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Frans de Waal

The origins and evolution of empathy

Here at Empathy Lab we’re aiming to imbue new models of intelligence (machine learning and AI) with empathy and prosociality. Our favorite thing to do is work with bright and like-hearted folks doing the same work in their own fields. Frans de Waal is a trailblazer in the origins and natural expressions of empathy, a hero of ours.

Time selected Frans as one of the world’s 100 Most Influential Humans, he’s also a leading primatologist, as the CH Candler Professor of Primate Behavior and director of the Living Links Center at Emory University. His scientific work has been featured in Science, Nature, and Scientific American, his books include Chimpanzee Politics, Our Inner Ape, Are We Smart Enough to Know How Smart Animals Are? and, of course, one that’s near and dear to the heart of the Empathy Lab: The Age of Empathy.

We were thrilled to host Frans and learn from him just how old, nuanced, and beautiful empathy is in our world and how many species we share this trait with.

Empathy Lab: Frans, we’re so thrilled to have you with us today. Just to set the stage for our conversation, are we going to focus on animal empathy or human empathy too?

Frans: I’m gonna be speaking purely about animals and you can make for yourself the connections with humans, which is not so difficult to do.

Got it. Can you tell us a bit about how your work started? What sparked your curiosity in animal empathy?

Frans: I started my work on reconciliation behavior. As a student I discovered that chimps embrace and kiss after fights and that got me really interested. At that time all the biologists only studied aggressive behavior in animals and aggressive behavior in humans, and I thought the other side was actually much more interesting…so that’s how I got involved in all these things like empathy, self awareness, pro social tendencies and fairness.

Can you give us an example of reconciliation in the animal world? How would you describe this discovery?

Frans: After a fight between two male chimpanzees…one of them holds out his hand and begs the other for contract and after this they come together and they kiss and embrace, and that’s the reconciliation. It sounds very counterintuitive to come together after a fight. At the time everyone said aggression is a dispersive mechanism, aggression drives individuals apart, but actually, aggression brings individuals together.

This is how we define reconciliation: a friendly union between former opponents. If you define it this way you can see it all the time. In another sequence [of behaviors], a male chimp attacks a female, the female comes back 10 minutes later, she offers her hand for a hand kiss, and then there’s the mouth kiss, which is the typical way chimpanzees reconcile.

How do those actions compare to other animals? Is this sort of behavior specific to chimps?

Frans: Since the gestures and the kissing and everything is very similar [to humans] that is, I think, why reconciliation was discovered in chimpanzees, and now we know that reconciliation occurs in all sorts of animals.

Monkeys do all these things without eye contact. There is what we call the “whole bottom ritual”: one of [the monkeys] presents, the other one holds the hips, and that’s the ritual that these monkeys use for reconciliation and for greeting. They can do reconciliation without eye contact. I don’t think humans can do that. People try it on email and I don’t think it’s a big success. Humans and apes, they need eye contact for reconciliation. The monkeys and many other animals don’t. And I think that’s because humans read emotions in the eyes much more than monkeys do.

How do you go about measuring something like reconciliation?

Frans: In a group of monkeys, for example, you wait for a spontaneous fight, you follow the animals for 10 minutes or sometimes half an hour, and you do a post-conflict observation and see how often they get together after a fight. Then you do a match control observation on a different day where they had no fight to see how often they get together, and you see a much lower percentage. This is called post-conflict attraction.

It’s exactly the opposite of the dispersive mechanism, and this has been found in many, many different species. Basically, these monkeys come together more often after aggression than without aggression, and this is related to the need for them to repair relationships.

There are all sorts of species now — elephants, dogs, wolves, dolphins, hyenas — in which reconciliation has been found. There’s only one mammal in which it has not been found: the domestic cat. The domestic cat is a solitary hunter, and so we think they probably don’t need reconciliation.

Reconciliation is related to depending on each other — you need each other for cooperation — and dolphins and elephants are very cooperative animals and so they absolutely need each other, so they reconcile.

