Sources of Reinforcement: Motivation, Opponent Process, Incentive Salience
Changes in the motivation for drugs and natural rewards are a key component of addiction (Table 1). Early work by Wikler (1952) stressed the function of changes in drive states associated with dependence (herein referred to as addiction. Subjects described withdrawal changes as a ?hunger' or primary need and the effects of morphine on such a state as ?satiation' or gratification of the primary need (Wikler, 1952). Although Wikler argued that positive reinforcement was retained even in heavily dependent subjects (eg, thrill of the intravenous opioid injection), addiction produced a new source of gratification, that of negative reinforcement (Table 1).
The concept of motivation was linked inextricably with hedonic, affective, or emotional states in the transition to addiction by Solomon's opponent process theory of motivation. Solomon and Corbit (1974) postulated that hedonic, affective, or emotional states, once initiated, are automatically modulated by the central nervous system with mechanisms that reduce the intensity of hedonic feelings. Positive hedonic responses in drug use occur shortly after presentation of a stimulus, correlate closely with the intensity, quality, and duration of the reinforcer, and show tolerance and affective or hedonic withdrawal (abstinence). In contrast, negative hedonic responses follow the positive hedonic responses, are sluggish in onset, slow to build up to an asymptote, slow to decay, and get larger with repeated exposure. The role of opponent processes begins early in drug-taking, reflects changes in the brain reward and stress systems, and later forms one of the major motivations for compulsivity in drug-taking in the form of a motivational withdrawal syndrome.
In this formulation, manifestation of a withdrawal syndrome after removal of chronic drug administration, either acute or protracted, is defined in terms of motivational aspects of dependence such as the emergence of a negative emotional state (eg, dysphoria, anxiety, irritability) when access to the drug is prevented (Koob and Le Moal, 2001), rather than on the physical signs of dependence, which tend to be of short duration. Indeed, some have argued that the development of such a negative affective state can define dependence relative to addiction (Russell, 1976; Baker et al, 1987) and that such a negative affective state contributes to compulsivity through negative reinforcement mechanisms (Koob and Le Moal, 2005).
Another conceptualization of the motivational changes associated with addiction is derived from early work on conditioned reinforcement, incentive motivation, behavioral sensitization, and maladaptive stimulus?response learning, all of which are subsumed under the motivational conceptualization of incentive salience. Drugs are hypothesized to usurp systems in the brain that are put in place to direct animals to stimuli with salience for preservation of the species. The incentive salience hypothesis has significant heuristic value as a common element of drug addiction because it narrows the focus to drug-seeking at the expense of natural rewards. The clinical observation that individuals with substance use disorders have an unusual focus on drug-seeking to the exclusion of natural rewards fits the incentive salience view.
The increase in incentive salience produced by psychostimulant drugs has early roots in the facilitation of conditioned reinforcement and drug-seeking (Robbins, 1976; Hill, 1970). Here, drug-seeking is controlled by a succession of drug-associated discriminative stimuli that can also function as conditioned reinforcers when presented as a consequence of instrumental responses (Everitt et al, 2008). Many have argued that by means of associative learning, the enhanced incentive salience state becomes oriented specifically toward drug-related stimuli, leading to escalating compulsion for seeking and taking drugs (Hyman et al, 2006; Kalivas and Volkow, 2005). The underlying activation of neural structures involved in maintaining the incentive salience state persists, making addicts vulnerable to long-term relapse.
Another view of incentive salience involved behavioral sensitization, usually measured as increased locomotor responses to repeated administration of a drug. The behavioral sensitization paradigm has provided a major impetus to exploring not only the neurocircuitry of addiction but also a model of the neuroplasticity that may occur during the transition from drug use to addiction. Here, a shift in an incentive salience state, described as ?wanting' linked to compulsive use, as opposed to ?liking' linked to hedonic responses, was hypothesized to be progressively increased by repeated exposure to drugs of abuse (Robinson and Berridge, 1993).