The evolution of self-control.

Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.

I can time with a little help from my friends: effect of social enrichment on timing processes in Pigeons (Columba livia).

There is evidence that impulsive decision-making is associated with errors in timing. However, there has been little attempt to identify the putative mechanism responsible for impulsive animals' timing errors. One means of manipulating impulsivity in non-human animals is providing different levels of access to conspecifics. These preclinical models have revealed that social isolation increases impulsive responding across a wide range of tasks. The goal of the present study was to determine whether social isolation modulates time perception in pigeons by inducing more variability or a bias to underestimate the passage of time in temporal judgments. A temporal bisection task was used to characterize time perception. One group of pigeons performed the bisection following social enrichment, and the remaining half of the pigeons were tested following social isolation. Results revealed pigeons in the social isolation condition categorized a temporal stimulus sample as "long" at shorter durations than pigeons in the social enrichment condition. These data highlight the mechanism(s) thought to underlie timing-based interventions aimed at reducing impulsivity in humans. Future work should consider whether impulsivity is produced by misperceptions of time or a reduced threshold for a response.

Who are the real bird brains? Qualitative differences in behavioral flexibility between dogs (Canis familiaris) and pigeons (Columba livia).

Pigeons given a simultaneous spatial discrimination reversal, in which a single reversal occurs at the midpoint of each session, consistently show anticipation prior to the reversal as well as perseveration after the reversal, suggesting that they use a less effective cue (time or trial number into the session) than what would be optimal to maximize reinforcement (local feedback from the most recent trials). In contrast, rats (Rattus norvegicus) and humans show near-optimal reversal learning on this task. To determine whether this is a general characteristic of mammals, in the present research, pigeons (Columba livia) and dogs (Canis familiaris) were tested with a simultaneous spatial discrimination mid-session reversal. Overall, dogs performed the task more poorly than pigeons. Interestingly, both pigeons and dogs employed what resembled a timing strategy. However, dogs showed greater perseverative errors, suggesting that they may have relatively poorer working memory and inhibitory control with this task. The greater efficiency shown by pigeons with this task suggests they are better able to time and use the feedback from their preceding choice as the basis of their future choice, highlighting what may be a qualitative difference between the species.

Alcohol cues impair learning inhibitory signals in beer drinkers.

BACKGROUND: Models of drug addiction emphasize the reciprocal influence of incentive-motivational properties of drug-related cues and poor impulse control resulting in drug use. Recent studies have shown that alcohol-related cues can impair response inhibition. What is unknown is whether these cues also disrupt learning of inhibitory associations. METHODS: Participants performed a conditioned inhibition (CI) task and were required to learn that a neutral image was a conditioned inhibitor when presented in the context of either an alcohol image intended to draw their attention away from the to-be-trained inhibitor, or a control condition in which the alcohol image was absent. After training, subjects in each condition rated the likelihood that the neutral image would signal the outcome. Eye tracking was used to verify that attention to the neutral image was in fact reduced when the alcohol image was present. RESULTS: Compared with controls those trained in the alcohol image condition reported a greater likelihood that the presence of the inhibitor would be followed by the outcome and thus were less able to acquire CI. Measures of eye tracking verified that attention to the alcohol cue was associated with this maladaptive behavior. CONCLUSIONS: When alcohol cues are present, there is a reduced ability to learn that such information is irrelevant to an outcome, and this impairs ones' ability to inhibit perseveration of a response. This has implications for persistence of a drinking episode.

Transitive inference by pigeons: does the geometric presentation of the stimuli make a difference?

In studies of transitive inference (TI), nonhuman animals are typically trained with the following 5-term task: A+B-, B+C-, C+D-, D+E- where the letters stand for arbitrary stimuli and [+] indicates that choice is reinforced and [-] indicates that choice is not reinforced. A TI effect is found when, given the untrained test pair BD, subjects choose B. TI effects have been found in many nonhuman species. Although reinforcement history has been posited as an account of the TI effect, it has failed to account for a variety of conditions under which TI effects have been found. A more cognitive account of TI is that organisms are able to form a representation of the series (A>B>C>D>E). In support of this hypothesis, Roberts and Phelps (Psychol Sci 5:368-374, 1994) found that presentation of the pairs of stimuli in a linear arrangement facilitated TI performance by rats, whereas presentation of the pairs of stimuli in a circular arrangement did not. Using methods adapted from Roberts and Phelps, we trained pigeons on either a linear or a circular arrangement of stimuli with the 5-term task. Results indicated that on the BD test pair, pigeons trained with a circular arrangement did not differ from those trained with a linear arrangement. Furthermore, we found that memory for training pairs was variable and was highly correlated with degree of TI. The results suggest that regardless of how pigeons are able to represent the stimuli, choice was not affected by the spatial arrangement of the stimuli during training.

