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The Evolution of Empathy

(Editor’s Note: we are pleased to bring you this arti­cle thanks to our col­lab­o­ra­tion with Greater Good Mag­a­zine).

The Evo­lu­tion of Empathy

Empathy’s not a uniquely human trait, explains pri­ma­tol­o­gist Frans de Waal. Apes and other ani­mals feel it as well, sug­gest­ing that empa­thy is truly an essen­tial part of who we are.

Once upon a time, the United States had a pres­i­dent known for a pecu­liar facial dis­play. In an act of con­trolled emo­tion, he would bite his lower lip and tell his audi­ence, “I feel your pain.” Whether the dis­play was sin­cere is not the issue here; how we are affected by another’s predica­ment is. Empa­thy is sec­ond nature to us, so much so that any­one devoid of it strikes us as dan­ger­ous or men­tally ill.

At the movies, we can’t help but get inside the skin of the char­ac­ters on the screen. We despair when their gigan­tic ship sinks; we exult when they finally stare into the eyes of a long-lost lover.

We are so used to empa­thy that we take it for granted, yet it is essen­tial to human soci­ety as we know it. Our moral­ity depends on it: How could any­one be expected to fol­low the golden rule with­out the capac­ity to men­tally trade places with a fel­low human being? It is log­i­cal to assume that this capac­ity came first, giv­ing rise to the golden rule itself. The act of perspective-taking is summed up by one of the most endur­ing def­i­n­i­tions of empa­thy that we have, for­mu­lated by Adam Smith as “chang­ing places in fancy with the sufferer.”

Even Smith, the father of eco­nom­ics, best known for empha­siz­ing self-interest as the lifeblood of human econ­omy, under­stood that the con­cepts of self-interest and empa­thy don’t con­flict. Empa­thy makes us reach out to oth­ers, first just emo­tion­ally, but later in life also by under­stand­ing their situation.

This capac­ity likely evolved because it served our ances­tors’ sur­vival in two ways. First, like every mam­mal, we need to be sen­si­tive to the needs of our off­spring. Sec­ond, our species depends on coop­er­a­tion, which means that we do bet­ter if we are sur­rounded by healthy, capa­ble group mates. Tak­ing care of them is just a mat­ter of enlight­ened self-interest.

Ani­mal empathy

It is hard to imag­ine that empathy—a char­ac­ter­is­tic so basic to the human species that it emerges early in life, and is accom­pa­nied by strong phys­i­o­log­i­cal reactions—came into exis­tence only when our lin­eage split off from that of the apes. It must be far older than that. Exam­ples of empa­thy in other ani­mals would sug­gest a long evo­lu­tion­ary his­tory to this capac­ity in humans.

Evo­lu­tion rarely throws any­thing out. Instead, struc­tures are trans­formed, mod­i­fied, co-opted for other func­tions, or tweaked in another direc­tion. The frontal fins of fish became the front limbs of land ani­mals, which over time turned into hoofs, paws, wings, and hands. Occa­sion­ally, a struc­ture loses all func­tion and becomes super­flu­ous, but this is a grad­ual process, and traits rarely dis­ap­pear alto­gether. Thus, we find tiny ves­tiges of leg bones under the skin of whales and rem­nants of a pelvis in snakes.

Over the last sev­eral decades, we’ve seen increas­ing evi­dence of empa­thy in other species. One piece of evi­dence came unin­ten­tion­ally out of a study on human devel­op­ment. Car­olyn Zahn-Waxler, a research psy­chol­o­gist at the National Insti­tute of Men­tal Health, vis­ited people’s homes to find out how young chil­dren respond to fam­ily mem­bers’ emo­tions. She instructed peo­ple to pre­tend to sob, cry, or choke, and found that some house­hold pets seemed as wor­ried as the chil­dren were by the feigned dis­tress of the fam­ily mem­bers. The pets hov­ered nearby and put their heads in their own­ers’ laps.

But per­haps the most com­pelling evi­dence for the strength of ani­mal empa­thy came from a group of psy­chi­a­trists led by Jules Masser­man at North­west­ern Uni­ver­sity. The researchers reported in 1964 in the Amer­i­can Jour­nal of Psy­chi­a­try that rhe­sus mon­keys refused to pull a chain that deliv­ered food to them­selves if doing so gave a shock to a com­pan­ion. One mon­key stopped pulling the chain for 12 days after wit­ness­ing another mon­key receive a shock. Those pri­mates were lit­er­ally starv­ing them­selves to avoid shock­ing another animal.

