Do animals feel emotions, or display empathy?

Since we can't know much of what's going on inside an animal's brain, can there still be a consensus among zoologists on whether animals feel emotions, display empathy, care, etc.? Or is it disputed? To clarify what I said since commenters asked for clarifications: Of course I don't mean SENSES (touch/vision/taste… ) and also I do NOT include humans.

A suggested definition of emotion:

An emotion is a feeling such as happiness, love, fear, anger, or hatred… the part of a person's character that consists of their feelings, as opposed to their thoughts. (Collins Dictionary)

An example of emotion:

No one doubts that dogs know when they are in trouble,” writes de Waal, “but whether they actually feel guilty is a point of debate.” According to a study by Alexandra Horowitz, the canine guilty look-“lowered gaze, ears pressed back, slumped body, averted head, tail rapidly beating between the legs-is… not about what they have done but about how their owner reacts. If the owner scolds them, they act extremely guilty. If the owner doesn't, everything is fine and dandy.” (Greater Good Magazine, University of California, Berkeley)


Empathy is the ability to share another person's feelings and emotions as if they were your own. (Collins Dictionary)

Example of empathy:

Macaques refused to pull a chain that delivered food to themselves if doing so also caused a companion to receive an electric shock.[48][49] This inhibition of hurting another conspecific was more pronounced between familiar than unfamiliar macaques, a finding similar to that of empathy in humans. (Wikipedia: Emotions in animals)

The answer to the question "Do scientists agree whether animals feel emotions, or display empathy?" is No. However, there are some animals that certainly do not have emotions or empathy (for example animals that have no substantial nervous system, like sponges), and some animals that certainly do have emotions. The only disagreement is in the middle and where to draw the boundaries. In reality, drawing such boundaries is probably not all that productive, coming down to semantic rather than biological argument.

As a citation for the disagreement, see the two answers and the comments here already.

The difficulty in assessing a statement like this is largely one of operational definition: you need to define what an emotion is, and depending on that definition you will get entirely different answers. You will also get different answers depending on your philosophical approach to questions like these and what the essence of experience is. I would also point out that emotion and empathy are vastly different levels of cognitive ability according to typical definitions of the two.

However, I will say that there are a lot of recent findings of higher cognitive abilities in non-human animals than previously thought. Rodents have long been used as generic models of mammalian nervous systems, including emotional processes. Mirror neurons in non-human primates respond to the behaviors and movements of conspecifics, which is an important aspect of both social learning and potentially empathy. Self-recognition is an important aspect of theory of mind, and has been observed in several species of different types. Mourning behaviors have been observed in various animals in response to death of conspecifics. There can be some disagreement of the interpretation of these findings, but I'll include some references here so readers can do some of their own research and find if they are convinced.

Note that these works study particular animal abilities in particular contexts, rather than making blanket judgments about "all animals" or anything like that. This is typically how science works, incrementally based on the results of individual experiments.


Gallese, V., & Goldman, A. (1998). Mirror neurons and the simulation theory of mind-reading. Trends in cognitive sciences, 2(12), 493-501.

Lane, R. D., & Nadel, L. (Eds.). (1999). Cognitive neuroscience of emotion. Oxford University Press.

Mulcahy, N. J., & Call, J. (2006). Apes save tools for future use. Science, 312(5776), 1038-1040.

Plotnik, J. M., De Waal, F. B., & Reiss, D. (2006). Self-recognition in an Asian elephant. Proceedings of the National Academy of Sciences, 103(45), 17053-17057.

Prior, H., Schwarz, A., & Güntürkün, O. (2008). Mirror-induced behavior in the magpie (Pica pica): evidence of self-recognition. PLoS biology, 6(8), e202.

Rajala, A. Z., Reininger, K. R., Lancaster, K. M., & Populin, L. C. (2010). Rhesus monkeys (Macaca mulatta) do recognize themselves in the mirror: implications for the evolution of self-recognition. PloS one, 5(9), e12865.

Roth, G., & Dicke, U. (2005). Evolution of the brain and intelligence. Trends in cognitive sciences, 9(5), 250-257.

Some animals definitely feel emotions, emotion are the basic workings of the brain, variations in neurotransmitters used to tag experiences and alter behavior. If it has a brain and is capable of learning it probably feels some emotions, even if only pleasure or discomfort because that is how learning affects ("codes") an experience for association.

Not every animal is going feel them all or feel them at all to the same degree. "Animal" covers a very wide group, a sponge is an animal and almost certainly feels no emotions as it lacks a brain or substantial nervous system, on the other hand humans are the group we use to define emotions are are also animals. But exactly which emotion each animal experiences is impossible to say at the moment, we can't even say for certain every human experiences every emotion. And currently discreet definitions for emotions elude us.

As for specific emotions, as far as we can tell every basic emotion humans experience is represented somewhere in the animal kingdom, and most the basic human emotions may be at least common to mammalia. It is extremely unlikely emotions arrose ex nihilo in humans given the similarities in behavior between humans and other animals, especially other primates. Bonobo's for instance even show similar emotional managements skills to humans.

A consensus is a tricky term, emotion in animals has been a taboo subject for a long time, and there are still pockets that assume animals experience no emotions and operate on pure logic like a robot, but this view has never withstood scrutiny. I tired to present the closest thing we have ot a consensus.

Mammalian associative learning has been the most well studied for obvious reasons, but there is no reason to believe it is unique given similarities in behavior with other groups.Yes the brain works like a different engine but emotions is how our difference engine operates, how it codes experiences.

Excerpt: The Emotional Lives of Animals

Many animals display their feelings openly, publicly, for anyone to see. And when we pay attention, what we see outside tells us lots about what’s happening inside an individual’s head and heart. As we’ll find, careful scientific research is validating what we intuitively understand: that animals feel, and their emotions areas important to them as ours are to us.

A few years ago my friend Rod and I were riding our bicycles around Boulder, Colorado, when we witnessed a very interesting encounter among five magpies. Magpies are corvids, a very intelligent family of birds. One magpie had obviously been hit by a car and was lying dead on the side of the road. The four other magpies were standing around him. One approached the corpse, gently pecked at it – just as an elephant noses the carcass of another elephant – and stepped back. Another magpie did the same thing. Next, one of the magpies flew off, brought back some grass, and laid it by the corpse. Another magpie did the same. Then, all four magpies stood vigil for a few seconds and one by one flew off.

Were these birds thinking about what they were doing? Were they showing magpie respect for their friend? Of were they merely acting as if they cared? Were they just animal automatons? I feel comfortable answering these questions, in order: yes, yes, no, no. Rod was astounded by how deliberate the birds were. He asked me if this was normal magpie behavior, and I told him that I’d never seen anything like this before and hadn’t read any accounts of grieving magpies. We can’t know what they were actually thinking of feeling, but reading their actions there’s no reason not to believe these birds were saying a magpie farewell to their friend.

Despite the more than three decades I’ve spent studying animal species, I never cease learning from the individuals I encounter. Red foxes live near my mountain home outside of Boulder, Colorado. As I stare into the eyes of a red fox sitting by my study and watching me type, or as I observe red fox pups playing with one another or a female red fox burying her mate, I can’t help but reflect deeply on what it’s like to be these individuals sharing my hillside. Many animals live on the surrounding land – coyotes, mountain lions, porcupines, raccoons, black bears, a wide variety of birds, and lizards, along with many dogs and cats. Through the years, they’ve been my friends and teachers.

In my musings about animal emotions I also can’t help wondering, What about the insects? Do even mosquitoes have emotional lives? Of course, mosquitoes have tiny brains and lack the neural apparatus necessary for the evolution of emotions, so it’s doubtful they do. But in truth, we just don’t know. One day, perhaps we’ll figure out a way to determine this. More important, however, would it make a difference to us if they did? It should, just as it should make a difference to us that other animals have emotions. Knowing that animals feel – and being able to understand them when they express joy, grief, jealousy, and anger – allows us to connect with them and also to consider their points of view when we interact with them. Knowledge about animal passions should make a difference in how we view, represent, and treat our fellow beings.


Babyl the Elephant and Her Unconditional Friends

A recent trip to Kenya and Tanzania opened my eyes to the world of elephants, who are some of the most amazing beings I’ve ever seen. Observing large groups of wild elephants close up I could feel their majestic presence, awareness, and emotions. These firsthand experiences were wholly different than seeing captive elephants, who often live alone, in the confines and unnatural settings of a zoo, and my visit was deeply spiritual, inspirational, and transformative.

While we were watching a group of wild elephants living in the Samburu Reserve in Northern Kenya, we noted that one of them, Babyl, walked very slowly. We learned that she was crippled and that she couldn’t travel as fast as the rest of the herd. However, we saw that the elephants in Babyl’s group didn’t leave her behind they waited for her. When I asked our guide, the elephant expert Iain Douglas-Hamilton, about this, he said that these elephants always waited for Babyl, and they’d been doing so for years. They would walk for a while, then stop and look around to see where Babyl was. Depending on how she was doing, they’d either wait or proceed. Iain said the matriarch even red her on occasion.

Why did the other elephants in the herd act this way? Babyl could do little for them, so there seemed no reason for or practical gain in helping her. The only obvious conclusion we could draw was that the other elephants cared for Babyl, and so they adjusted their behavior to allow her to remain with the group.

Friendship and empathy go a long way. And Babyl’s friends aren’t an isolated example. In October 2006 in a small village in eastern India, a group of fourteen elephants crashed through a village looking for a group member who had fallen into a ditch and drowned. Residents had already buried the seventeen-year-old female elephant, but still, thousands of people were forced to flee their homes as the other elephants searched and rampaged for more than three days.

In September 2006 there was a meeting about animal welfare called “The Heart of the Matter.” It’s nice to see scientists finally using the word heart, for the heart is the matter.

I study animal emotions and I love what I do. Over the course of my career, I’ve studied a wide variety of animals – coyotes, wolves, dogs, Adélie penguins, archer fish, western evening grosbeaks, and Steller’s jays – and I’ve tackled a wide range of questions, dealing with everything from social behavior, social organization, and social development to communication, play, antipredatory behavior, aggression, parental behavior, and morality. To me, the evidence for animal emotions is impossible to deny, and it is widely supported by our current knowledge in animal behavior, neurobiology, and evolutionary biology.

In fact, the study of animal emotions is a dynamic and rapidly developing field of science, and there’s no shortage of interest in animal emotions among scientists and average folks alike. In March 2005 about six hundred people from more than fifty nations gathered in London at a landmark meeting sponsored by the Compassion in World Farming Trust to learn more about animal sentience, animal consciousness, and the emotional lives of animals. In October 2006 the World Society for the Protection of Animals organized a conference in Rio de Janeiro to discuss how to improve animal welfare on farms and in research labs. Organizers expected about two hundred people, but twice that many attended, coming predominantly from Brazil and surrounding countries. The favorable response to the meetings in London and Rio is indicative that the time really has arrived for us to come to terms with the emotional lives of animals.

Stories about animal emotions and our complicated interrelationships with animals appear with increasing frequency in the press, from prestigious scientific journals like Science, Nature, Trends in Ecology and Evolution, and the Proceedings of the National Academy of Sciences to the New York Times, Psychology Today, Scientific American, Time, The Economist, and even Reader’s Digest. The emotional lives of animals was even the subject of a surprise hit movie, The March of the Penguins. Released in summer 2005, the documentary poignantly depicts penguin feelings and demonstrates how they experience suffering but also how they endure the most extreme challenges as they care for their eggs and their young.

Nevertheless, despite mounting scientific evidence and widespread popular belief, a decreasing few within the scientific community remain skeptical. Some still doubt that animal emotions even exist, and many who believe they do exist tend to think animal emotions must be lesser than human ones. This seems to me an outdated and even irresponsible point of view, and my main goal in this chapter – and indeed throughout the book – is to show that animal emotions exist, that they are important to humans, and that this knowledge should influence how we treat our fellow animals.

In discussing animal emotions, I focus mainly on behavioral data and anecdotal stories, weaving in recent discoveries in social neuroscience to show how a combination of common sense and scientific data – what I call “science sense” – makes a strong case for the existence of beastly passions. While stories drive much of my discussion, I bring in scientific studies as necessary for support.

However, once we agree that animal emotions exist and that they matter – which is what a great many people already believe – then what? Then we must consider ethics. We must look to our actions and see if they are consistent with our knowledge and beliefs. I feel strongly that ethics should always inform science. We should always strive to merge knowledge, action, and compassion. Indeed, that is always the heart of the matter.

It is very difficult to answer the question, “What are emotions?” Most of us know emotions when we see them but find it difficult to define them. Are they physical, mental, or both? As a scientist, I feel safe saying that emotions are psychological phenomena that help in behavioral management and control they are phenomena that emote us, that make us move. A distinction is often made between “emotional responses” to physical reactions and “feelings” that arise from thoughts. Emotional responses show that the body is responding to certain external stimuli. For example, we see an oncoming car about to hit us and we feel fear – increasing our heart rate, blood pressure, and body temperature. But actually, the fear isn’t felt until the brain responds to the physiological changes that were a reaction to seeing the oncoming car.

Feelings, on the other hand, are psychological phenomena, events that happen solely in an individual’s brain. An external event may trigger one emotion, such as anger or grief, but upon reflection we may decide we feel differently. We may interpret our emotions. Feelings express themselves as different moods. Feelings help us and influence how we interact with others in a wide variety of different social situations.

Charles Darwin, the first scientist to study animal emotions systematically, recognized six universal emotions: anger, happiness, sadness, disgust, fear, and surprise. He maintained these core emotions help us deal rapidly with a wide variety of circumstances and help us to get along in a complex social world. Others have since added to his list. Stuart Walton, in his book A Natural History of Human Emotions, adds jealousy, contempt, shame, and embarrassment to Darwin’s core group, while the neuroscientist Antonio Damasio (in Descartes’ Error) says that social emotions also include sympathy, guilt, pride, envy, admiration, and indignation. It’s interesting that none of these researchers mention love.

Which, if any, of these emotions do animals experience? And do animals experience any emotions that humans do not? This is a very interesting question. Ethologist Joyce poole, who has studied elephants for many years, states: “While I feel confident that elephants feel some emotions that we do not, and vice versa, I also believe that we experience many emotions in common.”

If Poole is right, then there may be some emotions that animals experience that humans will never understand, but there are many that we do. Aren’t animals, human and nonhuman alike, happy when playing of when reuniting with a loved one? Don’t animals become sad after losing a close friend? When wolves reunite, wagging their tails loosely to and fro in a circle, whining and jumping about, are they not displaying happiness? What about elephants who reunite in a greeting celebration, flapping their ears and spinning about and emitting a vocalization known as a “greeting rumble” – is this not happiness? Likewise, what name but grief can we give to the emotion that animals display when they remove themselves from their social group, sulk after the death of a friend, stop eating, and even die? Surely, despite differences, all species must share a similar core of emotions.


