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Who Needs Emotions? The Brain Meets the Robot

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56 brains<br />

serotonergic system is proposed to be “<strong>the</strong> most expansive neurochemical<br />

network in <strong>the</strong> vertebrate CNS” (Jacobs & Azmitia, 1992). It is a system that<br />

is highly conserved across phyla; serotonin is an important neurotransmitter<br />

in many invertebrates, and over 15 subtypes of 5-HT receptors have arisen<br />

through molecular evolution, most of which interact with G proteins<br />

(Peroutka & Howell, 1994; Saudou & Hen, 1994). This extensive development<br />

of receptor subtypes suggests a great diversity of signaling within<br />

serotonin systems across phyla. Serotonin is a particularly interesting and<br />

appropriate chemical signal to examine in <strong>the</strong> context of <strong>the</strong> evolution of<br />

motivated behavior and emotion. Over <strong>the</strong> past 25 years or so, its pharmacology,<br />

physiology, and molecular biology have been extensively studied in<br />

crustaceans, insects, mollusks, worms, and mammals. Moreover, dysfunction<br />

of serotonin in humans has been implicated in many psychiatric disorders,<br />

such as depression, anxiety, obsessive–compulsive behavior, and<br />

alcoholism (Nemeroff, 1998). Serotonin-selective reuptake inhibitors (SSRIs)<br />

are among <strong>the</strong> most commonly prescribed psychiatric drugs. Thus, understanding<br />

<strong>the</strong> functions of 5-HT through analysis of its role in behavior and<br />

brain function may provide important insights into <strong>the</strong> neurochemical basis<br />

of human emotion and its disorders.<br />

While serotonin clearly has roles of a very diverse nature, <strong>the</strong>re is an interesting<br />

common thread that weaves through much of <strong>the</strong> research on this<br />

substance. Work from a variety of approaches leads to <strong>the</strong> general consensus<br />

that 5-HT plays a critical role in <strong>the</strong> modulation of aggression and agonistic<br />

social interactions in many animals and possibly regulation of aggressive<br />

behavior and mood in nonhuman primates and humans (Insel & Winslow,<br />

1998). Experiments on crustaceans such as <strong>the</strong> lobster and crayfish clearly<br />

implicate serotonin, as well as octopamine (a phenol analogue of norepinephrine),<br />

in <strong>the</strong> control of behaviors concerned with <strong>the</strong> maintenance of social<br />

hierarchies. <strong>The</strong> fighting behavior of <strong>the</strong>se phylogenetically ancient animals,<br />

highly successful predators and scavengers, has been extensively studied in<br />

artificial aquatic environments.<br />

In a typical scenario, intensity of fighting increases in a step-wise<br />

fashion beginning with threat displays upon first contact, followed<br />

by phases of ritualized aggression, restrained use of claws, and in<br />

rare instances ending in periods of unbridled combat. <strong>The</strong> presence<br />

of such a structured behavioral system, combined with an opportunity<br />

to bring <strong>the</strong> analysis to <strong>the</strong> level of individual neurons, thus offers<br />

unique opportunities for exploring fundamental issues of interactions<br />

between aggression, dominance, and amine neurochemistry.<br />

(Huber et al., 2001, p. 272)

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