The anthro­poid apes, our clos­est rel­a­tives, are even more remark­able. In 1925, Robert Yerkes reported how his bonobo, Prince Chim, was so extra­or­di­nar­ily con­cerned and pro­tec­tive toward his sickly chim­panzee com­pan­ion, Panzee, that the sci­en­tific estab­lish­ment might not accept his claims: “If I were to tell of his altru­is­tic and obvi­ously sym­pa­thetic behav­ior towards Panzee, I should be sus­pected of ide­al­iz­ing an ape.”

Nadia Ladygina-Kohts, a pri­ma­to­log­i­cal pio­neer, noticed sim­i­lar empathic ten­den­cies in her young chim­panzee, Joni, whom she raised at the begin­ning of the last cen­tury, in Moscow. Kohts, who ana­lyzed Joni’s behav­ior in the minut­est detail, dis­cov­ered that the only way to get him off the roof of her house after an escape—much more effec­tive than any reward or threat of punishment—was by arous­ing sympathy:

If I pre­tend to be cry­ing, close my eyes and weep, Joni imme­di­ately stops his plays or any other activ­i­ties, quickly runs over to me, all excited and shagged, from the most remote places in the house, such as the roof or the ceil­ing of his cage, from where I could not drive him down despite my per­sis­tent calls and entreaties. He hastily runs around me, as if look­ing for the offender; look­ing at my face, he ten­derly takes my chin in his palm, lightly touches my face with his fin­ger, as though try­ing to under­stand what is hap­pen­ing, and turns around, clench­ing his toes into firm fists.

These obser­va­tions sug­gest that apart from emo­tional con­nect­ed­ness, apes have an appre­ci­a­tion of the other’s sit­u­a­tion and show a degree of perspective-taking. One strik­ing report in this regard con­cerns a bonobo female named Kuni, who found a wounded bird in her enclo­sure at Twycross Zoo, in Eng­land. Kuni picked up the bird, and when her keeper urged her to let it go, she climbed to the high­est point of the high­est tree, care­fully unfolded the bird’s wings and spread them wide open, one wing in each hand, before throw­ing it as hard as she could toward the bar­rier of the enclo­sure. When the bird fell short, Kuni climbed down and guarded it until the end of the day, when it flew to safety. Obvi­ously, what Kuni did would have been inap­pro­pri­ate toward a mem­ber of her own species. Hav­ing seen birds in flight many times, she seemed to have a notion of what would be good for a bird, thus giv­ing us an anthro­poid illus­tra­tion of Smith’s “chang­ing places in fancy.”

This is not to say that all we have are anec­dotes. Sys­tem­atic stud­ies have been con­ducted on so-called “con­so­la­tion” behav­ior. Con­so­la­tion is defined as friendly or reas­sur­ing behav­ior by a bystander toward a vic­tim of aggres­sion. For exam­ple, chim­panzee A attacks chim­panzee B, after which bystander C comes over and embraces or grooms B. Based on hun­dreds of such obser­va­tions, we know that con­so­la­tion occurs reg­u­larly and exceeds base­line lev­els of con­tact. In other words, it is a demon­stra­ble ten­dency that prob­a­bly reflects empa­thy, since the objec­tive of the con­soler seems to be to alle­vi­ate the dis­tress of the other. In fact, the usual effect of this kind of behav­ior is that it stops scream­ing, yelp­ing, and other signs of distress.

A bottom-up view of empathy

The above exam­ples help explain why to the biol­o­gist, a Russ­ian doll is such a sat­is­fy­ing play­thing, espe­cially if it has a his­tor­i­cal dimen­sion. I own a doll of Russ­ian Pres­i­dent Vladimir Putin, within whom we dis­cover Yeltsin, Gor­bachev, Brezh­nev, Kruschev, Stalin, and Lenin, in that order. Find­ing a lit­tle Lenin and Stalin within Putin will hardly sur­prise most polit­i­cal ana­lysts. The same is true for bio­log­i­cal traits: The old always remains present in the new.

This is rel­e­vant to the debate about the ori­gins of empa­thy, espe­cially because of the ten­dency in some dis­ci­plines, such as psy­chol­ogy, to put human capac­i­ties on a pedestal. They essen­tially adopt a top-down approach that empha­sizes the unique­ness of human lan­guage, con­scious­ness, and cog­ni­tion. But instead of try­ing to place empa­thy in the upper regions of human cog­ni­tion, it is prob­a­bly best to start out exam­in­ing the sim­plest pos­si­ble processes, some per­haps even at the cel­lu­lar level. In fact, recent neu­ro­science research sug­gests that very basic processes do under­lie empa­thy. Researchers at the Uni­ver­sity of Parma, in Italy, were the first to report that mon­keys have spe­cial brain cells that become active not only if the mon­key grasps an object with its hand but also if it merely watches another do the same. Since these cells are acti­vated as much by doing as by see­ing some­one else do, they are known as mir­ror neu­rons, or “mon­key see, mon­key do” neurons.