Researchers usually recognize two different types of emotions, primary and secondary emotions. Primary emotions are considered to be basic inborn emotions. These include generalized rapid, reflex-like (“automatic,” or hardwired) fear and fight-or-flight responses to stimuli that represent danger. They require no conscious thought and include Darwin’s six universal emotions: fear, anger, disgust, surprise, sadness, and happiness. Animals can perform a primary fear response, such as avoiding an object, almost unconsciously, before they have even recognized the object generating the reaction. Loud raucous sounds, certain odors, objects flying overhead: these and other such stimuli ate often inborn signals for “danger” that cause an automatic avoidance reaction. There’s little or no room for error when confronted with a dangerous stimulus, so natural selection has resulted in innate reactions that are crucial to individual survival.

Primary emotions are wired into the brain’s evolutionarily old limbic system (especially the amygdala) this is the “emotional” part of the brain (so named by Paul MacLean in 1952). The physical structures in the limbic system and similar emotional circuits are shared among many different species and provide a neural substrate for primary emotions. In his three-brains-in-one (or triune brain) theory, MacLean identifies the reptilian, or primitive, brain (possessed by fish, amphibians, reptiles, birds, and mammals) the limbic, or paleomammalian, brain (possessed by all mammals) and the neocortical, or “rational” neomammalian, brain (possessed by a few mammals, such as primates and humans) – all packaged into the cranium. Each is connected to the other two, but each also has its own capacities. While the limbic system seems to be the main area of the brain in which many emotions reside, current research now indicates that not all emotions are necessarily packaged into a single system, and there may be more than one emotional system in the brain.

Secondary emotions are more complex emotions, and they involve higher brain centers in the cerebral cortex. They could involve core emotions of fear and anger, or they could be more nuanced, involving such things as regret, longing, or jealousy. Secondary emotions are not automatic: they are processed in the brain, and the individual thinks about them and considers what to do about them – what action is the best one to perform in a certain situation. Conscious thought and secondary emotions can influence how we respond to situations that bring forth primary emotions: We may duck as an unseen object flies overhead, but as we recognize that it’s only a shadow, we will refrain from running and instead, feeling a twinge of embarrassment, quickly straighten up and pretend nothing is wrong.

Thinking about the emotion allows for flexibility of response in changing situations after evaluating which of a variety of actions would be the most appropriate to perform in the specific situation. Sometimes, if someone is bothering you, it might be appropriate to get away from them, and sometimes this might create an even worse social situation – depending on who the person is and what kind of consequences you fear. Although most emotional responses are unconsciously generated – they occur without thinking – we learn to try to think before acting. Thinking allows us to make connections between feelings and actions, and this allows for variability and flexibility in our behavior so that, depending on the social situation, we always do the right thing. In this way, evidence of emotions in any creature is also an important step in determining sentience and self-awareness.
From The Emotional Lives of Animals:A Leading Scientist Explores Animal Joy, Sorrow, and Empathy – and Why They Matter by Marc Bekoff. Excerpted by permission of Marc Bekoff.

Animals Don't Experience Emotions, Claims Texas Journalist

My email inbox was brimming over today with people asking me to respond to the most absurd claim I've heard in a long time. In an essay that somehow found it's way into the New York Times, a newspaper that has published numerous essays about the detailed scientific research that has been performed that shows that many nonhuman animals (animals) experience rich and deep emotional lives, Kate Murphy, a Texas journalist, claims "Eating is an orthodoxy you can practice three times a day. And because there is no definitive scientific evidence that animals experience emotions as we do, nor is there irrefutable proof that eating any particular diet is healthier than another, we are all free to use our individual feelings, desires and experiences to shape our eating ethos without fear that our beliefs can be effectively challenged. Just like you can't prove there is or isn't a God." Ms. Murphy's essay is titled "Blessed Be My Freshly Slaughtered Dinner," in which she tries to argue that it's perfectly fine to kill animals for food.

The New York Times should not have published this essay, not because Ms. Murphy favors meat diets and supports what has become known as "the eat-what-you-kill movement," but because she conveniently and extraordinarily ignores reams and reams of detailed data that show clearly that many nonhumans experience a wide variety of emotions "as we do." I often write about this research and you can find much more here and here. Ms. Murphy does attempt to appeal to authority by noting that Facebook's founder, Mark Zuckerberg, once said, "he would eat only meat he killed himself as part of a reported yearlong 'personal challenge.'" Who really cares about Mr. Zuckerberg's meal plans?

Animal welfare is based on what animals feel

Ms. Murphy's claim about the absence of data on the emotional lives of animals is absurd, incredibly uninformed, and thoroughly misleading. For example, we now know that rats, mice, and chickens, in addition to many other animals, display empathy, and that "food animals" also experience deep and rich emotions (see also). Indeed, animal welfarists across the board claim that the concern for the welfare of other animals is based on animal feelings and, for example, iconic world renowned animal welfarist Dr. Temple Grandin does what she does to reduce the pain and suffering of "food animals" -- to make their lives "relatively humane" -- based on what we know about the emotional lives of these sentient beings (see also an interview published in the New York Times called "Temple Grandin on Autism, Death, Celibacy and Cows" in which she talks about animal feelings). A thorough and current review of this literature can be found in the recently published fifth edition of Dr. Donald Broom and Dr. Andrew Fraser's book called Domestic Animal Behaviour and Welfare. The extremely significant Cambridge Declaration on Consciousness also recognized these scientific facts, as do a vast majority of scientists. The list goes on and on and the wealth of available scientific evidence about the emotional lives of animals makes utterly inane the claim that "there is no definitive scientific evidence that animals experience emotions as we do."

It's clear that researchers themselves know that other animals are emotional beings, and that is why there are numerous studies that attempt to enrich the lives of animals who are used by humans to reduce pain and suffering, and why there is legislation to protect other animals from pain and suffering. Some excellent examples, and there are many more, include a research paper titled "Assessment of positive emotions in animals to improve their welfare" and also essays that detail how a cow's ears tell us how they're feeling, how a cow's nose can reveal inner emotions, and how pigs display empathy. Donald Broom and Andrew Fraser's book mentioned above is a goldmine of current information on the emotional lives of "food" and other animals.

If one chooses to eat other animals even in the "eat-what-you-kill" movement, she or he must know they are killing sentient beings who care about what happens to themselves, families, and friends. And, the question is not "What are you eating," but rather "Whom are you eating." Animals are not disposable objects, but rather thinking and feeling beings, and those who choose to eat them can't conveniently ignore what we know about the cognitive and emotional lives of these individuals.

How this essay escaped the careful eyes of the editors at the New York Times baffles many others and me. If Ms. Murphy and others want to justify eating other animals they ought to come straight about what we know about the emotional lives of other animals from detailed comparative scientific research, and not make thoroughly vacuous claims to support their meal plans. As noted by Dr. Betty Moss, Ms. Murphy and others on whom she relies greatly fail on many different accounts by ignoring what we know. For more on the many misleading claims in Ms. Murphy's essay please also see Carol Adams' "How To Write An OpEd For The New York Times Defending Meat Eating And Get It Published."

It's surely not asking too much for those who write about animals to know about the vast amount of scientific research that has been done and is readily available.


The word "emotion" dates back to 1579, when it was adapted from the French word émouvoir, which means "to stir up". However, the earliest precursors of the word likely date back to the very origins of language. [10]

Emotions have been described as discrete and consistent responses to internal or external events which have a particular significance for the organism. Emotions are brief in duration and consist of a coordinated set of responses, which may include physiological, behavioural, and neural mechanisms. [11] Emotions have also been described as the result of evolution because they provided good solutions to ancient and recurring problems that faced ancestors. [12]

Laterality Edit

It has been proposed that negative, withdrawal-associated emotions are processed predominantly by the right hemisphere, whereas the left hemisphere is largely responsible for processing positive, approach-related emotions. This has been called the "laterality-valence hypothesis". [13]

Basic and complex human emotions Edit

In humans, a distinction is sometimes made between "basic" and "complex" emotions. Six emotions have been classified as basic: anger, disgust, fear, happiness, sadness and surprise. [14] Complex emotions would include contempt, jealousy and sympathy. However, this distinction is difficult to maintain, and animals are often said to express even the complex emotions. [15]

Behaviourist approach Edit

Prior to the development of animal sciences such as comparative psychology and ethology, interpretation of animal behaviour tended to favour a minimalistic approach known as behaviourism. This approach refuses to ascribe to an animal a capability beyond the least demanding that would explain a behaviour anything more than this is seen as unwarranted anthropomorphism. The behaviourist argument is, why should humans postulate consciousness and all its near-human implications in animals to explain some behaviour, if mere stimulus-response is a sufficient explanation to produce the same effects?

Some behaviourists, such as John B. Watson, claim that stimulus–response models provide a sufficient explanation for animal behaviours that have been described as emotional, and that all behaviour, no matter how complex, can be reduced to a simple stimulus-response association. [16] Watson described that the purpose of psychology was "to predict, given the stimulus, what reaction will take place or given the reaction, state what the situation or stimulus is that has caused the reaction". [16]

The cautious wording of Dixon exemplifies this viewpoint: [17]

Recent work in the area of ethics and animals suggests that it is philosophically legitimate to ascribe emotions to animals. Furthermore, it is sometimes argued that emotionality is a morally relevant psychological state shared by humans and non-humans. What is missing from the philosophical literature that makes reference to emotions in animals is an attempt to clarify and defend some particular account of the nature of emotion, and the role that emotions play in a characterization of human nature. I argue in this paper that some analyses of emotion are more credible than others. Because this is so, the thesis that humans and nonhumans share emotions may well be a more difficult case to make than has been recognized thus far.

Moussaieff Masson and McCarthy describe a similar view (with which they disagree): [18]

While the study of emotion is a respectable field, those who work in it are usually academic psychologists who confine their studies to human emotions. The standard reference work, The Oxford Companion to Animal Behaviour, advises animal behaviourists that "One is well advised to study the behaviour, rather than attempting to get at any underlying emotion. There is considerable uncertainty and difficulty related to the interpretation and ambiguity of emotion: an animal may make certain movements and sounds, and show certain brain and chemical signals when its body is damaged in a particular way. But does this mean an animal feels—is aware of—pain as we are, or does it merely mean it is programmed to act a certain way with certain stimuli? Similar questions can be asked of any activity an animal (including a human) might undertake, in principle. Many scientists regard all emotion and cognition (in humans and animals) as having a purely mechanistic basis.

Because of the philosophical questions of consciousness and mind that are involved, many scientists have stayed away from examining animal and human emotion, and have instead studied measurable brain functions through neuroscience.

Comparative approach Edit

In 1903, C. Lloyd Morgan published Morgan's Canon, a specialised form of Occam's razor used in ethology, in which he stated: [19] [20]

In no case is an animal activity to be interpreted in terms of higher psychological processes,
if it can be fairly interpreted in terms of processes which stand lower in the scale of psychological evolution and development.

Darwin's approach Edit

Charles Darwin initially planned to include a chapter on emotion in The Descent of Man but as his ideas progressed they expanded into a book, The Expression of the Emotions in Man and Animals. [21] Darwin proposed that emotions are adaptive and serve a communicative and motivational function, and he stated three principles that are useful in understanding emotional expression: First, The Principle of Serviceable Habits takes a Lamarckian stance by suggesting that emotional expressions that are useful will be passed on to the offspring. Second, The Principle of Antithesis suggests that some expressions exist merely because they oppose an expression that is useful. Third, The Principle of the Direct Action of the Excited Nervous System on the Body suggests that emotional expression occurs when nervous energy has passed a threshold and needs to be released. [21]

Darwin saw emotional expression as an outward communication of an inner state, and the form of that expression often carries beyond its original adaptive use. For example, Darwin remarks that humans often present their canine teeth when sneering in rage, and he suggests that this means that a human ancestor probably utilized their teeth in aggressive action. [22] A domestic dog's simple tail wag may be used in subtly different ways to convey many meanings as illustrated in Darwin's The Expression of the Emotions in Man and Animals published in 1872.

"Small dog watching a cat on a table"

"Dog approaching another dog with hostile intentions"

"Dog in a humble and affectionate frame of mind"

Anecdotal approach Edit

Evidence for emotions in animals has been primarily anecdotal, from individuals who interact with pets or captive animals on a regular basis. However, critics of animals having emotions often suggest that anthropomorphism is a motivating factor in the interpretation of the observed behaviours. Much of the debate is caused by the difficulty in defining emotions and the cognitive requirements thought necessary for animals to experience emotions in a similar way to humans. [15] The problem is made more problematic by the difficulties in testing for emotions in animals. What is known about human emotion is almost all related or in relation to human communication.

Scientific approach Edit

In recent years, the scientific community has become increasingly supportive of the idea of emotions in animals. Scientific research has provided insight into similarities of physiological changes between humans and animals when experiencing emotion. [23]

Much support for animal emotion and its expression results from the notion that feeling emotions doesn't require significant cognitive processes, [15] rather, they could be motivated by the processes to act in an adaptive way, as suggested by Darwin. Recent attempts in studying emotions in animals have led to new constructions in experimental and information gathering. Professor Marian Dawkins suggested that emotions could be studied on a functional or a mechanistic basis. Dawkins suggests that merely mechanistic or functional research will provide the answer on its own, but suggests that a mixture of the two would yield the most significant results.

Functional Edit

Functional approaches rely on understanding what roles emotions play in humans and examining that role in animals. A widely used framework for viewing emotions in a functional context is that described by Oatley and Jenkins [24] who see emotions as having three stages: (i) appraisal in which there is a conscious or unconscious evaluation of an event as relevant to a particular goal. An emotion is positive when that goal is advanced and negative when it is impeded (ii) action readiness where the emotion gives priority to one or a few kinds of action and may give urgency to one so that it can interrupt or compete with others and (iii) physiological changes, facial expression and then behavioural action. The structure, however, may be too broad and could be used to include all the animal kingdom as well as some plants. [15]

Mechanistic Edit

The second approach, mechanistic, requires an examination of the mechanisms that drive emotions and search for similarities in animals.

The mechanistic approach is utilized extensively by Paul, Harding and Mendl. Recognizing the difficulty in studying emotion in non-verbal animals, Paul et al. demonstrate possible ways to better examine this. Observing the mechanisms that function in human emotion expression, Paul et al. suggest that concentration on similar mechanisms in animals can provide clear insights into the animal experience. They noted that in humans, cognitive biases vary according to emotional state and suggested this as a possible starting point to examine animal emotion. They propose that researchers may be able to use controlled stimuli which have a particular meaning to trained animals to induce particular emotions in these animals and assess which types of basic emotions animals can experience. [25]

Cognitive bias test Edit

A cognitive bias is a pattern of deviation in judgment, whereby inferences about other animals and situations may be drawn in an illogical fashion. [26] Individuals create their own "subjective social reality" from their perception of the input. [27] It refers to the question "Is the glass half empty or half full?", used as an indicator of optimism or pessimism. To test this in animals, an individual is trained to anticipate that stimulus A, e.g. a 20 Hz tone, precedes a positive event, e.g. highly desired food is delivered when a lever is pressed by the animal. The same individual is trained to anticipate that stimulus B, e.g. a 10 Hz tone, precedes a negative event, e.g. bland food is delivered when the animal presses a lever. The animal is then tested by being played an intermediate stimulus C, e.g. a 15 Hz tone, and observing whether the animal presses the lever associated with the positive or negative reward, thereby indicating whether the animal is in a positive or negative mood. This might be influenced by, for example, the type of housing the animal is kept in. [28]

Using this approach, it has been found that rats which are subjected to either handling or tickling showed different responses to the intermediate stimulus: rats exposed to tickling were more optimistic. [6] The authors stated that they had demonstrated "for the first time a link between the directly measured positive affective state and decision making under uncertainty in an animal model".

