Seeing others react to threats triggers our own internal threat responses.

From Haaker et al. (open source): 

Significance

Social transmission of threat information by observation is effective in humans and other animals. However, it is unknown if such observation of others’ reacting to threats can retrieve memories that have been previously learned through direct, firsthand aversive experiences. Here, we show concordantly in humans and rats that observing a conspecific’s reactions to a threat is sufficient to recover associative memories of direct, firsthand aversive experiences, measured as conditioned threat responses (physiological responses and defensive behavior) in the observer. The reinstatement of threat responses by observation of others is specific to the context that is observed as being dangerous. Our findings provide cross-species evidence that observation of others’ threat reactions can recover associative memories of direct, firsthand aversive experiences.

Abstract

Information about dangers can spread effectively by observation of others’ threat responses. Yet, it is unclear if such observational threat information interacts with associative memories that are shaped by the individual’s direct, firsthand experiences. Here, we show in humans and rats that the mere observation of a conspecific’s threat reactions reinstates previously learned and extinguished threat responses in the observer. In two experiments, human participants displayed elevated physiological responses to threat-conditioned cues after observational reinstatement in a context-specific manner. The elevation of physiological responses (arousal) was further specific to the context that was observed as dangerous. An analogous experiment in rats provided converging results by demonstrating reinstatement of defensive behavior after observing another rat’s threat reactions. Taken together, our findings provide cross-species evidence that observation of others’ threat reactions can recover associations previously shaped by direct, firsthand aversive experiences. Our study offers a perspective on how retrieval of threat memories draws from associative mechanisms that might underlie both observations of others’ and firsthand experiences.

How mice feel each other's pain or fear

The abstract from Smith et al, who show the brain basis of empathetic behaviors in mice that mirror those in humans:

Empathy is an essential component of social communication that involves experiencing others’ sensory and emotional states. We observed that a brief social interaction with a mouse experiencing pain or morphine analgesia resulted in the transfer of these experiences to its social partner. Optogenetic manipulations demonstrated that the anterior cingulate cortex (ACC) and its projections to the nucleus accumbens (NAc) were selectively involved in the social transfer of both pain and analgesia. By contrast, the ACC→NAc circuit was not necessary for the social transfer of fear, which instead depended on ACC projections to the basolateral amygdala. These findings reveal that the ACC, a brain area strongly implicated in human empathic responses, mediates distinct forms of empathy in mice by influencing different downstream targets. 

Here is a summary graphic from a perspective by Klein and Gogolla (click to enlarge):

Mammalian empathy: neural basis and behavioral manifestations

I want to point to an interesting review by de Waal and Preston in Nature Reviews Neuroscience. Here are the Abstract and a few excerpts from the article:

Recent research on empathy in humans and other mammals seeks to dissociate emotional and cognitive empathy. These forms, however, remain interconnected in evolution, across species and at the level of neural mechanisms. New data have facilitated the development of empathy models such as the perception–action model (PAM) and mirror-neuron theories. According to the PAM, the emotional states of others are understood through personal, embodied representations that allow empathy and accuracy to increase based on the observer's past experiences. In this Review, we discuss the latest evidence from studies carried out across a wide range of species, including studies on yawn contagion, consolation, aid-giving and contagious physiological affect, and we summarize neuroscientific data on representations related to another's state.

Key points:

Observational and experimental studies dating back to the 1950s demonstrate that mammals spontaneously help distressed conspecifics. Research emphasizes the untrained, unrewarded nature of this behaviour, which is also biased towards familiar individuals, thus arguing against explanations that are exclusively based on associative learning or conditioning.

The perception–action model extends an existing motor theory on overlapping representations to emotional phenomena; it states that observers who attend to a target's state understand and 'feel into' it through personal distributed representations of the target, the state and the situation. Easily observed manifestations of this mechanism are emotional contagion and motor mimicry, which have been demonstrated in many animals. In cognitive forms of empathy, the same representations are accessed from the top-down.

Experiments on two common mammalian expressions of empathy — the consolation of distressed individuals and spontaneous assistance to those in need — support the crucial role of caught distress and arousal because these behaviours are suppressed by anti-anxiety medication and engage the same neuropeptide system that supports social attachment.

The Russian-doll model seeks to arrange forms of empathy into layers that are built on top of each other — with the layers ranging from emotional contagion to more cognitive forms of empathy — in a functionally integrated whole based on perception–action processes. Perspective-taking is well developed in some non-human species, as manifested by theory-of-mind and targeted helping.

One can segregate emotional and cognitive empathy (as well as felt and observed states) in the brains of observers, but all forms require some initial access to the observer's distributed, shared, personal representations of the target's state. At least in the initial phase of processing, this access helps to decode the target's state and provide subsequent processing with content and meaning, even if the shared state is not experienced, or is incomplete or inaccurate.

