Estimating the cardiac signals of chimpanzees using a digital camera: validation and application of a novel non-invasive method for primate research.

Cardiac measures such as heart rate measurements are important indicators of both physiological and psychological states. However, despite their extraordinary potential, their use is restricted in comparative psychology because traditionally cardiac measures involved the attachment of sensors to the participant's body, which, in the case of undomesticated animals such as nonhuman primates, is usually only possible during anesthesia or after extensive training. Here, we validate and apply a camera-based system that enables contact-free detection of animals' heart rates. The system automatically detects and estimates the cardiac signals from cyclic change in the hue of the facial area of a chimpanzee. In Study 1, we recorded the heart rate of chimpanzees using the new technology, while simultaneously measuring heart rate using classic PPG (photoplethysmography) finger sensors. We found that both methods were in good agreement. In Study 2, we applied our new method to measure chimpanzees' heart rate in response to seeing different types of video scenes (groupmates in an agonistic interaction, conspecific strangers feeding, nature videos, etc.). Heart rates changed during video presentation, depending on the video content: Agonistic interactions and conspecific strangers feeding lead to accelerated heart rate relative to baseline, indicating increased emotional arousal. Nature videos lead to decelerated heart rate relative to baseline, indicating a relaxing effect or heightened attention caused by these stimuli. Our results show that the new contact-free technology can reliably assess the heart rate of unrestrained chimpanzees, and most likely other primates. Furthermore, our technique opens up new avenues of research within comparative psychology and facilitates the health management of captive individuals.

Does quantity matter to a stingless bee?

Quantitative information is omnipresent in the world and a wide range of species has been shown to use quantities to optimize their decisions. While most studies have focused on vertebrates, a growing body of research demonstrates that also insects such as honeybees possess basic quantitative abilities that might aid them in finding profitable flower patches. However, it remains unclear if for insects, quantity is a salient feature relative to other stimulus dimensions, or if it is only used as a "last resort" strategy in case other stimulus dimensions are inconclusive. Here, we tested the stingless bee Trigona fuscipennis, a species representative of a vastly understudied group of tropical pollinators, in a quantity discrimination task. In four experiments, we trained wild, free-flying bees on stimuli that depicted either one or four elements. Subsequently, bees were confronted with a choice between stimuli that matched the training stimulus either in terms of quantity or another stimulus dimension. We found that bees were able to discriminate between the two quantities, but performance differed depending on which quantity was rewarded. Furthermore, quantity was more salient than was shape. However, quantity did not measurably influence the bees' decisions when contrasted with color or surface area. Our results demonstrate that just as honeybees, small-brained stingless bees also possess basic quantitative abilities. Moreover, invertebrate pollinators seem to utilize quantity not only as "last resort" but as a salient stimulus dimension. Our study contributes to the growing body of knowledge on quantitative cognition in invertebrate species and adds to our understanding of the evolution of numerical cognition.

Just kidding: the evolutionary roots of playful teasing.

Accounts of teasing have a long history in psychological and sociological research, yet teasing itself is vastly underdeveloped as a topic of study. As a phenomenon that moves along the border between aggression and play, teasing presents an opportunity to investigate key foundations of social and mental life. Developmental studies suggest that preverbal human infants already playfully tease their parents by performing 'the unexpected,' apparently deliberately violating the recipient's expectations to create a shared humorous experience. Teasing behaviour may be phylogenetically old and perhaps an evolutionary precursor to joking. In this review, we present preliminary evidence suggesting that non-human primates also exhibit playful teasing. In particular, we argue that great apes display three types of playful teasing described in preverbal human infants: teasing with offer and withdrawal, provocative non-compliance and disrupting others' activities. We highlight the potential of this behaviour to provide a window into complex socio-cognitive processes such as attribution of others' expectations and, finally, we propose directions for future research and call for systematic studies of teasing behaviour in non-human primates.

Are great apes able to reason from multi-item samples to populations of food items?

Inductive learning from limited observations is a cognitive capacity of fundamental importance. In humans, it is underwritten by our intuitive statistics, the ability to draw systematic inferences from populations to randomly drawn samples and vice versa. According to recent research in cognitive development, human intuitive statistics develops early in infancy. Recent work in comparative psychology has produced first evidence for analogous cognitive capacities in great apes who flexibly drew inferences from populations to samples. In the present study, we investigated whether great apes (Pongo abelii, Pan troglodytes, Pan paniscus, Gorilla gorilla) also draw inductive inferences in the opposite direction, from samples to populations. In two experiments, apes saw an experimenter randomly drawing one multi-item sample from each of two populations of food items. The populations differed in their proportion of preferred to neutral items (24:6 vs. 6:24) but apes saw only the distribution of food items in the samples that reflected the distribution of the respective populations (e.g., 4:1 vs. 1:4). Based on this observation they were then allowed to choose between the two populations. Results show that apes seemed to make inferences from samples to populations and thus chose the population from which the more favorable (4:1) sample was drawn in Experiment 1. In this experiment, the more attractive sample not only contained proportionally but also absolutely more preferred food items than the less attractive sample. Experiment 2, however, revealed that when absolute and relative frequencies were disentangled, apes performed at chance level. Whether these limitations in apes' performance reflect true limits of cognitive competence or merely performance limitations due to accessory task demands is still an open question.

Do monkeys compare themselves to others?

