135:, such as that taking place in fear conditioning, might occur. In this model of fear conditioning, strong depolarization of the lateral amygdala elicited by the stimulus leads to the strengthening of temporally and spatially relative conditioned stimulus inputs (that are coactive) onto the same neurons. Experimental data has been shown to support the idea that the plasticity and fear memory formation in the lateral amygdala are triggered by unconditioned stimulus-induced activation of the region's neurons. Thus, unconditioned stimulus-evoked depolarization is necessary for the enhancement of conditioned stimulus-elicited neural responses in this region after conditioned-unconditioned pairing and pairing a conditioned stimulus with direct depolarization of the lateral amygdala's pyramidal neurons as an unconditioned stimulus supports fear conditioning. It is also clear that synaptic plasticity at conditioned stimulus input pathways to the lateral amygdala does occur with fear conditioning.
327:, who had a rare bilateral amygdala damage, could not discern fear expressions because of her inability to look at the eye region of the face. When the subject was instructed to look directly at the eye region of faces with expression, the subject could recognize fear expressions of faces. Although the amygdala does play an important part in the recognition of fear, further research shows that there are alternate pathways that are capable to support fear learning in the absence of a functional amygdala. A study by Kazama also shows that although the amygdala may be damaged, it is still possible for patients to distinguish the difference between safety cues and fear.
301:(mGluRs). The proteins mGluRs likely serve a modulatory function and do not participate directly in Hebbian processes. This is because due to the fact these receptors do not contribute to depolarization during synapses. They are also not activated by receptors that participate in Hebbian processes. Finally, they do not detect pre- and postsynaptic neural activity. However, the activation of group I mGluRs in the lateral amygdala and basal nucleus enhances the acquisition, reduction, and amplification of fear conditioning by providing an influx of calcium ions.
93:
biochemistry (as mentioned below), chronic infusion of propranolol (beta-adrenergic receptor antagonist) prevented the behavioral changes following repeated stressor exposure thus halting long term potentiation. Some physiological changes also occurred including the decrease in body weight gain and adrenal hypertrophy observed in animals exposed to stress. Overall, the conditioned fear responses can contribute to behavioral changes in a repeated stress paradigm. This can be extended to correlate to other animals as well but with varying degrees of responses.
67:(LTP) and synaptic plasticity that enhances the response of lateral amygdala neurons to the conditioned stimulus occurs in the lateral amygdala. As a result, the conditioned stimulus is then able to flow from the lateral amygdala to the central nucleus of the amygdala. The basal and intercalated masses of the amygdala connect the lateral amygdala with the central nucleus of the amygdala directly and indirectly. Pathways from central nucleus of the amygdala to downstream areas then control defensive behavior (freezing) and
339:, where a neutral stimulus, such as a flash of light, is paired with a shock is given to a rat. The result of this conditioned stimulus is to provoke the unconditioned response, fear. The once neutral stimulus is given again to see if the rat would show the responses of fear. However, because fear responses involve many behaviors, it is important to see which behaviors are exhibited when the conditioned stimulus is given.
88:
altering an animal's (Fischer rat's) behavior in a repeated stress paradigm. Behavioral changes that are commonly referred to as depressive-like behaviors resulted from this model of testing. After setting a control and a valid experimental design, Fischer rats were exposed daily to different stressors in a complex environment. After four days of
411:
289:. They may also act in a parallel fashion with Hebbian mechanisms to implement synapses in the lateral amygdala and promote plasticity and fear learning through their respective signaling pathways. Accumulating evidence suggests that midbrain dopaminergic innervation of the basolateral amygdala facilitate the formation of fear memories.
