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The amygdala sends projections to the hypothalamusthe dorsomedial thalamus, the thalamic reticular nucleusthe nuclei of the trigeminal nerve and the facial nervethe ventral tegmental areathe locus coeruleusand the laterodorsal tegmental nucleus. The medial nucleus is involved in the sense of smell and pheromone -processing. It receives input from the olfactory bulb and olfactory cortex. The centromedial nuclei are the main outputs for the basolateral complexes, and are involved in emotional arousal in rats and cats.
In complex vertebrates, including humans, the amygdalae perform primary roles in the formation and storage of memories associated with emotional events. Research indicates that, during fear conditioningsensory stimuli reach the basolateral complexes of the amygdalae, particularly the lateral nuclei, where they form associations with memories of the stimuli. The association between stimuli and the aversive events they predict may be mediated by long-term potentiation  a sustained enhancement of signaling between affected neurons.
One study examined a patient with bilateral degeneration of the amygdala. He was told a violent story accompanied by matching pictures and was observed based on how much he could recall from the story.
The patient had less recollection of the story than patients with functional amygdala, showing that the amygdala has a strong connection with emotional learning. Emotional memories are thought to be stored in synapses throughout the brain. Fear memories, for example, are considered to be stored in the neuronal connections from the lateral nuclei to the central nucleus of the amygdalae and the bed nuclei of the stria terminalis part of the extended amygdala.
Of course, these connections are not the sole site of fear memories given that the nuclei of the amygdala receive and send information to other brain regions that are important for memory such as the hippocampus.
Some sensory neurons project their axon terminals to the central nucleus. Damage to the amygdalae impairs both the acquisition and expression of Pavlovian fear conditioning, a form of classical conditioning of emotional responses.
The amygdalae are also involved in appetitive positive conditioning. It seems that distinct neurons respond to positive and negative stimuli, but there is no clustering of these distinct neurons into clear anatomical nuclei.
Lesions of the basolateral regions do not exhibit the same effect. Thus, the acquired preference for male-derived volatiles reveals an olfactory-vomeronasal associative learning.
Moreover, the reward system is differentially activated by the primary pheromones and secondarily attractive odorants. Exploring the primary attractive pheromone activates the basolateral amygdala and the shell of nucleus accumbens but neither the ventral tegmental area nor the orbitofrontal cortex.
In contrast, exploring the secondarily attractive male-derived odorants involves activation of a circuit that includes the basolateral amygdala, prefrontal cortex and ventral tegmental area.
Therefore, the basolateral amygdala stands out as the key center for vomeronasal-olfactory associative learning. Glutamatergic neurons in the basolateral amygdala send projections to the nucleus accumbens shell and core.
Activation of these projections drive motivational salience. The ability of these projections to drive incentive salience is dependent upon dopamine receptor The Revealing Science Of The Amygdalae. The amygdala is also involved in the modulation of memory consolidation.
Following any learning event, the long-term memory for the event is not formed instantaneously. Rather, information regarding the event is slowly assimilated into long-term potentially lifelong storage over time, possibly via long-term potentiation. Recent studies suggest that the amygdala regulates memory consolidation in other brain regions.
Also, fear conditioninga type of memory that is impaired following amygdala damage, is mediated in part by long-term potentiation. During the consolidation period, the memory can be modulated. In particular, it appears that emotional arousal following the learning event influences the strength of the subsequent memory for that event.
Greater emotional arousal following a learning event enhances a person's retention of that event. Experiments have shown that administration of stress hormones to mice immediately after they learn something enhances their retention when they are tested two days later. The amygdala, especially the basolateral nuclei, are involved in mediating the effects of emotional arousal on the strength of the memory for the event, as shown by many laboratories including that of James McGaugh.
These laboratories have trained animals on a variety of learning tasks and found that drugs injected into the amygdala after training affect the animals' subsequent retention of the task. These tasks include basic classical conditioning tasks such as inhibitory avoidance, where a rat learns to associate a mild footshock with a particular compartment of an apparatus, and more complex tasks such as spatial or cued water maze, where a rat learns to swim to a platform to escape the water.
