Evan L Maclean

• PhD
• Professor (Assistant) at University of Arizona

Companion dogs are a powerful model for aging research given their morphologic and genetic variability, risk for age-related disease, and habitation of the human environment. In addition, the shorter life expectancy of dogs compared to human beings provides a unique opportunity for an accelerated timeline to test interventions that might extend hea...

Companion dogs are a valuable model for aging research, including studies of cognitive decline and dementia. With advanced age, some dogs spontaneously develop cognitive impairments and neuropathology resembling features of Alzheimer’s disease. These processes have been studied extensively in laboratory beagles, but the cognitive assays used in tha...

Importance Military members and veterans (hereafter, veterans) with posttraumatic stress disorder (PTSD) increasingly seek psychiatric service dogs as a complementary intervention, yet the effectiveness of service dogs is understudied. Objective To estimate the associations between psychiatric service dog partnership and self-reported and clinicia...

Introduction Service dogs are an increasingly popular complementary intervention for children with autism spectrum disorder. However, despite increasing demand, there remains a lack of empirical research on their potential benefits. The purpose of this study was to evaluate the effects of service dogs on children with autism and their caregivers....

A strong signature of selection in the domestic dog genome is found in a five-megabase region of chromosome six in which four structural variants derived from transposons have previously been associated with human-oriented social behavior, such as attentional bias to social stimuli and social interest in strangers. To explore these genetic associat...

To promote collaboration across canine science, address replicability issues, and advance open science practices within animal cognition, we have launched the ManyDogs consortium, modeled on similar ManyX projects in other fields. We aimed to create a collaborative network that (a) uses large, diverse samples to investigate and replicate findings,...

To promote collaboration across canine science, address replicability issues, and advance open science practices within animal cognition, we have launched the ManyDogs consortium, modeled on similar ManyX projects in other fields. We aimed to create a collaborative network that (a) uses large, diverse samples to investigate and replicate findings,...

Williams-Beuren Syndrome (WBS) is a neurodevelopmental disorder in humans caused by a hemizygous deletion of 28–30 genes and characterized by hypersociability and cognitive deficits. In canines, the homologous chromosomal region shows a strong signature of selection in domestic dogs relative to gray wolves, and four structural variants derived from...

Here, we address Hansen Wheat et al.’s commentary in this journal in response to Salomons et al. Current Biology, 31(14), 3137–3144.E11, (2021). We conduct additional analyses in response to Hansen Wheat et al.’s two main questions. First, we examine the claim that it was the move to a human home environment which enabled the dog puppies to outperf...

Canine cognitive dysfunction (CCD) is a form of dementia that shares many similarities with Alzheimer's disease. Given that physical activity is believed to reduce risk of Alzheimer's disease in humans, we explored the association between physical activity and cognitive health in a cohort of companion dogs, aged 6-18 years. We hypothesized that hig...

Although dogs have a special place in human history as the first domesticated species and play important roles in many cultures around the world, their role in scientific studies has been relatively recent. With a few notable exceptions (e.g., Darwin, Pavlov, Scott & Fuller), domestic dogs were not commonly the subject of rigorous scientific invest...

Psychiatric service dogs are an emerging complementary intervention for posttraumatic stress disorder (PTSD). Initial evidence suggests that partnership with a service dog may be related to less PTSD symptom severity. However, it remains unclear how or why this might occur. To address this gap, we conducted a longitudinal investigation of 82 post-9...

Oxytocin has become a popular analyte in behavioral endocrinology in recent years, due in part to its roles in social behavior, stress physiology, and cognition. Urine samples have the advantage of being non-invasive and minimally disruptive to collect, allowing for oxytocin measurements even in some wild populations. However, methods for urinary o...

During two retreats in 2017 and 2020, a group of international scientists convened to explore the Human-Animal Bond. The meetings, hosted by the Wallis Annenberg PetSpace Leadership Institute, took a broad view of the human-dog relationship and how interactions between the two may benefit us medically, psychologically or through their service as wo...

