subjects exhibited sub-clinical mood disturbance.
“We found that the tendency towards Pavlovian pruning was selectively predicted by the degree to which subjects exhibited sub-clinical mood disturbance, in accordance with theories that ascribe Pavlovian behavioural inhibition, via serotonin, a role in mood disorders.”
Quote in the paperSubstantial claim- Rests on an assumption: The BDI validly measures sub-clinical depressive symptoms/mood disturbance.
- Findingown data
Subjects were less willing to choose optimal sequences involving transitions through large losses than those that did not.
“Subjects were also less willing to choose optimal sequences involving transitions through large losses (shown in blue) than those that did not (shown in green).”
Quote in the paperSubstantial claim - Findingown data
Figure 4G shows that the deeper-tree effect can arise from the interaction of the particular sequence of reinforcements (and thus reinforcement depth) and the pruning and discount factors.
“Figure 4G shows that this can arise from the interaction of the particular sequence of reinforcements (and thus reinforcement depth) and the pruning and discount factors.”
Quote in the paperNot a substantial claim - Findingown data
In the first experimental group, subjects pruned extensively, tending to ignore subtrees lying beyond large losses.
“Subjects here pruned extensively: they tended to ignore subtrees lying beyond large losses.”
Quote in the paperSubstantial claim - Findingown data
Subjects pruned to a very substantial degree.
“First, subjects pruned to a very substantial degree.”
Quote in the paperSubstantial claim - Findingown data
When subjects had only a single choice left in state 6, on about 40% of such trials they were attracted to state 1, despite it no longer being optimal.
“Despite this, on about 40% of such trials, subjects were attracted to state 1 (Figure 4E).”
Quote in the paperSubstantial claim - Findingown data
Subjects chose to move from state 6 to state 1 on 90% of occasions when they had two or more choices left, in all three groups.
“Subjects duly chose to move to state 1 on 90% of these occasions in all three groups.”
Quote in the paperSubstantial claim - Findingown data
Subjects did so even at the expense of a substantial net loss in reward.
“Importantly, they did so even at the expense of a substantial net loss in reward.”
Quote in the paperSubstantial claim - Findingown data
The pruning strategy was evident above and beyond loss aversion.
“It was also evident above and beyond loss aversion.”
Quote in the paperSubstantial claim - Findingown data
Subjects' propensity to prune specifically in the face of negative valence was positively correlated with self-reported sub-clinical depressive symptoms.
“we found that subjects' propensity to prune specifically in the face of negative valence was positively correlated with self-reported sub-clinical depressive symptoms.”
Quote in the paperSubstantial claim - Findingreasoning
Loss models did not explain away pruning.
“However, this did not explain away pruning.”
Quote in the paperSubstantial claim - Findingown data
The sample of healthy volunteers reported only very mild depressive symptoms (mean BDI scores of , range ).
“our sample of healthy volunteers, which was thoroughly screened for past pathology, reported only very mild depressive symptoms (with mean BDI scores of , range )”
Quote in the paperNot a substantial claim - Findingown data
There were no significant correlations with NEO neuroticism.
“However, against our hypothesis, we did not identify any significant correlations with NEO neuroticism.”
Quote in the paperNot a substantial claim - Findingown data
Learned Pavlovian influence was associated with the learned attractiveness of previously rewarded states.
“The second, ‘Learned’, Pavlovian influence was associated with the learned attractiveness of previously rewarded states.”
Quote in the paperSubstantial claim - Findingreasoning
Pavlovian pruning appeared to have a particularly strong effect deeper in the tree.
“Pavlovian pruning thus appeared to have a particularly strong effect deeper in the tree.”
Quote in the paperNot a substantial claim - Findingown data
We found a relationship between the propensity to prune and sub-clinical mood disturbance.
“we found a relationship between the propensity to prune and sub-clinical mood disturbance”
Quote in the paperSubstantial claim - Findingown data
The restricted loss aversion models led to the same conclusions as the full loss aversion models.
