Empathy development throughout the lifespan

Abstract

Empathy is a cornerstone of social interaction from animals to humans. In this chapter we attempt to map the field of empathy research by discussing the different definitions of empathy and those of related concepts. In addition, we synthesize findings from social psychology, developmental psychology, neuroscience, and genetics into a coherent understanding of empathy throughout the lifespan, taking into account each field's methodological considerations, which influence interpretation of findings.

Full text

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Empathy development throughout the lifespan
Uzefovsky Florina1 & Knafo-Noam Ariel2
1Autism Research Centre, Department of Psychiatry, University of Cambridge, Cambridge, UK
2Psychology Department, The Hebrew University of Jerusalem, Israel
Abstract (70-100 words)
Empathy is a cornerstone of social interaction from animals to humans. In this chapter we attempt
to map the field of empathy research by discussing the different definitions of empathy and those of
related concepts. In addition, we synthesize findings from social psychology, developmental
psychology, neuroscience, and genetics into a coherent understanding of empathy throughout the
lifespan, taking into account each field’s methodological considerations, which influence
interpretation of findings.

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From the moment we enter the world as babies, we are dependent on others. This dependence
changes as we grow, but constantly we are reliant on others to fulfil our survival needs; from being
fed and held for physical and mental warmth when we are born to the need to be understood and
connected to others as we get older. At the outset, a caregiver’s ability to understand what the baby
needs, and later wants, is crucial for survival and for development. For human societies (as well as
for other animals) effective communication is the basis for any form of social living, and although
humans have a unique ability to communicate using words, much of our communication is not
verbal (Knapp, Hall, & Horgan, 2013). We express ourselves using facial expressions, body gestures,
and tone of voice – what we feel, what our true intentions are, what we desire and what we would
like to avoid. Much more outstanding is our ability to understand all these subtle cues automatically
and almost effortlessly. Not only survival, but also wellbeing is largely dependent on the ability to
communicate with others.
Theoretical and empirical research has, for many years, attempted to define and measure empathy.
Different disciplines such as social psychology, neuroscience and developmental psychology have
provided slightly different answers based on the different methodologies and theoretical
frameworks in each field. We will begin by presenting a definition of empathy, and discuss how it
has been measured in adulthood. We will then go on to present definitions of other concepts that
are closely related to empathy, and have been used in various fields of research. In the second part
of this chapter we discuss empathy measurement in childhood and its association with our
theoretical understanding of empathy. We distinguish between empathy in childhood and adulthood
not because they are essentially different concepts, but because they have been studied in separate
scientific literatures, often using very different methods. In the third section we will review research
into the later development of empathy, from adolescence into adulthood and old age. In the fourth
section we will review the neurobiological and genetic basis of empathy and how findings from these
field contribute to our understanding of the concept of empathy and its development.

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What is empathy?
Empathy in adults
In social psychology empathy has often been construed in terms of both a state and a trait. Some
research has focused on measuring empathy using questionnaires, and others manipulated different
factors such as interpersonal similarity, emotion intensity, and deservedness to measure its effects
on subsequent behaviours such as prosocial behaviour (Batson et al., 1988; Feather & Sherman,
2002; Greitemeyer, Osswald, & Brauer, 2010).
Empathy is a multifaceted concept with cognitive and emotional components. Adam Smith (1759)
was among the first to describe an automatic response of emotional resonance with another person
and distinguished that from the understanding of another's feelings, devoid of any emotional
response. More recent theorists have debated the centrality of one component over the other in
empathy. Some argue that the emotional component, or the immediate experience of another’s
emotion, is the most central aspect of empathy (e.g. Mehrabian & Epstein, 1972), while others
suggest that the cognitive aspect, defined as an intellectual experience of another’s experience, is
the most salient (e.g., Kohut, 1971). Yet others posit that both the cognitive and the emotional
aspects cannot be separated and both faces of empathy are integral parts to its understanding (e.g.,
Hoffman, 1977).
Based on this past research we define empathy here as the ability to recognize the emotions of
others and to share in those emotions while maintaining a self-other distinction (Davis, 1983;
Eisenberg & Eggum, 2009; Shamay-Tsoory, 2011).
Empathy types and measurement
Empathy can be seen as a process which reflects two principal components: a cognitive component,
namely, the ability to accurately recognize and understand what others feel. This component is
termed Cognitive empathy (CE) and is often measured in adults using self-report questionnaires, or

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by showing pictures of facial expression that participants are asked to label with mental state words
(e.g., Baron-Cohen, Wheelwright, Hill, Raste, & Plumb, 2001; Davis, 1983; Dziobek et al., 2008;
Reniers, Corcoran, Drake, Shryane, & Völlm, 2010). In both cases, the focus is usually placed on the
accuracy of emotion recognition.
The second component is an affective component, i.e. the ability to share the emotions of another,
while maintaining a self-other distinction (Davis, 1983; Zahn-Waxler, Radke-Yarrow, Wagner &
Chapman, 1992). This component is termed Emotional empathy (EE) and is more difficult to measure
than CE. EE is defined as an experience and therefore is not directly related to a specific observable
behaviour or state. Most often EE is measured using self-report questionnaires or self-reported
affect experienced due to perceiving others' emotions or situations (Davis, 1983; Dziobek et al.,
2008; Reniers et al., 2010). Another way of indirectly measuring EE is using psychophysiological or
neuroimaging measures. Several studies examined the association between observing emotionally
laden situations or pictures and indices of arousal such as heart rate or skin conductance. The
advantage of these measures is that they do not rely on self-report. On the other hand, the
association between these measures and specific emotions is unclear. For example, sadness or
concern for the other might evoke a decrease in heart rate, but low heart rate could also be a sign of
being unaffected by the other’s condition (Zahn-Waxler, Cole, Welsh, & Fox, 1995). There is not a
one to one association between a physical response and experience, which begs a careful approach
to interpretation of psychophysiological findings, especially when considering individual differences.
Another psychophysiological measure, is the contraction of specific facial muscles that were
associated with facial expressions of sadness or joy (sometimes termed motor empathy). These are
measured using facial electromyography (fEMG) and their association with specific emotions is more
straightforward. One interesting study examined the association between fEMG and state and trait
emotional and cognitive empathy in 379 adolescents (age range 15-20; Van der Graaff, Meeus, et al.,
2015). The study showed that fEMG responses to happy video clips was associated with state
affective empathy (measured as a self-report response to the video clips), which in turn predicted