Empathic pain does not usually include the peripheral sensation of the target's injury, but it can include sensory information when the stimuli and task instructions emphasize the specific nature of the feeling at the location of the injury.

The Russian Doll Model of the Evolution of Empathy

The evolution of dance

From Laland et al.:

Evidence from multiple sources reveals a surprising link between imitation and dance. As in the classical correspondence problem central to imitation research, dance requires mapping across sensory modalities and the integration of visual and auditory inputs with motor outputs. Recent research in comparative psychology supports this association, in that entrainment to a musical beat is almost exclusively observed in animals capable of vocal or motor imitation. Dance has representational properties that rely on the dancers’ ability to imitate particular people, animals or events, as well as the audience’s ability to recognize these correspondences. Imitation also plays a central role in learning to dance and the acquisition of the long sequences of choreographed movements are dependent on social learning. These and other lines of evidence suggest that dancing may only be possible for humans because its performance exploits existing neural circuitry employed in imitation.

This painting by Edgar Degas not only depicts a ballet rehearsal but also illustrates the roles of imitation and synchrony

Clips from the body of the article, with another figure:

...there can be no doubt that, compared to other animals, humans are exceptional imitators. It may be no coincidence that a recent PET scan analysis of the neural basis of dance found that foot movement to music excited regions of neural circuitry (e.g. the right frontal operculum) previously associated with imitation. Dancing may only be possible because its performance exploits the neural circuitry employed in imitation. Such reasoning applies equally where individuals dance alone; unlike much human behavior, dancing inherently seems to require a brain capable of solving the correspondence problem.

Dance often tells a story, and this representational quality provides another link with imitation. For instance, in the astronomical dances of ancient Egypt, priests and priestesses, accompanied by harps and pipes, mimed significant events in the story of a god or imitated cosmic patterns, such as the rhythm of night and day. Africa, Asia, Australasia and Europe all possess long-standing traditions for masked dances, in which the performers portray the character associated with the mask and enact religious stories. Native Americans have many animal dances, such as the Buffalo dance, which was thought to lure buffalo herds close to the village, and the eagle dance, which is a tribute to these revered birds. This tradition continues to the present. In 2009, Rambert (formerly the Rambert Dance Company), a world leader in contemporary dance, marked the bicentenary of Charles Darwin’s birth and 150th anniversary of his seminal work On the Origin of Species by collaborating with one of us (N.C.) to produce Comedy of Change (Figure above), which evoked animal behaviour on stage with spellbinding accuracy. In all such instances, the creation and performance of the dance requires an ability on the part of the dancer to imitate the movements and sounds of particular people, animals, or events. Such dances re-introduce the correspondence problem, as the dancer, choreographer and audience must be able to connect the dancers’ movements to the target phenomenon they represent.

How learning shapes the empathic brain.

From Hein et al.:

Abstract

Deficits in empathy enhance conflicts and human suffering. Thus, it is crucial to understand how empathy can be learned and how learning experiences shape empathy-related processes in the human brain. As a model of empathy deficits, we used the well-established suppression of empathy-related brain responses for the suffering of out-groups and tested whether and how out-group empathy is boosted by a learning intervention. During this intervention, participants received costly help equally often from an out-group member (experimental group) or an in-group member (control group). We show that receiving help from an out-group member elicits a classical learning signal (prediction error) in the anterior insular cortex. This signal in turn predicts a subsequent increase of empathy for a different out-group member (generalization). The enhancement of empathy-related insula responses by the neural prediction error signal was mediated by an establishment of positive emotions toward the out-group member. Finally, we show that surprisingly few positive learning experiences are sufficient to increase empathy. Our results specify the neural and psychological mechanisms through which learning interacts with empathy, and thus provide a neurobiological account for the plasticity of empathic reactions.

Knowing the internal states of ourselves and of others.

We frequently can infer the emotional state of someone from the style of their movement, distinguishing, for example, when they hand us something rudely versus gently. The time, space, force, trajectory, and direction of the handing movement differ, reflecting what Rizzolattia and colleagues call different 'vitality forms.' They find that vitality forms in the three different tasks of action observation, imagination, and self execution correlate with consistent activation of the dorsocentral sector of the insula:

Vitality form is a term that describes the style with which motor actions are performed (e.g., rude, gentle, etc.). They represent one characterizing element of conscious and unconscious bodily communication. Despite their importance in interpersonal behavior, vitality forms have been, until now, virtually neglected in neuroscience. Here, using the functional MRI (fMRI) technique, we investigated the neural correlates of vitality forms in three different tasks: action observation, imagination, and execution. Conjunction analysis showed that, in all three tasks, there is a common, consistent activation of the dorsocentral sector of the insula. In addition, a common activation of the parietofrontal network, typically active during arm movements production, planning, and observation, was also found. We conclude that the dorsocentral part of the insula is a key element of the system that modulates the cortical motor activity, allowing individuals to express their internal states through action vitality forms. Recent monkey anatomical data show that the dorsocentral sector of the insula is, indeed, connected with the cortical circuit involved in the control of arm movements.

