The DOI matches its Crossref record; 73 of the 73 references it cites were found in a registry or an index; none of the 65 sources its claims cite is checked for support yet.
Free Energy, Precision and Learning: The Role of Cholinergic Neuromodulation
Open accessAbstract
Acetylcholine (ACh) is a neuromodulatory transmitter implicated in perception and learning under uncertainty. This study combined computational simulations and pharmaco-electroencephalography in humans, to test a formulation of perceptual inference based upon the free energy principle. This formulation suggests that ACh enhances the precision of bottom-up synaptic transmission in cortical hierarchies by optimizing the gain of supragranular pyramidal cells. Simulations of a mismatch negativity paradigm predicted a rapid trial-by-trial suppression of evoked sensory prediction error (PE) responses that is attenuated by cholinergic neuromodulation. We confirmed this prediction empirically with a placebo-controlled study of cholinesterase inhibition. Furthermore, using dynamic causal modeling, we found that drug-induced differences in PE responses could be explained by gain modulation in supragranular pyramidal cells in primary sensory cortex. This suggests that ACh adaptively enhances sensory precision by boosting bottom-up signaling when stimuli are predictable, enabling the brain to respond optimally under different levels of environmental uncertainty.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
40 Not checked yet · 31 Own interpretation · 24 Own result · 18 No basis given · 3 Own method
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we found that drug-induced differences in PE responses could be explained by gain modulation in supragranular pyramidal cells in primary sensory cortex.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Abstract
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We confirmed this prediction empirically with a placebo-controlled study of cholinesterase inhibition.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Our null hypothesis was that Galantamine precluded precision updates. This produced a very different profile, driven by slower changes in extrinsic connections that underlie learning. Here the MMN on the second trial d2 (7.2 a.u.) was greater than on d1 (6.7 a.u.).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“A mismatch negativity was seen in the grand-averaged data – most prominently in trial d1 at 150 ms for both placebo and Galantamine conditions.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Data pre-processing and statistical analysis
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Figure 5b shows the results of a Bayesian model comparison – showing that the evidence for model 2 is greatest, with very strong evidence relative to the next best model – model 6 (log Bayes Factor = 153).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
boosted- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The direction of this effect is shown in Figure 5c and suggests that Galantamine enhances the gain in these bilateral sensory sources with a posterior probability of nearly 100%.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“while the placebo MMN reverses in polarity after the first oddball (Figure 3d).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plainn = 13- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we then tested for an interaction of repetition and drug – identifying a significant interaction in middle temporal gyri bilaterally (left MTG: x = −60, y = −14, z = −18, right MTG: x = 62, y = −14, z = −18; Figure 3b).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“After source localisation on a canonical mesh, we tested for a simple main effect of repetition (d10 – d1) in the placebo condition and found bilateral effects in middle temporal gyri (left MTG: x = −66, y = −30, z = −4; right MTG: x = 64, y = −16, z = −18) and inferior frontal gyri (left IFG: x = −46, y = 40, z = 2; right IFG: x = 36, y = 48, z = −2), (Figure 4b).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we observed an overall profile for the drug condition that is very similar to the simulated MMN when Galantamine enhanced precision.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plaincorrelationaln = 13- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In brief, we found a remarkable correspondence between simulations of Bayes optimal sensory learning and the empirical responses – at the level of scalp EEG and within the network of sources generating EEG signals.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results
boostedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Although we did not observe a significant drug × repetition interaction”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plainn = 13- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“we did find a significantly greater MMN under drug compared to placebo on the first and second presentation of a new tone (d1 and d2 relative to d10; p < 0.05, two-sample, one-tailed t test; Figure 3d).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plainn = 13 ·- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We observed a significant drug effect within the MMN mask in the middle temporal gyri bilaterally (Figure 4b).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Source reconstruction
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“One of the key features of the responses elicited by the roving paradigm is a repetition suppression – following a deviant stimulus – that recovers with subsequent presentations (see Figure 3).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“We simulated these electrophysiological responses by weighting prediction errors – during recognition dynamics – more highly under the influence of Galantamine. This results in exaggerated and prolonged MMN like responses over repetitions of a new stimulus.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The simulations and empirical results suggest that ACh biases perception towards bottom-up sensory processing – by boosting prediction errors at the lower (sensory) levels of the hierarchy (see Figure 1).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“The Helmholtzian notion of the brain as a statistical inference machine (Hemholtz 1866, von Helmholtz 1962) can be realised – under free energy formulations of perceptual inference – as a neurobiologically plausible implementation of predictive coding.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Hemholtz 1866
- Not checked yetcites von Helmholtz 1962
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“Under the free energy formulation, see Figure 1, ACh and other neuromodulators encode changes in the precision of (certainty about) prediction errors in sensory cortical hierarchies (Friston 2008).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Friston 2008
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“In this framework, the brain entails a generative model of its environment that reproduces the hierarchical and dynamic generation of sensory input.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Mumford 1992
- Not checked yetcites Rao and Ballard 1999
- Not checked yetcites Friston 2010
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“Put simply, to infer the causes of sensory input, the brain has to recognise when sensory information is noisy or uncertain and down-weight it suitably in relation to top-down predictions (Yu and Dayan 2005, Deco and Thiele 2009).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Yu and Dayan 2005