“The restricted models led to the same conclusions as the full loss aversion models and we thus do not report those results.”
Quote in the paperNot a substantial claim - Findingown data
Most subjects were unable to resist pruning even when it was disadvantageous.
“most subjects were unable to resist pruning even when it was disadvantageous”
Quote in the paperSubstantial claim - Interpretationown data
Pavlovian behavioural inhibition shapes highly flexible, goal-directed choices in a manner that may be important for theories of decision-making in mood disorders.
“We conclude that Pavlovian behavioural inhibition shapes highly flexible, goal-directed choices in a manner that may be important for theories of decision-making in mood disorders.”
Quote in the paperSubstantial claim- Rests on an assumption: The large losses in the task are aversive/punishing events capable of evoking Pavlovian behavioural inhibition.
- Rests on an assumption: The BDI validly measures sub-clinical depressive symptoms/mood disturbance.
- Study design and conductreasoning
The deeper-tree effect is not a feature explicitly built into the models.
“Although this makes intuitive sense, it is not a feature explicitly built into the models.”
Quote in the paperNot a substantial claim - Study design and conductreasoning
Including the second Pavlovian influence ensured that the pruning described was a pure influence on goal-based evaluation, not corrupted by intrinsic repulsion to punishment.
“Note that by including this second Pavlovian influence, we could be sure that the pruning described above was a pure influence on goal-based evaluation, and was not corrupted by an intrinsic repulsion to the punishment (which would have been ascribed to this second, Pavlovian, influence).”
Quote in the paperSubstantial claim - Interpretationown data
The best-performing model accounted so well for subjects' choices that it suggests it was a sufficient mechanism for the particular set of reinforcement sequences encountered here.
“Although this is not necessarily always the case, the fact that our best-performing model accounted so well for subjects' choices (Figure 4A) suggests that it was a sufficient mechanism for the particular set of reinforcement sequences encountered here.”
Quote in the paperSubstantial claim - Premisecitation
Orbitofrontal serotonin depletion impacts cognitive flexibility by impairing inhibitory control in monkeys.
“Next, orbitofrontal serotonin depletion impacts cognitive flexibility, or the adaptive ability to switch between contingencies, by impairing inhibitory control [37] in monkeys.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [37] HF Clarke, 2004, Cognitive inflexibility after prefrontal serotonin depletion., Science source
Not checked in the free run.
- Premisecitation
Mildly raised BDI scores do reveal a latent level of dysphoric symptoms amongst healthy participants.
“The mildly raised BDI scores do reveal a latent level of dysphoric symptoms amongst healthy participants [55].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [55] A Beck, 1996, Manual for the Beck Depression Inventory-II
Not checked in the free run.
- Premisecitation
A genetic polymorphism that downregulates the serotonin reuptake transporter predisposes towards depression and other related mood disorders.
“On the other hand, a genetic polymorphism that downregulates the very same serotonin reuptake transporter, thus acting in the same direction as the drugs, has the opposite effect on mood, predisposing towards depression and other related mood disorders ([30]; though see also [31] for a discussion of replication failures).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [30] A Caspi, 2003, Influence of life stress on depression: moderation by a polymorphism in the 5-HTT genes., Science source
Not checked in the free run.
- cites [31] M Wankerl, 2010, Current developments and controversies: does the serotonin transporter gene-linked polymorphic region (5-httlpr) modulate the association between stress and depression?, Curr Opin Psychiatry source
Not checked in the free run.
- Premisecitation
Serotonergic neurons project strongly to areas involved in goal-directed, affective choices including the medial prefrontal cortex.
“First, serotonergic neurons project strongly to areas involved in goal-directed, affective choices including the medial prefrontal cortex [35].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [35] K Varnäs, 2004, Autoradiographic distribution of serotonin transporters and receptor subtypes in human brain., Hum Brain Mapp source
Not checked in the free run.