Cognitive biases have been shown in a wide range of species including rats, dogs, rhesus macaques, sheep, chicks, starlings and honeybees. [6]

Self-medication with psychoactive drugs Edit

Humans can suffer from a range of emotional or mood disorders such as depression, anxiety, fear and panic. [29] To treat these disorders, scientists have developed a range of psychoactive drugs such as anxiolytics. Many of these drugs are developed and tested by using a range of laboratory species. It is inconsistent to argue that these drugs are effective in treating human emotions whilst denying the experience of these emotions in the laboratory animals on which they have been developed and tested.

Standard laboratory cages prevent mice from performing several natural behaviours for which they are highly motivated. As a consequence, laboratory mice sometimes develop abnormal behaviours indicative of emotional disorders such as depression and anxiety. To improve welfare, these cages are sometimes enriched with items such as nesting material, shelters and running wheels. Sherwin and Ollson [30] tested whether such enrichment influenced the consumption of Midazolam, a drug widely used to treat anxiety in humans. Mice in standard cages, standard cages but with unpredictable husbandry, or enriched cages, were given a choice of drinking either non-drugged water or a solution of the Midazolam. Mice in the standard and unpredictable cages drank a greater proportion of the anxiolytic solution than mice from enriched cages, indicating that mice from the standard and unpredictable laboratory caging may have been experiencing greater anxiety than mice from the enriched cages.

Spindle neurons Edit

Spindle neurons are specialised cells found in three very restricted regions of the human brain – the anterior cingulate cortex, the frontoinsular cortex and the dorsolateral prefrontal cortex. [31] The first two of these areas regulate emotional functions such as empathy, speech, intuition, rapid "gut reactions" and social organization in humans. [32] Spindle neurons are also found in the brains of humpback whales, fin whales, killer whales, sperm whales, [32] [33] bottlenose dolphin, Risso's dolphin, beluga whales, [34] and the African and Asian elephants. [35]

Whales have spindle cells in greater numbers and are maintained for twice as long as humans. [32] The exact function of spindle cells in whale brains is still not understood, but Hof and Van Der Gucht believe that they act as some sort of "high-speed connections that fast-track information to and from other parts of the cortex". [32] They compared them to express trains that bypass unnecessary connections, enabling organisms to instantly process and act on emotional cues during complex social interactions. However, Hof and Van Der Gucht clarify that they do not know the nature of such feelings in these animals and that we cannot just apply what we see in great apes or ourselves to whales. They believe that more work is needed to know whether emotions are the same for humans and whales.

Vocalizations Edit

Though non-human animals cannot provide useful verbal feedback about the experiential and cognitive details of their feelings, various emotional vocalizations of other animals may be indicators of potential affective states. [36] Beginning with Darwin and his research, it has been known that chimpanzees and other great apes perform laugh-like vocalizations, providing scientists with more symbolic self-reports of their emotional experiences. [37]

Research with rats has revealed that under particular conditions, they emit 50-kHz ultrasonic vocalisations (USV) which have been postulated to reflect a positive affective state (emotion) analogous to primitive human joy these calls have been termed "laughter". [38] [39] The 50 kHz USVs in rats are uniquely elevated by hedonic stimuli—such as tickling, rewarding electrical brain stimulation, amphetamine injections, mating, play, and aggression—and are suppressed by aversive stimuli. [6] Of all manipulations that elicit 50 kHz chirps in rats, tickling by humans elicits the highest rate of these calls. [40]

Some vocalizations of domestic cats, such as purring, are well known to be produced in situations of positive valence, such as mother kitten interactions, contacts with familiar partner, or during tactile stimulation with inanimate objects as when rolling and rubbing. Therefore, purring can be generally considered as an indicator of "pleasure" in cats. [41]

Low pitched bleating in sheep has been associated with some positive-valence situations, as they are produced by males as an estrus female is approaching or by lactating mothers while licking and nursing their lambs. [41]

Neurological Edit

Neuroscientific studies based on the instinctual, emotional action tendencies of non-human animals accompanied by the brains neurochemical and electrical changes are deemed to best monitor relative primary process emotional/affective states. [42] Predictions based on the research conducted on animals is what leads analysis of the neural infrastructure relevant in humans. Psycho-neuro-ethological triangulation with both humans and animals allows for further experimentation into animal emotions. Utilizing specific animals that exhibit indicators of emotional states to decode underlying neural systems aids in the discovery of critical brain variables that regulate animal emotional expressions. Comparing the results of the animals converse experiments occur predicting the affective changes that should result in humans. [42] Specific studies where there is an increase or decrease of playfulness or separation distress vocalizations in animals, comparing humans that exhibit the predicted increases or decreases in feelings of joy or sadness, the weight of evidence constructs a concrete neural hypothesis concerning the nature of affect supporting all relevant species. [42]

The argument that animals experience emotions is sometimes rejected due to a lack of higher quality evidence, and those who do not believe in the idea of animal intelligence often argue that anthropomorphism plays a role in individuals' perspectives. Those who reject that animals have the capacity to experience emotion do so mainly by referring to inconsistencies in studies that have endorsed the belief emotions exist. Having no linguistic means to communicate emotion beyond behavioral response interpretation, the difficulty of providing an account of emotion in animals relies heavily on interpretive experimentation, that relies on results from human subjects. [25]

Some people oppose the concept of animal emotions and suggest that emotions are not universal, including in humans. If emotions are not universal, this indicates that there is not a phylogenetic relationship between human and non-human emotion. The relationship drawn by proponents of animal emotion, then, would be merely a suggestion of mechanistic features that promote adaptivity, but lack the complexity of human emotional constructs. Thus, a social life-style may play a role in the process of basic emotions developing into more complex emotions.

Darwin concluded, through a survey, that humans share universal emotive expressions and suggested that animals likely share in these to some degree. Social constructionists disregard the concept that emotions are universal. Others hold an intermediate stance, suggesting that basic emotional expressions and emotion are universal but the intricacies are developed culturally. A study by Elfenbein and Ambady indicated that individuals within a particular culture are better at recognising other cultural members' emotions. [43]

Primates Edit

Primates, in particular great apes, are candidates for being able to experience empathy and theory of mind. Great apes have complex social systems young apes and their mothers have strong bonds of attachment and when a baby chimpanzee [44] or gorilla [45] dies, the mother will commonly carry the body around for several days. Jane Goodall has described chimpanzees as exhibiting mournful behavior. [46] Koko, a gorilla trained to use sign language, was reported to have expressed vocalizations indicating sadness after the death of her pet cat, All Ball. [47]

Beyond such anecdotal evidence, support for empathetic reactions has come from experimental studies of rhesus macaques. Macaques refused to pull a chain that delivered food to themselves if doing so also caused a companion to receive an electric shock. [48] [49] This inhibition of hurting another conspecific was more pronounced between familiar than unfamiliar macaques, a finding similar to that of empathy in humans.

Furthermore, there has been research on consolation behavior in chimpanzees. De Waal and Aureli found that third-party contacts attempt to relieve the distress of contact participants by consoling (e.g. making contact, embracing, grooming) recipients of aggression, especially those that have experienced more intense aggression. [50] Researchers were unable to replicate these results using the same observation protocol in studies of monkeys, demonstrating a possible difference in empathy between monkeys and apes. [51]

Other studies have examined emotional processing in the great apes. [52] Specifically, chimpanzees were shown video clips of emotionally charged scenes, such as a detested veterinary procedure or a favorite food, and then were required to match these scenes with one of two species-specific facial expressions: "happy" (a play-face) or "sad" (a teeth-baring expression seen in frustration or after defeat). The chimpanzees correctly matched the clips to the facial expressions that shared their meaning, demonstrating that they understand the emotional significance of their facial expressions. Measures of peripheral skin temperature also indicated that the video clips emotionally affected the chimpanzees.

Rodents Edit

In 1998, Jaak Panksepp proposed that all mammalian species are equipped with brains capable of generating emotional experiences. [53] Subsequent work examined studies on rodents to provide foundational support for this claim. [54] One of these studies examined whether rats would work to alleviate the distress of a conspecific. [55] Rats were trained to press a lever to avoid the delivery of an electric shock, signaled by a visual cue, to a conspecific. They were then tested in a situation in which either a conspecific or a Styrofoam block was hoisted into the air and could be lowered by pressing a lever. Rats that had previous experience with conspecific distress demonstrated greater than ten-fold more responses to lower a distressed conspecific compared to rats in the control group, while those who had never experienced conspecific distress expressed greater than three-fold more responses to lower a distressed conspecific relative to the control group. This suggests that rats will actively work to reduce the distress of a conspecific, a phenomenon related to empathy. Comparable results have also been found in similar experiments designed for monkeys. [56]

Langford et al. examined empathy in rodents using an approach based in neuroscience. [57] They reported that (1) if two mice experienced pain together, they expressed greater levels of pain-related behavior than if pain was experienced individually, (2) if experiencing different levels of pain together, the behavior of each mouse was modulated by the level of pain experienced by its social partner, and (3) sensitivity to a noxious stimulus was experienced to the same degree by the mouse observing a conspecific in pain as it was by the mouse directly experiencing the painful stimulus. The authors suggest this responsiveness to the pain of others demonstrated by mice is indicative of emotional contagion, a phenomenon associated with empathy, which has also been reported in pigs. [58] One behaviour associated with fear in rats is freezing. If female rats experience electric shocks to the feet and then witness another rat experiencing similar footshocks, they freeze more than females without any experience of the shocks. This suggests empathy in experienced rats witnessing another individual being shocked. Furthermore, the demonstrator's behaviour was changed by the behaviour of the witness demonstrators froze more following footshocks if their witness froze more creating an empathy loop. [59]

Several studies have also shown rodents can respond to a conditioned stimulus that has been associated with the distress of a conspecific, as if it were paired with the direct experience of an unconditioned stimulus. [60] [61] [62] [63] [64] These studies suggest that rodents are capable of shared affect, a concept critical to empathy.

Horses Edit

Although not direct evidence that horses experience emotions, a 2016 study showed that domestic horses react differently to seeing photographs of positive (happy) or negative (angry) human facial expressions. When viewing angry faces, horses look more with their left eye which is associated with perceiving negative stimuli. Their heart rate also increases more quickly and they show more stress-related behaviours. One rider wrote, 'Experienced riders and trainers can learn to read the subtle moods of individual horses according to wisdom passed down from one horseman to the next, but also from years of trial-and-error. I suffered many bruised toes and nipped fingers before I could detect a curious swivel of the ears, irritated flick of the tail, or concerned crinkle above a long-lashed eye.' This suggests that horses have emotions and display them physically but is not concrete evidence. [65]

Birds Edit

Marc Bekoff reported accounts of animal behaviour which he believed was evidence of animals being able to experience emotions in his book The Emotional Lives of Animals. [66] The following is an excerpt from his book:

A few years ago my friend Rod and I were riding our bicycles around Boulder, Colorado, when we witnessed a very interesting encounter among five magpies. Magpies are corvids, a very intelligent family of birds. One magpie had obviously been hit by a car and was laying dead on the side of the road. The four other magpies were standing around him. One approached the corpse, gently pecked at it-just as an elephant noses the carcass of another elephant- and stepped back. Another magpie did the same thing. Next, one of the magpies flew off, brought back some grass, and laid it by the corpse. Another magpie did the same. Then, all four magpies stood vigil for a few seconds and one by one flew off.

Bystander affiliation is believed to represent an expression of empathy in which the bystander tries to console a conflict victim and alleviate their distress. There is evidence for bystander affiliation in ravens (e.g. contact sitting, preening, or beak-to-beak or beak-to-body touching) and also for solicited bystander affiliation, in which there is post-conflict affiliation from the victim to the bystander. This indicates that ravens may be sensitive to the emotions of others, however, relationship value plays an important role in the prevalence and function of these post-conflict interactions. [67]

The capacity of domestic hens to experience empathy has been studied. Mother hens show one of the essential underpinning attributes of empathy: the ability to be affected by, and share, the emotional state of their distressed chicks. [68] [69] [70] However, evidence for empathy between familiar adult hens has not yet been found. [71]

Dogs Edit

Some research indicates that domestic dogs may experience negative emotions in a similar manner to humans, including the equivalent of certain chronic and acute psychological conditions. Much of this is from studies by Martin Seligman on the theory of learned helplessness as an extension of his interest in depression:

A dog that had earlier been repeatedly conditioned to associate an audible stimulus with inescapable electric shocks did not subsequently try to escape the electric shocks after the warning was presented, even though all the dog would have had to do is jump over a low divider within ten seconds. The dog didn't even try to avoid the "aversive stimulus" it had previously "learned" that nothing it did would reduce the probability of it receiving a shock. A follow-up experiment involved three dogs affixed in harnesses, including one that received shocks of identical intensity and duration to the others, but the lever which would otherwise have allowed the dog a degree of control was left disconnected and didn't do anything. The first two dogs quickly recovered from the experience, but the third dog suffered chronic symptoms of clinical depression as a result of this perceived helplessness.

A further series of experiments showed that, similar to humans, under conditions of long-term intense psychological stress, around one third of dogs do not develop learned helplessness or long-term depression. [72] [73] Instead these animals somehow managed to find a way to handle the unpleasant situation in spite of their past experience. The corresponding characteristic in humans has been found to correlate highly with an explanatory style and optimistic attitude that views the situation as other than personal, pervasive, or permanent.

