young - the social brain and the myth of empathy

Science in Context 25(3), 401424 (2012). Copyright C Cambridge University Pressdoi:10.1017/S0269889712000129

The Social Brain and the Myth of Empathy

Allan Young

McGill UniversityE-mail: [email protected]

Argument

Neuroscience research has created multiple versions of the human brain. The social brain isone version and it is the subject of this paper. Most image-based research in the field of socialneuroscience is task-driven: the brain is asked to respond to a cognitive (perceptual) stimulus.The tasks are derived from theories, operational models, and back-stories now circulating insocial neuroscience. The social brain comes with a distinctive back-story, an evolutionary historyorganized around three, interconnected themes: mind-reading, empathy, and the emergence ofself-consciousness. This paper focuses on how empathy has been incorporated into the socialbrain and redefined via parallel research streams, employing a shared, imaging technology.The concluding section describes how these developments can be understood as signaling theemergence of a new version of human nature and the unconscious. My argument is not thatempathy in the social brain is a myth, but rather that it is served by a myth consonant with thecanons of science.

Introduction

Empathy is today a popular subject among social neuroscientists, science journalists,and the consumers of this literature the curious public, social and behavioral scientists,ethicists, forensic psychiatrists, moral philosophers, other humanities scholars, cognitivepsychologists, and investigators working on psychopathy, autism spectrum disorder, andother developmental pathologies. In this paper, I describe the social neuroscience ofempathy and its object of inquiry, the social brain, from an ethnographic perspective. Thesocial brain is indissociable from the anatomical brain but likewise something differentfrom other scientific visions of the brain. It is called a social brain because of what itdoes (it permits minds and brains to read intentions and share feelings inside otherminds and brains), how it emerged (it is a product of hominid social evolution), andhow these presumptions have shaped research and discourse.

For three centuries, human nature and inter-subjectivity were similarly associatedwith the faculty of reasoning. The golden age of social anthropology, arguably the lastof the Enlightenment sciences, was pre-occupied with questions about the natureand universality of reason. Were mentalities and modes of reasoning around theworld essentially the same or different? Do human societies occupy positions along

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a single developmental continuum, whose bedrock is the psychic unity of mankind?Are magic and science cognate efforts to understand and manipulate the materialworld? These well-known debates, which included Frazer, Malinowski, Freud, andEvans-Pritchard, focused on mentalities and cultures, and, except for Freud, rarelyon biology and evolution (Tambiah 1990, 2). In the anglophone world at least,human nature is increasingly understood with reference to empathy, brain science,and evolutionary biology, rather than reason, and without recourse any longer to socialanthropology.

These developments include a widespread public misunderstanding. Neuroscienceknowledge about empathy and the social brain derives from two sources: research onmirror neurons and research based on the same technology but in combination witha neo-Darwinian back-story. The two streams are complementary and the boundariesare fuzzy. The misunderstanding is that the capacities and significance of mirror neuronsare grossly exaggerated in popular acounts, while the back-story and associated brainresearch, the focus of much interest among neuroscientists, are poorly understood orignored. In part one of this paper, I discuss research on mirror neurons, their popularappeal, and some reasons for the declining interest in them among neuroscientists. Parttwo is about the back-story and related research. The two parts run chronologically inparallel; my emphasis throughout is on empathy.

Part One: Mirror Neurons

Mirror neurons were initially discovered in premotor and inferior parietal regions ofrhesus monkey brains (Di Pellegrino et al. 1992). Evidence of human mirror neuronswas discovered soon afterward in homologous regions. Most neurons in these brainregions perform a single function: they communicate sensory information to the brainor transmit motor commands from the brain. Mirror neurons combine these functions:an individual observes goal directed behavior being performed by a second individual;the motor activation pattern in the observers brain mirrors (matches) the pattern inthe performers brain; the process is not conscious and the motor behavior is notperformed. Mirror neurons are multimodal: they respond to visual stimuli, auditorystimuli (e.g. the sound of a peanut shell being cracked open), imagined events (e.g.an athletic performance), and texts containing action verbs (Fadiga et al. 1996; Fogassiet al. 2002 and 2005; Kohler et al. 2002; Rizzolatti and Arbib 1998; Rizzolatti andCraighero 2004; and Tettamanti et al. 2005).

Mirror neuron research on monkey brains is based on an invasive procedure:microelectrodes are inserted into the brain at selected points. For ethical reasons,this technique was not permissible for human studies. Consequently, human researchwas based on non-invasive techniques, fMRI and PET. These technologies can imagethe activation of populations of neurons but not single neurons. The problem with thistechnology is that the existence of mirror neurons can be inferred but not demonstrated.





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The original conception was that every mirror neuron is strictly congruent withan identifiable goal-oriented action. The current conception divides mirror neuronsinto four sub-populations: strictly congruent neurons; broadly congruent neurons (thecorrespondence is not precise); canonical neurons (these fire when someone observeshimself performing a relevant action or observes an object to which this action mightbe directed, but not when he observes someone else performing the same action);and anti-mirror neurons (which inhibit the performance of mirrored action patterns)(Keysers et al. 2010).

Evidence of human mirror neurons is necessarily indirect and contested. Criticshave argued that the mirror effect that is observed via neuroimaging is the productof three sub-populations (sensory-motor, sensory only, motor only), no one of whichcorresponds to the original idea of mirror neurons (Dinstein 2008; Dinstein et al.2007; Dinstein et al. 2008; Jacob 2008; Jacob and Jeannerod 2005; Singer 2006;Turella et al. 2009). The very existence of human sensory-motor neurons has beenquestioned. But a recent and unprecedented single-neuron investigation (Mukamelet al. 2010), in which microelectrodes were inserted into the brains of surgical patientsdiagnosed with intractable epilepsy, is said to finally provide direct electrophysiologicalevidence that humans have mirror neurons (Keysers and Gazzola 2010,R353).

