Performance of Asian elephants (Elephas maximus) on a quantity discrimination task is similar to that of African savanna elephants (Loxodonta africana).

Using an object-choice task, we measured the relative quantity discrimination ability of Asian elephants. Two zoo-housed elephants were given auditory cues of food being dropped into two containers (Nonvisible condition), and in one condition they could also see the food on top of the containers (Visible condition). Elephants received sets of varying ratios and magnitudes. We found that the elephants chose the greater quantity of food significantly above chance in both the Visible and Nonvisible conditions. Additionally, we found the elephants' ability to discriminate between quantities decreased as the ratio, and not the absolute difference, between the quantities increased, which is predicted by the accumulator model. We also compare our findings to those from a study using the same methods with African savanna elephants and found that the two species performed at similar levels, but given our small sample size it is difficult to make strong species-level conclusions. In discussing our results, we consider differences between the two species' wild environments as well as the types of sensory cues provided in human care, and we provide recommendations for extensions of this work.

Irrational choice behavior in human and nonhuman primates.

Choice behavior in humans has motivated a large body of research with a focus on whether decisions can be considered to be rational. In general, humans prefer having choice, as do a number of other species that have been tested, even though having increased choice does not necessarily yield a positive outcome. Humans have been found to choose an option more often only because the opportunity to select it was diminishing, an example of a deviation from economic rationality. Here we extend this paradigm to nonhuman primates in an effort to understand the mechanisms underlying this finding. In this study, we presented two groups of laboratory monkeys, capuchins (Cebus apella) and rhesus macaques (Macaca mulatta), as well as human subjects, with a computerized task in which subjects were presented with two differently colored icons. When the subject selected an icon, differing numbers of food pellets were dispensed (or points were assigned), making each icon correspond to a certain level of risk (one icon yielded 1 or 4 pellets/points and the other yielded 2 or 3). Initially, both options remained constantly available and we established choice preference scores for each subject. Then, we assessed preference patterns once the options were not continuously available. Specifically, choosing one icon would cause the other to shrink in size on the screen and eventually disappear if never selected. Selecting it would restore it to its full size. As predicted, humans shifted their risk preferences in the diminishing options phase, choosing to click on both icons more equally in order to keep both options available. At the group level, capuchin monkeys showed this pattern as well, but there was a great deal of individual variability in both capuchins and macaques. The present work suggests that there is some degree of continuity between human and nonhuman primates in the desire to have choice simply for the sake of having choice.

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.

Waiting for what comes later: capuchin monkeys show self-control even for nonvisible delayed rewards.

Self-control tasks used with nonhuman animals typically involve the choice between an immediate option and a delayed, but more preferred option. However, in many self-control scenarios, not only does the more impulsive option come sooner in time, it is often more concrete than the delayed option. For example, studies have presented children with the option of eating a visible marshmallow immediately, or foregoing it for a better reward that can only be seen later. Thus, the immediately available option is visible and concrete, whereas the delayed option is not visible and more abstract. We tested eight capuchin monkeys to better understand this potential effect by manipulating the visibility of the response options and the visibility of the baiting itself. Monkeys observed two food items (20 or 5 g pieces of banana) each being placed either on top of or inside of one of the two opaque containers attached to a revolving tray apparatus, either in full view of monkeys or occluded by a barrier. Trials ended when monkeys removed a reward from the rotating tray. To demonstrate self-control, monkeys should have allowed the smaller piece of food to pass if the larger piece was forthcoming. Overall, monkeys were successful on the task, allowing a smaller, visible piece of banana to pass from reach in order to access the larger, nonvisible banana piece. This was true even when the entire baiting process took place out of sight of the monkeys. This finding suggests that capuchin monkeys succeed on self-control tasks even when the delayed option is also more abstract than the immediate one-a situation likely faced by primates in everyday life.

Rates of reinforcement and measures of compliance in free and protected contact elephant management systems.

Protected contact is an alternative to traditional captive elephant training techniques that emerged as a result of concerns for animal welfare and personnel safety. The present study documented the behavior of elephants and their animal care professionals to determine rates of reinforcement and measures of compliance under two handling systems. Behavioral data were collected from animal care professionals and elephants during the elephants' baths in both free contact (FC) and protected contact (PC). Positive reinforcement, in the form of food, was delivered, on average, nearly eight times more frequently in the PC condition. Further, the mean rate at which the animal care professionals used the ankus in the FC condition as negative reinforcement was similar to the mean rate at which they provided positive reinforcement to the elephants in the FC condition. Latencies between verbal commands and the elephants' behaviors demonstrated an inconsistent pattern, but were generally longer in the PC condition. The mean percent of "refusals" by the elephants was higher for most behaviors across elephants in the PC condition. The findings suggest that animal care professionals did not heavily rely on positive reinforcement in the FC condition to elicit desired behaviors from the elephants, but this was the case in the PC condition. We propose that longer latencies and higher mean percent of refusals by the elephants may indicate that they were exercising choice or control over their environment, which has been associated with improved well-being. Additional studies of this kind are needed to enable other institutions to make informed decisions about elephant management and welfare.