Canine Cognitive Dysfunction (CCD) is a form of dementia that shares many similarities with Alzheimer’s disease. Given that physical activity is believed to reduce risk of Alzheimer’s disease in humans, we explored the association between physical activity and cognitive health in a cohort of companion dogs, aged 6-18 years. We hypothesized that hig...

Oxytocin has become a popular analyte in behavioral endocrinology in recent years, due in part to its roles in social behavior, stress physiology, and cognition. Urine samples have the advantage of being non-invasive and minimally disruptive to collect, allowing for oxytocin measurements even in some wild populations. However, methods for urinary o...

As the most phenotypically diverse mammalian species that shares human environments and access to sophisticated healthcare, domestic dogs have unique potential to inform our understanding of the determinants of aging. Here we outline key concepts in the study of aging and illustrate the value of research with dogs, which can improve dog health and...

Hypotheses regarding the evolution of uniquely human social cognition often emphasize not only mental state representation, but also mental state sharing. Mental state sharing is evident in instances of joint intentionality – mutual understanding between individuals of each other's simultaneous and interdependent commitment to a shared activity or...

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...

The process of dog domestication likely involved at least two functional stages. The initial stage occurred when subpopulations of wolves became synanthropes, benefiting from life nearby or in human environments. The second phase was characterized by the evolution of novel forms of interspecific cooperation and social relationships between humans a...

Dogs are trained for a variety of working roles including assistance, protection, and detection work. Many canine working roles, in their modern iterations, were developed at the turn of the 20th century and training practices have since largely been passed down from trainer to trainer. In parallel, research in psychology has advanced our understan...

Canine science is rapidly maturing into an interdisciplinary and highly impactful field with great potential for both basic and translational research. The articles in this Frontiers Research Topic, Our Canine Connection: The History, Benefits and Future of Human-Dog Interactions, arise from two meetings sponsored by the Wallis Annenberg PetSpace L...

To promote collaboration across canine science, address reproducibility issues, and advance open science practices within animal cognition, we have launched the ManyDogs consortium, modeled on similar ManyX projects in other fields. We aimed to create a collaborative network that (a) uses large, diverse samples to investigate and replicate findings...

Love and strong social bonds are known buffers in the experience of adversity. Humans often form strong bonds with non-human animals. The human-animal bond refers to a mutually beneficial and dynamic relationship between humans and non-human animals. Previous research suggests that strong bonds with pets may promote resilience in the experience of...

Although non-human primates (NHPs) generally appear to predict how knowledgeable agents use knowledge to guide their behavior, the cognitive mechanisms that enable this remain poorly understood. We assessed the conditions under which NHPs' representations of an agent's awareness break down. Free-ranging rhesus macaques (Macaca mulatta) watched as a...

Although we know that dogs evolved from wolves, it remains unclear how domestication affected dog cognition. One hypothesis suggests dog domestication altered social maturation by a process of selecting for an attraction to humans.1, 2, 3 Under this account, dogs became more flexible in using inherited skills to cooperatively communicate with a new...

Oxytocin has garnered much interest due to its role in affective states, social behaviors, and diverse physiological functions. However, approaches for measuring endogenous oxytocin concentrations have generated considerable controversy and debate. Common procedures for measuring oxytocin often produce uncorrelated results, and the detected concent...

Human cognition is believed to be unique in part because of early-emerging social skills for cooperative communication.¹ Comparative studies show that at 2.5 years old, children reason about the physical world similarly to other great apes, yet already possess cognitive skills for cooperative communication far exceeding those in our closest primate...

Dogs perform a variety of integral roles in our society, engaging in work ranging from assistance (e.g., service dogs, guide dogs) and therapy to detection (e.g., search-and-rescue dogs, explosive detection dogs) and protection (e.g., military and law enforcement dogs). However, success in these roles, which requires dogs to meet challenging behavi...

Dogs exhibit similarities to humans in their sensitivity to cooperative-communicative cues, but the extent to which they are biologically prepared for communication with humans is heavily debated. To investigate the developmental and genetic origins of these traits, we tested 375 eight-week-old dog puppies on a battery of social-cognitive measures....

