Free Energy, Precision and Learning: The Role of Cholinergic Neuromodulation

May combine two claims.

Simulations of a mismatch negativity paradigm predicted rapid trial-by-trial suppression of evoked sensory prediction error responses, and that cholinergic neuromodulation attenuates this suppression.

“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.”

May combine two claims.

In the placebo simulation, the MMN difference between d10 and d1 was 8.1 a.u., reversed polarity and reduced to −1 a.u. on d2, then gradually increased to baseline −0.1 a.u. for d9.

“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.”

May combine two claims.

The precision estimates show biphasic changes: the initial oddball has a high precision estimate (d1 = 5.1 a.u.), followed by a large reduction (d2 = 2.1 a.u.) and a subsequent rebound (d10 = 4.7 a.u.).

“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).”

May combine two claims.

Empirical sensor-level evoked responses exhibited the same form of repetition suppression as the simulations, with trial d2 more similar to d10 and a deflection around 150 ms for the first oddball tone.

“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).”

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

The study observed a mismatch negativity in response to the onset of a deviant tone at about 150 ms, a timing consistent with earlier studies.

“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.”

May combine two claims.

Under Galantamine simulation, higher a priori precision on sensory prediction error increases the response to the first presentation of a new tone and attenuates the reduction of precision on subsequent presentations, leading to larger responses at the second deviant and more protracted repetition suppression.

“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.”

• AssumptionThe 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.

May combine two claims.

In the Galantamine simulation, the MMN difference between d10 and d1 was 34 a.u., remained positive at 2.5 a.u. on d2, and decreased slowly to baseline for d9.

“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.”

• AssumptionThe 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.

May combine two claims.

In the placebo simulation, a large precision-weighted prediction error occurred with a late deflection around 150 ms for the first deviant, and it almost disappeared on the second trial, which resembled d10.

“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).”

May combine two claims.

The empirical results show an enhanced MMN on Galantamine at d1 compared with placebo, and this enhancement persists at d2.

“Our empirical results show an enhanced MMN on Galantamine at d1 compared to placebo that persists at d2”

May combine two claims.

In the Galantamine simulations, 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.

“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).”

Probably not a substantial claim: may not pass S1 (asserted), S3 (load-bearing). May combine two claims.

Simulating Galantamine as reducing sensory precision produced a reduced MMN of 2.9 a.u. (d10−d1) that reversed polarity on d2 to −0.5.

“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.”

Probably not a substantial claim: may not pass S4 (based).

Optimal inference assigns greater or lesser precision (inverse variance) to different hierarchical levels.

“To weight hierarchical predictions with greater or lesser importance, optimal inference assigns greater or lesser precision (inverse variance) to different hierarchical levels.”

Classified as probably not a substantial claim.

Bottom-up signals are prediction errors that report discrepancies between top-down predictions and representations at each level.

“These signals are prediction errors that report discrepancies between top-down predictions and representations at each level (Kiebel, Daunizeau et al. 2009).”

• cites Kiebel, Daunizeau et al. 2009Not checked yet

  Not checked in the free run.

May combine two claims.

Neuronal responses are proposed to reflect inference on hidden causes of sensory stimuli, enabling the brain to predict its sensations.

“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).”

• cites Mumford 1992Not checked yet

  Not checked in the free run.

• cites Rao and Ballard 1999Not checked yet

  Not checked in the free run.

• cites Friston 2010Not checked yet

  Not checked in the free run.

May combine two claims.

Each level of a processing hierarchy sends predictions to the level below, and the lower level reciprocates with bottom-up signals.

“Each level of a processing hierarchy sends predictions to the level below, which reciprocate bottom-up signals.”

Classified as probably not a substantial claim.

Recurrent message passing continues until prediction errors are minimised throughout the hierarchy.

“This recurrent message passing continues until prediction errors are minimised throughout the hierarchy.”

Classified as probably not a substantial claim.

The free energy framework places predictive coding in a more general setting using dynamic hierarchical generative models of hidden environmental states and an approximate posterior density encoded by synaptic activity and weights.

“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.”

• cites Kiebel, Daunizeau et al. 2009Not checked yet

  Not checked in the free run.

May combine two claims.

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.

“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.”

Classified as probably not a substantial claim.

After model inversion, variational free energy can be used as an approximate log model evidence for Bayesian model comparison.

“After model inversion or optimisation, the variational free energy can then used as an approximate log model evidence for Bayesian model comparison.”

May combine two claims.

There is a remarkable convergence between the predictions of Bayes-optimal inference and learning under uncertainty and the empirical results from the roving mismatch negativity paradigm, both phenomenologically at the sensor level and mechanistically at the source level.

“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”

May combine two claims.

Simulation results suggest one-shot learning or repetition suppression, in which 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 generative model parameters are learned.

“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.”

Classified as probably not a substantial claim.

The canonical microcircuit model is based on a mesoscopic circuit description of cat primary visual cortex that represents cardinal features of functional macrocolumns throughout the cortical sheet.

“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).”

• cites Douglas and Martin 2004Not checked yet

  Not checked in the free run.

• cites Bastos, Usrey et al. 2012Not checked yet

  Not checked in the free run.

