The DOI matches its Crossref record; 33 of the 33 references it cites were found in a registry or an index; none of the 1 sources its claims cite is checked for support yet.
Distinct extended amygdala circuits for divergent motivational states
Open accessAbstract
The co-morbidity of anxiety and dysfunctional reward processing in illnesses such as addiction and depression suggests that common neural circuitry contributes to these disparate neuropsychiatric symptoms. The extended amygdala, including the bed nucleus of the stria terminalis (BNST), modulates fear and anxiety, but also projects to the ventral tegmental area (VTA), a region implicated in reward and aversion, thus providing a candidate neural substrate for integrating diverse emotional states. However, the precise functional connectivity between distinct BNST projection neurons and their postsynaptic targets in the VTA, as well as the role of this circuit in controlling motivational states, have not been described. Here we record and manipulate the activity of genetically and neurochemically identified VTA-projecting BNST neurons in freely behaving mice. Collectively, aversive stimuli exposure produced heterogeneous firing patterns in VTA-projecting BNST neurons. By contrast, in vivo optically identified glutamatergic projection neurons displayed a net enhancement of activity to aversive stimuli, whereas the firing rate of identified GABAergic (γ-aminobutyric acid-containing) projection neurons was suppressed. Channelrhodopsin-2-assisted circuit mapping revealed that both BNST glutamatergic and GABAergic projections preferentially innervate postsynaptic non-dopaminergic VTA neurons, thus providing a mechanistic framework for in vivo circuit perturbations. In vivo photostimulation of BNST glutamatergic projections resulted in aversive and anxiogenic behavioural phenotypes. Conversely, activation of BNST GABAergic projections produced rewarding and anxiolytic phenotypes, which were also recapitulated by direct inhibition of VTA GABAergic neurons. These data demonstrate that functionally opposing BNST to VTA circuits regulate rewarding and aversive motivational states, and may serve as a crucial circuit node for bidirectionally normalizing maladaptive behaviours.
Overview
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Written from the paper’s full text.
TL;DR
In mice, VTA-projecting BNST neurons comprise opposing glutamatergic and GABAergic subcircuits: aversive stimuli increase glutamatergic activity and suppress GABAergic activity, and optogenetic activation of these pathways produces aversive/anxiogenic versus rewarding/anxiolytic states, respectively. These findings identify a bidirectional BNST→VTA circuit node for motivational and emotional processing.
Key findings
- In vivo antidromic identification found 53/137 BNSTv→VTA projection neurons; aversive stimuli produced heterogeneous firing patterns across these neurons. Identified Vglut2BNSTv→VTA neurons (34/145) showed net enhanced firing during aversive events and shock-associated cues, whereas VgatBNSTv→VTA neurons (33/77) were principally suppressed.
- Circuit mapping showed that Vglut2BNSTv→VTA and VgatBNSTv→VTA terminals formed functional synapses primarily onto non-dopaminergic and medially located dopaminergic VTA neurons.
- Photostimulation of Vglut2BNSTv→VTA produced avoidance of a stimulation-paired chamber, reduced active reward seeking, and anxiety-like open-field behavior; the aversive phenotype was abolished by intra-VTA ionotropic glutamate receptor antagonists.
- Photostimulation of VgatBNSTv→VTA produced place preference, supported optical self-stimulation, and increased open-arm time in the elevated plus maze; these effects were prevented by an intra-VTA GABAA antagonist and recapitulated by direct inhibition of VTA GABAergic neurons. Concurrent VgatBNSTv→VTA stimulation during foot shock reduced freezing and increased later open-arm time/entries.
Methods
- Adult male C57BL/6J, Vgat-ires-cre, and Vglut2-ires-cre mice received stereotactic viral injections (ChR2-eYFP, NpHR) and optical fibers/MEA implants; experiments included anesthetized and behaving in vivo electrophysiology, patch-clamp recordings in VTA slices, unpredictable foot-shock sessions, and optogenetic behavioral assays.
- In vivo electrophysiology used a 16-channel MEA in BNSTv and optical fibers above VTA for antidromic identification; units were classified using spike latency, waveform correlation, and z-score firing changes during foot-shock sessions.
- Behavioral assays included real-time place preference, open field, sucrose self-administration, optical self-stimulation/self-inhibition, elevated plus maze, and foot-shock-induced freezing, with intra-VTA antagonist microinfusions in some experiments.
Limitations
- The paper states that shock artifacts prevented reliable quantification of neuronal firing 500–1000 ms after foot-shock onset, excluding approximately 0.8–1.66% of collected data during the foot-shock session.
- Evident from the text: all experiments used adult male mice, so sex differences and generalization to females are not addressed.
- Evident from the text: the paper does not include a dedicated limitations section; circuit specificity relies on viral targeting, optogenetic manipulation, slice physiology, and pharmacological blockade rather than fully selective circuit-silencing approaches.