But animals who don’t need each other, they probably don’t. Reconciliation is actually now so widespread that if we find an animal that is social and does not reconcile we are extremely surprised.

Is there a parallel for humans? Is reconciliation universal, or nuanced?

Frans: If you take in the same data [for humans] as you do with a monkey group, you get the same sort of graphs. Young children do [reconciliation] physically, older children do it verbally. This has been done in many different cultures in the world now, and there are big cultural differences. For example, Japanese children reconcile much more than American children and the people who do these studies think it is because Japanese teachers don’t interfere in fights, and American teachers tend to interfere in fights, which hampers the development of conflict resolution skills.

Empathy was once a controversial topic for animals, a hangover from Descartes. Can you share your specific understanding of empathy?

Frans: When I started doing studies on empathy, you could not mention the words “empathy” and “animal” in the same sentence because animals did not have empathy, and that’s because the psychologists think in cognitive terms, so they say empathy is when you imagine the situation of somebody else, which is a very cognitive view of empathy.

And if you take that view then most animals don’t have it, but this is a dictionary definition of empathy: the ability to understand and share the feelings of another. You see immediately the two components: the understanding, which is cognitive, and the feeling, which is emotional, and I would say the emotional part is extremely important.

So here we have the two channels of empathy. You have a body channel, which includes synchronisation, motor mimicry, and emotional contagion. You talk with someone who is sad, you will have a sad expression on your face; you talk to someone who’s happy and laughing, you will be laughing too. That’s the body channel of emotional contagion, which starts at day one of life. Babies cry when other babies cry.

Then you have the cognitive channel, which is more complex (and that’s why you don’t find it in all animals), which is where you make a self-others distinction. Instead of fusing with the other, you set yourself apart so that you can better understand the other. If you fuse, you’ll have the same emotions as the other, which is not necessarily the best way to understand the other. So you need the self-other distinction, and then you can take the perspective of somebody else. That’s the cognitive part of empathy. In humans that kicks in later in life.

What does cognitive empathy look like in action?

Frans: We look at body synchronization by looking at yawn contagion. All vertebrates yawn and no one knows exactly why. There’s no good theory about yawning, but all animals do it. We look at yawn contagion by presenting our chimps with videos on an iPod [of other chimps that are yawning]. This has been done on dogs. This has been done on birds. It’s a measure of body synchronization. In our study we had two categories: the “in group”, which are chimps that they know, and an “out group”, which are chimps that they don’t know, and the effect only occurs with the chimps that they know.

This has been found in humans too. There’s an Italian field study where they looked at waiting rooms and train stations and places like that, and if you stand next to a stranger who yawns you’re going to be unaffected. If you stand next to your wife who’s yawning, you’re going to be yawning.

There’s sort of a universal bias and all the empathy studies: the social bias that you are more affected by individuals who are similar and familiar. So that’s universal. It’s almost like a characteristic of empathy that empathy is most easily-generated between individuals who know each other, or who are socially close.

And the flip side of that is that we have trouble having empathy for individuals who are strangers, who we don’t know, or who are different from us. Empathy is a very biased mechanism.

You have a special affinity for bonobos, which you sometimes call “primate hippies”. We love that. Can you tell us a bit about them and your work with them?

Frans: I consider bonobos the most empathic apes.They’re often neglected because they are very peaceful. Anthropologists don’t like peaceful animals. They think humans got where they were by killing everybody…but the bonobos are totally peaceful. They don’t kill each other. They have a lot of sex, and they’re equally close to us as the chimpanzee, so there’s absolutely no reason to neglect them. Everyone is always going with the chimpanzee because chimpanzees kill each other and they have territorial warfare, and the anthropologists love that kind of stuff, but I think they should look more at the bonobo.

Bonobos do all these things that chimps do with kissing and embracing, only they do it with sex. People think they’re gonna make love the whole day, but 10 seconds is actually a long time for bonobos. So I call it a genital handshake. It’s a short genital contact. It is clearly sex and sometimes there’s more involved, but bonobos resolve all their issues with sex basically, and they’re very effective at it.