The evolution of gender effects on empathy.

Christov-Moore et al. offer a review making the point that differences in the capacity for empathy between males and females have deep evolutionary and developmental roots, in addition to any cultural expectations about gender roles. The review references a graphic summary of brain areas involved in experience sharing which I also pass on.

 Highlights

• Sex differences in empathy have phylogenetic and ontogenetic roots in biology. 

• As primary caregivers females evolved adaptations to be sensitive to infants’ signals. 

• Sex differences in empathy appear to be consistent and stable across the lifespan. 

• In affective empathy, females, compared to men, show higher emotional responsivity. 

• Males show greater recruitment of brain areas for the control of cognitive empathy.

Abstract

Evidence suggests that there are differences in the capacity for empathy between males and females. However, how deep do these differences go? Stereotypically, females are portrayed as more nurturing and empathetic, while males are portrayed as less emotional and more cognitive. Some authors suggest that observed gender differences might be largely due to cultural expectations about gender roles. However, empathy has both evolutionary and developmental precursors, and can be studied using implicit measures, aspects that can help elucidate the respective roles of culture and biology. This article reviews evidence from ethology, social psychology, economics, and neuroscience to show that there are fundamental differences in implicit measures of empathy, with parallels in development and evolution. Studies in nonhuman animals and younger human populations (infants/children) offer converging evidence that sex differences in empathy have phylogenetic and ontogenetic roots in biology and are not merely cultural byproducts driven by socialization. We review how these differences may have arisen in response to males’ and females’ different roles throughout evolution. Examinations of the neurobiological underpinnings of empathy reveal important quantitative gender differences in the basic networks involved in affective and cognitive forms of empathy, as well as a qualitative divergence between the sexes in how emotional information is integrated to support decision making processes. Finally, the study of gender differences in empathy can be improved by designing studies with greater statistical power and considering variables implicit in gender (e.g., sexual preference, prenatal hormone exposure). These improvements may also help uncover the nature of neurodevelopmental and psychiatric disorders in which one sex is more vulnerable to compromised social competence associated with impaired empathy.

The summary graphic by Zaki and Ochsner :

Neuroscientific approaches to studying experience sharing and mentalizing. (a) The experimental logic underlying first-person perception studies of experience sharing. The blue circle represents brain regions engaged by direct, first-person experience of an affective response, motor intention, or other internal state. The yellow circle represents regions engaged by third-person observation of someone else experiencing the same kind of internal state. To the extent that a region demonstrates neural resonance—common engagement by first- and third-person experience (green overlap)—it is described as supporting a perceiver's vicarious experience of a target's state (regions demonstrating such properties are highlighted in green in c). (b) Studies of mentalizing typically ask participants to make judgments about targets’ beliefs, thoughts, intentions and/or feelings, as depicted in highly stylized social cues, including vignettes (top left), posed facial expressions (right), or even more isolated nonverbal cues, such as target eye gaze (bottom left). Regions engaged by such tasks (blue in c) are described as contributing to perceivers’ ability to mentalize. (c) Brain regions associated with experience sharing (green) and mentalizing (blue). IPL, inferior parietal lobule; TPJ, temporoparietal junction; pSTS, posterior superior temporal sulcus; TP, temporal pole; AI, anterior insula; PMC, premotor cortex; PCC, posterior cingulate cortex; ACC, anterior cingulate cortex; MPFC, medial prefrontal cortex.

Cerebral coherence between communicators.

Stolk et al. have searched across the whole brain for a cerebral dynamics matching the behavioral dynamics of mutual understanding and note that brain activities in the right temporal lobes of individuals synchronize during communication in a way that reflects conceptualization of a signal's use apart from specific experiences of the signal:

How can we understand each other during communicative interactions? An influential suggestion holds that communicators are primed by each other’s behaviors, with associative mechanisms automatically coordinating the production of communicative signals and the comprehension of their meanings. An alternative suggestion posits that mutual understanding requires shared conceptualizations of a signal’s use, i.e., “conceptual pacts” that are abstracted away from specific experiences. Both accounts predict coherent neural dynamics across communicators, aligned either to the occurrence of a signal or to the dynamics of conceptual pacts. Using coherence spectral-density analysis of cerebral activity simultaneously measured in pairs of communicators, this study shows that establishing mutual understanding of novel signals synchronizes cerebral dynamics across communicators’ right temporal lobes. This interpersonal cerebral coherence occurred only within pairs with a shared communicative history, and at temporal scales independent from signals’ occurrences. These findings favor the notion that meaning emerges from shared conceptualizations of a signal’s use.