Prospective memory in children and chimpanzees.

Prospective memory (PM) involves remembering to do something at a specific time in the future. Here, we investigate the beginnings of this ability in young children (3-year-olds; Homo sapiens) and chimpanzees (Pan troglodytes) using an analogous task. Subjects were given a choice between two toys (children) or two food items (chimpanzees). The selected item was delivered immediately, whereas the unselected item was hidden in an opaque container. After completing an ongoing quantity discrimination task, subjects could request the hidden item by asking for it (children) or by pointing to the container and identifying the item on a symbol board (chimpanzees). Children and chimpanzees showed evidence of prospective-like memory in this task, as evidenced by successful retrieval of the item at the end of the task, sometimes spontaneously with no prompting from the experimenter. These findings contribute to our understanding of PM from an ontogenetic and comparative perspective.

Do monkeys choose to choose?

Both empirical and anecdotal evidence supports the idea that choice is preferred by humans. Previous research has demonstrated that this preference extends to nonhuman animals, but it remains largely unknown whether animals will actively seek out or prefer opportunities to choose. Here we explored the issue of whether capuchin and rhesus monkeys choose to choose. We used a modified version of the SELECT task-a computer program in which monkeys can choose the order of completion of various psychomotor and cognitive tasks. In the present experiments, each trial began with a choice between two icons, one of which allowed the monkey to select the order of task completion, and the other of which led to the assignment of a task order by the computer. In either case, subjects still had to complete the same number of tasks and the same number of task trials. The tasks were relatively easy, and the monkeys responded correctly on most trials. Thus, global reinforcement rates were approximately equated across conditions. The only difference was whether the monkey chose the task order or it was assigned, thus isolating the act of choosing. Given sufficient experience with the task icons, all monkeys showed a significant preference for choice when the alternative was a randomly assigned order of tasks. To a lesser extent, some of the monkeys maintained a preference for choice over a preferred, but computer-assigned, task order that was yoked to their own previous choice selection. The results indicated that monkeys prefer to choose when all other aspects of the task are equated.

Language-trained chimpanzees (Pan troglodytes) name what they have seen but look first at what they have not seen.

Metacognition can be defined as knowing what one knows, and the question of whether nonhuman animals are metacognitive has driven an intense debate. We tested 3 language-trained chimpanzees in an information-seeking task in which the identity of a food item was the critical piece of information needed to obtain the food. The chimpanzees could either report the identity of the food immediately or first check a container in which the food had been hidden. In two experiments, the chimpanzees were significantly more likely to visit the container first on trials in which they could not know its contents but were more likely to just name the food item without looking into the container on trials in which they had seen its contents. Thus, chimpanzees showed efficient information-seeking behavior that suggested they knew what they had or had not already seen when it was time to name a hidden item.

Chimpanzees (Pan troglodytes) transfer tokens repeatedly with a partner to accumulate rewards in a self-control task.

There has been extensive research investigating self-control in humans and nonhuman animals, yet we know surprisingly little about how one's social environment influences self-control. The present study examined the self-control of chimpanzees in a task that required active engagement with conspecifics. The task consisted of transferring a token back and forth with a partner animal in order to accumulate food rewards, one item per token transfer. Self-control was required because at any point in the trial, either chimpanzee could obtain their accumulated rewards, but doing so discontinued the food accumulation and ended the trial for both individuals. Chimpanzees readily engaged the task and accumulated the majority of available rewards before ending each trial, and they did so across a number of conditions that varied the identity of the partner, the presence/absence of the experimenter, and the means by which they could obtain rewards. A second experiment examined chimpanzees' self-control when given the choice between immediately available food items and a potentially larger amount of rewards that could be obtained by engaging the token transfer task with a partner. Chimpanzees were flexible in their decision-making in this test, typically choosing the option representing the largest amount of food, even if it involved delayed accumulation of the rewards via the token transfer task. These results demonstrate that chimpanzees can exhibit self-control in situations involving social interactions, and they encourage further research into this important aspect of the self-control scenario.

Can black-and-white ruffed lemurs (Varecia variegata) solve object permanence tasks?

