Audrey E. Parrish

Responses to the Assurance game in monkeys, apes, and humans using equivalent procedures

There is great interest in the evolution of economic behavior. In typical studies, species are asked to play one of a series of economic games, derived from game theory, and their responses are compared. The advantage of this approach is the relative level of consistency and control that emerges from the games themselves; however, in the typical experiment, procedures and conditions differ widely, particularly between humans and other species. Thus, in the current study, we investigated how three primate species, capuchin monkeys, chimpanzees, and humans, played the Assurance (or Stag Hunt) game using procedures that were, to the best of our ability, the same across species, particularly with respect to training and pretesting. Our goal was to determine what, if any, differences existed in the ways in which these species made decisions in this game. We hypothesized differences along phylogenetic lines, which we found. However, the species were more similar than might be expected. In particular, humans who played using “nonhuman primate-friendly” rules did not behave as is typical. Thus, we find evidence for similarity in decision-making processes across the order Primates. These results indicate that such comparative studies are possible and, moreover, that in any comparison rating species’ relative abilities, extreme care must be taken in ensuring that one species does not have an advantage over the others due to methodological procedures.

When less is more: like humans, chimpanzees (Pan troglodytes) misperceive food amounts based on plate size.

Abstract We investigated whether chimpanzees (Pan troglodytes) misperceived food portion sizes depending upon the context in which they were presented, something that often affects how much humans serve themselves and subsequently consume. Chimpanzees judged same-sized and smaller food portions to be larger in amount when presented on a small plate compared to an equal or larger food portion presented on a large plate and did so despite clearly being able to tell the difference in portions when plate size was identical. These results are consistent with data from the human literature in which people misperceive food portion sizes as a function of plate size. This misperception is attributed to the Delboeuf illusion which occurs when the size of a central item is misperceived on the basis of its surrounding context. These results demonstrate a cross-species shared visual misperception of portion size that affects choice behavior, here in a nonhuman species for which there is little experience with tests that involve choosing between food amounts on dinnerware. The biases resulting in this form of misperception of food portions appear to have a deep-rooted evolutionary history which we share with, at minimum, our closest living nonhuman relative, the chimpanzee.

Sequential responding and planning in capuchin monkeys (Cebus apella)

Previous experiments have assessed planning during sequential responding to computer generated stimuli by Old World nonhuman primates including chimpanzees and rhesus macaques. However, no such assessment has been made with a New World primate species. Capuchin monkeys (Cebus apella) are an interesting test case for assessing the distribution of cognitive processes in the Order Primates because they sometimes show proficiency in tasks also mastered by apes and Old World monkeys, but in other cases fail to match the proficiency of those other species. In two experiments, eight capuchin monkeys selected five arbitrary stimuli in distinct locations on a computer monitor in a learned sequence. In Experiment 1, shift trials occurred in which the second and third stimuli were transposed when the first stimulus was selected by the animal. In Experiment 2, mask trials occurred in which all remaining stimuli were masked after the monkey selected the first stimulus. Monkeys made more mistakes on trials in which the locations of the second and third stimuli were interchanged than on trials in which locations were not interchanged, suggesting they had already planned to select a location that no longer contained the correct stimulus. When mask trials occurred, monkeys performed at levels significantly better than chance, but their performance exceeded chance levels only for the first and the second selections on a trial. These data indicate that capuchin monkeys performed very similarly to chimpanzees and rhesus monkeys and appeared to plan their selection sequences during the computerized task, but only to a limited degree.

Delay of gratification by orangutans (Pongo pygmaeus) in the accumulation task.

