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.psychology | behavior | comparative methods | inhibitory control | executive function S ince Darwin, understanding the evolution of cognition has been widely regarded as one of the greatest challenges for evolutionary research (1). Although researchers have identified surprising cognitive flexibility in a range of species (2-40) and potentially derived features of human psychology (41-61), we know much less about the major forces shaping cognitive evolution (62-71). With the notable exception of Bitterman's landmark studies conducted several decades ago (63, 72-74), most research comparing cognition across species has been limited to small taxonomic samples (70, 75). With limited comparable experimental data on how cognition varies across species, previous research has largely relied on proxies for cognition (e.g., brain size) or metaanalyses when testing hypotheses about cognitive evolution (76-92). The lack of cognitive data collected with similar methods across large samples of species precludes meaningful species comparisons that can reveal the major forces shaping cognitive evolution across species, including humans (48,70,89,(93)(94)(95)(96)(97)(98). SignificanceAlthough 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 problemsolving tasks measuring self-control and evaluated the leading hypotheses regarding how ...

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.

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, in spite of the fact that the stimulus that predicts 0 pellets occurs four 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 to 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) and 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.

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.

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, 1203Society B, 278, -1208Society B, 278, , 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.Keywords decision making . suboptimal choice . gambling . humans . pigeons A subset of suboptimal decision making can be characterized as making a decision to choose a low-probability but high-payoff alternative (e.g., playing slot machines or buying lottery tickets) over a high-probability, low-payoff alternative (not gambling), such that the net expected return is less than what one has wagered. In this kind of gambling, over the long term, one is very likely to lose more than one wins, and having sufficient experience with the game to learn about the odds of winning does not appear to reduce its frequency. It appears that gamblers tend to over-attend to wins and discount losses (the availability heuristic; Tversky & Kahneman, 1974).Recently, Zentall and colleagues studied suboptimal decision-making processes in pigeons (Gipson, Alessandri, Miller, & Zentall, 2009;Stagner & Zentall, 2010;Zentall & Stagner, 2011). They asked whether pigeons would show choice behavior analogous to the suboptimal behavior shown by humans when humans purchase a lottery ticket or engage in casino gambling. To answer this question, Zentall and Stagner designed a task that may be considered analogous to human gambling (see Fig. 1a). In this task, pigeons are given a choice between one alternative reliably associated with three reinforcers (analogous to money in one's pocket) and a second alternative (analogous to gambling) associated with a low-probability, high-payoff stimulus on some trials (10 pellets on 20% of the trials) or a highprobability, zero-payoff stimulus on others. In one sense, the task is analogous to suboptimal human gambling, because the average payoff for the first alternative (with an overall mean reinforcement per trial of 3 pellets) is greater than that of the second alternative (with an overall mean reinforcement per trial of 2 pellets). Results show...

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 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 hrs 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.

Numerous functional magnetic resonance imaging (fMRI) studies have reported sex differences. To empirically evaluate for evidence of excessive significance bias in this literature, we searched for published fMRI studies of human brain to evaluate sex differences, regardless of the topic investigated, in Medline and Scopus over 10 years. We analyzed the prevalence of conclusions in favor of sex differences and the correlation between study sample sizes and number of significant foci identified. In the absence of bias, larger studies (better powered) should identify a larger number of significant foci. Across 179 papers, median sample size was n = 32 (interquartile range 23-47.5). A median of 5 foci related to sex differences were reported (interquartile range, 2-9.5). Few articles (n = 2) had titles focused on no differences or on similarities (n = 3) between sexes. Overall, 158 papers (88%) reached “positive” conclusions in their abstract and presented some foci related to sex differences. There was no statistically significant relationship between sample size and the number of foci (−0.048% increase for every 10 participants, p = 0.63). The extremely high prevalence of “positive” results and the lack of the expected relationship between sample size and the number of discovered foci reflect probable reporting bias and excess significance bias in this literature.