Synchrony between observers' brains during action observation.

Nummenmaa et al. (free access article, check out the nice graphics) have examined how we can understand what another person might be thinking or feeling just by observing their actions:

A frontoparietal action–observation network (AON) has been proposed to support understanding others' actions and goals. We show that the AON “ticks together” in human subjects who are sharing a third person's feelings. During functional magnetic resonance imaging, 20 volunteers watched movies depicting boxing matches passively or while simulating a prespecified boxer's feelings. Instantaneous intersubject phase synchronization (ISPS) was computed to derive multisubject voxelwise similarity of hemodynamic activity and inter-area functional connectivity. During passive viewing, subjects' brain activity was synchronized in sensory projection and posterior temporal cortices. Simulation induced widespread increase of ISPS in the AON (premotor, posterior parietal, and superior temporal cortices), primary and secondary somatosensory cortices, and the dorsal attention circuits (frontal eye fields, intraparietal sulcus). Moreover, interconnectivity of these regions strengthened during simulation. We propose that sharing a third person's feelings synchronizes the observer's own brain mechanisms supporting sensations and motor planning, thereby likely promoting mutual understanding.

Feeling the social touch being observed in others.

Interesting work by Bolognini et al. on our mirroring of the emotions of others :

Touch has an emotional and communicative meaning, and it plays a crucial role in social perception and empathy. The intuitive link between others’ somatosensations and our sense of touch becomes ostensible in mirror-touch synesthesia, a condition in which the view of a touch on another person’s body elicits conscious tactile sensations on the observer’s own body. This peculiar phenomenon may implicate normal social mirror mechanisms. Here, we show that mirror-touch interference effects, synesthesia-like sensations, and even phantom touches can be induced in nonsynesthetes by priming the primary somatosensory cortex (SI) directly or indirectly via the posterior parietal cortex. These results were obtained by means of facilitatory paired-pulse transcranial magnetic stimulation (ppTMS) contingent upon the observation of touch. For these vicarious effects, the SI is engaged at 150 ms from the onset of the visual touch. Intriguingly, individual differences in empathic abilities, assessed with the Interpersonal Reactivity Index, drive the activity of the SI when nonsynesthetes witness others’ tactile sensations. This evidence implies that, under normal conditions, touch observation activates the SI below the threshold for perceptual awareness; through the visual-dependent tuning of SI activity by ppTMS, what is seen becomes felt, namely, mirror-touch synesthesia. On a broader perspective, the visual responsivity of the SI may allow an automatic and unconscious transference of the sensation that another person is experiencing onto oneself, and, in turn, the empathic sharing of somatosensations.

Compassion - Bridging Practice and Science

Science magazine has an article by Kupferschmidt pointing to the work of Tania Singer, a director at the Max Planck Institute for Human Cognitive and Brain Sciences in Leipzig, who has embarked on an ambitious study involving 160 participants to find out whether meditation can make people more compassionate. He notes that meditation research does not have a very rigorous reputation, and some scientists are skeptical about the work, but Singer — who has long practiced meditation herself—hopes her study will be methodologically rigorous enough to withstand criticism. In 2004 Singer published a landmark paper in Science that showed that bilateral anterior insula (AI), rostral anterior cingulate cortex (ACC), brainstem, and cerebellum were activated when subjects received pain and also by a signal that a loved one experienced pain. AI and ACC activation correlated with individual empathy scores. Singer has just release a free 900-page e-book, entitled "Compassion - Bridging Practice and Science", that is quite amazing. I'm going to spend some time paging through it. Here is one clip from Kupferschmidt's commentary:

Numerous studies have shown that people can be "primed" to think more socially in various ways—from reading simple instructions to holding a warm cup of coffee. In one test, participants who listened to Bob Sinclar's hit song "Love Generation" were more likely to come up with words like "help" than those who listened to Sinclar's less uplifting song "Rock This Party." But Singer isn't interested in words; she wants to train people to act more socially in everyday life. And from personal experience, she believes meditation may be the way to do it. At its most basic, the technique simply involves focusing on a feeling. In one meditation exercise in her study, participants are told to imagine a person they love and to concentrate on positive feelings toward them. "May you be happy. May you be safe and sheltered. May you be healthy. May you live with a light heart," the teacher intones. Like bodybuilders increasing the weights they lift, meditators can intensify their compassionate feelings over time. Expert meditators can go very far, Singer says; rape victims may meditate on feeling compassion for their rapist, for instance. To measure meditation's effects, researchers in the ReSource Project determine the level of the stress hormone cortisol in participants' saliva, test their reaction times, have them fill out questionnaires, and shepherd them through virtual reality worlds while monitoring their heart rate. Each participant's brain is scanned for several hours five times over the course of the study. Participants also play computer games designed to evaluate their compassion level. In one of them, developed with Swiss economist Ernst Fehr, they have to guide a smiley along a winding path that leads to a treasure chest; they have blue or red keys to open gates of the same color. But another smiley is also wandering the screen, on its own quest to another treasure, and players have to decide whether to open gates for it, too. In a preliminary study in 2011, Singer showed that just one day of compassion meditation made people more likely to help the other smiley, whereas 1 day of memory training did not. Singer is also trying to better understand what goes on in the brain when it is feeling compassion. The activation patterns seen in the scanner leave open two possibilities: The feeling could be linked to the neurotransmitter dopamine and the brain's reward circuits (which, among many other things, makes you crave chocolate) or it could be linked to what she calls the affiliation network, which is activated for example when you view a picture of your partner or your own child, and is mediated by the neurotransmitters oxytocin or opioids. Singer admits that pinning down the neurobiology of compassion is difficult because the mental state it corresponds to remains fuzzy. A French Buddhist monk may have a very different concept of compassion than an African doctor or a British businessman, and there's friction between the classic third-person perspective of science and subjective experiences. "But we need the first-person experience as well as the third-person science," she says.

The limits of empathy

I thought I would follow up the Monday's post on well being, kindness, happiness and all that good stuff by noting a piece on how feel-good energy can lead us astray. Yale psychologist Paul Bloom has done an excellent article in the May 20 issue of the The New Yorker titled “The baby in the well - the limits of empathy.” Well meant feelings and actions of empathy can in some cases be counterproductive and blind us to more remote but statistically much more important hardships. Our evolved ability to feel what others are feeling (see numerous mindblog posts on mirror neurons, etc. ) is applied to very explicit and limited human situations, usually a specific individual (6 year old girl falls in well and nation focuses on watching the rescue) or defined and limited groups (mass shootings at Sandy Hook or Boston Marathon bombing). From Bloom:

In the past three decades, there were some sixty mass shootings, causing about five hundred deaths; that is, about one-tenth of one per cent of the homicides in America. But mass murders get splashed onto television screens, newspaper headlines, and the Web; the biggest ones settle into our collective memory —Columbine, Virginia Tech, Aurora, Sandy Hook. The 99.9 per cent of other homicides are, unless the victim is someone you’ve heard of, mere background noise.

After noting how empathy research is thriving, and several books arguing that more empathy has to be a good thing (with Rifkin, in “The Empathic Civilization” (Penguin), wanting us to make the leap to “global empathic consciousness”), Bloom notes:

This enthusiasm may be misplaced, however. Empathy has some unfortunate features—it is parochial, narrow-minded, and innumerate. We’re often at our best when we’re smart enough not to rely on it......the key to engaging empathy is what has been called “the identifiable victim effect.” As the economist Thomas Schelling, writing forty-five years ago, mordantly observed, “Let a six-year-old girl with brown hair need thousands of dollars for an operation that will prolong her life until Christmas, and the post office will be swamped with nickels and dimes to save her. But let it be reported that without a sales tax the hospital facilities of Massachusetts will deteriorate and cause a barely perceptible increase in preventable deaths—not many will drop a tear or reach for their checkbooks.”

You can see the effect in the lab. The psychologists Tehila Kogut and Ilana Ritov asked some subjects how much money they would give to help develop a drug that would save the life of one child, and asked others how much they would give to save eight children. The answers were about the same. But when Kogut and Ritov told a third group a child’s name and age, and showed her picture, the donations shot up—now there were far more to the one than to the eight.

In the broader context of humanitarianism, as critics like Linda Polman have pointed out, the empathetic reflex can lead us astray. When the perpetrators of violence profit from aid—as in the “taxes” that warlords often demand from international relief agencies—they are actually given an incentive to commit further atrocities.

A “politics of empathy” doesn’t provide much clarity in the public sphere, either. Typically, political disputes involve a disagreement over whom we should empathize with. Liberals argue for gun control, for example, by focussing on the victims of gun violence; conservatives point to the unarmed victims of crime, defenseless against the savagery of others.

