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A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP
Open accessAbstract
According to the Hebb rule, the change in the strength of a synapse depends only on the local interaction of presynaptic and postsynaptic events. Studies at many types of synapses indicate that the early phase of long-term potentiation (LTP) has Hebbian properties. However, it is now clear that the Hebb rule does not account for late LTP; this requires an additional signal that is non-local. For novel information and motivational events such as rewards, this signal at hippocampal CA1 synapses is mediated by the neuromodulator, dopamine. In this Review, we discuss recent experimental findings that support the view that this “neoHebbian” framework can account for memory behavior in a variety of learning situations.
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TL;DR
This review proposes a neoHebbian framework in which Hebbian early LTP at hippocampal CA1 synapses is supplemented by a third, non-local dopamine signal required for late LTP and stable episodic memory. It synthesizes rodent and human evidence that novelty, reward, and aversive/motivational events trigger dopamine release that selectively stabilizes memory at long delays and can enhance temporally nearby weak memories via a synaptic tag-and-capture “penumbra.”
Key findings
- Late LTP in CA1 requires D1 dopamine receptor signaling: D1 antagonists, D1 receptor deletion, or dopamine depletion block late LTP, while early LTP is less affected. In a rodent one-trial reward-location memory task, intrahippocampal D1/D5 antagonist SCH23390 before encoding impaired memory at 24 hours but not at 30 minutes.
- Dopamine neurons burst in response to reward, novelty, and aversive stimuli; bursts last about 200–500 msec, have interspike intervals <100 msec, and generally contain about 3–5 spikes, with up to 50% of dopamine neurons silent at baseline. In mice, 89% of VTA dopaminergic neurons responded with activation to a conditioned reward, the majority showed suppression to fearful events, and 25% responded with excitation to both positive and fearful events.
- Human fMRI studies show SN/VTA activation by reward, novelty, and aversive events. Reward prospect improved long-term episodic memory, with enhancement greater at late timepoints (3 weeks vs. 20 minutes) and consistently after 24-hour retention intervals.
- Novelty can create a behavioral “penumbra”: in rodents, a single reward-location exposure normally forgotten within hours persisted for at least 24 hours if preceded or followed within 30 minutes by novelty exploration; in humans, novel scenes presented 5 minutes before pre-familiarized words increased later recollection, with a stronger effect at 24 hours than at 30 minutes.
Methods
- Review article; no new experimental data are reported.
- Synthesizes electrophysiological, pharmacological, genetic, and behavioral studies in rodents, plus human fMRI, PET, and electrophysiological findings.
- Main analysis is a narrative integration of evidence around dopamine-dependent late LTP, dopamine neuron firing properties, and the synaptic tag-and-capture model.
Limitations
- The paper states that more human studies are needed, especially on dopaminergic manipulations and long-term memory; selective D1/D5 compounds are not readily available for healthy humans, and fMRI cannot distinguish SN/VTA subregions or tonic versus bursting modes. It also notes unresolved questions about the duration of hippocampal dopamine signals, penumbra properties, and the relative consolidation of rewarding versus aversive events in humans.
- Evident from the text: as a review, it does not use a systematic search or meta-analytic method, and some proposed circuitry—such as perirhinal cortical information generating phasic dopamine responses—is explicitly described as needing experimental testing.
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95 Not checked yet · 28 Own interpretation · 12 No basis given · 1 Own result
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In the paper
“We term this “neoHebbian” to indicate that in addition to the 2-factor Hebbian process, stable synaptic modification requires a third signal dependent on a systems level computation.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
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“Dopamine enhances protein synthesis within dendrites of hippocampal neurons [85].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [85] Smith WB 2005 Dopaminergic stimulation of local protein synthesis enhances surface expression of GluR1 and synaptic transmission in hippocampal neurons Neuron 45 765 779 15748851 source
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In the paper
“The ability to induce LTP at single visualizable synaptic connections by 2-photon uncaging of glutamate has directly confirmed the synapse specificity of LTP [5].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [5] Matsuzaki M 2004 Structural basis of long-term potentiation in single dendritic spines Nature 429 761 766 15190253 PMC4158816 source
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In the paper
“Hippocampal late LTP is blocked when ERK1/2 activation is blocked by a dominant-negative mutation of MEK1, the kinase that directly phosphorylates ERK1/2 [88].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [88] Kelleher RJ 3rd 2004 Translational control by MAPK signaling in long-term synaptic plasticity and memory Cell 116 467 479 15016380 source
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“Experiments show that genetic or pharmacological interference with NMDARs or CaMKII strongly interferes with memory formation [6].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [6] Lisman J 2002 The molecular basis of CaMKII function in synaptic and behavioural memory Nature reviews 3 175 190 11994750 source
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In the paper
“There is thus little doubt that a Hebbian form of LTP is important for memory formation.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
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In the paper
“there is converging evidence from fMRI studies that the human SN/VTA is also activated by novelty (see main text and Figure I).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Novelty
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In the paper
“Early experiments showed that LTP in CA1 is governed by the Hebb rule: the induction of LTP requires both presynaptic input and strong postsynaptic depolarization (the role of Na+ spikes remains unclear) [3].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [3] Wigstrom H Gustafsson B 1986 Postsynaptic control of hippocampal long-term potentiation J Physiol (Paris) 81 228 236 2883309
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In the paper
“Microdialysis [90, 91] and fast scan cyclic voltammetry [92] show that various types of novelty produce dopamine release in the rat striatum. These include social novelty [90], novel tastes [91], and novel environments [92].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Novelty
What the sources say
- Not checked yetcites [90] De Leonibus E 2006 Distinct kinds of novelty processing differentially increase extracellular dopamine in different brain regions The European journal of neuroscience 23 1332 1340 16553794 source
- Not checked yetcites [91] Bassareo V 2002 Differential Expression of Motivational Stimulus Properties by Dopamine in Nucleus Accumbens Shell versus Core and Prefrontal Cortex J Neurosci 22 4709 4719 12040078 PMC6758788 source
- Not checked yetcites [92] Rebec GV 1998 Real-time assessments of dopamine function during behavior: single-unit recording, iontophoresis, and fast-scan cyclic voltammetry in awake, unrestrained rats Alcohol Clin Exp Res 22 32 40 9514283 source
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In the paper
“studies of LTP (mostly in CA1) show that there is an additional factor: the persistence of LTP depends not just on the two factors of the Hebbian condition (glutamate release and postsynaptic depolarization), but also on a third, the action of the neurotransmitter dopamine [7]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [7] Frey U 1990 Dopaminergic antagonists prevent long-term maintenance of posttetanic LTP in the CA1 region of rat hippocampal slices Brain research 522 69 75 1977494 source
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In the paper
“Experimental results suggest that D1 receptor- and BDNF-mediated pathways interact in the activation of extracellular-signal regulated kinase (ERK) 1/2 [mitogen activated kinase (MAPK)] activity [87].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [87] Yoshii A Constantine-Paton M 2010 Postsynaptic BDNF-TrkB signaling in synapse maturation, plasticity, and disease Dev Neurobiol 70 304 322 20186705 PMC2923204 source
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In the paper
“Importantly, the dopamine release depends on systems-level processes that include motivation and stimulus novelty [8]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [8] Lisman JE Grace AA 2005 The hippocampal-VTA loop: controlling the entry of information into long-term memory Neuron 46 703 713 15924857 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Studies at many types of synapses indicate that the early phase of long-term potentiation (LTP) has Hebbian properties.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Abstract
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“For novel information and motivational events such as rewards, this signal at hippocampal CA1 synapses is mediated by the neuromodulator, dopamine.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Abstract
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In the paper
“Placing a rat in a novel environment leads to dopamine release in the hippocampus [19]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Novelty
What the sources say
- Not checked yetcites [19] Ihalainen JA 1999 Comparison of dopamine and noradrenaline release in mouse prefrontal cortex, striatum and hippocampus using microdialysis Neurosci Lett 277 71 74 10624812 source
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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 this Review, we discuss recent experimental findings that support the view that this “neoHebbian” framework can account for memory behavior in a variety of learning situations.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Two types of experiments demonstrate that dopamine can strengthen the synaptic potentiation produced by learning itself.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
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In the paper
“Together these results make a strong case for the involvement of dopamine in late LTP.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1
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In the paper
“It is therefore thought that dopamine antagonists prevent late LTP because they prevent the required protein synthesis (see Box 1).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1
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In the paper
“Protein synthesis-dependent forms of consolidation are not restricted to the hippocampus [127].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Does the neoHebbian framework apply to extrahippocampal MTL structures?
