Assessment of positive emotions in animals to improve their welfare.

Rewards are crucial objects that induce learning, approach behavior, choices, and emotions. Whereas emotions are difficult to investigate in animals, the learning function is mediated by neuronal reward prediction error signals which implement basic constructs of reinforcement learning theory. These signals are found in dopamine neurons, which emit a global reward signal to striatum and frontal cortex, and in specific neurons in striatum, amygdala, and frontal cortex projecting to select neuronal populations. The approach and choice functions involve subjective value, which is objectively assessed by behavioral choices eliciting internal, subjective reward preferences. Utility is the formal mathematical characterization of subjective value and a prime decision variable in economic choice theory. It is coded as utility prediction error by phasic dopamine responses. Utility can incorporate various influences, including risk, delay, effort, and social interaction. Appropriate for formal decision mechanisms, rewards are coded as object value, action value, difference value, and chosen value by specific neurons. Although all reward, reinforcement, and decision variables are theoretical constructs, their neuronal signals constitute measurable physical implementations and as such confirm the validity of these concepts. The neuronal reward signals provide guidance for behavior while constraining the free will to act.

Neuronal Reward and Decision Signals: From Theories to Data

Experiments on decision making by humans show that the choices that we make can be very labile. The magnitude of our preferences, and even our rank ordering of options, can vary according to the number and type of alternatives available for comparison. This apparent irrationality has been argued to result from our use of decision heuristics that have evolved to enable us to choose quickly and efficiently between options differing in multiple attributes. Here, we argue that, because there is also selective pressure for animals to make mating decisions quickly, and because potential mates also differ in multiple attributes, similar decision heuristics might have evolved for mate choice. Following this reasoning, the attractiveness of a given mate will depend on the others with whom he or she is being compared, rather than being an absolute function of his or her underlying quality. We describe some of the ramifications of such comparative evaluation, and argue that it could offer new insights into some of the biggest outstanding problems in mate choice and sexual selection.

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Comparative evaluation and its implications for mate choice.

Potential vulnerabilities of neuronal reward, risk, and decision mechanisms to addictive drugs

Stimuli associated with less effort or with shorter delays to reinforcement are generally preferred over those associated with greater effort or longer delays to reinforcement. However, the opposite appears to be true of stimuli thatfollow greater effort or longer delays. In training, a simple simultaneous discrimination followed a single peck to an initial stimulus (S+FR1 S−FR1) and a different simple simultaneous discrimination followed 20 pecks to the initial stimulus (S+FR20 S−FR20). On test trials, pigeons preferred S+FR20 over S+FR1 and S−FR20 over S−FR1. These data support the view that the state of the animal immediately prior to presentation of the discrimination affects the value of the reinforcement that follows it. This contrast effect is analogous to effects that when they occur in humans have been attributed to more complex cognitive and social factors.

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M. J. Zigler

Papers

Qualitative Differences between Binocular and Uniocular Impressions