The mirror phenomenon is a brain-to-brain product, unmediated by mental states,and different from situations where a match between brains (observer and performer)occurs in parallel and is routed through consciousness. Here is an example: Participantsin this experiment were inserted into an fMRI scanner, where they viewed photos ofhands and feet in painful situations. They were next asked to assess the level of pain beingexperienced by the anonymous individuals. Participants assessments and brain imageswere compared. High assessments correlated with high activations in brain regionsknown (from previous research) to play a significant role in pain processing (Jacksonet al. 2005, 771). In other words, the distinctiveness of the mirror phenomenon dependson more than demonstrating neural matching: it is necessary to likewise demonstratethe absence of mental processing. To understand the efforts of researchers to connectmirror neurons to empathy, it is important to carefully distinguish between mirrorstates and parallel states. There are social neuroscientists who insist on this distinction(e.g. Singer et al. 2004), but others who do not. For convenience, I will call the formerthe maximalists.

According to the maximalists, since the frontal and parietal regions that arecommonly associated with mirror neurons are connected to multiple brain regions,it can be presumed that mirror neurons extend to these regions as well (Keysers andGazzola 2010, R353). The relevant research has focused on the role of mirror neuronsin the social emotions, notably disgust. In the next paragraphs, I summarize the workpublished on this subject by maximalists (Vittoriao Gallese, Marco Iacoboni, GiacomoRizzolatti, Christian Keysers and colleagues) during the period of most intense interest,between 2002 and 2007.




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Mirror neurons and disgust

[T]he fundamental mechanism at the basis of the experiential understanding ofothers actions is the activation of the mirror neuron system. A similar mechanism,but involving the activation of viscero-motor centers, underlies the experientialunderstanding of the emotions of others. Evidence obtained from multiplesources (electrical stimulation of monkey brains, task-driven fMRI research, clinicalobservations) indicates that these neural populations intersect in the insula. The insulacontains neural populations active both when the participants directly experienceddisgust and when they understood it through the facial expression of others. It isa center of viscero-motor integration, and a relay between action representations(mirror neurons) and emotion (Gallese et al. 2004, 396, 398400).

This large-scale network, composed of the mirror neuron system, the insula, andlimbic structures including the amygdala, would permit an observer to empathizewith others through the representation and inner imitation of the actions (facialexpressions, body postures) of others (Iacoboni and Dapretto 2006, 942; also Carret al. 2003). (This inner imitation or action representation is synonymous withthe activation pattern that I have already mentioned.)

The process is demonstrated in an fMRI study in which participants inhaledodorants producing a strong feeling of disgust. The same participants observed videoclips showing the emotional facial expression of disgust. Observing such faces andfeeling of disgust activated the same sites in the anterior insula. . . . [Just] as observinghand actions activates the observers motor representation of that action, observingan emotion activates the neural representation of that emotion. Activation wasmore intense when participants imitated the facial emotion than when they observedit. An analogous effect is observed in monkey and mirror neuron studies of goaldirected behavior (Wicker et al. 2003, 655; also van der Gaag et al. 2007). In aninterview in the New York Times, Christian Keysers explained that this neural networkunderpins additional empathic social emotions his list includes guilt, shame, pride,embarrassment, humiliation, rejection, and lust and also empathic responses to pain(Blakeslee 2006).

These researchers (Keysers, Kass, and Gazzola 2010) welcomed the publicationof Mukamel et al.s single-neuron research, since it confirmed the existence ofhuman mirror neurons and, more specifically, located them in regions beyond thefrontal (premotor) and parietal regions of the brain, i.e. supplementary motorarea, and hippocampus and environs. While these findings are consonant with themaximalist vision of mirror neurons, there is only indirect evidence connecting mirrorneurons to the insula and, through this interface, connecting mirror neurons into alarge-scale network and social emotions. The reasoning is illustrated in an fMRIstudy (Carr et al. 2003) in which participants are asked to both imitate (a motoractivity) and observe (a visuo-sensory experience) emotional displays. These are thefindings:


1. Imitating and observing the same emotion are shown to activate the same premotorareas. This finding is necessary, but not sufficient, for showing the presence ofsensory-motor neurons and, more specifically, mirror neurons.

2. Imitating an emotion is shown to activate additional brain areas, insula andamygdala, that are relevant to (unconscious) action representations. This findingis necessary, but not sufficient, for showing the modulation of the actionrepresentation circuit onto limbic activity.

3. In other words: sensory stimulus activation of mirror neurons actionrepresentation insula interface limbic activation (includes the amygdala).

The lynchpin in this operation is the presence of action representations (Fadiga andCraighero 2004; Shmuelof and Zohary 2007). I will continue with this point in thefollowing section.

Mirror neurons and empathy

Although there is no standard definition of empathy among social neuroscientists, thereis a general understanding that empathy refers to a state that is shared by an observerand another individual, present or imagined. Empathic states include, singly or incombination, cognitive empathy, emotional empathy, and somatic empathy (mainlypain). Human mirror neurons are routinely depicted as being intrinsically empathic(e.g. Iacoboni and Dapretto 2006; Iacoboni 2009; Rizzolatti and Craighero 2004;Nummenmaa et al. 2008).

Motor theories of cognition date back to the nineteenth-century (James 1890, onideomotor theory); mirror neurons provide the latest chapter in this history (Jeannerod1994; Hickcok 2009). Until recently, interest in empathy has concentrated in thefields of developmental psychology, social psychology, counseling psychology, and inclinical investigations of certain psychiatric disorders, notably childhood autism andpersonality disorders. Biological research on empathy focused on limbic system andneuroendocrine responses to social signals such as expressed emotion (see Brothers1989 for a review). The discovery of the human mirror neuron system has expandedthe neurological frontiers of empathy by identifying a biological mechanism that appearsto illuminate, if not answer, one version of an enduring philosophical question, theproblem of other minds. How do minds infer intentions embedded in other minds(Custers and Aarts 2010, 49)?