- Premisecitation
People actively suffering from depression are impaired on the OTT.
“People actively suffering from depression are impaired on the OTT [48], [49].”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites [48] BC Beats, 1996, Cognitive performance in tests sensitive to fronal lobe dysfunction in the elderly depressed., Psychol Med source
Not checked in the free run.
- cites [49] R Elliott, 1996, Neuropsychological impairments in unipolar depression: the role of perceived failure on subsequent performance., Psychol Med source
Not checked in the free run.
- Premisecitation
The same group has also reported catastrophic breakdown in OTT performance in depressed subjects after negative feedback.
“Finally, the same group has also reported catastrophic breakdown in OTT performance in depressed subjects after negative feedback [52].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [52] R Elliott, 1997, Abnormal response to negative feedback in unipolar depression: evidence for a diagnosis-specific impairment., J Neurol Neurosurg Psychiatry source
Not checked in the free run.
- Interpretationown data
The process of pruning occurs by simple, Pavlovian, behavioural inhibition of ongoing thoughts in the face of punishments.
“supporting our hypothesis that this process occurs by simple, Pavlovian, behavioural inhibition of ongoing thoughts in the face of punishments [17].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [17] P Dayan, 2008, Serotonin, inhibition, and negative mood., PLoS Comput Biol source
Not checked in the free run.
- Interpretationreasoning
People who are suffering from depression have more difficulty searching a deep tree effectively.
“This suggests that people who are suffering from depression have more difficulty searching a deep tree effectively (possibly also captured by more general, superficial autobiographical recollections; [51]).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [51] JMG Williams, 2007, Autobiographical memory specificity and emotional disorder., Psychol Bull source
Not checked in the free run.
- Rests on an assumption: The one-touch Tower of London (OTT) task engages goal-directed processing.
- May combine two claims. 24
- Classified as probably not a substantial claim. 18
May combine two claims.
Findingown dataSpecific pruning exceeded general pruning across subjects, but there was no main effect of group and no interaction.
“Specific pruning exceeded general pruning across subjects, but there was no main effect of group and no interaction.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingown dataThere was a negative correlation between BDI score and reward sensitivity, although this did not survive correction for multiple comparisons.
“There was also a negative correlation between BDI score and reward sensitivity , although this did not survive correction for multiple comparisons ().”
Quote in the paperNot a substantial claim- Rests on an assumption: The BDI validly measures sub-clinical depressive symptoms/mood disturbance.
May combine two claims.
Findingown dataComputational models accounted for close to 90% of choices and verified that the nature of pruning corresponded to the Pavlovian reflexive account in detail.
“Computational models which accounted for close to 90% of choices verified that the nature of pruning corresponded to the Pavlovian reflexive account in detail.”
Quote in the paperSubstantial claimMay combine two claims.
FindingreasoningSubjects could transition through losses early on in the tree, but were more reluctant to do so when they appeared deeper in the tree.
“Next, subjects could transition through losses early on in the tree, but were more reluctant to do so when they appeared deeper in the tree.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe full look-ahead model ascribed the higher action value to subjects' actual choices 77% of the time, significantly better than chance.
“This model ascribed the higher action value to the subjects' actual choices a total of 77% of the time (fraction of choices predicted), which is significantly better than chance (fixed effect binomial ).”
Quote in the paperSubstantial claimMay combine two claims.
FindingreasoningIn terms of net earnings, pruning is never advantageous, but pruning does not always result in losses; it is most disadvantageous in the −70 group and never disadvantageous in the −140 group.
“In terms of net earnings, pruning is never advantageous (pruned values are never better than the optimal lookahead values); but pruning does not always result in losses (white areas). It is most disadvantageous in the −70 group, and it is never disadvantageous in the −140 group because there is always an equally good alternative choice sequence which avoids transitions through large losses.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataIn the second and third experimental groups, subjects incurred increasingly large costs for pruning yet continued to deploy it.