On many issues, empathy can pull us in the wrong direction. The outrage that comes from adopting the perspective of a victim can drive an appetite for retribution....In one study, conducted by Jonathan Baron and Ilana Ritov, people were asked how best to punish a company for producing a vaccine that caused the death of a child. Some were told that a higher fine would make the company work harder to manufacture a safer product; others were told that a higher fine would discourage the company from making the vaccine, and since there were no acceptable alternatives on the market the punishment would lead to more deaths. Most people didn’t care; they wanted the company fined heavily, whatever the consequence.

There’s a larger pattern here. Sensible policies often have benefits that are merely statistical but victims who have names and stories. Consider global warming—what Rifkin calls the “escalating entropy bill that now threatens catastrophic climate change and our very existence.” As it happens, the limits of empathy are especially stark here. Opponents of restrictions on CO2 emissions are flush with identifiable victims—all those who will be harmed by increased costs, by business closures. The millions of people who at some unspecified future date will suffer the consequences of our current inaction are, by contrast, pale statistical abstractions.

Moral judgment entails more than putting oneself in another’s shoes. “The decline of violence may owe something to an expansion of empathy,” the psychologist Steven Pinker has written, “but it also owes much to harder-boiled faculties like prudence, reason, fairness, self-control, norms and taboos, and conceptions of human rights.” A reasoned, even counter-empathetic analysis of moral obligation and likely consequences is a better guide to planning for the future than the gut wrench of empathy.

Newtown, in the wake of the Sandy Hook massacre, was inundated with so much charity that it became a burden. More than eight hundred volunteers were recruited to deal with the gifts that were sent to the city—all of which kept arriving despite earnest pleas from Newtown officials that charity be directed elsewhere....Meanwhile—just to begin a very long list—almost twenty million American children go to bed hungry each night, and the federal food-stamp program is facing budget cuts of almost twenty per cent.

Such are the paradoxes of empathy. The power of this faculty has something to do with its ability to bring our moral concern into a laser pointer of focussed attention. If a planet of billions is to survive, however, we’ll need to take into consideration the welfare of people not yet harmed—and, even more, of people not yet born. They have no names, faces, or stories to grip our conscience or stir our fellow-feeling. Their prospects call, rather, for deliberation and calculation. Our hearts will always go out to the baby in the well; it’s a measure of our humanity. But empathy will have to yield to reason if humanity is to have a future.

Social interactions prime us for motor empathy or resonance.

Hogeveen and Obhi1 find that recent experience tunes our mirroring systems to particular agent types. A bit from their introduction, followed by the abstract:

Detecting and responding to biological stimuli such as predators or potential mates is a fundamental and adaptive capability, supported by rain areas such as the posterior superior temporal sulcus (pSTS) which is biased for processing biological motion. The pSTS and the parietofrontal mirror system form part of a wider action observation network (AON), which is thought to underlie many social abilities. Motor resonance (MR) is the activation of matching motor representations during observation of action(s) made by others, and could index mirror activity within the wider AON.

Understanding the neural basis of social behavior has become an important goal for cognitive neuroscience and a key aim is to link neural processes observed in the laboratory to more naturalistic social behaviors in real-world contexts. Although it is accepted that mirror mechanisms contribute to the occurrence of motor resonance (MR) and are common to action execution, observation, and imitation, questions remain about mirror (and MR) involvement in real social behavior and in processing nonhuman actions. To determine whether social interaction primes the MR system, groups of participants engaged or did not engage in a social interaction before observing human or robotic actions. During observation, MR was assessed via motor-evoked potentials elicited with transcranial magnetic stimulation. Compared with participants who did not engage in a prior social interaction, participants who engaged in the social interaction showed a significant increase in MR for human actions. In contrast, social interaction did not increase MR for robot actions. Thus, naturalistic social interaction and laboratory action observation tasks appear to involve common MR mechanisms, and recent experience tunes the system to particular agent types.

We don't project our visceral states onto dissimilar others.

Interesting observations from O’Brien and Ellsworth on limits to the empathy of our embodied cognition:

What people feel shapes their perceptions of others. We have examined the assimilative influence of visceral states on social judgment. Replicating prior research, we found in a first experiment that participants who were outside during winter overestimated the extent to which other people were bothered by cold, and in a second study found that participants who ate salty snacks without water thought other people were overly bothered by thirst. However, in both studies, this effect evaporated when participants believed that the other people under consideration held political views opposing their own. Participants who judged these dissimilar others were unaffected by their own strong visceral-drive states, a finding that highlights the power of dissimilarity in social judgment. Dissimilarity may thus represent a boundary condition for embodied cognition and inhibit an empathic understanding of shared out-group pain. Our findings reveal the need for a better understanding of how people’s internal experiences influence their perceptions of the feelings and experiences of those who may hold values different from their own.