What the sources say
- Not checked yetcites [127] Balderas I 2008 The consolidation of object and context recognition memory involve different regions of the temporal lobe Learning & memory (Cold Spring Harbor, N Y 15 618 624 PMC2632790 18723431 source
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“The human lateral SN/VTA has been similarly shown to be activated during anticipation (i.e. before execution of motor responses) to acquire a reward or to avoid a punishment [53].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Interactions between valence and action
What the sources say
- Not checked yetcites [53] Guitart-Masip M 2011 Action dominates valence in anticipatory representations in the human striatum and dopaminergic midbrain Journal of Neuroscience 31 7867 7875 21613500 PMC3109549 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“LCholinergic [136] and noradrenergic [10] projections to MTL can also modulate LTP and long-term memory.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Do other neuromodulators contribute to neoHebbian plasticity?
What the sources say
- Not checked yetcites [136] Sarter M 2005 Unraveling the attentional functions of cortical cholinergic inputs: interactions between signal-driven and cognitive modulation of signal detection Brain Res Brain Res Rev 48 98 111 15708630 source
- Not checked yetcites [10] Frey S Frey JU 2008 ‘Synaptic tagging’ and ‘cross-tagging’ and related associative reinforcement processes of functional plasticity as the cellular basis for memory formation Progress in brain research 169 117 143 18394471 source
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“Indeed, sustained aversive stimuli or stressors (eg. repetitive footshocks or physical restraint), increase the number of dopamine neurons firing over longer periods of time [102].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Aversive events
What the sources say
- Not checked yetcites [102] Valenti O 2011 Aversive Stimuli Alter Ventral Tegmental Area Dopamine Neuron Activity via a Common Action in the Ventral Hippocampus J Neurosci 31 4280 4289 21411669 PMC3066094 source
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“It was found that D1 antagonists prevent this stabilization [20].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
What the sources say
- Not checked yetcites [20] Kentros CG 2004 Increased attention to spatial context increases both place field stability and spatial memory Neuron 42 283 295 15091343 source
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In the paper
“after learning several reward-location associations in an environment, rats rapidly (i.e., in only one trial) update their memory if one of the reward-location associations is changed [25].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > D1 antagonists block 1 day old memory without affecting early memory
What the sources say
- Not checked yetcites [25] Tse D 2007 Schemas and memory consolidation Science (New York, N Y 316 76 82 17412951 source
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In the paper
“This suggests that SN/VTA responses to aversive events are observed primarily under conditions that require active, rather than passive, avoidance.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Interactions between valence and action
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In the paper
“Plasticity in the hippocampus subserves the particular form of memory called episodic memory [24].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > D1 antagonists block 1 day old memory without affecting early memory
What the sources say
- Not checked yetcites [24] Tulving E 1985 Memory and consciousness Canadian Psychology 26 1 12 source
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“Subsequent work, however, showed that, with less strong induction protocols, there is a partial inhibition of early LTP [23].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
What the sources say
- Not checked yetcites [23] Otmakhova NA Lisman JE 1996 D1/D5 dopamine receptor activation increases the magnitude of early long-term potentiation at CA1 hippocampal synapses J Neurosci 16 7478 7486 8922403 PMC6579102 source
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In the paper
“Dopamine antagonists given only during such consolidation disrupt memory measured many hours later [144, 145].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > How does the neoHebbian framework interact with off-line replay?
