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Animal behaviour articles within Scientific Reports

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Article Contents

Why the gap, should animal behavior scientists concern themselves with animal ethics, how can animal behavior scientists engage with philosophy and animal ethics, conclusions, acknowledgments, references cited.

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Animal Ethics and Behavioral Science: An Overdue Discussion

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Christine E Webb, Peter Woodford, Elise Huchard, Animal Ethics and Behavioral Science: An Overdue Discussion, BioScience , Volume 69, Issue 10, October 2019, Pages 778–788, https://doi.org/10.1093/biosci/biz082

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Animal ethics—the field of philosophy concerned with the moral status of animals—is experiencing a momentum unprecedented in its history. Surprisingly, animal behavior science remains on the sidelines, despite producing critical evidence on which many arguments in animal ethics rest. In the present article, we explore the origins of the divide between animal behavior science and animal ethics before considering whether behavioral scientists should concern themselves with it. We finally envision tangible steps that could be taken to bridge the gap, encouraging scientists to be aware of, and to more actively engage with, an ethical revolution that is partly fueled by the evidence they generate.

The moral status of animals is a longstanding question dating back at least to Aristotelian philosophy (see Regan and Singer 1989 for an overview of historical and contemporary writings on the topic). However, it has been brought into especially acute focus in recent decades. The modern development of the animal ethics debate is fueled by many factors—among them, novel scientific insights into the complexities of animal minds and emotions (Bekoff and Pierce 2017 ); the ever-increasing scale of industrial farming (Harrison 2013 ); and the Anthropocene, an era heralded by unprecedented human-induced changes to the Earth's climate, environments, and resident wildlife (Ceballos et al. 2015 ). The cumulative impact of these trends has raised the urgency of moral concerns over the nature of human–animal relationships, particularly in the context of our use, overuse, and misuse of animals. The exploitation of animals for food and other products represents just part of the prevailing narrative: A major shift is taking place in how people view the role of animals in research, entertainment, and even companionship. This turn was detectable in early publications such as Ryder's ( 1975 ) Victims of Science and Singer's ( 1975 ) seminal Animal Liberation , and later built on by those of other, scholars (e.g., Regan 1983 , Rollin 1992 , Francione 1995 , Jamieson 2002 , Donaldson and Kymlicka 2011 , Korsgaard 2018 , Nussbaum 2018 ; see also Armstrong and Botzler 2017 for a comprehensive anthology of readings on animal ethics). The last two decades have witnessed an exponential increase in literature and journals focusing on animal ethics—the field of philosophy concerned with the moral status of animals (figure 1 ).

Web-based search results (the grey bars correspond to total number of hits) for the search terms (a) animal ethics as well as (b) human-animal relations in academic articles, and (c) animal rights in US state or federal case law. The ratio of the total number of hits for each search term to the total number of hits for the (control) search term animal during the same decade is illustrated by black lines (see the secondary y-axis) to account for growth in scientific knowledge over time. All information was obtained from Google Scholar on 17 May 2019.

Web-based search results (the grey bars correspond to total number of hits) for the search terms (a) animal ethics as well as (b) human-animal relations in academic articles, and (c ) animal rights in US state or federal case law. The ratio of the total number of hits for each search term to the total number of hits for the (control) search term animal during the same decade is illustrated by black lines (see the secondary y -axis) to account for growth in scientific knowledge over time. All information was obtained from Google Scholar on 17 May 2019.

This transition in ethical thinking about animals has been, at least in part, driven forward by evolutionary theory and discoveries made in the behavioral sciences. Darwin's theory of evolution by natural selection offered a new and powerful challenge to the anthropocentric assumption that humans are the pinnacle of creation—an assumption central to many historically and presently influential theological conceptions of the world (Rachels 1990 ). As the earlier scientific revolution guiding human understanding of the natural order showed that the Earth is not the center of the cosmos, this revolution more dramatically leveled human ontological status by insisting that humans are one of a countless variety of other animals. The disintegration of our pre-Darwinian understanding of nature, coupled today with the extent of anthropogenic changes faced by the environment and animals in the industrial world, has revealed deep-seated incompatibilities between dominant frameworks of value (still rooted in a pervasive sense of human superiority) and the current state of knowledge regarding the capacities of other species and their vulnerability to human actions (e.g., Rachels 1990 , Jones 2013 , Bekoff and Pierce 2017 ).

Recent progress in scientific research on animal behavior has provided evidence used in animal ethics by documenting previously unknown aspects of animal life that have fundamental ethical implications. Studies on the cognitive, emotional, and social capacities of other species have discredited long-held assumptions about capacities thought to be unique to humans. Nonexclusively, this list includes the design and use of tools (Sanz et al. 2013 ); the prevalence of animal cultures (Laland and Bennett 2009 ) and the capacity to innovate (Reader et al. 2016 ); the complexity and efficiency of animal vocal communication, including symbolic communication (Seyfarth et al. 1980 ) and forms of protosyntaxes (Ouattara et al. 2009 ); the capacity for self-awareness (Gallup et al. 2002 ), mental time-travel (Raby et al. 2007 ), and a wide range of emotional experiences, including joy and grief (de Waal 2019 ); reports of complex forms of consciousness, such as empathy (de Waal and Preston 2017 ), and of social intelligence, such as the formation of reciprocal alliances and the active management of long-term social relationships (Cheney and Seyfarth 2007 ), systems of conflict resolution (Aureli and de Waal 2000 ), and the ability to impute mental states to others (Call and Tomasello 2008 ), including the strategic adjustment of one's own knowledge of what others know (Emery and Clayton 2001 ). These findings have all blurred traditional divisions structuring historical discussions of human uniqueness—including the opposition between nature and culture, between animal objects and human subjects, and between instinctive and rational actions—consequently casting doubt on the anthropocentrism that has largely dominated the history of ethics as a field of philosophical inquiry.

Alternative systems of ethical values developed in contemporary animal ethics often rely on empirical evidence to demonstrate the possession (or lack thereof) by a nonhuman individual of the relevant attribute conferring moral consideration (Allen 2006 ). The main theories in animal ethics are pathocentric (i.e., centered on sentience and the capacity to suffer) and therefore hinge on empirical knowledge documenting the sentience of animals—such as recent work demonstrating that fish feel pain (Brown 2015 ). In addition, perceptions of animals as subjects-of-a-life are central to the deontological approach to animal ethics developed by Tom Regan that has also formed a critical part of the legal case for animal rights (Regan 1983 ). Studies revealing the existence of personalities (Sih et al. 2004 ), episodic memory (Clayton et al. 2001 ), intentionality (Allen and Bekoff 1995 ), and rationality (Hurley and Nudds 2006 ) have been instrumental in revealing that animals have a subjective life, personal history, interests, and goal-oriented agency (Jones 2013 ). Taken together, this constellation of results from scientific research on animals has paved the way to changed (and changing) perspectives on the moral status of animals.

Despite these critical contributions to animal ethics, animal behavior sciences such as ethology, behavioral ecology, and comparative psychology have played a rather passive role in the progression and expansion of this movement. In other words, although animal behavior scientists’ work has been integral, it is nonscientists who have primarily pioneered the integration of science and ethics. In the present article, we advance the argument that if the ethics of human–animal relationships are to be redefined, then more active participation on the part of animal behavior scientists has great potential—not just for moving animal ethics debates forward but for scientists themselves. To be clear, in attempting to bridge the study of animal behavior and animal ethics, we are not just referring to the ethics of using animals in behavioral sciences—which have already been the focus of thorough reviews (see box 1 ). Furthermore, although the scientific literature has recently highlighted how animal behavioral sciences can inform animal conservation (e.g., Caro 2007 , Greggor et al. 2016 ) and animal welfare science (e.g., Fraser 1999 , Dawkins 2006 ), it has not yet extended to engage with the full realm of issues debated in animal ethics, which include questions about the fundamental ground of moral status. Our primary purpose in this article is to make that extension by addressing three key questions: What are the primary reasons for a gap between animal behavior science and animal ethics? Should behavioral scientists feel concerned about this growing disconnect? And how could they more actively contribute to the development of animal ethics?

In this article, we emphasize potential interactions between the philosophical field of animal ethics and behavioral sciences. Ethical issues raised by research in behavioral sciences are a related but different and narrower issue. On top of legal requirements, professional organizations have taken further practical steps to ensure that ethical issues related to animal welfare are an integral part of the design of the research being conducted by setting up their own standards (see the Association for the Study of Animal Behaviour's 2012 Guidelines for the Treatment of Animals in Behavioural Research and Teaching, and the American Psychological Association's 2010 Guidelines for Ethical Conduct in the Care and Use of Nonhuman Animals in Research). The main scientific journals in psychology and behavioral ecology require that these standards be met to publish a paper. It certainly does not mean that all ethical issues associated with animal behavior sciences have been thoroughly resolved, and future work should strive to keep ethics questions central to its interests. Empirical work attempting to measure the stress, pain, and mortality caused by study protocols is an emerging field of research (e.g., Le Maho et al. 2011 , Hämäläinen et al. 2014 ), and several recent reviews have been dedicated to these and other ethical issues (Mackinnon and Riley 2010 , Costello et al. 2016 , Field et al. 2019 ).

A primary reason for a frequent lack of communication between animal ethicists and behavioral scientists may reflect traditional difficulties in crossing disciplinary boundaries. Contemporary scientific culture remains largely disconnected from philosophy, which—unfortunately, in our view—is not part of the regular academic training received by scientists; as a result, scientists may not be motivated or prepared to engage in broad ethical discussions that directly pertain to their scientific practice or results. The persistence of a gap is exacerbated institutionally by a lack of educational and career development opportunities that cross-over between behavioral science and philosophy. But it is also caused by fundamentally different theoretical and methodological orientations. Science aims to discover causal relationships between states of affairs and phenomena in the physical world, whereas ethics is an explicitly value-laden, normative field of inquiry that aims to defend our best judgments as to what we ought to do. We are not in the present article proposing a solution to the fact–value relationship or to the problem of whether there are normative facts and how they might fit into nature but, rather, proposing that differences in the basic questions and methods of scientists and ethicists underlie a prevalent, but divisive attitude that science is rigorous and objective, whereas ethical theorizing is more subjective. Acquainting scientists with rigorous debate in normative ethics and ethical theory about basic moral principles and their implications, as well as pointing them to the ways in which scientific research can be enmeshed within the values of the particular times and places in which it is carried out (Kincaid et al. 2007 ), may help emphasize the benefits of interdisciplinary dialogue and research into the complex historical and logical relationship between science and ethics. Greater awareness of the various conceptual and normative assumptions that may come along with different explanatory frameworks can only improve the quality of scientific thinking (Laplane et al. 2019 ).

Although scientists should all be aware of the spectrum of ethical discussions related to their daily scientific practice, they may sometimes fail to see that animal ethics is a broad and fast-growing area of philosophical inquiry and normative debate concerning the nature of human–animal relationships that is built on rational argumentation. It is important to realize that philosophers working on animal ethics may adopt a diversity of nuanced positions, and do not uniformly defend specific political or policy agendas. Scientists may sometimes lump the term animal ethics with other domains, in particular with the set of ethical regulations that rule their research activities (box 1 ), with the emergence of animal welfare or conservation as scientific fields using research to assess and improve the animal condition, or even with the activism incited by animal rights associations. Conversely, although ethicists may be more aware of scientists’ work than the reverse, they may not be up to date with the most current research and debates in the field. Nor do ethicists necessarily have experience rigorously observing animal behavior. Disciplinary segregations between animal welfare scientists, conservation biologists, and (some) animal ethicists are particularly telling examples of the oddity that the divide between these so-called two cultures persists even in the context of obviously shared ethical concerns (Fraser 1999 ). Although the integration of normative and empirical approaches to animal welfare and conservation sciences has eventually gained advocates (e.g., welfare: Fraser 1999 , Dawkins 2006 , Würbel 2009 ; conservation: Ramp and Bekoff 2015 ), scientific discussions of ethical issues have focused on a rather specific set of questions with limited attention to foundational reflection on ethical frameworks and on how normative and empirical approaches relate (Dawkins 2006 ).

The gap between ethicists and behavioral scientists has further been maintained by mutual defiance and skepticism. If and how animals should be used in science has been a core question of animal ethics since its inception, and behavioral research has immediately been the focus of severe criticism for conducting painful and unnecessary experiments (Ryder 1975 , Singer 1975 ). Ethicists subsequently became suspicious toward, or dismissive of, any scientific procedure involving animals to study their behavior (Fraser 1999 ). Meanwhile, animal behavior scientists naturally feared condemnation surrounding their research, and may have perceived the gradual development of ethical regulations on the use of animals in research (box 1 ) as an extra source of constraints and bureaucracy in their work. This divide has likely been furthered by several aspects of the predominant disciplinary culture of animal behavior. Many animal behavior researchers have traditionally adopted the attitude of stifling empathy toward their study subjects in the interest of preserving scientific objectivity and avoiding behavioral interferences with the study subject (Kennedy 1992 ). Such detachment in the name of objectivity may have contributed to the notion that an ethical sensibility toward subjects of research is “unscientific” and “subjective” and may still prevent many researchers from perceiving open engagement with current debates in animal ethics as an integral part of—or at least as compatible with—scientific thinking and practice.