It seems that devel­op­men­tally and evo­lu­tion­ar­ily, advanced forms of empa­thy are pre­ceded by and grow out of more ele­men­tary ones. Biol­o­gists pre­fer such bottom-up accounts. They always assume con­ti­nu­ity between past and present, child and adult, human and ani­mal, even between humans and the most prim­i­tive mammals.

So, how and why would this trait have evolved in humans and other species? Empa­thy prob­a­bly evolved in the con­text of the parental care that char­ac­ter­izes all mam­mals. Sig­nal­ing their state through smil­ing and cry­ing, human infants urge their care­giver to take action. This also applies to other pri­mates. The sur­vival value of these inter­ac­tions is evi­dent from the case of a deaf female chim­panzee I have known named Krom, who gave birth to a suc­ces­sion of infants and had intense pos­i­tive inter­est in them. But because she was deaf, she wouldn’t even notice her babies’ calls of dis­tress if she sat down on them. Krom’s case illus­trates that with­out the proper mech­a­nism for under­stand­ing and respond­ing to a child’s needs, a species will not survive.

Dur­ing the 180 mil­lion years of mam­malian evo­lu­tion, females who responded to their offspring’s needs out-reproduced those who were cold and dis­tant. Hav­ing descended from a long line of moth­ers who nursed, fed, cleaned, car­ried, com­forted, and defended their young, we should not be sur­prised by gen­der dif­fer­ences in human empa­thy, such as those pro­posed to explain the dis­pro­por­tion­ate rate of boys affected by autism, which is marked by a lack of social com­mu­ni­ca­tion skills.

Empa­thy also plays a role in coop­er­a­tion. One needs to pay close atten­tion to the activ­i­ties and goals of oth­ers to coop­er­ate effec­tively. A lioness needs to notice quickly when other lionesses go into hunt­ing mode, so that she can join them and con­tribute to the pride’s suc­cess. A male chim­panzee needs to pay atten­tion to his buddy’s rival­ries and skir­mishes with oth­ers so that he can help out when­ever needed, thus ensur­ing the polit­i­cal suc­cess of their part­ner­ship. Effec­tive coop­er­a­tion requires being exquis­itely in tune with the emo­tional states and goals of others.

Within a bottom-up frame­work, the focus is not so much on the high­est lev­els of empa­thy, but rather on its sim­plest forms, and how these com­bine with increased cog­ni­tion to pro­duce more com­plex forms of empa­thy. How did this trans­for­ma­tion take place? The evo­lu­tion of empa­thy runs from shared emo­tions and inten­tions between indi­vid­u­als to a greater self/other distinction—that is, an “unblur­ring” of the lines between indi­vid­u­als. As a result, one’s own expe­ri­ence is dis­tin­guished from that of another per­son, even though at the same time we are vic­ar­i­ously affected by the other’s. This process cul­mi­nates in a cog­ni­tive appraisal of the other’s behav­ior and sit­u­a­tion: We adopt the other’s perspective.

As in a Russ­ian doll, how­ever, the outer lay­ers always con­tain an inner core. Instead of evo­lu­tion hav­ing replaced sim­pler forms of empa­thy with more advanced ones, the lat­ter are merely elab­o­ra­tions on the for­mer and remain depen­dent on them. This also means that empa­thy comes nat­u­rally to us. It is not some­thing we only learn later in life, or that is cul­tur­ally con­structed. At heart, it is a hard-wired response that we fine-tune and elab­o­rate upon in the course of our lives, until it reaches a level at which it becomes such a com­plex response that it is hard to rec­og­nize its ori­gin in sim­pler responses, such as body mim­icry and emo­tional contagion.

On a leash

Biol­ogy holds us “on a leash,” in the felic­i­tous words of biol­o­gist Edward Wil­son, and will let us stray only so far from who we are. We can design our life any way we want, but whether we will thrive depends on how well that life fits human predispositions.

I hes­i­tate to pre­dict what we humans can and can’t do, but we must con­sider our bio­log­i­cal leash when decid­ing what kind of soci­ety we want to build, espe­cially when it comes to goals like achiev­ing uni­ver­sal human rights.

If we could man­age to see peo­ple on other con­ti­nents as part of us, draw­ing them into our cir­cle of reci­procity and empa­thy, we would be build­ing upon, rather than going against, our nature.

For instance, in 2004, the Israeli Min­is­ter of Jus­tice caused polit­i­cal uproar for sym­pa­thiz­ing with the enemy. Yosef Lapid ques­tioned the Israeli army’s plans to demol­ish thou­sands of Pales­tin­ian homes in a zone along the Egypt­ian bor­der. He had been touched by images on the evening news. “When I saw a pic­ture on the TV of an old woman on all fours in the ruins of her home look­ing under some floor tiles for her med­i­cines, I did think, ‘What would I say if it were my grand­mother?’” he said. Lapid’s grand­mother was a Holo­caust victim.