We examined object permanence in black-and-white-ruffed lemurs (Varecia variegata) at Zoo Atlanta. A series of visible and invisible displacement tasks with suitable controls were presented to five adult subjects. Subjects performed significantly above chance on all regular tasks, except for the double invisible displacements. Subjects failed visible and invisible controls. Failure on the control trials did not appear to be because subjects used the "last box touched" strategy (subjects did not choose the last box touched significantly more than expected by chance). However, a substantial percentage of choices was made to the last box touched by the experimenter. There was no significant difference between this percentage, and the percentage of choices made to the baited box (on both visible and invisible controls), which indicates that subjects were drawn to both boxes which the experimenter visited/touched, and thus failed the controls. Based on the results from the present study, we believe that there is no evidence that black-and-white ruffed lemurs understand visible and invisible tasks in the traditional object permanence battery.

Habitual prospective memory in preschool children.

Habitual prospective memory (PM) refers to situations in which individuals have to remember to perform a future task on a regular and frequent basis. Habitual PM tasks are ubiquitous and the ability to successfully complete these tasks (e.g., remembering to bring your lunch to school every day) is necessary for children as they begin to establish their own independence. The current investigation is the first to explore preschool children's ability to complete this kind of task. At the end of a regular testing session during which children engaged in a variety of unrelated cognitive tasks, participants were instructed to ask for a stamp on their card, which was sitting in a box on the table. Over the course of the first experiment, participants did this 13 times, spanning a time period of several months. The results demonstrated that children initially needed prompting from the experimenter to remember, but with experience, participants were able to retrieve this intention without assistance. Experiment 2 demonstrated that removing the box from participants' line of sight after numerous opportunities to perform the task did not negatively impact performance, although it did make a difference at the outset of this requirement to remember to ask for stamps. Together, these results indicate that with somewhat consistent and repeated practice, preschool children can fairly quickly demonstrate the ability to successfully perform future intentions that are likely to be repeated on numerous occasions.

Focality and prospective memory in preschool children.

We examined the influence of prospective memory (PM) cue focality in a sample of preschool children. Prior investigations in older populations indicated that focal targets were associated with enhanced PM performance, perhaps through more automatic retrieval processes. Importantly, this influential variable has not been thoroughly explored in younger samples. Over three test sessions, preschool children completed a memory task where they were shown a series of animals. During retrieval, participants were shown all of the animals except for one, and they had to name the missing animal. While engaged in this task, participants in the focal PM condition were instructed to remove particular animals (e.g., spider) from the game if they saw them. In the nonfocal condition, participants were told to remove any animal that was entirely one color (e.g., black) if they saw them during the game. The results demonstrated no difference in PM remembering between focal and nonfocal conditions. These results suggest that the effects of focality may not be present at the beginning stages of PM development. The implications for PM retrieval processes also are discussed.

Putting the elephant back in the herd: elephant relative quantity judgments match those of other species.

The ability to discriminate between quantities has been observed in many species. Typically, when an animal is given a choice between two sets of food, accurate performance (i.e., choosing the larger amount) decreases as the ratio between two quantities increases. A recent study reported that elephants did not exhibit ratio effects, suggesting that elephants may process quantitative information in a qualitatively different way from all other nonhuman species that have been tested (Irie-Sugimoto et al. in Anim Cogn 12:193-199, 2009). However, the results of this study were confounded by several methodological issues. We tested two African elephants (Loxodonta africana) to more thoroughly investigate relative quantity judgment in this species. In contrast to the previous study, we found evidence of ratio effects for visible and nonvisible sequentially presented sets of food. Thus, elephants appear to represent and compare quantities in much the same way as other species, including humans when they are prevented from counting. Performance supports an accumulator model in which quantities are represented as analog magnitudes. Furthermore, we found no effect of absolute magnitude on performance, providing support against an object-file model explanation of quantity judgment.

Capuchin monkeys (Cebus apella) let lesser rewards pass them by to get better rewards.

Self-control is defined as foregoing an immediate reward to gain a larger delayed reward. Methods used to test self-control comparatively include inter-temporal choice tasks, delay of gratification tasks, and accumulation tasks. To date, capuchin monkeys have shown different levels of self-control across tasks. This study introduced a new task that could be used comparatively to measure self-control in an intuitive context that involved responses that required no explicit training. Capuchin monkeys (Cebus apella) were given a choice between two food items that were presented on a mechanized, revolving tray that moved those foods sequentially toward the monkeys. A monkey could grab the first item or wait for the second, but was only allowed one item. Most monkeys in the study waited for a more highly preferred food item or a larger amount of the same food item when those came later, and they inhibited the prepotent response to grab food by not reaching out to take less-preferred foods or smaller amounts of food that passed directly in front of them first. These data confirm that the mechanisms necessary for self-control are present in capuchin monkeys and indicate that the methodology can be useful for broader comparative assessments of self-control.