Feeling the moves - motor empathy with expert performance

Jola et al. make the interesting observation that experienced viewers of ballet, even without physical training, covertly simulate the movements for which they have acquired visual experience, their empathic abilities heighten motor resonance during dance observation - activating the same brain motor pathways actually being used by the dancers:

The human “mirror-system” is suggested to play a crucial role in action observation and execution, and is characterized by activity in the premotor and parietal cortices during the passive observation of movements. The previous motor experience of the observer has been shown to enhance the activity in this network. Yet visual experience could also have a determinant influence when watching more complex actions, as in dance performances. Here we tested the impact visual experience has on motor simulation when watching dance, by measuring changes in corticospinal excitability. We also tested the effects of empathic abilities. To fully match the participants' long-term visual experience with the present experimental setting, we used three live solo dance performances: ballet, Indian dance, and non-dance. Participants were either frequent dance spectators of ballet or Indian dance, or “novices” who never watched dance. None of the spectators had been physically trained in these dance styles. Transcranial magnetic stimulation was used to measure corticospinal excitability by means of motor-evoked potentials (MEPs) in both the hand and the arm, because the hand is specifically used in Indian dance and the arm is frequently engaged in ballet dance movements. We observed that frequent ballet spectators showed larger MEP amplitudes in the arm muscles when watching ballet compared to when they watched other performances. We also found that the higher Indian dance spectators scored on the fantasy subscale of the Interpersonal Reactivity Index, the larger their MEPs were in the arms when watching Indian dance. Our results show that even without physical training, corticospinal excitability can be enhanced as a function of either visual experience or the tendency to imaginatively transpose oneself into fictional characters. We suggest that spectators covertly simulate the movements for which they have acquired visual experience, and that empathic abilities heighten motor resonance during dance observation.

Our mindreading of another person depends on how much skin we see!

This interesting piece in the Journal of Personality and Social Psychology makes observations on how our mindreading, or inferring someone's nature, depends on how much of them we are seeing. From the introduction:

Do people’s mental capacities fundamentally change when they remove a sweater? This seems absurd: How could removing a piece of clothing change one’s capacity for acting or feeling? In six studies, however, we show that taking off a sweater—or otherwise revealing flesh—can significantly change the way a mind is perceived. In this article, we suggest that the kind of mind ascribed to another person depends on the relative salience of his or her body—that the perceived capacity for both pain and planned action depends on whether someone wears a sweater or tank-top.

The abstract:

According to models of objectification, viewing someone as a body induces de-mentalization, stripping away their psychological traits. Here evidence is presented for an alternative account, where a body focus does not diminish the attribution of all mental capacities but, instead, leads perceivers to infer a different kind of mind. Drawing on the distinction in mind perception between agency and experience, it is found that focusing on someone's body reduces perceptions of agency (self-control and action) but increases perceptions of experience (emotion and sensation). These effects were found in three experiments when comparing targets represented by both revealing versus nonrevealing pictures or by simply directing attention toward physical characteristics. In two further experiments he effect of a body focus on mind perception also influenced moral intuitions, with those represented as a body seen to be less morally responsible (i.e., lesser moral agents) but more sensitive to harm (i.e., greater moral patients). These effects suggest that a body focus does not cause objectification per se but, instead, leads to a redistribution of perceived mind.

Heirarchies of empathy in the brain

Following the previous post I thought it would be useful to pass on more of Panskepp's review, which provides a more general description:

There is a growing recognition of how animals respond to the affective states of other animals, including the show of empathy, a state once thought to be unique to primates...A key question concerns the nature of the rats' motivations—the affective and cognitive underpinnings of their “empathy.” ...Future research needs to untangle whether empathic responses in mammals arise more from higher cognitive or lower affective brain functions, or some combination of these (see the figure). Human brain imaging studies of empathy suggest both are involved, especially in coping with the distress of others. But solid neurobiologically based evolutionary evidence, both bottom-up and top-down, is so far lacking.

Figure legend - One concept of how mammalian brains generate empathic responses at different levels is shown. Primary emotional processes, where sources of empathy may arise (i.e., feeling what other organisms are feeling), coordinate with secondary-process learning and memory mechanisms (i.e., knowing what others are feeling). Both of these then interact with higher mental processes, which can exert a variety of top-down influences on the regulation of empathic tendencies (i.e., desires to respond compassionately to others' distress).