319:
ranging from happiness to surprise to fear to sadness to disgust to anger. While control subjects classified these images to the nearest expression, subjects who had damage to the bilateral amygdala had problems with this task, especially with the recognition of facial expressions that show fear. The
238:
on other forms of learning, this effect is specific to only acquisition, as opposed to the posttraining processing or expression of fear memory. The activation of β-ARs in the lateral amygdala synergistically regulates
Hebbian processes to trigger the neuron's associative plasticity and fear learning
381:
was borrowed which states that the presentation of a neutral visual scene intensifies the percept of fear or suspense induced by a different channel of information, such as language. This principle has been applied in a way in which the percept of fear was present and amplified in the presence of a
87:
It has been observed that fear can contribute to behavioral changes. One way this phenomenon has been studied is on the basis of the repeated stress model done by Camp RM et al.(among others). In this particular study, it was examined that the contribution fear conditioning may play a huge role in
92:
exposure, both exploratory behavior and social interaction were tested on day 5 in either the same environment or a new environment. The rats showed much decreased exploration and social interaction when tested in different contexts compared to control rats. To further make a correlation to the
351:
and relayed to the amygdala for potential danger. The visual thalamus also relays the information to the visual cortex and is processed to see if the stimuli poses a potential threat. If so, this information is relayed to the amygdala and the muscle contraction, increased heart rate and blood
356:. A presentation of a neutral visual stimuli has been shown to intensify the percept of fear or suspense induced by a different channel of information, such as audition. From Le Doux's research, it shows that sound stimuli are not directly relayed from the auditory thalamus to the
382:
neutral visual stimuli. The main idea is that the visual stimuli intensify the fearful content of the stimuli (i.e. language) by subtly implying and concretizing what is described in the context (i.e. sentence). Activation levels in the right anterior
393:
also appear to influence pain through an interaction known as the valence-by-arousal interaction. During this reaction, negative emotions experienced by an individual with low levels of arousal tend to cause enhanced pain while negative
147:(NMDARs) and are located on postsynaptic neurons in the lateral amygdala. NMDARs are known to be coincidence detectors of presynaptic activity and postsynaptic depolarization. Auditory inputs are NMDARs in the lateral amygdala and use
907:
Liddell, Belinda J.; Brown, Kerri J.; Kemp, Andrew H.; Barton, Matthew J.; Das, Pritha; Peduto, Anthony; Gordon, Evian; Williams, Leanne M. (2005). "A direct brainstem–amygdala–cortical 'alarm' system for subliminal signals of fear".
368:
The perception of fear is elicited by many different stimuli and involves the process described above in biochemical terms. Neural correlates of the interaction between language and visual information has been studied by Roel
Willems
398:
emotions with higher levels of arousal have been observed to decrease the perception of pain. Low levels of arousal would include reactive emotions such as anxiety while higher levels of arousal include emotions such as fear.
314:
Research studies have shown that damage to the bilateral amygdala affects mostly the recognition of fear. In a specific study conducted by Andrew J. Calder and Andrew W. Young, they had subjects classify morphed images of
101:
Molecular mechanisms that have been linked directly to the behavioral expression of conditioning are easier to study in a clinical setting as opposed to mechanisms that underlie long-term potentiation (LTP), in which
106:
is induced by electrical or chemical stimulation of lateral amygdala circuits. LTP is important for fear processing because it strengthens the synapses in neural circuits. These strengthened synapses are how
520:
Camp, Robert M.; Remus, Jennifer L.; Kalburgi, Sahana N.; Porterfield, Veronica M.; Johnson, John D. (2012). "Fear conditioning can contribute to behavioral changes observed in a repeated stress model".
1375:; Young, Andrew W.; Calder, Andrew J.; Hellawell, Deborah J.; Aggleton, John P.; Johnsons, Michael (1997). "Impaired auditory recognition of fear and anger following bilateral amygdala lesions".
183:
also contribute to fear conditioning. The
Hebbian mechanisms contribute to plasticity in the lateral amygdala and fear learning. Other modulators apart from the Hebbian mechanisms include
1065:
Lutas, Andrew; Kucukdereli, Hakan; Alturkistani, Osama; Carty, Crista; Sugden, Arthur U.; Fernando, Kayla; Diaz, Veronica; Flores-Maldonado, Vanessa; Andermann, Mark L. (November 2019).
24:, has been hard-wired into almost every individual, due to its vital role in the survival of the individual. Researchers have found that fear is established unconsciously and that the
564:
Deandrade, Mark P.; Zhang, Li; Doroodchi, Atbin; Yokoi, Fumiaki; Cheetham, Chad C.; Chen, Huan-Xin; Roper, Steven N.; Sweatt, J. David; Li, Yuqing (2012). Di Cunto, Ferdinando (ed.).