Since these studies, symptoms analogous to clinical depression, neurosis, and other psychological conditions have also been accepted as being within the scope of emotion in domestic dogs. The postures of dogs may indicate their emotional state. [74] [75] In some instances, the recognition of specific postures and behaviors can be learned. [76]

Psychology research has shown that when humans gaze at the face of another human, the gaze is not symmetrical the gaze instinctively moves to the right side of the face to obtain information about their emotions and state. Research at the University of Lincoln shows that dogs share this instinct when meeting a human, and only when meeting a human (i.e. not other animals or other dogs). They are the only non-primate species known to share this instinct. [77] [78]

The existence and nature of personality traits in dogs have been studied (15,329 dogs of 164 different breeds). Five consistent and stable "narrow traits" were identified, described as playfulness, curiosity/fearlessness, chase-proneness, sociability and aggressiveness. A further higher order axis for shyness–boldness was also identified. [79] [80]

Dogs presented with images of either human or dog faces with different emotional states (happy/playful or angry/aggressive) paired with a single vocalization (voices or barks) from the same individual with either a positive or negative emotional state or brown noise. Dogs look longer at the face whose expression is congruent to the emotional state of the vocalization, for both other dogs and humans. This is an ability previously known only in humans. [81] The behavior of a dog can not always be an indication of its friendliness. This is because when a dog wags its tail, most people interpret this as the dog expressing happiness and friendliness. Though indeed tail wagging can express these positive emotions, tail wagging is also an indication of fear, insecurity, challenging of dominance, establishing social relationships or a warning that the dog may bite. [82]

Some researchers are beginning to investigate the question of whether dogs have emotions with the help of magnetic resonance imaging. [83]

Elephants Edit

Elephants are known for their empathy towards members of the same species as well as their cognitive memory. While this is true scientists continuously debate the extent to which elephants feel emotion. Observations show that elephants, like humans, are concerned with distressed or deceased individuals, and render assistance to the ailing and show a special interest in dead bodies of their own kind, [84] however this view is interpreted by some as being anthropomorphic. [85]

Elephants have recently been suggested to pass mirror self-recognition tests, and such tests have been linked to the capacity for empathy. [86] However, the experiment showing such actions did not follow the accepted protocol for tests of self-recognition, and earlier attempts to show mirror self-recognition in elephants have failed, so this remains a contentious claim. [ citation needed ]

Elephants are also deemed to show emotion through vocal expression, specifically the rumble vocalization. Rumbles are frequency modulated, harmonically rich calls with fundamental frequencies in the infrasonic range, with clear formant structure. Elephants exhibit negative emotion and/or increased emotional intensity through their rumbles, based on specific periods of social interaction and agitation. [87]

Cats Edit

It has been postulated that domestic cats can learn to manipulate their owners through vocalizations that are similar to the cries of human babies. Some cats learn to add a purr to the vocalization, which makes it less harmonious and more dissonant to humans, and therefore harder to ignore. Individual cats learn to make these vocalizations through trial-and-error when a particular vocalization elicits a positive response from a human, the probability increases that the cat will use that vocalization in the future. [88]

Growling can be an expression of annoyance or fear, similar to humans. When annoyed or angry, a cat wriggles and thumps its tail much more vigorously than when in a contented state. In larger felids such as lions, what appears to be irritating to them varies between individuals. A male lion may let his cubs play with his mane or tail, or he may hiss and hit them with his paws. [89] Domestic male cats also have variable attitudes towards their family members, for example, older male siblings tend not to go near younger or new siblings and may even show hostility toward them.

Hissing is also a vocalization associated with either offensive or defensive aggression. They are usually accompanied by a postural display intended to have a visual effect on the perceived threat. Cats hiss when they are startled, scared, angry, or in pain, and also to scare off intruders into their territory. If the hiss and growl warning does not remove the threat, an attack by the cat may follow. Kittens as young as two to three weeks will potentially hiss when first picked up by a human. [90]

Honeybees Edit

Honeybees ("Apis mellifera carnica") were trained to extend their proboscis to a two-component odour mixture (CS+) predicting a reward (e.g., 1.00 or 2.00 M sucrose) and to withhold their proboscis from another mixture (CS−) predicting either punishment or a less valuable reward (e.g., 0.01 M quinine solution or 0.3 M sucrose). Immediately after training, half of the honeybees were subjected to vigorous shaking for 60 s to simulate the state produced by a predatory attack on a concealed colony. This shaking reduced levels of octopamine, dopamine, and serotonin in the hemolymph of a separate group of honeybees at a time point corresponding to when the cognitive bias tests were performed. In honeybees, octopamine is the local neurotransmitter that functions during reward learning, whereas dopamine mediates the ability to learn to associate odours with quinine punishment. If flies are fed serotonin, they are more aggressive flies depleted of serotonin still exhibit aggression, but they do so much less frequently.

Within 5 minutes of the shaking, all the trained bees began a sequence of unreinforced test trials with five odour stimuli presented in a random order for each bee: the CS+, the CS−, and three novel odours composed of ratios intermediate between the two learned mixtures. Shaken honeybees were more likely to withhold their mouthparts from the CS− and from the most similar novel odour. Therefore, agitated honeybees display an increased expectation of bad outcomes similar to a vertebrate-like emotional state. The researchers of the study stated that, "Although our results do not allow us to make any claims about the presence of negative subjective feelings in honeybees, they call into question how we identify emotions in any non-human animal. It is logically inconsistent to claim that the presence of pessimistic cognitive biases should be taken as confirmation that dogs or rats are anxious but to deny the same conclusion in the case of honeybees." [8]

Crayfish Edit

Crayfish naturally explore new environments but display a general preference for dark places. A 2014 study [91] on the freshwater crayfish Procambarus clarkii tested their responses in a fear paradigm, the elevated plus maze in which animals choose to walk on an elevated cross which offers both aversive and preferable conditions (in this case, two arms were lit and two were dark). Crayfish which experienced an electric shock displayed enhanced fearfulness or anxiety as demonstrated by their preference for the dark arms more than the light. Furthermore, shocked crayfish had relatively higher brain serotonin concentrations coupled with elevated blood glucose, which indicates a stress response. [92] Moreover, the crayfish calmed down when they were injected with the benzodiazepine anxiolytic, chlordiazepoxide, used to treat anxiety in humans, and they entered the dark as normal. The authors of the study concluded ". stress-induced avoidance behavior in crayfish exhibits striking homologies with vertebrate anxiety."

A follow-up study using the same species confirmed the anxiolytic effect of chlordiazepoxide, but moreover, the intensity of the anxiety-like behaviour was dependent on the intensity of the electric shock until reaching a plateau. Such a quantitative relationship between stress and anxiety is also a very common feature of human and vertebrate anxiety. [93]

Are zoo animals happy? There's a simple empathy test we can apply

By Marc Bekoff - Jessica Pierce
Published April 16, 2017 3:00PM (EDT)

(Getty/Jean-Christophe Verhaegen)


Excerpted from "The Animals’ Agenda: Freedom, Compassion, and Coexistence in the Human Age" by Marc Bekoff and Jessica Pierce ( Beacon Press, 2017). Reprinted with Permission from Beacon Press.

The plight of animals in entertainment has gained unprecedented public attention over the past several years, and much of the consciousness-raising has occurred by way of a particular orca whale named Tilikum, known by his nickname, Tilly. Tilly was captured near Iceland in November 1983. When he was only two years old, he was torn away from his family and his ocean home. After a number of years of being transferred from one aquarium to another, Tilly was finally acquired by SeaWorld San Diego, and became one of the star attractions and moneymakers for the theme park. But the years of captivity and maltreatment took a toll on Tilly, and he started behaving erratically. He eventually killed one of his trainers, in front of a horrified audience. The details of Tilly’s tragic life and fateful end were beautifully captured in a documentary called "Blackfish" (2013). By weaving together ethological details about the cognitive, emotional, and social lives of orcas in the wild with a catalog of the abuses and deprivations experienced by Tilly, the film leaves the viewer in no doubt that SeaWorld is a living hell for these sensitive and intelligent creatures, who go crazy and must be pumped up with psychoactive drugs like Valium to control their behavior.

SeaWorld, for its part, has seen ticket sales plummet. In March 2016, SeaWorld announced that it will end its orca breeding program, won’t obtain new orcas from other sources, will begin replacing its theatrical orca shows with shows that exhibit the whales’ natural behaviors, and will have no orcas at all in any new parks around the world. Further, SeaWorld has pledged to invest millions of dollars for the rescue and rehabilitation of marine animals. (And in a nod to animal welfare more broadly, it also pledged to use cage-free eggs, gestation-crate-free pork, and sustainably sourced fishes at SeaWorld venues, and offer more vegetarian and vegan options.) This is a very good beginning, and we can look forward to the day when venues like SeaWorld, including terrestrial zoos, morph into sanctuaries in which the animals’ lives are put first and foremost. It is vitally important to keep working toward these goals. As the longtime animal advocate Gretchen Wyler once noted, “Cruelty can’t stand the spotlight.”

So how do animal well-being and freedom fare within the various entertainment venues that center on animals? Before we get to our main focus—zoos—consider a few instances of animals being used for entertainment.

Last Chain on Billie: Bridging the Empathy Gap

Billie, as she became known, was a majestic Asian elephant who was captured and shipped to America as a baby. She was taught to perform circus tricks such as standing on a small tub and balancing on one foot. Audiences clapped and marveled at her skills, but they didn’t know about Billie’s miserable behind-the-scenes life as she was hauled across the country and kept chained for hours on end when she wasn’t performing. Billie suffered enormous physical and psychological trauma but she finally got a lucky break when she was rescued and sent to a sanctuary for performing elephants in Tennessee. While she would never again be chained or beaten, her abuse had taken its toll, and Billie refused to interact with the other elephants or to allow anyone to remove the chain around her leg. For five years, Billie wouldn’t forget her past, and the chain remained. Finally, Billie allowed caregivers to remove it with a bolt cutter, from the other side of a fence. As Carol Bradley writes in her wonderful book "Last Chain on Billie," “It was almost as if Billie realized what they were attempting to do. The expression on her face softened and she stopped swinging at the fence. She lifted her foot again, this time higher than before, pushed up against the bars of the fence and rotated her ankle first one way and then another.” Billie didn’t give any indication that she understood the significance of being freed from the chain, and “headed out to the sand pile to wake [her elephant friend] Frieda from her nap.” When the chain finally fell from her leg, she “picked it up with her trunk, then dropped it and walked away.” Billie, Bradley writes, “had better things to do.”

Circuses involve humiliation, punitive training techniques, and poor living conditions for captive animals. Rodeos and dog- and cockfighting make spectacles of animals through overt violence. Dog- and horse racing are exploitative and often involve physical injury to the animals. Even venues in which humans can interact with “wild” animals and where the animals are not subjected to overt abuse can be harmful. Opportunities to swim with dolphins or pet wild tiger cubs involve a loss of freedom for the animals and a disruption of their lives, for the purpose of momentary human delight.

Billie’s story is hardly unique, and it captures just what happens to numerous entertainment animals whose hearts and spirits are broken as they are abused “in the name of entertainment.” A very useful thought experiment for bridging the empathy gap is to ask, “Would I do it to my dog?” This question brings home not only what entertainment animals endure, but also how our companion animals, who aren’t any more sentient than the animals used in entertainment, nevertheless enjoy a higher status. Most people would never imagine letting a dog be treated in such disrespectful and abusive ways.

The Case of Zooed Animals

Let’s now turn our attention to animals displayed at zoos and aquariums, where humans are simply observers, where an attempt is made to provide animals with suitable, even naturalistic, habitat, and where veterinary care and nutritious food are provided to all residents. Are animals living under these conditions happy and content, as zoo brochures would have us believe? The simple answer is no, they are not. Some zoos, particularly the thousands of roadside attractions, are shockingly mismanaged, and animals suffer from neglect, poor care, small, barren cages, and no attention to their species-specific or individual needs. But there are many zoos that seek to maintain the highest standards of care for animals, often through the application of animal welfare science. These are the focus of our attention. It is easy to see the problems for animals at the worst places it is more interesting to take a look at the better institutions and what they are doing to improve the lives and well-being of animals. We can see how welfare science can make an enormous difference in the lives of animals. We can also see why good welfare is not and can never be good enough and how the animals on display suffer from huge losses of freedom.

According to the Association of Zoos and Aquariums there are more than 10,000 zoos around the world. In the United States, as of September 2015, there were 230 AZA-accredited zoos and an estimated 2,400 “animal exhibitors” licensed by the USDA. (This number does not include individuals who own one or more exotic animals as pets.) AZA accreditation means that an institution has met certain standards for animal care and management, paying attention, for instance, to proper housing, nutrition, and social groupings. More than ten thousand different species are held in zoos around the world.

In zoos, welfarism and welfare science are hard at work. As with animal welfare on factory farms and in laboratories, science casts its sheen over our interactions with animals in zoos. A special issue of the Journal of Applied Animal Welfare Science (JAAWS) devoted to “issues in zoo animal management” suggests that welfare problems in zoos are issues of management, not ethics, and that good, scientifically informed management strategies can resolve welfare concerns. As in other realms, the welfarist paradigm controls the dialogue about animals. It focuses public and scientific attention on what we can do better for animals within the current paradigm. But it never pushes the questioning beyond the doors of the animals’ cages. Zoo workers justify the keeping of animals in captivity by appealing to the welfarist paradigm, saying that animals in captivity are safe and comfortable and better off, in some respects, than they would be in the wild. Yet as ethicist Koen Margodt puts it, zoos are essentially “welfare arks” in which animals are collected, purportedly to save them from extinction, but where human interests are put before the interests of individual animals.

Indeed, although zoos are open to the public, and sometimes even funded by local initiatives and taxes, what goes on behind the scenes is hidden from the public, unless and until problems become so severe that the bad news spills out, which it often does. The Smithsonian’s National Zoo in Washington, DC, for instance, was recently the subject of an investigation into mismanagement and animal welfare problems related to the zoo’s Cheetah Conservation Station. The zoo decided to double the number of animals in its CCS exhibits, but without increasing the amount of space allocated to the animals or preparing in advance where and how the animals would be housed. The results, not surprisingly, were tragic. Because there was no exhibit space ready for them, two newly acquired hornbill birds were kept in a shack for seven months. When zoo workers complained to management, the birds were put into the wallaby exhibit, which stressed out the wallaby, who bloodied his nose on the enclosure wall when he tried to run away. Holly, one of two newly acquired red river hogs, became so malnourished she died. A dama gazelle and a kudu both died after running into barriers in their enclosures and breaking their necks. A young Przewalski’s horse also died after he broke his neck in a cage at the zoo’s Conservation Biology Institute in Front Royal, Virginia. Przewalski’s horses are listed as endangered animals, and in the recent past were listed as critically endangered, so this loss is not only tragic for the individual, but also for this species as a whole.

Zoo officials insisted there were no problems, and the animals were doing just fine. Steve Feldman, a spokesman for the AZA, said in a news interview that the zoo had recently been evaluated and had met the AZA’s rigorous criteria for animal welfare, and that the events at the National Zoo were just natural. “The circle of life that occurs for all living things repeats itself in zoos on a daily basis,” he said. “Just as with humans, we notice more when it happens to celebrities, we tend to take more notice when these things happen at a place like the National Zoo because their zoo animals are the celebrities of the animal world.”