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21 Own method · 15 Own result · 8 Own interpretation · 1 Not checked yet
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“in vivo optically-identified glutamatergic projection neurons displayed a net enhancement of activity to aversive stimuli”
- Also stated in: § Abstract · plain
“Collectively, exposure to the aversive event or associated cues alone enhanced the firing of Vglut2BNSTv→VTA neurons”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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- Also stated in: § Abstract · plain
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Collectively, aversive stimuli exposure produced heterogeneous firing patterns in VTA-projecting BNST neurons.”
- Also stated in: § Abstract · boosted
“Identified BNSTv→VTA neurons segregated into three functionally distinct classes based on changes in their normalized firing rates throughout the foot-shock session (Supplementary Fig. 3), demonstrating that BNSTv→VTA neurons differentially encode information related to aversive stimuli and their associated contextual cues.”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
boostedcorrelational- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“whereas the firing rate of identified GABAergic projection neurons was suppressed”
- Also stated in: § Abstract · plain
“while simultaneously suppressing the activity of VgatBNSTv→VTA neurons.”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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- Also stated in: § Abstract · plain
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“activation of BNST GABAergic projections produced rewarding and anxiolytic phenotypes”
- Also stated in: § Abstract · hedged
“Photostimulation of the VgatBNSTv→VTA pathway and direct inhibition of VgatVTA neurons significantly increased time spent in the open arms of an elevated plus maze, indicative of anxiolysis”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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- Also stated in: § Abstract · hedged
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“VgatBNSTv→VTA::ChR2 mice spent significantly less time frozen (Fig. 5h)”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“as well as significantly more open-arm time and entries in the elevated-plus maze relative to controls (Supplementary Fig. 18)”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plain- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In vivo photostimulation of BNST glutamatergic projections resulted in aversive and anxiogenic behavioral phenotypes.”
- Also stated in: § Abstract · hedged
“inescapable activation of this pathway for 20 min in an open field resulted in significantly less center- and more corner-time in Vglut2BNSTv→VTA::ChR2 mice in the 10 min following stimulation offset compared to controls”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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- Also stated in: § Abstract · hedged
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“VTA infusions of a GABAA receptor antagonist prevented the VgatBNSTv→VTA mediated place preference compared to saline injections”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Vglut2BNSTv→VTA and VgatBNSTv→VTA terminals formed functional synapses primarily onto non-dopaminergic and medially located dopaminergic neurons”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaincorrelational- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“activation of BNST GABAergic projections produced rewarding and anxiolytic phenotypes”
- Also stated in: § Abstract · plain
“20 Hz photostimulation in VgatBNSTv→VTA::ChR2 mice resulted in a significant place preference”
- Also stated in: § Abstract · plain
“VgatBNSTv→VTA::ChR2 mice readily nose poked to receive photostimulation”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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- It may state two separate claims as one. 9
- Probably not a substantial claim. 19
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“photostimulation of the CaMKIIaBNSTv→VTA pathway produced both glutamatergic and γ-aminobutyric acid (GABAergic) currents in VTA neurons (Supplementary Fig. 4), demonstrating that neurochemically distinct BNSTv neurons project to the VTA.”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
boosteddirect- Asserted: passes — The authors put it forward as true.
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In the paper
“photostimulation of ChR2-containing fibers originating from Vglut2 (Vglut2BNSTv→VTA::ChR2) or Vgat (VgatBNSTv→VTA::ChR2) expressing BNSTv neurons produced excitatory or inhibitory postsynaptic currents respectively”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Thus, data collected during these time epochs were excluded from analysis.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“into the VTA in a counter balanced design (all drugs from Tocris).”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Intra-VTA injection of antagonists and photostimulation of Vglut2BNSTv→VTA::ChR2 and VgatBNSTv→VTA::ChR2 projections during real-time place preference
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“Together, these results show that reward-related responses to VgatBNSTv→VTA activation are recapitulated by directly inhibiting VgatVTA neurons, thus providing a circuit mechanism for the VgatBNSTv→VTA pathway to regulate motivated behavior.”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
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In the paper
“Briefly, discrimination of individual units was performed offline using principal component analysis to separate individual units from the same electrode. In addition, auto- and cross-correlograms, firing characteristics, and inter-spike interval distributions were examined to ensure units were well-isolated.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“Placements of recording electrode tips within the BNSTv and optical fibers within the VTA were verified with histological examination of brain tissue following the experiments.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo anesthetized electrophysiology
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“This represents an approximate loss of 0.8 – 1.66% of the collected data during the foot-shock session.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
hedged- Asserted: passes — The authors put it forward as true.
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“During the time epoch surrounding the individual delivered foot shocks, large shock artifact were readily apparent.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“Resulting data was binned in 30 s epochs in order to minimize the skewing of our results due to data lost by the shock artifacts.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Electrical stimulation of the BNST produces both excitatory and inhibitory responses in VTA neurons in vivo6 implying that distinct subcircuits may exist.”
§ Distinct extended amygdala circuits for divergent motivational states > Abstract
What the sources say
- Not checked yetcites [6] Georges, F. & Aston-Jones, G. Potent regulation of midbrain dopamine neurons by the bed nucleus of the stria terminalis. J. Neurosci. 21, RC160 (2001) source
hedgedconditional- Asserted: passes — The authors put it forward as true.