- Not checked yetcites Deco and Thiele 2009
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“The implicit Bayes-optimal recognition requires statistical distributions over hidden states and the environment to be encoded by neuronal activity in cortical hierarchies; where these representations are updated until they mirror real-world statistics through accurate predictions of sensory input.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: unclear — It is unclear why the paper says it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The empirical responses – and associated changes in postsynaptic gain – provide strong support for a specific role of ACh during perceptual inference under uncertainty.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“This formulation suggests that acetylcholine enhances the precision of bottom-up synaptic transmission in cortical hierarchies by optimising the gain of supragranular pyramidal cells.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Abstract
What the sources say
- Not checked yetcites Friston 2008
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“These data suggest a selective cholinergic modulation of sensory cortex (Nelson, Sarter et al. 2005, Sarter, Hasselmo et al. 2005, Benarroch 2010, Hasselmo and Sarter 2010, Fadel 2011).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Nelson, Sarter et al. 2005
- Not checked yetcites Sarter, Hasselmo et al. 2005
- Not checked yetcites Benarroch 2010
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“The ensuing Bayes optimal perception rests on optimising precision at each level of the hierarchy that is commensurate with the environmental statistics they represent.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“Predictive coding schemes (Rao and Ballard 1999) invoke recurrent neural message passing to simultaneously predict sensory stimuli and report prediction errors”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
What the sources say
- Not checked yetcites Rao and Ballard 1999
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“This three-level hierarchy is consistent with previous models of the MMN (Garrido, Kilner et al. 2007, Garrido, Kilner et al. 2007, Garrido, Friston et al. 2008, Garrido, Kilner et al. 2009, Garrido, Kilner et al. 2009).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
What the sources say
- Not checked yetcites Garrido, Kilner et al. 2007
- Not checked yetcites Garrido, Friston et al. 2008
- Not checked yetcites Garrido, Kilner et al. 2009
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our simulations and empirical results suggest that when stimuli are repeated in a predictable way, the statistical regularity leads to an adaptive increase in the expected precision of sensory prediction errors.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“Galantamine increases the availability of ACh in cholinergic synapses by competitive inhibition of acetylcholinesterase – the enzyme responsible for its breakdown.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Subjects and pharmacological manipulation
What the sources say
- Not checked yetcites Coyle and Kershaw 2001
- Not checked yetcites Samochocki, Höffle et al. 2003
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This withdrawal of a neuromodulatory boost leads to a rapid repetition suppression during successive presentations of the same (initially surprising and then increasingly predictable) stimulus.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These results are consistent with a key role for acetylcholine in modulating the gain of superficial pyramidal cells in early sensory regions that may encode prediction error in predictive coding formulations of hierarchical inference.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“After the first deviant, sensory precision is reduced (by the high prediction errors in the previous trial), rendering subsequent prediction errors less precise and thereby attenuating neuronal responses.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Mathematically, precision increases the influence of prediction errors, when they are precise.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“By assigning ACh the role of signalling sensory precision, we have shown how its augmentation can reduce adaptation to surprising stimuli in sensory cortices.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These empirical response profiles are inconsistent with either of the alternative models of Galantamine effects (Figure 3b).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plainn = 13 ·- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“By changing the precision (synaptic gain) at a particular level of cortical hierarchy one can simulate the effects of manipulating cholinergic neuromodulation at this level, on the ensuing inference (Figure 1).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“Crucially, the process of free energy minimisation – in networks implementing predictive coding – can be used to predict the neuronal dynamics one would expect to measure using EEG – see Figure 1 (Friston and Kiebel 2009).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
What the sources say
- Not checked yetcites Friston and Kiebel 2009
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This could be mediated (in part) by increased cholinergic neurotransmission that is suppressed after an unpredicted or surprising (deviant) stimulus.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“These models have been validated using animal electrophysiological recordings (Moran, Stephan et al. 2008, Moran, Jung et al. 2011).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites Moran, Stephan et al. 2008
- Not checked yetcites Moran, Jung et al. 2011
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This difference in the optimisation of (cholinergic) precision or gain is reflected in the simulated evoked responses (Figure 3a), with enhanced and prolonged precision-weighted prediction errors over repetitions, compared to placebo.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Optimal changes in synaptic efficacy and gain
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Crucially, these Bayesian updates involve not only the parameters responsible for generating sensory predictions but also the precision encoding the uncertainty about those predictions.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“From Figure 3a, we see that the corresponding evoked responses reflect these estimates of precision, because they are based on precision-weighted prediction errors.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Optimal changes in synaptic efficacy and gain
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In brief, our results are consistent with a role for ACh in setting the gain of prediction error units implied by predictive coding in the brain.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“Crucially the parameters of the model can be inferred from empirical data, using standard Bayesian inversion techniques (Kiebel at al 2006).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites Kiebel at al 2006
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“Using standard univariate analyses, the neural network responsible for these – possibly reciprocal – adjustments under ACh has remained unclear (Thiel and Fink 2008), given the difficulty of specifying how a “bottom-up” enhancement and “top-down” reduction would manifest in measurements of brain responses such as fMRI.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Thiel and Fink 2008