175:
is called heterosynaptic plasticity. Homosynaptic plasticity is also prevalent which consists solely of the
Hebbian plasticity. In a variety of model systems, it has been shown that
230:(β-ARs) in the lateral nucleus of the amygdala interferes with the acquisition of fear learning when given pretraining stimuli but has no effect when applied posttraining or before
55:
In fear conditioning, the main circuits that are involved are the sensory areas that process the conditioned and unconditioned stimuli, certain regions of the amygdala that undergo
813:
Sigurdsson, Torfi; Doyère, Valérie; Cain, Christopher K.; Ledoux, Joseph E. (2007). "Long-term potentiation in the amygdala: A cellular mechanism of fear learning and memory".
320:
subjects with the damaged bilateral amygdala had no problems differentiating happiness from sadness, but they could not differentiate the expression of anger from fear.
151:
as a transmitter. Furthermore, it was tested that when the region's neurons that received auditory inputs also received unconditioned stimulus inputs and broad spectrum
285:(Gi-coupled) and stimulate adenylate cyclase (Gs-coupled), respectively. Just like β-ARs, dopamine receptors may modulate Hebbian processes directly by reducing
386:
were selectively increased and is believed to serve as a binding function of emotional information across domains such as visual and linguistic information.
676:"Neural substrates for the distinct effects of presynaptic group III metabotropic glutamate receptors on extinction of contextual fear conditioning in mice"
239:
in the lateral nucleus of the amygdala. One theory suggests that the mechanism of β-AR involvement in the acquisition of fear learning is that they act on
856:
Nader, Karim; Schafe, Glenn E.; Le Doux, Joseph E. (2000). "Fear memories require protein synthesis in the amygdala for reconsolidation after retrieval".
246:
to suppress feed-forward inhibition and enhance
Hebbian plasticity. β-ARs are found on GABAergic interneurons as well as in the lateral amygdala's
623:
Rogan, Michael T.; Stäubli, Ursula V.; Ledoux, Joseph E. (1997). "Fear conditioning induces associative long-term potentiation in the amygdala".
167:
It is believed that monoamine transmitters such as norepinephrine and dopamine that are released in emotional situations function in regulating
674:
Dobi, Alice; Sartori, Simone B.; Busti, Daniela; Van Der Putten, Herman; Singewald, Nicolas; Shigemoto, Ryuichi; Ferraguti, Francesco (2012).
323:
However, in an experiment conducted by Ralph
Adolphs, it elucidated the mechanism of the impaired fear recognition. Adolphs found that his
155:
in the lateral amygdala resulted in the disruption of the acquisition of fear learning. Therefore, these receptors are crucial to the
281:
subtypes) in the amygdala contributes to the acquisition of fear conditioning. D1 and D2 receptors are G protein coupled and inhibit
1500:"Add a picture for suspense: Neural correlates of the interaction between language and visual information in the perception of fear"
1174:
Calder, Andrew J. (1996). "Facial
Emotion Recognition after Bilateral Amygdala Damage: Differentially Severe Impairment of Fear".
1566:
298:
258:(PKA). This activation can elicits the phosphorylation of NMDARs as well as the ser845 site on GluA1, which could facilitate
79:
in fear expression as well, possibly by way of its connections to the basal and then to the central nucleus of the amygdala.
44:
35:
By understanding how fear is developed within individuals, it may be possible to treat human mental disorders such as
1201:
Adolphs, Ralph; Gosselin, Frederic; Buchanan, Tony W.; Tranel, Daniel; Schyns, Philippe; Damasio, Antonio R. (2005).
953:"A VTA to Basal Amygdala Dopamine Projection Contributes to Signal Salient Somatosensory Events during Fear Learning"
353:
1546:
Rhudy, JL. Williams, AE. "Gender differences in pain: do emotions play a role?" Gender
Medicine, 2005. p. 208-226.