Midsession reversals with pigeons: visual versus spatial discriminations and the intertrial interval.

Discrimination reversal learning has been used as a measure of species flexibility in dealing with changes in reinforcement contingency. In the simultaneous-discrimination, midsession-reversal task, one stimulus (S1) is correct for the first half of the session, and the other stimulus (S2) is correct for the second half. After training, pigeons show a curious pattern of choices: They begin to respond to S2 well before the reversal point (i.e., they make anticipatory errors), and they continue to respond to S1 well after the reversal (i.e., they make perseverative errors). That is, pigeons appear to be using the passage of time or the number of trials into the session as a cue to reverse, and are less sensitive to the feedback at the point of reversal. To determine whether the nature of the discrimination or a failure of memory for the stimulus chosen on the preceding trial contributed to the pigeons' less-than-optimal performance, we manipulated the nature of the discrimination (spatial or visual) and the duration of the intertrial interval (5.0 or 1.5 s), in order to determine the conditions under which pigeons would show efficient reversal learning. The major finding was that only when the discrimination was spatial and the intertrial interval was short did the pigeons perform optimally.

Environmental enrichment affects suboptimal, risky, gambling-like choice by pigeons.

Pigeons prefer a risky option with a low probability of a high payoff over a less risky option that results in more food. This finding is analogous to suboptimal human monetary gambling because in both cases there appears to be an overemphasis of the occurrence of the winning event and an underemphasis of the losing event. In the present research, we found that pigeons that were exposed to an enriched environment (a large cage with three other pigeons for 4 h a day) were less likely to show this suboptimal choice behavior compared with typically housed laboratory pigeons in a control group. These results have implications for the mechanisms underlying suboptimal choice by humans (e.g., problem gamblers), and they suggest that a enriched environment may allow for enhanced self-control.

Midsession reversal learning: why do pigeons anticipate and perseverate?

Past research has shown that when given a simultaneous visual-discrimination midsession reversal task, pigeons typically anticipate the reversal well before it occurs and perseverate after it occurs. It appears that they use the estimation of time (or trial number) into the session, rather than (or in addition to) the more reliable cue, the outcome from the previous trial (i.e., a win-stay/lose-shift response rule), to determine which stimulus they should choose. In the present research, we investigated several variables that we thought might encourage pigeons to use a more efficient response strategy. In Experiment 1, we used a treadle-stepping response, rather than key pecking, to test the hypothesis that reflexive key pecking may have biased pigeons to estimate the time (or trial number) into the session at which the reversal would occur. In Experiment 2, we attempted to make the point of reversal in the session more salient by inserting irrelevant trials with stimuli different from the original discriminative stimuli, and for a separate group, we added a 5-s time-out penalty following incorrect choices. The use of a treadle-stepping response did not improve reversal performance, and although we found some improvement in reversal performance when the reversal was signaled and when errors resulted in a time-out, we found little evidence for performance that approached the win-stay/lose-shift accuracy shown by rats.

Decision making by humans in a behavioral task: do humans, like pigeons, show suboptimal choice?

Consistent with human gambling behavior but contrary to optimal foraging theory, pigeons show a strong preference for an alternative with low probability and high payoff (a gambling-like alternative) over an alternative with a greater net payoff (Zentall & Stagner, Proceedings of the Royal Society B, 278, 1203-1208, 2011). In the present research, we asked whether humans would show suboptimal choice on a task involving choices with probabilities similar to those for pigeons. In Experiment 1, when we selected participants on the basis of their self-reported gambling activities, we found a significantly greater choice of the alternative involving low probability and high payoff (gambling-like alternative) than for a group that reported an absence of gambling activity. In Experiment 2, we found that when the inhibiting abilities of typical humans were impaired by a self-regulatory depletion manipulation, they were more likely to choose the gambling-like alternative. Taken together, the results suggest that this task is suitable for the comparative study of suboptimal decision-making behavior and the mechanisms that underlie it.