New points of tension have arisen in the course of contemporary discussions in animal ethics, which have for the most part been dominated by two competing approaches: Utilitarian welfare-based and deontological rights-based approaches. Whereas both approaches share the idea that animal welfare is worthy of protection for its own sake and not for the sake of humans, the welfare approach insists that moral duties related to the humane treatment of animals come from animals’ capacity to feel pain and pleasure. It stems from a utilitarian and consequentialist approach to animal ethics, according to which the aggregate benefits of any intervention into animal lives must exceed any harmful costs (Singer 1975 ). In contrast, theories of animal rights are based in deontological ethics, pursuant to which duties to animals come from the respect that they deserve as agents with their own unique interests, aims, and goals. Theorists in this school consider animals’ lives to be intrinsically valuable and propose to grant them basic rights—such as the right to life, freedom, and not to be tortured—to prevent them from being treated as mere means, such that their interests are sacrificed to human interests (Regan 1983 , Francione 1995 , Donaldson and Kymlicka 2011 ). This generates a critical tension with welfare approaches, according to which animals retain an instrumental value in situations in which benefits (to the human community) might outweigh harm (to the animals). It is clear that when animal scientists do engage with ethical debates, the prevailing utilitarian, welfare-based approach is often adopted by default, probably due—at least in part—to the use of animals in scientific research. However, many ethicists have instead favored theories of animal rights (Donaldson and Kymlicka 2011 ), although they have not uniformly condemned the use of animals in research within this framework. Therefore, although the practical implications of such divergences in underlying ethical theory may be profound, rights-based theories do not necessarily exclude the possibility of research on animals. For example, just as human volunteers can participate in scientific experiments, it may be possible to envision a research protocol that respects the dissent of a nonhuman subject (e.g., Fenton 2014 ), especially in behavioral research where experiments can be designed in which animals are free to participate.

A greater integration between animal ethics and the animal behavior scientific community is desirable for ethical and pragmatic reasons. Foremost, there is an ethical reason in that scientists fulfill a social responsibility when they engage with and help others understand the ethical implications of research. However, there are also pragmatic benefits for science, including helping scientists examine sources of historical and cultural bias that may limit scientific questions and approaches, and so further enrich and broaden scientific understanding. Some of these benefits may admittedly arise from interactions with philosophical discussions about the nature of animals that are broader than animal ethics, for example philosophical work on animal minds, perception and representation, social learning and culture, altruism and cooperation, and rationality (Andrews 2015 , Andrews and Beck 2018 ). Nonetheless, the recent renewal of the philosophy of animals as a subfield within the philosophy of science has played and continues to play a major role in the expansion of animal ethics insofar as it has also challenged anthropocentric approaches that have dominated classical philosophy. Although the recognition of animal consciousness and subjectivity is growing, it is not unanimous in philosophy (cf., e.g., Carruthers 2000 , Tye 2016 ). These important debates that are relevant to animal ethics, but are also broader, draw on and require science and should, therefore, be inspiring to animal scientists as well.

Fulfilling a social responsibility

Many scientists are naturally interested in how their results inform and inspire societal debates—one obvious reason for animal behavioral scientists to engage with the literature on animal ethics. In addition to this natural curiosity, and despite commonly holding the view that scientific findings have no intrinsic normative value, scientists still usually support ideas of moral progress that follow from scientific progress in our rational understanding of the natural world. For example, where progress in understanding the neurologic development of infants uncovered the capacity for pain (Anand and Hickey 1987 ), it became an ethical duty for scientists to advocate against neonatal surgeries without anesthesia. The parallels to our understanding of animal pain are obvious, and scientists could play an important role in advocating against farming or research practices that involve suffering in the form of pain as well. More generally, ethics makes a claim on scientists to engage with public debates on ethical issues that are related to their scientific activities (and sometimes even raised by their results; Siekevitz 1970 , Pain 2013 ). At a time when researchers in science and technology are often consulted to set the direction and values of society, and often occupy leadership roles on decision-making bodies, this obligation must increasingly be emphasized. For example, a communication from Mark S. Frankel, director of the Scientific Responsibility, Human Rights, and Law Program at the American Association for the Advancement of Science (AAAS), argues that students and scientists should put less emphasis on their internal responsibility regarding how research should be conducted, and more on their external responsibility by being “vitally concerned” with the influence that their work and knowledge can have on society (Pain 2013 ). Scientists are increasingly required to justify the benefits of their research to society—this is notably the case for individual applications to research positions or funding, as well as for research evaluations at the institutional level—and growing debates on animal moral status spark public interest in animal behavior science. In this context, active engagement with animal ethics could translate to a new and promising applied dimension of scientists’ work—one that is both instrumentally beneficial and aligned with social responsibilities.

Opening the black box of animal minds

Following the vast accumulation of knowledge on animal behavior, scientists are faced with new questions about the nature of animal minds, a crucial topic also at the center of philosophical debate today (Lurz 2009 , Andrews 2015 , Andrews and Beck 2018 ). Nonetheless, the lingering conviction that animal mental states are unknowable—a black box that is inaccessible to science—or irrelevant to the explanation of behavior (e.g., Dawkins 2015 ) has limited scientifically-informed ethical reasoning about animals. For example, Griffin's ( 1998 ) call to bring the study of consciousness to the fore of ethology nearly two decades ago is continually met with considerable resistance (for a historical overview on cognitive ethology, see Allen and Bekoff 2007 ), and some contemporary scientists deny that documenting the degree of animal consciousness is useful in the science of comparative cognition (Shettleworth 2010 ). In addition to this fundamental debate on whether animal consciousness can and should be studied by scientists, some leading approaches in animal behavior science downplay the explanatory significance of animal mental and emotional lives. In particular, behavioral ecologists are traditionally trained to focus on the adaptive value of a trait, favoring ultimate over proximate explanations for behavior. As one example, infanticide is often framed exclusively in terms of evolutionary costs and benefits rather than any underlying emotion or proximate motivation (e.g., see van Schaik and Janson 2009 ). Although these functional evolutionary explanations are valuable in their own right, they offer only a very limited view of animal emotions, capabilities, and agency, and little to no insight into perceptions, intentionality, rationality, or consciousness residing in animal minds. This poses a deeper, more fundamental epistemological problem in the sense that building a whole field of scientific inquiry around what is currently a black box inherently hampers ultimate explanatory and predictive efforts. This shortcoming in turn reveals how explanatory frameworks in the behavioral sciences can quickly overlook or render invisible the very object of moral concern—the organism itself as a potentially sentient entity that can be benefited or harmed—or, at the very least, relegate the organism to secondary status (Walsh 2015 ).

The language that animal behavior scientists habitually employ reflects this deeply entrenched practice (Crist 1999 ). Reducing animal behavior to mechanistic, causal descriptions has reinforced the view of animals as mere objects or vehicles of their genes and environment, preempting any inferences to their mental life or agency (it is noteworthy that in the writings of early naturalists such as Darwin, animals were commonly portrayed as individuals with an array of meaningful subjective experiences and aims). For example, scientists have traditionally used terms such as innate releasing mechanism while habitually relegating complex behavioral phenomena—usually those linked with cognitive or affective capacities—to more parsimonious explanations, further distancing themselves from the animals they study. However, this presupposes that such technical, parsimonious descriptions are also unbiased, and it would behoove scientists to realize that the theoretical language they employ is built on an inherently skeptical bias toward animal subjective and agential traits. The animal ethics literature, which puts animal subjectivity and agency at the heart of its argumentation, places an ethical urgency and burden of proof on mechanistic views of animal behavior in the behavioral sciences to show that animals are not sentient (Birch, 2017 , 2018 ), and to develop more solid inferences about the existence and character of animal subjectivity (e.g., see Smuts 2001 , Godfrey-Smith 2016 ).

Questioning the anthropocentric legacy of behavioral studies

The slow development of cognitive ethology is not merely a consequence of empirical limitations in accessing animal minds or a predominant focus on ultimate explanation in studies of animal behavior. The avoidance of attributing—or even studying—morally-relevant traits such as agency, interests, or motivations and goals to nonhuman animals reflects a more pervasive bias, namely the perceived dangers of anthropomorphism (e.g., Wynne 2004 ). In addition to shaping research questions, experimental settings and interpretations of results traditionally tend to disfavor anthropomorphic hypotheses, according to which similar mechanisms underlie the behavioral similarities observed between humans and nonhumans. This occurs even when studying species that are closely related to us, a revelatory context regarding such a bias, referred to as anthropodenial by de Waal ( 1999 ). According to basic evolutionary principles, the most parsimonious explanation in such cases is the one assuming that similar processes in closely related species emerge from common ancestry (phylogenetic parsimony). A scenario in which the evolution of distinct cognitive processes generates similar behavioral manifestations in closely related species is, in fact, improbable. It is also revealing to note that simple mechanistic explanations are generally favored over phylogenetic parsimony when discussing cognitive capacities, as opposed to physiological or anatomical traits, for which scientists have no problem invoking human–animal similarity (de Waal 1999 ). This bias appears to be a direct, pervasive legacy of the famous Morgan's Canon proposed at the end of the 19th century, which states that “In no case may we interpret an action as the outcome of the exercise of a higher psychical faculty, if it can be interpreted as the outcome of the exercise of one which stands lower in the psychological scale” (Morgan 1894 , p. 53). A large philosophical literature has recently accumulated around the notion of parsimony and related methodological issues (Keeley 2004 , Fitzpatrick 2008 , Sober 2012 , Mikhalevich 2014 , Halina 2015 , Buckner 2017 ), and scientists can benefit from deeper reflection on any bias toward simplicity that is motivated by worries about the dangers of anthropomorphism.

Another upshot of this approach is that the threshold of evidence needed to provide support for a particular cognitive or emotional faculty in other species is much higher than in our own. For example, the definition of animal teaching initially proposed by Caro and Hauser ( 1992 ) has proven so strict that it would exclude many occurrences of human teaching as employed in common parlance (Laland and Hoppitt 2003 ). This anthropocentric perspective has also figured prominently in debates about animal emotions (Bekoff 2009 , de Waal 2019 ); it is not just a remnant of the behaviorist era but still alive today in the form of categorical rejections of anthropomorphism and anecdote. However, when the animal's perspective is carefully considered, anthropomorphic and anecdotal accounts have an important role to play in informing and inspiring rigorous science (Burghardt 1991 , de Waal 1999 , Bates and Byrne 2007 , Godfrey-Smith 2016 ), particularly when it comes to animal mental capacities and emotions (Bekoff 2009 ). Although this form of anthropocentric reductionism is very entrenched in the Western scientific culture in animal behavior, an independent academic tradition emerged in Japan, where anecdotes were valued and where anthropomorphism was not considered a threat (Asquith 1996 , de Waal 2003 ). Despite intense criticisms by Western scientists, Japanese primatologists used individually-based observations—which are now the standard in ethological studies—and made fundamental discoveries in socioecology, such as the existence of tight family bonds structuring animal societies, and the diffusion of socially-learned behaviors throughout animal groups, long before these questions crystallized interest in Western research (Asquith 1996 , de Waal 2003 ). This example illustrates the potential benefits of raising scientists’ awareness of the cultural and cognitive biases that may hamper progress in their discipline. And even though the fear of anthropomorphism may be less present in today's scientific culture than it used to be, the critical stance adopted by thinkers in animal ethics regarding anthropocentric values, combined with their fresh and attentive eye toward animal minds and subjectivity, encourages behavioral scientists who have not already done so to inspect the deeply entrenched sources of biases that inevitably affect their discipline.

Enriching scientific practices

A greater consideration of animal interests and subjectivity may be beneficial pragmatically by changing the way scientists ask questions, design protocols, and interpret animal reactions to experimental conditions. Integrating information about the first-person perspective of the animal is increasingly recognized as important in evolutionary modeling of the effects of natural selection on behavior (e.g., Akçay et al. 2009 ). When designing experiments, careful attention to the animal's perspective on a proposed task can reduce some biases—such as experimenter effects (Despret 2015 ). For example, laboratory mice perceive gender of the experimenter and may consequently modify their behavioral response in an experiment, with male experimenters eliciting a greater stress response than females (Sorge et al. 2014 ). Along similar lines, earlier scholars appreciated that animals live in meaningful and complex worlds, and that adopting the animal's sensory perspective was a necessary precondition for the successful study of behavior (von Uexküll 1992 ). In contrast, subsequent behavioral studies have sometimes failed to adopt such a perspective by designing studies linked to species-specific daily environmental challenges, and so are at risk of making erroneous inferences about animal capacities. For example, dogs were once thought to lack self-awareness because of their failure to pass the mirror self-recognition task, which is strongly biased toward visual species, but they were subsequently found to succeed in passing an olfactory mirror test (e.g., Gatti 2016 ). The role of perspective-taking in animal behavior research is also central to the influential work of philosopher Merleau-Ponty ( 1998 ), who first established a link between animal behavior and phenomenology—which can arguably make a major contribution to both animal ethics (Painter and Lotz 2007 ) and scientific research on animal behavior (Ruonakoski 2007 ) by offering additional insights into animal subjectivity. When it comes to studying animal behavior, Merleau-Ponty ( 1998 ) questions the behaviorist way of interpreting the scientist's role, requiring the scientist's detachment from the study subject. Rather than rejecting anthropomorphism and denying their own sensitivity toward the behavior of study subjects, scientists could acknowledge that human experience, careful observation, and even engaged interaction with animals are the only possible starting points for their investigations, because absolute detachment is impossible in practice (e.g., see Smuts 2001 , Ruonakoski 2007 , Candea 2010 ).

An interdisciplinary dialogue between philosophers, ethicists, and scientists may promote changes in paradigms that could usefully complement traditional approaches and open productive, more holistic avenues to study and understand animal behavior without compromising scientific rigor. In particular, research in cognitive ethology on concepts rooted in classical, anthropocentric philosophy (e.g., self-awareness, empathy, free will, or culture) would benefit from such discussions, which may facilitate the establishment of more inclusive definitions (i.e., applicable to studying nonhuman animals) that retain theoretical and empirical traction. It would further encourage reflection on the most efficient research approaches and the criteria that would provide supporting evidence for the existence of such phenomena in other species. As one example, some philosophers reexamined the state of knowledge regarding behaviors long thought to be unique to humans, such as the capacity to commit suicide (e.g., Peña-Guzmán 2017 ), by lending more weight to animal subjectivity than many scientists traditionally have. These exercises illustrate the potential power of such interdisciplinary dialogues for enriching the perspectives of scientists working on animal behavior while making them more aware of the fact that a collection of scientific observations can lead to divergent interpretative frameworks.

