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Who Needs Emotions? The Brain Meets the Robot

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asic principles for emotional processing 89<br />

Thalamus<br />

CS<br />

(tone)<br />

Auditory Cortex<br />

Behavior<br />

Figure 4.2. <strong>The</strong> neural pathways involved in fear conditioning are well<br />

characterized. When <strong>the</strong> conditioned stimulus (CS) is acoustic, <strong>the</strong> pathways<br />

involve transmission to <strong>the</strong> lateral nucleus of <strong>the</strong> amygdala (LA) from<br />

auditory processing areas in <strong>the</strong> thalamus and auditory cortex. LA, in turn,<br />

projects to <strong>the</strong> central nucleus of <strong>the</strong> amygdala (CE), which controls <strong>the</strong><br />

expression of fear responses by way of projections to brain-stem areas<br />

controlling <strong>the</strong> autonomic nervous system, <strong>the</strong> production of hormones, and<br />

<strong>the</strong> appropriate behavior.<br />

1997). Although some lesion studies have questioned <strong>the</strong> ability of <strong>the</strong> thalamic<br />

pathway to mediate conditioning (Shi & Davis, 1999), recordings from<br />

single neurons show that <strong>the</strong> cortical pathway conditions more slowly over<br />

trials than <strong>the</strong> thalamic pathway (Quirk, Armony, & LeDoux, 1997), thus<br />

indicating that <strong>the</strong> association between CS and US in <strong>the</strong> amygdala occurs<br />

initially through <strong>the</strong> thalamic pathway. Recent functional magnetic resonance<br />

imaging (fMRI) studies in humans have found that <strong>the</strong> amygdala shows<br />

activity changes during conditioning that correlate with activity in <strong>the</strong> thalamus<br />

but not <strong>the</strong> cortex (Morris, Ohman, & Dolan, 1999), fur<strong>the</strong>r emphasizing<br />

<strong>the</strong> importance of <strong>the</strong> direct thalamo-amygdala pathway.<br />

In addition to expressing fear responses to <strong>the</strong> CS, rats exhibit <strong>the</strong>se when<br />

returned to <strong>the</strong> chamber in which <strong>the</strong> tone and shock were paired or a chamber<br />

in which shocks occur alone. This is called “contextual fear conditioning,”<br />

where context refers to <strong>the</strong> various visual and olfactory aspects of <strong>the</strong> chamber,<br />

and requires both <strong>the</strong> amygdala and hippocampus, a brain structure<br />

know to enable long-term memories (Kim & Fanselow, 1992; Phillips &<br />

LeDoux, 1992; Maren, Aharonov, & Fanselow, 1997; Frankland et al.,<br />

1998). Areas of <strong>the</strong> hippocampus project to B and AB in <strong>the</strong> amygdala<br />

(Canteras & Swanson, 1992), and damage to <strong>the</strong>se areas interferes with<br />

contextual conditioning (Maren & Holt, 2000). Hippocampal projections<br />

to B and AB thus seem to be involved in contextual conditioning (for a<br />

LA<br />

CE<br />

Autonomic<br />

system<br />

Hormones

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