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Mammalian empathy: behavioural manifestations and neural basis

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.

Abstract

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.

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Figure 1: The Russian-doll model of the evolution of empathy.
Figure 2: Behavioural manifestations of animal empathy.
Figure 3: From affect transfer to altruism.
Figure 4: Both cognitive and affective empathy access distributed, person-specific affective representations.
Figure 5: Neural regions that participate in human empathy.

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Acknowledgements

The authors thank their sources of funding during the preparation of this manuscript, including support for the Living Links Center from Emory College, Georgia, USA (to F.B.M.d.W.) and a grant from Rackham Graduate School at the University of Michigan, USA (to S.D.P.). The authors also thank Y. Lei for assistance with manuscript preparation, and L. Bickel for creating the original brain figures.

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Glossary

Empathy

Any process that emerges from the fact that observers understand others' states by activating personal, neural and mental representations of that state, including the capacity to be affected by and share the emotional state of another; assess the reasons for the other's state; and identify with the other, adopting his or her perspective.

Theory-of-mind

The ability to attribute mental states to others, such as knowledge, intentions and beliefs.

Affective empathy

Also known as emotional empathy. Empathy that is directly affected by the emotional state of another by matching or 'feeling with' it, as a result of perceiving this state.

Cognitive empathy

Empathy derived from a top-down process in which the observer imagines how the target feels, even if the target is not present or their feelings cannot be directly observed.

Empathic perspective-taking

The capacity to take another's affective perspective: for example, understanding their specific situation and needs, separate from one's own, which still requires access to personal representations of the other's state.

False-belief task

A crucial theory-of-mind task that determines whether an observer knows what another knows, even if this knowledge is incongruent with the observer's own.

Targeted helping

Assistance and care based on a cognitive appreciation of the other's specific need or circumstances.

Tinbergian framework

A series of 'why' questions that we may ask about any observed behaviour, as proposed by Niko Tinbergen. The questions concern different levels of causation: for example, why did a behaviour evolve (what are its benefits), why does the behaviour occur (what caused it) and what is the behaviour's phylogenetic origin?

Emotional contagion

Emotional state matching between a target and an observer.

Consolation behaviour

Reassurance behaviour directed at a distressed party, such as a victim of aggression (also see the definition of 'empathic concern').

Empathic concern

Also known as sympathetic concern. Concern about another's state, and attempts to ameliorate this state (also see the definition of 'consolation behaviour').

Perception–action mechanism

Spontaneous activation of an individual's own personal representations for a target, their state and their situation when perceiving the target's state.

Emotion self-regulation

Control over one's own emotions to promote adaptive responding, including response delay, recovery from upsets and selective attention.

Directed altruism

Helping or comforting behaviour directed at an individual in need, pain or distress.

Convergent evolution

A process in which unrelated species independently evolve similar traits or capacities in response to similar environmental pressures.

Ideomotor action

An action that is covertly mimicked when another's action or movement of an object is being observed, such as moving your arm when you watch someone bowl or tilting your head in synchrony with a pendulum swing.

Motor imitation

Re-enactment by the observer of a target's motor movements or facial expressions.

Affordances

Motor or action properties of objects that are activated by their percepts, which are intrinsic to the mental representation of the object.

Simon effect

A behavioural effect in which motor actions are facilitated when they are consistent with the spatial location of the stimulus, and slowed when they are inconsistent or opposing the location.

Bottom-up

Describes a neural and mental process that is stimulus-driven on the basis of directly observed information without requiring explicit cognitive processes or capacities. Empathy arises from bottom-up processes that are shared across species such as motor mimicry, emotional contagion and state matching.

Top-down

Describes a neural and mental process that requires a conscious, cognitive evaluation to take into account information that is not directly observable, such as taking another's perspective or reasoning about their state on the basis of conceptual knowledge. These processes participate in more advanced forms of empathy that are more sophisticated in humans and with age, but they are not required to simply understand how someone feels.

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de Waal, F., Preston, S. Mammalian empathy: behavioural manifestations and neural basis. Nat Rev Neurosci 18, 498–509 (2017). https://doi.org/10.1038/nrn.2017.72

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