You mentioned consolation as another aspect of your work. What is consolation and how is it expressed by the animals you study?

Frans: Consolation is a very interesting mechanism because consolation is how empathy studies got started in humans. Carolyn Zon Waxler was the first one who studied empathy in humans and in the beginning she was not respected because empathy was not an official topic. She was told not to publish in scientific journals, and to publish in women’s magazines. So she started to study empathy in children and the way she did that is that she would ask a family member to cry — an adult or a family member — and then she would look at how very young children would react to that. And she found that even two-year-old children who can barely walk, walk up to this person and they touch them and stroke them and if they can talk they talk to them and she called that empathic concern.

When I heard her talk about that I said, “Well if that’s empathy, then I have tons of empathy in my animals because they do that kind of stuff all the time!” There’s now also dog studies, and actually Carolyn Zon Waxler in her studies of children discovered by accident that the pets in the home were doing the same thing. They would also approach people who were distressed. And in the apes it is very human-like. The apes do very human-like behavior. With bonobos, one of them has lost the fight and the other one puts the arms around it, and that’s the consolation behavior of bonobos.

In humans, we take it for granted. Anyone who goes to a picnic with young children is in the consolation business the whole day. We take it for granted in humans, but in animals, it’s actually not that common. For example, monkeys do very little of it, but the apes do it all the time, so you can actually quantify it and study it.

We’d love for you to share about your study on consolation in orphaned bonobos, the tender relationship between trauma and consolation.

Frans: We study bonobos who live in a sanctuary and it’s very sad that the sanctuary exists, because there’s bushmeat hunting going on in Africa, so all these bonobos are traumatized orphans, except for a few who are born in the colony. So we have a study of traumatized orphans, which is really, in a way, interesting. There are [also] studies of human orphans. The Romanian orphanages have traumatized orphans and people have studied these kids, and they have a lot of trouble with empathy. They have a lot of trouble with emotion regulation, which is essential for empathy. You need to be able to down-regulate your emotions. Let’s say I’m affected by someone who’s distressed, you need to be able to down-regulate that in order to respond appropriately. And so orphans have a lot of trouble with empathy. And so it’s an interesting comparison. We have a few mother-reared bonobos and a lot of traumatized orphans.

And if you look at the data, what we find is that the young bonobos show quite a bit more consolation behavior than the adults. Twice as much. Recently we found the same thing in chimpanzees, that it decreases with age. We think it is because the adults become more selective. It’s not that they have less empathy than the young ones, but they are going to apply it more selectively and not for everybody like the young ones do. If you look at mother-reared ones, they do far better than everybody else. This is similar to the human studies where empathy is compromised in traumatized orphans.. So empathy requires a balanced emotional upbringing, which is what [the mother-reared bonobos] have and [the orphans] don’t have. We’ve been working recently on emotion regulation, which is much better developed in the juveniles who are raised by their own mothers than by the orphans.

Are there other animals with poignant behaviors of consolation or other acts of empathy?

Frans: I do a lot of elephant studies at the moment with Josh Plotnick. The elephants rumble with each other and they stick their trunks in each other’s mouths, which is a very vulnerable thing to do because if there’s one thing important for an elephant, it’s its trunk, and it’s a very sensitive organ. These are called vulnerable contact behaviors. Chimps do the same thing. One chimp will stick his hand in the mouth of another when they are consoling and reconciling. In a sense you make yourself vulnerable by sticking your hand in the mouth of another, it’s actually a stupid thing to do, nut we do these kinds of things. I heard a story once that the words “testament” and “testimony” come from the fact that people used to touch each other’s testicles when they went into a contract. Vulnerable contact behavior I would say.