“In contrast, in the second and third experimental groups subjects incurred increasingly large costs for this pruning strategy (Figure 2D,E); yet, they continued to deploy it.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe pruning model did not predict the attraction to state 1 when only a single choice remained; paired t-tests showed significant differences between empirical and predicted choice probabilities for each of the three groups.
“This was not predicted by the pruning model: paired t-tests showed significant differences between empirical and predicted choice probabilities for each of the three groups: , ; , ; and , , for groups −70, −100 and −140 respectively.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe model with an added Learned Pavlovian component provided the most parsimonious account of the data despite two additional parameters and correctly predicted 91% of choices.
“Figure 4B,C show that this model (Pruning and Learned) provided the most parsimonious account of the data despite two additional parameters, and Figures 4D–E show that the addition of the Learned parameters allowed the model to capture more faithfully the transition probabilities out of state 6. The blue bars in Figure 4A display the probability that this model chose the same action as subjects (c”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataWe did not observe correlations with NEO neuroticism, which is an established risk factor for depression.
“we did not observe correlations with NEO neuroticism, which is an established risk factor for depression [47].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [47] KS Kendler, 1993, The prediction of major depression in women: toward an integrated etiologic model., Am J Psychiatry source
Not checked in the free run.
May combine two claims.
Findingown dataWe used a novel sequential decision-making task in conjunction with a sophisticated computational analysis that fitted a high proportion of healthy subjects' choices.
“We used a novel sequential decision-making task in conjunction with a sophisticated computational analysis that fitted a high proportion of healthy subjects' choices.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationLoss aversion is a specific phenomenon in behavioural economics whereby subjects are more strongly opposed to a given probability of losing a certain amount than to winning that same amount.
“Loss aversion is a specific phenomenon in behavioural economics, whereby subjects are more strongly opposed to a given probability of losing a certain amount than to winning that same amount [26].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [26] A Tversky, 1991, Loss aversion in riskless choice: A reference-dependent model., Q J Econ source
Not checked in the free run.
Classified as probably not a substantial claim.
InterpretationreasoningIf the pruning observed was just a signature of loss aversion, one would have expected the extent of pruning not to be the same across groups.
“Also, if the pruning we observed was just a signature of loss aversion, one would have expected the extent of pruning not to be the same across groups.”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationown dataThe pruning strategy was Pavlovian: it was reflexively evoked by large losses and persisted even when overwhelmingly counterproductive.
“This pruning strategy was Pavlovian: it was reflexively evoked by large losses and persisted even when overwhelmingly counterproductive.”
Quote in the paperSubstantial claim- Rests on an assumption: The large losses in the task are aversive/punishing events capable of evoking Pavlovian behavioural inhibition.
May combine two claims.
Interpretationown dataThis persistence suggests that pruning was reflexively evoked in response to punishments and relatively insensitive to the ultimate outcomes.
“This persistence suggests that pruning was reflexively evoked in response to punishments and relatively insensitive to the ultimate outcomes.”
Quote in the paperSubstantial claim- Rests on an assumption: The large losses in the task are aversive/punishing events capable of evoking Pavlovian behavioural inhibition.
Classified as probably not a substantial claim. May combine two claims.
PremisecitationDrugs that putatively increase serotonin by inhibiting serotonin reuptake are effective for both acutely treating and preventing relapse of depression.
“On the one hand, drugs that putatively increase serotonin by inhibiting the serotonin reuptake mechanism are effective for both acutely treating [28], and preventing relapse of [29], depression.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [28] A Cipriani, 2009, Comparative efficacy and acceptability of 12 new-generation antidepressants: a multiple-treatments meta-analysis., Lancet source
Not checked in the free run.
- cites [29] JR Geddes, 2003, Relapse prevention with antidepressant drug treatment in depressive disorders: a systematic review., Lancet source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
InterpretationcitationLoss aversion may represent a third instance of Pavlovian responses to losses interfering with goal-directed decisions in this task.