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higher cognitive state empathy. For sadness, fEMG responses similarly predicted state affective
empathy and state cognitive empathy, indirectly. In addition, affective state empathy was predictive
of affective trait empathy, but not cognitive trait empathy, which was predicted by cognitive state
empathy. To summarize, this study shows that fEMG is a valid measure of state empathy and is also
associated with trait empathy (note that the study reports interesting sex differences as well).
Related concepts
Many different concepts concerning how others' psychological states are perceived or interpreted
have been developed theoretically and studied empirically. Some of them overlap with empathy to
some extent. We present these concepts here to elucidate the similarities and differences between
them and empathy, in the hope of promoting a clearer understanding of empathy as well as these
other phenomena, by creating a glossary of the relevant concepts (see summary in Error! Reference
source not found.). It is important to note here regarding the overlap among cognitive empathy,
affective theory of mind (ToM) and perspective taking. Affective ToM is defined very similarly to
cognitive empathy (Shamay-Tsoory & Judith Aharon-Peretz, 2007; Shamay-Tsoory et al., 2007;
Shamay-Tsoory, Tibi-Elhanany, & Aharon-Peretz, 2006; Walter, 2012). The different names are
largely the product of the fact that they were researched by different disciplines. This in turn caused
methodological differences in the way affective ToM and cognitive empathy (CE) were measured.
Traditionally, affective ToM is measured using vignettes and comic strips where the participant is
asked to infer the emotion of the other using situational cues, while cognitive empathy is typically
measured using pictures of facial expressions, where the participant is asked to accurately name the
emotion depicted, or using questionnaires, where the participant is asked to report on her ability to
accurately perceive the emotions of others in everyday life.
Another important issue to clarify is the definitional association between empathy and prosocial
behaviour. Prosocial behaviour is defined as a behaviour that increases the benefit of the other and
may include different types of behaviours such as helping, sharing and comforting. Prosocial

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behaviour cannot be interpreted as an overt form of empathy or sympathy, because (1) the absence
of prosocial behaviour does not necessarily mean that empathy or sympathy were not felt and (2)
prosocial behaviour may be driven by other motivations such as conformity to norms or social
desirability. Notably, however, research has shown that empathy seems to motivate prosocial
behaviour even when other motivations (such as reduction of personal distress) are accounted for,
such when personal distress is reduced (Batson et al., 1988; Batson et al., 1991, and see recent
review in Decety, Ben-Ami Bartal, et al., in press).
-------Table 1 here -------
Empathy in Childhood
Traditionally, the concepts of emotion contagion, personal distress and empathic concern have been
studied primarily in childhood. Before describing the way they are measured, we will present the
three most influential definitions of empathy in developmental psychology. Similar to the definition
we have adopted in this chapter, Eisenberg defined empathy as an emotional response that stems
from the apprehension of another’s emotional state or condition, and that is congruent
with it (Eisenberg & Miller, 1987). According to Eisenberg, empathic arousal can lead to either
sympathy (and this, in turn, could lead to prosocial behaviour) or to personal distress, when one is
unable to regulate the empathic arousal (Eisenberg, Spinrad, & Sadovsky, 2006).
Hoffman defined empathy as the observer's vicarious affective response to another person
(Hoffman, 1977), focusing on the developmental stages of empathy. According to that view, an
empathic response entails affective arousal which is similar to that of the target both in quality and
in direction.

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Zahn-Waxler, on the other hand, focused on empathic concern, i.e., feelings of concern for another
in distress, and refers to the physical aspects of the response as well (Zahn-Waxler, Robinson, &
Emde, 1992).
Empathy measures in childhood. In early childhood empathy is primarily measured as the response
to a simulation of the other in distress (e.g., Knafo, Zahn-Waxler, et al.,, 2008; Zahn-Waxler, Radke-
Yarrow, et al., 1992; Zahn-Waxler, Robinson, & Emde, 1992). In a typical experiment, children view
an experimenter, or their mother, simulate pain (caused by bumping her knee/finger/elbow into
something). For the first 30 seconds a moderate intensity pain is simulated (whimpering, saying
“ouch, this hurts”, rubbing the hurt organ, etc.), for an additional 30 seconds low intensity pain is
simulated (“ouch”, rubbing the hurt organ, etc.). Afterwards the experimenter (or parent) returns to
baseline, showing the child that they are no longer in pain (“I am okay now”). Children are filmed
during this episode and their facial expressions and behaviours are coded on several indices. The
most relevant to our discussion are three codes, all usually scored on a scale of intensity (absent,
slight, moderate, substantial): (1) empathic concern – facial expressions (e.g., brow furrow), body
gestures (e.g. tension), vocalizations (e.g., “ouch”) and verbalizations (e.g., “are you okay?”)
expressing concern for the other in pain. (2) Hypothesis testing – attempts to understand the
situation as evident in gestures (e.g., looking interchangeably at the hurt organ, the simulator’s facial
expression and the hurting object), vocalizations (e.g., “huh?”, that is a sound with a questioning
intonation), and verbalizations (e.g., “are you okay”?). (3) Personal-distress – anxiety and fear, or
sadness as evident in facial expressions (e.g., grimacing, lips turned down), body gestures (e.g.
fidgeting, hand wringing), and vocalizations, including whimpering and crying. It is important to note
here that empathic distress is measured similarly for children and adults. Not so for the cognitive
measure, hypothesis taking. Although hypothesis taking is measured independently from empathic
concern in order to capture non-emotional attempts to understand the situation (i.e., “did you
stumble into the chair?” does not necessitate an emotional response, but rather an interest in
understanding what is happening), it is not the same as the definition of cognitive empathy in adults.