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our DCM analysis helps to resolve previous ambiguities regarding the effects of ACh on top-down vs. bottom-up processing in human neuroimaging experiments.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Using precision-weighted prediction errors from our Bayes optimal (predictive coding) scheme, we could predict suppression of the MMN response under the different drug conditions (Figure 3).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“In general, hierarchical inference using empirical Bayes, as implemented in our simulations, provides a mechanism by which neural circuits can selectively up-weight and down-weight particular features in the sensorium.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“Previous research has shown that MMN responses are enhanced following cholinergic stimulation, particularly nicotinic stimulation in healthy controls (Baldeweg, Wong et al. 2006) and patient populations (Engeland, Mahoney et al. 2002).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Baldeweg, Wong et al. 2006
- Not checked yetcites Engeland, Mahoney et al. 2002
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Crucially, this effect can be simulated easily, when sensory precision remains higher for longer periods.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We extend these findings to show that this short-term plasticity may be modulated by ascending neuromodulatory systems.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“When making perceptual inferences about the states of the world and their causes, the brain should up-weight reliable sensory evidence by increasing the precision of signals from lower-level cortical regions; e.g., primary sensory areas.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“Our findings fit comfortably with previous theoretical and empirical accounts; namely, that ACh boosts bottom-up signals in response to uncertainty (Yu and Dayan 2002, Yu and Dayan 2003, Bentley, Husain et al. 2004).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Yu and Dayan 2002
- Not checked yetcites Yu and Dayan 2003
- Not checked yetcites Bentley, Husain et al. 2004
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our DCM results suggest that under cholinesterase inhibition, sensory cortices respond to incoming stimuli with exaggerated and prolonged trial-by-trial responses at superficial pyramidal cells – that are the source of bottom-up or forward projections.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This enhanced bottom-up effect is mediated via tonic increases in ACh in our simulations and experiments.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- It may state two separate claims as one. 37
- Probably not a substantial claim. 29
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Simulations of a mismatch negativity paradigm predicted a rapid trial-by-trial suppression of evoked sensory prediction error (PE) responses that is attenuated by cholinergic neuromodulation.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“For the placebo MMN, when comparing trial d10 and trial d1, we observe a MMN of 8.1 a.u. that reverses polarity and reduces in size to −1 a.u. on trial d2. The MMN effect then gradually increases again to baseline (−0.1 a.u.) for d9.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our precision estimates show biphasic changes, where the initial oddball coincides with a high precision estimate (d1 = 5.1 a.u.) that undergoes a large reduction (d2 = 2.1 a.u.) and a subsequent rebound (d10 = 4.7 a.u.); Figure 4a (left).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Optimal changes in synaptic efficacy and gain
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These responses exhibited the same form of repetition suppression that we saw in the simulations – with trial d2 (green curve; Figure 1c) more similar to d10 (red curves) and a deflection at around 150 ms for the first oddball tone (blue curves).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“As shown in Figure 3c, we observed an MMN in response to the onset of a deviant tone at a time commensurate with previous studies – at ~150 msec.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plainn = 13- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Under Galantamine (Figure 3a right), the high a priori precision on the sensory prediction error increases the response elicited by the first presentation of a new tone (blue curve), and attenuates the reduction of precision on subsequent presentations – leading to larger responses at the second deviant (green curve) and a more protracted repetition suppression of subsequent responses.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- The authors state this as an assumption: Galantamine increases both the brain's prior expectation about the precision of sensory input and the confidence about that expectation.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Comparing trial d10 and trial d1, we observe a difference of magnitude of 34 a.u. which remains positive at 2.5 a.u. on the subsequent trial, d2. The MMN effect decreases slowly to a baseline for d9.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- The authors state this as an assumption: Galantamine increases both the brain's prior expectation about the precision of sensory input and the confidence about that expectation.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Figure 3a (left) shows the (precision weighted) prediction error in the placebo condition for the first deviant. A large precision-weighted prediction error is observed with a late deflection at around 150 ms (blue curve). This almost disappears on the second trial (green curve), which has a form similar to d10 (red curve).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our empirical results show an enhanced MMN on Galantamine at d1 compared to placebo that persists at d2”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Empirical Evoked Brain Responses and the MMN under Placebo and Galantamine
plaindirectn = 13 ·- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In the Galantamine simulations (Figure 4a – right), however, there was a profound difference in the log-precision updates, compared to placebo. The log-precision at the first deviant was higher (8.1 a.u.) and remained relatively high after the second tone repetition (d2 = 6.1 a.u.), reaching a minimum at d3 (Figure 4a).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Optimal changes in synaptic efficacy and gain
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Our first alternate hypothesis was that Galantamine reduced – rather than increased – sensory precision. This assumption resulted in a reduced MMN response of 2.9 a.u. (d10−d1) that reduces polarity on d2 to −0.5.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plaindirect- Asserted: fails — The authors may not put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The authors may not put it forward as true. The paper’s conclusions may not depend on it. It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“To weight hierarchical predictions with greater or lesser importance, optimal inference assigns greater or lesser precision (inverse variance) to different hierarchical levels.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds.