What the sources say
- Not checked yetcites [144] Rossato JI 2009 Dopamine controls persistence of long-term memory storage Science (New York, N Y 325 1017 1020 19696353 source
- Not checked yetcites [145] Guzman-Ramos K 2010 Off-line concomitant release of dopamine and glutamate involvement in taste memory consolidation Journal of neurochemistry 114 226 236 20412388 source
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In the paper
“As a result of this inhibition, up to 50% of dopamine neurons are in the silent mode at baseline.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
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In the paper
“Indeed, according to the neoHebbian framework, activating the SN/VTA by a reward should enhance hippocampal-dependent episodic memory of information present at the time of reward.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
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“In contrast, higher-concentration phasic dopamine signals activate D1 receptors and thereby enhance subicular input [121].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [121] Goto Y Grace AA 2005 Dopaminergic modulation of limbic and cortical drive of nucleus accumbens in goal-directed behavior Nature neuroscience 8 805 812 15908948 source
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In the paper
“Tonic dopamine, which works through the higher-affinity D2 receptors, attenuates medial PFC (mPFC) inputs.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
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In the paper
“The novelty processing hierarchy of the MTL (Figure I) is part of the circuitry that can regulate dopamine responses to novelty.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
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In the paper
“conversely, simulating burst firing using channelrhodopsin is sufficient to induce behavioral conditioning [42].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [42] Tsai HC 2009 Phasic firing in dopaminergic neurons is sufficient for behavioral conditioning Science (New York, N Y 324 1080 1084 PMC5262197 19389999 source
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In the paper
“Burst firing in dopaminergic neurons is potently driven by the brainstem pedunculopontine tegmentum (PPTg), which provides a glutamatergic input that acts via NMDARs ([39, 40] see Box 3).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [39] Floresco SB 2003 Afferent modulation of dopamine neuron firing differentially regulates tonic and phasic dopamine transmission Nature neuroscience 6 968 973 12897785 source
- Not checked yetcites [40] Lodge DJ Grace AA 2006 The hippocampus modulates dopamine neuron responsivity by regulating the intensity of phasic neuron activation Neuropsychopharmacology 31 1356 1361 16319915 source
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In the paper
“Despite these difficulties, there is converging evidence for a close relationship between the fMRI response in the SN/VTA and dopamine release (reviewed in [50]).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
What the sources say
- Not checked yetcites [50] Duzel E 2009 Functional imaging of the human dopaminergic midbrain Trends in neurosciences 32 321 328 19446348 source
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In the paper
“Strong evidence for hippocampal involvement in novelty-dependent dopamine release comes from experiments showing that this release is blocked by inactivating the subiculum, an output structure of the hippocampus that projects, by a polysynaptic pathway, to the SN/VTA (Figure I).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
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In the paper
“The PPTg regulates burst firing of dopamine cells rather than their tonic resting activity [[39]].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [39] Floresco SB 2003 Afferent modulation of dopamine neuron firing differentially regulates tonic and phasic dopamine transmission Nature neuroscience 6 968 973 12897785 source
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In the paper
“Such studies (e.g. [48, 51]) have demonstrated that the human SN/VTA is activated by rewards, a finding confirmed by a recent study using electrophysiological recordings [52].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
What the sources say
- Not checked yetcites [48] D’Ardenne K 2008 BOLD responses reflecting dopaminergic signals in the human ventral tegmental area Science (New York, N Y 319 1264 1267 18309087 source
- Not checked yetcites [51] Guitart-Masip M 2010 Contextual novelty changes reward representations in the striatum J Neurosci 30 1721 1726 20130181 PMC2838369 source
- Not checked yetcites [52] Zaghloul KA 2009 Human substantia nigra neurons encode unexpected financial rewards Science (New York, N Y 323 1496 1499 PMC2839450 19286561 source
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In the paper
“learning of responses to aversive stimuli in rodents (ie. during fear conditioning) is inhibited by dopaminergic antagonists [66–68].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [66] Fadok JP 2009 Dopamine is necessary for cue-dependent fear conditioning J Neurosci 29 11089 11097 19741115 PMC2759996 source
- Not checked yetcites [67] Ortiz O 2010 Associative learning and CA3-CA1 synaptic plasticity are impaired in D1R null, Drd1a−/− mice and in hippocampal siRNA silenced Drd1a mice J Neurosci 30 12288 12300 20844125 PMC6633447 source
- Not checked yetcites [68] Shen YL 2010 Dopamine receptor antagonists impair place conditioning after acute stress in rats Behav Pharmacol 21 77 82 20038833 source
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In the paper
“the observation that there are dopaminergic responses to aversive events (see Box 2)”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites Box 2
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In the paper
“dopamine is required for association between stimuli and aversive events in the amygdala [69].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [69] Rosenkranz JA Grace AA 2002 Dopamine-mediated modulation of odour-evoked amygdala potentials during pavlovian conditioning Nature 417 282 287 12015602 source
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In the paper
“suggests that dopaminergic consolidation mechanisms similar to those seen with novelty and reward may apply.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
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In the paper
“It is noteworthy that earlier cognitive psychology experiments on the effects of reward on memory used shorter retention intervals than the aforementioned studies (e.g. [58, 59]).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
What the sources say
- Not checked yetcites [58] Craik FIM Tulving E 1975 Depth of Processing and the Retention of Words in Episodic Memory Journal ol Experimental Psychology: General 3 268 294 source
- Not checked yetcites [59] Ngaosuvan L Mantyla T 2005 Rewarded remembering: dissociations between self-rated motivation and memory performance Scandinavian journal of psychology 46 323 330 16014076 source
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In the paper
“stronger stimulation produces the dopamine-dependent protein synthesis that allows late LTP.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
What the sources say
- Not checked yetcites [9] Frey U Morris RG 1997 Synaptic tagging and long-term potentiation [see comments] Nature 385 533 536 9020359 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“There is now converging evidence that different forms of novelty signals are computed along the medial temporal lobe (MTL) processing hierarchy from the parahippocampal and perirhinal cortices to the entorhinal cortex and from there to the hippocampus (see Box 3).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
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In the paper
“Interestingly, the personality trait of novelty seeking is correlated with the magnitude of the SN/VTA novelty fMRI response even when novel stimuli predict the absence of reward [56]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [56] Krebs RM 2009 Personality traits are differentially associated with patterns of reward and novelty processing in the human substantia nigra/ventral tegmental area Biological psychiatry 65 103 110 18835480 source
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In the paper
“Memory enhancement after long retention intervals (e.g. 24 hours) has been consistently found [56, 57].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
What the sources say
- Not checked yetcites [56] Krebs RM 2009 Personality traits are differentially associated with patterns of reward and novelty processing in the human substantia nigra/ventral tegmental area Biological psychiatry 65 103 110 18835480 source
- Not checked yetcites [57] Wittmann B 2011 Behavioural specifications of reward-associated long-term memory enhancement in humans Learning and Memory 18 296 300 21502336 PMC3465832 source
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In the paper
“Moreover, the enhancement was greater at late timepoints than at early intervals (ie. 3 weeks vs. 20 minutes) [47].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
What the sources say
- Not checked yetcites [47] Wittmann BC 2005 Reward-related FMRI activation of dopaminergic midbrain is associated with enhanced hippocampus-dependent long-term memory formation Neuron 45 459 467 15694331 source
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In the paper
“This reward-related memory enhancement was associated with a co-activation of SN/VTA, striatum, and hippocampus, as detected by fMRI [46, 47].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
What the sources say
- Not checked yetcites [46] Adcock RA 2006 Reward-motivated learning: mesolimbic activation precedes memory formation Neuron 50 507 517 16675403 source
- Not checked yetcites [47] Wittmann BC 2005 Reward-related FMRI activation of dopaminergic midbrain is associated with enhanced hippocampus-dependent long-term memory formation Neuron 45 459 467 15694331 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“According to this model, weak stimulation induces only early LTP.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
What the sources say
- Not checked yetcites [9] Frey U Morris RG 1997 Synaptic tagging and long-term potentiation [see comments] Nature 385 533 536 9020359 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“It was found that the prospect of receiving reward (e.g., monetary gain) improved long-term episodic memory for novel stimuli in incidental [47, 55–57] and intentional [46] encoding paradigms.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
What the sources say
- Not checked yetcites [47] Wittmann BC 2005 Reward-related FMRI activation of dopaminergic midbrain is associated with enhanced hippocampus-dependent long-term memory formation Neuron 45 459 467 15694331 source
- Not checked yetcites [55] Callan DE Schweighofer N 2008 Positive and negative modulation of word learning by reward anticipation Human brain mapping 29 237 249 17390317 PMC6870695 source
- Not checked yetcites [56] Krebs RM 2009 Personality traits are differentially associated with patterns of reward and novelty processing in the human substantia nigra/ventral tegmental area Biological psychiatry 65 103 110 18835480 source
- Not checked yetcites [57] Wittmann B 2011 Behavioural specifications of reward-associated long-term memory enhancement in humans Learning and Memory 18 296 300 21502336 PMC3465832 source
- Not checked yetcites [46] Adcock RA 2006 Reward-motivated learning: mesolimbic activation precedes memory formation Neuron 50 507 517 16675403 source
plaindirect- Asserted: passes — The authors put it forward as true.