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Whereas in adults cognitive empathy captures primarily accuracy in understanding and recognizing
the emotions of others, in the case of hypothesis testing it captures primarily expressed interest in
the situation. It is possible that a child accurately understands the situation and the emotional
response of the experimenter/parent and therefore does not ask the questions or makes the
gestures that are coded as hypothesis testing. That is, hypothesis testing capture cognitive aspects of
empathy that are different to those that most studies in adult define as cognitive empathy.
Therefore, the interpretation of these studies with respect to lifetime cognitive empathy should be
done carefully.
Another way of measuring empathy in young children is by other-report, i.e., by parent or teacher.
Several scales have empathy related items in them but no distinction between emotional and
cognitive components was made, or distinction from prosocial behaviours. Recently, a questionnaire
was designed to assess empathy in toddlers and young children (validated on children 1-5 years old),
measuring three subscales: emotion contagion, attention to others’ feelings, and prosocial actions
(Rieffe, Ketelaar, & Wiefferink, 2010), based on Hoffman’s theory of empathy development
(Hoffman, 1975). The EmQue consists of 20 items rated on a 3-point scale (never/sometimes/often),
and has good concordant validity with the prosocial behaviour scale of the Strengths and Difficulties
Questionnaire (SDQ; Goodman, 1997) as well as children’s responses to pain simulations (as
described above), which were coded using the same three subscales (Rieffe et al., 2010). Another
scale, the Griffith Empathy Measure (GEM) has been developed for the measurement of empathy
across middle childhood and into adulthood (Dadds et al., 2008). This scale is based on the Bryant
empathy questionnaire (Bryant, 1982), which was initially designed as a self-report measure (see
below). The GEM measures both the cognitive and emotional aspects of empathy using 23 items
rated on a 9-point scale.
For older children, empathy has been measured using different tasks including emotion recognition
in face and voice (Nowicki & Duke, 1994), which primarily taps into cognitive empathy; similarly

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emotionally laden picture stories are used to either ask regarding the characters’ emotional
response (cognitive empathy) or the evoked emotional response in the child (emotion contagion;
Feshbach & Roe, 1968; Knafo et al., 2011). Additionally, psychophysiological measures, such as
different indices of heart rate and skin conductance, have been used to indirectly assess the
influence of another’s emotion on the autonomic nervous system. The specific patterns of
physiological responses are interpreted as reflecting either sympathy, self-distress or disregard and
callousness (Fabes, Eisenberg, & Eisenbud, 1993; Van Hulle et al., 2013; Zahn-Waxler et al., 1995).
As children grow older and are able to read, write and report on their experience, self-report
measures can be used. Thus, school aged children’s empathy can be measured with questionnaires
such as the Bryant questionnaire which is a simplified and shortened version of the Questionnaire
Measure of Emotional Empathy which was designed for adults (Mehrabian & Epstein, 1972). The
Bryant questionnaire consists of 22 yes/no items (Bryant, 1982) and was used with children and
adolescents. In recent years several other self-report measures were developed. For example the
Children's Empathic Attitudes Questionnaire (Funk, Fox, Chan, & Curtiss, 2008), which was based on
the Bryant questionnaire and the Interpersonal Reactivity Index (Davis, 1983), consists of 16 items
rated on a 3-point scale (yes/maybe/no), and has 3 subscales, mostly related to emotional empathy.
Importantly, the association between empathy as indexed by observational and questionnaire
measures and empathy as indexed by brain activation (see also section empathy in the brain) is not
clear or direct. Several studies find associations between the two types of measures (e.g., Schulte-
Rüther, Markowitsch, et al., 2008; Shamay-Tsoory, Aharon-Peretz, & Perry, 2009; Singer, Seymour,
et al., 2004). However, one study which was uniquely conducted in a sample of children aged 4-17,
showed that sex differences in reported affective empathy (measured by the Bryant questionnaire)
do not translate to sex differences in empathy-related brain activation (Michalska, Kinzler & Decety,
2013). This brings into focus the fact that the brain mechanism underlying empathy is still far from
understood, and that the inferences we make about empathy depend upon the type of