There is considerable evidence indicating that chimpanzees can delay gratification for extended time intervals, particularly in the accumulation task in which food items accumulate within a participants reach until the participant begins to consume them. However, there is limited evidence that other ape species might also exhibit this capacity, despite there being a number of similar studies indicating that nonape species (e.g., monkeys and birds) can delay gratification, but not for nearly as long as chimpanzees. To help define the taxonomic distribution of delay of gratification behavior in the order Primates, we tested 6 orangutans in the current experiments and compared their performance with comparable data from a previous study with capuchin monkeys. We varied delay length and visibility of the items that were still available for accumulation to determine the impact of these factors on performance. Species differences on the accumulation task emerged when comparing the current data to data from a previous study. Orangutans outperformed capuchin monkeys, suggesting that ape species may generally show better delay of gratification and delay maintenance abilities than monkeys. However, more studies are necessary to rule out alternative hypotheses on the distribution of delay maintenance abilities across primate species.

Do primates see the solitaire illusion differently? A comparative assessment of humans (Homo sapiens), chimpanzees (Pan troglodytes), rhesus monkeys (Macaca mulatta), and capuchin monkeys (Cebus apella).

An important question in comparative psychology is whether human and nonhuman animals share similar principles of perceptual organization. Despite much empirical research, no firm conclusion has been drawn. The Solitaire illusion is a numerosity illusion in humans that occurs when one misperceives the relative number of 2 types of items presented in intermingled sets. To date, no study has investigated whether nonhuman animals perceive the Solitaire illusion as humans do. Here, we compared the perception of the Solitaire illusion in human and nonhuman primates in 3 experiments. We first observed (Experiment 1) the spontaneous behavior of chimpanzees when presented with 2 arrays composed of a different number of preferred and nonpreferred food items. In probe trials, preferred items were presented in the Solitaire pattern in 2 different spatial arrangements (either clustered centrally or distributed on the perimeter). Chimpanzees did not show any misperception of quantity in the Solitaire pattern. Next, humans, chimpanzees, rhesus monkeys, and capuchin monkeys underwent the same testing of relative quantity judgments in a computerized task that also presented the Solitaire illusion (Experiments 2 and 3). Unlike humans, chimpanzees did not appear to perceive the illusion, in agreement with Experiment 1. The performance of rhesus monkeys and capuchin monkeys was also different from that of humans, but was slightly more indicative of a potential Solitaire illusion. On the whole, our results suggest a potential discontinuity in the visual mechanisms underlying the Solitaire illusion between human and nonhuman primates.

Visual nesting of stimuli affects rhesus monkeys’ (Macaca mulatta) quantity judgments in a bisection task

Nonhuman animals are highly proficient at judging relative quantities presented in a variety of formats, including visual, auditory, and even cross-modal formats. Performance typically is constrained by the ratio between sets, as would be expected under Weber’s law and as is described in the approximate number system (ANS) hypothesis. In most cases, tests are designed to avoid any perceptual confusion for animals regarding the stimulus sets, but despite this, animals show some of the perceptual biases that humans show based on organization of stimuli. Here, we demonstrate an additional perceptual bias that emerges from the illusion of nested sets. When arrays of circles were presented on a computer screen and were to be classified as larger than or smaller than an established central value, rhesus monkeys (Macaca mulatta) underestimated quantities when circles were nested within each other. This matched a previous report with adult humans (Chesney & Gelman, Attention, Perception, & Psychophysics 24:1104–1113, 2012), indicating that macaques, like humans, show the pattern of biased perception predicted by ANS estimation. Although some macaques overcame this perceptual bias, demonstrating that they could come to view nested stimuli as individual elements to be included in the estimates of quantity used for classifying arrays, the majority of the monkeys showed the bias of underestimating nested arrays throughout the experiment.

The elusive illusion: Do children (Homo sapiens) and capuchin monkeys (Cebus apella) see the Solitaire illusion?