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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
“Only if a systems-wide computation determines that there is a high level of novelty or motivational salience will dopamine be released, allowing the biochemical processes of late LTP to incorporate the information into long-term memory.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
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In the paper
“Studies of LTP have shown that strong stimuli can enhance the encoding of weak stimuli even if the strong stimulus is given many minutes before or after the weak one.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“pre-exposure to familiar, but emotionally very negative, photographs did not cause a comparable enhancement.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [74] Fenker DB 2008 Novel scenes improve recollection and recall of words J Cogn Neurosci 20 1250 1265 18284351 source
plain- Asserted: passes — The authors put it forward as true.
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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
“Generally, this newly stored information will fade away as early LTP declines.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“In aging, there is now converging evidence from human imaging studies for a structural degeneration of the VTA/SN (for reviews, see [54, 81]).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [54] Duzel E 2010 NOvelty-related motivation of anticipation and exploration by dopamine (NOMAD): implications for healthy aging Neuroscience and biobehavioral reviews 34 660 669 19715723 source
- Not checked yetcites [81] Backman L 2010 Linking cognitive aging to alterations in dopamine neurotransmitter functioning: recent data and future avenues Neuroscience and biobehavioral reviews 34 670 677 20026186 source
boostedcorrelational- Asserted: passes — The authors put it forward as true.
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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
“It is therefore unlikely that affective arousal can account for the memory enhancement after novelty exposure.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
hedged- Asserted: passes — The authors put it forward as true.
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In the paper
“However, the synapses that produce the PRPs can be different from those at which the tag is set.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
What the sources say
- Not checked yetcites [9] Frey U Morris RG 1997 Synaptic tagging and long-term potentiation [see comments] Nature 385 533 536 9020359 source
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“This effect was stronger after 24 hours as compared to 30 minutes after encoding [74].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [74] Fenker DB 2008 Novel scenes improve recollection and recall of words J Cogn Neurosci 20 1250 1265 18284351 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“intrahippocampal application of the D1/D5 receptor antagonist SCH23390 during novelty exploration diminishes memory retention [72].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [72] Wang SH 2010 Relevance of synaptic tagging and capture to the persistence of long-term potentiation and everyday spatial memory Proceedings of the National Academy of Sciences of the United States of America 107 19537 19542 20962282 PMC2984182 source
plaindirect- Asserted: passes — The authors put it forward as true.
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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
“so the duration of dopamine elevations may be relatively long.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
hedged- Asserted: passes — The authors put it forward as true.
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In the paper
“Behavioral experiments suggest that there are differences in the penumbra duration for novel as compared to rewarding events [57]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [57] Wittmann B 2011 Behavioural specifications of reward-associated long-term memory enhancement in humans Learning and Memory 18 296 300 21502336 PMC3465832 source
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“24hr memory is enhanced in rats for rewarded locations (even after only a single exposure)”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [72] Wang SH 2010 Relevance of synaptic tagging and capture to the persistence of long-term potentiation and everyday spatial memory Proceedings of the National Academy of Sciences of the United States of America 107 19537 19542 20962282 PMC2984182 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“It is thus understandable that severe episodic memory problems are not a symptom of early PD [80].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [80] Perry RJ Hodges JR 1996 Spectrum of memory dysfunction in degenerative disease Current opinion in neurology 9 281 285 8858186 source
plain- Asserted: passes — The authors put it forward as true.
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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
“Hence, the behavioral penumbra seen after exposure to novelty in humans is compatible with the synaptic tag and capture model.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
plaincorrelational- Asserted: passes — The authors put it forward as true.
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None of the sources cited for this claim have been checked yet.
In the paper
“exploration of a novel environment can make a memory for a ‘weak’ event more persistent [70–72].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [70] Moncada D Viola H 2007 Induction of long-term memory by exposure to novelty requires protein synthesis: evidence for a behavioral tagging J Neurosci 27 7476 7481 17626208 PMC6672624 source
- Not checked yetcites [71] Ballarini F 2009 Behavioral tagging is a general mechanism of long-term memory formation Proceedings of the National Academy of Sciences of the United States of America 106 14599 14604 19706547 PMC2732837 source
- Not checked yetcites [72] Wang SH 2010 Relevance of synaptic tagging and capture to the persistence of long-term potentiation and everyday spatial memory Proceedings of the National Academy of Sciences of the United States of America 107 19537 19542 20962282 PMC2984182 source
plaindirect- Asserted: passes — The authors put it forward as true.
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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 results reviewed here indicate that the neoHebbian framework has explanatory power in the understanding of memory.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
hedged- Asserted: passes — The authors put it forward as true.
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In the paper
“The idea that long-term memory depends on more than the Hebbian condition makes sense, given the likelihood that too much information written into memory might lead to overwriting (and thus degradation) of pre-existing memories [77].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [77] Blumenfeld B 2006 Dynamics of memory representations in networks with novelty-facilitated synaptic plasticity Neuron 52 383 394 17046699 source
hedgeddirect- Asserted: passes — The authors put it forward as true.
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In the paper
“this long-term memory can be blocked by a D1/D5 receptor antagonist at the time of encoding [72].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [72] Wang SH 2010 Relevance of synaptic tagging and capture to the persistence of long-term potentiation and everyday spatial memory Proceedings of the National Academy of Sciences of the United States of America 107 19537 19542 20962282 PMC2984182 source
plaindirect- Asserted: passes — The authors put it forward as true.
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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
“It therefore seems likely that these principles may apply across all mammals.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
hedged- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: The conditions that lead to activation of the human dopamine system, as judged by fMRI signals, are similar to the conditions leading to activation in non-primate mammals.
In the paper
“The result was a higher proportion of recollected (as opposed to those recognized on the basis of familiarity) words when compared to individuals who were exposed to familiar pictures prior to encoding.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [74] Fenker DB 2008 Novel scenes improve recollection and recall of words J Cogn Neurosci 20 1250 1265 18284351 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Consistent findings have also been recently reported in aged rats [82].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [82] Sanchez HL 2008 Dopaminergic mesencephalic systems and behavioral performance in very old rats Neuroscience 154 1598 1606 18554807 source
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“The dopamine transporter (DAT), which rapidly clears dopamine from the synaptic cleft, is much less abundant in the hippocampus than in the striatum [75]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [75] Schott BH 2006 The dopaminergic midbrain participates in human episodic memory formation: evidence from genetic imaging J Neurosci 26 1407 1417 16452664 PMC6675495 source
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“However, at late stages, when all types of dopaminergic projection neurons are affected, memory problems do become severe.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [80] Perry RJ Hodges JR 1996 Spectrum of memory dysfunction in degenerative disease Current opinion in neurology 9 281 285 8858186 source
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
“Taken together, these findings suggest that there is a “penumbra” surrounding events that cause dopamine release; the memory for events that occur before or after the dopamine release would depend not only on their own properties, but also on whether they fell within the penumbra of a dopamine-releasing stimulus.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
hedgedconditional- Asserted: passes — The authors put it forward as true.