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measurement was used. Therefore, it is prudent to qualify such inferences based on the specific
characteristics of the measures used.
The development of empathy
The research into the development of empathy is highly dependent upon the available measures. In
the same way, theory dictates the design and application of the measures. This interdependency is
evident in the research and theory of empathy development. We will begin by describing a theory of
empathy development, as has been proposed by Hoffman (1982) and then discuss the bidirectional
implications of measurement and theory on the field.
Hoffman (1982) was influenced by cognitive theories of development and building on that suggested
that empathy, at its mature form, co-develops with other cognitive-based functions. Thus, empathy
development is seen as following a route going from purely emotional to cognitively controlled, from
an intuitive and automatic response to a complex process that relies on top-down control and
regulation.
According to Hoffman’s influential theory, self-other differentiation is the driving force behind the
maturation of empathic responses from early infancy through early childhood. According to this
theory, we are born with a rudimentary ability to feel others, but with no ability to distinguish
between self and other. For example, at this stage and as early as a few hours after birth, infants
respond in crying to the sound of another infant's cry (Sagi & Hoffman, 1976). This stage, which is
similar to emotional contagion, lasts, according to Hoffman, for most of the first year of life. That is,
emotional empathy appears developmentally earlier, and as a sense of self develops and higher
cognitive functions become active, cognitive empathy comes into play. Thus, during the second and
third year of life, children reach the ‘veridical empathic distress’ stage, at which they are able to
empathize with others even when they are not physically present, and later on (in adolescence) with
an entire, even abstract, group of people (Eisenberg, Fabes, & Spinrad, 1998).

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Based on this theory most of the research on empathy in childhood has focused on early childhood,
and empathy in infancy was rarely studied (Davidov, Zahn-Waxler, Roth-Hanania, & Knafo, 2013).
One of prerequisites for the ability to empathize with others in Hoffman’s theory is the existence of
self-knowledge, which supports self-other differentiation. Research suggests that although explicit
self-knowledge may develop later in life, implicit self-knowledge, which is based on infants’ sensory
perception and actions in the world exists from birth and perhaps even before (Castiello et al., 2010;
Rochat & Striano, 2000). Indeed, a recent study measured empathy in response to a simulation of
distress in infants aged 8- to 16-months (Roth-Hanania, Davidov, & Zahn-Waxler, 2011). The
prediction stemming from Hoffman’s theory is that infants during the first year of life faced with the
distress of others would exhibit self-distress, yet self-distress was very rare. On the other hand,
infants showed moderate levels of empathic concern as well as hypothesis testing, a cognitive
component that reflects attempts to understand the distress of the other and the cause of the
distress. Thus, the study provided a proof-of-concept that both emotional and cognitive components
of empathy manifest already during the first year of life. Importantly, these predicted prosocial
behaviour in the second year of life, affirming that the responses coded during the first year of life
reflect stable empathic tendencies. Similarly, another study examined three-, six-, and nine-month
old infants’ response to an almost 3-minute audio recording of another’s infant crying in pain
(Geangu, Benga, Stahl, & Striano, 2011). Importantly, infants did not start crying in pain immediately
when presented with the stimulus, and self-soothing behaviours in three months olds were related
to greater latency until crying commenced.
Davidov and colleagues (2013) suggest that the limiting factor for expressing an empathic response
is not self-other differentiation, but the ability to self-regulate the emotional arousal that is brought
on by witnessing another’s distress. Thus, it is the developmental stage of self-regulatory abilities
that will dictate to what stimulus an infant will respond with empathic concern versus personal-
distress (Davidov et al., 2013).

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So far we have discussed the development of empathy as a single factor, however the development
of emotional and cognitive empathy seems to follow different trajectories. As described above,
emotional empathy appears very early on, and remains stable or increases only slightly during the
second year of life, while cognitive aspects of empathy, as well as prosocial behaviour, increase over
the second and third year of life (reviewed in Davidov et al., 2013). The same pattern was identified
using the GEM in a cross sectional study of 2612 children aged 4 to 16; whereas cognitive empathy
increased with age, there was no association between age and emotional empathy (Dadds et al.,
2008). Finally, a longitudinal study of 14-17 year old adolescents' self-reported empathy found that
empathic concern was highly stable, substantially more so than perspective taking. Moreover,
empathic concern longitudinally predicted changes in perspective taking, and not the other way
around, which suggests that the affective component of empathy develops earlier than, and perhaps
contributes to, the cognitive aspect of empathy (Van Lissa et al., 2014).
Nevertheless, studies of eye-tracking with young infants suggest that infants have some cognitive
empathy abilities from early on as well, as they are able to discern between different facial
emotional expressions. For example, four to six months olds show discrimination between different
basic emotions as measured by visual fixation, and by their behaviour when observing facial
expressions, measured as approach vs. avoidant movements and gestures (Serrano, 1995). Similarly,
four and seven month old infants tended to use avoidant looking patterns at faces expressing
threatening emotions (angry and fearful) when compared with other emotions (happy sad and
neutral; Hunnius, de Wit, Vrins, & von Hofsten, 2010).
It is important here to note that cognitive empathy has also been studied as part of the research on
affective Theory of Mind (ToM), as the definitions of both concepts are very similar. Therefore ToM
studies can inform our understanding of cognitive empathy development. Theory of Mind was
thought to develop much later than empathy, at around age 4, yet recent research with young
infants suggests that rudimentary forms of ToM appear much earlier, during the second year of life.