The layering of evolutionary progressions is evident in the human brain. The deepest midbrain and hypothalamic regions mediate primary-process, instinctual affects. More recently evolved subcortical regions, among them basal ganglia, amygdala, and nucleus accumbens, help promote higher cognitive activities through learning and memory. Although we currently look to mirror-neuron zones of the neocortex for evidence of the highest mind functions such as compassion, empathic tendencies are surely also promoted by the more ancient primary-process emotional networks that are essential foundations for mental life. For example, a primal form of “empathy” is mothers' exquisite sensitivity to crying babies. Might crying access those systems in mothers' brains that are known to mediate separation anxiety in young animals? Perhaps affective urges for maternal caregiving are triggered as mothers' brains experience psychological pain engendered by their infants' cries. It may be that empathic coordination of social motivations is mediated by emotional resonances among nearby animals, allowing receivers to experience the emotions of transmitters. At such deep affective levels, emotional states may reverberate among animals, with no need for learned rerepresentations arising from mirror neurons. Mammals may have intrinsic abilities to resonate with the pains and joys of nearby others through primal emotional contagion. 

Much deep-brain research remains to be done to understand the degree to which mammalian empathy is achieved more through higher social-cognitive processes or primal affective processes in the brain. Simplified models of empathy, as in mice and rats, offer new inroads for understanding our own social-emotional nature and nurture. Such knowledge may eventually help us promote nurturant behaviors in humans.

When it's an error to mirror...

Mimicry and imitation can facilitate cultural learning, maintenance of culture, and group cohesion, and individuals must competently select the appropriate models and actions to imitate. Mimicry and imitation also play an important role in dyadic social interactions. People mimic their partners’ mannerisms, which increases rapport and the partners’ liking of the mimickersA collaboration between psychologists and philosophers at the Univ. of California, San Diego asks whether and how mimicry unconsciously influences evaluations made by third-party observers of dyadic interactions. Their results indicate that third-party observers make judgments about individuals’ competence on the basis of their decisions concerning whether and whom to mimic. Contrary to the notion that mimicry is uniformly beneficial to the mimicker, people who mimicked an unfriendly model were rated as less competent than nonmimics. Thus, a positive reputation depends not only on the ability to mimic, but also on the ability to discriminate when not to mimic. Here is their experimental setup (click on figure to enlarge):

Figure: Illustration of the experimental paradigm and experimental results. Subjects watched two videos, in each of which an interviewer (model) interacted with an interviewee. After each video, subjects rated the interviewee’s competence, trustworthiness, and likeability. For each subject, one video showed a mimicking interviewee, and the other showed a nonmimicking interviewee. In Experiment 1, video frames were uncropped, so subjects could see the interviewer; in Experiment 2, video frames were cropped, so subjects could not see the interviewer, and mimicry was obscured. The interviewer’s attitude varied between subjects; some subjects saw videos with a cordial interviewer, and other subjects saw videos with a condescending interviewer. The graph shows the difference in average competence ratings between the cordial- and condescending-model conditions as a function of whether or not the interviewee mimicked the interviewer, separately for Experiments 1 and 2. Error bars represent standard errors of the difference between conditions.

The Empathetic Brain

Christian Keyser, who does research on mirror neurons at The Netherlands Institute for Neuroscience, sent me an email yesterday pointing out his new book on human empathy. My scan of the copy he sent to me finds the book to written in a friendly, engaging, and accessible style for a non-technical audience. I point it out here because it is available as a Kindle Book at Amazon, and the price is right ($3.00).

Speakers and Listeners - fMRI shows coupled brains

As a followup to Monday's post on mirror neurons, this fascinating study by Stephens et al. shows that brain activities in a speaker-listener pair are tightly coupled, and that the magnitude of activity in areas exhibiting predictive anticipatory responses correlates with understanding. The graphic summaries of fMRI data in this open access article are quite nice, and you might want to check them out. The MindBlog reader who pointed out this early PNAS publication to me wonders "Could their findings open a new window of how to interpret the "function" of "conscious self", with the conscious self as the evaluating "outpost" of the coupled companion.?"

Verbal communication is a joint activity; however, speech production and comprehension have primarily been analyzed as independent processes within the boundaries of individual brains. Here, we applied fMRI to record brain activity from both speakers and listeners during natural verbal communication. We used the speaker's spatiotemporal brain activity to model listeners’ brain activity and found that the speaker's activity is spatially and temporally coupled with the listener's activity. This coupling vanishes when participants fail to communicate. Moreover, though on average the listener's brain activity mirrors the speaker's activity with a delay, we also find areas that exhibit predictive anticipatory responses. We connected the extent of neural coupling to a quantitative measure of story comprehension and find that the greater the anticipatory speaker–listener coupling, the greater the understanding. We argue that the observed alignment of production- and comprehension-based processes serves as a mechanism by which brains convey information.