Extended treatment for cigarette smoking cessation: a randomized control trial.

AIM: To test the potential benefit of extending cognitive-behavioral therapy (CBT) relative to not extending CBT on long-term abstinence from smoking. DESIGN: Two-group parallel randomized controlled trial. Patients were randomized to receive non-extended CBT (n = 111) or extended CBT (n = 112) following a 26-week open-label treatment. SETTING: Community clinic in the United States. PARTICIPANTS: A total of 219 smokers (mean age: 43 years; mean cigarettes/day: 18). INTERVENTION: All participants received 10 weeks of combined CBT + bupropion sustained release (bupropion SR) + nicotine patch and were continued on CBT and either no medications if abstinent, continued bupropion + nicotine replacement therapy (NRT) if increased craving or depression scores, or varenicline if still smoking at 10 weeks. Half the participants were randomized at 26 weeks to extended CBT (E-CBT) to week 48 and half to non-extended CBT (no additional CBT sessions). MEASUREMENTS: The primary outcome was expired CO-confirmed, 7-day point-prevalence (PP) at 52- and 104-week follow-up. Analyses were based on intention-to-treat. FINDINGS: PP abstinence rates at the 52-week follow-up were comparable across non-extended CBT (40%) and E-CBT (39%) groups [odds ratio (OR) = 0.99; 95% confidence interval (CI) = 0.55, 1.78]. A similar pattern was observed across non-extended CBT (39%) and E-CBT (33%) groups at the 104-week follow-up (OR = 0.79; 95% CI= 0.44, 1.40). CONCLUSION: Prolonging cognitive-behavioral therapy from 26 to 48 weeks does not appear to improve long-term abstinence from smoking.

Drivers who self-estimate lower blood alcohol concentrations are riskier drivers after drinking.

RATIONALE: Alcohol increases the tendency for risky driving in some individuals but not others. Little is known about the factors underlying this individual difference. Studies find that those who underestimate their blood alcohol concentration (BAC) following a dose of alcohol tend to be more impulsive and report greater willingness to drive after drinking than those who estimate their BACs to be greater than their actual BAC. BAC underestimation could contribute to risky driving behavior following alcohol as such drivers might perceive little impairment in their driving ability and thus no need for caution. OBJECTIVES: This study was designed to test the relationship between drivers' BAC estimations following a dose of alcohol or a placebo and the degree of risky driving they displayed during a simulated driving test. METHODS: Forty adult drivers performed a simulated driving test and estimated their blood alcohol concentration after receiving a dose of alcohol (0.65 g/kg for men and 0.56 g/kg for women) or a placebo. RESULTS: Alcohol increased risk-taking and impaired driving skill. Those who estimated their BAC to be lower were the riskiest drivers following both alcohol and placebo. CONCLUSIONS: The tendency to estimate lower BACs could support a series of high-risk decisions, regardless of one's actual BAC. This could include the decision to drive after drinking.

Simulated driving performance under alcohol: Effects on driver-risk versus driver-skill.

BACKGROUND: Those who place their vehicles closer to others on the roadway are said to have high risk acceptance, and this contributes to motor vehicle crashes. However, the effect of alcohol on this risky driving behavior is understudied. Behavioral mechanisms that contribute to risky driving are also not well understood. Further, whether increased risk-taking behavior in a driver co-occurs with pronounced impairment in the driver's skill is unknown. METHODS: The study examined the effect of alcohol on driver risk and skill and whether riskier drivers were also those who showed high skill impairment. The relationship between driving behavior and inhibitory control was also tested. Participants completed two driving simulations. In the first drive test, risky driving was encouraged and in the second test, skill-based performance was emphasized. The cued go/no-go task provided a measure inhibitory control. Tests were completed under a 0.65g/kg alcohol and 0.0g/kg (placebo) dose of alcohol. RESULTS: Alcohol impaired a measure of driving skill and increased driver risk taking. It was also found that riskier drivers were not necessarily those who showed the greatest impairments in skill. Poorer inhibitory control was associated with greater driver risk in the sober state. CONCLUSIONS: Alcohol-induced risk-taking behaviors can be dissociable from impairing effects on driver skill and poor inhibitory control is selectively related to risky driving. As such, a distinction between driver risk and driver skill needs to be made in the investigation of problems concerning DUI-related accidents and fatalities in future research.

Self-regulatory depletion in dogs: insulin release is not necessary for the replenishment of persistence.