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In the paper
“BNSTv neurons were classified as antidromic-responsive, if the following 3 criteria were met: 1) stable antidromic spike latency (< 0.2 ms), 2) ability to respond reliably to high frequency photostimulation, 3) collision between orthodromic- and antidromic-evoked spikes.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo anesthetized electrophysiology
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In the paper
“Neuronal units were included in the data if the signal-to-noise ratio was high, and the mean firing rate was between 0.5 – 25 Hz during baseline recording periods.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“Mice had 2 days off between each VTA microinjection.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Intra-VTA injection of antagonists and photostimulation of Vglut2BNSTv→VTA::ChR2 and VgatBNSTv→VTA::ChR2 projections during real-time place preference
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In the paper
“In addition, recording sites was verified histologically using electrolytic lesions at 200 µA for 5 s.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“neuronal firing responses in the 500 – 1000 ms following foot shock onset could not be reliably quantified”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“All mice were infused at a rate of 0.1 µl per minute. The injector remained in place for approximately 2 min following infusion to ensure proper diffusion of drug into the VTA.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Intra-VTA injection of antagonists and photostimulation of Vglut2BNSTv→VTA::ChR2 and VgatBNSTv→VTA::ChR2 projections during real-time place preference
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In the paper
“Recorded BNSTv units were classified as light-responsive, and thus VTA-projecting, if they met both of the following two criteria. 1) The latency of the first spike after light stimulation onset was less than 20 ms for ≥ 20% of the trials. 2) Light-evoked and spontaneous waveform shapes had a correlation coefficient of > 0.90.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“Neural activity was recorded from the same mice during the cue exposure, foot shock, and cue re-exposure sessions, therefore we were able to reliably record activity from the same population of neurons during each of the three sessions.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“In addition, timestamp data to signify the start and end of foot-shock sessions, and the delivery of light pulses to optical fibers was synchronized with electrophysiological data.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“Following the 1 day 20 Hz training session, mice were run daily at each photostimulation frequency (1, 5, 10, 20, 40 Hz) in a counter-balanced design.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Optical self-stimulation of VgatBNSTv→VTA::ChR2 projections
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“While these were easily isolated and excluded from the analyzed waveform data using offline spike sorting”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo electrophysiology
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In the paper
“The injector needle (33 gauge steel tube, McMasters-Carr) extended approximately 1 mm past the cannula to ensure drug delivery 0.5 mm below the optical fiber.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Intra-VTA injection of antagonists and photostimulation of Vglut2BNSTv→VTA::ChR2 and VgatBNSTv→VTA::ChR2 projections during real-time place preference
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In the paper
“Once stable nose-poking behavior for 15% sucrose was observed (approx. 100 active nose pokes on at least 2 consecutive days), all mice received constant 20 Hz photostimulation during the entire 30 min sucrose session.”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Photostimulation of Vglut2BNSTv→VTA::ChR2 projections during sucrose self-administration
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Claims other papers make about this one
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The cited source says what the claim attributes to it.
In the citing paper
“Our results provide further evidence of the importance of long-range GABAergic projection neurons in the control of anxiety (Jennings et al., 2013; Kim et al., 2013).”
From Control of Stress-Induced Persistent Anxiety by an Extra-Amygdala Septohypothalamic Circuit (2014) · cites this paper as Jennings et al., 2013
The cited source says what the claim attributes to it.
In the citing paper
“This is supported by optogenetic studies showing the projection-specific effects of BNST neurons on behavior (87–90).”
From Sexual differentiation of neural mechanisms of stress sensitivity during puberty (2023) · cites this paper as [89]
In the citing paper
“Moreover, individual subregions of the BNST have been shown to differentially regulate separable features of the anxiety phenotype31,85 (Fig. 3).”
From Resolving the neural circuits of anxiety (2015) · cites this paper as [85]
In the citing paper
“activity in glutamatergic ventral BNST neurons projecting to the VTA produces anxiety-like behaviors, whereas GABAergic neurons in the ventral BNST exert anxiolysis via a parallel pathway85.”
From Resolving the neural circuits of anxiety (2015) · cites this paper as [85]
In the citing paper
“It has recently been shown that the BNST contains distinct anxiogenic and anxiolytic GABAergic neuronal subsets with unique patterns of long-range connections (Jennings et al., 2013; Kim et al., 2013).”
From Control of Stress-Induced Persistent Anxiety by an Extra-Amygdala Septohypothalamic Circuit (2014) · cites this paper as Jennings et al., 2013
In the citing paper
“In contrast, photostimulation of glutamatergic projections from ventral BNST to the ventral tegmental area (VTA) (Jennings et al., 2013) was sufficient to induce a persistent elevation in anxiety.”
From Control of Stress-Induced Persistent Anxiety by an Extra-Amygdala Septohypothalamic Circuit (2014) · cites this paper as Jennings et al., 2013
In the citing paper
“Notably, BNST neurons have been shown to inhibit dopamine neurons involved in conventional reward processing by activating local GABAergic neurons63–65, suggesting that anxiety-related circuits may interfere with appetitive learning and behavior.”