** Full-text view-only SharedIt link: https://rdcu.be/b9o1a ** While our understanding of adult dog cognition has grown considerably over the past 20 years, relatively little is known about the ontogeny of dog cognition. To assess the development and longitudinal stability of cognitive traits in dogs, we administered a battery of tasks to 160 cand...

While we know that dogs evolved from wolves through a process of domestication, it remains unclear how this process may have affected dog cognitive development. Here we tested dog (N=44) and wolf (N=37) puppies, 5-18 weeks old, on a battery of temperament and cognition tasks. Dog puppies were more attracted to humans, read human gestures more skill...

Oxytocin is a pleiotropic, peptide hormone with broad implications for general health, adaptation, development, reproduction, and social behavior. Endogenous oxytocin and stimulation of the oxytocin receptor support patterns of growth, resilience, and healing. Oxytocin can function as a stress-coping molecule, an anti-inflammatory, and an antioxida...

Trait heritability is necessary for evolution by both natural and artificial selection, yet we know little about the heritability of cognitive traits. Domestic dogs are a valuable study system for questions regarding the evolution of phenotypic diversity due to their extraordinary intraspecific variation. While previous studies have investigated mo...

To characterize the early ontogeny of dog cognition, we tested 168 domestic dog, Canis familiaris, puppies (97 females, 71 males; mean age = 9.2 weeks) in a novel test battery based on previous tasks developed and employed with adolescent and adult dogs. Our sample consisted of Labrador retrievers, golden retrievers and Labrador × golden retriever...

Synopsis Given their remarkable phenotypic diversity, dogs present a unique opportunity for investigating the genetic bases of cognitive and behavioral traits. Our previous work demonstrated that genetic relatedness among breeds accounts for a substantial portion of variation in dog cognition. Here, we investigated the genetic architecture of breed...

Across mammals, increased body size is positively associated with lifespan. However, within species, this relationship is inverted. This is well illustrated in dogs (Canis familiaris), where larger dogs exhibit accelerated life trajectories: growing faster and dying younger than smaller dogs. Similarly, some age-associated traits (e.g., growth rate...

Over two decades of research have produced compelling evidence that non-human primates understand some psychological states in other individuals but are unable to represent others’ beliefs. Recently, three studies employing anticipatory looking (AL) paradigms reported that non-human primates do show hints of implicitly understanding the beliefs of...

In Horschler et al. (Anim Cognit 22(2):187–198, 2019), we found that two components of executive function (short-term memory and self-control) were strongly associated with estimated absolute brain weight across dog breeds, and argued that dogs present a powerful model for studying evolutionary links between cognition and neuroanatomy due to their...

Variation across dog breeds presents a unique opportunity to investigate the evolution and biological basis of complex behavioural traits. We integrated behavioural data from more than 14 000 dogs from 101 breeds with breed-averaged genotypic data (n = 5697 dogs) from over 100 000 loci in the dog genome. We found high levels of among-breed heritabi...

Non-human primates can often predict how another agent will behave based on that agent's knowledge about the world. But how do non-human primates represent others' knowledge states? Researchers have recently proposed that non-human primates form "awareness relations" to attribute objectively true information to other minds, as opposed to human-like...

Since its discovery more than a century ago, oxytocin has become one of the most intensively studied molecules in behavioral biology. In the last five years, Psychoneuroendocrinology has published more than 500 articles with oxytocin in the title, with many of these articles including measures of endogenous oxytocin concentrations. Despite longstan...

Large-scale phylogenetic studies of animal cognition have revealed robust links between absolute brain volume and species differences in executive function. However, past comparative samples have been composed largely of primates, which are characterized by evolutionarily derived neural scaling rules. Therefore, it is currently unknown whether posi...

Assistance dogs can greatly improve the lives of people with disabilities. However, a large proportion of dogs bred and trained for this purpose are deemed unable to successfully fulfill the behavioral demands of this role. Often, this determination is not finalized until weeks or even months into training, when the dog is close to 2 years old. Thu...

Variation across dog breeds presents a unique opportunity for investigating the evolution and biological basis of complex behavioral traits. We integrated behavioral data from more than 17,000 dogs from 101 breeds with breed-averaged genotypic data (N = 5,697 dogs) from over 100,000 loci in the dog genome. Across 14 traits, we found that breed diff...