It has been hypothesized that self-control is constrained by a limited energy resource that can be depleted through exertion. Once depleted, this resource can be replenished by the consumption or even the taste of glucose. For example, the need to inhibit reduces subsequent persistence at problem solving by humans and dogs, an effect that is not observed when a glucose drink (but not a placebo) is administered following initial inhibition. The mechanism for replenishment by glucose is currently unknown. Energy transfer is not necessary, although insulin secretion may be involved. This possibility was investigated in the current study by having dogs exert self-control (sit-stay) and subsequently giving them (1) glucose that causes the release of insulin, (2) fructose that does not result in the release of insulin nor does it affect glucose levels (but it is a carbohydrate), or (3) a calorie-free drink. Persistence measures indicated that both glucose and fructose replenished canine persistence, whereas the calorie-free drink did not. These results indicate that insulin release is probably not necessary for the replenishment that is presumed to be responsible for the increase in persistence. This article is part of a Special Issue entitled: Canine Behavior.

Six-term transitive inference with pigeons: successive-pair training followed by mixed-pair training.

In nonhuman animals, the transitive inference (TI) task typically involves training a series of four simultaneous discriminations involving, for example, arbitrary colors in which choice of one stimulus in each pair is reinforced [+] and choice of the other color is nonreinforced [-]. This can be represented as A+B-, B+C-, C+D-, D+E- and can be conceptualized as a series of linear relationships: A > B > C > D > E. After training, animals are tested on the untrained non-endpoint pair, BD. Preference for B over D is taken as evidence of TI and occurs because B is greater than D in the implied series. In the present study we trained pigeons using a novel training procedure-a hybrid of successive pair training (training one pair at a time) and mixed-pair training (training all pairs at once)-designed to overcome some of the limitations of earlier procedures. Using this hybrid procedure, we trained five premise pairs (A+B-, B+C-, C+D-, D+E-, and E+F-) which allowed us to test three untrained non-endpoint pairs (BD, CE, and BE). A significant TI effect was found for most subjects on at least two out of three test pairs. Different theories of TI are discussed. The results suggest that this hybrid training is an efficient procedure for establishing mixed-pair acquisition and a TI effect.

Suboptimal choice by pigeons may result from the diminishing effect of nonreinforcement.

Pigeons prefer an alternative that provides them with a stimulus 20% of the time that predicts 10 pellets of food and a different stimulus 80% of the time that predicts 0 pellets, over an alternative that provides them with a stimulus that always predicts 3 pellets of food, even though the preferred alternative provides them with considerably less food. It appears that the stimulus that predicts 10 pellets acts as a strong conditioned reinforcer, despite the fact that the stimulus that predicts 0 pellets occurs 4 times as often. In the present research, we tested the hypothesis that early in training conditioned inhibition develops to the 0-pellet stimulus, but later in training it dissipates. We trained pigeons with a hue as the 10-pellet stimulus and a vertical line as the 0-pellet stimulus. To assess the inhibitory value of the vertical line, we compared responding to the 10-pellet hue with responding to the compound of the 10-pellet hue and the vertical line early in training and once again late in training, using both a within-subject design (Experiment 1) and a between-groups design (Experiment 2). We found that there was a significant reduction in inhibition between the early test (when pigeons chose optimally) and late test (when choice was suboptimal). Thus, the increase in suboptimal choice may result from the decline in inhibition to the 0-pellet stimulus. Implications for human gambling behavior are considered.

Impulsivity affects suboptimal gambling-like choice by pigeons.

Pigeons prefer a low-probability, high-payoff but suboptimal alternative over a reliable low-payoff optimal alternative (i.e., one that results in more food). This finding is analogous to suboptimal human monetary gambling because in both cases there appears to be an overemphasis of the occurrence of the winning event (a jackpot) and an underemphasis of losing events. In the present research we found that pigeons chose suboptimally to the degree that they were impulsive as indexed by the steeper slope of the hyperbolic delay-discounting function (i.e., the shorter the delay they would accept in a smaller-sooner/larger-later procedure). These correlational findings have implications for the mechanisms underlying suboptimal choice by humans (e.g., problem gamblers) and they suggest that high baseline levels of impulsivity can enhance acquisition of a gambling habit.

Less means more for pigeons but not always.