From Dopamine dynamics as a regulatory mechanism for shifting between defensive and reward-seeking behaviors (2026) · cites this paper as [63]
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Each contradiction between two passages listed here was confirmed by an independent check.
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Experimental subjects and stereotactic surgery
“For all in vivo electrophysiology experiments, mice were implanted with a 16-wire (4 × 4 configuration, wire diameter∼30µm) tungsten multielectrode array (MEA)”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo anesthetized electrophysiology
“Extracellular neural activity was recorded using a glass recording electrode (5 − 10 MΩ: and filled with 0.5 M NaCl).”
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > Experimental subjects and stereotactic surgery
“For all in vivo electrophysiology experiments, male mice were unilaterally injected with 0.5 µl of purified and concentrated adeno-associated virus (AAV)”
§ Distinct extended amygdala circuits for divergent motivational states > Online methods > In vivo anesthetized electrophysiology
“C57BL/6J mice were bilaterally injected with 0.3 µl of AAV5-CaMKIIa-ChR2-eYFP into the BNSTv.”
Citations
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“Recorded vBNST units were classified as light-responsive, and thus VTA-projecting, if they met both of the following two criteria: (1) the latency of the first spike after light stimulation onset was less than 20ms for$20% of the trials; and (2) light-evoked and spontaneous waveform shapes had a correlation coefficien…”
BackgroundMethodThe mesolimbic dopamine reward circuit in depression
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“Recorded vBNST units were classified as light-responsive, and thus VTA-projecting, if they met both of the following two criteria: (1) the latency of the first spike after light stimulation onset was less than 20ms for$20% of the trials; and (2) light-evoked and spontaneous waveform shapes had a correlation coefficien…”
BackgroundMethodPhasic vs Sustained Fear in Rats and Humans: Role of the Extended Amygdala in Fear vs Anxiety
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Items 501–600 of 703
Neural Contributions of the Hypothalamus to Parental Behaviour
Orikasa, C. (2021) · International Journal of Molecular Sciences · doi:10.3390/ijms22136998 · Cited by 8
“In the amygdala neural circuits, GABAergic and glutamatergic neurons in the VTA specifically tune each rewarding and aversive motivational predicaments [79].”
BackgroundDo oxytocin neurones affect feeding?
Worth, A. A., & Luckman, S. M. (2021) · Journal of Neuroendocrinology · doi:10.1111/jne.13035 · Cited by 7
Kourosh‐Arami, M., Gholami, M., Alavi-Kakhki, S. S., & Komaki, A. (2022) · Neuropeptides · doi:10.1016/j.npep.2022.102259 · Cited by 8
Automated and parallelized spike collision tests to identify spike signal projections
Mitani, K., Kawabata, M., Isomura, Y., & Sakai, Y. (2022) · iScience · doi:10.1016/j.isci.2022.105071 · Cited by 7
Gain in Translation: Is It Time for Thigmotaxis Studies in Humans?
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Visceral hypersensitivity induced by optogenetic activation of the amygdala in conscious rats
Johnson, A. C., Latorre, R., Ligon, C. O., & Meerveld, B. G. (2018) · American Journal of Physiology-Gastrointestinal and Liver Physiology · doi:10.1152/ajpgi.00370.2017 · Cited by 7
“yellow laser at 589 nm (1), we used a more versatile green 532-nm laser that can activate both halorhodopsin and other optogenetic vectors (15), based on multiple studies in the literature (4, 20, 37).”
MethodHara, R., Takahashi, D., Takehara, T., Amano, T., & Minami, M. (2020) · Molecular Brain · doi:10.1186/s13041-020-00684-4 · Cited by 8
Domschke, K., Kapfhammer, H., & Deckert, J. (2017) · Springer eBooks · doi:10.1007/978-3-662-49295-6_69 · Cited by 9
Separable dorsal raphe dopamine projections mimic the facets of a loneliness-like statePreprint
Lee, C. R., Matthews, G. A., Lemieux, M., Wasserlein, E. M., Borio, M., Miranda, R. L., Keyes, L. R., Schneider, G. P., Jia, C., Tran, A., et al. (2025) · eLife · doi:10.7554/elife.105955 · Cited by 9
The regulative role and mechanism of BNST in anxiety disorder
Xie, M., Xiong, Y., & Wang, H. (2024) · Frontiers in Psychiatry · doi:10.3389/fpsyt.2024.1437476 · Cited by 7
Furman, K. L., Baron, L., Lyons, H. I., Cha, T., Evans, J. R., Manna, J., Zhu, L., Mattis, J., & Burgess, C. R. (2025) · Journal of Neuroscience · doi:10.1523/jneurosci.1725-24.2024 · Cited by 7
Luo, L., Jing, W., Guo, Y., Liu, D., He, A., & Lu, Y. (2025) · Nature Communications · doi:10.1038/s41467-025-58591-y · Cited by 8
Ly, A., Barker, A., Hotchkiss, H., Prévost, E. D., McGovern, D. J., Kilpatrick, Z. P., & Root, D. H. (2023) · European Journal of Neuroscience · doi:10.1111/ejn.16137 · Cited by 7
“Notably, different subregions of the BNST have opposing effects on anxiety-like behaviour that appear highly dependent on the projection site (Giardino et al., 2018; Jennings et al., 2013; Kim et al., 2013).”