Then we did a study on voles. I had a lot of behavioral data on empathy expression in lots of animals, but the mechanism behind it we never could put our finger on. Then James Birkhoff came to me and he said I have consolation behavior from voles, which immediately opened the door to look at the neuroscience behind it. And what James did with the voles is he would separate the male and the female — and these voles are monogamous, so males or females are very attached to each other. He would separate them, which is already stressful, and then bring them back together. He would sometimes separate them and in another room the partner would get some electrical shocks, and then be brought back. So you have two conditions. One is a separation reunion. The other one is a separation, plus stress, and a reunion. And what we found is that the partner who comes back when the other partner has been stressed is going to groom far more. So the one picks up on the stress of the other, and the one is just as equally stressed. You can measure the stress hormones and that one is physiologically just as stressed as the partner, even though it has not been shocked at all, and has just been sitting in its cage. So that’s emotional contagion.

Are there any limits to emotional contagion? And how does that work for humans?

Frans: We found first of all a social bias: It only works between voles who know each other, like mates or siblings. Then there’s emotional contagion, which you can measure physiologically. And there’s oxytocin involved. If you block the oxytocin receptors, the mechanism doesn’t occur, the consolation doesn’t occur. So the conclusion from our study was that empathy in voles works essentially the same as in humans. I think we may need more data to reach that conclusion but there’s no reason to think that it’s a different mechanism that’s going on.

There’s a study that was done at Michigan University where they would bring five students together, and then after a while they would tell one of them, “You’re going to give a talk to the rest,” which is very stressful. And then they would have 10 minutes to prepare and then they would give their talk. Then they measure the stress level of all the students involved. And if the speaker was very nervous, the others would be very nervous. If the speaker was very relaxed, the other ones would be very relaxed in terms of stress level. So that’s emotional contagion right here at the physiological level, almost identical to what we saw in the voles.

So we’re seeing the mechanisms behind empathy, also the neural mechanism for empathy in animals is actually very similar. We know that for voles. We will probably never know that in such detail for elephants or apes, but we think it’s a mammalian characteristic.

Wow. So these studies explore the emotional aspects of empathy. Can you share some studies around the cognitive side?

Frans: On the more cognitive side is perspective taking. In humans, there’s actually quite a bit of evidence that this comes up at the same time that mirror self-recognition comes up. At the age of two years, children have the self-other distinction that is required to take the perspective of somebody else…We think there is a connection between mirror self-recognition and empathy at the child level, and it has been proposed that at the species level the same thing is true, and very few species recognize themselves in the mirror, so let me say a few things about the measures that we have for perspective-taking.

One of the dimensions (and this is spontaneous behavior) is targeted helping. Targeted helping is helping that you provide to somebody else based on an understanding of the situation of somebody else, and very few animals do that kind of thing. Most animals are not good at taking the perspective of somebody else. A picture was sent to me of wild bonobos. And one of them has a poacher snare around her finger, which is extremely dangerous. And you can see how everyone [the other bonobos] is concerned. This is very serious and something needs to be done to help this female, and that’s targeted helping.

I [showed] you a video clip of a young elephant who has been struggling to get out of a ditch and keeps slipping back in. And an adult elephant then reacts to it. That requires that the adult understands the problem, which is not really an easy thing to do, you need to understand the goals of another individual, the situation, you have to see all the attempts that are being done, there’s maybe some distress calls involved, etc. That whole conglomeration of things needs to be understood by the other.

We have experiments done in Kyoto, on targeted helping. So this is an experiment in which one chimp has a choice of seven different tools, and another chimp in another compartment has a situation for which it needs one tool to collect food. So this chimp has to look at the situation of the other and judge the situation and decide which tool is needed and then pick the right tool and then be willing to hand it to the other [chimp], even though she’s not going to get any food for it. And they do that and that means that this chimp can take the perspective of the other one and understands what the other one needs under these circumstances. And I’m convinced that chimps and some other animals like elephants have that capacity of perspective taking.

How does mirror self-recognition play a role in this?

