There is disagreement in the literature about the exact nature of the phenomenon of empathy. There are emotional, cogni- tive, and conditioning views, applying in varying degrees across species. An adequate description of the ultimate and proximate mecha- nism can integrate these views. Proximately, the perception of an object's state activates the subject's corresponding representations, which in turn activate somatic and autonomic responses. This mechanism supports basic behaviors (e.g., alarm, social facilitation, vicar- iousness of emotions, mother-infant responsiveness, and the modeling of competitors and predators) that are crucial for the reproduc- tive success of animals living in groups. The Perception-Action Model (PAM), together with an understanding of how representations change with experience, can explain the major empirical effects in the literature (similarity, familiarity, past experience, explicit teach- ing, and salience). It can also predict a variety of empathy disorders. The interaction between the PAM and prefrontal functioning can also explain different levels of empathy across species and age groups. This view can advance our evolutionary understanding of empa- thy beyond inclusive fitness and reciprocal altruism and can explain different levels of empathy across individuals, species, stages of de- velopment, and situations.

/pdf/empathy-its-ultimate-and-proximate-bases-3gnr9okjki.pdf

Empathy: Its ultimate and proximate bases.

Here we provide a review of the animal and human literature concerning the role of the amygdala in fear conditioning, considering its potential influence over autonomic and hormonal changes, motor behavior and attentional processes. A stimulus that predicts an aversive outcome will change neural transmission in the amygdala to produce the somatic, autonomic and endocrine signs of fear, as well as increased attention to that stimulus. It is now clear that the amygdala is also involved in learning about positively valenced stimuli as well as spatial and motor learning and this review strives to integrate this additional information. A review of available studies examining the human amygdala covers both lesion and electrical stimulation studies as well as the most recent functional neuroimaging studies. Where appropriate, we attempt to integrate basic information on normal amygdala function with our current understanding of psychiatric disorders, including pathological anxiety.

The amygdala: vigilance and emotion

The mentalizing (theory of mind) system of the brain is probably in operation from ca. 18 months of age, allowing implicit attribution of intentions and other mental states. Between the ages of 4 and 6 years explicit mentalizing becomes possible, and from this age children are able to explain the misleading reasons that have given rise to a false belief. Neuroimaging studies of mentalizing have so far only been carried out in adults. They reveal a system with three components consistently activated during both implicit and explicit mentalizing tasks: medial prefrontal cortex (MPFC), temporal poles and posterior superior temporal sulcus (STS). The functions of these components can be elucidated, to some extent, from their role in other tasks used in neuroimaging studies. Thus, the MPFC region is probably the basis of the decoupling mechanism that distinguishes mental state representations from physical state representations; the STS region is probably the basis of the detection of agency, and the temporal poles might be involved in access to social knowledge in the form of scripts. The activation of these components in concert appears to be critical to mentalizing.

/pdf/development-and-neurophysiology-of-mentalizing-3mxlcttihw.pdf

Development and neurophysiology of mentalizing.

https://www.ece.uvic.ca/~bctill/papers/learning/Kringelbach_Rolls_2004.pdf

The functional neuroanatomy of the human orbitofrontal cortex: evidence from neuroimaging and neuropsychology

Neural systems for recognizing emotion.

Face processing impairments after encephalitis: amygdala damage and recognition of fear.

https://www.mrc-cbu.cam.ac.uk/personal/rik.henson/personal/MaratosEtAl_N_01.pdf

Neural activity associated with episodic memory for emotional context.

http://people.uncw.edu/hakanr/documents/emotionwordcohesion.pdf

Recognition memory for emotionally negative and neutral words: An ERP study

In two experiments, words were presented in negatively or neutrally valenced sentences. At test, subjects made old/new recognition judgments to these words. In Experiment 2 only, for words judged old, subjects also indicated whether the words had been studied in a neutral or a negative context. In Experiment 1, left parietal old/new event-related brain potential (ERP) effects were larger and more sustained when elicited by words that had been studied in negative sentences, and a right frontal old/new effect was elicited by these words exclusively. In Experiment 2, the left parietal and right frontal effects elicited by old words correctly assigned to their study context were equivalent in size regardless of the nature of the context; a third ERP old/new effect, maximal over posterior scalp regions, was seen only for words from negative contexts. The findings indicate that incidental retrieval of emotional context gives rise to greater activation in neural systems supporting conscious recollection than does retrieval of nonemotional context. When contextual retrieval is intentional, recollection of emotional and non-emotional information are associated with equivalent engagement of these systems. The findings from Experiment 2 suggest that additional neural circuitry may be activated selectively by emotionally valenced episodic information.

Elizabeth J. Maratos

Papers

Face processing impairments after encephalitis: amygdala damage and recognition of fear.

Neural activity associated with episodic memory for emotional context.

Recognition memory for emotionally negative and neutral words: An ERP study

Electrophysiological Correlates of the Retrieval of Emotional and Non-emotional Context