In summary, interactions between the science of animal behavior and animal ethics could have a greater and mutually beneficial scope, addressing questions about what animals are, how we should treat them, and how to envision potential futures for human–animal interactions. The possibility of such a productive exchange between science and philosophy has a strong precedent in the relationship between the science of ecology and environmental philosophy. By generating new scientific knowledge on the interconnectedness and dependence among various forms of life, the field of ecology has also influenced ethical thought. Although it is not uniformly defended by ethicists, ecology has led to calls to regard supraindividual processes, such as ecosystems themselves, as intrinsically valuable and as objects of ethical concern—particularly concerning planetary health criteria grounded in the capacity to sustain and generate biodiversity. Just as in the case of animal behavior science and animal ethics, ecology too has been a source for combating anthropocentrism and generating a more balanced, indeed scientifically-informed, worldview regarding the place of humans as one species embedded within deeply interconnected, interdependent ­living systems (Callicott 1990 ).

In turn, animal behavior scientists can contribute to animal ethics in various ways. Their potential contributions to animal ethics span the full spectrum of scientific ­activities—not only in offering original evidence that fuels theoretical progress in animal ethics, but shaping its practical applications, lending pertinent expertise, and communicating ­effectively with the wider public. There are, however, boundary conditions to what they can offer to ethicists; one prerequisite is that some research involving animals is tolerated—itself a source of disagreement even within the animal ethics community—provided that its costs are minimal and outweighed by clear benefits. As a result, behavioral scientists should carefully take ethical considerations into account when designing their research (box 1 ).

Foster productive interdisciplinary exchanges

An essential first step in this integration could be for behavioral scientists to familiarize themselves with the field of animal ethics (Armstrong and Botzler 2017 provided a comprehensive anthology of readings on animal ethics), which will also cultivate mutual respect and awareness across fields. However, at least to our knowledge, animal ethics, and philosophy more generally, are often absent from animal behavior educational programs and curricula. Reciprocally, academic departments in animal studies are typically housed in social science or humanities faculties, and often lack scientists. This structural separation limits cross-disciplinary exchanges, which could be encouraged by the development of joint teaching, reading groups, research programs, and conferences. Mutual engagement and integrative theory-building could be further fostered by hosting philosophers and ethicists in scientific labs and research groups. Furthermore, several interdisciplinary journals now provide a forum wherein scholars across these disparate fields can comment on topics of mutual interest ranging from animal emotions to the most sound approaches to animal protection legislation (e.g., see Birch 2017 along with associated commentaries).

On gaining meaningful exposure to the basic purview of animal ethics, scientists of animal behavior can further update some of their conceptual frameworks and research practices (as elaborated in the previous section), which may simultaneously foster the endorsement of their findings by nonscientists pursuing related questions. Among the most notable successes in this regard is pioneering work in the area of compassionate conservation, which attempts to appease tensions between scientists who conventionally focus on species and populations and ethicists who typically focus on individuals (Ramp and Bekoff 2015 ).

Produce relevant original evidence

As highlighted above, original evidence stemming from the natural course of animal behavior science has already played a role in inspiring important developments in animal philosophy. Of course, although detailed knowledge concerning the cognitive, affective, and social lives of animals can contribute to our understanding of what is painful to an animal, its degree of sentience and consciousness, the optimal environments in which it thrives, and so on, it certainly cannot tell us what is right or wrong—the central concepts that structure ethical theory and practice. Nevertheless, biological knowledge on the natural behavior of different species, in relation to their phylogenetic position and ecology, can help in setting species-specific criteria for animal ethics agendas. It also has a hand in proposing modes of interactions with animals that are respectful of their physiology and psychology, consistent with a new theory of animal rights that borrows concepts from political philosophy (including citizenship or sovereignty) to envision a new legal frame applicable to the complexity and diversity of animal–human relationships (Donaldson and Kymlicka 2011 ).

In addition, as has already occurred in conservation practice, there is growing pressure for policy decisions concerning animal ethics to be evidence-based, and animal behavior scientists are positioned to contribute data and knowledge that can, at a minimum, inform political decisions regarding the assignment of diverse taxonomic groupings to particular moral categories (Jones 2013 ). The diversity of species that animal behavior scientists study—many of which are beneath the radar of philosophers in favor of a focus on higher vertebrates (with notable exceptions, e.g., Godfrey-Smith 2016 , Tye 2016 )—can raise new ethical concerns and priorities. For example, combined with novel insights on behavioral and cognitive complexity, accumulating scientific evidence that fish feel pain has supported the argument that they be granted similar legal protections to other vertebrates (Brown 2015 ). Similarly, legal personhood campaigns, generally devised to grant legal protections to large mammals, are contingent on evidence concerning capacities such as self-awareness and autonomy (Wise 2000 , Andrews et al. 2018 ). Thompson ( 2019 ) recently outlined how scientists’ work could better position lawyers to build personhood cases, citing four domains—innovativeness, altruism, self-control, and defiance—that would more demonstrably provide evidence for autonomy to the court system. Despite philosophical disagreement over the personhood defense of animal rights (e.g., see Korsgaard 2018 , Nussbaum 2018 ), such communications are important given that scientists do not naturally design their research in light of legal principles and questions.

The need for evidence-based animal ethics frameworks is not to overlook the veritable limits of scientific knowledge. It is important to note that the pace and reach of scientific progress is not always compatible with more immediate ethical decisions, necessitating guidance on what to do in the absence of convincing scientific evidence for aspects of animal sentience (e.g., see Birch 2017 ). Furthermore, burden of proof frameworks must weigh the relative consequences of under- versus overattributing particular mental states to animals, as such estimations can immediately affect welfare and related policy decisions—in particular, the implications of our systematic use of skepticism as the default position should be carefully evaluated (Birch, 2017 , 2018 ). It is also important to acknowledge the diversity of ethical stances toward the weight of scientific evidence—not merely in terms of what is accepted or tolerated, but what is encouraged as the optimal way to understand the complexity of the world around us. There are important philosophical discussions about the sources of our judgment as to whether or not animals have minds and mental lives, with some defending noninferential approaches on the basis of direct experience (see Jamieson 2012 ; see also Bekoff 2009 for an interesting discussion of scientific versus common-sense approaches, which are likely best considered in tandem when it comes to animal ethics).

Regardless of one's position, many scientists in the field of animal behavior spend considerable time observing animals and, therefore, have a wealth of direct, real-world experiences in this regard. A deep understanding of evolutionary theory, allied with the intimate experiences that people who work extensively with animals have, can translate to a unique perspective on animals and human–­animal relationships that ethical debate should capitalize on (see Smuts 2001 , Godfrey-Smith 2016 for pertinent examples). Phenomenologists are particularly interested in such perspectives given their potential to elucidate new realms of being and experience, challenging traditional philosophical views on animal natures and intersubjectivity (Merleau-Ponty 1998 ).

Provide scientific expertise

Scientists’ ability to synthesize and scrutinize academic knowledge has the potential to further guide the public and policymakers in their interpretation of scientific evidence. For example, The Cambridge Declaration on Consciousness (Low et al. 2012 )—prompted by the accumulation of data revealing that humans are not unique in possessing the neurological substrates that generate consciousness—was written by a group of neuroscientists to challenge previously held standards. Animal behavior scientists could similarly consider synthesizing information about species’ intellectual, emotional, and social lives in a format that can be used by decision-makers when drafting or updating ethics policies and legislation, preferably through quantitative meta-analyses and systematic reviews. Conservation and animal behavior scientists recently convened to identify research priorities in animal behavior that promote progress in applied conservation (Greggor et al. 2016 ); a similar exercise could benefit the translation of animal ethics into practical actions. Court cases on animal legal personhood are arising in a growing number of countries, providing a new context in which animal behavior scientists may be expected to act as experts. However, unlike scientists who regularly intervene in court cases and are well aware of the legal culture, such as criminologists or psychiatrists, animal behavioral scientists are generally unprepared for such an exercise. At a smaller scale (and if not already the case), behavioral scientists can join ethics committees to ensure independent representation of animal interests in other scientific fields, and ascertain that ethical concerns are carefully weighed when reviewing articles and grant applications in their own field.

Add a scientific credit to animal ethics in outreach efforts

inally, scientists are often perceived as the authorities on animal behavior, and therefore have the opportunity to inform and engage the public about animal interests. Although it is commonplace for animal behavior scientists to emphasize the conservation implications of their work, other broader impacts related to the moral standing of animals are emphasized relatively less in their public outreach. Increasingly, research on animal behavior has mass public appeal, which opens the door for animal behavior scientists to more actively engage with contemporary animal ethical or philosophical debates and discussions—following the recent tracks of some behavioral scientists (Smuts 2001 , Balcombe 2006 , Bekoff 2009 , Brown 2015 ).

Animal behavioral scientists have much to gain from their academic community's engagement with animal ethics. By jumping into the discussion, scientists also engage more directly with a revolution that has been in part stimulated by their work. Given the rapid rise and foreseeable progress of debates around animal ethics, it is certain that the current generation of animal behavior scientists will have to confront the questions that it raises in the coming decade(s), both as scientists and as citizens. Developing a stronger, more informed and engaged stance that aims to build consensus surrounding questions raised in animal ethics becomes critical to ensure the long-term importance and contribution of their scientific field, to fulfill their moral obligations, and to meet societal expectations by taking part in debates that they are well-positioned to inform. We hope that the present article will encourage this pressing and overdue discussion.

We would like to extend our thanks to many colleagues for insightful comments at various stages of this manuscript's preparation, especially Alice Baniel, Alecia Carter, Marie Charpentier, Becca Franks, Jennifer Jacquet, Alex Lee, and Harry Marshall. We are further grateful to several anonymous reviewers, who provided thorough and constructive feedback, as well as Marc Bekoff for his advice and encouragement. Finally, we would like to express our gratitude to the nonhuman animals with whom we have worked over the years. Those experiences inspired the present article and, most importantly, cultivated a deep personal and professional concern for animal ethics. Contribution ISEM 2019–114.

Author Biographical

Christine E. Webb ( [email protected] ) is an animal behavior scientist in the Department of Human Evolutionary Biology at Harvard University in Cambridge, Massachusetts. Peter Woodford is an assistant professor in religious studies and philosophy at Union College in Schenectady, New York. Elise Huchard is a behavioral ecologist at the Institut des Sciences de l’Evolution de Montpellier, Centre National de la Recherche Scientifique, Université de Montpellier, France.

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A Cognitive Revolution in Animal Research

Animal ‘personalities’ are forcing scientists to rethink basic research.

Six crows wearing six different outfits

This article was originally published in Knowable Magazine .

Several years ago, Christian Rutz started to wonder whether he was giving his crows enough credit. Rutz, a biologist at the University of St. Andrews, in Scotland, and his team were capturing wild New Caledonian crows and challenging them with puzzles made from natural materials before releasing them again. In one test , birds faced a log with drilled holes that contained hidden food; they could get the food out by bending a plant stem into a hook. If a bird didn’t try within 90 minutes, the researchers removed it from the data set.

But, Rutz says, he soon began to realize that he was not, in fact, studying the skills of New Caledonian crows. He was studying the skills of a subset of New Caledonian crows that quickly approached a weird log they’d never seen before—maybe because they were especially brave or reckless.

The team changed their protocol: They gave the more hesitant birds an extra day or two to get used to their surroundings, then tried the puzzle again. “It turns out that many of these retested birds suddenly start engaging,” Rutz says. “They just needed a little bit of extra time.”

More and more scientists are realizing that animals, like people, are individuals: They have distinct tendencies, habits, and life experiences that may affect how they perform in an experiment. That means, some researchers argue, that much published research on animal behavior may be biased. Studies claiming to show something about a species as a whole—the distance that green sea turtles migrate, for example, or how chaffinches respond to the song of a rival—may say more about individual animals that were captured or housed in a certain way, or that share certain genetic features. That’s a problem for researchers who seek to understand how animals sense their environments, gain new knowledge, and live their lives.

“The samples we draw are quite often severely biased,” Rutz says. “This is something that has been in the air in the community for quite a long time.”

In 2020, Rutz and his colleague Michael Webster, also at the University of St. Andrews, proposed a way to address this problem. They called it STRANGE.

Why “STRANGE”? In 2010, an article in Behavioral and Brain Sciences suggested that the people studied in much of published psychology literature are WEIRD—drawn from Western, educated, industrialized, rich, and democratic societies—and are “among the least representative populations one could find for generalizing about humans.” Researchers might draw sweeping conclusions about the human mind when, really, they’ve studied only the minds of, say, undergraduates at the University of Minnesota.

A decade later, Rutz and Webster, drawing inspiration from WEIRD, published a commentary in the journal Nature called “ How STRANGE Are Your Study Animals? ”

Read: Is ‘instinct’ really keeping Flaco the owl alive?

They proposed that their fellow behavior researchers consider several factors about their study animals: social background, trappability and self-selection, rearing history, acclimation and habituation, natural changes in responsiveness, genetic makeup, and experience.

“I first began thinking about these kinds of biases when we were using mesh minnow traps to collect fish for experiments,” Webster says. He suspected—and then confirmed in the lab —that more active sticklebacks were more likely to swim into these traps. “We now try to use nets instead,” Webster says, to catch a wider variety of fish.

That’s trappability. Other factors that might make an animal more trappable than its peers, besides its activity level, include a bold temperament, lack of experience, or simply being hungrier for bait.

Other research has shown that adult female pheasants housed in groups of five performed better on a learning task (figuring out which hole contained food) than those housed in groups of three—that’s social background. Jumping spiders raised in captivity were less interested than wild spiders in videos of prey (rearing history), and honeybees learned best in the morning (natural changes in responsiveness). And so on.

It might be impossible to remove every bias from a group of study animals, Rutz says. But he and Webster want to encourage other scientists to think through STRANGE factors with every experiment, and to be transparent about how those factors might have affected their results.

“We used to assume that we could do an experiment the way we do chemistry—by controlling a variable and not changing anything else,” says Holly Root-Gutteridge, a postdoctoral researcher at the University of Lincoln, in the United Kingdom, who studies dog behavior. But research has uncovered individual patterns of behavior—scientists sometimes call it “personality ”—in all kinds of animals, including monkeys and hermit crabs .