The introduction to the JAAWS special issue on management of zoos, by Cynthia Bennett, tells us that zoos and aquariums are committed to enhancing the welfare of nonhuman animals in captivity. Bennett admits, as does nearly everyone writing in the zoo-animal welfare literature, that our techniques and practices are not ideal and that animals are suffering. Animal welfare science was developed within the context of agricultural animals, and it isn’t all that easy to transfer welfare considerations from one venue to another, because the goals and focus are different. Within agriculture, the focus is on the productivity of animals over a very short time period. This doesn’t translate seamlessly into a zoo setting, where animals are expected to live out their natural life span and where the goal is to keep them healthy, not necessarily to make them fat. Furthermore, welfare science in agriculture is focused on identifying and responding to poor welfare states, and not to actually optimizing happiness or quality of life. And finally, zoos generally forgo common domestic animals like pigs and cows, about whose welfare we have a great deal of information, and instead hope to house exotic, endangered, or charismatic species about whose welfare in captivity we know very little to nothing. Zoo administrators and managers simply can’t know what each of the thousands of different species of animal need and want within the captive environment. As a result, not all of these various creatures “are managed in fully evidenced-based, optimized ways.”

Bennett concludes, “Acceptable standards and best practices remain elusive and are often subject to debate.” Yet she takes for granted that holding animals in captivity is perfectly acceptable and normal. Welfare science deflects attention away from moral concerns over animal captivity, because it stays focused on immediate welfare challenges and provides a nice veneer of scientific and thus ethical acceptability to the overall endeavor. When Bennett poses questions that might shape future discussion, she asks whether we might rethink the accepted rules of facility design “that give people more room and freedom to move than the captive animals they come to see.” This is a great question, because it exposes the irony and the fundamental insult of zoos: animals have had their basic freedoms stolen and are viewed as commodities, to feed human pockets.

What Kinds of Violations of Freedom Do Zoo Animals Experience?

The basic welfare problem of zoos, and the violation that causes the most misery, is the loss of the big F, Freedom. Attempts to provide bits and pieces of the Five Freedoms are of no consolation to an animal who has lost his most cherished possession.

What Do Zooed Animals Want?

This is easy, and we don’t need any preference studies to figure it out. They want to be free from captivity. Whether they are born into captivity or “born free,” wild animals “want” to live in a setting in which they can engage the repertoire of evolved behaviors that define them as a species.

Zoo Management: Welfare for Profit

In some respects, zooed animals are like farmed animals, because every aspect of their lives is managed by their human keepers. In welfarist lingo, this is “zoo animal management.” Indeed, running a successful zoo is much like running a successful business, balancing the costs of procuring and managing inventory against the profits from sales.

Zoo animals are carefully bred using studbooks and genetic analysis social groupings are manipulated animals are fed and watered at set times deaths are scheduled and orchestrated by veterinarians or zoo managers reproductive cycles are watched, sometimes controlled birth is carefully managed at some zoos, newborns (the core genetic inventory) are taken from their mothers and killed if not needed, or sold off, if necessary to optimize the zoo’s holdings and maximize profit. The costs of providing for animal welfare are always balanced against profit and optimization.

As in the other animal-centered venues we’ve discussed so far, money is one of the key drivers of zoo-animal welfare. Zoos have discovered that good welfare is more profitable, at least on the whole, than poor welfare. The one-hundred-million-odd people who visit zoos each year in the United States, for example, want to view animals who are active and engaged. It is much more fun to watch a lion stalking past the viewing window than to watch a lion sleeping or pacing back and forth, and far more entertaining to watch penguins dive and swim than to see them huddled on a fake glacier in a refrigerated cage.

Just as welfare of food animals is measured in productivity (number of eggs, liters of milk, quantity of muscle), so too is the welfare of zoo animals often measured not by how happy they are, but by how well they are serving the needs of zoo administrators and the public and how well they are generating money. What are the measures of welfare success used by zoos? One is fecundity the more fertile a given animal the better they are thought to be doing. There is some science behind this, of course. Stress has a pronounced effect on reproduction. In particular, the stress hormone cortisol suppresses the secretion of reproductive hormones and can disrupt estrus cycles. So, poor reproduction is a sign of poor welfare. The corollary is assumed also to be true: good reproduction equals good welfare. And reproduction is one of the key drivers of a healthy zoo industry. A successful breeding program allows a zoo to build the “captive gene pool” of a given species, which the zoo industry can then use to further replicate animals. Zoos make money selling animals to other zoos and live breeding animals—itinerant sperm donors—or their semen are traded around among zoos, as zoos seek to create the best collection. It is a great big game of musical animals and musical semen, though not a game that’s very much fun for the animals involved. (Marc first heard the term “musical semen” used by Julie Woodyer of Zoocheck Canada.)

One of the more nettlesome management dilemmas faced by zoos is the best way to deal with overly successful breeding and limited zoo space. And on this issue, American and European zoos take different approaches. In American zoos, the preferred approach is to make active use of contraceptives. All manner of animals, from chimpanzees to small rodents, are fed hormones to control ovulation and prevent pregnancies. The upside is that unwanted offspring don’t have to be killed. The downside is that many animals never get to engage in some of the most basic natural behaviors: giving birth and raising young. European zoos are less keen on denying their zoo animals these fundamental experiences, and will often allow animals to mate and bear offspring. As Bengt Holst, director of conservation for the Copenhagen Zoo, said in an interview, “We’d rather they have as natural behavior as possible. We have already taken away their predatory and anti-predatory behaviors. If we take away their parenting behavior, they have not much left.” Once the young reach the age at which, in the wild, they would become independent of their parents, they are removed and, if they aren’t needed for future breeding, they are killed.

Life span is another measure of good management/good welfare. Zoo managers often say things like, “Look! Our animals live even longer than they would have in the wild, which means they are safe and happy.” But for the animals themselves, of course, quantity isn’t necessarily the same as quality. There is the huge issue of having to live one’s entire life in captivity. But there are other problems, too. Because their lives are fairly sedentary, animals in zoos develop some of the same diseases as overfed and underexercised humans. Obesity can be a serious problem, leading some zoos to pursue absurd fixes like a treadmill built for Maggie, a fat elephant in the Alaska Zoo. She refused to use it. Animals fed a suboptimal diet over many years can develop other nutritional problems, such as hyperparathyroidism in large cats and cholesterol granulomas in meerkats. Hsing-Hsing, the famous panda housed at the National Zoo in Washington, DC, had a favorite treat, which his keepers kindly provided nearly every day: the waist-expanding Starbucks blueberry muffin.

When individuals in captivity live to be senior citizens, unique welfare problems can emerge. One study, for example, found that aged zoo animals, despite appearing healthy, often suffer from chronic health conditions, such as painful osteoarthritis, that go undiagnosed by zoo veterinary staff. On the other hand, over the past several years, as researchers and zoo staff have become better at recognizing the welfare challenges of geriatric animals, some zoos actually spend considerable energy and resources taking care of elderly or ill animals who cannot be displayed. Thus, it is not always true that animals living for a long time is good for a zoo’s bottom line. Moreover, although some species and some individuals may have longer lifespans in captivity, there are many species and individuals who do not.

Girafficide, aka Zoothanasia

One of the management tools of zoos is so-called management euthanasia, or what Marc has termed “zoothanasia.” Sometimes, of course, zoo animals are killed to relieve suffering. Animals become very old or terminally ill, and their pain cannot be adequately addressed with veterinary care. These account for some of the animals euthanized. But healthy animals are also killed, and for much more mercenary reasons. They are considered disposable when breeding programs are too successful or when a species is overrepresented.

The poster child for zoothanasia is Marius the giraffe. Marius lived at the Copenhagen Zoo until 2014, when he ceased to be genetically valuable. The zoo’s stated reason for “euthanizing” the eighteen-month-old giraffe was that he was not going to be a good breeder, because his genes were already well represented in the giraffe breeding program. Several other zoos and sanctuaries offered to take Marius in, but Copenhagen Zoo proceeded with the killing. As further public spectacle—or, as the zoo saw it, as an educational opportunity par excellence—Marius was publicly dismembered and fed, in large pieces, to some of the zoo’s carnivores. The most shocking thing is that Marius’s case is not an isolated one. Animals considered unwanted, surplus, or genetically unfit are routinely killed at zoos around the country and around the world. It is part of normal zoo-management practices. So too is what zoo managers call “breed and cull,” where animals are allowed to reproduce but the offspring are killed.

One of the core ideologies of animal welfare science is that euthanasia of animals is acceptable. There is a strong preference for death over discomfort, and an unwillingness to accept death as a harm or even as a significant event in the lives of animals. This attitude detracts from our commitment to positive welfare, because we fail to acknowledge that the potential for future enjoyment might be valuable to animals. The acceptability of killing as a management strategy must be challenged. In April 2016 some good news came out of the Antwerp Zoo. It has introduced a no-kill policy for surplus animals. Healthy individuals will be allowed to live, even if the zoo doesn’t have space for them.

Captivity Is Harder on Some Than Others

As University of Guelph’s Georgia Mason notes, the diverse species held in zoos vary “in the propensities for good captive health and welfare.” Some species tend to be healthier, longer-lived, and more fecund than their wild counterparts others survive and breed less well in captivity, and seem to suffer more psychologically. Why?

One risk factor that is very clear, and for which there are excellent data, is home-range size. Mason analyzed home-range size in relation to risk of developing stereotypies in zoos in several species. She found that animals with large home-ranges are at high risk of developing welfare problems in captivity. Polar bears, for example, have an average home range of 80,000 square kilometers. The average size of a polar bear enclosure in a zoo is one-millionth of this. Polar bears in captivity have high rates of stereotypies such as repetitive pacing and route tracing. Lions have home ranges of 350–900 square kilometers rates of stereotypies in zoos are 48 percent. Elephant home range is 1,500–3,700 square kilometers. Zoo managers and welfare researchers have concluded that optimal size for an elephant zoo enclosure is 1.24 acres, which is about 0.005 square kilometers. Almost two-thirds of elephants in captivity develop stereotypies.

Another risk factor is what a species of animal eats and how they have evolved to procure food, and how well their natural foraging behaviors can be replicated in captivity. According to Mason’s research, “prey-chase” distances can predict stereotypic pacing in carnivores, as for wolves, who tend to chase prey over long distances, and are likely to develop stereotyped pacing. Another interesting set of research data on ruminant species suggests that browsers, who feed on woody shrubs, don’t do as well in captivity as grazers, perhaps because the behavioral needs of grazers are easier to simulate. She also predicts that dietary specialists (who eat a very narrow range of food) may be at greater risk for poor welfare and stereotypies than generalists (who can eat a variety of foods and are flexible in foraging style).

One of the factors that seems to protect captive animals is phenotypic plasticity, or the ability to alter behavior to suit current conditions. For example, what are sometimes called “weed species” (such as deer and coyotes) seem to do well just about anywhere. Also, being sedentary and of limited range (like sloths and koalas) seems to “preadapt” a species to captivity, as does being gregarious (like flamingos), as long as they have friends with whom to interact. Sometimes the factors that allow animals to do well in captivity are mysterious to zoo managers and researchers. For example, ring-tailed lemurs seem to thrive in all sorts of captive environments, but nobody is quite sure why.

Understanding the factors that predispose animals to poor welfare in captivity can help zoos develop effective enrichment strategies. For example, mimicking prey-chase behaviors or foraging styles can allow animals to engage in at least a small range of food-related behaviors. It can also help zoo managers and animal advocates establish a list of animals that simply should not be held captive. This list must include large mammals such as elephants, dolphins, and whales, whose social organization and territories cover vast expanses in the wild. It must include large carnivores such as lions, polar bears, and wolves, whose prey-chase behavior cannot be mimicked in captivity.

Making Zoo Animals Happy

Zoos love to showcase charismatic polar bears, because they are moneymakers who attract a lot of visitors. But the bears don’t fare well, and visitors often become concerned when they see the bears behaving strangely, such as rocking or pacing back and forth. Whether or not they understand what they are seeing, visitors find stereotypies disturbing to watch.

Just as for laboratory animals and farm animals, stereotypies are common in animals held in zoos and aquariums. One study, for example, found that polar bears spend an average of 11 percent of their day engaged in stereotypic behavior, and that stereotypy was correlated with higher fecal glucocorticoid concentrations, which mean higher levels of stress. Stereotypies are the physical manifestations of a pathology caused by confinement. In fact, this pathology has been given a name: zoochosis. The term was first used by Born Free Foundation’s Bill Travers, who rightly identified the abnormal repetitive, obsessive behaviors of zoo animals as a form of psychosis.1 Captivity literally drives animals mad. Animals in zoos can be seen pacing back and forth, tracing and retracing a particular route through their enclosure, plucking out all their feathers or pulling out all their hair, scratching or rubbing or licking themselves to the point of serious self-injury, biting the bars of their cages for hours on end, and engaging in what zoo managers euphemistically call “regurgitation and reingestion” (eating their own vomit).

Zoos can meet some of the needs of animals, but they surely cannot meet them all, and some animals’ needs are harder to provide than others or are simply impossible to provide. When an animal’s needs are not being adequately met, or when she faces “adverse conditions” such as frustration at not being able to perform a highly motivated behavior such as hunting or scent-marking, the frustrated behavior “spills over” into stereotypy. Stereotypies are also thought to be caused by brain dysfunction brought on by stress-induced damage to the central nervous system.

This stress-induced damage may be caused by the animal’s current environment, or might be a result of earlier trauma. Either way, stereotypies are widely regarded as a serious welfare concern.

As we noted above, stereotypies are common in captive polar bears, though absent from the wild. Giving polar bears certain “enrichments” seems to help. For example, giving bears more dry land area and visual access beyond their enclosure have both been shown to reduce pacing. Often, the more naturalistic and free-living a captive animal’s environment, the fewer problems with stereotypies a zoo will see. Enrichment studies emerged primarily in response to the prevalence of stereotyped behaviors. How can we make bored and frustrated animals stop engaging in abnormal behaviors? By giving them stuff to do. But enrichment is also part of another related trend in animal welfare science, specifically paying more attention to positive welfare states. “Welfare” is being redefined as not merely the absence of or ability to successfully cope with negative experiences, but the promotion of states of enjoyment and happiness. This is an important shift in perspective.

Enrichment emerged as a topic in animal welfare science in the early 1980s, in relation to how the quality of an animal’s housing affects health and well-being—what is called “environmental enrichment.” Enrichment research fell into some disrepute among scientists during the 1990s, over concerns that animal interests are not necessarily served by enrichments. Since then, attention to enrichment has grown. Zoos are starting to have enrichment programs and departments, and to keep people on staff as enrichment managers. Some examples of zoo enrichments give a sense of what they try to accomplish. One of the most popular trends is to provide naturalistic zoo enclosures, which aim to give animals a tiny slice of home. A barren cage might be replaced with an “African savanna” full of tall grass and a few plastic “trees.” Cement “rocks” might be added to a penguin exhibit. Monkeys might be given climbing structures, “vines” on which they can swing, and platforms that provide some visual diversity. Elephants might be provided a mud pit in which they can cool off. One study showed that giving animals the choice to go outside, whether or not they opted to take advantage of it, had a positive behavioral effect. At the Brookfield Zoo in Chicago, the red panda exhibit is built around a giant magic tree. At semi-random intervals, steel cups emerge from “knotholes” in the tree. Sometimes the cups contain food and sometimes they don’t, which the zoo’s curator says is just like nature: sometimes food just appears. At the Woodland Park Zoo in Seattle, the orangutans are given cut branches or hay or banana leaves so they can build nests. For jaguars, food and spices are hidden inside logs, so that the animals will be stimulated to search their exhibit for the scents. Big Cat Rescue in Tampa rotates animals into a “vacation enclosure,” which is much bigger and more interesting than the normal enclosures.