Social comparisons are a fundamental characteristic of human behaviour, yet relatively little is known about their evolutionary foundations. Adapting the co-acting paradigm from human research (Seta in J Pers Soc Psychol 42:281-291, 1982. doi: 10.1037//0022-3514.42.2.281), we examined how the performance of a partner influenced subjects' performance in long-tailed macaques (Macaca fascicularis). Using parallel testing in touch screen setups in which subjects had to discriminate familiar and novel photographs of men and women, we investigated whether accuracy and reaction time were influenced by partner performance and relationship quality (affiliate vs. non-affiliate). Auditory feedback about the alleged performance of the co-actor was provided via playback; partner performance was either moderately or extremely better or worse than subject performance. We predicted that subjects would assimilate to moderately different comparison standards as well as to affiliates and contrast away from extreme standards and non-affiliates. Subjects instantly generalized to novel pictures. While accuracy was not affected by any of the factors, long reaction times occurred more frequently when subjects were tested with a non-affiliate who was performing worse, compared to one who was doing better than them (80% quantile worse: 5.1, better: 4.3 s). For affiliate co-actors, there was no marked effect (worse: 4.4, better: 4.6 s). In a control condition with no auditory feedback, subjects performed somewhat better in the presence of affiliates (M = 77.8% correct) compared to non-affiliates (M = 71.1%), while reaction time was not affected. Apparently, subjects were sensitive to partner identity and performance, yet variation in motivation rather than assimilation and contrast effects may account for the observed effects.

Great ape cognition is structured by stable cognitive abilities and predicted by developmental conditions.

Great ape cognition is used as a reference point to specify the evolutionary origins of complex cognitive abilities, including in humans. This research often assumes that great ape cognition consists of cognitive abilities (traits) that account for stable differences between individuals, which change and develop in response to experience. Here, we test the validity of these assumptions by assessing repeatability of cognitive performance among captive great apes (Gorilla gorilla, Pongo abelii, Pan paniscus, Pan troglodytes) in five tasks covering a range of cognitive domains. We examine whether individual characteristics (age, group, test experience) or transient situational factors (life events, testing arrangements or sociality) influence cognitive performance. Our results show that task-level performance is generally stable over time; four of the five tasks were reliable measurement tools. Performance in the tasks was best explained by stable differences in cognitive abilities (traits) between individuals. Cognitive abilities were further correlated, suggesting shared cognitive processes. Finally, when predicting cognitive performance, we found stable individual characteristics to be more important than variables capturing transient experience. Taken together, this study shows that great ape cognition is structured by stable cognitive abilities that respond to different developmental conditions.

Long-tailed macaques (Macaca fascicularis) can use simple heuristics but fail at drawing statistical inferences from populations to samples.

Human infants, apes and capuchin monkeys engage in intuitive statistics: they generate predictions from populations of objects to samples based on proportional information. This suggests that statistical reasoning might depend on some core knowledge that humans share with other primate species. To aid the reconstruction of the evolution of this capacity, we investigated whether intuitive statistical reasoning is also present in a species of Old World monkey. In a series of four experiments, 11 long-tailed macaques were offered different pairs of populations containing varying proportions of preferred versus neutral food items. One population always contained a higher proportion of preferred items than the other. An experimenter simultaneously drew one item out of each population, hid them in her fists and presented them to the monkeys to choose. Although some individuals performed well across most experiments, our results imply that long-tailed macaques as a group did not make statistical inferences from populations of food items to samples but rather relied on heuristics. These findings suggest that there may have been convergent evolution of this ability in New World monkeys and apes (including humans).

Intuitive statistical inferences in chimpanzees and humans follow Weber's law.

Humans and nonhuman great apes share a sense for intuitive statistical reasoning, making intuitive probability judgments based on proportional information. This ability is of fundamental importance, in particular for inferring general regularities from finite numbers of observations and, vice versa, for predicting the outcome of single events using prior information. To date it remains unclear which cognitive mechanism underlies and enables this capacity. The aim of the present study was to gain deeper insights into the cognitive structure of intuitive statistics by probing its signatures in chimpanzees and humans. We tested 24 sanctuary-living chimpanzees in a previously established paradigm which required them to reason from populations of food items with different ratios of preferred (peanuts) and non-preferred items (carrot pieces) to randomly drawn samples. In a series of eight test conditions, the ratio between the two ratios to be discriminated (ROR) was systematically varied ranging from 1 (same proportions in both populations) to 16 (high magnitude of difference between populations). One hundred and forty-four human adults were tested in a computerized version of the same task. The main result was that both chimpanzee and human performance varied as a function of the log(ROR) and thus followed Weber's law. This suggests that intuitive statistical reasoning relies on the same cognitive mechanism that is used for comparing absolute quantities, namely the analogue magnitude system.

Chimpanzees Consider Humans' Psychological States when Drawing Statistical Inferences.

Great apes have been shown to be intuitive statisticians: they can use proportional information within a population to make intuitive probability judgments about randomly drawn samples [1, J.E., J.C., J.H., E.H., and H.R., unpublished data]. Humans, from early infancy onward, functionally integrate intuitive statistics with other cognitive domains to judge the randomness of an event [2-6]. To date, nothing is known about such cross-domain integration in any nonhuman animal, leaving uncertainty about the origins of human statistical abilities. We investigated whether chimpanzees take into account information about psychological states of experimenters (their biases and visual access) when drawing statistical inferences. We tested 21 sanctuary-living chimpanzees in a previously established paradigm that required subjects to infer which of two mixed populations of preferred and non-preferred food items was more likely to lead to a desired outcome for the subject. In a series of three experiments, we found that chimpanzees chose based on proportional information alone when they had no information about experimenters' preferences and (to a lesser extent) when experimenters had biases for certain food types but drew blindly. By contrast, when biased experimenters had visual access, subjects ignored statistical information and instead chose based on experimenters' biases. Lastly, chimpanzees intuitively used a violation of statistical likelihoods as indication for biased sampling. Our results suggest that chimpanzees have a random sampling assumption that can be overridden under the appropriate circumstances and that they are able to use mental state information to judge whether this is necessary. This provides further evidence for a shared statistical inference mechanism in apes and humans.