“Just because we haven’t previously given animals the credit for their individuality or distinctiveness doesn’t mean that they don’t have it,” Root-Gutteridge says.

This failure of human imagination or empathy mars some classic experiments, Root-Gutteridge and co-authors noted in a 2022 paper focused on animal-welfare issues. For example, experiments by the psychologist Harry Harlow in the 1950s involved baby rhesus macaques and fake mothers made from cloth or wire. They allegedly gave insight into how human infants form attachments. But given that these monkeys were torn from their mothers and kept unnaturally isolated, the authors ask whether the results are really generalizable. Or do Harlow’s findings apply only to his uniquely traumatized animals?

“All this individual-based behavior, I think this is very much a trend in behavioral sciences,” says Wolfgang Goymann, a behavioral ecologist at the Max Planck Institute for Biological Intelligence and the editor in chief of Ethology . The journal officially adopted the STRANGE framework in early 2021, after Rutz, who is one of the journal’s editors, suggested it to the board.

Goymann didn’t want to create new hoops for already overloaded scientists to jump through. Instead, he says, the journal simply encourages authors to include a few sentences in their methods and discussion sections addressing how STRANGE factors might bias their results (or how they’ve accounted for those factors).

“We want people to think about how representative their study actually is,” Goymann says.

Several other journals have recently adopted or recommended using the STRANGE framework, and since their 2020 paper, Rutz and Webster have run workshops, discussion groups, and symposia at conferences. “It’s grown into something that is bigger than we can run in our spare time,” Rutz says. “We are excited about it, really excited, but we had no idea it would take off in the way it did.”

His hope is that widespread adoption of STRANGE will lead to findings in animal behavior that are more reliable. The problem of studies that can’t be replicated has lately received much attention in certain other sciences—human psychology in particular.

Read: Junk food is bad for you. Is it bad for raccoons?

The psychologist Brian Nosek, the executive director of the Center for Open Science, in Charlottesville, Virginia, and a co-author of the 2022 paper “ Replicability, Robustness, and Reproducibility in Psychological Science ” in the Annual Review of Psychology , says that animal researchers face similar challenges as those who focus on human behavior. “If my goal is to estimate human interest in surfing, and I conduct my survey on a California beach, I am not likely to get an estimate that generalizes to humanity,” Nosek says. “When you conduct a replication of my survey in Iowa, you may not replicate my finding.”

The ideal approach, Nosek says, would be to gather a study sample that’s truly representative—but that can be difficult and expensive. “The next-best alternative is to measure and be explicit about how the sampling strategy may be biased,” he says.

That’s just what Rutz hopes STRANGE will achieve. If researchers are more transparent and thoughtful about the individual characteristics of the animals they’re studying, he says, others might be better able to replicate their work—and be sure that the lessons they’re taking away from their study animals are meaningful, not quirks of experimental setups. “That’s the ultimate goal,” Rutz says.

In his own crow experiments, he doesn’t know whether giving shyer birds extra time changed his overarching results. But it did give him a larger sample size, which can mean more statistically robust results. And, he says, if studies are better designed, it could mean that fewer animals need to be caught in the wild or tested in the lab in order to reach firm conclusions. Overall, he hopes that STRANGE will be a win for animal welfare.

In other words, what’s good for science could also be good for the animals—seeing them “not as robots,” Goymann says, “but as individual beings that also have a value in themselves.”

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Animal Behavior

Reviewed by Psychology Today Staff

The study of animal behavior is a cornerstone of psychology for several reasons. Ethology, or the study of animals in their natural habitats, sheds light on how animals interact with each other and their environments, and why they behave the way they do. By studying animal behavior, humans can also learn more about their own behavior—a field known as comparative psychology.

  • Do Animals Have Thoughts and Emotions?
  • Understanding Animal Behavior
  • Are Pets Good For Your Health?

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Many researchers who study animal cognition agree that animals “think”—that is, they perceive and react to their environment, interact with one another, and experience different emotions, like stress or fear . Whether they are “conscious” in the same way that humans are, however, has been widely debated in both the fields of ethology (the study of animal behavior) and psychology.

Animals can communicate emotion to one another, but this does not qualify as language. Language is an exchange of information using non-fixed symbols (speech). Animals produce innate signals to warn or manipulate other animals (such as the screech of an eagle when it encounters predators). They cannot vary these sounds to create new signals that are arbitrary and content-rich, as do humans.

Charles Darwin with his theory of evolution was one of the first scientists to acknowledge animals’ mental and emotional capacities. Since then, there have been many discoveries of animals that can think : Chimpanzees can make tools and help each other, parrots can talk, newborn chicken can calculate, dolphins can recognize themselves in the mirror, and scrub jays can plan for the future.

Some animal species, such as chimpanzees and goats , are self-aware. They have clearly demonstrated a Theory of Mind —they understand that others have different perspectives, beliefs, and desires, and they can attribute mental states to others as well as themselves.

Among the most intelligent non-human species are chimpanzees, great apes, elephants, New Caledonian Crows, and dolphins.

While scientists haven’t proven conclusively whether animals love, the evidence that they feel grief suggests they can form attachments. Mammals have the same brain areas required to feel emotions as humans do, and bird brains contain similar structures for thinking and feeling. Animals may also go out of their way to spend time with specific individuals when it’s not necessary for their survival—a possible indication of affection.

Many animals will make vocalizations that sound like laughter while playing or for the purpose of social bonding. For instance, domesticated foxes can laugh , a trick they learned by observing people. Additionally, some dog breeds appear to have a sense of humor and will exhibit playful behaviors to amuse humans.

Practically all living creatures shed tears to clear debris and other irritants from their eyes; however, there is some debate over whether non-human animals cry to express emotions, like sadness or grief . Some experts claim that wild animals who cry make themselves vulnerable, so they are more likely to mask their emotions.   

Animals demonstrate through their actions that they are impacted by the loss of a loved one, but it’s unclear whether they understand death or know they’re going to die. Anecdotally, there are examples of animals that hide themselves when it’s time to die, as well as individual animals that kill themselves shortly after a great loss (raising questions about animal suicide ).

A wide range of animal species—including whales, dolphins, horses, cats, dogs, rabbits, birds, elephants, monkeys, and chimpanzees— exhibit grieving behavior after the death of a mate or other member of their family or social group. They might sit motionless, withdraw or seek seclusion, lose interest in food or sex , or remain with the carcass for days.

Tory Kallman/Shutterstock

Animal behavior research is particularly relevant to the study of human behavior when it comes to the preservation of a species, or how an animal’s behavior helps it survive. The behavior of animals in stressful or aggressive situations can be studied to help find solutions for humans in similar circumstances; it may also provide insight for dealing with depression, anxiety , or similar mental health disorders.

Animal-assisted therapy , in which dogs, horses, and other domestic animals help facilitate different forms of therapy, can be helpful for individuals who are socially isolated, living with a diagnosis on the autism spectrum, or suffering from a mood disorder or post- traumatic stress. Interacting with animals has been found to increase humans' levels of oxytocin , a hormone that enhances social bonding. Animal behaviorists are also interested in the ways in which animals themselves may benefit from relationships with humans.

Animal behavior is a result of biology and environment. Behavioral changes are triggered by an internal or external cue, such as the appearance of a threat nearby. Animal responses are driven by the primal urges to survive and reproduce. While animal behavior can vary widely based on the individual, certain behavioral traits, like attention seeking and chasing prey, are genetically inherited, as with dog behavior.

While some animal behavior scholars perform experiments and study animals in a laboratory setting, others advocate watching animals in their natural habitats to get a clearer sense of what they do and how they allocate their time.

Innate behaviors are genetically hardwired and can be performed in response to stimuli without any prior experience. Learned behaviors are acquired by social learning , often by watching and imitating adult members of their species. Through natural selection, animals are more likely to pass on skills that will help their young survive and thrive.

Animals are motivated to fulfill basic survival needs for shelter, food, warmth, and community. Through a combination of genetics and social learning , they acquire skills based on their species’ preferences (e.g., some animals forage, while others hunt). Other animal behaviors include migrating to warmer climates during the winter, establishing a group pecking order, and imprinting on a parental figure. 

Humans share planet Earth with other non-human animals, many of whom are in danger of going extinct . Learning more about animal behavior can help people conserve nature and better coexist with animals. Additionally, observations about animal behavior may provide fresh insights on why people behave the way they do, and how they can change for the better.

Like humans, animals acquire the necessary skills to survive by watching and imitating adult members of their species. Social learning is quicker and more effective than having to figure out how to do something through trial-and-error, and it gives individuals and the species as a whole a better chance at survival.

Inherited behaviors may vary between species and even among individuals. In dogs, for example, many behaviors are strongly inherited , including trainability, aggression towards strangers, attachment , and attention-seeking.

Animals learn from the behavior of more experienced individuals in their family or social group to figure out which behaviors are likely to be punished and which rewarded. They are motivated to avoid pain and seek out pleasure. They can also be conditioned by people to behave in a certain way using a system of rewards and punishments.

Animal sampling is taking a group of animals from a larger population for measurement. The findings are then used to make generalized conclusions about the whole population. Smaller sample sizes tend to be more problematic and prone to error than larger ones.

DoraZett/Shutterstock

Humans and house pets such as dogs have co-evolved ever since humans first domesticated animals some 14,000 years ago. Dogs and cats are beloved creatures the world over and are the lynchpin of a multi-billion-dollar pet product industry.

The so-called pet effect is the widespread belief that owning a pet will make one healthier and happier. This effect may be more anecdotal than reality-based, as many studies find no support or even counter-evidence for the idea that living with a pet enhances human quality of life. In rare cases, pets can transmit serious disorders, such as toxoplasmosis via cat's litter boxes or autoimmune disorders associated with pet birds.

That said, in an era when contact with the natural world is on the decline for many, humans' complex and loving relationships with house pets will endure.

Generally yes. Despite some mixed results in studies, kids with pets seem mostly better off. They have fewer behavioral and learning problems, are less moody, are more physically active, are more obedient, and have improved health overall. They also tend to come from wealthier families and enjoy other socioeconomic advantages.

Pets can provide affection and positive interactions that alleviate feelings of loneliness and isolation. Having a pet can teach children about responsibility and caretaking , while also offering an instant friend and playmate. Pet love can be a powerful emotional resource, particularly during periods of insecurity and self-doubt.

While pets are not a panacea, evidence shows that companion animals can help people who are struggling with mental health problems. They can provide comfort, relieve worry, and ward off a sense of loneliness or isolation; they also prove a source of physical activity and social interactions. Furthermore, caring for a pet can build up a person’s identity and self-worth .

The “Pet Effect” is the rapidly spreading notion that people who have pets live longer and healthier lives. Many people credit their pets with an increase in well-being and health benefits, which include reduced stress, lower levels of cortisol, and higher levels of dopamine and oxytocin, aka the “love hormone.” However, research aimed at proving whether the “Pet Effect” is real remains inconclusive.

When pets are unruly, a person may feel guilty or frustrated that they can’t manage the undesirable behavior. Pets also restrict one’s freedom, making it more logistically difficult to travel or take spontaneous outings. There can be heavy financial and emotional costs to sharing one’s life with a companion animal, including coping with their eventual loss.

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Media and literature tend to focus on high-flying and wildly successful business ventures such as unicorns or gazelles.

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Birds can turn you into an optimist—they can train your mind to wake up every morning expecting to see something beautiful, something miraculous, anywhere and everywhere.

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How to socialize your puppy and to give them safe play opportunities.

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A preschool teacher brings her classroom pet in for emergency veterinary care and explains what her students learn from the unusual animal.

research paper on animal behavior

Dogs want to have fun and teasing is a good way to break the ice and to get to know one another before play-fighting, zooming frenetically here and there, and enjoying themselves.

research paper on animal behavior

Personal Perspective: I feel scared and sad when I watch my rescue dog experience apparent night terrors.

research paper on animal behavior

Miriam Darlington's "Otter Country" is a wonderful book of hope as she reveals the scientific, environmental, and cultural importance of these graceful and elusive beings.

research paper on animal behavior

Assessing potential ADHD in dogs is complex. The use of rating scales can evaluate a dog's temperament, ADHD behaviors, and anxiety levels.

research paper on animal behavior

Factional hatred has a powerful emotional hold over us even though it hurts us practically, psychologically, and somatically.

research paper on animal behavior

Many of the risky behaviors of both men and small animals can be attributed to human inventions—such as roads separating living things from the things they need to survive.

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Perspective article, the new era of canine science: reshaping our relationships with dogs.

research paper on animal behavior

  • 1 School of Anthropology, University of Arizona, Tucson, AZ, United States
  • 2 College of Veterinary Medicine, University of Arizona, Tucson, AZ, United States
  • 3 Cognitive Science, University of Arizona, Tucson, AZ, United States
  • 4 California State Polytechnic University, Pomona, CA, United States
  • 5 Department of Psychology, Western Carolina University, Cullowhee, NC, United States
  • 6 Center for Urban Resilience, Loyola Marymount University, Los Angeles, CA, United States
  • 7 Animal Welfare Science Centre, Faculty of Veterinary and Agricultural Sciences, The University of Melbourne, Melbourne, VIC, Australia

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 Leadership Institute, which convened experts from diverse areas of canine science to assess the state of the field and challenges and opportunities for its future. In this final Perspective paper, we identify a set of overarching themes that will be critical for a productive and sustainable future in canine science. We explore the roles of dog welfare, science communication, and research funding, with an emphasis on developing approaches that benefit people and dogs, alike.