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These studies use longer looking time, a measure of surprise, as well as other implicit measures to
investigate infants’ false-belief and desire understanding (reviewed in Baillargeon, Scott, & He, 2010;
Slaughter, 2015). In addition, more complex forms of ToM, including affective ToM, differ across
individuals and develop into middle childhood. Moreover, they are related to more basic forms of
ToM, such as false belief tasks (reviewed in Hughes & Devine, 2015), suggesting an association
between ToM in all its forms and empathy.
This distinction between the developmental trajectories of emotional and cognitive aspects of
empathy is in line with one of the most influential theories of empathy evolution, proposed by
Preston and de Waal (2003). This theory suggests that empathy evolved from a coupling between
representations of perception and action, and turned to a coupling of the perceived emotions of
others and our own experience of the same emotions. Thus, emotional aspects of empathy are
considered to be early appearing in evolution and related to the function limbic brain areas. Later on
in evolution, the cognitive aspects of empathy developed, allowing to control our response to
others’ emotional expression. Moreover, they theory suggests that from a developmental
perspective, the default is an automatic response to others' emotions, which is later on moderated
through development of knowledge and regulatory mechanisms (such as the possibility of
responding differently to the plight of friends and foes). The end result of this developmental
process is a relatively governed response to others that involves both emotional and cognitive
components (Preston & De Waal, 2002).
From adolescence to adulthood
We know very little about the course of empathy development from childhood into adolescence, yet
slightly more is known about the development of empathy from adolescence into adulthood. For
example, one recent study investigated empathy in a large sample (N=2,054) of adolescents,
measuring empathy (using an eight item scale; no distinction between cognitive and emotional
empathy) yearly from 12 to 16 years of age. The same individuals reported on their empathy again

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(using a three item scale) at age 35 (N=1,527). Findings from this study showed that empathy
increases throughout adolescence, and higher empathy in adolescence predicts higher empathy at
35, as well as higher communicative skills. In addition, those who reported on increases in empathy
during adolescence also perceived themselves as being more empathic and better socially integrated
than adolescents whose empathy scores decreased during adolescence (Allemand, Steiger, & Fend,
2015). Similarly, another study measured cognitive empathy using the facial scale from the
Cambridge Mindreading Face-Voice Battery, wherein participants are asked to infer mental states
from dynamic facial and body expressions. Participants in this study were adolescents and young
adults (12-23 year olds), and findings showed a positive correlation between performance on this
task and age, possibly at least partially explained by age-related increases in inhibition abilities,
which is a component of executive function (Vetter, Altgassen, Phillips, Mahy, & Kliegel, 2013).
Another interesting study examined empathy longitudinally in 467 Dutch adolescents (Van Lissa,
Hawk, et al., 2014). The adolescents reported on their empathic concern and perspective taking
yearly, for six years, starting from a mean age of 13. Using a cluster analysis approach, the authors
showed that adolescents clustered into three groups of low, medium and high empathy (combining
empathic concern and perspective taking). The classes were primarily differentiated by an increase
in perspective taking for the high and medium empathy groups and a decrease for the low empathy
group throughout adolescence. In addition, the low empathy group showed a temporary decrease in
empathic concern around age 16. Additionally, the low empathy group was associated with higher
conflict with parents, suggesting a familial effect (whether the association between family conflict
and low empathy is causal in any direction, or a genetic link can explain both conflict and low
empathy is yet to be determined). This study emphasizes the importance of investigating
developmental trajectories of empathy, as the effects may not be linear.
Few studies examined associations between age and empathy across adulthood in cross sectional
samples. Some studies found no association with age (e.g., Eysenck, Pearson, Easting, & Allsopp,
1985) and others report of a negative association. For example, one study examined the association

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between age and empathy in a large sample (N=1,567) of adult individuals aged 22-92 years, with
and without physical disabilities (Schieman & Gundy, 2000). Empathy was measured using 8 items
that tap into primarily emotional and motivational aspects of empathy. Findings show a negative
association between age and empathy. This negative association is explained by an age-associated
decrease in income and education, an increase in widowhood status (as well as retirement, but to a
lesser extent), increase in physical impairment (primarily through effects on mastery). In addition,
age associated differences in personal attributes such as mastery, and introspectiveness. These
negative effects were found to be ameliorated by positive relationships and religiosity. Conversely,
another study used a similar age group (18-90 year olds) to investigate empathy across the life span
in three different cohorts (O’Brien, Konrath, Grühn, & Hagen, 2013). This study used two of the
subscales of the IRI (Davis, 1983) to measure empathic concern (emotional empathy) and
perspective taking (cognitive empathy) in a total of 75,263 participants aged 18-90 years old.
Empathic concern and perspective taking both showed increase with age, yet a quadratic effect of
age was also observed, with an increase in empathy until middle-age, followed by a decrease
empathy.
When considering cross sectional research one must take into account that the effects of age are
intermixed with cohort effects. That is, the effect of being born and raised in different social
climates. A study using a longitudinal assessment may be able to answer this question. Indeed the
negative association between empathy and age is supported by one such longitudinal study (Helson,
Jones, & Kwan, 2002); empathy was assessed four times throughout adulthood in two different
samples. Empathy was measured by a subscale of the California Psychological Inventory (CPI; Gough
& Bradley, 1996). In this study as well a negative correlation between age and empathy emerged.
However, another longitudinal study provides a different answer. Grühn and colleagues examined
data from a longitudinal study of individuals aged 10-87 who were followed up for 12 years.
Empathy was again measured using the CPI at four time points. Examination of the cross sectional
effects revealed a negative correlation between age and empathy, yet no correlation between age

