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Who Needs Emotions? The Brain Meets the Robot

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organization of motivational–emotional systems 47<br />

Walker, Brooks, & Holden–Dye, 1996). Neurotransmitters are released<br />

from axon terminals, cross <strong>the</strong> synapse, and bind to postsynaptic receptor<br />

sites to effect a cascade of intracellular biochemical events. Uptake sites<br />

on presynaptic terminals are proteins that regulate <strong>the</strong> synaptic level of<br />

neurotransmitter by binding released transmitter and transporting it back<br />

into <strong>the</strong> terminal. <strong>The</strong>se molecules play a role in adaptive behaviors to a<br />

surprisingly conserved degree across species and phyla. Subjective states<br />

in humans which are associated with such feelings as joy, fear, anxiety, and<br />

maternal love are derived from <strong>the</strong> actions of truly primordial chemical<br />

systems. Following <strong>the</strong> origins of bacterial life, eukaryotic cells appeared<br />

approximately 2 billion years ago, primitive multicelled organisms appeared<br />

around 800 million years ago, and vertebrates are estimated to have diverged<br />

from invertebrates around 500–600 million years ago. All extant mammals,<br />

birds, and reptiles are derived from stem reptiles that lived approximately<br />

200–300 million years ago. Neurotransmitter development followed this<br />

evolutionary path. All neurons, throughout <strong>the</strong> animal kingdom, contain<br />

at least one releasable substance (usually an amine, peptide, amino acid,<br />

or acetylcholine) and utilize ei<strong>the</strong>r ligand-gated ion channels or second<br />

messengers such as G proteins, AMP, phospholipase C, and calcium to<br />

communicate <strong>the</strong>ir signal postsynaptically. Second-messenger systems<br />

appeared quite early in evolution, perhaps to add a longer time scale and<br />

greater flexibility in neural communication.* For example, <strong>the</strong> yeast alphamating<br />

factor (a peptide pheromone) is a member of <strong>the</strong> G protein–coupled<br />

receptor superfamily (Darlison & Richter, 1999), and G protein–coupled<br />

receptors are found throughout arthropods, flatworms, and mollusks<br />

(Walker, Brooks, & Holden-Dye, 1996).† Calcium, a ubiquitous second<br />

messenger, plays this role even in bacteria (Tisa & Adler, 1992). Ligandgated<br />

channels, complex membrane-bound proteins that allow fast chemical<br />

transmission via gating of <strong>the</strong> flow of cations and anions in and out of <strong>the</strong><br />

cell (such as that involving g-aminobutyric acid, acetylcholine, and<br />

glutamate), are present in all animal species studied thus far. Chemical<br />

compounds can act in several ways: strictly as transmitters that convey<br />

specific information via <strong>the</strong>ir effect on postsynaptic receptors, as modulators<br />

of <strong>the</strong> postsynaptic receptor so as to alter o<strong>the</strong>r incoming signals, or as<br />

signals acting at sites distal from release sites, thus acting as neurohormones.<br />

*Ligand-gated ion channels are proteins that allow rapid flux of ions such as<br />

sodium or potassium in and out of <strong>the</strong> neuron, depending on <strong>the</strong> binding of neurotransmitter<br />

to its receptor. Second messengers are molecules that aid in <strong>the</strong> transduction<br />

of <strong>the</strong> chemical signal to an electrical signal.<br />

†G protein–coupled receptors are receptors for neurotransmitters that utilize<br />

specific membrane-bound proteins—G proteins—that activate certain critical intracellular<br />

second messenger enzymes, such as cyclic AMP.

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