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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
“According to the neoHebbian model, the default condition is that information is stored automatically in the hippocampus by early LTP based on a local Hebbian process.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
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- Probably not a substantial claim. 16
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In the paper
“However, it is now clear that the Hebb rule does not account for late LTP; this requires an additional signal that is non-local.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Abstract
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In the paper
“The opening of the NMDAR is Hebbian: the channel opens only if there is both presynaptic glutamate release and strong postsynaptic depolarization.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
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In the paper
“In primates, novel stimuli can trigger burst firing of dopamine cells which habituates as stimuli become familiar [44].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Novelty
What the sources say
- Not checked yetcites [44] Ljungberg T 1992 Responses of monkey dopamine neurons during learning of behavioral reactions J Neurophysiol 67 145 163 1552316 source
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In the paper
“Furthermore, preventing strong depolarization by injecting negative current prevents the induction of LTP [4].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction
What the sources say
- Not checked yetcites [4] Malinow R Miller JP 1986 Postsynaptic hyperpolarization during conditioning reversibly blocks induction of long-term potentiation Nature 320 529 530 3008000
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Whereas the Hebbian NMDAR-dependent processes in the slice occur only during the initial induction of LTP, events in vivo also involve NMDAR-dependent processes triggered by hippocampal replay processes [142].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > How does the neoHebbian framework interact with off-line replay?
What the sources say
- Not checked yetcites [142] Shimizu E 2000 NMDA receptor-dependent synaptic reinforcement as a crucial process for memory consolidation Science (New York, N Y 290 1170 1174 11073458
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In the paper
“the observation that novelty-related activation of the SN/VTA has a long (ca. 30 minutes) penumbra that affects memory for unrelated information (e.g. exposure to novel scenes can affect memory for words) [74]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Is there a tension between the penumbra and reward-related memory enhancement by dopamine?
What the sources say
- Not checked yetcites [74] Fenker DB 2008 Novel scenes improve recollection and recall of words J Cogn Neurosci 20 1250 1265 18284351
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Based on these results, one would predict that dopamine antagonists would produce a weak reduction of memory at short times after learning but would produce a strong reduction at later times.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
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In the paper
“Early experiments revealed that dopamine antagonists applied during the LTP induced by multiple tetani (generally 100Hz for 1s) had an effect that was strongly time dependent [7]. Whereas the first phase of LTP was not strongly affected, the potentiation measured an hour after induction was strongly attenuated.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1
What the sources say
- Not checked yetcites [7] Frey U 1990 Dopaminergic antagonists prevent long-term maintenance of posttetanic LTP in the CA1 region of rat hippocampal slices Brain research 522 69 75 1977494
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In the paper
“Another quandary was posed by early work showing that dopamine agonists alone could produce potentiation [21].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
What the sources say
- Not checked yetcites [21] Huang YY Kandel ER 1995 D1/D5 receptor agonists induce a protein synthesis-dependent late potentiation in the CA1 region of the hippocampus Proceedings of the National Academy of Sciences of the United States of America 92 2446 2450 7708662 PMC42234
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In the paper
“Results such as those shown in Fig. 1 (top traces) suggest that early LTP is completely unaffected by dopamine antagonist.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
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In the paper
“Late LTP is blocked by D1 antagonists [14] (Fig. 1A) and by genetic deletion of D1 receptors (Fig. 1C) [15, 16].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1
What the sources say
- Not checked yetcites [14] O’Carroll CM Morris RG 2004 Heterosynaptic co-activation of glutamatergic and dopaminergic afferents is required to induce persistent long-term potentiation Neuropharmacology 47 324 332 15275821
- Not checked yetcites [15] Matthies H 1997 Dopamine D1-deficient mutant mice do not express the late phase of hippocampal long-term potentiation Neuroreport 8 3533 3535 9427321
- Not checked yetcites [16] Granado N 2008 D1 but not D5 dopamine receptors are critical for LTP, spatial learning, and LTP-Induced arc and zif268 expression in the hippocampus Cereb Cortex 18 1 12 17395606
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Thus, although 89% of VTA dopaminergic neurons respond with activation to a conditioned reward, they also respond to fearful events, with the majority of the neurons showing suppression. Interestingly, 25% respond with excitation to both positive and fearful events [101].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Aversive events
What the sources say
- Not checked yetcites [101] Wang DV Tsien JZ 2011 Convergent processing of both positive and negative motivational signals by the VTA dopamine neuronal populations PloS one 6 e17047 21347237 PMC3039659 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“LTP at these later times (termed late LTP) is dependent on protein synthesis, whereas the LTP at early times is not [13].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1
What the sources say
- Not checked yetcites [13] Frey U 1993 Effects of cAMP simulate a late stage of LTP in hippocampal CA1 neurons Science (New York, N Y 260 1661 1664 8389057 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“It is now well-established that dopamine neurons can also fire in response to unexpected aversive outcomes and cues predicting punishments [45].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Aversive events
What the sources say
- Not checked yetcites [45] Bromberg-Martin ES 2010 Dopamine in motivational control: rewarding, aversive, and alerting Neuron 68 815 834 21144997 PMC3032992 source
boosteddirect- Asserted: passes — The authors put it forward as true.
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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
“Thus, both early and late LTP are dopamine-dependent, but late LTP has a stronger dependence.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Furthermore, depletion of dopamine selectively blocks late LTP [18], a block that can be reversed by a D1 agonist (Fig. 1D).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1
What the sources say
- Not checked yetcites [18] Yang HW 2002 Change in bi-directional plasticity at CA1 synapses in hippocampal slices taken from 6-hydroxydopamine-treated rats: the role of endogenous norepinephrine The European journal of neuroscience 16 1117 1128 12383241
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“This one-trial memory for novel associations is critically dependent on the hippocampus and on glutamatergic neurotransmission during encoding [25, 26].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > D1 antagonists block 1 day old memory without affecting early memory
What the sources say
- Not checked yetcites [25] Tse D 2007 Schemas and memory consolidation Science (New York, N Y 316 76 82 17412951 source
- Not checked yetcites [26] Bethus I 2010 Dopamine and memory: modulation of the persistence of memory for novel hippocampal NMDA receptor-dependent paired associates J Neurosci 30 1610 1618 20130171 PMC6633999 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“In reinforcement learning, dopamine responses conform with a signed prediction error; they increase for better than expected (reward) outcomes and decrease for worse than expected (aversive) outcomes [43].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Reward prediction errors
What the sources say
- Not checked yetcites [43] Schultz W 2007 Multiple dopamine functions at different time courses Annual review of neuroscience 30 259 288 17600522 source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Reward-related SN/VTA activation improves memory for the rewarded stimulus, but not for the non-rewarded stimuli given in close temporal proximity [57]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Is there a tension between the penumbra and reward-related memory enhancement by dopamine?