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and empathy was found for the longitudinal analysis, suggesting that the observed negative
association is the product of cohort effects and not of age (Grühn, Rebucal, Diehl, Lumley, &
Labouvie-Vief, 2008). This view is further supported by a meta-analysis of self-reported empathy, as
measured by the IRI (Davis, 1983) in college students between 1979 and 2009 (Konrath, O'Brien, &
Hsing, 2011). The meta-analysis included 72 samples representing a total of 13,737 participants. The
results suggest that college students from later birth-cohorts report having lower empathic concern
and, to a lesser extent, lower perspective taking, than college students from earlier birth-cohorts.
These findings could be the result of changes in reporting tendencies over the years rather than a
true decline in levels of empathy. Taken together, most studies reviewed here suggest a cohort
effect, but its influence seems to be in opposite direction to that suggested by the findings of the
meta-analysis. The cross-sectional studies suggest that earlier-born individuals tend to report having
lower empathy than later born-individuals, yet the meta-analysis suggests the opposite. Further
research is needed to elucidate the question of age effects in adulthood, but it seems that both age
and cohort effects are at play.
Biological basis of empathy
Empathy in the brain
The theoretical understanding of what empathy is can be enriched by understanding the
mechanisms by which empathic feelings and responses come to be. In recent years, neurobiological
studies of empathy provided some insight into the mechanism of empathy across development. It is
important to note here that most studies have been conducted with adults.
According to the above definition, emotional empathy would entail some correspondence (although
not complete) between the experience of the other and of oneself. Therefore, the human Mirror
Neuron System (hMNS) has been suggested as a mechanism for the phenomenon of emotional

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empathy (Gallese, Keysers, & Rizzolatti, 2004; Iacoboni & Dapretto, 2006; Oberman, Pineda, &
Ramachandran, 2007, but see Decety, 2011). The hMNS has been proposed to include brain areas
that are activated both when we experience a certain emotion and when we perceive others
experiencing the same emotion. For example, the insula is activated both when one feels disgust and
when one is shown a disgusted face (Phillips et al., 1997; Wicker et al., 2003). The same is true for
the perception of other’s pain, which activates the pain matrix (Corradi-Dell'Acqua, Hofstetter, &
Vuilleumier, 2011; Jackson, Rainville, & Decety, 2006; Singer et al., 2004, see a meta-analysis in
Lamm, Decety, & Singer, 2011) and for the perception of other’s fear which activates the amygdala
(Whalen et al., 1998). A similar activation pattern was observed for 7-12 year old children who
observed others experiencing pain (Decety, Michalska, & Akitsuki, 2008). Moreover, the work done
by Singer and colleagues showed that reported levels of emotional empathy were positively
correlated with activation of the ACC and left anterior insula in response to observed pain (Singer et
al., 2004). That is, a connection was found between experienced emotional empathy and brain
structures that are thought to be involved in emotional empathy.
Although the above evidence supports the idea that overlapping brain regions are activated when
experiencing distress and when perceiving another’s distress, there is less evidence for a neuron-
level overlap in activation. Recent studies which examined these activation patterns using higher
spatial resolution methods show that the overlap between areas activated for self vs. other
experience do not completely overlap (Lamm et al., 2011). That is, different neurons, though
spatially close, react to self vs. other experience. This might mean that the hypothesized mirror
properties are incomplete or absent. It might also mean that nearby areas of emotion processing
interact closely to process emotionally valenced stimuli. Moreover, studies conducted with
psychopaths (Cheng, Hung, & Decety, 2012; Decety, Lewis, & Cowell, 2015; Fecteau, Pascual-Leone,
& Théoret, 2008), a condition characterized by deficits in emotional empathy, suggest increased
activation in the so-called ‘mirror’ areas. Similar findings are reported when comparing individuals
with very high psychopathic traits and very low psychopathic traits (both groups were drawn from

18
the typical population; Marcoux, Michon, et al., 2013). These findings are contradictory to the
hypothesis that these brain areas underlie emotional empathy. Similarly, autism is a condition
characterized by deficits in cognitive empathy and an apparently intact emotional empathy, yet
studies find decreased activation in the mirror system as compared to controls (e.g., Bernier,
Dawson, et al., 2007; Oberman, Hubbard, et al., 2005).
On the other hand, a larger consensus surrounds the properties of brain areas that are activated
when participants are specifically asked to think of others’ emotions. Studies which specifically
looked at cognitive empathy have emphasized the role of the ventromedial prefrontal cortex
(vmPFC; Hynes, Baird, & Grafton, 2006; Shamay-Tsoory, Aharon-Peretz, & Perry, 2009), as well as
the temporoparietal junction (TPJ; Saxe & Kanwisher, 2003; Saxe & Wexler, 2005) in understanding
others’ emotions. Interestingly, a study which compared adolescents (11-16 year olds) to adults (24-
40 year olds) found that the vmPFC is activated to a greater extent in adolescents as compared to
adults when preforming an affective ToM task (Sebastian et al., 2012)
A double dissociation of areas involved in emotional and cognitive empathy is further supported by
the extant lesions studies. In one study, Shamay-Tsoory and colleagues compared empathy deficits
in patients with lesions in the vmPFC and patients with lesions in the Inferior Frontal Gyrus (IFG, an
area considered as part of the hMNS). Patients with vmPFC lesions exhibited lower cognitive, but not
lower emotional empathy, while patients with IFG lesions exhibited lower emotional, but not
cognitive, empathy (Shamay-Tsoory et al., 2009). Other studies by the same group repeatedly
showed that patients with vmPFC lesions had specific difficulties in CE tasks (Shamay-Tsoory &
Aharon-Peretz, 2007; Shamay-Tsoory, Tomer, Berger, Goldsher, & Aharon-Peretz, 2005). Another
way of investigating the mechanism of emotional and cognitive empathy is by investigating dynamic
patterns of activation in the brain. One such study found that emotional empathy was associated
with stronger within-network functional connectivity in social-emotional networks, while cognitive
empathy was associated with stronger within-network functional connectivity in social-cognitive and