The simplest form of mirror neuron activation (and action representation) is referredto as resonance and it is unconscious. The human mirror neuron system is capableof de-coupling an action representation so that it can be projected (attributed) backto its source, the performer, thereby differentiating between self and other. Recentresearch suggests that sometimes de-coupling is spontaneously reversed during memoryreconsolidation (retrieval). This occurs in instances where an observer remembers







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someone elses performance (via an action representation) as having been his ownexperience a phenomenon that is conventionally called source amnesia and falsememory (Lindner et al. 2010).

The mirror neurons and action representation are sufficient for inferring theperformers intention. I put it this way failing to indicate who or what is doingthe inferring to make a point. Many theorists presume that mirror neurons possessthis capacity: that is, they can interpret action representations and detect intentionswithout cognitive processing by other parts of the brain. It is possible that certainnon-human species are capable of doing this, but there is much disagreement onthis point. It is clear that only humans are capable of the final stage, in which de-coupled representations can be objectified, i.e. detached from their source (performersand original performances) and stored for future use as templates and prototypesfor imitation, emulation, analogical reasoning, and sophisticated forms of mind-reading. While language obviously facilitates objectification, researchers are dividedas to whether it was essential for the emergence of this capacity, since pre-linguisticbabies are demonstrably capable of performing these functions.

The final process, the basis of cognitive empathy, operates with different levels ofaction representation, from the motor intention that drives a given chain of motor actsto the propositional attitudes (beliefs, desires and so on) that . . . explain the observedbehaviour in terms of its plausible psychological reasons. The end-product is anover-arching mentalizing network that connects the mirror neuron system whoseaction representations provide a subpersonally instantiated common space betweenobserver and performer and a scaffold for bootstrapping mutual intelligibility (Galleseand Goldman 1998; Gallese 2001 and 2003) to high-level operations (reasoning incortical areas without demonstrable mirror properties (Corrado 2010, 264; also Kilnerand Frith 2008). Evidence connecting mirror neurons to the putative network andother brain regions is, once again, indirect, e.g. bolstered by efforts to discovermirror neuron abnormalities in people diagnosed with autism spectrum disorders, adiagnosis associated with poor mind-reading abilities and other deficits in cognitiveempathy (Hadjikhani et al. 2006; Oberman et al. 2005; Rizzolatti and Fabbri-Destro2010).

Controversy

There are obvious similarities between human mirror neurons, as I have describedthem, and the mental modules delineated by Jerry Fodor (1983) and later elaboratedby evolutionary psychologists (Barkow et al. 1992). The modules and neuronsrespond spontaneously to domain-specific sensory inputs that require no cognitiveprocessing. Their operations are depicted as automatic (obligatory), rapid (unimpededby consciousness), and encapsulated (invisible to reflective consciousness). Eachis associated with a dedicated neural architecture and a characteristic pattern of







decomposition (pathology). And they are both products of evolution and adaptation.Commonality is exemplified by the so-called intentionality detector (a functionrequired for mind reading) that evolutionary psychologists conceive as a mental moduleand neuroscientists attribute to human mirror neurons. The intentionality detector alsohighlights an important difference, affecting the way in which the two perspectivesare connecting the mind to the brain: mental modules are connected to the brainmetaphorically (e.g. via inference engines and mental organs); mirror neuronsare biological mechanisms that uniquely bridge the gap between mind and brain (viaaction representations).

While the discovery of a visible bridge connecting mind to brain and the possibility,perhaps inevitability, of reducing mind to brain, has stimulated intense popularinterest in mirror neurons, these claims have likewise drawn the attention of skepticalresearchers. Skeptics have three concerns. Do mirror neurons exist? (Mukamel et al.2010 has resolved this concern for the time being). If they do, do they possess thislist of module-like features? Is there convincing evidence that mirror neurons canindependently infer intentions?

To answer these questions, one should begin with the lynchpin of mirror neuronoperations, the notion of action representations. The cognitive science view ofthe central nervous system is that it is a dynamic apparatus comprising parts andprocesses that respond adaptively to a dynamic external world, a process facilitatedby a sensory-motor loop. Adaptation is constant and requires the nervous system toconstruct and continually update its representations of the external world (Boden2006, 11781179). Mirror neurons are presumed, by advocates, to be pre-adaptedcomponents, and their action representations are static, unaffected by experience ormemory. During interactions with the external environment observing behavior mirror neurons form logically related chains of representations that, in turn, make itpossible to represent performers intentions (Iacoboni et al. 2005). The process takesplace outside of consciousness, without the interference of mentalizing structures. Thislogic is unlike propositional logic, and Gallese identifies it with the brains forwardmodel architecture. This model is a theoretical system that describes how motoroutput and bodily movement is regulated in vertebrate species. Thus:

When I am going to stretch my arm to grasp a handle in front of me, the resultingpostural perturbation that would follow, causing my body to bend [forward], is canceledby a forward signal sent to the posterior muscles of my leg, which stabilize my standingposture. The muscles . . . contract before my arm is set in motion. The contraction . . .anticipates, predicts the outcome of the programmed action of the arm, [the] perturbation,[thus] preventing it. . . . Neither overt knowledge nor conscious inference is involved.(Gallese 2001, 38)

The job of the system represented in the forward model is to compare itsestablished predictions/representations with internal sensory feedback from the external




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environment (inverse model representations). When a mismatch between predictionsand real-time feedback exceeds a predetermined limit, the forward model readjusts themotor command. Following sufficient cycles, a revised prediction/representation isestablished. The forward model logic thus logical relations among mirror neuronrepresentations has been appropriated from the field of control engineering,where the model is employed to manage dynamic constellations of internal forces,resistances (intrinsic to all mechanical systems), and feedback from the external world(Schwartz 1999; Kosslyn 2005; Boden 2008, 1184; de Vignemont and Haggard 2008)systematically decomposes the non-propositional logic attributed to mirror neurons.