In the paper
“These signals are prediction errors that report discrepancies between top-down predictions and representations at each level (Kiebel, Daunizeau et al. 2009).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Kiebel, Daunizeau et al. 2009
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Neuronal responses are proposed to reflect inference on the (hidden) causes of sensory stimuli that enable the brain to predict its sensations (Mumford 1992; Rao and Ballard 1999: Friston 2010).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
What the sources say
- Not checked yetcites Mumford 1992
- Not checked yetcites Rao and Ballard 1999
- Not checked yetcites Friston 2010
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Each level of a processing hierarchy sends predictions to the level below, which reciprocate bottom-up signals.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“This recurrent message passing continues until prediction errors are minimised throughout the hierarchy.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Introduction
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
In the paper
“The free energy framework places predictive coding in a more general setting, using dynamic and hierarchical generative models of hidden environmental states (causes of sensory inputs) and an approximate posterior density on the states and parameters of the generative model, encoded by synaptic activity and weights respectively.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
What the sources say
- Not checked yetcites Kiebel, Daunizeau et al. 2009
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This suggests that acetylcholine adaptively enhances sensory precision by boosting bottom-up signalling when stimuli are predictable, enabling the brain to respond optimally under different levels of environmental uncertainty.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Abstract
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“After model inversion or optimisation, the variational free energy can then used as an approximate log model evidence for Bayesian model comparison.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In summary, there is a remarkable convergence between the predictions of Bayes-optimal inference and learning under uncertainty and the empirical results obtained with a roving mismatch negativity paradigm. This convergence was both phenomenologically, in terms of repetition suppression at the sensor level, and mechanistically, in terms of underlying changes in synaptic gain at the source level (a”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
boostedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These simulation results suggest a type of one-shot learning or repetition suppression – where the brain responds to a new tone with a precise prediction error and then reduces the precision of subsequent error-related responses – as precision is updated and new parameters of the generative model are learned.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“This ensemble model is based on a mesoscopic circuit description of cat primary visual cortex, which represents the cardinal features of functional macrocolumns throughout the cortical sheet (Douglas and Martin 2004, Bastos, Usrey et al. 2012).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites Douglas and Martin 2004
- Not checked yetcites Bastos, Usrey et al. 2012
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
In the paper
“This learning also corresponds to a gradient descent on free energy and – under the simple linear model above – reduces to associative plasticity (see Friston 2008 for details).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
What the sources say
- Not checked yetcites Friston 2008
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“This CMC model (Figure 4c) was used to model explicitly the laminar specificity of forward and backward connections that are central to predictive coding – where forward connections convey prediction errors to higher levels and backward connections send predictions back to lower areas.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The DCM generates a predicted ERP as the response of a network of coupled sources to sensory (thalamic) input – where each source corresponds to a neural mass model of several neuronal subpopulations.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“where greater ACh levels enhance postsynaptic responses in the superficial layers of primary auditory cortex.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds.
In the paper
“Importantly, acetylcholine modulates this process; e.g., by activation of muscarinic receptors suppresses sIAHP through a cGMP-dependent mechanism (Krause and Pedarzani 2000).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Krause and Pedarzani 2000
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“In brief, this scheme encodes beliefs about the world by associating synaptic activity with expected states of the world, causal structure (parameters controlling transitions among states) with synaptic connection strengths and the precision (inverse variance) of random fluctuations of hidden states with synaptic gain: see (Friston 2008) for details.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
What the sources say
- Not checked yetcites Friston 2008
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
In the paper
“Extrinsic connections in these models follow the same rules on which predictive coding schemes are based: forward connections from superficial pyramidal cells synapse on granular layer IV and deep cell layers, while backward connections from deep pyramidal cells terminate outside layer IV (Felleman and Van Essen 1991).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites Felleman and Van Essen 1991
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Subjects were not asked to attend to the auditory stimuli, since this type of auditory oddball paradigm elicits mismatch potentials that are automatic or pre-attentive (Garrido, Kilner et al. 2009, Näätänen, Astikainen et al. 2010).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Task and EEG acquisition
What the sources say
- Not checked yetcites Garrido, Kilner et al. 2009
- Not checked yetcites Näätänen, Astikainen et al. 2010
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This means that the effect of Galantamine can be best explained as a change in the gain of superficial pyramidal cells in primary auditory cortex.”
- Also stated in: § Discussion · hedged
“Moreover, dynamic causal modelling of a temporo-frontal network suggested that the observed differences in event-related responses can be attributed to cholinergic gain modulation of supragranular pyramidal cells in early auditory cortex (Figure 5).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > DCM results and their relationship to theoretical predictions
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- The authors state this as an assumption: Model parameters such as C are encoded by the strength of extrinsic connections between brain regions, while log-precisions are encoded by the gain of prediction error units, i.e. the intrinsic excitation of supragranular pyramidal cells.
- The authors state this as an assumption: In predictive coding formulations of free energy minimisation, gain corresponds to the precision of prediction errors.