Frans: Now to move from emotional contagion, which is universal in all mammals and has been found also in birds, to perspective taking and targeted helping, which requires a self-other distinction that we think correlates with mirror self-recognition, this is what a chimp does with a mirror. Our chimp Raveena has a little hole in her head, and uses our iPhone as a mirror, and she inspects that hole in a very coordinated way. And chimps do this all the time. This is a natural thing for them. If I come with my sunglasses on, they use my glasses to look inside their mouth, which is a part of the body they want to see and never get to see. Monkeys don’t have mirror self-recognition. The monkey is very interested in the mirror, but the monkey does not use the mirror to inspect itself.

Because elephants are so empathic and altruistic we decided we needed some tests of mirror self-recognition. On one side of the [elephant’s] head he puts [white] paint and on the other side of the head he puts the same paint without the color, basically an invisible mark so the elephant feels being marked on both sides, but only one of them is visible. Just like the chimps, he wants to look inside his mouth, the inside of the mouth is a part of your body that you never get to see. You feel it the whole day, but you never get to see it. So we found that the elephants pass the mark test and so we think that actually the support for the idea that species who pass the mark test and children who pass the mark test, that’s a special moment as far as self-other distinction is concerned.

Circling back to targeted helping, is that the same as caring for another’s well-being? Or is well-being something else?

Frans: The well-being of others is a related topic. We started doing experiments on whether chimpanzees care about the well-being of others in a time where people had concluded that only humans care about the well-being of others. People were making big conclusions based on a few negative experiments. On this particular experiment, we put two chimps side-by-side, and we have a little table and a token. The chimps have been trained on tokens and they know that these tokens give them food. So there’s 30 of these tokens in a bucket. There’s a little table, there’s a partner, and two colors of tokens. If a chimp gives us a red token, which is the selfish token, they only feed her for this token. The partner gets nothing. And the partner is going to express her opinion about the test. If the chimp picks a green token we reward both of them. And that’s the whole experiment. So for the chimp who makes the choices, it doesn’t make any difference, he gets rewarded for every choice he makes. The only difference is, does my partner get something yes or no? And then if you look at the data, what we find is that they have a tendency to choose the prosocial token most of the time. If the partner draws attention to herself by climbing around and vocalizing and things like that, they make more prosocial choices. If the partner puts pressure on them by banging on the window and spitting and yelling, make less prosocial choices. Basically what we found is that the chimps prefer a situation where both of them get food. It’s not costly altruism.

Is there an experiment where the altruism was more costly, where the prosocial choice of one didn’t result in equal reward?

Frans: We did an experiment on elephants in which two elephants need to work together on a task. The elephants need to be able to pull a table closer to gain access to sunflower seeds, and they need a rope to do that. But if only one of them pulls the rope, then they both go hungry. This task has been tried on many different animal species and chimps can do it and elephants can do it. A lot of animals cannot do this because it requires that a response that has been rewarded many times now all of a sudden needs to be suppressed based on social information. Elephants have a lot of emotional control. I’m very interested in the issue of emotional control because I think all animals need to have it and do have it to some degree. Elephants are particularly good at that.

These concepts of cooperation and reciprocity get into the fabric of our interspecies social web. What are some other examples, for fun?

Frans: I’ve worked for 25 years with capuchin monkeys, and by accident we discovered that they care about what the other one is getting. This is sort of ridiculous. People have never thought that animals might care about the rewards that somebody else gets, because they’ve always tested animals singly. And since we always test our animals in pairs, we discovered that if the other one gets better food they get upset about that. And we started testing this out and the way we test it out is by putting two monkeys side by side, and giving them both grapes or both cucumber, or giving one of them grapes and the other one cucumber. Now grapes are 10 times better than cucumber for the monkeys. If you give them a choice, it’s always the grape they go for. If you just feed chimps different foods nothing happens. It needs to be a reward for performed actions. If both monkeys get cucumber, they perform the task at 100% almost. If the partner is getting grape, and you’re still getting cucumber, you really don’t want to do it anymore so you drop to 50%, and you start to refuse the food and you start to throw the food out, which is what economists call an” irrational response”, and it’s irrational because you always need to take what you can get. If I give you $1 and I give you a neighbor $1,000, you’re probably pissed off, but you still need to take the dollar, but you probably won’t because you would be so pissed off that you refuse the money. That’s an irrational response. Back to the experiment, if you give the grape to the partner and the partner doesn’t even need to work for the grape, he gets it for free then you really don’t want to do it anymore. And this shows that effort is a part of the equation. In a recent study on crows they found that effort was even more important than the reward itself.