There are prominent neuroscientists who reject this account, on the grounds thatmirror neurons cannot independently infer intentions because they do not possessthe modular properties that make this feat conceivable. Thus according to Heyes andcolleagues, humans are not born with mirror neurons; while human mirror neuronsdo emerge, they are not pre-adapted to anything in particular. Evolution provides uswith motor neurons that become mirror neurons, but it has not selectively establishedlinks between visual and motor neurons coding the same action (Heyes 2010, 790).These neurons form during each individuals development via a familiar Hebbianprocess, the idea that neurons that fire together wire together (Heyes 2010, 789;Hickok 2009). Similarly, it is mistaken to suppose that these neurons are capable ofrepresenting a movement or object without a context. Hume correctly argued thatmeaningful perceptions presuppose an act of interpretation and contextualization, andthis act relies on memory and associations acquired through experience (Brass etal. 2007; Csibra and Gergely 2007). Consider an experiment conducted by Gergelyet al. (2002). Preverbal babies observed a goal-directed action: a seated adult activateda light-box by leaning forward and touching the box with her forehead. At onesession (context), her hands were encumbered and unavailable; at a second session, herhands were free. Two-thirds of the babies imitated the unusual head-down behaviorafter watching the hands-free demonstration; just one-fifth imitated this behavior afterobserving the hands-occupied demonstration. In the first context, babies inferred thatthe head action offered some advantage, since the demonstrator could have turned onthe light in the familiar way, using her hands, if she had wished. In the second context,most babies chose to emulate motor actions in their repertoire (using their hands), afterpresumably concluding that that the head action was not the most rational.

Conclusion to Part One

There is a consensus among social and cognitive neuroscientists that human mirrorneurons exist in one form or another. Opinion is divided on whether mirror neuronsare pre-adapted to recognize (match) and represent specific goal directed actionsand identify performers intentions. One opinion is that mirror neurons performboth functions. An alternative view among neuroscientists is that mirror neurons






exist but are not innate and perform neither function. A third view questions theexistence of mirror neurons, but doubts seem untenable in the light of recent single-neuron research. Mirror neurons are now widely identified with embodied empathy(biologized intersubjectivity). This is one way for fMRI images to represent embodiedempathy. Embodied empathy is also represented in research that shows matching brainactivations in observer and performer, but without the involvement of mirror neurons.The distinction between the two demonstrations between mirrored and parallel kindsof activations is very often blurred in journalistic accounts, leaving the mistakenimpression that mirror neurons are a default mechanism rather than just one possibility.

Part Two: A neo-Darwinian back-story

A narratives back-story is a fictive or notional history that precedes the eventsdescribed in the narrative. An effective back-story makes fictional charactersrecognizable, and creates expectations in the reader that the author can manipulateas the narrative develops. A prologue is an overt back-story; the introduction sectionin science publications performs a similar function. The invocation can be didactic, aswhen a fictional character overtly reflects on his past, or subtle, as when back-storiesunfold in a series of clues and strings of bibliographic citations. The effectiveness of theseclues for creating a sub-text depends partly on the readers background knowledge.

This neo-Darwinian back-story is a plausible genealogy of the social brain. Bythe end of the Paleolithic era, the course of social and biological evolution hasproduced the psychologically modern human and the cognitively demanding kindsof reciprocity that are both source and product of the social brain. This story has been(and continues to be) erected piece-meal: the work of multiple authors, each tacklinga discrete puzzle, with no collective goal in mind. The story is compelling because itis consonant with the available empirical evidence and because the constituent puzzlesand solutions are logically connected. The story is also implicit, in that it is part ofthe collective consciousness of social brain researchers, but there is (as yet) no occasionwhen it is retold in its entirety. Individual investigators are not necessarily familiarwith, or subscribe to, all of the episodes. While individual episodes can be (and are)operationalized through empirical research, the entirety extending perhaps sevenmillion years can only be inferred, and a researcher can justifiably claim that herwork is confined to the topic that currently attracts her attention, e.g. the intersectionof affective empathy and perceived fairness. One might compare this back-story withthe idea of myth described in Paul Veynes book, Did the Greeks Believe in their Myths?An Essay on the Constitutive Imagination:

[N]othing shines in the night of the world. The materiality of things has no naturalluminescence. . . . But the accidents of human history, as erratic and unplanned as thesuccessive hands in a poker game, lead men to shine an endlessly changing lighting on



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their affairs. Only then is the materiality of things reflected in the light. This lighting. . . begins to make a certain world exist. It is a spontaneous creation, the product of animagination. When a lighted clearing appears in this way, it is generally taken for the verytruth, since there is nothing else to see. (Veyne 1988, 125)

Before proceeding to the back-story, I want to clarify what neo-Darwinianmeans in this context of the social brain. Beginning in the 1950s, John MaynardSmith, an evolutionary biologist, pioneered an empiricist approach to natural selectionthat combined the methods and perspectives of population genetics, game theory,and cost-benefit analysis. The approach was the basis for a dialectical understandingof human biological evolution and, we shall see, a back-story for the neurosciencerevolution (Maynard Smith 1979). In an article, Reconciling Marx and Darwin,published shortly before his death in 2004, Maynard Smith recalls Marxs thesis onhuman consciousness thoughts, inferences, perceptions, and desires the view thatconsciousness is determined by the material conditions of social life. Marx mightreasonably claim to be a materialist, Maynard Smith writes, but he was not a reductionist,someone interested in investigating the biological evolution of social life and the brainthat made human consciousness possible in the first place. Marxs vision of humannature as malleable a view shared by many cultural anthropologists during the1950s was unscientific and proved to be tragically flawed and a manifest failurewhen put into action by communist regimes.

