Jennifer R. Laude

The Evolution of Self-Control

Significance Although scientists have identified surprising cognitive flexibility in animals and potentially unique features of human psychology, we know less about the selective forces that favor cognitive evolution, or the proximate biological mechanisms underlying this process. We tested 36 species in two problem-solving tasks measuring self-control and evaluated the leading hypotheses regarding how and why cognition evolves. Across species, differences in absolute (not relative) brain volume best predicted performance on these tasks. Within primates, dietary breadth also predicted cognitive performance, whereas social group size did not. These results suggest that increases in absolute brain size provided the biological foundation for evolutionary increases in self-control, and implicate species differences in feeding ecology as a potential selective pressure favoring these skills. 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.

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.

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.

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.

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.

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.

Do Pigeons Gamble? I Wouldn’t Bet Against It

Human gambling generally involves suboptimal choice because the expected return is usually less than the investment. We have found that animals, too, choose suboptimally under similar choice conditions. Pigeons, like human gamblers, show an impaired ability to objectively assess overall probabilities and amounts of reinforcement when a rare, high-value outcome (analogous to a jackpot in human gambling) is presented in the context of more frequently occurring losses. More specifically, pigeons prefer a low-probability, high-reward outcome over a guaranteed low-reward outcome with a higher overall value. Furthermore, manipulations assumed to increase impulsivity (pigeons maintained at higher levels of motivation for food and pigeons housed in individual cages) result in increased suboptimal choice. They do so presumably because they function to increase attraction to the signal for the low-probability, high-reward outcomes rather than consider the more global probability of reinforcement associated with each alternative.

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.

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 drivers 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.

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.