Luyck, K., & Luyten, L. (2015) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2015.00069 · Cited by 7
“In addition, Jennings et al. demonstrated that photostimulation of the glutamatergic connections between the ventral BST and ventral tegmental area was anxiogenic, whereas activation of the GABAergic projections in the same area resulted in anxiolytic effects (Jennings et al., 2013).”
BackgroundHessel, M., Pape, H., & Seidenbecher, T. (2020) · European Neuropsychopharmacology · doi:10.1016/j.euroneuro.2020.08.006 · Cited by 7
Optimization of whole-brain rabies virus tracing technology for small cell populations
Roelofs, T. J. M., Menting-Henry, S., Gol, L. M., Speel, A. M., Wielenga, V. H., Garner, K. M., Luijendijk, M. C. M., Hennrich, A. A., Conzelmann, K., & Adan, R. A. H. (2021) · Scientific Reports · doi:10.1038/s41598-021-89862-5 · Cited by 7
Origins of Addiction Predictably Embedded in Childhood Trauma: A Neurobiological Review
Wiet, S. (2017) · Journal of korean Academy of Child and Adolescent Psychiatry · doi:10.5765/jkacap.2017.28.1.4 · Cited by 7
Minami, S., Nomura, H., & Minami, M. (2018) · Neuropsychopharmacology Reports · doi:10.1002/npr2.12036 · Cited by 7
Banasikowski, T. J., & Hawken, E. R. (2019) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2019.00170 · Cited by 8
Bed nucleus of the stria terminalis regulates fear to unpredictable threat signalsPreprint
Goode, T. D., Ressler, R. L., Acca, G. M., Miles, O. W., & Maren, S. (2018) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/376228 · Cited by 8
“Increases in immediate early gene expression, metabolic activity, and changes in 441 electrophysiological firing patterns have been observed in the BNST in response to direct exposure 442 to conditioned contextual cues (Campeau et al., 1997; Daldrup et al., 2016; Jennings et al., 2013; 443 Lemos et al., 2010; Luyten e…”
Control of defensive behavior by the nucleus of Darkschewitsch GABAergic neurons
Zhao, H., Liu, J., Shao, Y., Feng, X., Zhao, B., Sun, L., Liu, Y., Zeng, L., Li, X., Yang, H., et al. (2024) · National Science Review · doi:10.1093/nsr/nwae082 · Cited by 7
“Conditioning paradigms were used to demonstrate that animals avoid contextual cues previously associated with fear [22 –26 ].”
Lubianiker, N., Koren, T., Djerassi, M., Sirotkin, M., Singer, N., Jalon, I., Lerner, A., Sar-El, R., Sharon, H., Shahar, M., et al. (2026) · Nature Medicine · doi:10.1038/s41591-025-04140-5 · Cited by 11
“…and vice versa. indeed, it is known that the VTA in general, and specifically reward-related VTA, is functionally connected to other regions except the Nac via mesocortical and mesolimbic connections, such as the dorsolateral and medial prefrontal cortex 24 , the amygdala 25 and hippocampus 41 .”
InfluentialKikuchi, E., Inui, T., Su, S., Sato, Y., & Funahashi, M. (2022) · Behavioural Brain Research · doi:10.1016/j.bbr.2022.114253 · Cited by 6
Miura, Y., Shanley, M. R., Sanchez, A. S., & Friedman, A. K. (2023) · Frontiers in Neural Circuits · doi:10.3389/fncir.2022.1081099 · Cited by 7
“They are thought to be projecting to the VTA GABAergic interneurons to disinhibit VTA dopaminergic neurons (Kudo et al., 2012, 2014; Jennings et al., 2013; Soden et al., 2020).”
Huijgens, P. T., Heijkoop, R., Vanderschuren, L. J. M. J., Lesscher, H. M. B., & Snoeren, E. M. S. (2024) · Psychopharmacology · doi:10.1007/s00213-024-06561-5 · Cited by 6
Contemporary strategies for dissecting the neuronal basis of neurodevelopmental disorders
Seo, D.-O., Motard, L. E., & Bruchas, M. R. (2018) · Neurobiology of Learning and Memory · doi:10.1016/j.nlm.2018.03.015 · Cited by 7
Alcohol Cues, Craving, and Relapse: Insights from Animal Models
Pina, M. M., & Williams, A. R. (2016) · InTech eBooks · doi:10.5772/63105 · Cited by 7
“Of note, more recent evidence derived from studies using more advanced and selective tools suggests that a BNST GABA input to VTA GABA is the predominant source of the modulation of the BNST of the VTA [129].”