- Also stated in: § Discussion · hedged
In the paper
“Interestingly, DCMs are inverted using exactly the same variational scheme (variational Laplace) that underlies predictive coding (Friston et al 2007).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites Friston et al 2007
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Mathematically, this optimisation involves minimising a variational free energy bound on log evidence (henceforth free energy) which, when minimised, provides a recognition density that approximates the true posterior – this is also known as approximate Bayesian inference.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Each source includes four neuronal subpopulations: infra- and supragranular pyramidal cell populations, a granular layer of stellate cells, and an inhibitory interneuron population.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites Douglas and Martin 2004
- Not checked yetcites Bastos, Usrey et al. 2012
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“Note that we report uncorrected results at p < 0.05 because these analyses only served to identify candidate sources in the subsequent DCM analysis.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Source reconstruction
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Following an oral dose, the peak plasma concentration is attained within 2 hours, and declines with a half-life of about 7 hours (Huang, Lasseter et al. 2002).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Subjects and pharmacological manipulation
What the sources say
- Not checked yetcites Huang, Lasseter et al. 2002
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“This activation may maintain a high postsynaptic gain and facilitate oscillatory activity, with a shift towards higher frequencies (desynchronisation) in the power spectra of neuronal recordings (Liljenstrom and Hasselmo 1995).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Liljenstrom and Hasselmo 1995
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“This Bayesian filtering or predictive coding can be formulated in terms of Bayesian updates that are driven by precision-weighted prediction errors.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“The precision weighting of these prediction errors units depends on their postsynaptic gain (c.f., Kalman gain).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In other words, the first deviant stimulus produces large amplitude prediction errors that are boosted by an inappropriately high precision – established by the preceding sequence of predictable standards. This produces the mismatch negativity.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Simulating Perception under Placebo and Galantamine
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Several intracellular processes may underlie such repetition suppression, including slow after-hyperpolarizing currents (sIAHP) that are mediated by calcium dependent potassium channels and decrease neuronal excitability and firing rate: for a review, see (Faber and Sah 2003).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Faber and Sah 2003
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“In neurobiological terms, it is generally assumed that prediction errors are encoded by pyramidal cells in the superficial layers of cortex that receive top-down predictions from deeper laminae in higher regions (Mumford 1992, Cauller 1995, Bar 2003).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
What the sources say
- Not checked yetcites Mumford 1992
- Not checked yetcites Cauller 1995
- Not checked yetcites Bar 2003
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“We studied 13 right-handed, healthy volunteers (5 female, aged 25 ± 7 yrs), using a within-subject crossover placebo-controlled double-blind design.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Subjects and pharmacological manipulation
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“In DCM, event related potentials are modelled as the response of a dynamic input-output system to exogenous (experimental) inputs (David, Kiebel et al. 2006, Kiebel, David et al. 2006).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Dynamic Causal Modelling
What the sources say
- Not checked yetcites David, Kiebel et al. 2006
- Not checked yetcites Kiebel, David et al. 2006
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Galantamine also enhances cholinergic neurotransmission by sensitizing nicotinic receptors (it both increases the probability of channel opening induced by acetylcholine and slows down receptor desensitization) (Coyle and Kershaw 2001, Samochocki, Höffle et al. 2003).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Subjects and pharmacological manipulation
What the sources say
- Not checked yetcites Coyle and Kershaw 2001
- Not checked yetcites Samochocki, Höffle et al. 2003
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In short, there is a wealth of theoretical and empirical evidence for a computational role of ACh in promoting the influence of sensory evidence in perception and attention – which our study supplements with a functional anatomy at the systems level.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In summary, pharmacologically augmenting cholinergic neuromodulation appears to boost event related responses to deviant stimuli and attenuate their subsequent repetition suppression.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“ACh has long been known to enhance the firing of cortical pyramidal neurons (Krnjevi and Phillis 1963), inducing a decrease in spike frequency adaptation via several mechanisms – including the reduction of hyperpolarizing potassium currents (Benardo and Prince 1982, Cole and Nicoll 1984, Huang, Morielli et al. 1993).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Krnjevi and Phillis 1963
- Not checked yetcites Benardo and Prince 1982
- Not checked yetcites Cole and Nicoll 1984
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“In summary, the proposition that neuromodulators encode the precision or predictability of environmental states is supported here by empirical evidence that sensory regions respond with enhanced gain under cholinergic enhancement – producing exaggerated and prolonged mismatch responses that localise to superficial pyramidal populations encoding prediction error (Mumford 1992).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Mumford 1992
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“Of course, our EEG measurements do not allow us to characterise the source of cholinergic modulation per se – e.g. between prefrontal regions and basal forebrain cholinergic neurons, which may serve as a route through which enhanced precision is initiated.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Previous theoretical circuit level proposals have suggested that ACh supports learning and memory processes by switching the cortex from a ‘read-out’ to ‘read-in’ mode – providing an enhancement of sensory-evoked afferent responses and suppression of internal cortical processing (Hasselmo and McGaughy 2004).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Hasselmo and McGaughy 2004
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The resulting adaptive sensory bias may be implemented via gain control in a manner suggested by previous physiological accounts of cholinergic modulation.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: fails — It may be too vague to check against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“We show here an equivalent and complementary effect, where ACh boosts bottom-up sensory evidence in the auditory hierarchy.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Indeed, it has been demonstrated in the auditory domain that a sensory “memory” relies upon cholinergic modulation (Miasnikov, Chen et al. 2008).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Miasnikov, Chen et al. 2008
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our analysis of the simulated and empirical responses recapitulates some of these established ideas about the role of cholinergic signalling in perception and attention.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In short, these findings suggest that acetylcholine mediates the representation of precision and acts to facilitate the bottom-up signalling of sensory information (through precise prediction errors).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Similarly, in human studies with cholinergic agonists, the specificity of perceptual learning for behaviourally-relevant features is enhanced (Rokem and Silver 2010), while validity effects in spatial cueing tasks are diminished (Bentley, Husain et al. 2004, Vossel, Thiel et al. 2007, Thiel and Fink 2008), indicative of selective boosting of stimulus-related information.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Rokem and Silver 2010
- Not checked yetcites Bentley, Husain et al. 2004
- Not checked yetcites Vossel, Thiel et al. 2007
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“The framework of Yu and Dayan (Yu and Dayan 2002) proposes that ACh modulates uncertainty about high-level representations.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Yu and Dayan 2002
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This proposal is consistent with the current study.”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
plain- Asserted: passes — The authors put it forward as true.