19
interoceptive networks (Cox et al., 2011). These type of studies add insight regarding the brain
dynamics of empathic response, of which we know very little.
It is important to note, that though there may be distinct areas in the brain that are associated with
emotional or cognitive empathy, it is also clear that normally these two networks work together. In
typical situations and for typical individuals, seeing another in distress would evoke ‘mirror’ brain
activation as well as cognitive appraisal mechanisms involved in better understanding the situation
and its meaning. Similarly, recognizing that one is upset through a descriptive story would evoke
cognitive appraisal mechanisms, and often an emotional response would be evoked simultaneously.
Indeed, functional imaging studies that analysed brain activation in response to CE evoking stimuli
compared with cognitive ToM evoking stimuli (as opposed to the studies comparing cognitive and
emotional empathy that were described previously) find activation in emotion processing areas of
the brain such as the insula and the amygdala (Schnell, Bluschke, Konradt, & Walter, 2011; Völlm et
al., 2006). That is, functional imaging studies typically compare between two situations (i.e., task
and control) and the resulting pattern of activation is the product of a comparison between the two
conditions, such that comparisons against different control conditions would elicit different patterns
of activation. This emphasizes the importance of understanding the comparison stimuli used in each
study for the interpretation of results.
As described earlier, the development of emotional and cognitive empathy follows different
trajectories, and the interaction between the emotional and cognitive empathy systems may change
with age. This view is supported by a unique imaging study of individuals across a large
developmental range, 7-40 years old (Decety & Michalska, 2010). In this study, participants viewed
two types of videos. The first type of videos depicted individuals harmed unintentionally, a situation
that was thought to elicit empathy. The second type of videos depicted individuals being
intentionally harmed, a situation that was thought to elicit sympathy and a motivation to help the
injured other. Younger participants rated the situations as more painful overall, and these ratings

20
were associated with increased activation of the amygdala, posterior insula, and somatosensory
cortex. The pattern of activation was associated with age in a way that supports the theoretical and
behavioural observation of emotional aspects of empathy developing earlier than cognitive-
evaluative aspects (Decety & Michalska, 2010; Knafo et al., 2008). An additional study from the same
group (Cheng, Chen, & Decety, 2014) investigated the association between empathy facets and
electroencephalographic (EEG) activity. The study evaluated brain activity in response to pictures of
limbs in painful situations in both children (N=57, 2-9 year olds) and adults (N=15, 23-25 year olds).
Several indices were examined in this study. An early automatic component (EAC, N200), as
reflecting empathic arousal, a late positive potential (LPP) as reflecting processes of cognitive
appraisal, and mu suppression as reflecting the activity of the mirror network. All these indices
showed age-related differences, so that EAC decreases with age, the LPP increased with and mu
suppression was stronger for the pain stimulus only in adults but not in children. This study again
suggests that empathic arousal (an affective component) is present early in life, and cognitive
processes develop later in life.
Importantly, the earlier development of emotional empathy does not mean that later in life, once
both facets are fully developed, the first must always be utilized before the other, or take
precedence over the other facet. It does probably mean that the two facets may activate one
another. Thus, for example, when we hear about a natural disaster in another part of the world we
may first think about what it must be like to lose your home or loved ones (cognitive empathy) and
only then may feel for their pain (emotional empathy).
So which area in the brain acts as the relay station? There is evidence to support the role of the TPJ
as connecting the emotional empathy and the cognitive empathy networks as it is close to the
inferior parietal lobule (IPL; Shamay-Tsoory, 2011; Van Overwalle & Baetens, 2009). This view is
based mainly on studies of ToM (often with no distinction of cognitive and affective ToM), where
activation of the TPJ is often observed. Others point to the possible role of the insula (specifically

21
the anterior insula) as the region in the brain where cognitive and emotional empathy meet, where
the cognitive appraisal of information is relayed to the emotional centres of the brain (Gu, Liu, Van
Dam, Hof, & Fan, 2012). Indeed, in a review of studies which employed imaging techniques to study
emotion processing in the brain it was found that 60% of all studies reported insula activation (Phan,
Wager, Taylor, & Liberzon, 2004). See a schematic representation of the process in
.
------Figure 1 here------
Empathy genetics
Genetics provide another avenue for research on the structure of empathy. Quantitative genetic
studies compare the similarity in a certain trait of identical (monozygotic; MZ) twins to the similarity
of non-identical (dizygotic; DZ) twins, who share 50% of their DNA. Higher similarity between MZ as
compared to DZ twins suggests that genetic factors contribute to individual differences in the
examined trait. Additional sources of variance are the shared environment (SE; that makes twins
more similar to each other) and the non-shared environment (NSE: that makes twins less similar to
each other and includes a component of error; Plomin, DeFries, Knopik, & Neiderheiser, 2013).
Only four studies, that we are aware of, examined the heritability of empathy in adults (heritability is
an estimate of the relative contribution of genetic factors to individual differences in a certain