BackgroundResultEvidence for the involvement of a fronto-striatal pathway in the processing of social reward
Lin, K., & Coutellier, L. D. (2025) · Progress in Neurobiology · doi:10.1016/j.pneurobio.2025.102834 · Cited by 6
Bed nuclei of the stria terminalis: A key hub in the modulation of anxiety
Wang, X., Ge, S., & Zhang, C. (2023) · European Journal of Neuroscience · doi:10.1111/ejn.15926 · Cited by 5
Traumatic brain injury-induced anxiety: Injury and plasticity of the central noradrenergic system
Tsuda, S., Hou, J., Thompson, F. J., & Bose, P. (2025) · Experimental Neurology · doi:10.1016/j.expneurol.2025.115182 · Cited by 6
Chronic Ethanol Exposure Modulates Periaqueductal Gray to Extended Amygdala Dopamine Circuit
Pati, D., Downs, A. M., McElligott, Z. A., & Kash, T. L. (2022) · Journal of Neuroscience · doi:10.1523/jneurosci.1219-22.2022 · Cited by 5
Self-citationZhu, K., Tao, G., Huang, Z., Qu, W., & Wang, L. (2023) · European Journal of Neuroscience · doi:10.1111/ejn.16068 · Cited by 5
“Based on our results and previous studies in mice (Giardino et al., 2018; Jennings et al., 2013; Kim et al., 2013), we divided the BNST into eight subregions (Figure 5): The adlBNST, admBNST, F I GURE 4 The average proportion of whole-brain monosynaptic inputs to the bed nucleus of the stria terminalis (BNST) calretin…”
BackgroundResultLi, Y., Huang, Y., Chen, J. J., Hyland, B. I., & Wickens, J. R. (2024) · European Journal of Neuroscience · doi:10.1111/ejn.16269 · Cited by 5
“The GABA interneurons themselves also receive GABA synapses from the nucleus accumbens (Yang et al., 2018), the bed nucleus of the stria terminalis (Jennings et al., 2013) and the lateral hypothalamus (Nieh et al., 2016).”
BackgroundHartsock, M. J., Levy, C. T., Navarro, M. J., Saddoris, M. P., & Spencer, R. L. (2024) · Journal of Neuroscience · doi:10.1523/jneurosci.0878-24.2024 · Cited by 6
Optogenetics: illuminating the neural bases of rodent behavior
Lobo, M. K., Francis, C., & Chaudhury, D. (2014) · Open Access Animal Physiology · doi:10.2147/oaap.s42339 · Cited by 6
“136 Optogenetic activation of BNST GABA-ergic projections to the lateral hypothalamus enhance feeding while inhibition is aversive and suppresses feeding.”
InfluentialHellsten, K., Linden, A.-M., & Korpi, E. R. (2015) · Neuroscience · doi:10.1016/j.neuroscience.2015.02.052 · Cited by 5
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Stress History Modulates Corticotropin-Releasing Factor Neurons to Establish Resilience
Haynes, S. E., Lacagnina, A. F., Seo, H. S., Fang, L., Yang, X., Afzal, M. F., Morel, C., Menigoz, A., Rajan, K., Clem, R. L., et al. (2025) · Biological Psychiatry Global Open Science · doi:10.1016/j.bpsgos.2025.100656 · Cited by 5
“Previous work has shown that resiliency is influenced by dopaminergic VTA neurons in the nucleus accumbens (NAc) (27), in part through the actions of brain-derived neurotrophic factor (BDNF) (19,44,85).”
Ventral striatum links motivational and motor networks during operant-conditioned movement in rats
Hori, Y., Ihara, N., Sugai, C., Ogura, J., Honda, M., Kato, K., Isomura, Y., & Hanakawa, T. (2018) · NeuroImage · doi:10.1016/j.neuroimage.2018.10.018 · Cited by 5
“The limbic system plays a central role in motivational processes (Ikemoto et al., 2015; Ikemoto and Panksepp, 1996; Jennings et al., 2013; Koob, 1992; Wise, 1978; Wise and Bozarth, 1987).”
Rivera-Irizarry, J. K., Hámor, P. U., Rowson, S. A., Asfouri, J. J., Liu, D., Zallar, L. J., Garcia, A. F., Skelly, M. J., & Pleil, K. E. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.07.03.547570 · Cited by 6
Self-citationUntangling the multifaceted VTA responses to stress
Skupio, U., Harris, A. Z., & Polter, A. M. (2025) · Trends in Neurosciences · doi:10.1016/j.tins.2025.06.004 · Cited by 5
Li, W., Huang, X., Xu, X., Lv, Q., Wu, J., Hu, Z., Li, R., Yang, Y., Yu, L., Liu, B., et al. (2025) · Nature Neuroscience · doi:10.1038/s41593-025-02156-1 · Cited by 4
Laing, B. T., Anderson, M. S., Jayan, A., Park, A. S., Erbaugh, L. J., Solís, Ó., Wilson, D. J., Michaelides, M., & Aponte, Y. (2025) · eNeuro · doi:10.1523/eneuro.0417-24.2024 · Cited by 4
“Moreover, self-grooming behavior is observed following stimulation of glutamatergic neurons in the amygdala (Jennings et al., 2013).”