So, when we first published this we got complaints from people. The media immediately said this was a fairness study. We never used the word fairness, we called it inequity aversion. But they say it’s a fairness study and we got angry emails. One was a guy who wrote that we must be communists, because we were interested in fairness. Now I look at my monkeys as capitalistic. My monkeys work for food and compare their income, but we would get that. We also had a philosopher who wrote that it’s impossible for monkeys to have a sense of fairness because fairness was invented during the French Revolution.

And this shows a little bit that people think that morality is a sort of top down system, and that’s how morality works. The monkey level of fairness, we call first order fairness, which is that you care about getting. You’re basically envious of what somebody else is getting. This is also what young children have. The chimpanzees go a step further when you do this kind of test. The one who gets the grape may refuse the grape till the other one also gets a grape. So chimpanzees care about equalizing outcomes and we play the ultimate end game with chimpanzees, which is the gold standard of the human sense of fairness. If you ask me what is the difference between the chimpanzee’s sense of fairness and that of humans, I don’t know anymore. I’m not sure that there is much difference.

Would you say the desire for fairness is a mark of empathy?

Frans: We think that all these tendencies have to have to do with cooperation. Fairness is sometimes presented as a form of empathy, but I don’t think it is that. I think in cooperative animals you need to watch what you get. If I work with you every day and you take much more than me of the benefits that we together derive, I’m being screwed. And so I need to pay attention to that. I either need to protest against you or I need to look for a different partner.

So fairness has to do with equalizing outcomes so that cooperation runs smoothly, and what chimpanzees do (and humans) is they can anticipate the envy reaction. So chimpanzees think ahead, and they understand that if they create a kind of reaction they undermine their own cooperation, and that’s why they try to equalize outcomes. In the end I think it’s actually beneficial behavior in a cooperative society.

One last thing, of all these behaviors, what do you feel is instinct, and what is learned?

Frans: Everything that animals do and humans do is subject to learning and experience. People sometimes say things like, “Okay, reconciliation is an instinct” or “aggression is an instinct”. The word instinct is not used by people who work with animals anymore. We don’t use it because it has this suggestion of “genetically programmed”, which really things aren’t. So for example, we’ve done experiments where we bring two species together. You have some species that are very conciliatory and easily reconcile after fights and other species like the rhesus monkey who almost never reconciles after a fight. We brought juveniles of the two together and influence the behavior so in this case, the older juveniles were the ones of the very conciliatory species, and the young rhesus monkeys started to learn to reconcile and started to reconcile three times more than a rhesus monkey normally does. So I think the same thing can probably be done with empathy. All these tendencies are inborn tendencies to develop relationships and inborn tendencies to care about others, but they’re all subject to environmental influences, so nothing is sort of written in stone. And I would say the same thing for empathy. Empathy is trainable and can be expanded.

After getting to engage with Frans and some of his wealth of research and insight, we opened the floor to Googlers for Q&A.

Is there some kind of evolutionary cost to being empathic?

Frans: Yeah, because empathy often leads to altruistic behavior, and altruistic behavior is costly. So you don’t want to do that for everyone. We think that empathy started with maternal care, and that would explain why females have more of it and why oxytocin is involved, oxytocin being basically a maternal hormone.