- Checkable: fails — It may be too vague to check against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it.
In the paper
“This profile has been reported previously in intrinsic (within-source) DCM connections exhibiting biphasic changes that were much faster than changes in extrinsic (between-source) connections (Garrido, Kilner et al. 2009).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Discussion
What the sources say
- Not checked yetcites Garrido, Kilner et al. 2009
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
The authors state this as an assumption.
The hierarchical organisation of cortex implies a hierarchical model of the sensorium that can be described by stochastic differential equations.
“we assume that the hierarchical nature of cortical organisation (Felleman and Van Essen 1991) implies a hierarchical model of the sensorium that can be described by a set of stochastic differential equations of the following general form:”
The authors state this as an assumption.
The brain uses the same hierarchical model that generated the stimuli, but does not know the frequency modulation parameter C.
“we assumed that the brain was equipped with the same hierarchical model used to generate the stimuli, but did not know the parameter controlling frequency modulation over peristimulus time (the C parameter above).”
The authors state this as an assumption.
Prediction errors are reported by superficial pyramidal cells, which are the principal contributors to electrophysiological measurements.
“Assuming that prediction errors are reported by superficial pyramidal cells – that are the principal contributors to electrophysiological measurements – we can treat the precision weighted prediction errors as event related potentials (ERPs).”
The authors state this as an assumption.
Galantamine increases both the brain's prior expectation about the precision of sensory input and the confidence about that expectation.
“we assumed that Galantamine increases both the brain’s prior expectation about the precision of sensory input and the confidence about that expectation (Figure 1; upper panel).”
The authors state this as an assumption.
Model parameters such as C are encoded by the strength of extrinsic connections between brain regions, while log-precisions are encoded by the gain of prediction error units, i.e. the intrinsic excitation of supragranular pyramidal cells.
“Under our biological implementation of Bayes-optional perception, model parameters (like C in Equation 2) are encoded by the strength of extrinsic connections between brain regions; while log-precisions are encoded by the gain of prediction error units (i.e. intrinsic excitation of supragranular pyramidal cells).”
The authors state this as an assumption.
In predictive coding formulations of free energy minimisation, gain corresponds to the precision of prediction errors.
“In predictive coding formulations of free energy minimisation, this gain corresponds to the precision of prediction errors.”
The authors state this as an assumption.
The first presentation of the deviant (d1) is not confounded by sensory learning under the different drug conditions.
“To test for the predicted differences in placebo and Galantamine, we focused on the first presentation of the deviant (d1), because this is not confounded by sensory learning – under the different drug conditions – in subsequent repetitions.”
The paper relies on this without support.
When making perceptual inferences about the states of the world and their causes, the brain should up-weight reliable sensory evidence by increasing the precision of signals from lower-level cortical regions such as primary sensory areas.
“When making perceptual inferences about the states of the world and their causes, the brain should up-weight reliable sensory evidence by increasing the precision of signals from lower-level cortical regions; e.g., primary sensory areas.”
The paper relies on this without support.
Hierarchical inference using empirical Bayes provides a mechanism by which neural circuits can selectively up-weight and down-weight particular features in the sensorium.
“In general, hierarchical inference using empirical Bayes, as implemented in our simulations, provides a mechanism by which neural circuits can selectively up-weight and down-weight particular features in the sensorium.”
The paper relies on this without support.
Microdialysis measures of ACh in behaving animals show increases in modality-specific sensory cortex and hippocampus during the acquisition of novel, behaviourally relevant stimuli.
“Microdialysis measures of ACh in behaving animals support this view – with increases observed in modality-specific sensory cortex and hippocampus during the acquisition of novel, behaviourally-relevant stimuli.”