This inci­dent shows how a sim­ple emo­tion can widen the def­i­n­i­tion of one’s group. Lapid had sud­denly real­ized that Pales­tini­ans were part of his cir­cle of con­cern, too. Empa­thy is the one weapon in the human reper­toire that can rid us of the curse of xenophobia.

Empa­thy is frag­ile, though. Among our close ani­mal rel­a­tives, it is switched on by events within their com­mu­nity, such as a young­ster in dis­tress, but it is just as eas­ily switched off with regards to out­siders or mem­bers of other species, such as prey. The way a chim­panzee bashes in the skull of a live mon­key by hit­ting it against a tree trunk is no adver­tise­ment for ape empa­thy. Bono­bos are less bru­tal, but in their case, too, empa­thy needs to pass through sev­eral fil­ters before it will be expressed. Often, the fil­ters pre­vent expres­sions of empa­thy because no ape can afford feel­ing pity for all liv­ing things all the time. This applies equally to humans. Our evo­lu­tion­ary back­ground makes it hard to iden­tify with out­siders. We’ve evolved to hate our ene­mies, to ignore peo­ple we barely know, and to dis­trust any­body who doesn’t look like us. Even if we are largely coop­er­a­tive within our com­mu­ni­ties, we become almost a dif­fer­ent ani­mal in our treat­ment of strangers.

This is the chal­lenge of our time: glob­al­iza­tion by a tribal species. In try­ing to struc­ture the world such that it suits human nature, the point to keep in mind is that polit­i­cal ide­o­logues by def­i­n­i­tion hold nar­row views. They are blind to what they don’t wish to see. The pos­si­bil­ity that empa­thy is part of our pri­mate her­itage ought to make us happy, but we are not in the habit of embrac­ing our nature. When peo­ple kill each other, we call them “ani­mals.” But when they give to the poor, we praise them for being “humane.” We like to claim the lat­ter ten­dency for our­selves. Yet, it will be hard to come up with any­thing we like about our­selves that is not part of our evo­lu­tion­ary back­ground. What we need, there­fore, is a vision of human nature that encom­passes all of our ten­den­cies: the good, the bad, and the ugly.

Our best hope for tran­scend­ing tribal dif­fer­ences is based on the moral emo­tions, because emo­tions defy ide­ol­ogy. In prin­ci­ple, empa­thy can over­ride every rule about how to treat oth­ers. When Oskar Schindler kept Jews out of con­cen­tra­tion camps dur­ing World War II, for exam­ple, he was under clear orders by his soci­ety on how to treat peo­ple, yet his feel­ings interfered.

Car­ing emo­tions may lead to sub­ver­sive acts, such as the case of a prison guard who dur­ing wartime was directed to feed his charges only water and bread, but who occa­sion­ally sneaked in a hard-boiled egg. How­ever small his ges­ture, it etched itself into the pris­on­ers’ mem­o­ries as a sign that not all of their ene­mies were mon­sters. And then there are the many acts of omis­sion, such as when sol­diers could have killed cap­tives with­out neg­a­tive reper­cus­sions but decided not to. In war, restraint can be a form of compassion.

Emo­tions trump rules. This is why, when speak­ing of moral role mod­els, we talk of their hearts, not their brains (even if, as any neu­ro­sci­en­tist will point out, the heart as the seat of emo­tions is an out­dated notion). We rely more on what we feel than what we think when solv­ing moral dilemmas.

It’s not that reli­gion and cul­ture don’t have a role to play, but the build­ing blocks of moral­ity clearly pre­date human­ity. We rec­og­nize them in our pri­mate rel­a­tives, with empa­thy being most con­spic­u­ous in the bonobo ape and reci­procity in the chim­panzee. Moral rules tell us when and how to apply our empathic ten­den­cies, but the ten­den­cies them­selves have been in exis­tence since time immemorial.

Frans B. M. de Waal, Ph.D., a Dutch-born pri­ma­tol­o­gist, is the C. H. Can­dler Pro­fes­sor at Emory Uni­ver­sity and direc­tor of the Liv­ing Links Cen­ter at the Yerkes National Pri­mate Research Cen­ter in Atlanta. This essay is adapted from his book, Our Inner Ape: A Lead­ing Pri­ma­tol­o­gist Explains Why We Are Who We Are. His lat­est book is The Age of Empa­thy. Greater Good Mag­a­zine, based at UC-Berkeley, is a quar­terly mag­a­zine that high­lights ground break­ing sci­en­tific research into the roots of com­pas­sion and altruism.

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