Dogs have played important roles in the lives of humans for millennia ( 1 , 2 ). However, throughout much of scientific history they have been dismissed as an artificial species with little to contribute to our understanding of the natural world, or our place within it. During the last two decades, this sentiment has changed dramatically; canine science is rapidly maturing into an established, impactful, and highly interdisciplinary field ( Figure 1 ). Canine scientists, who previously occupied relatively marginalized roles in academic research, are increasingly being hired at major research universities, and centers devoted to the study of dogs and their interactions with humans are proliferating around the world. The factors underlying dogs' newfound popularity in science are diverse and include (1) increased interest in understanding dog origins, behavior, and cognition; (2) diversification in our approaches to research with non-human animals; (3) recognition of dogs' value as a unique biological model with relevance for humans; and (4) growth in research on the nature and consequences of dog-human interactions, in their myriad forms, from working dog performance to displaced canines living in shelters.

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Figure 1 . Canine science is an interdisciplinary field with connections to other traditional and emerging areas of research. The specific fields shown overlap in ways not depicted here and are not an exhaustive list of disciplines contributing to canine science. Rather, they are included as examples of the diversity of scholarship in canine science.

This Perspective represents the final article in a collection of manuscripts arising from two workshops sponsored by the Wallis Annenberg PetSpace Leadership Institute. Leadership Fellows from around the world gathered in 2017 and 2020 to discuss the state of research and future directions in canine science. The individual articles in this collection provide a detailed treatment of key topics discussed at these events. In this final article, we identify a set of overarching challenges that emerge from this work and identify priorities and opportunities for the future of canine science.

The rise of canine science has benefited substantially from public interest and participation in the research process. Unlike many research studies, which unfold quietly in the ivory towers of research universities, the new era of canine science is intentionally public facing. The dogs being studied are not laboratory animals, bred and housed for research purposes, but rather are companions living in private homes, or assisting humans in capacities ranging from assistance for people with disabilities, to medical and explosives detection. Campus-based research laboratories have opened their doors to members of the public who bring their dogs to participate in problem-solving tasks, social interactions, and sometimes even non-invasive neuroimaging studies. Increasingly, dog owners themselves play a significant role in the scientific process, serving as community scientists who contribute to the systematic gathering of data from the convenience of their homes.

This new research model in conjunction with emerging technologies, makes canine science a highly visible field that engages public stakeholders in unprecedented ways. From a scientific perspective, society has become the new laboratory, and in doing so, has facilitated research with dogs of a scope and scale that was heretofore unthinkable. As tens of thousands of dogs contribute to research on topics ranging from cognition and genetics ( 3 , 4 ) to aging and human loneliness ( 5 ), canine science is entering the realm of “big data” and eclipsing many traditional research approaches. Importantly, these advances are occurring simultaneously across diverse fields of science, creating powerful new opportunities for consilience that will make canine science even more valuable in the years ahead. However, maturing this model toward a sustainable future that serves its diverse stakeholders—who include scientists, research funders, members of the public, and dogs themselves—will require careful navigation of key challenges related to dog welfare, science communication, and financial support ( Figure 2 ).

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Figure 2 . Visual summary of the key issues identified in this Perspective . A sustainable future in canine science will require (1) research approaches that prioritize and monitor the welfare of dogs, (2) improved science communication to avoid incorrect reporting of study results, and to translate research findings to meaningful change in practices relating to dogs, and (3) availability of research funding that is not tied exclusively to studying the possible benefits of dogs for humans.

Dog Welfare

Globally, animal welfare has been linked to the public acceptability that underpins sustainable animal interactions and partnerships ( 6 ). Where human-animal interactions have failed to meet community expectations, practices and in some case entire industries, have been disrupted or ceased. Recent examples include whaling for profit and greyhound racing ( 6 , 7 ). Science is not exempt from this necessity to meet with public expectations and the new era of canine science must place canine welfare at the forefront. Considering dogs as individuals and co-workers, rather than tools for work or subjects, reflects a community moral and ethical paradigm shift that is currently underway. Reimagining our relationship with domestic dogs in research will also help inform our treatment of other animals. In this way, studies of dogs and our interactions with them can serve as a pioneering new model for many areas of science.

As scientists advocate for the revision of community and industry practices with dogs in light of new evidence, we must apply the same criteria to the conduct of our research. This includes adjusting canine research and training methods to acknowledge the sentience of dogs, and the importance of the affective experience for dogs in both research and community settings ( 8 – 11 ). The discipline of animal welfare science has progressed rapidly over the last two decades, and we have many animal-based, welfare-outcome measures available to us ( 6 , 11 ). Ensuring the well-being of the dogs we study will be as critical to ongoing social license to operate (i.e., community approval) for canine science as it is for working dog interests ( 12 ). Being transparent about the issues of animal consent and vulnerability, as well as offering animals agency with regard to their participation in science are valuable suggestions offered within this special issue. We encourage our colleagues to not just consider this paradigm shift, but to effect it through prioritizing and representing the dog's perspective and welfare in their research.

Although increasingly, researchers may include a single or limited set of canine stress measures in studies exploring dogs' potential benefits to humans, this approach alone does not fill the need for studies that prioritize an understanding of canine welfare as their central focus. Canine welfare should be considered not just as an emergent population-level measure ( 13 ) but rather with respect to the way in which it is experienced: from the perspectives of individual dogs. Commonly used statistical methods from human research, such as group-based trajectory analysis ( 14 ) may offer proven techniques that allow meaningful reporting on populations while reflecting the nuance of shared, sub-group patterns. Such approaches will better reflect individual differences, for example variations in canine personality, social support and relationship styles, as well as other significant factors. One impediment to robust measurement of animal welfare in canine science has been limited funding.

We believe that all granting bodies who fund exploration of the possible benefits to people from dogs should also fund and require the canine perspective to be robustly monitored and reported. Impediments to this work arise not from lack of researcher interest or access to dogs, but rather from challenges to securing funding that is independent from a focus on human health outcomes, or other tangible outcomes of work that dogs perform. To be able to optimize canine welfare, there is an urgent need for increased funding specifically to study the welfare of dogs, in all their diversity. The new era of canine science will identify what dogs need to thrive, propelling us toward a mutually sustainable partnership between people and dogs.

Communication

One area that has not received much attention in relation to canine science is the way in which research findings are communicated outside the empirical literature. Fueled by media reports, interest in canine science and the impact of dogs on human health and well-being has grown substantially in the last 10 years. A survey by the Human-Animal Bond Research Institute found that 71% of pet owners were aware of studies demonstrating that pets improve mental and physical health. Some of these claims are justified. For example, many studies have found that interacting with therapy dogs reduces stress and anxiety and increases positive emotional states in a variety of settings including hospitals, schools and nursing homes ( 15 , 16 ). In other cases, high public expectations about the healing power of pets are not matched by the results of empirical studies. For instance, while the Human-Animal Bond Research Institute survey found that 86% of pet owners believe pets relieve depression, the majority of studies on pet-ownership and depression do not support these conclusions ( 17 ).

Because so many people have extensive personal experiences with dogs, investigators face unique challenges in sharing research results with the public. In their hearts, dog owners believe that their canine companions make them feel less depressed, or that dogs feel guilty when they've eliminated indoors or explored the kitchen garbage—even though research might suggest otherwise. In addition, when it comes to animal companions, people much prefer to read a news article in which visits with a therapy dog improved the well-being of a child undergoing chemotherapy than an article about a randomized clinical trial which found no differences between the well-being of children in a therapy dog group and a control group ( 18 ). Nor is there likely to be much press coverage devoted to methodological issues such as small effect sizes and inappropriate attributions of causality to the results of correlational studies.

Canine scientists and scholars of human-animal interactions (anthrozoologists) are fortunate that the public is intrinsically interested in our research. We feel that it is critical for investigators to make efforts to communicate the findings of important studies to the public. We caution however, that researchers should not overstate the implications of their findings in press releases and conversations with journalists, despite frequent pressure to do so. These distortions could have a negative impact on misleading the public and misrepresenting the actual findings, a problem that is particularly acute in canine science where well-intentioned pet owners may eagerly adopt practices based on media coverage of scientific studies. The now-established discipline of science communication offers guidance for how best to engage with community and research stakeholders in meaningful ways.

Traditionally, science communication has relied on the knowledge deficit model of communication ( 19 ). Directionally one-way, the deficit model operates on the assumption that ignorance is the reason for a lack of community support and application of scientific evidence. Examples where practices have not been updated in response to research findings include dog training methodology ( 9 ) and breeding selection for extreme body types, such as brachycephaly in pugs and bulldogs, even though the health and welfare impacts are scientifically well understood ( 20 ). Scientists who share their research results thinking that knowledge disseminated—to “educate” the public—is enough to result in different dog care decisions, industry practices or legislation, will generally find this to be ineffective ( 21 ). This is because the deficit model overlooks the underlying beliefs, existing attitudes and motivations for current practices. We now recognize that the deficit model is not the most effective way to communicate, engage stakeholders and effect change ( 22 , 23 ).

Further exploration of the effect of targeted and intentional science communication, informed by human behavior change research, will improve the translation of canine science to meaningful outcomes for dogs and people alike ( 12 ). This is important, as many studies in canine science have applied aims designed to inform global policies and the creation of best practices ( 24 , 25 ). Applied research from the livestock and farming sector suggests that coordinating human behavior change strategies from social and psychological sciences can influence beliefs and attitudes to motivate changes in the ways people behave toward animals, resulting in improved animal welfare ( 26 – 28 ). In the era of attention economics, where scientists are competing for public attention alongside other diverse media, it is vital that the communication of our work is honest, relevant, and effective, to ensure that our field stays on the radar of key stakeholders, funding bodies and change agents.

A third key challenge in the future of canine science concerns research funding and a careful balancing of the priorities of scientists and funding agencies. In the last decade, canine science has received considerable support from the pet care sector, as well as human health and defense agencies [e.g., ( 29 )]. Fine and Andersen ( 30 ) stress that although funding is still a challenge in human-animal interaction research, there are now more options to be found. In 2008, the Waltham Petcare Science Institute initiated a public-private partnership with the Eunice Kennedy Shriver National Institute of Child Health and Human Development. Over the past decade, this partnership has provided funding for research aimed at measuring the impact of specific Animal-assisted interventions. Since 2014, the Human Animal Bond Research Institute has funded a total of 35 academic research grants investigating the health outcomes of pet ownership and/or human-animal interaction, both for the people and non-human animals involved. Despite clear benefits for enabling research, there remains a limited group of agencies responsible for funding this work. This has potential to constrain the range of topics being studied. In addition, scientists may feel compelled to support the agendas of industry groups, such as those in the pet sector, who often encourage research that will demonstrate the benefits of pets and human-animal interactions.

These constraints were recognized by Wallis Annenberg PetSpace in 2017 when they envisioned their Leadership Institute Program with a mission to promote interdisciplinary scholarship and convene meetings to accelerate research and policy development ( https://www.annenbergpetspace.org/about/leadership ). This model for engagement inspired the organization to offer two invited retreats (2017, 2020) for a total of 33 experts in the field that provided opportunities for open ended and frank discussion about the nature of human-animal interaction research, and the maturing field of canine science. By providing the space and financial support, plus the opportunity to work together and publish, Annenberg PetSpace provided a way to both illuminate current limitations, and to identify priorities for the future, free of constraints from outside interest groups. These intellectual salons have no specific agenda other than to consider the future of the field and what kinds of questions need to be asked based on what we already know. The results of these two retreats include 14 published refereed papers, plus a suite of collaborations that might otherwise not have happened. We hope that these fellowships and retreats continue and inspire others to support similar initiatives so that scholars across multiple disciplines have the opportunity to experience the transformational exchanges that occur during these programs. The new era of canine science will require diverse funding that is not limited to how dogs can benefit humans, from health, safety and economic perspectives. This change will enable researchers the freedom to further our understanding of dogs and their needs for optimized welfare. In turn, this will allow us to identify how dogs and people can thrive together.

Looking Ahead

We hope that the publications emerging from these retreats will reach a diverse community of stakeholders, including students, early career researchers, animal welfare and advocacy groups, legislators and policy makers, philanthropies, and traditional agency funders. The goal of these papers is to spark imagination for projects not yet engaged and to help set the agenda for future research that can enhance our understanding of human-dog interactions and identify paths to ensure a future of symbiotic relationships between these species.

The vision of this collective group of scholars includes the goal of establishing studies with dogs as a sustainable and broad-reaching research focus. Although dogs provide many advantages as a “model species” —including their phenotypic diversity, and shared environments and evolutionary history with humans—a research model centered around dogs has many additional benefits. Dogs provide a rich, interactive and sentient model with deep implications for the way scientists approach animal research, and animal welfare. Dogs also increase the accessibility of research, both literally, due to their ubiquity and opportunities for large-scale public participation in research ( 31 , 32 ), and figuratively, through a body of work with appeal to the broader public.

The field of canine science has much in common with a similar emerging science, that of urban ecology. Humans are historically at the core of the subject material, but non-human elements are often the focus of the study. As such, the work is always culturally embedded, relevant to a variety of stakeholders, and ultimately expected to improve quality of life. The urban ecologists coined a term Use-Inspired Research ( 33 ) from modifying the existing idea of Pasteur's Quadrant which organizes research questions across the axes of fundamental understanding and considerations of use ( 34 ). Both canine research and urban ecology seek fundamental understanding, but also expect to directly apply the knowledge gained to improve outcomes for their subjects and stakeholders.

By including the public in canine science we not only increase the quantity of the data that we can gather, we serve as ambassadors for a new model of responsible animal research. The result increases the value of human-animal interaction research and creates opportunities for the next generation of interdisciplinary scientists. The goal of this collection has been both to highlight specific recent advances in canine science as well as to identify emerging and overarching issues that will shape the future of this field. The multidisciplinary nature of our work with dogs allows scientists to contribute to a robust research agenda, enhancing our understanding of canines and their impact on society. Ultimately, the nexus of our discoveries should have profound effects on reshaping and enriching our relationships with dogs.

Data Availability Statement

The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author/s.

Author Contributions

All authors listed have made a substantial, direct and intellectual contribution to the work, and approved it for publication.

We thank Wallis Annenberg PetSpace for supporting the open-access publishing fees associated with this manuscript.