Technology at the zoo: the influence of a touchscreen computer on orangutans and zoo visitors.

A computer-controlled touchscreen apparatus (hereafter referred to as "touchscreen") in the orangutan exhibit at Zoo Atlanta provides enrichment to the animals and allows cognitive research to take place on exhibit. This study investigated the impact of the touchscreen on orangutan behavior and visibility, as well as its impact on zoo visitors. Despite previous research suggesting that providing a single computer system may negatively affect orangutan behavior, there was not a significant increase in aggression, stereotypic, or distress-related behaviors following the activation of the on-exhibit touchscreen. We also investigated the possibility that zoo visitors may be negatively affected by technology because it deviates from naturalism. However, we did not find a change in stay time or overall experience rating when the computer was turned on. This research was the first to assess visitor attitudes toward technology at the zoo, and we found that visitors report highly positive attitudes about technology for both animals and visitors. If subjects visited the exhibit when the computer was turned on, they more strongly agreed that orangutans benefit from interacting with computerized enrichment. This study is the first investigation of an on-exhibit touchscreen in group-housed apes; our findings of no negative effects on the animals or zoo visitors and positive attitudes toward technology suggest a significant value of this practice.

The evolution of quantitative sensitivity.

The ability to represent approximate quantities appears to be phylogenetically widespread, but the selective pressures and proximate mechanisms favouring this ability remain unknown. We analysed quantity discrimination data from 672 subjects across 33 bird and mammal species, using a novel Bayesian model that combined phylogenetic regression with a model of number psychophysics and random effect components. This allowed us to combine data from 49 studies and calculate the Weber fraction (a measure of quantity representation precision) for each species. We then examined which cognitive, socioecological and biological factors were related to variance in Weber fraction. We found contributions of phylogeny to quantity discrimination performance across taxa. Of the neural, socioecological and general cognitive factors we tested, cortical neuron density and domain-general cognition were the strongest predictors of Weber fraction, controlling for phylogeny. Our study is a new demonstration of evolutionary constraints on cognition, as well as of a relation between species-specific neuron density and a particular cognitive ability. This article is part of the theme issue 'Systems neuroscience through the lens of evolutionary theory'.

Sex differences in spatial ability: a test of the range size hypothesis in the order Carnivora.

Sex differences in spatial cognition have been reported for many species ranging from voles to humans. The range size hypothesis predicts that sex differences in spatial ability will only occur in species in which the mating system selects for differential range size. Consistent with this prediction, we observed sex differences in spatial ability in giant pandas, a promiscuous species in which males inhabit larger ranges than females, but did not observe sex differences in Asian small-clawed otters, a related monogamous species in which males and females share home ranges. These results provide the first evidence of sex differences in spatial ability in the order Carnivora, and are consistent with the range size hypothesis.

The use of technology to enhance zoological parks.

Technology can be used in a zoological setting to improve visitor experience, increase research opportunities, and enhance animal welfare. Evaluating the quality of these technological innovations and their use by nonhuman and human counterparts is a critical part of extending the uses of technology to enhance animal welfare and visitor experience at zoological parks. Survey data from a small sample of institutions housing primates suggest that computers, television, radio, and sprinklers are the most prevalent types of technological enrichment currently used. Survey respondents were positive about the technology implemented, stating a desire to increase its use.

Factors affecting aggression in a captive flock of Chilean flamingos (Phoenicopterus chilensis).

The influence of pair bond status, age and sex on aggression rates in a flock of 84 captive Chilean flamingos at Zoo Atlanta was examined. Analysis showed no difference between aggression rates of male and female flamingos, but adult flamingos had higher rates of aggression than juveniles. There were also significant differences in aggression depending on pair bond status (single, same-sex pair, male-female pair or group). Bonded birds were significantly more aggressive than single birds, which is consistent with the concept that unpaired birds are not breeding and do not need to protect pair bonds or eggs. Birds in typical pair bonds (male-female) and atypical pair bonds (same-sex pairs or groups) exhibited similar rates of aggression. These results contribute to the existing body of research on aggression in captive flamingos.

Spatial memory recall in the giant panda (Ailuropoda melanoleuca).

The current study tested spatial memory recall in 1 male and 1 female giant panda (Ailuropoda melanoleuca). The task required subjects to make a delayed response to a previously lighted location, with varying lengths of delay between the observation phase and the test phase. The male subject reached criterion at 2-, 3-, 4-, 5-, 6-, and 10-s delays. The female subject reached criterion at 2-, 3-, 4-, 5-, 6-, 10-, and 15-s delays. The results support the hypothesis that giant pandas demonstrate significant working memory for spatial location in the absence of external cues, which may be an important mechanism for survival in the wild.