If a drug that activates the amygdalae is injected into the amygdalae, the animals had better memory for the training in the task. In rats, DNA damage was found to increase in the amygdala immediately after exposure to stress. By seven days after exposure to these stresses, increased DNA damage was no longer detectable in the amygdala, probably because of DNA repair. Buddhist monks who do compassion meditation have been shown to modulate their amygdala, along with their temporoparietal junction and insuladuring their practice.
Amygdala activity at the time of encoding information correlates with retention for that information. However, this correlation depends on the relative "emotionalness" of the information. More emotionally arousing information increases amygdalar activity, and that activity correlates with retention.
Amygdala neurons show various types of oscillation during emotional arousal, such as theta activity. These synchronized neuronal events could promote synaptic plasticity which is involved in memory retention by increasing interactions between neocortical storage sites and temporal lobe structures involved in declarative memory. Research using Rorschach test blot 03 finds that the number of unique responses to this random figure links to larger sized amygdalae.
The researchers note, "Since previous reports have indicated that unique responses were observed at higher frequency in the artistic population than in the nonartistic normal population, this positive correlation suggests that amygdalar enlargement in the normal population might be related to creative mental activity.
Early research on primates provided explanations as to the functions of the amygdala, as well as a basis for further research. As early asrhesus monkeys with a lesioned temporal cortex including the amygdala were observed to have significant social and emotional deficits.
Some monkeys also displayed an inability to recognize familiar objects and would approach animate and inanimate objects indiscriminately, exhibiting a loss of fear towards the experimenters. Monkey mothers who had amygdala damage showed a reduction in maternal behaviors towards their infants, often physically abusing or neglecting them. With advances in neuroimaging technology such as MRIneuroscientists have made significant findings concerning the amygdala in the human brain.
A variety of data shows the amygdala has a substantial role in mental states, and is related to many psychological disorders. Some studies have shown children with anxiety disorders tend to have a smaller left amygdala.
In the majority of the cases, there was an association between an increase in the size of the left amygdala with the use of SSRIs antidepressant medication or psychotherapy. The left amygdala has been linked to social anxiety, obsessive and compulsive disordersand post traumatic stressas well as more broadly to separation and general anxiety. Some borderline patients even had difficulties classifying neutral faces or saw them as threatening. Patients with severe social phobia showed a correlation with increased response in the amygdala.
This hyperactivity was normalized when patients were administered antidepressant medication. A study found that adult and adolescent bipolar patients tended to have considerably smaller amygdala volumes and somewhat smaller hippocampal volumes. Studies in and showed that normal subjects exposed to images of frightened faces or faces of people from another race will show increased activity of the amygdala, The Revealing Science Of The Amygdalae if that exposure is subliminal.
Recent research suggests that parasites, in particular toxoplasmaform cysts in the brain of rats, often taking up residence in the amygdala. This may provide clues as to how specific parasites may contribute to the development of disorders, including paranoia. Conscious control of brain function towards positive brain response with accompanying changes in amygdala activity was suggested in the early s by independent behaviorist T. Lingo  and this possibility has been corroborated by later research, such as that carried on by Sara W.
Lazar,  Herbert Benson . Future studies have been proposed to address the role of the amygdala in positive emotions, and the ways in which the amygdala networks with other brain regions.
Recent studies have suggested possible correlations between brain structure, including differences in hemispheric ratios and connection patterns in the amygdala, and sexual orientation. Homosexual men tend to exhibit more The Revealing Science Of The Amygdalae patterns in the amygdala than heterosexual males do, just as homosexual females tend to show more masculine patterns in the amygdala than heterosexual women do.
It was observed that amygdala connections were more widespread from the left amygdala in homosexual males, as is also found in heterosexual females. Amygdala connections were more widespread from the right amygdala in homosexual females, as in heterosexual males. Amygdala volume correlates positively with both the size the number of contacts a person has and the complexity the number of different groups to which a person belongs of social networks, The Revealing Science Of The Amygdalae.
