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Who Needs Emotions? The Brain Meets the Robot

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94 brains<br />

<strong>the</strong> amygdala to determine whe<strong>the</strong>r danger is present in <strong>the</strong> sensory world,<br />

but in addition to processing <strong>the</strong> significance of external stimuli, <strong>the</strong> amygdala<br />

can influence sensory processing occurring in cortical areas. <strong>The</strong> amygdala<br />

receives inputs only from <strong>the</strong> late stages of cortical sensory processing, but<br />

it projects back to <strong>the</strong> earliest stages (Amaral, Price, Pitkanen, & Carmichael,<br />

1992). Thus, once <strong>the</strong> amygdala is activated by a sensory event from <strong>the</strong><br />

thalamus or cortex, it can begin to regulate <strong>the</strong> cortical areas that project to<br />

it, controlling <strong>the</strong> kinds of input it receives from <strong>the</strong> cortex (Fig. 4.4). <strong>The</strong><br />

amygdala also influences cortical sensory processes indirectly, by way of<br />

projections to various “arousal” networks, including <strong>the</strong> basal forebrain cholinergic<br />

system, <strong>the</strong> brain-stem cholinergic system, and <strong>the</strong> locus coeruleus<br />

noradrenergic system, each of which innervates widespread areas of <strong>the</strong><br />

Sensory Cortex<br />

Thalamus<br />

Stimulus<br />

Behavior<br />

mPFC dlPFC<br />

Working memory<br />

arousal<br />

Basal forebrain<br />

<strong>Brain</strong>stem<br />

Locus coeruleus<br />

bodily feedback<br />

Hormones<br />

Proprioception<br />

Figure 4.4. <strong>The</strong> amygdala (A) receives inputs only from <strong>the</strong> late stages of<br />

cortical sensory processing (thick arrow) but projects back to <strong>the</strong> earliest<br />

stages. Once <strong>the</strong> amygdala is activated by a sensory event from <strong>the</strong> thalamus<br />

or cortex, it can begin to regulate <strong>the</strong> cortical areas that project to it. <strong>The</strong><br />

amygdala also influences cortical sensory processes indirectly by way of<br />

projections to various “arousal” networks, including <strong>the</strong> basal forebrain<br />

cholinergic system, <strong>the</strong> brain-stem cholinergic system, and <strong>the</strong> locus coeruleus<br />

noradrenergic system, each of which innervates widespread areas of <strong>the</strong><br />

cortex. Thus, once <strong>the</strong> amygdala detects danger, it can activate <strong>the</strong>se arousal<br />

systems, which can <strong>the</strong>n influence sensory processing. <strong>The</strong> bodily responses<br />

initiated by <strong>the</strong> amygdala can also influence cortical areas by way of feedback<br />

ei<strong>the</strong>r from proprioceptive or visceral signals or hormones. <strong>The</strong> amygdala also<br />

interacts with <strong>the</strong> medial prefrontal cortex (mPFC), which toge<strong>the</strong>r with <strong>the</strong><br />

dorsolateral prefrontal cortex (dlPFC) has widespread influences on cognition<br />

and behavior and sends connections to several amygdala regions, allowing<br />

cognitive functions organized in prefrontal regions, especially working<br />

memory, to regulate <strong>the</strong> amygdala and its fear reactions. Modulatory/<br />

regulatory inputs are marked with dashed lines.<br />

A

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