1325:"Effects of neonatal amygdala lesions on fear learning, conditioned inhibition, and extinction in adult macaques"
200:
180:
226:
is a huge player in fear memory formation. Recent studies have demonstrated that the blockade of norepinephrine
1571:
68:
59:(or long-term potentiation) during learning, and the regions that bear an effect on the expression of specific
227:
286:
64:
60:
373:. The study consisted of observing how visual and linguistic information interact in the perception of
1127:"The Lateral Amygdaloid in Fear Conditioning Nucleus: Sensory Interface Amygdala in Fear Conditioning"
725:"Extinction of Fear-potentiated Startle: Blockade by Infusion of an NMDA Antagonist into the Amygdala"
1384:
1269:
1214:
865:
632:
577:
395:
274:
132:
1430:; Calder, A. J.; Andrew, C.; Giampietro, V.; Williams, S. C.; Bullmore, E. T.; Brammer, M. (1998).
336:
124:
103:
1408:
1287:
1238:
1202:
1008:
Cardozo Pinto, Daniel F.; Taniguchi, Lara; Norville, Zane C.; Pomrenze, Matthew B. (2020-09-30).
933:
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838:
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76:
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29:
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231:
192:
152:
108:
357:
172:
119:
Synaptic input can be strengthened when activity in the presynaptic neuron co-occurs with
56:
1388:
1273:
1218:
1067:"State-specific gating of salient cues by midbrain dopaminergic input to basal amygdala"
869:
636:
581:
495:
1524:
1499:
1464:
1431:
1349:
1324:
1151:
1142:
1126:
1099:
1066:
1042:
1009:
985:
774:"Post-training unilateral amygdala lesions selectively impair contextual fear memories"
749:
740:
724:
700:
675:
600:
565:
324:
247:
223:
204:
128:
120:
1125:
Ledoux, Joseph E.; Cicchetti, Piera; Xagoraris, Andrew; Romanski, Lizabeth M. (1990).
179:
modulate plasticity underlying memory formation such as a heightened percept of fear.
1561:
1555:
1300:
1257:
951:
Tang, Wei; Kochubey, Olexiy; Kintscher, Michael; Schneggenburger, Ralf (2020-05-13).
921:
826:
691:
383:
188:
171:
transmission and
Hebbian plasticity. The modulation of all of the different types of
168:
144:
1010:"A New Look at the Role of Mesoamygdaloid Dopamine Neurons in Aversive Conditioning"
937:
842:
1427:
1412:
1372:
1242:
1025:
968:
893:
660:
550:
465:
235:
566:"Enhanced Hippocampal Long-Term Potentiation and Fear Memory in Btbd9 Mutant Mice"
590:
378:
278:
243:
952:
1082:
534:
449:
406:
1455:
1090:
1033:
976:
1323:
Kazama, Andy M.; Heuer, Eric; Davis, Michael; Bachevalier, Jocelyne (2012).
1187:
251:
240:
184:
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148:
72:
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1234:
1108:
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994:
929:
885:
834:
799:
709:
609:
542:
457:
131:. This hypothesis is especially appealing as an explanation for how simple
1515:
1473:
1404:
1309:
1160:
790:
773:
758:
652:
503:
348:
270:
212:
89:
25:
1226:
20:
interprets stimuli and how animals develop fear responses. The emotion,
390:
374:
196:
36:
1486:
1432:"Neural responses to facial and vocal expressions of fear and disgust"
436:
Ledoux, Joseph (2003). "The
Emotional Brain, Fear, and the Amygdala".
1396:
1340:
1291:
1258:"A Subcortical Pathway to the Right Amygdala Mediating 'Unseen' Fear"
877:
208:
40:
723:
Falls, William A.; Miserendino, Mindy J. D.; Davis, Michael (1992).
644:
259:
17:
1203:"A mechanism for impaired fear recognition after amygdala damage"
482:
Davis, M (1992). "The Role of the Amygdala in Fear and Anxiety".
21:
293:
Metabotropic glutamate receptor-mediated neuromodulation during
347:
Initially, the visual stimuli is first received by the visual
250:. The process of activation of β-ARs start off by coupling to
215:) but the role of these compounds are not fully understood.
335:
There has been a substantial amount of research done on
389:
Exposure to different types of emotion and levels of
159:
of processing and eliciting for the percept of fear.