What the sources say
- Not checked yetcites [57] Wittmann B 2011 Behavioural specifications of reward-associated long-term memory enhancement in humans Learning and Memory 18 296 300 21502336 PMC3465832
plaindirect- Asserted: passes — The authors put it forward as true.
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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
“Thus, it is now clear that dopamine by itself does not produce potentiation; rather, it acts as a permissive signal for the late-phase LTP induced by a NMDAR-dependent process.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
boosteddirect- Asserted: passes — The authors put it forward as true.
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In the paper
“In monkeys, a substantial fraction of dopamine neurons in the dorsolateral SN/VTA are responsive to conditioned cues predicting aversive events; in contrast, those in the more ventromedial portions of the SN/VTA are inhibited by aversive stimuli [97]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Introduction > Aversive events
What the sources say
- Not checked yetcites [97] Matsumoto M Hikosaka O 2009 Two types of dopamine neuron distinctly convey positive and negative motivational signals Nature 459 837 841 19448610 PMC2739096 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“However, it was subsequently shown that this potentiation depended on the action of test pulses given during agonist application and that activation of NMDARs by the test pulses was required [22].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
What the sources say
- Not checked yetcites [22] Navakkode S 2007 Synergistic requirements for the induction of dopaminergic D1/D5-receptor-mediated LTP in hippocampal slices of rat CA1 in vitro Neuropharmacology 52 1547 1554 17433377
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“In vivo recordings show that CA3/CA1 synapses are strengthened over time during trace eyeblink conditioning but that this strengthening is greatly reduced either in D1 receptor knockout mice or after reducing D1 receptors by virally delivered siRNA [16].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Neohebbian synaptic plasticity: role of dopamine in late LTP in CA1 > Where is dopamine released in the hippocampus?
What the sources say
- Not checked yetcites [16] Granado N 2008 D1 but not D5 dopamine receptors are critical for LTP, spatial learning, and LTP-Induced arc and zif268 expression in the hippocampus Cereb Cortex 18 1 12 17395606
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Intrahippocampal infusion of the D1/D5 dopaminergic antagonist SCH23390 just before encoding causes impaired memory for the new reward-location association measured 24 hours later, but has no effect on early memory measured at 30 minutes [26]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > D1 antagonists block 1 day old memory without affecting early memory
What the sources say
- Not checked yetcites [26] Bethus I 2010 Dopamine and memory: modulation of the persistence of memory for novel hippocampal NMDA receptor-dependent paired associates J Neurosci 30 1610 1618 20130171 PMC6633999 source
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Although the SN and VTA are known as dopaminergic regions, some cells in these regions are GABAergic or glutamatergic, and others release both glutamate and dopamine [31, 32].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [31] Nair-Roberts RG 2008 Stereological estimates of dopaminergic, GABAergic and glutamatergic neurons in the ventral tegmental area, substantia nigra and retrorubral field in the rat Neuroscience 152 1024 1031 18355970 PMC2575227
- Not checked yetcites [32] Hnasko TS 2010 Vesicular glutamate transport promotes dopamine storage and glutamate corelease in vivo Neuron 65 643 656 20223200 PMC2846457 source
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Studies performed both in vivo and in vitro show that the tonic mode is driven by intrinsic pacemaker conductances [34, 36].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [34] Grace AA Bunney BS 1983 Intracellular and extracellular electrophysiology of nigral dopaminergic neurons--1. Identification and characterization Neuroscience 10 301 315 6633863
- Not checked yetcites [36] Grace AA Onn SP 1989 Morphology and electrophysiological properties of immunocytochemically identified rat dopamine neurons recorded in vitro J Neurosci 9 3463 3481 2795134 PMC6569889
plaindirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“Activation of the ventral subiculum (an output structure of the hippocampus) excites the nucleus accumbens (Nac), which, in turn, inhibits the ventral pallidum. This reduces the inhibition of dopamine cells and thereby increases the number of tonically firing cells [38, 39].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [38] Floresco SB 2001 Glutamatergic afferents from the hippocampus to the nucleus accumbens regulate activity of ventral tegmental area dopamine neurons Journal of Neuroscience 21 4915 4922 11425919 PMC6762358
- Not checked yetcites [39] Floresco SB 2003 Afferent modulation of dopamine neuron firing differentially regulates tonic and phasic dopamine transmission Nature neuroscience 6 968 973 12897785
plaindirect- Asserted: passes — The authors put it forward as true.
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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 results indicate that burst firing controls some downstream functions, but others may depend on how many cells are in the tonic mode (see Boxes 3 and 4).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
hedgedconditional- Asserted: passes — The authors put it forward as true.
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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
“Thus, novelty, reward stimuli and aversive stimuli are all able to activate the dopamine system in both monkeys and humans.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“The perirhinal cortex [65], for instance, shows much stronger responses to novel items than familiar items and may form connects to the PPTg (Figure I), a structure that regulates the SN/VTA.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [65] Xiang JZ Brown MW 1998 Differential neuronal encoding of novelty, familiarity and recency in regions of the anterior temporal lobe Neuropharmacology 37 657 676 9705004
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“Classical work showed the importance of reward as a stimulus for the bursting of these cells [43]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
What the sources say
- Not checked yetcites [43] Schultz W 2007 Multiple dopamine functions at different time courses Annual review of neuroscience 30 259 288 17600522 source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“selective genetic inactivation of NMDARs in dopaminergic cells blocked burst production and prevented conditioned behavioral responses that were dependent on dopamine [41]”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [41] Zweifel LS 2009 Disruption of NMDAR-dependent burst firing by dopamine neurons provides selective assessment of phasic dopamine-dependent behavior Proceedings of the National Academy of Sciences of the United States of America 106 7281 7288 19342487 PMC2678650 source
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Other work using fMRI has demonstrated that activation can also occur due to aversive events [53] as well as novel stimuli [54].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
What the sources say
- Not checked yetcites [53] Guitart-Masip M 2011 Action dominates valence in anticipatory representations in the human striatum and dopaminergic midbrain Journal of Neuroscience 31 7867 7875 21613500 PMC3109549
- Not checked yetcites [54] Duzel E 2010 NOvelty-related motivation of anticipation and exploration by dopamine (NOMAD): implications for healthy aging Neuroscience and biobehavioral reviews 34 660 669 19715723 source
boosteddirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Midbrain dopamine neurons consist of three largely contiguous cell groups: the retrorubral field (cell group A8 in the rat nomenclature), the substantia nigra pars compacta (SNc, A9), and the ventral tegmental area (VTA, A10) [28, 29].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [28] Dahlström A Fuxe K 1964 Evidence for the existence of monoamine containing neurons in the central nervous system. I. Demonstration of monoamines in the cell bodies of brain stem neurons Acta Physiol Scand 62 1 80 14229500
- Not checked yetcites [29] Beckstead RM 1979 Efferent connections of the substantia nigra and ventral tegmental area in the rat Brain research 175 191 217 314832 source
plain- Asserted: passes — The authors put it forward as true.