The neo-Darwinian back-story for the social brain begins with two puzzles. Itis propelled forward, from the emergence earliest hominids (perhaps five millionyears ago) to the advent and dispersion of psychologically modern humans, by adialectical logic. I will consider each puzzle in turn. The first puzzle concerns altruism.Population geneticists define altruism as behavior that transfers some or all of thealtruists reproductive potential (fitness) to a beneficiary. In the most extreme case, analtruist dies so that a beneficiary might live and reproduce. Altruistic behavior is oftenobserved in animal populations, and presumed to be genetically determined. It is apuzzle because it gives the non-altruist recipients (lacking altruism genes) a reproductiveadvantage. They will eventually outbreed altruists, and altruists and altruism genes willeventually disappear from the population. This does not happen however and this is thepuzzle. W.D. Hamilton solved the puzzle in the 1960s with the theory of kin selection(inclusive fitness). The theory says that altruistic behavior, including self-sacrifice, hasfavorable cost-benefits if the beneficiaries are closely related to the altruists. Whenthis happens, altruism preserves a homogeneous gene pool and a continuing supply ofaltruism genes for future generations.

Hamiltons solution leads to a new puzzle, once biologists discover populationsin which altruistic behavior is common but the altruists and beneficiaries areoften unrelated or too distantly related for Hamiltons solution. Robert Trivers, ananthropologist, solved this puzzle in 1971 by demonstrating that the arrangementworks (benefits are greater than costs) if altruism is reciprocal. The recipient repays the











altruist. Reciprocal behavior is common among non-human primates, but limited toindividuals living in close proximity. It is typically restricted to grooming and, lessoften, food-sharing, and the interval between gift-giving and repayment is brief, a fewseconds in the case of capuchin monkeys and just minutes with chimpanzees. Morecomplex forms of reciprocity would involve dispersed partners, and long intervalsbetween gifts and repayment. In time, multiple local groups would be connected, andextensive social networks and coalitions would develop. But these developments dependon the ability of the individuals to keep track of their exchanges and the reputations ofpotential exchange partners. Thus memory-based reciprocity represented a great leapforward, but was cognitively demanding, and presupposed various developments in thenervous system, including the capacity to inhibit the archaic impulse for immediategratification.

When altruism and reciprocity are limited to a genetically homogeneous population,cheating is impossible. Of course there will be non-reciprocators and their reproductivefitness will benefit from the costs absorbed by the altruists. But the costs and benefitscirculate within a closed system. The term cheater is reserved for geneticallyheterogeneous populations, in which costs and benefits can accumulate withindifferent lineages circumstances that emerge together with memory-based reciprocity.Once again, there is a solution (reciprocity) that creates a problem (cheaters) and apuzzle. Cheaters are inevitable because all organisms are driven to maximize theirreproductive interests and the interests of close kin a neo-Darwinian premise.The cost-benefits of cheating always favor cheaters. All things being equal, cheaters(non-reciprocators) will outbreed the reciprocators; the nascent social networks willcollapse; human social evolution will progress no farther (Nowak and Sigmund 2005;Rosas 2008). Of course the collapse did not occur. This is a puzzle. There is aneo-Darwinian solution, and it leads to a further contradiction. Puzzle solution contradiction puzzle etc.: this is the dialectic that glues the back-storytogether.

In this case, the solution is in two parts. The social fabric is preserved by theevolution of positive sentiments that include friendship, gratitude, and sympathy. Fastforward to the present time: evidence from game-playing experiments, mathematicalmodeling and simulation, and ethnographies of small-scale, pre-industrial communitiesindicate that positive sentiments are insufficient to prevent the collapse. Somethingstronger is needed and the evidence points to a gene-driven impulse to punish cheaters.The solution is efficient except in one respect: it cannot pay for itself. The fitnesscosts of being an enforcer exceed the benefits. Punishment consumes resources (e.g.energy) and can be terminally expensive if cheaters retaliate violently. For this reason,the enforcers behavior is labeled altruistic punishment. There is a further problemor contradiction, in that punishment creates a new class of cheaters, namely friendsand neighbors who are good reciprocators but unwilling to be enforcers. These aresecond order cheaters; they get the cost-free benefits of punishment and eventuallythey should outbreed gene-driven enforcers (Boyd et al. 2003).


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Human nature and empathic cruelty

Punishment can prevent entropy, but why would a rational individual someoneinnately self-interested and able to estimate cost-benefits become an enforcer?Benefits are often hypothetical (scheduled to arrive in the distant future) and indirect(dissuading potential cheaters), and costs are unpredictable (perhaps bringing retaliationby the cheater or his kin). Even when an enforcer gets his fair share, he cannotknow whether this would happen without his intervention. Thus material rewards canprovide only a weak motive for altruistic punishment.

A solution is described in The neural basis of altruistic punishment, publishedin Science (de Qervain et al. 2004). The experiment is bit of ontological theater,based on the trust game. The game comprises interaction between two anonymousparticipants, A and B. Individual A plays the game seven times, facing seven differentBs. Each participant is given 10 money units, convertible into Swiss francs. The ruleis that A can transfer all or none of his units to B. If he gives 10 units to B, they arequadrupled, so that B now has 50 units (40 plus his original 10). B can now send 25units to A (fair behavior), or he can keep the entire 50 for himself (unfair). Ais now given an additional 20 units. If A now wants to punish B, he can do this inthree ways: 1. he can buy penalty points (each point costs A one unit, and subtractstwo units from B); 2. he can reduce Bs sum cost-free (A does not have to pay for thepenalty points); or 3. he can reduce Bs sum symbolically (B is informed of As decisionbut loses no units). Each game is limited to just one formula (1, 2, or 3), designatedby the researchers. There is an additional variable: in some games, a random devicedetermines whether Bs choice will be fair or unfair. A knows the rules obtainingduring each game: that is, whether his option is 1, 2, or 3, and whether Bs choicewas decided by the random device. Once A learns Bs choice, he has one minute tomake a decision whether to punish, how much to punish. During this one-minuteinterval, his brain is scanned via PET. As brain images reflect his deliberations and hisanticipation (imagining) of Bs response.