When humans are asked to judge the value of a set of objects of excellent quality, they often give this set higher value than those same objects with the addition of some of lesser quality. This is an example of the affect heuristic, often referred to as the less-is-more effect. Monkeys and dogs, too, have shown this suboptimal effect. But in the present experiments, normally hungry pigeons chose optimally: a preferred food plus a less-preferred food over a more-preferred food alone. In Experiment 2, however, pigeons on a less-restricted diet showed the suboptimal less-is-more effect. Choice on control trials indicated that the effect did not result from the novelty of two food items versus one. The effect in the less-food-restricted pigeons appears to result from the devaluation of the combination of the food items by the presence of the less-preferred food item. The reversal of the effect under greater food restriction may occur because, as motivation increases, the value of the less-preferred food increases faster than the value of the more-preferred food, thus decreasing the difference in value between the two foods.

Pigeons show near-optimal win-stay/lose-shift performance on a simultaneous-discrimination, midsession reversal task with short intertrial intervals.

Discrimination reversal tasks have been used as a measure of species flexibility in dealing with changes in reinforcement contingency. The simultaneous-discrimination, midsession reversal task is one in which one stimulus (S1) is correct for the first 40 trials of an 80-trial session and the other stimulus (S2) is correct for the remaining trials. After many sessions of training with this task, pigeons show a curious pattern of choices. They begin to respond to S2 well before the reversal point (they make anticipatory errors) and they continue to respond to S1 well after the reversal (they make perseverative errors). That is, they appear to be using the passage of time or number of trials into the session as a cue to reverse. We tested the hypothesis that these errors resulted in part from a memory deficit (the inability to remember over the intertrial interval, ITI, both the choice on the preceding trial and the outcome of that choice) by manipulating the duration of the ITI (1.5, 5, and 10 s). We found support for the hypothesis as pigeons with a short 1.5-s ITI showed close to optimal win-stay/lose-shift accuracy.

Pigeons prefer discriminative stimuli independently of the overall probability of reinforcement and of the number of presentations of the conditioned reinforcer.

When pigeons are given a choice between two alternatives, one leading to a stimulus 20% of the time that always signals reinforcement (S+) or another stimulus 80% of the time that signals the absence of reinforcement (S-) and the other alternative leading to one of two stimuli each signaling reinforcement 50% of the time, the 20% reinforcement alternative is preferred although it provides only 40% as much reinforcement. In Phase 1 of the present experiment, we tested the hypothesis that pigeons compare the S+ associated with each alternative and ignore the S- by giving them a choice between two pairs of discriminative stimuli (20% S+, 80% S- and 50% S+, 50% S-). Reinforcement theory suggests that the alternative associated with more reinforcement should be preferred but the pigeons showed indifference. In Phase 2, the pigeons were divided into two groups. For one group, the discriminative function was removed from the 50% reinforcement alternative and a strong preference for the 20% reinforcement alternative was found. For the other group, the discriminative function was removed from both alternatives and a strong preference was found for the 50% reinforcement alternative. Thus, the indifference found in Phase 1 was not due to the absence of discriminability of the differential reinforcement associated with the two alternatives (20% vs. 50% reinforcement); rather, the indifference can be attributed to the pigeons' insensitivity to the differential frequency of the two S+ and two S- stimuli. The relevance to human gambling behavior is discussed.

Hungry pigeons make suboptimal choices, less hungry pigeons do not.

Hungry animals will often choose suboptimally by being attracted to reliable signals for food that occur infrequently (they gamble) over less reliable signals for food that occur more often. That is, pigeons prefer an option that 50 % of the time provides them with a reliable signal for the appearance of food but 50 % of the time provides them with a reliable signal for the absence of food (overall 50 % reinforcement) over an alternative that always provides them with a signal for the appearance of food 75 % of the time (overall 75 % reinforcement). The pigeons appear to choose impulsively for the possibility of obtaining the reliable signal for reinforcement. There is evidence that greater hunger is associated with greater impulsivity. We tested the hypothesis that if the pigeons were less hungry, they would be less impulsive and, thus, would choose more optimally (i.e., on the basis of the overall probability of reinforcement). We found that hungry pigeons choose the 50 % reinforcement alternative suboptimally but less hungry pigeons prefer the more optimal 75 % reinforcement. Paradoxically, pigeons that needed the food more received less of it. These findings have implications for how level of motivation may also affect human suboptimal choice (e.g., purchase of lottery tickets and playing slot machines).