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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
“Furthermore, this improvement should be small with short delays when early LTP is present, but large at much longer delays when late LTP is present.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“More recent work has demonstrated conditions under which dopamine cells can be driven by aversive stimuli (reviewed in [45]; see also Box 2).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
What the sources say
- Not checked yetcites [45] Bromberg-Martin ES 2010 Dopamine in motivational control: rewarding, aversive, and alerting Neuron 68 815 834 21144997 PMC3032992 source
boostedconditional- 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
“Dopamine has no effect on the previously formed memories and is not required at the time of retrieval [27].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > D1 antagonists block 1 day old memory without affecting early memory
What the sources say
- Not checked yetcites [27] O’Carroll CM 2006 Dopaminergic modulation of the persistence of one-trial hippocampus-dependent memory Learning & memory (Cold Spring Harbor, N Y 13 760 769 PMC1783630 17142305
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 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 data demonstrate that D1/D5 receptor-mediated effects do not strongly affect encoding or early memory, but do affect memory at later times.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > D1 antagonists block 1 day old memory without affecting early memory
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.
It may state two separate claims as one.
In the paper
“However, in the absence of an expected reward, or if a worse than expected outcome occurs (i.e., a negative prediction error), the consequent depression of dopaminergic neuron activity would attenuate dopamine-mediated potentiation of the ventral subiculum while at the same time removing inhibition of the mPFC input. Consequently the mPFC would drive the animal to switch response strategies [122].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [122] Goto Y Grace AA 2008 Limbic and cortical information processing in the nucleus accumbens Trends in neurosciences 31 552 558 18786735 PMC2884964
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.
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 a condition in which the animal’s behavior leads to a better than expected outcome and phasic release (i.e. positive prediction error), the subicular input would be potentiated, causing the animal to maintain focus on the current task, thus allowing rewarded behaviors to be maintained.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
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
“while subsequent studies have demonstrated that a novel stimulus (in the absence of reward) can also produce burst firing [44].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Dopamine release by reward, novelty and aversive stimuli
What the sources say
- Not checked yetcites [44] Ljungberg T 1992 Responses of monkey dopamine neurons during learning of behavioral reactions J Neurophysiol 67 145 163 1552316 source
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.
In the paper
“The lateral habenula provides SN/VTA dopamine neurons with a negative reward-prediction signal through the fasciculus retroflexus fiber bundles [[119]].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Firing properties of dopaminergic neurons
What the sources say
- Not checked yetcites [119] Matsumoto M Hikosaka O 2007 Lateral habenula as a source of negative reward signals in dopamine neurons Nature 447 1111 1115 17522629 source
plaindirect- 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
“Frey and Morris propose that this specificity occurs because LTP immediately produces a synapse-specific “tag” and that PRPs can only produce late LTP at tagged synapses.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
What the sources say
- Not checked yetcites [9] Frey U Morris RG 1997 Synaptic tagging and long-term potentiation [see comments] Nature 385 533 536 9020359
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.
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 hippocampus, on the other hand, integrates information about stimuli and their spatial location into a contextual representation and can compute various types of novel changes in these relationships.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
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.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Indeed, fMRI studies (for examples see Box 2 Figure 1) confirm SN/VTA activation by associative novelty [60] and stimulus novelty [61] but less by other salient events such as rareness or negative emotional content [61].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [60] Schott BH 2004 Activation of midbrain structures by associative novelty and the formation of explicit memory in humans Learning & memory (Cold Spring Harbor, N Y 11 383 387 15254215
- Not checked yetcites [61] Bunzeck N Duzel E 2006 Absolute stimulus-novelty is coded by the human substantia nigra/VTA Neuron 3 369 379 16880131 source
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
“positron emission tomography (PET) studies in humans show that novelty-seeking traits are inversely associated with D2-like receptor availability in the SN/VTA region [62] and are positively correlated with white matter connectivity between the hippocampus and the ventral striatum [63].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [62] Zald DH 2008 Midbrain dopamine receptor availability is inversely associated with novelty-seeking traits in humans J Neurosci 28 14372 14378 19118170 PMC2748420
- Not checked yetcites [63] Cohen MX 2009 Connectivity-based segregation of the human striatum predicts personality characteristics Nature neuroscience 12 32 34 19029888
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.
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
“From the perspective of the neoHebbian framework, this negative result is understandable; i.e., the SN/VTA activity-related enhancement of memory would be more apparent if testing is conducted after long retention intervals.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Effect of Motivation (Reward) on Behavioral Memory
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.
In the paper
“In the perirhinal cortex, for instance, stimulus-related incoming information can be compared to stored information and a stimulus-related novelty signal is computed [65].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [65] Xiang JZ Brown MW 1998 Differential neuronal encoding of novelty, familiarity and recency in regions of the anterior temporal lobe Neuropharmacology 37 657 676 9705004
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
“The human SN/VTA is also activated by cues that predict novel images, or by unexpected novel images that followed familiarity-predictive cues (ie. an ‘unexpected novelty’ response) [64].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Importance of the novelty and aversive signals for memory
What the sources say
- Not checked yetcites [64] Wittmann BC 2007 Anticipation of novelty recruits reward system and hippocampus while promoting recollection Neuroimage 38 194 202 17764976 PMC2706325
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.
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
“Moreover, other properties of LTP (weak/strong; penumbra) appear also to have predictive power in explaining properties of memory.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
hedgedcorrelational- 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: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
In the paper
“In schizophrenia, the enhanced activation of the hippocampal system may increase dopamine release and skew behavioral choice towards repetitive behavior (see Box 3).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites Box 3
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.
In the paper
“That this drug-induced memory impairment is not due to an effect of dopamine on encoding is nicely demonstrated by the rescue of long-term memory if there is a preceding exposure to a novel environment [72].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [72] Wang SH 2010 Relevance of synaptic tagging and capture to the persistence of long-term potentiation and everyday spatial memory Proceedings of the National Academy of Sciences of the United States of America 107 19537 19542 20962282 PMC2984182
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.
It may state two separate claims as one.
In the paper
“Parkinson’s disease (PD) usually begins with loss of nigrostriatal dopamine cells in the ventral tier SNc, leaving the dorsal tier projection neurons to the hippocampus, ventral striatum, and cortex intact [79].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [79] Fearnley JM Lees AJ 1991 Ageing and Parkinson’s disease: substantia nigra regional selectivity Brain 114 Pt 5 2283 2301 1933245 source
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
“long-term retention is not helped by simple re-exposure to recently learned material but is greatly helped by retesting even when subjects already know the answer [83].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
What the sources say
- Not checked yetcites [83] Karpicke JD Roediger HL 3rd 2008 The critical importance of retrieval for learning Science (New York, N Y 319 966 968 18276894 source
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
“a single exposure to the location of a small reward is rapidly forgotten in rodents within a few hours, but persists for at least 24 hours if it is preceded or followed (within 30 minutes) by novelty exploration [72].”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Testing the synaptic tag and capture model at the behavioral level
What the sources say
- Not checked yetcites [72] Wang SH 2010 Relevance of synaptic tagging and capture to the persistence of long-term potentiation and everyday spatial memory Proceedings of the National Academy of Sciences of the United States of America 107 19537 19542 20962282 PMC2984182
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
“It appears that both the tag and the PRPs have considerable lifetimes.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
hedged- 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
“stimulation of protein synthesis and the induction of Hebbian plasticity do not need to be coincident; indeed, the trigger for protein synthesis can occur either before or after Hebbian plasticity (with intervening periods of many minutes).”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > The synaptic tag and capture model: role of plasticity-related proteins
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
“A key insight to emerge from physiological experiments is that LTP at short times is relatively insensitive to dopamine, whereas LTP with longer delays is sensitive.”