Conflict of Interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Keywords: canine science, dog, animal welfare, human-animal interaction, science communication, funding, sustainability

Citation: MacLean EL, Fine A, Herzog H, Strauss E and Cobb ML (2021) The New Era of Canine Science: Reshaping Our Relationships With Dogs. Front. Vet. Sci. 8:675782. doi: 10.3389/fvets.2021.675782

Received: 03 March 2021; Accepted: 11 June 2021; Published: 15 July 2021.

Reviewed by:

Copyright © 2021 MacLean, Fine, Herzog, Strauss and Cobb. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY) . The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Evan L. MacLean, evanmaclean@arizona.edu

This article is part of the Research Topic

Our Canine Connection: The History, Benefits and Future of Human-Dog Interactions

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AP®︎/College Biology

Course: ap®︎/college biology   >   unit 8, intro to animal behavior.

  • Innate behaviors
  • Learned behaviors
  • Animal communication
  • Animal behavior: foraging
  • Responses to the environment
  • Animal behavior includes all the ways animals interact with other organisms and the physical environment.
  • Behavior can also be defined as a change in the activity of an organism in response to a stimulus , an external or internal cue or combo of cues.
  • To fully understand a behavior, we want to know what causes it, how it develops in an individual, how it benefits an organism, and how it evolved.
  • Some behaviors are innate , or genetically hardwired, while others are learned , or developed through experience. In many cases, behaviors have both an innate component and a learned component.
  • Behavior is shaped by natural selection. Many behaviors directly increase an organism's fitness, that is, they help it survive and reproduce.

Introduction

What is behavior.

  • Ethology is a field of basic biology, like ecology or genetics. It focuses on the behaviors of diverse organisms in their natural environment.
  • Comparative psychology is an extension of work done in human psychology. It focuses largely on a few species studied in a lab setting.

Four questions to understand a behavior

  • Causation—What causes the behavior? What triggers the behavior, and what body parts, functions, and molecules are involved in carrying it out? Example: Singing is triggered in zebra finches by social cues, such as the proximity of a potential mate, as well as the appropriate hormonal state. The ability to produce songs is influenced by male hormones and occurs mainly in male birds. Songs are produced when air flows from air sacs in the bronchii through an organ called the syrinx. Certain parts of the brain control song production and are well-developed in male zebra finches.
  • Development—How does the behavior develop? Is the behavior present early in life? Does it change over the course of the organism's lifetime? What experiences are necessary for its development? Example: Young male zebra finches first listen to the songs of nearby males of their species, particularly their fathers. Then, they start to practice singing. By adulthood, male zebra finches have learned to produce their own songs, which are unique but often have similarities to those of their fathers. Once a finch has perfected its song, the song remains fixed for life. A diagram of 3 zebra finches. On the bottom of the diagram is a zebra finch labeled male and to the right of it there is a zebra finch labeled female. There is a red arrow pointing from the female towards the male with the label mate above the arrow. Above the male zebra finch is a blue arrow pointing upwards towards a zebra finch with a musical note above its beak. The blue arrow is labeled tutor. From the zebra finch on the top of the diagram there is a black arrow pointing back to a musical note that is above the male finches head. Image credit: modified from Songbird species recognition by Petra Deane, CC BY 1.0
  • Function/adaptive value—How does the behavior affect fitness? How does the behavior affect an organism's chances of survival and reproduction? Example: Singing helps male zebra finches attract mates, increasing the chances that they will reproduce. Singing is part of an elaborate courtship ritual that entices the female to choose the male.
  • Phylogeny—How did the behavior evolve? How does the behavior compare to those of related species? Why might it have evolved as it did? Example: Almost all species of birds can make vocal sounds, but only those in the suborder Passeri are songbirds. Relative to the zebra finch, other songbird species differ in the timing of their listening and practicing phases, the plasticity of song over their lifetimes, the extent to which the song is similar among individuals of the species, and the way that singing is used—for example, for defense of territory vs. courtship of mates.

Cues that trigger behavior

  • In hibernation , an animal goes into a den or burrow, reduces its metabolic rate, and enters a state of inactivity during the winter, conserving resources while conditions are harsh and food is scarce. Environmental cues often trigger hibernation behavior. For instance, brown bears enter their den and hibernate when temperature drops to 0 o ‍   C and snowfall begins. 2 ‍  
  • Estivation is similar to hibernation, but it occurs during the summer months. Some desert animals estivate in response to dry conditions. This shift helps them survive the harshest months of the year. 3 ‍   The snails in the photo below climb to the tops of fence posts to estivate. A photograph of a row of fence posts along a gravel road. At the top of the fence posts are several snails attached. Image credit: Kadina snails climb fence by Vladimir Menkov, CC BY-SA 4.0
  • Migration is a behavior in which animals move from one location to another in a seasonal pattern. For instance, monarch butterflies living in the northern and central United States migrate to Mexico in the autumn, where they spend the winter. Environmental cues that trigger the autumn migration include air temperature, day length, and food availability. 4 ‍  

Innate vs. learned behaviors

  • Innate behaviors are genetically hardwired and are inherited by an organism from its parents.
  • Learned behaviors are not inherited. They develop during an organism's lifetime as the result of experience and environmental influence.

Mostly innate behaviors

Partly innate, partly learned behaviors, mostly learned behaviors, check your understanding.

  • (Choice A)   Innate A Innate
  • (Choice B)   Learned B Learned
  • (Choice C)   Mostly learned, with a small innate component C Mostly learned, with a small innate component

Natural selection shapes behavior.

  • Baby birds of many species instinctively open their mouths for food when the mother returns to the nest. 8 ‍   Birds with this heritable behavior will tend to get fed more—and thus survive to adulthood more—than those that don’t.
  • Mother greylag geese instinctively roll eggs back into the nest if they fall out. 8 ‍   Geese with this heritable behavior will tend to have more offspring that survive to hatch than geese without the behavior.
  • Zebra finch males learn songs while they are juveniles, young birds, and they use these songs in courtship rituals. Birds with the heritable tendency to learn a song will obtain a mate more often than those that don't.

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Searching for Animal Sentience: A Systematic Review of the Scientific Literature

Simple summary.

The emotional lives of animals is often doubted and questioned. Due to the subjective nature of animal emotions, many think that they are out of the reach of scientific measurement. In this systematic review, of over two decades of scientific literature, we found that this was not actually the case. By using a list of keywords, formed of both positive and negative emotions, and terminology relating to animal sentience, we reviewed the scientific literature. We found that the subjective lives of animals are not only a vital part of human medical research but are regularly measured and studied with scientific rigor.

Knowledge of animal sentience is fundamental to many disciplines and imperative to the animal welfare movement. In this review, we examined what is being explored and discussed, regarding animal sentience, within the scientific literature. Rather than attempting to extract meaning from the many complex and abstract definitions of animal sentience, we searched over two decades of scientific literature using a peer-reviewed list of 174 keywords. The list consisted of human emotions, terminology associated with animal sentience, and traits often thought to be indicative of subjective states. We discovered that very little was actually being explored, and instead there was already much agreement about what animals can feel. Why then is there so much scepticism surrounding the science of animal sentience? Sentience refers to the subjective states of animals, and so is often thought to be impossible to measure objectively. However, when we consider that much of the research found to accept and utilise animal sentience is performed for the development of human drugs and treatment, it appears that measuring sentience is, after all, not quite as impossible as was previously thought. In this paper, we explored what has been published on animal sentience in the scientific literature and where the gaps in research lie. We drew conclusions on the implications for animal welfare science and argued for the importance of addressing these gaps in our knowledge. We found that there is a need for more research on positive emotional states in animals, and that there is still much to learn about taxa such as invertebrates. Such information will not only be useful in supporting and initiating legislative amendments but will help to increase understanding, and potentially positive actions and attitudes towards animals.

1. Introduction

“Animals are like robots: they cannot reason or feel pain” (Descartes, 1596–1650). This quote may seem outdated when we consider both when it was said and what contradictory scientific evidence we have garnered since. However, when you consider that many non-human animals (hereafter referred to as animals) are treated inhumanely on a daily basis for the purpose of food, entertainment, research, and profit, the quote still seems relevant today. What stops us from taking the humane approach to agriculture, and what stops us from banning animal cruelty for entertainment? The arguments are often multi-faceted; disbelief or unawareness of animal suffering, lust for profit, or lack of empathy brought about by historical processes and layers of discourse around the moral value of animals [ 1 ]. Developing and sharing knowledge of animal sentience are key to addressing these arguments. Animal sentience refers to the ability of animals to feel and experience emotions such as joy, pleasure, pain and fear. It is animals’ capacity to feel both positive and negative states that drives the animal welfare movement and is the reason why animal protection laws exist [ 2 , 3 , 4 ].

Originally, concern for animals focused primarily on the animals’ physical health, with little thought for their mental well-being [ 5 ]. However, scientific interest in the subjective experience of animals has noticeably increased in the last 10 to 20 years (see [ 4 ] for a review). Animal sentience is sometimes dismissed due to the subjective nature of emotions and feelings; the building blocks of animal sentience, e.g., [ 6 , 7 , 8 ]. Whereas others argue that the complex and subjective nature of sentience should not be reason for its denial or dismissal as a robust science [ 4 , 9 , 10 ]. We feel that although sentience refers to subjective states it is not alone, as so does much of human psychology. The emotional experience of humans is both a personal experience and subject to false reporting [ 4 ]. We do not deny that humans are sentient because of this, but many do question animal sentience on the same basis. It appears therefore, that animal sentience is an unlucky victim of this scientific paradox. Whilst other areas of science will often make do with imperfect data, animal sentience is required to buck the trend and provide unequivocal proof [ 11 , 12 ]. Neuroscientist Donald Griffin coined the term “Paralytic perfectionism” to describe this contradictory way in which scientists still demand absolute certainty before they can accept animal sentience. He argued that the successful interpretation of mental states in others is a vital tool for social interactions, for both humans and animals [ 13 ].

Despite being subject to debate, descriptions of animal sentience, albeit in various forms, exist throughout the scientific literature. In fact, many experiments rely upon their animal subjects being sentient [ 14 ]. Analgesia studies for example, require animal models to feel pain, and animal models of schizophrenia are tested for a range of emotions such as fear and anxiety. Furthermore, there is a wealth of scientific studies, laws and policies which look to minimise suffering in the very animals whose sentience is so often questioned [ 15 , 16 , 17 ]. To overcome the paradoxical nature of the science of animal sentience, we sought to understand what is accepted and known about animal sentience in the scientific literature. The first challenge was to address the lack of consensus in regards to the definition of sentience. There is no universally accepted definition of sentience, and there are many different opinions as to where sentience exists in the animal kingdom [ 2 , 18 ]. We dealt with this by aiming to be as holistic as possible. The result was a peer-reviewed list of keywords comprised of primary and secondary emotions, technical terms, and traits commonly thought to be indicative of sentience. We were not intending to prove the strength or validity of these keywords in defining or proving animal sentience, but we instead wished to review what has been explored and discussed regarding the subjective states of animals.

1.1. The Positive Side of Sentience

Although today, the subjective experiences of animals receive considerably more attention than 50 or even 20 years ago, research is still focused on the negative experiences of animals [ 19 ]. Whilst this research has been fundamental in improving many practices involving animals, it has failed to take into account the importance of positive experiences and emotions to the well-being of animals [ 20 ]. In more recent years, scientists have slowly begun to recognise that positive emotions and experiences are also a fundamental area of animal welfare science and key to ensuring a good state of animal welfare [ 5 , 20 , 21 , 22 , 23 , 24 ]. The emergence of new disciplines such as ‘Positive Psychology’ [ 25 , 26 ] and ‘Affective Neuroscience’ [ 27 ], which refers to both positive and negative effects, is evidence of this new focus. Progress continues to be slow however, and scientific understanding of negative emotions far outweighs that of positive emotions, both in animals and humans [ 20 ]. In this study we aimed to review what is assumed and explored in the scientific literature in regards to the positive and negative aspects of animal sentience and the impact this has on animal welfare.

1.2. Mammalcentrism

Animal sentience research is often accused of being mammal-centric. This is primarily due to the similarity of physiology and neurology in humans and other mammals, and the relative ease of drawing conclusions from argument-by-analogy [ 12 , 28 ]. In addition, attitudes to animals may be affected by innate human tendencies to sympathise with animals depending on their status, use, attractiveness, or believed intelligence [ 29 , 30 ]. Yue-Cottee for example, describes how cold-bloodedness is often used as a reason for the denial of subjective feelings to fish. She argues that a metabolic difference should not be used as a reason for denying them concern or protection, particularly in light of the contradictory scientific evidence [ 12 ]. There is hope, however, and science is slowly moving away from this dominant, mammalcentric perspective. For instance, in recent years we have seen a growing focus on the subjective minds of invertebrates such as cephalopods and decapod crustaceans [ 31 , 32 , 33 ]. As the field of animal sentience research continues to grow, scientists should be able to further develop the methodologies used to explore the affective states of animals. The resulting increase in scientific knowledge on the abilities of animals will hopefully help to change people’s perceptions of animals and will have varying implications for practices and industries. In this study we have reviewed articles published from 1990 to 2012 and identified the taxa being studied. This has allowed us to evaluate the progression of research and knowledge of animal sentience, understand what is known about the different taxa, and to identify the remaining gaps in our knowledge.

1.3. Humane Research

Whether a study design impairs the welfare of the animal subjects remains one of the greatest ethical paradoxes of animal sentience research. Although many studies using animals will have been subjected to some level of ethical review, this does not necessarily mean the study has not significantly impaired the welfare of the animals involved. For example, methodologies involving inhumane procedures can be approved due to the potential of the results to justify the suffering [ 34 ]. Furthermore, a lot of un-moderated animal research still continues around the world [ 35 , 36 ]. It is likely that this situation will improve as the focus of animal sentience and welfare research shifts on to the study of positive emotional states. The objective of such studies would encourage the promotion and evaluation of positive emotions, rather than negative ones.