The amygdala is responsible for facial recognition and allows others to respond appropriately to different emotional expressions. It does not, however, process the direction of the gaze of the person being perceived. The amygdala is also thought to be a determinant of the level of a person's emotional intelligence.
It is particularly hypothesized that larger amygdalae allow for greater emotional intelligence, enabling greater societal integration and cooperation with others. The amygdala processes reactions to violations concerning personal space.
These reactions are absent in persons in whom the amygdala is damaged bilaterally. Animal studies have shown that stimulating the amygdala appears to increase both sexual and aggressive behavior. Likewise, studies using brain lesions have shown that harm to the amygdala may produce the opposite effect. Thus, it appears that this part of the brain may play a role in the display and modulation of aggression. There are cases of human patients with focal bilateral amygdala lesions due to the rare genetic condition Urbach-Wiethe disease.
This finding reinforces the conclusion that the amygdala "plays a pivotal role in triggering a state of fear". The amygdala appears to play a role in binge drinkingbeing damaged by repeated episodes of intoxication and withdrawal.
The protein is involved in controlling the function of other proteins and plays a role in development of the ability to consume a large amount of ethanol.
There may also be a link between the amygdala and anxiety. In an experiment, degu pups were removed from their mother but allowed to hear her call. In response, the males produced increased serotonin receptors in the amygdala but females lost them. This led to the males being less affected by the stressful situation. The clusters of the amygdala are activated when an individual expresses feelings of fear or aggression.
This occurs because the amygdala is the primary structure of the brain responsible for fight or flight response. Anxiety and panic attacks can occur when the amygdala senses environmental stressors that stimulate fight or flight response.
The amygdala is directly associated with conditioned fear. Conditioned fear is the framework used to explain the behavior produced when an originally neutral stimulus is consistently paired with a stimulus that evokes fear.
The amygdala represents a core fear system in the human body, which is involved in the expression of conditioned fear. Fear is measured by changes in autonomic activity including increased heart rate, increased blood pressure, as well as in simple reflexes such as flinching or blinking.
The central nucleus of the amygdala has direct correlations to the hypothalamus and brainstem — areas directly related to fear and anxiety. This connection is evident from studies of animals that have undergone amygdalae removal. Such studies suggest that animals lacking an amygdala have less fear expression and indulge in non-species-like behavior.
Many projection areas of the amygdala are critically involved in specific signs that are used to measure fear and anxiety. Mammals have very similar ways of processing and responding to danger. Scientists have observed similar areas in the brain — specifically in the amygdala — lighting up or becoming more active when a mammal is threatened or beginning to experience anxiety.
Similar parts of the brain are activated when rodents and when humans observe a dangerous situation, the amygdala playing a crucial role in this assessment. By observing the amygdala's functions, people can determine why one rodent may be much more anxious than another. There is a direct relationship between the activation of the amygdala and the level of anxiety the subject feels.
Feelings of anxiety start with a catalyst — an environmental stimulus that provokes stress. This can include various smells, sights, and internal sensations that result in anxiety. The amygdala reacts to this stimuli by preparing to either stand and fight or to turn and run.
This response is triggered by the release of adrenaline into the bloodstream. Consequently, blood sugar rises, The Revealing Science Of The Amygdalae immediately available to the muscles for quick energy.
Shaking may occur in an attempt to return blood to the rest of the body. A better understanding of the amygdala and its various functions may lead to a new way of treating clinical anxiety. There seems to be a connection with the amygdalae and how the brain processes posttraumatic stress disorder. Multiple studies have found that the amygdalae may be responsible for the emotional reactions of PTSD patients.
One study in particular found that when PTSD patients are shown pictures of faces with fearful expressions, their amygdalae tended to have a higher activation than someone without PTSD. Amygdala dysfunction during face emotion processing is well-documented in bipolar disorder.
The study indicates that the changes in amygdala volumes as well as changes in binding potentials, functional connectivity, regional homogeneity and regional cerebral blood flow were found to have various impacts on emotionality in people with Parkinson's disease.