“To the extent to which loss aversion can be described as an inflexible, reactive, response to an aversive stimulus, it may represent a third instance of Pavlovian responses to losses interfering with goal-directed decisions in this task [27].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [27] M Guitart-Masip, 2010, Conditioned associations and economic decision biases., Neuroimage source
Not checked in the free run.
Classified as probably not a substantial claim. May combine two claims.
PremisecitationThe model [17] suggested that subjects prone to depression exhibit decision making that is more reliant on serotonergic function, expressed as excess pruning, but that the depressed state itself is characterised by a low serotonin state and thus a loss of pruning.
“The model suggested that subjects prone to depression exhibit decision making that is more reliant on serotonergic function, expressed as excess pruning, but that the depressed state itself is characterised by a low serotonin state and thus a loss of pruning.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [17] P Dayan, 2008, Serotonin, inhibition, and negative mood., PLoS Comput Biol source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationLearned helplessness depends critically on pre- and infralimbic cortex in rats, and is known to be mediated by serotonin.
“Third, learned helplessness, which can be interpreted in goal-directed terms [17], depends critically on pre- and infralimbic cortex in rats [38], and is known to be mediated by serotonin [39].”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites [17] P Dayan, 2008, Serotonin, inhibition, and negative mood., PLoS Comput Biol source
Not checked in the free run.
- cites [38] J Amat, 2005, Medial prefrontal cortex determines how stressor controllability affects behavior and dorsal raphe nucleus., Nat Neurosci source
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- cites [39] SF Maier, 2005, Stressor controllability and learned helplessness: the roles of the dorsal raphe nucleus, serotonin, and corticotropin-releasing factor., Neurosci Biobehav Rev source
Not checked in the free run.
Classified as probably not a substantial claim.
PremisecitationDayan and Huys [17] explained the serotonin-depression paradox by suggesting that people who experienced high levels of serotonin and exaggerated Pavlovian behavioural inhibition during early development would be most sensitive to effects of any interference with this inhibition in adulthood secondary to a drop in serotonin levels.
“Dayan and Huys [17] explained this paradox by suggesting that people who experienced high levels of serotonin and thus exaggerated Pavlovian behavioural inhibition during early development [32] would be most sensitive to the effects of any interference with this inhibition in adulthood secondary to a drop in serotonin levels [33], [34].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [17] P Dayan, 2008, Serotonin, inhibition, and negative mood., PLoS Comput Biol source
Not checked in the free run.
- cites [32] MS Ansorge, 2004, Early-life blockade of the 5-HT transporter alters emotional behavior in adult mice., Science source
Not checked in the free run.
- cites [33] JP Roiser, 2006, Serotonin transporter polymorphism mediates vulnerability to loss of incentive motivation following acute tryptophan depletion., Neuropsychopharmacology source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
InterpretationreasoningThe OTT requires sequences of moves to be evaluated, but there is no obvious aversive point at which Pavlovian pruning might be invoked.
“One possible explanation for this apparent discrepancy is that although the OTT requires sequences of moves to be evaluated, there is no obvious aversive point at which Pavlovian pruning might be invoked.”
Quote in the paperNot a substantial claim- Rests on an assumption: The one-touch Tower of London (OTT) task engages goal-directed processing.
Classified as probably not a substantial claim. May combine two claims.
InterpretationcitationThe inhibitory consequences of serotonin could account for both its predisposing qualities on a developmental time-scale and more acute relief during depressive episodes.
“Thus, the inhibitory consequences of serotonin could account for both its predisposing qualities on a developmental time-scale, and more acute relief during depressive episodes.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationThe impairment relative to controls grows with the difficulty of the problem; and depressed subjects also spend increasing amounts of time thinking about the harder problems, without showing improved choices.