At the Beijing Zoo, officials have undertaken an environmental-enrichment initiative that involves allowing animals access to privacy. As one zoo official said in a news interview, “Animals have the right to be seen as well as not to be seen . . . Beijing zoo is continuing to add animals while allowing them the freedom ‘not to be seen.’ The plan is so effective that some unknowing tourists have even been ‘tricked’ into complaining that the zoo has fewer animals than previously.” It may not seem like simply being watched would be a significant concern for animals, but research suggests that the mere presence of zoo visitors can be stressful for animals. For example, a recent study explored so-called “visitor effects” on little penguins and found that after exposure to visitors the birds showed increased aggression, vigilance, and avoidance behaviors such as huddling. Enrichments are designed to improve animal welfare, and sometimes even to make animals happy. But they have a secondary purpose, which is to increase the entertainment value of the animals on display. Bored animals are boring to watch. People want to see the otters and seals swimming they want to see the lions moving around, not just lying on a rock or hiding behind a tree. So keepers try to get the animals busy doing things that are fun and interesting to watch. According to a newspaper report, the Maryland Zoo has a full-time employee dedicated to enrichment, and one of this employee’s tricks is to spray Calvin Klein Obsession perfume—which is made with a synthetic form of civetone, a civet cat pheromone—on surfaces in the big cats’ enclosures, to encourage tracking and scent-marking behaviors.

Dr. Hal Markowitz, a longtime advocate of enrichment, marvels, in his book "Enriching Animal Lives," at how far we have come conceptually in understanding the reasons for enrichment and the ways to provide it. Yet he is at the same time dismayed at how little financial support is actually provided for enrichment. It remains “something of a nicety rather than a fundamental need in the eyes of many.” If we are to continue keeping animals in confinement, this needs to change. Making animals happier must be a top priority, and written into the budgets of zoo managers. Nevertheless, we need to remember that enrichment is just a Band-Aid solution. It serves, like the Valium given to SeaWorld’s whales, to manage the symptoms. But it can’t treat the underlying disease. Only freedom from captivity can really resolve the illness.

Research raises new questions about animal empathy

The emotions of rats and mice and the mental infrastructure behind them promise to illuminate the nature of human emotions, including empathy and nurturance, a Washington State University neuroscientist writes in this Friday's issue of the journal Science.

Jaak Panksepp, Baily Endowed Chair of Animal Well-Being Science and a professor of Veterinary and Comparative Anatomy, Pharmacy and Physiology, makes his case in a Perspectives column responding to research in which rats helped other rats with no explicit rewards at stake. The research, Panksepp writes, "raises questions about the affective experiences of animals other than humans."

Panksepp, who has pioneered work in how core emotions stem from deep, ancient parts of the brain, said there remains a good deal of resistance in the scientific community towards the notion that "nonhuman animals have affective experiences, and that these can and should be studied in empirical ways."

But he argues that recent advances in neuroscience are letting researchers look at how animal affect, or emotions, control learning, memory and behavior.

"Simplified models of empathy, as in mice and rats, offer new inroads for understanding our own social-emotional nature and nurture," he writes. "Such knowledge may eventually help us promote nurturant behaviors in humans."

Panksepp elaborated on his essay in a recent correspondence with the Washington State University News Center:

Q: Humans are under the impression that they are the animal with the greatest feelings and certainly have the greatest capacity to empathize with other creatures. Is this a mistaken assumption? Why?

Panksepp: There is no question that all other animals have emotional feelings. The science is strong for that. And all our strongest basic emotional feelings come from brain networks all mammals share. Unfortunately, currently we can't scientifically compare the intensity or greatness of feelings across species.

However, because we have a greater capacity to think than most, we can do more with our emotions than other animals. We can write music. Create poetry. And because of our higher mental abilities, we also have greater capacities for both empathy among strangers and cruelty. There are hints that across modern history empathy has been winning out over cruelty. But then one looks at the 20th century and wonders.

Still, the only way that empathy will continue to grow is if our higher mind gets in touch with the better angels of our lower minds—with maternal care and social joy being among the most important.

Q: If I read you correctly, the logic of attributing empathy to other, lower order animals grows out of the way our brain reflects our evolution, with higher order thinking and feeling on the more recently evolved outer layers but key, core emotions lying deep in the center. So while an animal may have a more rudimentary brain, its brain still has core functions that can include empathy. Right?

Panksepp: Indeed, we mammals share the basic tools for feeling and learning and perhaps even thinking. And empathy is reflected at all these levels. But our capacity for empathy would probably collapse without the basic emotions we share with other mammals.

Emotional contagion, a primitive form of empathic feelings, seems universal among mammals. Thinking about what others are thinking about and feeling seems much more developed in us than any other creature, except perhaps those with brains as big and complex as ours, like dolphins. Indeed, dolphins have certain brain areas that are more enlarged than ours—higher emotional regions of the brain that probably are needed for higher forms of empathy and positive fellow feelings.

Q: Why are people resisting the notion that nonhumans can have affective experiences?

Panksepp: I don't think animal lovers have much doubt about the fact that animals have emotional feelings. Many scientists have little more than doubts. Thus, science has not yet reached agreement on how to study the many kinds of basic feelings we have, and that many other animals surely have.

It is clear that when we finally understand their emotions, we will begin to have lasting scientific knowledge about our own. Only modern brain science can give us answers to questions such as, 'What are emotions?' and 'What are affective feelings?' It is clear that we can have many types of affective experiences—feeling good (positive) and bad (negative) in various ways. Certain positive and negative feelings are aroused by our sensory channels, like various forms of pain and taste. Others arise from inside our bodies, like hunger and thirst signals to the brain. And then there are emotional feelings that arise largely from complex networks that reside completely within our brains, but which move our bodies intensely in various ways.

These last kinds of feelings are most important for understanding our moods and psychiatric disorders. We now have a great deal of knowledge about which brain systems generate various basic emotional feelings—experiences like desire, anger, fear, lust, motherly love, grief and playfulness. Once we understand the brain chemistries that control these kinds of emotional feelings in animals, we will better understand ourselves, as well as develop much better medicines for human emotional problems.

Q: You have a zinger of an ending. If we better understand the affective processes of mouse and rat brains, we might be better able to help humans be more nurturing. I read it this way: Humans may have the greatest capacity for compassion and empathy on earth, owing in part to our consciousness, but at times we behave worse than rats. If we understand the core, instinctual capacity for empathy among all animals, we might be better humans in the humanistic sense.

Panksepp: Yes, I think the more we know about the emotions of other animals, the more we will understand our own emotions. Without the ancient emotional systems that all mammals share, our ability to be conscious is drastically impaired. The more we know about our animal emotions, which support the rest of our mental apparatus, the more ideas we will have about how to be better people. As we follow the old philosophical advice to, "know thyself," the more options we will have for being good to others and the world.

But until quite recently, an enormous gap in our knowledge has been any solid scientific knowledge about our emotional nature. Neuroscience is changing that. And when we really know ourselves, we will be able to think about ourselves more clearly as creatures of the world. What we do with this knowledge will vary from one mind to another. My hope is that our desire to care about others will grow. To do that well is one of the best ways to take care of yourself. . .and the world.

Do Animals Have Emotions?

One of the hottest questions in the study of animal behavior is, “Do animals have emotions?” And the simple and correct answer is, “Of course they do.” Just look at them, listen to them and, if you dare, smell the odors that pour out when they interact with friends and foes. Look at their faces, tails, bodies and, most importantly, their eyes. What we see on the outside tells us a lot about what’s happening inside animals’ heads and hearts. Animal emotions aren’t all that mysterious.

When I first began my studies three decades ago—asking the question, “What does it feel like to be a dog or a wolf?”—researchers were almost all skeptics who spent their time wondering if dogs, cats, chimpanzees and other animals felt anything. Since feelings don’t fit under a microscope, these scientists usually didn’t find any, and, as I like to say, I’m glad I wasn’t their dog!

But now there are far fewer skeptics prestigious scientific journals publish essays on joy in rats, grief in elephants and empathy in mice and no one blinks. The question of real importance is not whether animals have emotions, but why animal emotions have evolved. Simply put, emotions have evolved as adaptations in numerous species. They serve as a social glue to bond animals with one another and also catalyze and regulate a wide variety of social encounters among friends and foes.

Emotions permit animals to behave adaptively and flexibly, using various behavior patterns in a wide variety of venues. Research has shown that mice are empathic rodents, but it turns out they’re fun-loving as well. We also read accounts of pleasure-seeking iguanas amorous whales angry baboons elephants who suffer from psychological flashbacks and post-traumatic stress disorder (PTSD—elephants have a huge hippocampus, a brain structure in the limbic system that’s important in processing emotions) grieving otters, magpies and donkeys sentient fish and a sighted dog who serves as a “seeing-eye dog” for his blind canine buddy. Today, the paradigm has shifted to such an extent that the burden of “proof” now falls on those who still argue that animals don’t experience emotions.


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Many researchers also recognize that we must be anthropomorphic (attribute human traits to animals) when we discuss animal emotions, but that if we do it carefully, we can still give due consideration to the animals’ points of view. No matter what we call it, researchers agree that animals and humans share many traits, including emotions. Thus, we’re not inserting something human into animals rather, we’re identifying commonalities and then using human language to communicate what we observe. Being anthropomorphic is doing what’s natural and necessary to understand animal emotions.

We might expect to find close, enduring and endearing emotional relationships between members of the same species, but improbable relationships also occur between animals of wildly different species, even between animals who are normally predator and prey! Such is the case for Aochan, a rat snake, who befriended a dwarf hamster named Gohan at Tokyo’s Mutsugoro Okoku Zoo, and a lioness in northern Kenya who adopted a baby oryx (usually an appetizer before a larger meal) on five different occasions.

It’s bad biology to argue against the existence of animal emotions. Scientific research in evolutionary biology, cognitive ethology (the study of animal minds) and social neuroscience support the view that numerous and diverse animals have rich and deep emotional lives. (Here I focus on mammals, although there are data showing that birds and perhaps fish experience various emotions as well as pain and suffering.)

Charles Darwin’s well-accepted ideas about evolutionary continuity—that differences among species are differences in degree rather than kind—argue strongly for the presence of animal emotions, empathy and moral behavior. Continuity allows us to connect the “evolutionary dots” among different species to highlight similarities in evolved traits, including individual feelings and passions. All mammals (including humans) share neuroanatomical structures, such as the amygdala and neurochemical pathways in the limbic system that are important for feelings.

Mirror neurons help explain feelings such as empathy. Research on these neurons supports the notion that individuals can feel the feelings of others. Mirror neurons allow us to understand another individual’s behavior by imagining ourselves performing the same behavior and then mentally projecting ourselves into the other individual’s shoes.

To what degree various species share this capability remains to be seen, but there is compelling evidence that humans are not alone in possessing it. Diana monkeys and chimpanzees help one another acquire food, and elephants comfort others in distress. Mirror neurons also help explain observations of rhesus monkeys who won’t accept food if another monkey suffers when they do so, and empathic mice who react more strongly to painful stimuli after they observed other mice in pain.

The borders between “them” and “us” are murky and permeable, and the study of animal emotions helps inform the big question of just who we are. Another big question for which answers are revealed by studying animal passions is, “Can animals be moral beings?” In my development of the phenomenon that I call “wild justice,” I argue that they can. Many animals know right from wrong and live according to a moral code.

When people tell me that they love animals because they’re feeling beings and then go on to abuse them, I tell them that I’m glad they don’t love me. I often ask researchers who conduct invasive work with animals or people who work on factory farms, “Would you do that to your dog?” Some are startled to hear this question, but if people won’t do something to their own dog that they do daily to other dogs or to mice, rats, cats, monkeys, pigs, cows, elephants or chimpanzees, we need to know why. There’s no doubt whatsoever that, when it comes to what we can and cannot do to other animals, it’s their emotions that should inform our discussions and our actions on their behalf.

Emotions are the gifts of our ancestors. We have them, and so do other animals. We must never forget this. When it comes to animal welfare, we can always do better. Most of the time, “good welfare” is not good enough.

Do Insects Have Emotions and Empathy?

A recent survey suggests that for most of us that have ever had a pet companion, it’s a no-brainer that mammals and birds are emotional creatures, sharing emotions with multiple species and not just their own. Yet despite the thousands of YouTube videos and hundreds of recent scientific studies presenting easily accessible evidence and examples, not everyone thinks so.

It was only in 2012 that scientists finally agreed that nonhuman animals are conscious beings. Meanwhile, in the laboratory, we have only just discovered that dogs display immensely complex human-like emotions like jealousy, and we are only just deciphering how cows express positive emotions through the white of their eyes. But what about insects?

What are emotions?

Briefly, let’s touch on the definition of what an emotion is, which is still largely debated even today. While most of us are pretty convinced that emotions exist (seeing as we experience them ourselves), defining emotions is easier said than done.

A universal definition for emotions that is equally applicable across all academic fields, from neuroscience to psychology to philosophy, has been near impossible to achieve. In fact, the well-known neuroscientist, Professor Joseph LeDoux, went so far as to suggest that we erase the word “emotion” from our scientific vocabulary entirely.

Although there are literally hundreds of different definitions for emotion, the most universal definition we could find originates from an article in Science called Emotion, cognition and behavior:

“…emotions include (but are not limited to) certain expressive behaviors that are associated with internal brain states that we, as humans, subjectively experience as ‘feelings’.”

Pretty vague! And due to the subjective nature of it all, it confounds making comparisons between species as we can’t simply ask other species what they feel, and as such, may wrongfully interpret or not even notice key emotional behaviors.

Another way of looking at things is that emotions are essentially ways for an organism to interpret the neural map of their body and surrounding environment in the part of their brain that monitors homeostasis. As Wikipedia puts it:

“Homeostasis is the property of a system in which variables are regulated so that internal conditions remain stable and relatively constant.”

Basically, an organism experiences too much or too little of something either within them or around them in the environment (i.e. something deviates from neutrality or optimum balance), which is then detected by our brains (i.e. via neural maps of the body). We then experience our brain’s recognition of these changes or differences through two main avenues: As subjective feelings that motivate and influence our perceptions, actions and behaviors, and as emotional behavior to express and communicate our state to others. This goes for primordial emotions like sexual desire, as well as more complex social emotions like embarrassment.

An experiment with honeybees

A fantastic example of an insect emotion experiment was carried out on our all-important flower loving friends, honeybees.