Thus the experiment reproduces moments in the evolutionary narrative thepunishment of free-loaders by enforcers is simulated when B is unfair and A penalizeshim; altruistic punishment is simulated by option 1 (A buys Bs penalty points). ThePET images of enforcers brains show activation of the caudate nucleus of the dorsalstriatum, a region is associated with dopamine excretion and part of the brains rewardcenter. The images also indicate that the intensity of the activation correlates positivelywith the severity of the punishment the number of penalty points that A assigns to B.Thus the images show that when A is paying out units to punish B, As brain is payingitself back with pleasure. In which case, (altruistic) punishment is its own reward.

The Concise Oxford Dictionary defines cruelty as having pleasure in anotherssuffering. If so, the back-story permits the conclusion that cruelty entered humannature along with our felicitous pro-social disposition to altruistic punishment. (Thisis an ethnographic observation, not a moral judgment.) The enforcers in the experiment




actively imagined the disappointment that will be experienced by the defectors; theyvicariously participated in the defectors state of mind. This is not simple cruelty but,more precisely, empathic cruelty.

Research by Takahashi et al. (2009) on Schadenfreude is evenmore explicit. Accordingto Takahashi, Schadenfreude (pleasurable response to news that a misfortune has fallento a person who is envied or resented) and envy (a painful feeling of inferiority andresentment that results from awareness of someone elses superior quality, achievement,or possessions) are two sides of one coin. Students in the experiment read descriptionsof three fictive students A, B, and C and were instructed to see B and Cfrom As perspective. Student A had average abilities, achievements, possessions, socialendowments and prospects. Student B was superior and successful in each respect andalso in life domains important to A. Student C was superior and successful but indomains not important to A. Participants silently read additional texts pertaining to A,B, and C while their brains were scanned (fMRI). In phase one, the texts describedthe successes of B and C, and participants reported how envious the descriptions madethem feel. In phase two, texts described misfortunes that spoiled events and prospectsfor A, B, and C. Participants reported the intensity of their pleasure (Schadenfreude)regarding each of the events. Brain images and self-reports were compared.

Empathic cruelty has three elements: the targets present or anticipated distress, theperpetrators pleasure (reward), and empathy (shared pain, distress, etc.). In Takahashisexperiment, brain images demonstrated the pleasure part directly, via the activationof the participants reward center (dorsal and ventral striatum and medial orbitofrontalcortex). Evidence of empathy is more complicated. Prior research shows that cognitiveconflicts and social pain are processed in the dorsal anterior cingulate cortex (dACC),part of the brains pain matrix. Brain images of the participants in Takahashisexperiment showed activation of the dACC after reading the texts. This activationhad two sources: the participants envy (phase one) and his internal representations(imagination) of the targets distress (phase two). Thus the brain images are evidenceof shared affect identified with empathy.

To the extent that envy is an expression of frustrated entitlement, the enviedperson (target) in Takahashis experiment resembles the unfair person in de Quervainsexperiment. No data relating to empathic affect was collected by de Quervain et al.Participants were simply asked to describe their own feelings while deciding to punishunfair participants (for additional research relating to empathic cruelty, see Fehr andCamerer 2007; Fliessbach et al. 2007; Knoch et al. 2006; Lanzetta and Englis 1989;Shamay-Tsoory et al. 2009; Singer et al. 2006).

Empathy and evolution

The neo-Darwinian back-story of the social brain starts with two puzzles: one puzzleconcerns altruism, the second concerns the brain. The size of the human brain is

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an evolutionary puzzle. Our ancestors split from the great apes six million years ago.During this period, the ancestral human brain quadrupled in volume. The metaboliccost of the human brain is enormous: it constitutes 2 per cent of total body weightand consumes 15 per cent of cardiac output and 20 per cent of body oxygen, andthese demands are ceaseless and inflexible. We can assume that the bigger brain paid foritself, yielding favorable cost-benefits. However efforts to model these developmentsindicate that increasing metabolic cost would eventually exceed benefits. Why didthe brain continue to grow (in size and power) when further growth was no longeradaptive? This is the puzzle, and its solution is a story about how brains adapted to otherbrains.

The process is a cognitive arms race (Byrne andWhiten 1988; Barton andDunbar 1997;Dunbar 2003). It begins with the emergence of a mind-reading capacity: the abilityto detect the intentions and predict the behavior of other individuals. Mind-readingfacilitates more complex social relations, but it also facilitates cheaters, individualswho use mind-reading to manipulate and deceive other individuals. All things beingequal, cheating is cost-effective: it gives cheaters a reproductive advantage. As cheatersincrease as a proportion of a population, social life grows more unpredictable, thusundermining the stability of social relations. Entropy looms. This might have been theevolutionary fate of our hominid ancestors, but it was not. Entropy was avoided througha further improvement to the neural hardware: the emergence of a cheater detector.This could be no more than a temporary solution however. The next generation ofopportunists used their improved brains to subvert the function (or adaptation) of thecheater-detector. Entropy was avoided by the evolution of an improved version ofthe cheater-detector that would, of course, facilitate the emergence of a cohort ofimproved cheaters. And so on over millions of years, following the dialectical logicof the neo-Darwinian back-story.

This part of the story begins with mind-reading. Opinion in neuroscience isdivided on the biological basis of this ability. There are rival explanatory accountsof mind-reading: a version based on mirror neurons and a version based on cognitivemechanisms, including analogical reasoning, as mentioned by Hume. In both versions,the evolution of empathy is understood to be a pro-social development: the glueof the social world, drawing us to help others and stopping us from hurting others(Baron-Cohen andWheelwright 2004, 163; see also Lawson et al. 2004; Baron-Cohenet al. 2005; Wheelright et al. 2006; Williams et al. 2001; Iacoboni and Dapretto 2006).According to Simon Baron-Cohen, an authority on autism, human evolution producedtwo (polar) kinds of brains: a female brain with highly developed empathic capacitiesand a male brain adapted to manipulating objects and creating systems. Empathyoriginated as a pro-social adaptation allowing females to detect the wants of pre-verbalchildren and the moods of the potentially dangerous males with whom they lived(see also Hrdy 2009). Autistic individuals are characteristically poor empathizers, andautisms epidemiology is biased towards males: 10 to 1, in high functioning autisticdisorder.