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 source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“In rodent models, superficial pyramidal cell gain is modulated by currently licensed cholinergic treatments for AD. 5”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“During mismatch negativity paradigms, cholinergic agonists increase 5 , 7 and cholinergic antagonists decrease 7 parameters for the gain modulation of superficial pyramidal cells.”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
Experimental — accuracy not yet measuredThe cited source says what the claim attributes to it.
In the citing paper
“Empirical evidence suggests that this precision-weighting is a generic computational process throughout the brain [39] and may be instantiated through neuromodulatory mechanisms of gain control at a synaptic level [42].”
From Active interoceptive inference and the emotional brain (2016) · cites this paper as [42]
The cited source says what the claim attributes to it.
In the citing paper
“I further hypothesize that the salience network tunes the internal model by predicting which prediction errors to pay attention to [i.e. those errors that are likely to be allostatically relevant and therefore worth the cost of encoding and consolidation; called precision signals (Feldman and Friston, 2010; Clark, 2013a,b; Moran et al., 2013; Shipp et al., 2013)].”
From The theory of constructed emotion: an active inference account of interoception and categorization (2016) · cites this paper as Moran et al., 2013
The cited source says what the claim attributes to it.
In the citing paper
“Prediction errors are weighted by precision, reflecting their reliability. 5”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
The cited source says what the claim attributes to it.
In the citing paper
“For example, galantamine modulates activity of superficial pyramidal cells, leading to changes in neurophysiological responses to rapidly changing stimuli.”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
The cited source says what the claim attributes to it.
In the citing paper
“The precision of prediction errors is proposed to be encoded by the gain modulation of pyramidal cells in superficial layers of cortex. 5 , 7”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
The cited source says what the claim attributes to it.
In the citing paper
“Superficial pyramidal cell gain modulation is proposed to encode the precision of prediction errors. 5 , 7”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
The cited source says what the claim attributes to it.
In the citing paper
“This is suggested to encode superficial pyramidal cell gain, 5 , 7 which is reduced by AD (Figure 3).”
From Alzheimer's disease and its progression reduce pyramidal cell gain and connectivity (2025) · cites this paper as [5]
In the citing paper
“Validation studies in humans and animals have shown that DCM for EEG is capable of capturing short-term changes in synaptic efficacy, such as neuromodulation of glutamatergic receptor conductances, and distinguish different types of synaptic plasticity in cortical microcircuits (66, 132, 138–140)”
From Can Bayesian Theories of Autism Spectrum Disorder Help Improve Clinical Practice? (2016) · cites this paper as [140]
In the citing paper
“This broadcasting of precision-weighted prediction errors may rest on neuromodulatory gain control mechanisms at a synaptic level [24].”
From Cerebral hierarchies: predictive processing, precision and the pulvinar (2015) · cites this paper as [24]
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Results > Optimal changes in synaptic efficacy and gain
“model parameters (like C in Equation 2)”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
“This model was used to simulate tones of different frequencies (by changing the parameter C in Equation 3).”
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
“ωv(i)∼N(0,Πx(i)−1)Πx(i)=exp(γx(i))⋅IΠv(i)=exp(γv(i))⋅I”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
“To model a placebo condition we used a Gaussian prior on the sensory log precision (Figure 1): Πv(1)=exp(γ)⋅I”
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
“To model the effects of Galantamine we used a prior with a higher mean and log-precision: p(γ) = N(μt−1 + 2, exp(−4+2))).”
§ Free energy, precision and learning: the role of cholinergic neuromodulation > Materials and Methods > Simulating Brain Responses using Variational Free Energy Minimisation
“To model a placebo condition we used a Gaussian prior on the sensory log precision (Figure 1): Πv(1)=exp(γ)⋅I, with p(γ) = N(μt−1, exp(−4))”
Citations
Cites
The free-energy principle: a unified brain theory?
Friston, K. (2010) · Nature Reviews Neuroscience · doi:10.1038/nrn2787 · Cited by 8,120 · cited 2×
“This type of precision modulation, i.e., state-induced changes in sensory precision, can be included in generalized Bayesian schemes to provide a compelling metaphor for attentional modulation (Feldman and Friston, 2010).”
BackgroundSelf-citationDistributed hierarchical processing in the primate cerebral cortex
Felleman, D. J., & Van Essen, D. C. (1991) · Cerebral Cortex · doi:10.1093/cercor/1.1.1 · Cited by 7,783
Marr, D. (1982) · Cumulative Index of Computer Aided Architectural Design · Cited by 5,733
Rao, R. P. N., & Ballard, D. H. (1999) · Nature Neuroscience · doi:10.1038/4580 · Cited by 5,483 · cited 3×
“Neuronal responses are proposed to reflect inference on the (hidden) causes of sensory stimuli that enable the brain to predict its sensations (Mumford, 1992; Rao and Ballard, 1999; Friston, 2010).”
BackgroundA theory of cortical responses.