§ A neoHebbian framework for episodic memory; role of dopamine-dependent late LTP > Concluding Comments
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.
The paper relies on this without support.
There is relatively little work comparing the consolidation of rewarding and aversive events in humans.
“Surprisingly, there is relatively little work comparing the consolidation of rewarding and aversive events (e.g. losses) in humans (although there is extensive work on the consolidation of emotionally negative stimuli).”
The paper relies on this without support.
The conditions that lead to activation of the human dopamine system, as judged by fMRI signals, are similar to the conditions leading to activation in non-primate mammals.
“The conditions that lead to activation of the human dopamine system, as judged by fMRI signals, are similar to the conditions leading to activation in non-primate mammals.”
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?
In the citing paper
“The results support the hypothesis that D1/D5 neurotransmission is important for tagging synapses in the CA3-CA1 circuit for L-LTP (Frey and Morris, 1997; Lisman et al., 2011).”
From Dopamine regulates aversive contextual learning and associated in vivo synaptic plasticity in the Hippocampus (2016) · cites this paper as Lisman et al., 2011
Citations
Cites
The Organization of Behavior; A Neuropsychological Theory.
Attneave, F., B., M., & Hebb, D. O. (1950) · The American Journal of Psychology · doi:10.2307/1418888 · Cited by 5,146
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Tulving, E. (1985) · Canadian Psychology / Psychologie canadienne · doi:10.1037/h0080017 · Cited by 4,638
Aston-Jones, G., & Cohen, J. D. (2005) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.28.061604.135709 · Cited by 4,472
The Organization of Behavior: A Neuropsychological Theory
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Depth of Processing and the Retention of Words in Episodic Memory
Craik, F. I. M., & Tulving, E. (1975) · Journal of Experimental Psychology: General · doi:10.1037/0096-3445.104.3.268 · Cited by 4,121
The reward circuit: linking primate anatomy and human imaging
Haber, S. N., & Knutson, B. (2010) · Neuropsychopharmacology · doi:10.1038/npp.2009.129 · Cited by 3,718
AGEING AND PARKINSON'S DISEASE: SUBSTANTIA NIGRA REGIONAL SELECTIVITY
FEARNLEY, J. M., & LEES, A. J. (1991) · Brain · doi:10.1093/brain/114.5.2283 · Cited by 3,529
Are the Dorsal and Ventral Hippocampus Functionally Distinct Structures?
Fanselow, M. S., & Dong, H.-W. (2010) · Neuron · doi:10.1016/j.neuron.2009.11.031 · Cited by 3,358
Structural basis of long-term potentiation in single dendritic spines
Matsuzaki, M., Honkura, N., Ellis-Davies, G. C. R., & Kasai, H. (2004) · Nature · doi:10.1038/nature02617 · Cited by 2,483
Research works printed but not linked: 1
- Watson, J.S., trans. and ed. (1986) Cicero on Oratory and Orators (Landmarks in Rhetoric and Public Address), Southern Illinois University Press
Cited by
Sorted by citations
How schema and novelty augment memory formation
van Kesteren, M. T. R., Ruiter, D. J., Fernández, G., & Henson, R. N. (2012) · Trends in Neurosciences · doi:10.1016/j.tins.2012.02.001 · Cited by 860
States of Curiosity Modulate Hippocampus-Dependent Learning via the Dopaminergic Circuit
Gruber, M. J., Gelman, B. D., & Ranganath, C. (2014) · Neuron · doi:10.1016/j.neuron.2014.08.060 · Cited by 812
“…to memory formation during curious states strongly parallel theoretical accounts and findings in rodents showing that dopaminergic activity can modulate hippocampus-dependent learning (for reviews, see D üzel et al., 2010; Lisman and Grace, 2005; Lisman et al., 2011; Shohamy and Adcock, 2010).”
InfluentialWhat Learning Systems do Intelligent Agents Need? Complementary Learning Systems Theory Updated
Kumaran, D., Hassabis, D., & McClelland, J. L. (2016) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2016.05.004 · Cited by 759
The synaptic plasticity and memory hypothesis: encoding, storage and persistence
Takeuchi, T., Duszkiewicz, A. J., & Morris, R. (2013) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2013.0288 · Cited by 712
Lachaux, J.-P., Axmacher, N., Mormann, F., Halgren, E., & Crone, N. E. (2012) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2012.06.008 · Cited by 535
“the effect of dopamine release on hippocampal activity (Lisman and Grace, 2005; Axmacher et al., 2010b; Lisman et al., 2011), but it also raises very complex safety issues that should be resolved before use.”
Background
Metadata
- Authors
- Lisman, J., Grace, A. A., & Duzel, E.
- Affiliations
- Brandeis University (US)
- University of Pittsburgh (US)
- Corresponding author
- Lisman, John
- Venue
- Trends in Neurosciences
- Publisher
- Elsevier BV
- Published
- 2011-10
- Volume
- 34
- Issue
- 10
- Pages
- 536-547
- ISSN
- 0166-2236 (ISSN-L)
- PMC
- PMC3183413
- OpenAlex
- W2060322330
- Semantic Scholar
- 2d3befef55d10b6b1538c81c9214e8c2a0fd75fc
- Cited by
- 465
- References
- 146
- Citations by year
- 2026: 27 · 2025: 25 · 2024: 39 · 2023: 29 · 2022: 36 · 2021: 31 · 2020: 31 · 2019: 33 · 2018: 36 · 2017: 30 · 2016: 41 · 2015: 32 · 2014: 22 · 2013: 29 · 2012: 22
- FWCI
- 1.96
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Trends in Neurosciences (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- UCL Discovery (University College London) (submittedVersion)
- DZNE Pub (submittedVersion, other-oa)
- Funders
- National Institutes of Health (2r01mh057440-26a1)
- National Institute of Mental Health (MH-57440, R01-MH057440, R37MH057440)
- National Institute on Drug Abuse (DA 15408, R01DA027807)
- National Institute of Neurological Disorders and Stroke (R01NS027337)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Keywords
- episodic memory, long-term potentiation, dopamine, hippocampal CA1, Hebbian rule, synaptic plasticity, reward, memory formation
- MeSH terms
- Animals, Dopamine, Neuronal Plasticity, Synapses, Long-Term Potentiation, Memory, Episodic
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 2026-10-10, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
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