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Who Needs Emotions? The Brain Meets the Robot

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356 conclusions<br />

<strong>The</strong> Motivated Toad<br />

Earlier, we examined <strong>the</strong> basic, overlapping circuitry for fleeing and feeding<br />

in <strong>the</strong> toad but did not show how this circuitry could be modulated by<br />

motivational systems. Toads can be conditioned to treat objects that normally<br />

elicit escape behavior as prey. Toads which are allowed to eat mealworms<br />

out of an experimenter’s hand can be conditioned to respond to <strong>the</strong><br />

moving hand alone (Brzoska & Schneider, 1978). Heatwole and Heatwole<br />

(1968) showed that <strong>the</strong> upper size threshold for acceptable prey increases<br />

with long-term food deprivation while <strong>the</strong> lower size threshold remains<br />

constant. In spring, prey-catching behavior decreases or fails to occur. Indeed,<br />

prey recognition is “exchanged” for female recognition, introducing<br />

mating behavior. Moving females release orienting, approaching, and clasping<br />

behaviors in <strong>the</strong> male (Heusser, 1960; Kondrashev, 1976). In many species,<br />

males will attempt to clasp practically any moving object including o<strong>the</strong>r<br />

males during mating season (Wells, 1977).<br />

Betts (1989) carried forward <strong>the</strong> modeling of tectal–pretectal interactions<br />

(reviewed in Arbib, 1987) to include <strong>the</strong> effects of motivation. <strong>The</strong> basic idea<br />

is that perceptual schemas can be modulated by broadcast signals for drive<br />

levels such as those for feeding or mating, thus shifting <strong>the</strong> balance of behavior.<br />

For example, standard models of prey catching address <strong>the</strong> finding that<br />

ablation of <strong>the</strong> pretectum results in disinhibition of prey catching, with animals<br />

snapping at objects much larger than normal prey (Ewert, 1984); modulating<br />

<strong>the</strong> level of pretectal inhibition can thus shift <strong>the</strong> balance between feeding<br />

and fleeing. Betts (1989) fur<strong>the</strong>r suggested parallels between <strong>the</strong> effects of<br />

pretectal ablation and <strong>the</strong> conditioning results and changes that occur during<br />

<strong>the</strong> mating season. T5 neurons in <strong>the</strong> tectum have a variety of responses, including<br />

those which we classify as prey recognition. Betts modeled T5 neuron<br />

function by distinguishing <strong>the</strong> T5 base of T5 cells within <strong>the</strong> tectum from <strong>the</strong><br />

pretectal inhibition which modulates it. A T5 neuron <strong>the</strong>n has <strong>the</strong> potential<br />

to be, for example, ei<strong>the</strong>r a prey or mate feature detector depending on this<br />

inhibition. Betts suggests that <strong>the</strong> subclasses of T5 neurons with different<br />

detector properties should be regarded as more or less stable states of a modulated<br />

system capable of adaptability and changeability.<br />

In summary, <strong>the</strong> frog has basic systems for feeding, fleeing, and mating;<br />

but we can distinguish circuitry that provides basic “subroutines” from circuitry<br />

that uses motivational factors to bias <strong>the</strong>ir deployment. This separates<br />

<strong>the</strong> motivation from <strong>the</strong> behavior. From my point of view, fear is not a behavior,<br />

such as freezing, but ra<strong>the</strong>r a process that biases <strong>the</strong> system to be<br />

more likely to emit such a behavior. Freezing is one of <strong>the</strong> many possible<br />

behaviors that express fear. Which one is chosen depends on learning, species,<br />

and all kinds of o<strong>the</strong>r bias. Part of <strong>the</strong> task of a model of emotion and

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