“The impairment relative to controls grows with the difficulty of the problem; and depressed subjects also spend increasing amounts of time thinking about the harder problems, without showing improved choices [50].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [50] GM Goodwin, 1997, Neuropsychological and neuroimaging evidence for the involvement of the frontal lobes in depression., J Psychopharmacol source
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe stronger dependence on serotonin in at-risk subjects would explain why only they are sensitive to the mood effects of tryptophan depletion, and why individuals with a polymorphism in the serotonin transporter gene that reduces serotonin uptake are more liable to develop mood disturbance, especially following serotonin depletion.
“The stronger dependence on serotonin in at-risk subjects would explain why only they are sensitive to the mood effects of tryptophan depletion [34], and why individuals with a polymorphism in the serotonin transporter gene that reduces serotonin uptake are more liable to develop mood disturbance, especially following serotonin depletion [41], [42].”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [34] HG Ruhé, 2007, Mood is indirectly related to serotonin, norepinephrine and dopamine levels in humans: a meta-analysis of monoamine depletion studies., Mol Psychiatry source
Not checked in the free run.
- cites [41] JP Roiser, 2006, Serotonin transporter polymorphism mediates vulnerability to loss ofincentive motivation following acute tryptophan depletion., Neuropsychopharmacology source
Not checked in the free run.
- cites [42] A Neumeister, 2002, Association between serotonin transporter gene promoter polymorphism (5HTTLPR) and behavioral responses to tryptophan depletion in healthy women with and without family history of depression., Arch Gen Psychiatry source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationMood manipulation, but not acute tryptophan depletion, impairs processing on the one-touch Tower of London (OTT) task.
“Contrary to this, there is a recent report that mood manipulation, but not acute tryptophan depletion, impairs processing on the one-touch Tower of London (OTT) task [40], which should certainly engage goal-directed processing.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [40] OJ Robinson, 2009, A double dissociation in the roles of serotonin and mood in healthy subjects., Biol Psychiatry source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationGenetic variation in the serotonin transporter allele modulates functional coupling between amygdala and rostral cingulate cortex.
“Genetic variation in the serotonin transporter allele modulates functional coupling between amygdala and rostral cingulate cortex [36].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [36] L Pezawas, 2005, 5-HTTLPR polymorphism impacts human cingulate-amygdala interactions: a genetic susceptibility mechanism for depression., Nat Neuosci source
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe theory predicts excessive pruning to occur in subjects at risk for depression, and reduced pruning to occur during a depressive episode.
“That is, this theory predicts excessive pruning to occur in subjects at risk for depression, and reduced pruning to occur during a depressive episode.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductown dataOne subject's age information, and one subject's STAI information were lost; these subjects were excluded from the psychometric correlation analyses.
“One subjects' age information, and one subject's STAI information were lost. These subjects were excluded from the psychometric correlation analyses.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable).
Interpretationown dataThis allowed us to unpack a central facet of effective computation, pruning.
“This allowed us to unpack a central facet of effective computation, pruning.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductnone givenReaching training criterion was mandatory to move on to the main task.
“Reaching training criterion was mandatory to move on to the main task.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductnone givenThe study was conducted in accord with the Helsinki declaration and approved by the UCL Graduate School Ethics Committee.
“The study was conducted in accord with the Helsinki declaration and approved by the UCL Graduate School Ethics Committee.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Study design and conductcitationThe absence of axis-I psychopathology and alcohol- or substance abuse/dependence was confirmed with the Mini International Neuropsychiatric Inventory.
“The absence of axis-I psychopathology and alcohol- or substance abuse/dependence was confirmed with the Mini International Neuropsychiatric Inventory [53].”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [53] DV Sheehan, 1998, The mini-international neuropsychiatric interview (m.i.n.i.): the development and validation of a structured diagnostic psychiatric interview for dsm-iv and icd-10., J Clin Psychiatry
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown dataSubjects assigned to the different groups were matched for age, IQ and sex.
“Subjects who were assigned to the different groups, were matched for age, IQ and sex (all , one-way ANOVA).”