May combine two claims.

Learning under the linear model corresponds to gradient descent on free energy and reduces to associative plasticity.

“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).”

• cites Friston 2008Not checked yet

  Not checked in the free run.

May combine two claims.

The CMC model explicitly models the laminar specificity of forward and backward connections central to predictive coding, with forward connections conveying prediction errors to higher levels and backward connections sending predictions to lower areas.

“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.”

Classified as probably not a substantial claim.

The DCM generates a predicted ERP from a network of coupled sources responding to sensory thalamic input, with each source a neural mass model of several neuronal subpopulations.

“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.”

Probably not a substantial claim: may not pass S4 (based).

Greater acetylcholine levels enhance postsynaptic responses in the superficial layers of primary auditory cortex.

“where greater ACh levels enhance postsynaptic responses in the superficial layers of primary auditory cortex.”

May combine two claims.

Acetylcholine modulates this process; for example, activation of muscarinic receptors suppresses sIAHP through a cGMP-dependent mechanism.

“Importantly, acetylcholine modulates this process; e.g., by activation of muscarinic receptors suppresses sIAHP through a cGMP-dependent mechanism (Krause and Pedarzani 2000).”

• cites Krause and Pedarzani 2000Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim. May combine two claims.

The predictive coding scheme encodes beliefs about the world by associating synaptic activity with expected states, synaptic connection strengths with causal structure, and synaptic gain with the precision of random fluctuations.

“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.”

• cites Friston 2008Not checked yet

  Not checked in the free run.

May combine two claims.

Extrinsic connections in these models follow predictive coding rules: 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.

“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).”

• cites Felleman and Van Essen 1991Not checked yet

  Not checked in the free run.

May combine two claims.

The roving-oddball paradigm elicits mismatch potentials that are automatic or pre-attentive, so subjects need not attend to auditory stimuli.

“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).”

• cites Garrido, Kilner et al. 2009Not checked yet

  Not checked in the free run.

• cites Näätänen, Astikainen et al. 2010Not checked yet

  Not checked in the free run.

May combine two claims.

The effect of Galantamine is best explained as a change in the gain of superficial pyramidal cells in primary auditory cortex, and DCM of a temporo-frontal network attributes the observed differences in event-related responses to cholinergic gain modulation of supragranular pyramidal cells in early auditory cortex.

“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).”

• AssumptionThe 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.
• AssumptionThe authors state this as an assumption: In predictive coding formulations of free energy minimisation, gain corresponds to the precision of prediction errors.

Probably not a substantial claim: may not pass S3 (load-bearing).

DCMs are inverted using the same variational scheme (variational Laplace) that underlies predictive coding.

“Interestingly, DCMs are inverted using exactly the same variational scheme (variational Laplace) that underlies predictive coding (Friston et al 2007).”

• cites Friston et al 2007Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim. May combine two claims.

Minimising variational free energy provides a recognition density that approximates the true posterior, a process known as approximate Bayesian inference.

“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.”

Classified as probably not a substantial claim.

The canonical microcircuit model used for each source includes infra- and supragranular pyramidal populations, a granular layer of stellate cells, and an inhibitory interneuron population.

“Each source includes four neuronal subpopulations: infra- and supragranular pyramidal cell populations, a granular layer of stellate cells, and an inhibitory interneuron population.”

• cites Douglas and Martin 2004Not checked yet

  Not checked in the free run.

• cites Bastos, Usrey et al. 2012Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Uncorrected results at p < 0.05 were reported because the source analyses only served to identify candidate sources for the subsequent DCM analysis.

“Note that we report uncorrected results at p < 0.05 because these analyses only served to identify candidate sources in the subsequent DCM analysis.”

Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

After an oral dose, Galantamine reaches peak plasma concentration within about 2 hours and declines with a half-life of about 7 hours.

“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).”

• cites Huang, Lasseter et al. 2002Not checked yet

  Not checked in the free run.

May combine two claims.

This activation may maintain a high postsynaptic gain and facilitate oscillatory activity, with a shift towards higher frequencies in the power spectra of neuronal recordings.

“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).”

• cites Liljenstrom and Hasselmo 1995Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

Bayesian filtering or predictive coding can be formulated as Bayesian updates driven by precision-weighted prediction errors.

“This Bayesian filtering or predictive coding can be formulated in terms of Bayesian updates that are driven by precision-weighted prediction errors.”

Probably not a substantial claim: may not pass S4 (based).

The precision weighting of prediction error units depends on their postsynaptic gain.

“The precision weighting of these prediction errors units depends on their postsynaptic gain (c.f., Kalman gain).”

May combine two claims.

The first deviant stimulus produces large prediction errors boosted by an inappropriately high precision established by the preceding predictable standards, producing the mismatch negativity.

“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.”

May combine two claims.

Slow after-hyperpolarizing currents (sIAHP) mediated by calcium-dependent potassium channels decrease neuronal excitability and firing rate.

“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).”

• cites Faber and Sah 2003Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

Prediction errors are generally assumed to be encoded by pyramidal cells in superficial cortical layers that receive top-down predictions from deeper laminae in higher regions.

