Joshua M. Plotnik

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.

Elephants know when they need a helping trunk in a cooperative task

Elephants are widely assumed to be among the most cognitively advanced animals, even though systematic evidence is lacking. This void in knowledge is mainly due to the danger and difficulty of submitting the largest land animal to behavioral experiments. In an attempt to change this situation, a classical 1930s cooperation paradigm commonly tested on monkeys and apes was modified by using a procedure originally designed for chimpanzees (Pan troglodytes) to measure the reactions of Asian elephants (Elephas maximus). This paradigm explores the cognition underlying coordination toward a shared goal. What do animals know or learn about the benefits of cooperation? Can they learn critical elements of a partners role in cooperation? Whereas observations in nature suggest such understanding in nonhuman primates, experimental results have been mixed, and little evidence exists with regards to nonprimates. Here, we show that elephants can learn to coordinate with a partner in a task requiring two individuals to simultaneously pull two ends of the same rope to obtain a reward. Not only did the elephants act together, they inhibited the pulling response for up to 45 s if the arrival of a partner was delayed. They also grasped that there was no point to pulling if the partner lacked access to the rope. Such results have been interpreted as demonstrating an understanding of cooperation. Through convergent evolution, elephants may have reached a cooperative skill level on a par with that of chimpanzees.

Asian elephants (Elephas maximus) reassure others in distress

Contact directed by uninvolved bystanders toward others in distress, often termed consolation, is uncommon in the animal kingdom, thus far only demonstrated in the great apes, canines, and corvids. Whereas the typical agonistic context of such contact is relatively rare within natural elephant families, other causes of distress may trigger similar, other-regarding responses. In a study carried out at an elephant camp in Thailand, we found that elephants affiliated significantly more with other individuals through directed, physical contact and vocal communication following a distress event than in control periods. In addition, bystanders affiliated with each other, and matched the behavior and emotional state of the first distressed individual, suggesting emotional contagion. The initial distress responses were overwhelmingly directed toward ambiguous stimuli, thus making it difficult to determine if bystanders reacted to the distressed individual or showed a delayed response to the same stimulus. Nonetheless, the directionality of the contacts and their nature strongly suggest attention toward the emotional states of conspecifics. The elephants’ behavior is therefore best classified with similar consolation responses by apes, possibly based on convergent evolution of empathic capacities.

Self‐recognition in the Asian elephant and future directions for cognitive research with elephants in zoological settings

The field of animal cognition has grown steadily for nearly four decades, but the primary focus has centered on easily kept lab animals of varying cognitive capacity, including rodents, birds and primates. Elephants (animals not easily kept in a laboratory) are generally thought of as highly social, cooperative, intelligent animals, yet few studies-with the exception of long-term behavioral field studies-have been conducted to directly support this assumption. In fact, there has been remarkably little cognitive research conducted on Asian (Elephas maximus) or African (Loxodonta africana or L. cyclotis) elephants. Here, we discuss the opportunity and rationale for conducting such research on elephants in zoological facilities, and review some of the recent developments in the field of elephant cognition, including our recent study on mirror self-recognition in E. maximus.

Visual Cues Given by Humans Are Not Sufficient for Asian Elephants (Elephas maximus) to Find Hidden Food

Recent research suggests that domesticated species – due to artificial selection by humans for specific, preferred behavioral traits – are better than wild animals at responding to visual cues given by humans about the location of hidden food. \Although this seems to be supported by studies on a range of domesticated (including dogs, goats and horses) and wild (including wolves and chimpanzees) animals, there is also evidence that exposure to humans positively influences the ability of both wild and domesticated animals to follow these same cues. Here, we test the performance of Asian elephants (Elephas maximus) on an object choice task that provides them with visual-only cues given by humans about the location of hidden food. Captive elephants are interesting candidates for investigating how both domestication and human exposure may impact cue-following as they represent a non-domesticated species with almost constant human interaction. As a group, the elephants (n = 7) in our study were unable to follow pointing, body orientation or a combination of both as honest signals of food location. They were, however, able to follow vocal commands with which they were already familiar in a novel context, suggesting the elephants are able to follow cues if they are sufficiently salient. Although the elephants’ inability to follow the visual cues provides partial support for the domestication hypothesis, an alternative explanation is that elephants may rely more heavily on other sensory modalities, specifically olfaction and audition. Further research will be needed to rule out this alternative explanation.

Extraordinary elephant perception

When it comes to people, elephants can be very discriminating, especially those under human care in Southeast Asia. There is the story of the orphaned elephant who, at the age of 10, pulled her drowning mahout out of a lake after hearing his cries for help a kilometer away, or the dangerous 3-m-tall bull who would charge anyone who approached except the 160-cm wife of the village elder, who he would caress with his trunk as she fed him. This behavioral connection between elephants and ourselves is not the product of domestication—in the sense of artificial genetic selection—but born from individual life histories involving near constant human contact (1).

Visual Field Information in the Face Perception of Chimpanzees (Pan troglodytes)

Abstract: Evidence for a visual field advantage (VFA) in the face perception of chimpanzees was investigated using a modification of a free‐vision task. Four of six chimpanzee subjects previously trained on a computer joystick match‐to‐sample paradigm were able to distinguish between images of neutral face chimeras consisting of two left sides (LL) or right sides (RR) of the face. While an individuals ability to make this distinction would be unlikely to determine their suitability for the VFA tests, it was important to establish that distinctive information was available in test images. Data were then recorded on their choice of the LL vs. RR chimera as a match to the true, neutral image; a bias for one of these options would indicate an hemispatial visual field advantage. Results suggest that chimpanzees, unlike humans, do not exhibit a left visual field advantage. These results have important implications for studies on laterality and asymmetry in facial signals and their perception in primates.

Elephants know when their bodies are obstacles to success in a novel transfer task

The capacity to recognise oneself as separate from other individuals and objects is difficult to investigate in non-human animals. The hallmark empirical assessment, the mirror self-recognition test, focuses on an animal’s ability to recognise itself in a mirror and success has thus far been demonstrated in only a small number of species with a keen interest in their own visual reflection. Adapting a recent study done with children, we designed a new body-awareness paradigm for testing an animal’s understanding of its place in its environment. In this task, Asian elephants (Elephas maximus) were required to step onto a mat and pick up a stick attached to it by rope, and then pass the stick forward to an experimenter. In order to do the latter, the elephants had to see their body as an obstacle to success and first remove their weight from the mat before attempting to transfer the stick. The elephants got off the mat in the test significantly more often than in controls, where getting off the mat was unnecessary. This task helps level the playing field for non-visual species tested on cognition tasks and may help better define the continuum on which body- and self-awareness lie.