16:
Many experiments have been done to find out how the
63:. These pathways converge in the lateral amygdala.
297:Plasticity and learning can also be modulated by
1498:Willems, R. M.; Clevis, K.; Hagoort, P. (2010).
1262:Proceedings of the National Academy of Sciences
1120:
1118:
515:
513:
477:
475:
163:Monoamine neuromodulatory-dependent mechanisms
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429:
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8:
1504:Social Cognitive and Affective Neuroscience
1256:Morris, J. S.; Ohman, A; Dolan, RJ (1999).
1523:
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1348:
1299:
1281:
1150:
1098:
1041:
984:
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748:
699:
599:
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143:Hebian plasticity is believed to involve
254:signaling cascades, which then activate
139:NMDA-type ionotropic glutamate receptors
111:is developed and how fear is developed.
75:responses. Recent studies implicate the
423:
772:Flavell, C. R.; Lee, J. L. C. (2012).
352:pressure begins, thus activating the
7:
496:10.1146/annurev.ne.15.030192.002033
438:Cellular and Molecular Neurobiology
1436:Proceedings of the Royal Society B
1143:10.1523/JNEUROSCI.10-04-01062.1990
741:10.1523/JNEUROSCI.12-03-00854.1992
14:
922:10.1016/j.neuroimage.2004.08.016
827:10.1016/j.neuropharm.2006.06.022
692:10.1016/j.neuropharm.2012.05.025
409:
299:metabotropic glutamate receptors
1426:Phillips, M. L.; Young, A. W.;
1026:10.1523/JNEUROSCI.1483-20.2020
969:10.1523/JNEUROSCI.1796-19.2020
145:N-methyl-d-aspartate receptors
1:
484:Annual Review of Neuroscience
45:posttraumatic stress disorder
591:10.1371/journal.pone.0035518
354:sympathetic neuronal pathway
234:. In contrast to effects of
1131:The Journal of Neuroscience
1014:The Journal of Neuroscience
729:The Journal of Neuroscience
377:. A common phenomenon from
343:Visual and auditory stimuli
129:Hebbian synaptic plasticity
115:Hebbian synaptic plasticity
1595:
523:Behavioural Brain Research
273:receptor activation (both
262:insertion at the synapse.
1176:Cognitive Neuropsychology
1083:10.1038/s41593-019-0506-0
535:10.1016/j.bbr.2012.05.040
201:gastrin-releasing peptide
1567:Behavioral neuroscience
1329:Behavioral Neuroscience
1188:10.1080/026432996381890
957:Journal of Neuroscience
450:10.1023/A:1025048802629
287:feed-forward inhibition
1448:10.1098/rspb.1998.0506
1283:10.1073/pnas.96.4.1680
236:β-AR receptor blockade
228:β-adrenergic receptors
65:Long-term potentiation
51:Neuronal fear pathways
791:10.1101/lm.025403.111
778:Learning & Memory
61:conditioned responses
133:associative learning
1516:10.1093/scan/nsq050
1442:(1408): 1809–1817.
1389:1997Natur.385..254S
1274:1999PNAS...96.1680M
1227:10.1038/nature03086
1219:2005Natur.433...68A
1071:Nature Neuroscience
870:2000Natur.406..722N
637:1997Natur.390..604R
582:2012PLoSO...7E5518D
337:conditioned stimuli
331:Conditioned stimuli
127:. This is known as
125:postsynaptic neuron
104:synaptic plasticity
317:facial expressions
1077:(11): 1820–1833.
1020:(40): 7590–7592.
963:(20): 3969–3980.
815:Neuropharmacology
680:Neuropharmacology
417:Philosophy portal
283:adenylate cyclase
157:metabolic pathway
153:NMDAR antagonists
30:fear conditioning
28:is involved with
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1397:10.1038/385254a0
1373:Scott, Sophie K.
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310:Fear recognition
256:protein kinase A
232:memory retrieval
193:endocannabinoids
109:long-term memory
83:Behavioral basis
77:prelimbic cortex
1594:
1593:
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1572:Neuropsychology
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1497:
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1485:
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1425:
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1383:(6613): 254–7.