“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).”

• cites Mumford 1992Not checked yet

  Not checked in the free run.

• cites Cauller 1995Not checked yet

  Not checked in the free run.

• cites Bar 2003Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

The study used a within-subject crossover placebo-controlled double-blind design.

“We studied 13 right-handed, healthy volunteers (5 female, aged 25 ± 7 yrs), using a within-subject crossover placebo-controlled double-blind design.”

Classified as probably not a substantial claim.

DCM models ERPs as the response of a dynamic input-output system to exogenous experimental inputs.

“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).”

• cites David, Kiebel et al. 2006Not checked yet

  Not checked in the free run.

• cites Kiebel, David et al. 2006Not checked yet

  Not checked in the free run.

May combine two claims.

Galantamine also enhances cholinergic neurotransmission by sensitising nicotinic receptors, increasing the probability of acetylcholine-induced channel opening and slowing receptor desensitisation.

“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).”

• cites Coyle and Kershaw 2001Not checked yet

  Not checked in the free run.

• cites Samochocki, Höffle et al. 2003Not checked yet

  Not checked in the free run.

May combine two claims.

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, and the study supplements this with a functional anatomy at the systems level.

“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.”

May combine two claims.

Pharmacologically augmenting cholinergic neuromodulation appears to boost event-related responses to deviant stimuli and attenuate their subsequent repetition suppression.

“In summary, pharmacologically augmenting cholinergic neuromodulation appears to boost event related responses to deviant stimuli and attenuate their subsequent repetition suppression.”

May combine two claims.

ACh has long been known to enhance the firing of cortical pyramidal neurons and to induce a decrease in spike frequency adaptation via several mechanisms, including reduction of hyperpolarizing potassium currents.

“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).”

• cites Krnjevi and Phillis 1963Not checked yet

  Not checked in the free run.

• cites Benardo and Prince 1982Not checked yet

  Not checked in the free run.

• cites Cole and Nicoll 1984Not checked yet

  Not checked in the free run.

May combine two claims.

The proposition that neuromodulators encode the precision or predictability of environmental states is supported by empirical evidence that sensory regions respond with enhanced gain under cholinergic enhancement, producing exaggerated and prolonged mismatch responses localised to superficial pyramidal populations encoding prediction error.

“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).”

• cites Mumford 1992Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

The EEG measurements do not allow the study to characterise the source of cholinergic modulation per se, such as between prefrontal regions and basal forebrain cholinergic neurons.

“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.”

Classified as probably not a substantial claim. May combine two claims.

Previous theoretical circuit-level proposals suggest that ACh supports learning and memory by switching the cortex from a read-out to a read-in mode, enhancing sensory-evoked afferent responses and suppressing internal cortical processing.

“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).”

• cites Hasselmo and McGaughy 2004Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

The resulting adaptive sensory bias may be implemented via gain control in a manner suggested by previous physiological accounts of cholinergic modulation.

“The resulting adaptive sensory bias may be implemented via gain control in a manner suggested by previous physiological accounts of cholinergic modulation.”

May combine two claims.

The study shows an equivalent and complementary effect, where ACh boosts bottom-up sensory evidence in the auditory hierarchy.

“We show here an equivalent and complementary effect, where ACh boosts bottom-up sensory evidence in the auditory hierarchy.”

Probably not a substantial claim: may not pass S3 (load-bearing).

It has been demonstrated in the auditory domain that a sensory memory relies upon cholinergic modulation.

“Indeed, it has been demonstrated in the auditory domain that a sensory “memory” relies upon cholinergic modulation (Miasnikov, Chen et al. 2008).”

• cites Miasnikov, Chen et al. 2008Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S3 (load-bearing).

The analysis of simulated and empirical responses recapitulates some established ideas about the role of cholinergic signalling in perception and attention.

“Our analysis of the simulated and empirical responses recapitulates some of these established ideas about the role of cholinergic signalling in perception and attention.”

May combine two claims.

The findings suggest that acetylcholine mediates the representation of precision and acts to facilitate the bottom-up signalling of sensory information through precise prediction errors.

“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).”

May combine two claims.

Human studies with cholinergic agonists show enhanced specificity of perceptual learning for behaviourally relevant features and diminished validity effects in spatial cueing tasks, indicating selective boosting of stimulus-related information.

“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.”

• cites Rokem and Silver 2010Not checked yet

  Not checked in the free run.

• cites Bentley, Husain et al. 2004Not checked yet

  Not checked in the free run.

• cites Vossel, Thiel et al. 2007Not checked yet

  Not checked in the free run.

Classified as probably not a substantial claim.

The framework of Yu and Dayan proposes that ACh modulates uncertainty about high-level representations.

“The framework of Yu and Dayan (Yu and Dayan 2002) proposes that ACh modulates uncertainty about high-level representations.”

• cites Yu and Dayan 2002Not checked yet

  Not checked in the free run.

Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

This proposal is consistent with the current study.

“This proposal is consistent with the current study.”

Probably not a substantial claim: may not pass S3 (load-bearing).

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.

“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).”

• cites Garrido, Kilner et al. 2009Not checked yet

  Not checked in the free run.