As just discussed, emotions influence our perceptions and behavior. So, imagine that your house has just been ransacked by burglars and you are feeling shocked, upset and really, really, REALLY angry. You’re so mad in fact that despite your friends trying to do and say everything possible to cheer you up, you feel so pessimistic that you simply see the downside in everything. In fact, you’re so upset that even your favorite food seems totally unappetizing.

Well, this is exactly what happened with the bees. The poor guys were shoved into a vortex (a machine used to vigorously mix chemicals) for one minute to simulate a badger attack on their hive and presumably cause them to feel complete and utter bee rage.

The bees were then presented with different solutions containing different proportions of two smelly chemicals: Octanone that the bees had been trained to associate with a yummy sugary treat and hexanol that they had been trained to associate with a bitter nasty taste.

Bees that had been shaken became pessimistic, glass half-empty characters that were more likely to react to the nasty smell in the mixtures and recoil as opposed to being attracted to the yummy smell — a result of presumably being pretty irritated. Unshaken bees on the other hand remained their more optimistic, glass half-full selves and were more likely to see the mixtures as half-appetizing, as opposed to half-disgusting like their bad-tempered counterparts did. Moreover, there were emotionally relevant changes in neurotransmitter levels in the shaken up bees, like serotonin and dopamine.

This can scientifically be interpreted as the act of shaking the bees creating an internal neurological state in them that affected their subsequent behavior that was associated with changes in brain chemistry. More specifically, this implies that agitated honeybees exhibit pessimistic cognitive biases.

Yet the authors were reluctant to say that this anger-like state was a definitive emotion. It is interesting to note that if dogs did the same thing and turned down their food after their owner had just died for example, many wouldn’t even question that the behavior was emotional.

An experiment with drosophila flies

A similar experiment was conducted with hungry fruit flies. This time the experimenters tried to induce primal fear by casting a shadow over them to mimic the presence of an overhead predator. This was much like the fear we experience when we hear an unexpected gunshot, making us feel and behave apprehensive until we consider the coast is clear and manage to calm down. And this is exactly what seemed to happen with the fruit flies.

When the fake predator was introduced and then removed, potentially freaked out and hungry flies ignored their food until many minutes later when they eventually calmed down. This suggests that an emotion-like state affected their behavior even after the stimulus was gone. Other key building blocks of emotions like scalability were also demonstrated, i.e. repeating the predator’s shadow simulation multiple times making the flies even more freaked out, taking them longer to calm down and dig into their food.

However, the authors of the study made it clear that although the flies’ responses were more complex than a simple avoidance reflex, they will not take the next leap and classify it as a bona fide emotion. What they did say was in the title of the study itself: “Behavioral responses to a repetitive visual threat stimulus express a persistent state of defensive arousal in Drosophila.”

Do insects have empathy?

As we mentioned earlier, a second aspect of emotions is the expression of emotional behavior that allows other individuals to be aware of our emotions and respond to them. As such, in order to detect and understand those emotions we have the ability to empathize and respond in kind.

In an experiment hot off the press, woodlice have shown empathic-like behavior. The researchers demonstrated that calm woodlice reduced their more excited neighbors causing them to also become calm.

One can argue that this is simply the mimicking of behaviors, as opposed to recognizing and then matching emotions. Again however, remember that if one dog barks in what we interpret as an upset nervous manner, and causes the other dog to do the same, we would tend to automatically assume that the first dog passed on its emotion to the second one if they adopt the same postures and emotional behavior. Moreover, a study published this year quite clearly stated that emotional contagion was observed in pigs as a form of empathy.

Do insects experience emotions?

To be strictly honest we still can’t say to what degree insects experience emotions on par with our own yet, although these early experiments are certainly setting the foundations for a future where we recognize that all animals have emotions of some sort.

Hopefully, by mapping the neural circuitry that underlies the fear-like behavior in flies, anger-like behavior in bees or empathic-like behavior in woodlice we may be one step closer to comparing insects’ experiences of feelings with our own. With insect brains surprising even entomology experts in their extraordinary similarities with our own brains despite marked differences, the similarities may be more profound than we would like to think.

Thankfully, we are beginning to stop with our centuries long obsession with human species superiority. It can be considered part of what has led us to ravage and raid the earth like parasites and to downplay emotions felt by other animals. Humans feel love, other animals merely bond. Humans feel jealous, but other animals merely guard resources.

While we cannot experience what it feels like for a bee to have a bee in its bonnet, a fly to feel like a bundle of nerves, or for a woodlouse to chill-out with its buddies, neither can we experience other human’s emotions… it is only because we can communicate (to a degree) that we know other humans have emotions too. Bear in mind that emotions are so subjective that we aren’t especially accurate at understanding other humans’ emotions at the best of times, never mind another species!

As we continually delve deeper into the genetic and neurological basis of emotions, the evolutionary origins of emotions will undoubtedly be gradually unearthed. Perhaps this may take us one step closer to letting go of our overinflated humanist ego and realize that ALL animals experience emotions of some kind.

“Even insects express anger, terror, jealousy and love, by their stridulation.”

Anderson DJ, & Adolphs R (2014). A framework for studying emotions across species. Cell, 157 (1), 187-200 PMID: 24679535

Current biology : CB, 21 (12), 1070-3 PMID: 21636277

Gibson WT, Gonzalez CR, Fernandez C, Ramasamy L, Tabachnik T, Du RR, Felsen PD, Maire MR, Perona P, & Anderson DJ (2015). Behavioral responses to a repetitive visual threat stimulus express a persistent state of defensive arousal in Drosophila. Current biology : CB, 25 (11), 1401-15 PMID: 25981791

Harris, C., & Prouvost, C. (2014). Jealousy in Dogs PLoS ONE, 9 (7) DOI: 10.1371/journal.pone.0094597

Proctor HS, & Carder G (2015). Measuring positive emotions in cows: Do visible eye whites tell us anything? Physiology & behavior, 147 , 1-6 PMID: 25862928

Reimert I, Bolhuis JE, Kemp B, & Rodenburg TB (2015). Emotions on the loose: emotional contagion and the role of oxytocin in pigs. Animal cognition, 18 (2), 517-32 PMID: 25385575

Walker, J., Mcgrath, N., Handel, I., Waran, N., & Phillips, C. (2014). Does owning a companion animal influence the belief that animals experience emotions such as grief? Animal Welfare, 23 (1), 71-79 DOI: 10.7120/09627286.23.1.071


The difference between animal cognition and animal emotion is recognized by ethicists. Animal cognition covers all aspects related to the thought processes in animals. Though the topics related to cognition such as self-recognition, memory, other emotions and problem-solving have been investigated, the ability to share the emotional state of another has now been established in hens. [4] [5] [6] [7]

"We found that adult female birds possess at least one of the essential underpinning attributes of 'empathy' the ability to be affected by, and share, the emotional state of another." [4]

Chickens have the basic foundations of emotional empathy. [8] [9] [10] [11] Empathy is sometimes regarded as a form of emotional intelligence and is demonstrated when hens display signs of anxiety when they observed their chicks in distressful situations. The hens have been said to "feel their chicks' pain" and to "be affected by, and share, the emotional state of another." [2]

A study funded by the BBSRC and published in 2011 [10] was the first to demonstrate that chickens possess empathy and the first study to use both behavioral and physiological methods to measure these traits in birds. [7] Chicks were exposed to a puff of air, which they find mildly distressing. During the exposure, their mother's behaviour and physiological responses were monitored non-invasively. The hens altered their behaviour by decreased preening, increased alertness, and an increased numbers of vocalisations directed to their chicks — behaviours interpreted as a demonstration of concern. Furthermore, the hens' heart rate increased and eye temperature decreased. [4] [12]

In a 2012 follow-up study, the same researchers repeated the air-puff procedure with hens observing familiar-but-unrelated adults rather than chicks. They found that the adult hens "actually produced slightly more alarm and warning vocalisations than the chicks in the previous study produced distress vocalisations", but that "the only response detected in the observer hens was an increase in sitting. . There was no indication of heightened physiological arousal and the sitting posture more likely indicated that hens were in a calmer, more ‘relaxed’ state." They concluded that the "result clearly shows that empathic responses in hens are not facilitated by warning or alarm vocalisations". They felt that further study was warranted, perhaps with related adults, or with stimuli that prompted distress signals rather than alarm or warning signals. [13]

Fear Edit

Previous investigations established the indicators of an emotional response in chickens. Domestic chickens can be observed to have different states of alertness. Hens exhibit fear by increasing the time spent standing alert and increased preening. Before empathy in chickens was reported, other investigations demonstrated that hens avoid environments associated with higher preening rates and standing. [4]

Stress and empathy Edit

Empathetic response by hens is preceded by the determination that hens recognize distress in their chicks. Assessing the distress of chicks and the effect of the presence of their mother has been investigated using an air puff treatment. Each treatment chick and control were exposed to puffs of airs applied to their eyes in the presence and absence of their mothers. The responses interpreted as distress in the chicks were:

  • reduced temperature of the eye of the chick
  • increased ground pecking
  • increased preening
  • increased standing (compared to normal rates of movement) [14]

The response of the hens to the apparent distress of their chicks differed to those not exhibiting distress. When distress was recognized by a hen, her heart rate increased. This is correlated to the degree of distress exhibited by the chicks. If the hen is present, distress associated behaviours are less in the chick. Demonstrating empathy by hens toward their chicks is accompanied by the reduction of distress in the chicks. [14]

The specific emotional attribute of empathy in chickens has not been only investigated in terms of its existence but it has applications that have resulted in the designed reduction of stress in farm-raised poultry. [4]

At one time, a Virginia prison was planned to be transformed into 'Chicken empathy museum' by PETA to raise awareness of the emotional, empathetic nature of chickens. [15] [16] [17] [18]

Wild Justice

Let&rsquos get right to the point. In Wild Justice, we argue that animals feel empathy for each other, treat one another fairly, cooperate towards common goals, and help each other out of trouble. We argue, in short, that animals have morality.

Both popular and scientific media constantly remind us of the surprising and amazing things animals can do, know, and feel. However, when we pay careful attention to the ways in which animals negotiate their social environments, we often come to realize that what we call surprises aren&rsquot really that surprising after all. Take, for example, the story of a female western lowland gorilla named Binti Jua, Swahili for &ldquodaughter of sunshine,&rdquo who lived in the Brookfield Zoo in Illinois. One summer day in 1996, a three-year-old boy climbed the wall of the gorilla enclosure at Brookfield and fell twenty feet onto the concrete floor below. As spectators gaped and the boy&rsquos mother screamed in terror, Binti Jua approached the unconscious boy. She reached down and gently lifted him, cradling him in her arms while her own infant, Koola, clung to her back. Growling warnings at the other gorillas who tried to get close, Binti Jua carried the boy safely to an access gate and the waiting zoo staff.

This story made headlines worldwide and Binti Jua was widely hailed as an animal hero. She was even awarded a medal from the American Legion. Behind the splashy news, the gorilla&rsquos story was adding fuel to an already smoldering debate about what goes on inside the mind and heart of an animal like Binti Jua. Was Binti Jua&rsquos behavior really a deliberate act of kindness or did it simply reflect her training by zoo staff?

Even in the mid-1990s there was considerable skepticism among scientists that an animal, even an intelligent animal like a gorilla, could have the cognitive and emotional resources to respond to a novel situation with what appeared to be intelligence and compassion. These skeptics argued that the most likely explanation for Binti Jua&rsquos &ldquoheroism&rdquo was her particular experience as a captive animal. Because Binti Jua had been hand raised by zoo staff, she had not learned, as she would have in the wild, the skills of gorilla mothering. She had to be taught by humans, using a stuffed toy as a pretend baby, to care for her own daughter. She had even been trained to bring her &ldquobaby&rdquo to zoo staff. She was probably simply replaying this training exercise, having mistaken the young boy for another stuffed toy.

A few scientists disagreed with their skeptical colleagues and argued that at least some animals, particularly primates, probably do have the capacity for empathy, altruism, and compassion, and could be intelligent enough to assess the situation and understand that the boy needed help. They pointed to a small but growing body of research hinting that animals have cognitive and emotional lives rich beyond our understanding.

We&rsquoll never know why Binti Jua did what she did. But now, years later, the staggering amount of information that we have about animal intelligence and animal emotions brings us much closer to answering the larger question raised by her behavior: can animals really act with compassion, altruism, and empathy? The skeptics&rsquo numbers are dwindling. More and more scientists who study animal behavior are becoming convinced that the answer is an unequivocal &ldquoYes, animals really can act with compassion, altruism, and empathy.&rdquo Not only did Binti Jua rescue the young boy, but she also liberated some of our colleagues from the grip of timeworn and outdated views of animals and opened the door for much-needed discussion about the cognitive and emotional lives of other animals.

Wild Justice: What Are We Really Talking About?

Even a decade ago, at the time that Binti Jua rescued the injured boy, the idea of animal morality would have been met with raised eyebrows and a &ldquosurely you must be joking!&rdquo dismissal. However, recent research is demonstrating that animals not only act altruistically, but also have the capacity for empathy, forgiveness, trust, reciprocity, and much more as well. In humans, these behaviors form the core of what we call morality. There&rsquos good reason to call these behaviors moral in animals, too. Morality is a broadly adaptive strategy for social living that has evolved in many animal societies other than our own.

Our argument relies upon well-established and mostly uncontroversial research. We simply suggest that the many parts, taken together, represent an interesting and provocative pattern. Our most controversial move, of course, is to use the label &ldquomorality&rdquo to describe what we see going on in animal societies. This jump is controversial not for scientific reasons so much as philosophical ones, and we will keep these philosophical concerns in the foreground of our discussion.

Let us take you through the evidence. We invite you to enter into the lives of social animals. We show that these animals have rich inner worlds&mdashthey have a complex and nuanced repertoire of emotions as well as a high degree of intelligence and behavioral flexibility. They&rsquore also incredibly adept social actors. They form and maintain complex networks of relationships, and live by rules of conduct that maintain a delicate balance, a finely tuned social homeostasis.

Looking for the Bad, Looking for the Good: The More We Look the More We See

Here&rsquos a common distillation of Charles Darwin&rsquos theory of evolution. Natural selection, to borrow a popular metaphor from biology, is an evolutionary arms race. Life is a war of all against all, a ruthless and bloody battle, usually over sex and food. Mothers eat their young and siblings fight to the death against siblings (a phenomenon called siblicide). When we look at nature through this narrow lens we see animals eking out a living against the glacial forces of evolutionary conflict. This scenario makes for great television programming, but it reflects only a small part of nature&rsquos ineluctable push. For alongside conflict and competition there is a tremendous show of cooperative, helpful, and caring behavior as well.