Quote in the paperNot a substantial claim
The paper relies on this without support.
The large losses in the task are aversive/punishing events capable of evoking Pavlovian behavioural inhibition.
“We term this heuristic pruning a “Pavlovian” response because it is invoked, as an immediate consequence of encountering the large loss, when searching the tree in one's mind.”
The paper relies on this without support.
The BDI validly measures sub-clinical depressive symptoms/mood disturbance.
“We found that BDI was positively correlated with the specific pruning parameter ().”
The authors state this as an assumption.
Actions are independent given the stimuli, and thus factorize over trials.
“We assume that actions are independent (given the stimuli, which we omit for notational clarity), and thus factorize over trials.”
The authors state this as an assumption.
The value of each choice depends on how many choices are left after it, but not on the choices preceding it.
“we emphasize that the value of each choice depends on how many choices are left after , but not on the choices preceding it.”
The paper relies on this without support.
The one-touch Tower of London (OTT) task engages goal-directed processing.
“Contrary to this, there is a recent report that mood manipulation, but not acute tryptophan depletion, impairs processing on the one-touch Tower of London (OTT) task [40], which should certainly engage goal-directed processing.”
The paper relies on this without support.
The pruning parameter mediates the inhibition of iterative thought processes.
“if serotonin indeed mediates behavioural inhibition in the face of punishments [10], [12]–[14] then it is a strong prediction that the pruning parameter , which mediates the inhibition of iterative thought processes, should be related to, and modulated by, serotonergic activity.”
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?
Other related search tasks have found behavioral evidence for these calculations and have started to look at heuristics for pruning the tree, a necessity when it gets too wide or deep.
“Other related search tasks have found behavioral evidence for these calculations and have started to look at heuristics for pruning the tree, a necessity when it gets too wide or deep (Huys et al., 2012).”
From Goals and Habits in the Brain (2013) · cites this paper as Huys et al., 2012
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- Authors
- Huys, Q. J. M., Eshel, N., O'Nions, E., Sheridan, L., Dayan, P., & Roiser, J. P.
- Affiliations
- Wellcome Centre for Human Neuroimaging (GB)
- University College London (GB)
- Gatsby Computational Neuroscience Unit (GB)
- Corresponding author
- Huys, Quentin J. M.
- Venue
- PLoS Computational Biology
- Publisher
- Public Library of Science (PLoS)
- Published
- 2012-03-08
- Volume
- 8
- Issue
- 3
- Pages
- e1002410
- ISSN
- 1553-734X (ISSN-L)
- 1553-7358
- PMC
- PMC3297555
- OpenAlex
- W2059569317
- Semantic Scholar
- 09c66a4231400ba276c92700fe993d6c185bc4ac
- Cited by
- 439
- References
- 70
- Citations by year
- 2026: 16 · 2025: 35 · 2024: 32 · 2023: 42 · 2022: 38 · 2021: 30 · 2020: 33 · 2019: 32 · 2018: 44 · 2017: 34 · 2016: 27 · 2015: 27 · 2014: 29 · 2013: 11 · 2012: 5
- FWCI
- 17.75
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- PLoS Computational Biology (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Staff Publications Online (The Tavistock and Portman) (acceptedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- UCL Discovery (University College London) (submittedVersion)
- MPG.PuRe (Max Planck Society) (submittedVersion)
- Funders
- Gatsby Charitable Foundation
- Topics
- Social Sciences › Decision Sciences › General Decision Sciences › Decision-Making and Behavioral Economics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural and Behavioral Psychology Studies
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Keywords
- decision tree pruning, goal-directed behavior, sequential decision making, loss aversion, serotonin, mood disorders, reinforcement learning, human decision-making
- MeSH terms
- Algorithms, Computer Simulation, Decision Making, Goals, Humans, Models, Neurological, Reinforcement, Psychology, Task Performance and Analysis
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Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-09.
Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-09.
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