When research must involve animals, one possible change is to address how the animals are housed for these studies. The issue of housing has received a lot of attention in terms of enriched cages and naturalistic settings [ 37 , 38 ], but there is even greater scope for improvement when you address the issue of housing and breeding as a whole. For instance, when research aims to explore animal behaviour for greater ethological understanding, there are many alternatives to laboratories that should at least be explored. For example, existing populations of pet animals or animals in shelters, zoos, farms, or in the wild can often provide the subjects required for research. In fact, such populations can provide a more realistic model of the species than a laboratory bred animal [ 39 ]. In this study we documented where the animals were housed or where the studies took place, for example, were they zoo or laboratory animals? We also recorded the main purpose of the study, for instance, did the study seek to develop knowledge of animal behaviour or improve animal welfare? We then examined the relationship between these data to understand how the animals were housed for each of the main purposes and we drew conclusions regarding the potential for welfare improvements.

1.4. The Importance of Animal Sentience Research

Understanding animal sentience has many benefits to humans, animals and science. Too much scepticism, particularly when unfounded, hinders scientific process and positive change for animals [ 40 , 41 ]. Furthermore, accepting the existence of affective states in animals can be an important step towards tackling other key problems in neuroscience [ 14 ]. The many parallels between the subjective experiences of animals and humans are clearly utilised in research that requires animal models for human afflictions [ 27 ]. Most importantly, knowledge of what animals experience, what is important to them, and what constitutes a good life for them, is key to truly improving their welfare. Just like for humans the experience of positive emotions, such as joy and pleasure, has meaningful bearings on the mental and physical welfare of animals [ 16 , 42 ].

We sought to address the lack of consensus on the prevalence of animal sentience by extensively reviewing the scientific literature. We analysed the progression of published research discussing and exploring various aspects of animal sentience over a focal period of 22 years. The results highlight what is being explored and what is already assumed in regards to animal sentience, and in which taxa. As the human population continues to grow so does the number of animals we use for our own means. Understanding the subjective minds of animals is therefore of utmost importance to their welfare. We hope that the findings of this paper can highlight where future research is needed in the field of animal sentience and the importance of what we already know.

2. Materials and Methods

2.1. keywords.

We compiled a list of emotions, traits, and terminology associated with or indicative of animal sentience using three existing lists of human emotions [ 43 , 44 , 45 ], and 22 keywords specific to animals and animal sentience ( Appendix Table A1 ). These words were derived from literature reviews performed prior to the start of the study. Each keyword was extensively defined to ensure only reference to the subjective experiences of animals was considered in the review. The final list of 174 keywords was then peer-reviewed and approved by a scientist in the field of animal sentience [ 46 ].

2.2. Literature Search

We searched two journal databases; Science Direct and Ingenta Connect, for articles from peer reviewed journals, indexed since 1990, containing both the keyword, and the word ‘animal’ in the abstract, title or keywords. The focal period of 1990 to 2012 was chosen because it allowed for a large and recent study period, yet it was still feasible given our time restraints. We then filtered the results according to the following criteria. Firstly, we removed any books, short communications, letters, non-English articles, review papers, and articles without abstracts, leaving only original, full research articles. Secondly, we removed any articles that were not using animals but were only referring to previous studies or findings from animal research. Finally, we only retained articles that used the keyword in line with the detailed definition and in reference to the animals’ subjective state. For example, stress as an emotional state was recorded, whereas reference to stress as a physiological state, such as heat stress, was omitted.

Each of the authors took part in collecting the data, and so to ensure consistency, each keyword and category used in the study was fully defined with working examples to reduce the degree of subjectivity. Furthermore, inter-observer reliability tests were performed for each aspect of the data collection (e.g., article selection and categorisation) throughout the study period. Reliability exceeded 95% agreement upon each of the tests.

2.3. Research Questions

After the initial sorting phase, we answered a number of questions for each article abstract. To start with we looked at whether the study assumed or explored the existence of the keyword in the animal subjects. For example, a study could explore whether rats can experience pain, or it could measure the pain experienced by rats following analgesia. The latter accepts that rats can feel pain and uses that knowledge, whereas the former is exploring whether or not rats can experience pain at all. Both types of study were reviewed, in order to measure the acceptance of animal emotions in the scientific literature and to establish which aspects of animal sentience have been experimentally explored.

To determine the number of articles referring to positive and negative keywords, we labelled each of the keywords as positive, negative or neutral, depending on the valence of the emotion or trait depicted. For example, the keyword pain was labelled as negative, whereas the keyword pleasure was positive. For the neutral keywords the valence was defined at the individual article level wherever appropriate. For example, the use of the term ‘affective state’ in a study could have referred to either a negative or positive affective state, or both, whereas the keyword ’theory of mind’ had no valence and remained neutral.

We then asked which year the article was published. When analysing this question we only looked at the data returned from the years 1990 to 2011. This was because the 2012 results were not representative of the entire year due to the timings of the data collection, which took place in mid-2012. All of the other questions looked at the entire 1990–2012 period.

To determine whether any observed differences were unique to the articles reviewed or merely reflective of the general trends in publication numbers, we looked at the total number of articles published in Ingenta Connect and Science Direct in the years 1990 and 2011. For consistency we used the same search criteria as before but without the keyword. For example, an advanced search was performed in both databases to determine the total number of papers published in 1990 with the word ‘animal’ in the title, abstract or keywords. We then determined the percentage increase or decrease between these years for both the total number of papers published and for our reviewed papers.

The remaining questions probed for further details of the animals used in the study. We looked at which taxa were being studied, recording the sub-phylum, order, class and species or common name of the animals used in each study. When possible we identified the experimental setting of the study from the article abstract. For example, did the research take place in a laboratory, a zoo, or on a farm? Research farms were labelled as ‘farms’, due to the similarity in the housing environment for the animals. Finally, we determined what the primary purpose of the study was, recording whether the research was performed for human benefit, such as a pharmaceutical study, to advance knowledge of animal behaviour, to further knowledge of animal sentience, or to improve animal welfare.

2.4. Data Analysis

We organised the data into two spreadsheets; version one (V1) was the original intact spreadsheet, and version two (V2) had the duplicate articles removed (some articles referred to more than one keyword). We used V1 for the analyses that looked at individual keywords, such as the number of articles returned for each keyword. Finally, we used V2 for the analyses that required us to look at the data set as a whole without the duplicate entries. For example, the number of articles published in 1990 vs. 2011.

The primary analysis was descriptive to allow us to review the relationships between the different research questions and to identify appropriate sample sizes for statistical analysis. Following this we used the chi-square goodness of fit test to identify significant differences between the number of assumed and explored articles, the numbers recorded for each sub-phylum, the purpose of the studies, the experimental setting, and the numbers of articles published in 1990 compared to 2011. All analyses were performed using Statistical Package for the Social Sciences (SPSS) version 21 for Windows. Statistical significance was indicated by P < 0.05.

We collected a total of 2,804 papers from all of the searches performed; dropping to 2,562 once the duplicate entries were removed. Forty-three keywords out of the total 174 returned suitable results, ranging in number from one to 635 articles per keyword. From these keywords, eight were labelled positive, 23 were negative, and 12 were either neutral or dependent upon the individual article.

Animal sentience was not the primary reason for why any of the studies were performed, and it was only deemed to be a secondary or subsequent purpose for five of the articles we reviewed. Instead, we found there to be three over-arching reasons for the studies, and these were; human benefit, animal welfare and animal behaviour. Significantly more studies were performed for human benefit (e.g., pharmaceutical development), than there were for either animal welfare or animal behaviour reasons ( X 2 = 1,462.34, df = 2, P < 0.001). There were also significantly more studies performed for animal welfare reasons than there were for animal behaviour reasons ( X 2 = 9.94, df = 1, P < 0.05).

We captured detailed information about the animals used for each article, and found that, overall, vertebrates (n = 2,519) were used significantly more than invertebrates (n = 32, X 2 = 2,424.61, df = 1, P < 0.001). These two sub-phyla were comprised of 12 taxonomical classes; six vertebrate and six invertebrate. Mammalia was the most popular class of animals used (n = 2,346, 91.89%), followed by Aves (n = 116, 4.54%), and Actinopterygii (n = 45, 1.76%). Climbing down the taxonomical tree we found that these classes gave way to 57 orders, 11 of which were invertebrates, and the remaining 46 were vertebrates. The top five orders and species are shown in Figure 1 , Figure 2 .

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The number of reviewed articles using each of the top five orders. Data labels refer to the percentage of the total articles.

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The number of reviewed articles using each of the top five species or common names. Data labels refer to the percentage of the total articles.

Because the human benefit studies comprised the majority of the articles we reviewed (n = 1,765), we also looked at the results with those articles removed to see whether there were any differences in the returned results. We found no differences in the use of vertebrates and invertebrates, with the majority of studies still using vertebrates (vertebrates: n = 766, 96.47%, invertebrates: n = 28, 3.66%). Mammalia, Aves, and Actinopterygii were still the most popular classes used (Mammalia: n = 610, 76.73%, Aves: n = 110, 13.84%, Actinopterygii: n = 36, 4.53%). However, there was a difference for the orders; Rodentia, which were used for 69.07% of the articles overall, were only used for 9.91% of the articles once the human benefit studies were removed. The top five orders changed to Artiodactyla (n = 277, 35.24%), Carnivora (n = 110, 13.99%), Primates (n = 90, 11.45%), Rodentia (n = 79, 9.91%), and Galliformes (n = 60, 7.53%). The top five species changed to pigs (n = 100, 12.55%), cows (n = 73, 9.16%), sheep (n = 67, 8.41%), chickens (n = 48, 6.02%) and rats (n = 48, 6.02%).

3.3. Assumed or Explored?

Of the 2,562 articles we reviewed, 2,546 of them referred to a keyword as assumed; an accepted trait or emotion already deemed to be present in the animal subjects. A further 16 of the articles explored whether or not the animals experienced the trait or emotion. There were significantly more articles assuming the keywords (n = 2,546) than there were studies exploring their existence (n = 16) ( X 2 = 2,497.4, df = 1, P < 0.001). Looking more closely we found that the vertebrate bias was apparent in both the explored and assumed studies. Out of the 16 explored articles only two were studying invertebrates and only 29 of the 2534 assumed articles looked at invertebrates.

3.4. Keywords

We found that 74% of the articles arose from just five keywords. These were fear (n = 636, 22.68%), stress, (n = 607, 21.65%), pain (n = 305, 10.88%), anxiety (n = 267, 9.52%), and depression (n = 222, 7.92%). These words also posed data collection difficulties. Each of these keywords returned between 1,409 to 2,026 results from the initial Science Direct search and unfortunately Science Direct only allows you to view the first 1,000 returned articles. These searches were therefore clipped at 1,000 articles, compared with the other keywords that returned less than 1,000 articles. Had the data been collected from the full list of returned articles these keywords would still remain the top five. It is expected however, that there would have been a higher number of returned articles for each of these keywords, and they would not necessarily remain in the same order. When we removed the human benefit studies from the analysis we found that the top keywords differed. The top five keywords changed to stress (n = 223, 27.98%), fear (n = 142, 17.82%), aggressiveness (n = 139, 17.44%), play (n = 60, 7.53%), and distress (n = 42, 5.27%). It is possible that the sampling issue may have also affected these figures.

Some of the keywords with returned results were assumed in a range of species and orders. For example, the keyword ‘aggressiveness’, which referred to the emotional state, rather than simply aggressive behaviour, was assumed in 34 out of 57 orders. Seven of these were invertebrate orders, which meant that ‘aggressiveness’ was assumed in 63.64% of the invertebrate orders recorded in the review. The keyword ‘stress’, which referred to emotional stress, was assumed for 31 different orders, 29 of which were vertebrates and two were invertebrates. ‘Fear’ was an assumed emotion for 17 of the orders, one invertebrate and 16 vertebrates. None of the keywords were both explored and assumed for the same species or order, within a two year period of publication.

3.5. Positive or Negative?

There appears to be a greater tendency for studies to assume the existence of negative states in animals than positive ones. Out of the 2,546 ‘assumed’ articles, only 154 of them referred to positive states or experiences in animals, compared to 2,359 articles which referred to negative keywords. The remaining 31 articles were classed as neutral and discussed keywords that had no valence, such as theory of mind or consciousness. In the ‘exploring’ studies we found the opposite to be the case, with 11 out of 16 articles looking at positive keywords, compared to just five articles looking at negative ones, however the sample size was too small for any analysis. When we removed the human benefit articles from both the explored and assumed studies we found the negative bias was still present. There were only 149 articles referring to positive states that were performed for animal welfare or behaviour reasons, compared to 625 articles referring to negative states. Furthermore, studies looking at positive emotions and keywords were more likely to be performed to develop knowledge of animal behaviour (n = 99, 29.29%), compared to animal welfare (n = 49, 11.32%), or human benefit reasons (n = 15, 0.85%).

3.6. Where?

We noted 10 different types of experimental or observational settings in the review. From these, laboratories were used the most (n = 2,018, 78.92%), followed by farms (n = 323, 12.63%), the wild (n = 109, 4.26%), zoos (n = 43, 1.68%), and pet households (n = 33, 1.29%). The remaining five categories ranged in number from one to 20 articles and comprised of stables, circuses, shelters, sanctuaries, and stray animals (domestic). Laboratories were clearly used the most, but both laboratories and farms were recorded significantly more than the other eight categories ( X 2 = 2,497.4, df = 1; P < 0.001). When we removed the human benefit studies we found similar results, although laboratories were less likely to be used for these studies (farm: n = 320, 40.40%, laboratory: n = 257, 32.45%, wild: n = 13.13%, zoo: n = 40, 5.05%, and pets: n = 28, 3.54%). When we looked at what type of keywords were being studied, we found that pet and zoo animals were more likely to be studied for positive keywords (pets: n = 12, 35.29%, zoo: n = 15, 34.88%) than laboratory (n = 73, 3.64%), farm (n = 41, 13.36%), or wild animals (n = 20, 18.02%).