Working dogs play a variety of important roles, ranging from assisting individuals with disabilities, to explosive and medical detection work. Despite widespread demand, only a subset of dogs bred and trained for these roles ultimately succeed, creating a need for objective measures that can predict working dog aptitude. Most previous research has...

The cover image, by Kelsey Lucca et al., is based on the Short Report The development and flexibility of gaze alternations in bonobos and chimpanzees, DOI: 10.1111/desc.12598. Photo Credit: Jingzhi Tan.

Theory of mind-the ability to reason about the thoughts and emotions of others-is central to what makes us human. Chimpanzees too appear to understand some psychological states. While less is known about bonobos, several lines of evidence suggest that the social-cognitive abilities of the two sister taxa may differ in key respects. This chapter out...

Aggressive behavior in dogs poses public health and animal welfare concerns, however the biological mechanisms regulating dog aggression are not well understood. We investigated the relationships between endogenous plasma oxytocin (OT) and vasopressin (AVP)—neuropeptides that have been linked to affiliative and aggressive behavior in other mammalia...

Oxytocin (OT) and vasopressin (AVP) are neuropeptides with diverse effects on social behavior, cognition and stress responses. Recent studies suggest that OT facilitates and responds to affiliative forms of human–animal interaction (HAI). However, previous studies measuring OT and AVP in dogs have been limited to measures from blood or urine, which...

Background: Oxytocin (OT) and Vasopressin (AVP) are phylogenetically conserved neuropeptides with effects on social behavior, cognition and stress responses. Although OT and AVP are most commonly measured in blood, urine and cerebrospinal fluid (CSF), these approaches present an array of challenges including concerns related to the invasiveness of...

Aggressive behavior in dogs poses public health and animal welfare concerns, however the biological mechanisms regulating dog aggression are not well understood. We investigated the relationships between endogenous plasma oxytocin (OT) and vasopressin (AVP) – neuropeptides that have been linked to affiliative and aggressive behavior in other mammal...

Oxytocin (OT) and Vasopressin (AVP) are neuropeptides with diverse effects on social behavior, cognition and stress responses. Recent studies suggest that OT facilitates and responds to affiliative forms of human-animal interaction (HAI). However, previous studies measuring OT and AVP in dogs have been limited to measures from blood or urine, which...

Background Oxytocin (OT) and Vasopressin (AVP) are phylogenetically conserved neuropeptides with effects on social behavior, cognition and stress responses. Although OT and AVP are most commonly measured in blood, urine and cerebrospinal fluid (CSF), these approaches present an array of challenges including concerns related to the invasiveness of s...

By 2.5 years of age humans are more skilful than other apes on a set of social, but not nonsocial, cognitive tasks. Individual differences in human infants, but not chimpanzees, Pan troglodytes, are also explained by correlated variance in these cooperative communicative skills. Relative to nonhuman apes, domestic dogs, Canis familiaris, perform mo...

Among some haplorhine primates, including humans, relaxed yawns spread contagiously. Such contagious yawning has been linked to social bonds and empathy in some species. However, no studies have investigated contagious yawning in strepsirhines. We conducted an experimental study of contagious yawning in strepsirhines, testing ring-tailed and ruffed...

The neuropeptide oxytocin (OT) is associated with a plethora of social behaviors, and is a key topic at the intersection of psychology and biology. However, tools for measuring OT are still not fully developed. We describe a robust nano liquid chromatography-mass spectrometry (nanoLC-MS) platform for measuring the total amount of OT in human plasma...

A satisfactory account of human cognitive evolution will explain not only the psychological mechanisms that make our species unique, but also how, when, and why these traits evolved. To date, researchers have made substantial progress toward defining uniquely human aspects of cognition, but considerably less effort has been devoted to questions abo...

Previous research has shown that chimpanzees exploit the behavior of humans and conspecifics more readily in a competitive than a cooperative context. However, it is unknown whether bonobos, who outperform chimpanzees in some cooperative tasks, also show greater cognitive flexibility in competitive contexts. Here we tested the cooperative-competiti...