Friston, K. (2005) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2005.1622 · Cited by 4,578
Self-citationPreface: Cerebral Cortex Has Come of Age
Goldman, P. S., & RAKIC, P. T. (1991) · Cerebral Cortex · doi:10.1093/cercor/1.1.1-a · Cited by 3,537
Canonical Microcircuits for Predictive Coding
Bastos, A. M., Usrey, W. M., Adams, R. A., Mangun, G. R., Fries, P., & Friston, K. J. (2012) · Neuron · doi:10.1016/j.neuron.2012.10.038 · Cited by 2,421
Self-citationNEURONAL CIRCUITS OF THE NEOCORTEX
Douglas, R. J., & Martin, K. A. C. (2004) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.27.070203.144152 · Cited by 1,886
The free-energy principle: a rough guide to the brain?
Friston, K. (2009) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2009.04.005 · Cited by 1,831
Self-citationUncertainty, Neuromodulation, and Attention
Yu, A. J., & Dayan, P. (2005) · Neuron · doi:10.1016/j.neuron.2005.04.026 · Cited by 1,774
Cited by
Sorted by citations
The theory of constructed emotion: an active inference account of interoception and categorization
Barrett, L. F. (2016) · Social Cognitive and Affective Neuroscience · doi:10.1093/scan/nsw154 · Cited by 1,426
“those errors that are likely to be allostatically relevant and therefore worth the cost of encoding and consolidation; called precision signals (Feldman and Friston, 2010; Clark, 2013a,b; Moran et al., 2013; Shipp et al., 2013)].”
The theory of constructed emotion: an active inference account of interoception and categorization
Barrett, L. F. (2017) · Social Cognitive and Affective Neuroscience · doi:10.1093/scan/nsx060 · Cited by 1,009
“…relevant and therefore worth the cost of encoding and consolidation; called precision signals (Feldman and Friston, 2010; Clark, 2013a,b; Moran et al., 2013; Shipp et al., 2013)].20 Specifically, I hypothesize that precision signals optimize the sampling of the sensory periphery for…”
Active interoceptive inference and the emotional brain
Seth, A. K., & Friston, K. J. (2016) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2016.0007 · Cited by 945
Self-citationPrecise minds in uncertain worlds: Predictive coding in autism.
Van de Cruys, S., Evers, K., Van der Hallen, R., Van Eylen, L., Boets, B., de-Wit, L., & Wagemans, J. (2014) · Psychological Review · doi:10.1037/a0037665 · Cited by 885
An aberrant precision account of autism
Lawson, R. P., Rees, G., & Friston, K. J. (2014) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2014.00302 · Cited by 641
Self-citation
Metadata
- Authors
- Moran, R. J., Campo, P., Symmonds, M., Stephan, K. E., Dolan, R. J., & Friston, K. J.
- Affiliations
- Wellcome Centre for Human Neuroimaging (GB)
- Corresponding author
- Moran, Rosalyn J.
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2013-05-08
- Volume
- 33
- Issue
- 19
- Pages
- 8227-8236
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC4235126
- OpenAlex
- W2013554506
- Cited by
- 312
- References
- 72
- Citations by year
- 2026: 19 · 2025: 19 · 2024: 25 · 2023: 31 · 2022: 25 · 2021: 22 · 2020: 29 · 2019: 19 · 2018: 39 · 2017: 18 · 2016: 22 · 2015: 22 · 2014: 17 · 2013: 5
- FWCI
- 12.02
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Journal of Neuroscience (publishedVersion, cc-by-nc-sa)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- LA Referencia (Red Federada de Repositorios Institucionales de Publicaciones Científicas) (publishedVersion, other-oa)
- Bristol Research (University of Bristol) (publishedVersion)
- Tesis Doctorals en Xarxa (Consorci de Serveis Universitaris de Catalunya) (publishedVersion)
- Zurich Open Repository and Archive (University of Zurich) (acceptedVersion, other-oa)
- UCL Discovery (University College London) (submittedVersion)
- Max Planck Digital Library (submittedVersion, other-oa)
- Hispana (submittedVersion)
- Research Portal (King's College London) (submittedVersion)
- MPG.PuRe (Max Planck Society) (submittedVersion, other-oa)
- Biblos-e Archivo (Universidad Autónoma de Madrid) (submittedVersion, public-domain)
- Explore Bristol Research (submittedVersion)
- Zurich Open Repository and Archive (University of Zurich) (submittedVersion, other-oa)
- Zurich Open Repository and Archive (University of Zurich)
- Funders
- Wellcome Trust (091593, 098362, 098362/Z/12/Z, 088130)
- Ministerio de Ciencia e Innovación
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Embodied and Extended Cognition
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- acetylcholine, cholinergic modulation, free energy principle, perceptual inference, prediction error, sensory precision, gain modulation, cholinesterase inhibition, mismatch negativity, primary sensory cortex, uncertainty, learning
- MeSH terms
- Acetylcholine, Acoustic Stimulation, Adolescent, Adult, Algorithms, Brain, Brain Mapping, Cholinesterase Inhibitors, Computer Simulation, Double-Blind Method, Electroencephalography, Evoked Potentials, Auditory, Female, Galantamine, Humans, Learning, Male, Models, Neurological, Neuropsychological Tests, Perception, Predictive Value of Tests, Young Adult
About these checks
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.
A language model drew the claims from the paper on 2026-10-04, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
A second model labelled each claim on 2026-10-04 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review