To offer a particularly striking example, after carefully analyzing the social interactions of various primate species, primatologists Robert Sussman and Paul Garber and geneticist James Cheverud came to the conclusion that the vast majority of social interactions are affiliative rather than agonistic or divisive. Grooming and bouts of play predominate the social scene, with only an occasional fight or threat of aggression. In prosimians, the most ancestral of existing primates, an average of 93.2 percent of social interactions are affiliative. Among New World monkeys who live in the tropical forests of southern Mexico and Central and South America, 86.1 percent of interactions are affiliative, and likewise for Old World monkeys who live in South and East Asia, the Middle East, Africa, and Gibraltar, among whom 84.8 percent of interactions are affiliative. Unpublished data for gorillas show that 95.7 percent of their social interactions are affiliative. After about twenty-five years of research on chimpanzees, Jane Goodall noted in her book The Chimpanzees of Gombe, &ldquoit is easy to get the impression that chimpanzees are more aggressive than they really are. In actuality, peaceful interactions are far more frequent than aggressive ones mild threatening gestures are more common than vigorous ones threats per se occur much more often than fights and serious, wounding fights are very rare compared to brief, relatively mild ones.&rdquo These don&rsquot appear to be animals whose social lives are defined only by conflict.

The social lives of numerous animals are strongly shaped by affiliative and cooperative behavior. Consider wolves. For a long time researchers thought that pack size was regulated by available food resources. Wolves typically feed on prey such as elk and moose, both of which are bigger than an individual wolf. Successfully hunting such large ungulates usually takes more than one wolf, so it makes sense to postulate that wolf packs evolved because of the size of wolves&rsquo prey. However, long-term research by David Mech shows that pack size in wolves is regulated by social and not food-related factors. Mech discovered that the number of wolves who can live together in a coordinated pack is governed by the number of wolves with whom individuals can closely bond (the &ldquosocial attraction factor&rdquo) balanced against the number of individuals from whom an individual could tolerate competition (the &ldquosocial competition factor&rdquo). Packs and their codes of conduct break down when there are too many wolves.

As we begin to look at the &ldquogood&rdquo side of animal behavior, at what animals do when they&rsquore not eating each other or committing siblicide, we begin to take in just how rich the social lives of many animals are. Indeed, the lives of animals are shaped at a most basic level by &ldquogood&rdquo&mdashor what biologists call prosocial&mdashinteractions and relationships. Even more, it seems that at least some prosocial behavior is not a mere byproduct of conflict, but may be an evolutionary force in its own right. Within biology, early theories of kin selection and reciprocal altruism have now blossomed into a much wider inquiry into the many faces and meanings of prosocial behavior. And, it seems, the more we look, the more we see. There&rsquos now an enormous body of research on prosocial behavior, and new research is being published all the time on cooperation, altruism, empathy, reciprocity, succorance, fairness, forgiveness, trust, and kindness in animals ranging from rats to apes.

Even more striking, within this huge repertoire of prosocial behaviors, particular patterns of behavior seem to constitute a kind of animal morality. Mammals living in tight social groups appear to live according to codes of conduct, including both prohibitions against certain kinds of behavior and expectations for other kinds of behavior. They live by a set of rules that fosters a relatively harmonious and peaceful coexistence. They&rsquore naturally cooperative, will offer aid to their fellows, sometimes in return for like aid, sometimes with no expectation of immediate reward. They build relationships of trust. What&rsquos more, they appear to feel for other members of their communities, especially relatives, but also neighbors and sometimes even strangers&mdashoften showing signs of what looks very much like compassion and empathy.

It is these &ldquomoral&rdquo behaviors in particular that are our focus in Wild Justice. Here is just a sampling of some of the surprising things research has revealed about animal behavior and more specifically about animal morality in recent years.

Some animals seem to have a sense of fairness in that they understand and behave according to implicit rules about who deserves what and when. Individuals who breach rules of fairness are often punished either through physical retaliation or social ostracism. For example, research on play behavior in social carnivores suggests that when animals play, they are fair to one another and only rarely breach the agreed-upon rules of engagement&mdashif I ask you to play, I mean it, and I don&rsquot intend to dominate you, mate with you, or eat you. Highly aggressive coyote pups, to give just one example, will bend over backwards to maintain the play mood with their fellows, and when they don&rsquot do this they&rsquore ignored and ostracized.

Fairness also seems to be a part of primate social life. Researchers Sarah Brosnan, Frans de Waal, and Hillary Schiff discovered what they call &ldquoinequity aversion&rdquo in capuchin monkeys, a highly social and cooperative species in which food sharing is common. These monkeys, especially females, carefully monitor equity and fair treatment among peers. Individuals who are shortchanged during a bartering transaction by being offered a less preferred treat refuse to cooperate with researchers. In a nutshell, the capuchins expect to be treated fairly.

Many animals have a capacity for empathy. They perceive and feel the emotional state of fellow animals, especially those of their own kind, and respond accordingly. Hal Markowitz&rsquos research on captive diana monkeys strongly suggests a capacity for empathy, long thought to be unique to humans. In one of his studies, individual diana monkeys were trained to insert a token into a slot to obtain food. The oldest female in the group failed to learn how to do this. Her mate watched her unsuccessful attempts, and on three occasions he approached her, picked up the tokens she had dropped, inserted them into the machine, and then allowed her to have the food. The male apparently evaluated the situation and seemed to understand that she wanted food but could not get it on her own. He could have eaten the food, but he didn&rsquot. There was no evidence that the male&rsquos behavior was self-serving. Similarly, Felix Warneken and Michael Tomasello at the Max-Planck Institute for Evolutionary Anthropology in Leipzig, Germany, discovered that captive chimpanzees would help others get food. When a chimpanzee saw that his neighbor couldn&rsquot reach food, he opened the neighbor&rsquos cage so the animal could get to it.

Even elephants rumble onto the scene. Joyce Poole, who has studied African elephants for decades, relates the story of a teenage female who was suffering from a withered leg on which she could put no weight. When a young male from another group began attacking the injured female, a large adult female chased the attacking male, returned to the young female, and touched her crippled leg with her trunk. Poole believes that the adult female was showing empathy. There is even evidence for empathy in rats and mice.

Altruistic and cooperative behaviors are also common in many species of animal. One of the classic studies on altruism in animals comes from Gerry Wilkinson&rsquos work on bats. Vampire bats who are successful in foraging for blood that they drink from livestock will share their meal with bats who aren&rsquot successful. And they&rsquore more likely to share blood with those bats who previously shared blood with them. In a recent piece of surprising research, rats appear to exhibit generalized reciprocity they help an unknown rat obtain food if they themselves have been helped by a stranger. Generalized reciprocity has long been thought to be uniquely human.

The presence of these behaviors may seem puzzling to scientists or lay readers who still view animals from the old &ldquonature red in tooth and claw&rdquo framework. But puzzling or not, moral behaviors can be seen in a wide variety of species in a spectrum of different social contexts. And the more we look, the more we see.

What Is Morality and What Moral Behaviors Do Animals Exhibit?

Before we can discuss the moral behaviors that animals exhibit, we need to provide a working definition of morality. We define morality as a suite of interrelated other-regarding behaviors that cultivate and regulate complex interactions within social groups. These behaviors relate to well-being and harm, and norms of right and wrong attach to many of them. Morality is an essentially social phenomenon, arising in the interactions between and among individual animals, and it exists as a tangle of threads that holds together a complicated and shifting tapestry of social relationships. Morality in this way acts as social glue.

Animals have a broad repertoire of moral behaviors. It&rsquos sloppy business trying to squeeze these diverse behaviors into structured categories, but we need some way to organize and present a picture of moral behavior in animals. We envision a suite of moral behavior patterns that falls into three rough categories, around which we have organized our book. We call these rough categories &ldquoclusters,&rdquo a cluster being a group of related behaviors that share some family resemblances, and we identify three specific such clusters: the cooperation cluster, the empathy cluster, and the justice cluster. Wild justice is shorthand for this whole suite.

The cooperation cluster includes behaviors such as altruism, reciprocity, trust, punishment, and revenge. The empathy cluster includes sympathy, compassion, caring, helping, grieving, and consoling. The justice cluster includes a sense of fair play, sharing, a desire for equity, expectations about what one deserves and how one ought to be treated, indignation, retribution, and spite. We devote separate chapters to exploring each of these clusters in detail (cooperation in chapter 3, empathy in chapter 4, and fairness in chapter 5).

Forcing structure in this way raises many questions. Do the behaviors that we cluster together really belong in the same group? For example, is consolation behavior an example of an empathic response, or is it more closely related to cooperation and reciprocity? Are some behaviors more basic than others? For example, is empathy a necessary precursor to fairness? What are the interrelationships between and among behaviors, both evolutionarily and physiologically? Have these behaviors co-evolved? And are we correct in our claim that moral animals will have a behavioral repertoire that spans all three clusters?

Who Are the Moral Animals? Penciling in a Shifting Line

Many people will immediately want to know who the moral animals are. Can we draw a line that separates species in which morality has evolved from those in which it hasn&rsquot? Given the rapidly accumulating data on the social behavior of numerous and diverse species, drawing such a line is surely an exercise in futility, and the best we can offer is that if you choose to draw a line, use a pencil. For the line will certainly shift &ldquodownwards&rdquo to include species to which we would never have dreamed of attributing such complex behaviors, such as rats and mice.

Taking animal-behavior research as it stands now, there&rsquos compelling evidence for moral behavior in primates (particularly the great apes, but also at least some species of monkey), social carnivores (most well studied are wolves, coyotes, and hyenas), cetaceans (dolphins and whales), elephants, and some rodents (rats and mice, at the very least). This isn&rsquot a comprehensive catalogue of all animals with moral behavior it simply represents the animals whose social behavior has been studied well enough to provide ample data to draw conclusions. There are other species, such as many ungulates and cats, for which data are simply lacking. But it would not be surprising to discover that they, too, have evolved moral behaviors.

Research on primates currently provides the most robust account of moral behavior in animals. Given our evolutionary kinship with other primates, it seems reasonable to suppose that these species will have the most behavioral continuity with humans. And indeed, Jessica Flack and Frans de Waal have argued that nonhuman primates are the most likely animals to show precursors of human morality. Yet looking for &ldquoprecursors&rdquo of human morality, though interesting, is not the same as looking for moral behavior in animals. Furthermore, the assumption that primate behavior will be most similar to human behavior may actually prove incorrect. For example, Nobel Prizeûwinning ethologist Niko Tinbergen and renowned field biologist George Schaller have suggested that we might learn a lot about the evolution of human social behavior by studying social carnivores, species whose social behavior and organization resemble that of early hominids in a number of ways (divisions of labor, food sharing, care of young, and intersexual and intrasexual dominance hierarchies). For these reasons, we&rsquore interested in extending the research paradigm on animal morality well beyond primates.

Morality may be exclusive to mammals, and mammals are our focus in this book. At this point, however, it would be premature to pronounce other species lacking in moral behaviors. We simply do not have enough data to make hard and fast claims about the taxonomic distribution among different species of the cognitive skills and emotional capacities necessary for being able to empathize with others, behave fairly, or be moral agents. All must remain quite tentative at this point. It is possible, for example, that some birds, such as the highly intelligent corvids, have a kind of morality. In his book Mind of the Raven, biologist and raven expert Bernd Heinrich observed that ravens remember an individual who consistently raids their caches if they catch him in the act. Sometimes a raven will join in an attack on an intruder, even if he did not see the cache being raided. Is this moral? Heinrich seems to think it is. He says of this behavior, &ldquoIt was a moral raven seeking the human equivalent of justice, because it defended the group&rsquos interest at a potential cost to itself.&rdquo In two subsequent experiments, Heinrich confirmed that group interests could drive what an individual raven decides to do.

There is abundant evidence for the range of behaviors we&rsquore exploring in this book, so much so that the basic claim that these behavioral clusters are present to some degree in some animals isn&rsquot really controversial at all. But why take the further step and call these behavioral clusters moral, a label bound to raise hackles, rather than sticking to the seemingly more objective term prosocial?

Challenging and Revising Stereotypes about Animals: Bad Habits are Hard to Break

So far, very few scientists and other academics have been willing to use the term moral in relation to animal behavior without protective quotation marks (which signal a kind of &ldquowink, wink: we don&rsquot really mean æmoral&rsquo as in human morality&rdquo) or without some other modifying trick, as in the term proto-morality (read: &ldquothey may have some of the seeds of moral behavior, but obviously not morality per se&rdquo). Indeed, there is strong resistance to the use of the term &ldquomoral&rdquo in relation to the behavior of nonhuman animals, both from scientists and philosophers.

The belief that humans have morality and animals don&rsquot is such a longstanding assumption it could well be called a habit of mind, and bad habits, as we all know, are damned hard to break. A lot of people have caved in to this assumption because it is easier to deny morality to animals than to deal with the complex reverberations and implications of the possibility that animals have moral behavior. The historical momentum, framed in the timeworn dualism of us versus them, and the Cartesian view of animals as nothing more than mechanistic entities, is reason enough to dismissively cling to the status quo and get on with the day&rsquos work. Denial of who animals are conveniently allows for retaining false stereotypes about the cognitive and emotional capacities of animals. Clearly a major paradigm shift is needed, because the lazy acceptance of habits of mind has a strong influence on how science and philosophy are done and how animals are understood and treated.

The irony, of course, is that the field of animal behavior is already bursting with terminology that has moral color: altruism, selfishness, trust, forgiveness, reciprocity, and spite. All of these terms and more are used by scientists to describe the behavior of animals. Certain words like altruism, selfishness, and spite have been ascribed specific and carefully circumscribed meanings within the field of animal behavior&mdashmeanings that diverge from, and even sometimes contradict common usage. Other moral terms such as forgiveness, fairness, retribution, reciprocity, and empathy have joined the animal behavior lexicon, and retain, for now, their connection to the morality we know and live. Lay readers and even scientists are bound to be confused by this apparent lack of consistency. We plan to clear up some of this mess.

We could have coined a new word or phrase to describe our particular suite of prosocial behaviors in animals. The phrase &ldquoanimal morality&rdquo will certainly strike some people as odd, and perhaps even as an oxymoron. And in some respects, morality is not the most solicitous term. Morality is notoriously hard to define and there is disagreement about how best to understand what morality is. On the other hand, morality is a very useful term, because &ldquoanimal morality&rdquo challenges some stereotypes about animals and, as we&rsquoll see, about humans. It also emphasizes evolutionary continuity between humans and other animals, not only in anatomical structure, but also in behavior. And this emphasis, in our view, is important. Finally, morality is also a useful term because the root meaning&mdashmore, or custom&mdashcaptures an essential element of animal morality.

We need to be quite explicit that the meaning of morality is itself under consideration, and we&rsquore suggesting a shift in meaning. How we define morality will, of course, determine whether and to what extent animals have it. And yes, we&rsquore defining morality in such a way as to lend credence to our argument for evolutionary continuity between humans and animals. But this is not sleight of hand: our definition of morality is well supported both scientifically and philosophically and also by &ldquounscientific&rdquo common sense. We want to detach the word morality from some of its moorings, allowing us to rethink what it is in light of a huge pile of research from various fields that speaks to the phenomenon. We ask that you let us play freely with the term and, in the end, you can decide if you think &ldquoanimal morality&rdquo makes sense.

Watch the video: Do animals have empathy? (December 2021).