Amygdala size has been correlated with cognitive styles with regard to political thinking. A study found that "greater liberalism was associated with increased gray matter volume in the anterior cingulate cortex, whereas greater conservatism was associated with increased volume of the right amygdala. From Wikipedia, the free encyclopedia. For other uses, see Amygdala disambiguation. Each of two small structures deep within the temporal lobe of complex vertebrates.
See also: Neuroscience of sex differences. Main article: Emotion and memory. Archived from the original on 18 October Retrieved 9 November Archived from the original on 31 March Retrieved 15 March Anatomy and Embryology.
Frontiers in Neuroanatomy. Brain imaging studies of healthy humans people without brain damage suggest something similar. When they are exposed to threats, neural activity in the amygdala increases and body responses like sweating or increased heart rate result.
This is true even if the threatening stimuli are presented subliminally, such that the person is not consciously aware that the threat is present and does not consciously experience feel "fear. The conclusion that the amygdala is the brain's fear center wrongly assumes that the feelings of "fear" and the responses elicited by threats are products of the same brain system. How did the interpretation come to be? We humans frequently feel afraid when we find ourselves freezing or fleeing when in harm's way.
In other words, these two things the feeling and the body responses tend to be tightly correlated in our conscious introspections. These introspections are talked about and become shared experiences that are ingrained as natural truths. Most people thus believe that the feeling of fear is the reason an animal or person runs from danger; or that the classic facial expression we know as "fear" is driven by feeling afraid. But when it comes to the brain, what is obvious is not always what is the case.
The purpose of science is to go beyond the obvious to reveal the deeper truths that cannot be gleaned simply from observing nature. One of the first things a scientist learns is that a correlation does not necessarily reveal causation. The interpretation that the amygdala is the brain's fear center confuses correlation and causation. Actually, there are two confusions involved: 1 because we often feel afraid when we are responding to danger, fear is the reason we respond the way we do; and 2 because the amygdala is responsible for the response to danger, it must also be responsible for the feeling of fear.
From the beginning, my research suggested that the amygdala contributes to non-conscious aspects of fear, by which I meant the detection of threats and the control of body responses that help cope with the threat. Conscious fear, I argued in my books The Emotional Brain Simon and Schuster, and Synaptic Self Viking,and most recently in Anxious Viking,is a product of cognitive systems in the neocortex that operate in parallel with the amygdala circuit. But that subtlety the distinction between conscious and non-conscious aspects of fear was lost on most people.
When one hears the word "fear," the pull of the vernacular meaning is so strong that the mind is compelled to think of the feeling of being afraid. For this reason, I eventually concluded that it is not helpful to talk about conscious and non-conscious aspects of fear.
A feeling like "fear" is a conscious experience. To use the word "fear" in any other way only leads to confusion. The amygdala has a role in fear, but it is not the one that is popularly described. It's role in fear is more fundamental and also more mundane. It is responsible for detecting and responding to threats, and only contributes to feelings of fear indirectly.
For example, the amygdala outputs driven by threat detection alter information processing in diverse regions of the brain. One important set of outputs result in the secretion of chemicals throughout the brain norepinephrine, acetylcholine, dopamine, serotonin and body hormones such as adrenalin and cortisol. In situations of danger, these chemicals alert the organism that something important is happening. As a result, attention systems in the neocortex guide the perceptual search the environment for an explanation for the highly aroused state.
The meaning of the environmental stimuli present is added by the retrieval of memories. If the stimuli are known sources of danger, "fear" schema are retrieved from memory. My hypothesis, then, is that the feeling of "fear" results when the outcome of these various processes attention, perception, memory, arousal coalesce in consciousness and compel one to feel "fear.
There's nothing wrong with speculation in science I just speculated about how feelings come about. But when a speculative interpretation becomes ingrained in the culture of science, and the culture at large, as an unquestioned fact, we have a problem.
This is problem is especially acute in neuroscience, where we start from mental state words like fear that have historical meanings, and treat the words as if they are entities that live in brain areas like the amygdala. In sum, there is no fear center out of which effuses the feeling of being afraid.
Postscript: Be suspicious of any statement that says a brain area is a center responsible for some function.
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