Mechanisms of impulsive choice: Experiments to explore and models to map the empirical terrain.

Impulsive choice is preference for a smaller-sooner (SS) outcome over a larger-later (LL) outcome when LL choices result in greater reinforcement maximization. Delay discounting is a model of impulsive choice that describes the decaying value of a reinforcer over time, with impulsive choice evident when the empirical choice-delay function is steep. Steep discounting is correlated with multiple diseases and disorders. Thus, understanding the processes underlying impulsive choice is a popular topic for investigation. Experimental research has explored the conditions that moderate impulsive choice, and quantitative models of impulsive choice have been developed that elegantly represent the underlying processes. This review spotlights experimental research in impulsive choice covering human and nonhuman animals across the domains of learning, motivation, and cognition. Contemporary models of delay discounting designed to explain the underlying mechanisms of impulsive choice are discussed. These models focus on potential candidate mechanisms, which include perception, delay and/or reinforcer sensitivity, reinforcement maximization, motivation, and cognitive systems. Although the models collectively explain multiple mechanistic phenomena, there are several cognitive processes, such as attention and working memory, that are overlooked. Future research and model development should focus on bridging the gap between quantitative models and empirical phenomena.

Generalizability of time-based interventions: Effects of choice procedure and smaller-sooner delay.

Interventions exposing rats to delayed-reward contingencies attenuate suboptimal impulsive choices, a preference for a smaller-sooner (SS) over a larger-later (LL) reward. Interventions may potentially improve delay-tolerance, timing of delays, and/or discrimination of reward magnitudes. Generalization from the intervention to impulsive choice under different procedures can provide insights into the processes that underlie the intervention effects. Experiment 1 tested intervention effects on systematic-delay (SYS) and adjusting-delay (ADJ) procedures, predicting that intervention effects would be more effective on the SYS procedure with predictable delays. The ADJ procedure did not benefit significantly from intervention, but the SYS procedure, unexpectedly, showed greater impulsive choices following intervention. Experiment 2 tested whether short (5 s) SS intervention delays may have promoted greater impulsivity in the SYS impulsive choice procedure in Experiment 1. Short SS delays in choice and intervention procedures increased impulsive choices in comparison to longer (10 s) delays. Incongruent SS delays in the intervention/choice procedures resulted in negative intervention effects. The results suggest that short SS delays are detrimental to self-control and that specific temporal information generalizes from the intervention to the SYS choice task, but not the ADJ choice task.

Capuchin monkeys (sometimes) go when they know: Confidence movements in Sapajus apella.

To test for evidence of metacognition in capuchin monkeys (Sapajus apella), we analyzed confidence movements using a paradigm adapted from research with chimpanzees. Capuchin monkeys provide an interesting model species for the comparative assessment of metacognition as they show limited evidence of such cognitive-monitoring processes in a variety of metacognition paradigms. Here, monkeys were presented with a computerized delayed matching-to-sample (DMTS) memory test in one location but were rewarded for correct responses in a separate location. Movements could be made from one location to the other at any time, but movements between a response and reward feedback may reflect confidence in the accuracy of the response. Critically, DMTS tests included occasional "no sample" trials where monkeys' performance was at chance when the trial started without a sample and a 1-s interval to the response options. We predicted that monkeys would (1) perform less accurately (and less confidently) at longer retention intervals, (2) move to the dispenser early more often on trials completed correctly than incorrectly, and (3) show a relation between faster response latency and early movements. Analyses of response times and "go" or "no go" confidence movements before feedback to the reward location suggested that the monkeys were capable of monitoring confidence in their responses. However, their confidence movements were less precise and less flexible than chimpanzees. Overall, this paradigm can reveal potential metacognitive abilities in nonhuman animals that otherwise demonstrate these abilities inconsistently.

Outcome expectancy and suboptimal risky choice in nonhuman primates.