The number of published articles discussing the sentience-related keywords has increased over the past two decades ( Figure 3 ). We compared the number of articles published in 1990 and 2011 and found there were significantly more articles published in 2011 than in 1990 ( X 2 = 166.88, df = 1, P < 0.001). This represented a 693.54% increase in articles published in 2011 compared to 1990. In comparison, there was a 249.25% increase in the number of articles published in Science Direct and Ingenta Connect in 2011 compared to 1990, with the word ‘animal’ in the abstract, title or keywords. The increase in publications is also consistent for both the positive ( Figure 4 ) and negative articles ( Figure 5 ). There were significantly more articles published in the year 2011 compared to 1990, for both the positive ( X 2 = 15.7, df = 1, P < 0.001) and negative studies ( X 2 = 141.788, df = 1, P < 0.001). Studies being performed for each of the three ‘why’ categories also significantly increased from 1990 to 2011 (animal behaviour; X 2 = 33.62, df = 1; P < 0.001; animal welfare; X 2 = 30.19, df = 1, P < 0.001; and human benefit; X 2 = 104.26, df = 1, P < 0.001).

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The total number of reviewed articles published from 1990 to 2011. The 242 articles published in 2012 were not included in this analysis as the data collection period did not account for the entire year.

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The number of reviewed articles containing positive keywords, published between 1990 and 2011.

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The number of reviewed articles containing negative keywords, published between 1990 and 2011.

4. Discussion

Animal sentience is often thought of as a complex, poorly defined, subjective, and abstract concept, raising as many questions as it does answers. If you talk to different people about their views on animal sentience you will undoubtedly get various responses. If you asked a pet owner whether animals have feelings, they would regale you with stories of when their dog comforts them, or is proud of himself when he opens a closed door. A farmer who works closely with his or her animals may tell you about how cow number 19 likes being tickled behind her ears, and how cow number 25 is the shy and cautious one. But then you may talk to someone who sees animals only in terms of their monetary value. Animals to such people are not living, sentient beings, they represent commodities. It is far easier for them to see animals in this way but far less convenient for them to consider their ability to suffer or their need to experience positive emotions like pleasure. How people perceive animals is never black and white, attitudes may depend on the species in question, and the animals perceived mental ability [ 47 ]. However, when you consider how we treat the animals we farm for food, experiment upon, or use for entertainment, you can clearly see how important these perspectives are, and which the dominant ones are.

Although our review recorded the use of a wide range of species and keywords, the majority of articles referred to the top five keywords; fear, stress, pain, anxiety, and depression, were performed for human benefit, and used rats and mice. Such a result is unsurprising when you consider the dependence of research upon rodents, and that their ability to feel and experience emotions is often both beneficial and essential to animal model research. This is interesting however, when you consider that much of the criticism around animal sentience science is concerned with the inability to measure subjective states [ 6 , 7 , 8 ]. In the 1,765 studies performed for human benefit, the subjective states of animals were not only measured but were often fundamental to the research being performed. The primary aim of these studies may not have been to measure animal sentience, but the scientists responsible have, perhaps unwittingly, assumed and measured its existence for the purposes of their studies. Given that this type of research is often looking at the development of drugs for human use, the scientific standards for this research should be of a very high standard and subject to extensive scrutiny. It could therefore be safely assumed that their measures of subjective states in animals are not leaps of faith but are instead based upon robust, empirical data. If this is the case, then it would provide strong evidence for the credible and objective nature of animal sentience research, and offer powerful rebuttals to criticisms which maintain the opposite to be true.

We can see from the results that industry and human medical progress are major factors influencing which species are studied. The pharmaceutical industry relies heavily upon rodents to act as animal models for human disorders, such as depression and anxiety [ 48 , 49 ]. Moreover, the billions of animals used in agriculture every year further outweighs the recorded numbers of research animals [ 36 , 50 ]. It is therefore unsurprising that rats, mice, pigs, cows, sheep and chickens were the top species used in the studies reviewed. However, very few of the studies looked at fish. Fish are increasingly being farmed and billions are wild-caught every year [ 51 ], and they are increasingly being used in experimentation [ 52 ]. As a result, we would have expected fish to feature more frequently than the 45 times we recorded in the review. The shortage of research on fish may be a result of the lack of consensus around fish sentience. Despite fish often being protected in legislation and in research regulations, some still argue that they are incapable of feeling pain [ 6 , 8 ]. In recent years, several studies have suggested that fish do have the capacity to feel pain, despite claims that their neurology renders them incapable of such experience [ 8 , 53 , 54 , 55 ]. We hope therefore, that future reviews will feature fish more frequently as a result of the growing understanding of their subjective states.

Invertebrates are used and managed on a considerable scale. They are killed during pest control, experimented upon, and both consumed and farmed on an increasing scale every year [ 56 ]. In addition, human reliance on invertebrates is expected to intensify, as they are increasingly being viewed as a viable and sustainable food source for the growing human population [ 57 , 58 ]. Considering the increased impact we have on invertebrates, and the fact that invertebrate species comprise 99% of the world’s animals [ 59 ], we were disappointed to see how little they featured within the scientific literature. The treatment of invertebrates differs greatly to that of vertebrates, due to the difference in attitudes towards these animals, and the lack of understanding about their capacity for subjective feelings [ 28 , 59 ]. In recent years, as a result of increased understanding of the subjective states of invertebrates, several positive developments regarding their protection have come about. For example, the UK’s Animals (Scientific Procedures) Act (1986) was updated in 2012 to include all cephalopods and New Zealand’s Animal Welfare Act (1999) includes both crabs and crayfish. Research into the subjective states of invertebrates must continue to ensure that all sentient invertebrate species are protected.

4.3. Assumed or Explored?

We performed this review to scratch beneath the surface of animal research, to understand what is being explored, and in whom. What we found surprised us; very little is actually being explored. A lot of these traits and emotions are in fact already being accepted and utilised in the scientific literature. Indeed, 99.34% of the studies we recorded assumed these sentience related keywords in a number of species. In comparison, there were only 16 studies exploring the existence of these traits in animals, and these took place across the entire study focal period and were not seen to increase in recent years. The small number of studies exploring the capacity of emotions in animals suggests that such explorative studies are not increasing, as has previously been suggested, e.g., [ 12 , 20 ]. In view of the importance of animal sentience research to the welfare of animals, we hope that we will see an increase in the future, as more scientists continue to explore animal sentience.

4.4. Positive or Negative?

Each of the top five keywords were negative, and there were far fewer articles discussing the positive keywords than the negative ones. Each of the top keywords referred to states in animals that are intrinsic and necessary for fitness and survival, but extended experience of them can be detrimental to their welfare. The large number of studies discussing the negative keywords is still a positive outcome though, as good animal welfare is dependent upon the absence of these. It is however, increasingly being recognised that good animal welfare also requires the promotion of positive states such as pleasure [ 5 , 23 , 24 ]. By looking at each ‘why’ category separately we can present some possible explanations for the overwhelming bias for negative states. The human benefit studies in this review were mostly performing research into human physical and mental health. Human research has the same bias for focusing on negative emotions as animal sentience research does [ 20 ]. It is therefore unsurprising that the majority of animal research performed for human benefit has the same negative bias. For the animal welfare and behavioural studies the lack of discussion and exploration of positive emotions is a greater concern and we had hoped for a more balanced focus. The bias in these sectors may be reflective of the historical focus on negative states and the relative recent shift in attention towards the promotion of positive states in animals. The discussion of positive keywords did increase over the 21 year focal period, and comparisons between 1990 and 2011 showed a significant increase. This is an encouraging result and shows that reference to positive states is increasing. However, Figure 2 shows that it has not been a steady or consistent increase. These results were disappointing but not unanticipated, as the bias towards negative states in animals has been discussed before [ 4 , 20 , 23 , 60 ]. There are associated difficulties with measuring positive states in animals, which may give further indication as to why the focus is so biased towards negative states. For example, emotions such as fear and pain are often far more intensely communicated and expressed than positive emotions, making them easier to identify [ 22 , 53 , 54 ]. This in turn often creates a sense of importance and urgency to the issue. Fortunately there is success from those scientists seeking to tackle these issues, and new approaches for measuring positive emotions are appearing, e.g., [ 16 , 61 , 62 , 63 ].

Animal welfare science needs to move away from the bias towards negative states. Although addressing negative states is a fundamental step in addressing animal welfare, failing to recognise the importance of positive experiences and emotions can have detrimental effects on both the science of animal welfare and the well-being of the animals we use. When we focus on negative states we are only addressing half of the problem. Animals have an interest in positive experiences in the same way humans do, and so positive experiences and emotions warrant much more consideration than they currently receive [ 4 ]. This one-sided approach to welfare over-simplifies the motivations and needs of animals [ 5 ] and fails to recognise some of the benefits that positive emotions may have on the animals’ mental and physical health. For example, in humans it is thought that humour and laughter may benefit health, and humour is increasingly being incorporated into human medical care [ 20 , 64 ]. Furthermore, a more holistic knowledge of animals’ emotional state may be helpful in predicting the responses of animals to certain situations [ 20 ]. Knowledge such as this would have significant practical applications to many situations where animal welfare needs to be improved.

4.5. Where?

Of the 10 experimental settings recorded, laboratories were used the most, coming only second to farms once the human benefit studies were removed. This is unsurprising considering the number of human benefit studies performed where laboratory settings are the standard. The animal behaviour category consisted of 106 laboratory studies out of a possible 350. This was surprising given that these studies were performed primarily to further ethological knowledge. We do not wish to criticise such research or question its value, after all, knowledge of animal behaviour is integral to understanding animal sentience. We also acknowledge that laboratories offer the standardised settings that are sometimes required for such studies. We would however, like to highlight that there are also a number of other suitable settings where research can be performed. Moreover, on many occasions these can provide a truer representation of the species behaviour than an artificial laboratory setting can. Breeding animals for a laboratory existence should always be seriously considered given the welfare implications of laboratory research and housing. Wherever possible, existing populations such as farm, wild, zoo, or pet animals should be utilised to avoid the unnecessary over-breeding and discomfort laboratories often inflict [ 65 ].

Overall, the number of published articles reviewed had increased from 1990 to 2011 ( Figure 1 ). When we compared the percentage increase of the reviewed papers to that of the total number of papers published, we found that the increase was far greater for the studies reviewed (693.54% vs. 249.25%). This suggests that the observed increase in papers referring to the keywords can be attributed to a specific increase in the use of these sentience related keywords and not attributable to a general increase in publication. This is a positive result, and we hope that as acknowledgment of animal sentience increases, this will in turn have a positive impact on how we view and treat animals.

4.7. Limitations and Future Research

Our results have provided a beneficial and original insight into the issue, but because we only looked at two journal databases they are not inclusive of the entire body of scientific literature. Future work would therefore benefit from incorporating other databases and also the non-English literature, which was excluded in this study. Our results provide information about articles published in 1990 through to mid-2012, and we would like to continue to review future research on a bi-annual basis.

Due to time and budgetary constraints we were only able to review the abstracts of the articles and not the full papers. Although we could identify the information we required for most of the time via this method, there were a few instances when we could not determine which species were being used. Furthermore, the keyword searches performed in the journal databases only searched the abstract, title and keywords. As a result, some articles which only used the keyword or the word ‘animal’ in the main text and not the abstract, title or keywords, would have been excluded from the review. Sample searches performed in the pilot phase of the study showed minimal differences in the number of valid articles returned from this method, vs. searches performed using the entire article. Future work could look at analysing the entire papers to confirm these sample findings. In addition, by only looking at the abstracts we were unable to evaluate whether or not the study’s methodology caused any pain or discomfort to the animals used. Should future research be performed that looks at the entire article, the inclusion of such criteria would make an interesting addition. One other limitation was the inability to view more than 1,000 abstracts from Science Direct for the five searches that returned more than 1,000 results. As these words were still the top five keywords it appears that this limitation had little impact, other than potentially affecting the order and number of returned results for these keywords.

4.8. Emotions Count

Knowledge of whether animals can experience emotions or possess certain traits seen in humans, gives further weight to their value as sentient, emotional beings. We humans continuously seek to compare animals against our own abilities, whether it is by training chimps to use sign-language or making animals do arithmetic. This anthropocentric view is often why we dismiss animal emotions , as we do not recognise their emotional experiences or we consider them to significantly differ from ours and be of less importance. The list of 174 keywords used in this review was not meant to represent a catalogue of sentience indicators. It was however, developed to capitalise upon humans’ anthropocentric nature and accommodate the innate tendency of humans to evaluate and measure animals against our human values. Each of the words included in the list has meaning and value either in terms of human sentience and emotions, or in regards to existing work in the animal sentience field. We hope, therefore, that by using these as a benchmark for measuring the prevalence of sentience and related concepts, we have garnered a greater insight into what is considered important by scientists performing animal research. This in turn provides a powerful tool for animal advocates, advisors and animal welfare scientists, helping us to improve the well-being of the animals in our care.

Animal sentience is often thought to be an abstract concept, something without real definition or tangible indicators. We hope that this review has gone some way towards dispelling some of these misconceptions by approaching the matter in a new way. Animal sentience forms the foundation of animal welfare science and it is why animals need protection. The results clearly show there are fundamental areas which are not yet being considered. Future research must continue to fill these gaps, particularly for those taxa that we use so much yet know so little about. We have shown how little is known about the experience and promotion of positive emotions in animals, and this is an area of utmost importance to the field of animal welfare. By ignoring positive emotions we are ignoring a valuable part of what it means to be alive. With so much to learn about the subjective minds of animals and the challenges this brings, the future of animal sentience science is certainly an exciting one.

Acknowledgements

We would like to thank the World Society for the Protection of Animals for their support of this research project. We would like to thank Mark Kennedy and Maisie Tomlinson for their helpful comments, Jessica Wilkinson for her assistance during the data collection, Marc Bekoff for peer reviewing the key words and the anonymous reviewers for their helpful and insightful comments.

The keywords used in the study, including details of their source, valence and whether they returned suitable results.

Conflicts of interest

The authors declare no conflict of interest.

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