Family dogs and dog owners offer a potentially powerful way to conduct citizen science to answer questions about animal behavior that are difficult to answer with more conventional approaches. Here we evaluate the quality of the first data on dog cognition collected by citizen scientists using the Dognition.com website. We conducted analyses to und...

The emotional-reactivity hypothesis proposes that problem-solving abilities can be constrained by temperament, within and across species. One way to test this hypothesis is with the predictions of the Yerkes-Dodson law. The law posits that arousal level, a component of temperament, affects problem solving in an inverted U-shaped relationship: Optim...

Oxytocin facilitates social connections between humans and dogs [Also see Report by Nagasawa et al. ]

We tested five lemur species-ring-tailed lemurs, ruffed lemurs, mongoose lemurs, black lemurs, and Coquerel's sifakas-(N = 52) in an experiment that evaluated skills for inhibitory control in a social context. First, two human experimenters presented identical food rewards; the "generous" experimenter allowed the subject to eat from her hand, where...

Previous research has shown that chimpanzees exploit the behavior of humans and conspecifics more readily in a competitive than a cooperative context. However, it is unknown whether bonobos, who outperform chimpanzees in some cooperative tasks, also show greater cognitive flexibility in competitive contexts. Here we tested the cooperative-competiti...

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-con...

In a series of four experiments we investigated whether dogs use information about a human's visual perspective when responding to pointing gestures. While there is evidence that dogs may know what humans can and cannot see, and that they flexibly use human communicative gestures, it is unknown if they can integrate these two skills. In Experiment...

The social intelligence hypothesis suggests that living in large social networks was the primary selective pressure for the evolution of complex cognition in primates. This hypothesis is supported by comparative studies demonstrating a positive relationship between social group size and relative brain size across primates. However, the relationship...

The procedure for Experiment 1. (MOV)

The procedure for Experiment 2. (M4V)

Supplemental analyses, Table S1, and Figure S1. (DOCX)

Across three experiments, we explored whether a dog's capacity for inhibitory control is stable or variable across decision-making contexts. In the social task, dogs were first exposed to the reputations of a stingy experimenter that never shared food and a generous experimenter who always shared food. In subsequent test trials, dogs were required...

Humans are believed to have evolved a unique motivation to participate in joint activities that first develops during infancy and supports the development of shared intentionality. We conducted five experiments with bonobos (Pan paniscus) and chimpanzees (Pan troglodytes) (Total n = 119) to assess their motivation to spontaneously participate in jo...

We examined the ability of bonobos, Pan paniscus (N = 39), and chimpanzees, Pan troglodytes (N = 74), to infer the target of an experimenter's visual attention in a series of three experiments. In each experiment subjects were first introduced to a novel object while an experimenter's (E1) visual access to this object was manipulated by (1) having...

Now more than ever animal studies have the potential to test hypotheses regarding how cognition evolves. Comparative psychologists have developed new techniques to probe the cognitive mechanisms underlying animal behavior, and they have become increasingly skillful at adapting methodologies to test multiple species. Meanwhile, evolutionary biologis...

Numerous studies have demonstrated that animals' tolerance for risk when foraging can be affected by changes in metabolic state. Specifically, animals on a negative energy budget increase their preferences for risk, while animals on a positive energy budget are typically risk-averse. The malleability of these preferences may be evolutionarily advan...

Many haplorhine primates flexibly exploit social cues when competing for food. Whether strepsirrhine primates possess similar abilities is unknown. To explore the phylogenetic origins of such skills among primates, we tested ringtailed lemurs, Lemur catta, for their ability to exploit social cues while competing for food. We found that in two conte...

We investigated numerical discrimination and numerical rule-learning in ring-tailed lemurs (Lemur catta). Two ring-tailed lemurs were trained to respond to two visual arrays, each of which contained between one and four elements, in numerically ascending order. In Experiment 1, lemurs were trained with 36 exemplars of each of the numerosities 1–4 a...

The social brain hypothesis proposes that haplorhine primates have evolved relatively large brains for their body size primarily as an adaptation for living in complex social groups. Studies that support this hypothesis have shown a strong relationship between relative brain size and group size in these taxa. Recent reports suggest that this patter...