Animals will favor a risky option when a stimulus signaling reward bridges the choice and the outcome. The present experiments investigated signal-induced risky choices and reward-outcome expectations in rhesus and capuchin monkeys. Risky choice was assessed by preference for a large-probabilistic reward over a modest-certain reward. Outcome expectancy was assessed by providing a truncation-response to shorten the delay period. In Experiment 1 both species generally favored the risky option compared to a safe option when the outcomes were signaled and generally shortened the delays except when a signaled-loss stimulus was presented. The use of the delay-truncation response suggested that the monkeys were sensitive to the information conveyed by the stimulus. Experiments 2 and 3 were designed to investigate whether the delay-truncation response used by capuchin monkeys was strategically used reflecting explicit decision-making versus a conditioned response to reward stimuli. A perceptual judgment task was included and the selective use of the delay-truncation response on unsignaled correct trials may suggest the involvement of metacognitive processes. The capuchin monkeys generally truncated the delays except under conditions where reward would not be expected (risky-loss or incorrect-judgment). When the outcomes were unsignaled during the delay some capuchin monkeys were less likely to truncate the delay following an incorrect task response. Overall, the monkeys: (1) made more risky choices when the outcomes were signaled - consistent with gambling-like behavior. (2) selectively truncated the unsignaled delays when rewards could be anticipated (even when metacognitive-like awareness guided anticipation) - suggesting that delay truncation responses reflect explicit outcome expectancy.

Effects of opioid receptor ligands in rats trained to discriminate 22 from 2 hours of food deprivation suggest a lack of opioid involvement in eating for hunger.

It is well accepted that opioids promote feeding for reward. Some studies suggest a potential involvement in hunger-driven intake, but they suffer from the scarcity of methodologies differentiating between factors that intersect eating for pleasure versus energy. Here, we used a unique food deprivation discrimination paradigm to test a hypothesis that, since opioids appear to control feeding reward, injection of opioid agonists would not produce effects akin to 22 h of food deprivation. We trained rats to discriminate between 22 h and 2 h food deprivation in a two-lever, operant discrimination procedure. We tested whether opioid agonists at orexigenic doses produce discriminative stimulus effects similar to 22 h deprivation. We injected DAMGO, DSLET, or orphanin FQ in the paraventricular hypothalamic nucleus (PVN), a site regulating hunger/satiety, and butorphanol subcutaneously (to produce maximum consumption). We assessed the ability of the opioid antagonist, naltrexone, to reduce the discriminative stimulus effects of 22 h deprivation and of the 22 h deprivation-like discriminative stimulus effects of PVN-injected hunger mediator, neuropeptide Y (NPY). In contrast to PVN NPY, centrally or peripherally injected opioid agonists failed to induce discriminative stimuli similar to those of 22 h deprivation. In line with that, naltrexone did not reduce the hunger discriminative stimuli induced by either 22 h deprivation or NPY administration in 2 h food-restricted subjects, even though doses used therein were sufficient to decrease deprivation-induced feeding in a non-operant setting in animals familiar with consequences of 2 h and 22 h deprivation. We conclude that opioids promote feeding for reward rather than in order to replenish lacking energy.

Limited evidence of number-space mapping in rhesus monkeys (Macaca mulatta) and capuchin monkeys (Sapajus apella).

Humans exhibit evidence of a mental number line that suggests a left-to-right, or sometimes right-to-left, representation of smaller to larger numbers. The Spatial Numerical Association of Response Codes (SNARC) effect is one example of this mental number line and has been investigated extensively in humans. Less research has been done with animals, and results have been inconclusive. Rugani, Vallortigara, Priftis, and Regolin (2015) found that young chicks showed a bias to respond to small quantities presented to their left and large quantities presented to their right when forced to move toward those stimuli to gain food reward. We replicated this design with rhesus macaques and capuchin monkeys using a computerized task, but we did not find this outcome. We also trained monkeys to choose between 2 arrays of dots, and then assessed biases in terms of choice location and response latency on trials with a numerical difference and on trials with equal numbers of items in both sets. There was no evidence of SNARC-like effects in equal trials, although when arrays differed in number, 12 of 19 monkeys showed differential performance depending on whether the smaller array was at the left or at the right onscreen. These results indicate that SNARC-like effects may not emerge in all contexts and may not be phylogenetically widespread. More effort is needed to broaden the number of species assessed and match other methods that are used with human participants so that we can better define the presence and extent of such effects. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Task switching in rhesus macaques (Macaca mulatta) and tufted capuchin monkeys (Cebus apella) during computerized categorization tasks.