In that context, obviously empathy is very beneficial whether you’re a mouse or an elephant, you need to pay attention to the distress of your offspring when they’re hungry, when they’re cold, you need to pay attention and be sensitive to their distress. In species that have bigger societies, it started to expand to other relationships where it’s not just between mother and offspring but also between let’s say friends who help each other. And so that’s also why males also have empathy, because if you live in a society where you depend on each other, let’s say lionesses who hunt together or male chimpanzees who help each other, you need to care about the other because if you lose the other individual, you lose a partner. And so, empathy started to play a role in the bigger society, but only really between individuals who need each other or depend on each other. You don’t invest in individuals you don’t care about.

But I think it can be expanded. For example we humans can have empathy for a stranded whale. Now I can guarantee you, our empathy did not evolve to take care of stranded whales, and if we are hungry, as people and the whale strands, I’m sure we do very different things than pushing it back into the ocean. So we have conditional empathy, so to speak, that we can apply to other species, and we can apply it to strangers, but that’s all conditional on the circumstances. So the empathy that our model has for our offspring is sort of unconditional because we have to invest in offspring. But as soon as it gets to strangers or other species, it depends on the circumstances. So it’s beautiful that we can have empathy for individuals who are not even related to us and don’t live in our society. We have that capacity. But I do think it’s fragile in the sense that if tomorrow there would be a drought, and we would all go hungry, we would have much less empathy I think for individuals who are far from us.

If empathy and prosocial behavior are inborn tendencies in mammals, where do you suppose that selfishness and greed comes from and why is it that in some instances we see greed outplaying empathy?

Frans: Humans and other primates are by default cooperators within their own ingroup and there are some brain studies now that indicate that the first choice is a cooperative choice and secondarily we start to think about the selfish choices. And I think the same applies to many animals within their groups, but human society has gotten so big, not just with thousands but millions of people, that social distance is disrupted, the social closeness of small-scale societies has been lost in our current societies, and that’s where greed comes in because now I can start to exploit people left and right on the internet or wherever I do that, because I don’t know these people. We live in societies that are largely anonymous, so the greed that you see and the inequity that we see in our current societies are aroused because the face-to-face contact is gone.

When reconciliation is happening, is it a rational decision that I must reconcile so that I can cooperate and get resources?

In biology, we usually think of the benefits of a behavior, but that doesn’t mean that animals are necessarily motivated by the benefits. A lot of the things that we do and the animals do, we do without giving thought to their function. For example, sex serves reproduction and so if you ask a biologist like why do we have sex, we come up with genetic arguments, we come up with rubber duck reproductive arguments, but we don’t think animals know the connection between sexual reproduction. So for them, that is not the motivating part, and actually the reason that we humans have the morning after pill is because we don’t always think of the consequences of sex. I would say the same is true for altruistic behavior. The same is true for reconciliation behavior. We do a lot of things on a sort of emotional reaction basis, and biologists will tell you what the benefits are of it because we think in those terms, the animals themselves do not necessarily [think in those terms].

Are there any studies about developmental periods in which empathy capacities and learning capacities form, physiologically and cognitively? What can we do about that?

Frans: I’m not an expert on humans, but I do know that the age keeps creeping up earlier and earlier. So for example, theory of mind, meaning that you can take the perspective of somebody else, we used to say that kids have that at five years of age. I think many of us now believe that at two years of age they have some of that already. So in human development the opinions keep shifting.

For other species, generally speaking, we first discover something in the chimps because we do work on chimps and chimps are most like us so we know how to test them. But within five years or 10 years the same phenomenon is going to be found in monkeys and dogs and dolphins and certain large-brained birds and I call that the cognitive ripple effect. Things are almost never limited to the primates, and the whole idea that capacities like theory of mind are limited to humans is now being abandoned.

Can you tell us more about the studies that look at the developmental aspects of empathy in individuals?

Frans: In general, people who work on animal behavior aren’t very good at the developmental part. I think many of them ignore the developmental part because they’re so eager to demonstrate a certain capacity in a certain species that they go with the smartest subjects and they test them in the smartest possible way. How it develops and how it builds up, they don’t know.