In both versions, there is the further assumption that empathy is intrinsically amorally positive disposition. According to Baron-Cohen, we respond to suffering inthree ways: the response mirrors the sufferers distress (we experience it); the responseis culturally appropriate (e.g. pity) but does not mirror the suffering; or the observertakes pleasure in the sufferers condition. Baron-Cohen equates empathy with thefirst two responses, and explicitly excludes the third. De Quervains research vindicatesa further possibility empathic cruelty in which the observers brain mirrors thesufferers distress and also takes pleasure in the sufferers condition.

This is an unexpected twist in the evolutionary back-story: evidence of an evolvedempathic disposition that is simultaneously pro-social and cruel. Human nature inthe age of neuroscience grows morally complex: a theme that recurs in the storyof the cognitive arms race, where an uncomplicated kind of cheating, the refusal toreciprocate, evolves, via mind-reading, into deception, the effort to represent the currentsituation as something different from the reality. Deception puts great demands on thebrain, which is now required to do two things simultaneously. An individual mustconstruct a lie and also withhold (inhibit) the truth. Experimental and clinical evidencesuggests that telling the truth is the brains default response, a legacy of a pro-socialevolutionary adaptation. Responding with a lie demands something extra, and . . .will engage executive prefrontal systems [responsible for planning, decision-making,and monitoring] more than does telling the truth (Spence 2004, 8; see also Spenceet al. 2004). Thus a deceiver is in constant danger of signaling the truth and betrayinghimself, for instance, by involuntarily and detectable hesitation preceding a lie. Tosucceed, deception requires a capacity for self-deception, the ability to conceal ones trueintentions and facts from oneself (Trivers 1971). At this point, two unprecedentedobjects are created there is the other (created via projection) and the self (refracted in adoppelganger, the fraudulent self).

Empathic time travel

The evolutionary origins of the self are brought to life in research on mental timetravel (Suddendorf et al. 2009). Mental time travel is the capacity to project onesself into situations in the past, future, and subjunctive (an alternative scenario to theactual past or present). Travel to the past evolved first and provided a prototype (anepisodic memory in which the thinker is spectator or protagonist) for constructingmental representations of possible futures and also alternative presents and pasts.1 Theemergence of language notably pronouns and verb forms and empathy were

1 See Ingvar 1985 for initial appearance of memory of the future; see Busby and Suddendorf 2005, andSchacter et al. 2007, on the role of the prospective brain in facilitating strategic planning and behavioralflexibility in new situations; and see Addis et al. 2007, Okuda et al. 2003, and Szpunar et al. 2007, on sharedand non-shared neural substrates of memories of the past and future.











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prerequisite for the evolution of mental time travel (Corballis 2009). The idea thatwe simply project ourselves wholesale into the past or future oversimplifies timetravel. It is a first-person experience that requires the splitting of the self: one mustbe both here-and-now and there-and-then at the same time. The bond between thesplit-selves is empathic, but not necessarily positive. (In this sense, time travel parallelsTrivers account of self-deception.)

Neuroscience makes it possible to see time travel: pain provides an efficientmodality. Jean Decety and his collaborators conducted fMRI experiments in whichthey asked participants to imagine themselves and others in painful situations(Decety and Gre`zes 2006). In other words, participants traveled to the subjunctive(an alternative present-time). One expects that, in some participants, the targetedsituations stimulated spontaneous time travel to other places as well notably travel tointensely empathic memories involving loved ones in pain. The imagined situationsactivated one might say mirrored brain regions reliably associated withexperiencing the emotional content of pain in present-time. There was nophenomenological confusion between the mental act and the experience however.Neuroimages in the self vs. other scenarios were similar but, as one might expect, therewere discernible differences which is what one would expect, given that a minimaldistinction between self and other is essential for social interaction in general and forempathy in particular.

In the neo-Darwinian back-story, human evolutionary history begins with a greatleap forward. Up to this point, social exchange is based on altruism and simple kindsof reciprocity between genetically similar individuals. Afterwards, relations take theform of networks of exchange (reciprocity) among individuals genetically unrelated oronly distantly related. Many problems were encountered along the dialectical road tomodern times. The future was one of these problems. Where reciprocity entails longdelays between giving and repayment, exchange partners must share some awarenessof time, as a continuum that connects past-time to future-time and is the sine quanon for debt. It is assumed that the concept of time would emerge from incessanttravels between memories of the past and the future.

The benefits of having or knowing time are obvious. Time travel promotesbehavioral flexibility in novel situations and it is the basis for long-term strategicplanning targeted to pre-selected goals. With the emergence of language, transientmemories of individuals could be transformed into reproducible narratives that couldbe accumulated and circulated within groups, creating a powerful collective memory.But human nature can be uncooperative, since there is a demonstrable tendency forpeople to treat present-time and future-time unequally when they calculate costs andbenefits: they discount future benefits while inflating costs incurred in the present. Thusself-interest is intrinsically impulsive and opportunistic, going for immediate gain. Ifunrestrained, it limits reciprocity and would have curtailed the dialectical developmentsdescribed in the back-story. We have already seen this work in the genesis ofpunishment: the cost-benefits of being an enforcer and the benefits (rewards) provided




by empathic cruelty and time travel to the future. A suitable countermotivationdevice was needed and there was one available:

Memory for emotions . . . does not align with our current goals. This is striking in thecommon phenomenon of rumination, the unwanted but persistent activation of thoughtsconcerning an unpleasant past situation. . . . Time travel . . . provides emotions that bypasscurrent goals, as well as time discounting and . . . provides us with immediate counter-rewards against opportunistic motivation. (Boyer 2010, 222; see above, the evolutionaryorigins of empathic cruelty)