One approach to gaining a better understanding of how we perceive the world is to assess the errors that human and nonhuman animals make in perceptual processing. Developmental and comparative perspectives can contribute to identifying the mechanisms that underlie systematic perceptual errors often referred to as perceptual illusions. In the visual domain, some illusions appear to remain constant across the lifespan, whereas others change with age. From a comparative perspective, many of the illusions observed in humans appear to be shared with nonhuman primates. Numerosity illusions are a subset of visual illusions and occur when the spatial arrangement of stimuli within a set influences the perception of quantity. Previous research has found one such illusion that readily occurs in human adults, the Solitaire illusion. This illusion appears to be less robust in two monkey species, rhesus macaques and capuchin monkeys. We attempted to clarify the ontogeny of this illusion from a developmental and comparative perspective by testing human children and task-naïve capuchin monkeys in a computerized quantity judgment task. The overall performance of the monkeys suggested that they perceived the numerosity illusion, although there were large differences among individuals. Younger children performed similarly to the monkeys, whereas older children more consistently perceived the illusion. These findings suggest that human-unique perceptual experiences with the world might play an important role in the emergence of the Solitaire illusion in human adults, although other factors also may contribute.

Go when you know: Chimpanzees' confidence movements reflect their responses in a computerized memory task.

Three chimpanzees performed a computerized memory task in which auditory feedback about the accuracy of each response was delayed. The delivery of food rewards for correct responses also was delayed and occurred in a separate location from the response. Crucially, if the chimpanzees did not move to the reward-delivery site before food was dispensed, the reward was lost and could not be recovered. Chimpanzees were significantly more likely to move to the dispenser on trials they had completed correctly than on those they had completed incorrectly, and these movements occurred before any external feedback about the outcome of their responses. Thus, chimpanzees moved (or not) on the basis of their confidence in their responses, and these confidence movements aligned closely with objective task performance. These untrained, spontaneous confidence judgments demonstrated that chimpanzees monitored their own states of knowing and not knowing and adjusted their behavior accordingly.

Rhesus macaques (Macaca mulatta) exhibit the decoy effect in a perceptual discrimination task.

The asymmetric dominance effect (or decoy effect) is a form of context-dependent choice bias in which the probability of choosing one of two options is impacted by the introduction of a third option, also known as the decoy. Decoy effects are documented widely within the human consumer choice literature, and even extend to preference testing within nonhuman animals. Here, we extended this line of research to a perceptual discrimination task with rhesus monkeys to determine whether decoy stimuli would impact size judgments of rectangular stimuli. In a computerized task, monkeys attempted to choose the larger of two rectangles that varied in size and orientation (horizontally or vertically oriented). In probe trials, a third stimulus (the decoy) was presented that was smaller than the other two rectangles but matched the orientation of one of them. On half of the probe trials, the presented decoy matched the orientation of the larger stimulus, and on the other half, the decoy matched the orientation of the smaller stimulus. Monkeys rarely selected the decoy stimulus. However, their performance (selection of the largest rectangle) increased relative to the baseline trials (with only two choices) when the decoy was congruent in its orientation with the largest rectangle, but decreased relative to baseline when the decoy was incongruent with the largest rectangle. Thus, a decoy stimulus impacted monkeys’ perceptual choice behavior even when it was not a viable choice option itself. These results are explained with regard to comparative evaluation mechanisms.

Do rhesus monkeys (Macaca mulatta) perceive the Zöllner illusion

A long-standing debate surrounds the issue of whether human and nonhuman animals share the same perceptual mechanisms. In humans, the Zöllner illusion occurs when two parallel lines appear to be convergent when oblique crosshatching lines are superimposed. Although one baboon study suggests that they too might perceive this illusion, the results of that study were unclear, whereas two recent studies suggest that birds see this illusion in the opposite direction from humans. It is currently unclear whether these mixed results are an artifact of the experimental design or reflect a peculiarity of birds’ visual system or, instead, a wider phenomenon shared among nonhuman mammals. Here, we trained 6 monkeys to select the narrower of two gaps at the end of two convergent lines. Three different conditions were set up: control (no crosshatches), perpendicular (crosshatches not inducing the illusion), and Zöllner (crosshatches inducing the illusion in humans). During training, the degrees of convergence between the two lines ranged from 15° to 12°. Monkeys that reached the training criterion were tested with more difficult discriminations (11°–1°), including probe trials with parallel lines (0°). The results showed that monkeys perceived the Zöllner illusion in the same direction as humans. Comparison of these data with the data from bird studies points toward the existence of different orientation-tuned mechanisms between primate and nonprimate species.