In reference to this study where you have one chimp choosing a token, and that’s going to determine whether that chimp gets fed and whether the other chimp gets fed, you had a breakdown of two different kinds of pleading that happen, where one is more attention grabbing and the other is kind of being pushy. Is that a sharp distinction, are there other kinds of display behavior that are involved? And are there other kinds of background causes, for example, social status in regards to the relationship that will determine whether the chimp succeeds?

Frans: The behavioral part, it’s really your protester you don’t protest, and I’m not sure that the protesting really helps us. As you saw in the data, it actually seems not to help in the sense that the partner is actually less willing to share with someone who puts pressure on them.

The relational part we investigated, and we actually found in both the monkey studies and the chimpanzee studies, that the most generous individuals are the most dominant individuals and the highest-ranking individuals, the top male/female in the group, are by far the most generous. This raises a very interesting chicken and egg question. Are they dominant because they are such generous individuals, which is possible, given the structure of their societies, or is it that once you have reached a position you can be generous with everybody? There was for example, a chimpanzee male in the wild who stayed in power for 12 years as an alpha male, and he was extremely generous and he had a whole bribery system down in the sense that he would appropriate meat that other chimps had hunted, and he would hand it out to his friends and supporters and not to individuals who did not support him. And so this is an indication that it’s basically used as a sort of political tool. Generosity can be used as a political tool.

Parents have expressed that they want AI like Siri, Alexa and Google Assistant to respond to children politely. They basically want the kids to say please and thank you. And the prosocial experiment seems to suggest that chimps are punishing other chimps when they bang on the walls and use pressurizing behavior to get somebody to do something. Is that punishing behavior a contributing element to enforce that non-banking pressuring behavior? And should we really implement that so we don’t have a being that allows people to just have pressuring behavior?

Frans: I’ve done tons of studies on spontaneous food sharing in chimpanzees. The juvenile chimps are like children. They’re very impolite, and they try to steal things, and if they’re very young, they get away with it. If they are two or three years old no one’s going to bother with them. When they get older, [the older chimps] start to punish them or they start to withhold food from them if they behave like that. And I do think that the civility that we see among the adults is taught this way. I don’t think it’s an intentional teaching process necessarily, but the young ones learn that if they behave in a particular way and they have reached a certain age, they won’t get away with it. And so in a sense there is this aspect of training them to behave.

How far back in our evolution does empathy and cooperation go? Does it go into marsupials or how far beyond that does it go?

Frans: I used to say it’s a mammalian characteristic, and so I already felt extremely generous towards the rodents, but I have friends who study fish and friends who study birds and are now involved in an empathy study on fish. There are certain fish who love body contact so for example you have cleaner fish who clean a big fish. And if the big fish gets upset because the cleaning can be rough, they give them a massage. The cleaner fish, which is a little fish, goes around to big fish, and the big fish calms down if he has some body contact. So, there are indications that body contact is important in some fish at least.

So I’m now involved in repeating the vole study on cleaner fish. For birds we definitely have evidence in certain species that some sort of empathy mechanism is involved. So I don’t know if it relates a bit to what I said before that usually we discover things first in the chimpanzee and then very soon afterwards we find it in other species and many of these things are much older than we think. So, for example, the oxytocin vasopressin system is much older than just mammals. It’s a very old system.

Is the evolution convergent? Did it evolve multiple times, or do we think it started in one place?

Frans: Many things evolve multiple times. Recently there was a study about the appendix, which we long thought was useless. The appendix has evolved 26 times independently in different animals, and so this is used as an argument to say that the appendix is probably very important. Another example is tool use in apes, tool use in capuchin monkeys, tool use in crows, these are probably independent evolutions. That doesn’t mean there aren’t some basic mechanisms involved that are shared across the board, and that’s why I’m saying the oxytocin vasopressin system is very ancient. It goes back to the mollusks even, and it’s probably involved in all these so-called independent empathies that we see in different branches, and so that whole distinction that we used to make between homology and analogy is sort of breaking apart, with the current knowledge I would say. Things are never fully independent.

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