The present experiments extended to monkeys a previously used abstract categorization procedure (Castro & Wasserman, 2016) where pigeons had categorized arrays of clipart icons based upon two task rules: the number of clipart objects in the array or the variability of objects in the array. Experiment 1 replicated Castro and Wasserman by using capuchin monkeys and rhesus monkeys and reported that monkeys' performances were similar to pigeons' in terms of acquisition, pattern of errors, and the absence of switch costs. Furthermore, monkeys' insensitivity to the added irrelevant information suggested that an associative (rather than rule-based) categorization mechanism was dominant. Experiment 2 was conducted to include categorization cue reversals to determine (a) whether the monkeys would quickly adapt to the reversals and inhibit interference from a prereversal task rule (consistent with a rule-based mechanism) and (b) whether the latency to make a response prior to a correct or incorrect outcome was informative about the presence of a cognitive mechanism. The cue reassignment produced profound and long-lasting performance deficits, and a long reacquisition phase suggested the involvement of associative learning processes; however, monkeys also displayed longer latencies to choose prior to correct responses on challenging trials, suggesting the involvement of nonassociative processes. Together these performances suggest a mix of associative and cognitive-control processes governing monkey categorization judgments. (PsycINFO Database Record

Gambling in rhesus macaques (Macaca mulatta): The effect of cues signaling risky choice outcomes.

Preference for a larger-variable "risky" option over a smaller-reliable "safe" option often depends upon the likelihood that the risky option will deliver a sufficiently sized reward to have an equivalent or superior expected value. However, preference for the risky option has been shown to increase under conditions where informative stimuli signaling the outcome of a risky choice is included between the choice and the outcome and this risk-prone preference persists even when the risky option has a lower expected value than the alternative safe option. In the present study, rhesus macaques chose between a risky option and a safe option across two experimental phases to determine whether the outcome signal affected the degree of preference for the risky option. Overall, six out of seven macaques showed a greater preference for the risky option in the signaled condition than in the unsignaled condition. The macaques' risky choices were sensitive to the expected value of the risky option and the signaled condition produced a general increase in risky choices independently of the expected value of the risky outcome. Overall, these results are consistent with those obtained with other animals, and this may relate to a process where animals show a biased preference for "good news." This process may model some of the relevant factors that explain the psychology of gambling in humans.

Simple tone discriminations are disrupted following experimental frontal traumatic brain injury in rats.

PRIMARY OBJECTIVE: To assess cognitive deficits in a rat model of brain injury. RESEARCH DESIGN: Cognitive deficits are some of the most pervasive and enduring symptoms of frontal traumatic brain injury (TBI) in human patients. In animal models, the assessment of cognitive deficits from TBI has primarily been limited to tests of spatial learning. Recently, simple discrimination performance has been shown to be sensitive to frontal brain damage. The current study provides a detailed characterization of deficits in a two-choice tone discrimination following a bilateral frontal controlled cortical impact injury. METHODS AND PROCEDURES: Rats were trained on a two-tone discrimination task in a standard operant chamber, then either a frontal brain injury was delivered or sham procedures performed. Following recovery, they were re-tested on the discrimination task and then tested on a reversal of the discrimination. MAIN OUTCOMES AND RESULTS: Frontal injury caused substantial deficits in responding and discrimination accuracy as well as an increase in side bias. CONCLUSIONS: Based on the outcomes seen in this study, discrimination and other operant tasks may provide a sensitive tool to assess the effect of therapeutic agents on cognitive deficits in animal models, which could lead to improved characterization of deficits and yield an improved assessment tool to aid in drug discovery.

Effects of sibutramine and rimonabant in rats trained to discriminate between 22- and 2-h food deprivation.

OBJECTIVE: The objective of the study was to evaluate whether sibutramine and rimonabant, drugs that decrease food intake in human and non-human animals, affect the discriminative stimulus effects associated with acute food deprivation ("hunger"). MATERIALS AND METHODS: Rats were trained to discriminate between 22- and 2-h food deprivation in a two-lever choice procedure. After rats acquired the discrimination, subjects were food-restricted for 22 h and administered with sibutramine (0.32-10 mg/kg, p.o.) or rimonabant (0.32-10 mg/kg, s.c.) before a generalization test session. RESULTS: Sibutramine (3.2 mg/kg) produced significant decreases in 22-h deprivation-appropriate responding, response rates (resulting in lever pressing rates similar to those following 2-h food deprivation), and food intake measured 1 h after the generalization test. A larger sibutramine dose eliminated responding and significantly reduced food intake. Rimonabant did not alter the discriminative stimulus effects of 22-h food deprivation, but rimonabant did significantly reduce both response rates and food intake. CONCLUSION: Sibutramine appears to decrease food intake by reducing hunger sensations associated with food deprivation. In contrast, rimonabant does not alter the discrimination of acute food deprivation. The use of food-deprivation discrimination techniques may be useful in identifying the role of specific neuroactive compounds in eating stimulated by a sense of hunger and may aid in medication development for more effective treatments for obesity and other eating-related conditions.