Worth, A. A., Feetham, C. H., Morrissey, N. A., & Luckman, S. M. (2024) · Frontiers in Endocrinology · doi:10.3389/fendo.2024.1449326 · Cited by 4
Pérez‐Garci, E., Pysanenko, K., Rizzi, G., Studer, F., Ulrich, D., Fritzius, T., Früh, S., Porcu, A., Besseyrias, V., Melichar, A., et al. (2025) · Neurobiology of Disease · doi:10.1016/j.nbd.2025.106831 · Cited by 5
“Aversive stimuli activate GABA VTA neurons (Jennings et al., 2013; Mitten et al., 2024; Tan et al., 2012), and knockdown of HCN2 in the VTA (Zhong et al., 2018), along with decreased excitability in DA VTA neurons results in anxiety-like behavior (Zweifel et al., 2011).”
Patrono, E., Nishijo, H., Gasbarri, A., Pompili, A., & Tomaz, C. A. B. (2017) · Springer eBooks · doi:10.1007/978-3-319-53126-7_16 · Cited by 4
Synaptic Regulation of Metabolism
Xu, J., Bartolome, C. L., & Kong, D. (2018) · Advances in experimental medicine and biology · doi:10.1007/978-981-13-1286-1_4 · Cited by 4
Balancing anxiety and social desire
Blackman, D. R., & Falkner, A. L. (2021) · Nature Neuroscience · doi:10.1038/s41593-021-00812-w · Cited by 5
MouseCircuits.org: An online repository to guide the circuit era of disordered affectPreprint
Anderson, K. R., & Dumitriu, D. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.02.16.951608 · Cited by 3
Erasing motion: Scrambling direction selectivity in visual cortex during saccadesPreprint
Miura, S. K., & Scanziani, M. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.03.30.437338 · Cited by 4
Pross, A., Metwalli, A. H., Abellán, A., Desfilis, E., & Medina, L. M. (2023) · The Journal of Comparative Neurology · doi:10.1002/cne.25569 · Cited by 3
“, 2006) could lead to activation of HPA or other targets of BSTL (such as the ventral tegmental area, involved in both reward and aversion; Riters et al., 2012; Jennings et al., 2013; Holy and Miczek, 2016), but their inhibition by GABAergic inputs from the central amygdala or from other cells of BSTL could have the o…”
BackgroundThe link between early-life adversity and later alcohol use disorder: A role for microglia?
Lichtenstein, H. D., Sequeira, M. K., & Bolton, J. L. (2025) · Neurobiology of Stress · doi:10.1016/j.ynstr.2025.100714 · Cited by 3
“For example, during adolescence, the brain ’ s “ go system ” , consisting of the mesolimbic reward system (NAc, VTA) are more mature at this time than the brain ’ s “ stop system ” , which involves top-down control from brain regions such as the PFC and orbitofrontal cortex (Melbourne et al., 2021).”
InfluentialWittenberg, R. E., Yun, S., Yang, K., Swanson, O. K., Wolfman, S. L., Colón, L. M., Eisch, A. J., & Dani, J. A. (2025) · Biological Psychiatry · doi:10.1016/j.biopsych.2025.05.027 · Cited by 3
Miller, C. N., Li, Y., Beier, K. T., & Aoto, J. N. (2025) · Nature Communications · doi:10.1038/s41467-025-60512-y · Cited by 4
“The aBNST is a highly sexually dimorphic nucleus that controls stress andanxietystates 52,53 .”
Matsuda, T. (2025) · Neuroscience Research · doi:10.1016/j.neures.2025.104941 · Cited by 3
The distinct subpopulations in parafascicular thalamic nucleus encoding innate emotional valence
Chen, P.-P., Wang, J.-G., Teng, S.-W., Yao, Z.-X., Che, Y.-Y., Chen, X.-L., Gao, S.-B., Chi, R.-F., Zhang, D., & Chen, Z.-Y. (2025) · Cell Reports · doi:10.1016/j.celrep.2025.116482 · Cited by 2
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Contrôle de l’anxiété par les récepteurs de la sérotonine 5-HT2c de la strie terminale
Bockaert, J., & Bécamel, C. (2017) · médecine/sciences · doi:10.1051/medsci/20173301015 · Cited by 3
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Anxiety as a Positive Epistemic Emotion in Politics
Brasi, L. D., Guglielmetti-Serrano, F., & Rosati, A. (2021) · Critical Review · doi:10.1080/08913811.2021.1924994 · Cited by 3
Bed Nucleus of Stria Terminalis (BNST) circuits
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Antagonistic Neural Circuits Drive Opposing Behaviors towards the Young in FemalesPreprint
Mei, L., Yan, R., Yin, L., Sullivan, R. M., & Lin, D. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.03.12.532288 · Cited by 3
Tsuruga, R., Tajika, Y., Minami, M., & Tsutsui-Kimura, I. (2026) · Communications Biology · doi:10.1038/s42003-026-10485-5 · Cited by 2
Serotonergic Modulation of the BNST-CeA Circuit Promotes Sex Differences in Fear Learning
Ravenelle, R., Fernandes-Henriques, C., Lee, J., Liu, J., Likhtik, E., & Burghardt, N. S. (2024) · Biological Psychiatry · doi:10.1016/j.biopsych.2024.02.038 · Cited by 2
What’s the occasion? Phasic dopamine signaling and interoception
Roitman, M. F., & McCutcheon, J. E. (2025) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2025.103074 · Cited by 3
“One possibility is via the bed nucleus of the stria terminalis (BNST) d which plays a role in eating and drinking and modulates dopamine cell body activity [53].”