Clinical psychiatry was acquainted with mental time travel avant la lettre, as earlyas the 1880s. Today, posttraumatic stress disorder (PTSD) is the most widely knowntime travel syndrome (Young 2004). PTSD comprises an etiological event, a distressfuland intrusive memory of this event, and a behavioral syndrome that represents anadaptation to the memory. Traumatic memories are a pathological expression of thephenomenon of rumination mentioned by Boyer (Berntsen and Hall 2004). In thelanguage of psychiatry, traumatic memories are re-experiences and their exemplaris the flashback. Schreckneurose or fright neurosis is especially interesting in thisregard. The disorder, the German variation of shellshock during World War One, wascharacterized by the victims terrifying dreams of a traumatizing experience. In theview of influential German doctors, the syndrome could be caused by re-experiences(memories, nightmares) of the future in combination with the past. The theory isthat the soldiers were fixated on visions of their deaths. The man is overcome by anempathic tenderness for himself the subject in the nightmares and intrusive images. Theremembered event is a composite of two events: a real past event and an imagined futureevent. The past is re-enacted in the future but with a significant change. The past eventwas harmless; the future event is fatal. He experiences two events as a single, etiologicalevent, in the past. His abnormality is a weakness of will and an excess of self-empathy.His symptoms, which often include psychogenic paralysis, are an unconscious effort tohide from the future. His true defect is moral, not medical. A real man (a soldier) livesin the present moment. The doctors job is to terminate to the patients pathogenictime travel with the most effective means, including electrical torture (Lerner 2003;Young 1995).2

Post-Paleolithic developments

Human nature, as portrayed in the back-story, was fully formed during the upperPaleolithic Period. The story tells us that we are innately empathic, but that

2 See also Young 2002, on self-traumatized perpetrators, a clinical phenomenon that intersects pathologicaltime travel and empathic cruelty.








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empathy may include unexpected and undesirable attitudes: dissimulation, self-deceit, spitefulness, Schadenfreude, and cruelty. Western normative institutions religion, secular ethics, clinical psychology regard these attitudes as anti-social andself-destructive. The dialectical history in this chapter views these attitudes from adifferent perspective, as the causes and consequences of human social evolution and self-awareness. Leaving aside the professional cynics, no one is claiming that these attitudesform the core or essence of human nature.Mind-reading, perspective-taking, hormonalresponsiveness, and mental time travel were likewise responsible for psychologicalaltruism, the propensity to adjust ones desires and intentions to the perceived needsor wishes of others. (In contrast, the starting point for the neo-Darwinian back-story isbiological altruism, which is defined by fitness costs and unconcerned with altruistsperceptions and intentions.) Human nature, as described in the back-story, is morallycomplex, even contradictory, certainly inclined to read and share the concern ofothers (Hrdy 2009), but likewise prepared for cruel pleasures.

Conclusion

A recent article in the New Yorker magazine celebrates a revolution in consciousnessattributed to neuroscience (Brooks 2011a). The author, David Brooks, is a columnistfor the New York Times and author of a recent book on brain science, human nature,and public policy (Brooks 2011b). Revolutions have winners and losers, and Brooksloser is the idea of consciousness that we inherited from the Enlightenment. Hiswinner is a new vision of the unconscious. According to Brooks, the revolutionelevates emotion over pure reason, social connections over individual choice, moralintuition over abstract logic, [and] perceptiveness over I.Q. The biases, longings,[and] predispositions . . . about which our culture has least to say now command ourrespect and not, as in the past, contempt. Where shall we find our ontological bearingsin this new world? Neuroscience is the source of the problem (the primacy of theunconscious) and also the source of the solution: Brain science helps fill the hole leftby the atrophy of theology and philosophy (Brooks 2011a, 26).

Multiple versions of the unconscious have circulated for two centuries. Theyshare the idea that some determinants and contents of mental life are located beyondconsciousness, and that certain kinds of puzzling behavior, mainly connected withintentionality, are attributable to these hidden elements. The notable puzzles includebehavior that occurs in the absence of conscious intentions, and occasions when peopleerroneously believe that their intentions (reasons) are also the causes of their behavior.Freuds Psychopathology of Everyday Life is of course about actions without intentions:slips of the tongue, forgetting proper names, mistakes made while reading and writing,and erroneously performed actions. Likewise his writing on Oedipal desire is aboutconfusing reasons with causes. The Freudian unconscious is obsolescent, as interest hasmoved from forces hidden in the mind to forces sequestered in the brain.





The new unconscious originates in cognitive science, in experiments published inthe 1980s. The emblematic experiment (Libet et al. 1983) challenged the idea thatour decisions to perform our actions necessarily precede the brains preparation tomake these actions happen. Libets experiment showed that cerebral initiation of aspontaneous, freely voluntary act can begin unconsciously (ibid., 399400). The oldand new unconscious are obviously unlike: Freuds self-confirming style of reasoning,based on his clinical observations, is now replaced by brain imaging, experimentation,and falsifiable hypotheses (Hassan, Uleman, and Bargh 2005; Bargh andMorsella 2008).The old and new versions are similar in one notable respect: they are organized aroundtheories and myths about evolutionary origins. Myth is not antithetical to science,and the myth of empathy, the twists and turns that lead dialectically from altruismto empathic cruelty, are not antithetical to the emergence of a science of empathy.Freuds evolutionary myth was the work of just one man, writing and revising hisaccounts over the course of three decades, in Totem and Taboo (1913), Overview of theTransference Neurosis (1915), Group Psychology (1921), andMoses and Monotheism (1939).The neo-Darwinian myth, the back-story to the social brain, began sixty years ago. It isa collective effort, combining population biology, evolutionary science, mathematicalmodeling, anthropology, primatology, experimental economics, and brain science, andit is still unfinished.

Acknowledgments

This paper has been supported by a research grant provided by the Social Science andHumanities Research Council of Canada.

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