1371:
1370:
1366:
1322:
1321:
1317:
1255:
1254:
1250:
1213:(7021): 68–72.
1200:
1199:
1195:
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1172:
1168:
1124:
1123:
1116:
1064:
1063:
1059:
1007:
1006:
1002:
950:
949:
945:
906:
905:
901:
864:(6797): 722–6.
855:
854:
850:
812:
811:
807:
771:
770:
766:
722:
721:
717:
673:
672:
668:
631:(6660): 604–7.
622:
621:
617:
563:
562:
558:
519:
518:
511:
481:
480:
473:
444:(4/5): 727–38.
435:
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415:
410:
408:
405:
366:
358:central nucleus
345:
333:
312:
307:
295:
268:
248:pyramidal cells
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181:Neuromodulators
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97:Molecular basis
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12:
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5:
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1539:
1490:
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1364:
1335:(3): 392–403.
1315:
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1193:
1182:(5): 699–745.
1166:
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1000:
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899:
848:
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764:
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666:
615:
556:
509:
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305:Fear circuitry
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267:
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224:Norepinephrine
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219:Norepinephrine
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195:, and various
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121:depolarization
116:
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1275:
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1268:(4): 1680–5.
1267:
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1232:
1228:
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1137:(4): 1062–9.
1136:
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1128:
1121:
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1487:Willems lab
379:film theory
279:D2 receptor
1556:Categories
910:NeuroImage
686:: 274–89.
490:: 353–75.
403:References
364:Perception
177:monoamines
173:plasticity
57:plasticity
1456:0962-8452
1091:1546-1726
1034:0270-6474
977:0270-6474
252:G protein
241:GABAergic
199:(such as
185:serotonin
149:glutamate
73:endocrine
69:autonomic
1577:Amygdala
1534:20530540
1359:22642884
1235:15635411
1109:31611706
1052:32998956
995:32277045
938:18969739
930:15588615
886:10963596
843:14407392
835:16919687
800:22615481
710:22643400
610:22536397
570:PLOS ONE
543:22664265
458:14514027
396:valenced
349:thalamus
271:Dopamine
266:Dopamine
213:oxytocin
197:peptides
90:stressor
26:amygdala
1525:3150851
1474:9802236
1465:1689379
1413:4332467
1405:9000073
1385:Bibcode
1350:3740331
1310:9990084
1270:Bibcode
1243:2139996
1215:Bibcode
1161:2329367
1152:6570227
1100:6858554
1043:7531542
986:7219297
894:4420637
866:Bibcode
759:1347562
750:6576037
701:3557389
661:4310181
653:9403688
633:Bibcode
601:3334925
578:Bibcode
551:8144171
504:1575447
466:3216382
391:arousal
375:emotion
209:opiates
123:in the
37:anxiety
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211:, and
43:, and
41:phobia
1409:S2CID
1301:15559
1292:47262
1288:JSTOR
1239:S2CID
934:S2CID
890:S2CID
839:S2CID
657:S2CID
547:S2CID
462:S2CID
371:et al
260:AMPAR
18:brain
1562:Fear
1530:PMID
1470:PMID
1452:ISSN
1401:PMID
1355:PMID
1306:PMID
1231:PMID
1157:PMID
1105:PMID
1087:ISSN
1048:PMID
1030:ISSN
991:PMID
973:ISSN
926:PMID
882:PMID
831:PMID
796:PMID
755:PMID
706:PMID
649:PMID
606:PMID
539:PMID
500:PMID
454:PMID
277:and
71:and
22:fear
1520:PMC
1512:doi
1460:PMC
1444:doi
1440:265
1393:doi
1381:385
1345:PMC
1337:doi
1333:126
1296:PMC
1278:doi
1223:doi
1211:433
1184:doi
1147:PMC
1139:doi
1095:PMC
1079:doi
1038:PMC
1022:doi
981:PMC
965:doi
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874:doi
862:406
823:doi
786:doi
745:PMC
737:doi
696:PMC
688:doi
641:doi
629:390
596:PMC
586:doi
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527:233
492:doi
446:doi
205:NPY
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