Advances in the study of pharmacotherapy for addiction to naturally-derived psychoactive substances
Xie, K., Xiao, D., Xu, P., Shen, H., & Di, B. (2025) · Chinese Journal of Natural Medicines · doi:10.1016/s1875-5364(25)60831-4 · Cited by 3
Reduced oxytocin signaling in the dBNST drives the transition from acute pain to persistent anxiety
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Laing, B. T., Anderson, M. S., Jayan, A., Park, A. S., Erbaugh, L. J., Solís, Ó., Wilson, D. J., Michaelides, M., & Aponte, Y. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.09.06.506804 · Cited by 2
Punishment Risk Task: Monitoring anxiogenic states during goal directed actions in micePreprint
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Acute and chronic alcohol modulation of extended amygdala calcium dynamicsPreprint
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“In a similar vein, the BNST exhibits complementary ventral tegmental area (VTA) outputs distinguished by expression of either GABA or glutamate, which convey reward and aversion, respectively (14).”
Self-citationParker, K. E., Kuo, C., Buckley, A. R., Patterson, A. P., Duong, V., Hunter, S. C., & McCall, J. G. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.10.04.615025 · Cited by 3
“We used custom-made unbiased, balanced two-compartment conditioning apparatus (52.5 x 25.5 x 25.5 cm) as described previously 16,23,86,87 .”
MethodInfluentialDevelopmental Shifts in Amygdala Function
Rosenkranz, J. A. (2024) · Current topics in behavioral neurosciences · doi:10.1007/7854_2024_538 · Cited by 1
Separable dorsal raphe dopamine projections mimic the facets of a loneliness-like state
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Self-citationVGluT3 BNST neurons transmit GABA and restrict sucrose consumption
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“The VTA GABA neurons are believed to play a critical regulatory role in anxiety (Jennings et al, 2013; McHenry et al, 2017; Nieh et al, 2016).”
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“However, like other nearby hypothalamic and extended amygdala regions (Barker et al., 2017;Jennings et al., 2013 ; Lammel et al., 2012; Nieh et al., 2016), VP also contains glutamatergic populations that instead mediate aversion, in part via their direct projections to lateral habenula (LHb) (Haber…”
Endocannabinoid modulation of a reciprocal fronto-coerulear connection in contextual discrimination
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“Previous attempts at answering these questions have used antidromic activation of axons within the VTA to identify neurons in an input region that project to the midbrain (Jennings et al., 2013; Hong and Hikosaka, 2014).”
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Metadata
- Authors
- Jennings, J. H., Sparta, D. R., Stamatakis, A. M., Ung, R. L., Pleil, K. E., Kash, T. L., & Stuber, G. D.
- Affiliations
- University of North Carolina at Chapel Hill (US)
- Corresponding author
- Stuber, Garret D.
- ORCID
- Venue
- Nature
- Published
- 2013-03-20
- Volume
- 496
- Issue
- 7444
- Pages
- 224-228
- ISSN
- 0028-0836 (ISSN-L)
- 1476-4687
- PMC
- PMC3778934
- OpenAlex
- W2067203693
- Semantic Scholar
- b094f8c7bc24b675130d787985a6c38dc2555363
- Cited by
- 704
- References
- 33
- Citations by year
- 2026: 34 · 2025: 53 · 2024: 30 · 2023: 44 · 2022: 35 · 2021: 65 · 2020: 43 · 2019: 55 · 2018: 57 · 2017: 55 · 2016: 78 · 2015: 66 · 2014: 58 · 2013: 29
- FWCI
- 37.77
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Licence
- www.springer.com/tdm (tdm)
- Locations
- Nature (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- RePEc: Research Papers in Economics (submittedVersion)
- Funders
- Foundation for the National Institutes of Health
- Whitehall Foundation
- National Institute on Alcohol Abuse and Alcoholism (F32AA018610, AA011605, AA 007573, F32AA021043, AA018610, AA021043, P60AA011605, T32AA007573)
- National Institute on Drug Abuse (F31DA034472, R01DA032750, DA034472, DA029325, R21DA029325, DA032750, R37DA032750)
- National Institute of Neurological Disorders and Stroke (P30NS045892, NS 007431, T32NS007431)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neurotransmitter Receptor Influence on Behavior
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Keywords
- bed nucleus of the stria terminalis, ventral tegmental area, extended amygdala, glutamatergic projections, GABAergic projections, anxiety, reward, aversion, motivational states, depression, channelrhodopsin-2, aversive stimuli
- MeSH terms
- Channelrhodopsins, Amygdala, Animals, Anxiety, Avoidance Learning, Behavior, Animal, Cues, Electroshock, Glutamine, Male, Mice, Inbred C57BL, Motivation, Phenotype, Reward, Septal Nuclei, Ventral Tegmental Area, Mice, GABAergic Neurons, Optogenetics
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