22
population and context). One study measured empathy using a subscale of the Temperament and
Character Inventory (TCI; Cloninger, Przybeck, & Svrakic, 1994) in a sample of 617 twins aged 15-30
years old (Ando et al., 2004). The study reported similar correlations for MZ and DZ twins (r=.29 and
r=.24, respectively) suggesting no genetic contribution to individual differences in empathy.
However other studies arrived at different conclusions. The first study measured empathic concern
in 42-57 year old twins using 13 items (e.g. sympathetic, kind) selected from the Adjective Check List
(ACL; Gough & Heilbrun, 1965) as reflecting empathic concern (Matthews, Batson, Horn, &
Rosenman, 1981). The heritability estimates stemming from this study were high (72%). Another
study used the Questionnaire Measure of Emotional Empathy (Mehrabian & Epstein, 1972) in 19 to
over 60 year old twins and estimated that genes explain 68% of the variance in empathy (Rushton,
Fulker, Neale, Nias, & Eysenck, 1986). The third study (Davis, Luce, & Kraus, 1994) examined three
facets of empathy: empathic concern, personal distress and perspective taking. Scores on these
scales were created by selecting 28 adjectives that were judged as reflecting at least one of the three
empathy facets. This study reported heritability estimates of 28%, 32%, and 20% for empathic
concern, personal distress and perspective taking, respectively. This last study shows that different
empathy facets may be influenced by genetic factors to a different magnitude, or even by different
genetic effects, although this last option was not examined in this paper.
Studies of empathy heritability in children show similar effects. The studies used observational
measures of empathy (as described above; Knafo, Zahn-Waxler, et al., 2008, 2009; Volbrecht,
Lemery-Chalfant, et al., 2007) with heritability increasing with age, and different estimates of the
contribution of genetic and environmental effects for emotional and cognitive empathy (when these
were examined separately).
Further evidence for the distinction between emotional and cognitive aspects of empathy comes
from molecular genetic studies of empathy. Not many studies have investigated the genetics of
empathy, and of those that did only few examined emotional and cognitive aspects separately. The

23
first ever study to examine an empathy component was a study by Rodrigues and colleagues (2009).
These authors examined the association between the rs53576 Single Nucleotide Polymorphism
(SNP) in oxytocin receptor (OXTR) gene and general empathy as measured by the IRI (Davis, 1983)
and emotion recognition (part of cognitive empathy) as measured by Reading the Mind in the Eyes
Test (Baron-Cohen et al., 2001). The study found that carriers of the A allele tended to have lower
scores on both measures (Rodrigues, Saslow, Garcia, John, & Keltner, 2009). Another study
examined the association between emotional and cognitive empathy and the same SNP in the OXTR
in addition to a polymorphic repeat region in the promotor of the vasopressin receptor 1a (AVPR1a)
gene. The findings showed that the rs53576 OXTR SNP was associated with emotional but not
cognitive empathy, whereas the AVPR1a repeat polymorphism was associated with cognitive but not
emotional empathy (Uzefovsky et al., 2015). These findings seem to be somewhat at odds with the
former study and this brings into focus the question of measurement. The former study measured
emotion recognition in a task which asks participants to infer mental states from pictures of the eye
area only, while the latter used both cognitive subscales if the IRI. These are very different
measurements of cognitive empathy. In addition, it is possible that participants rely, at least to some
extent, on emotional resonance, or emotional empathy, in order to infer mental states from images,
due to the unavailability of other cues in the task. On the other hand this also supports the notion
that for normal functioning, people rely on both emotional and cognitive processes in order to
connect with others.
Supporting the role of the rs53576 OXTR SNP in emotional empathy is a study that used the IRI as
well as measures of electrodermal activity of participants while watching videos of violent
encounters (mixed martial arts; MMA). Carriers of the A allele had lower empathic concern scores,
and had a less intense skin conductance response to watching the violent movies (Smith, Porges,
Norman, Connelly, & Decety, 2014). In this case, although two very different measures were used,
the findings were similar. In a similar vein, a recent study (Christ, Carlo, & Stoltenberg, 2015) report

24
an association between another SNP in the OXTR (rs2268498) and empathic concern, but not
perspective taking as measured by the IRI.
An additional recent study reported an association between emotional but not cognitive empathy,
as measured by the IRI, with the rs28373064 SNP in the gene coding for the Vasopressin 1b Receptor
(AVPR1b) in a Han-Chinese population (Wu, Shang, & Su, 2015).
Another system that was investigated in relation to emotional and cognitive empathy is the
dopaminergic system. One study examined, in a population of Han Chinese university students, the
dopamine beta-hydroxylase (DBH) gene which codes for an enzyme that converts dopamine to
norepinephrine. Out of two examined polymorphisms, one SNP was found to be associated with
both the RMET and the empathic concern subscale of the IRI, and the other polymorphism, an
insertion/deletion, was associated with the perspective taking subscale of the IRI (Gong, Liu, Li, &
Zhou, 2014). One additional study of particular interest here examined both emotional and cognitive
empathy with relation to a polymorphic repeat region in exon 3 of the Dopamine D4 Receptor
(DRD4). This study found no association between the gene and emotional empathy, but found an
effect of a DRD4 by sex interaction on cognitive empathy, so that male carriers of the 7-repeat allele
scored lower than female carriers of the same allele (Uzefovsky et al., 2014; Study 1 partially
overlapping withUzefovsky et al., 2015). This finding may mean that either gene by gene (epistasis)
or gene by environment interaction (GxE) effects contribute to the individual differences in empathy.
The first option would suggest that DRD4 interacts with sex genes to affect empathy, the second
option would suggest two possible interpretations of the environmental factor, either the internal,
hormonal environment, or the social environment. According to the first interpretation the finding is
caused by biological interactions between DRD4 and sex hormones such as androgens and
oestrogens. The second interpretation taps into a larger literature suggesting that some genetic
variations, as the DRD4 7-repeat allele, confer higher susceptibility to the effects of the
environment. Thus, DRD4-7 carriers are more influenced by gender stereotypes according to which

25
empathy is a female characteristic (see discussion in (Uzefovsky et al., 2014). The actual mechanism
may involve some or all of the suggested alternatives and further research is needed to determine
the relative contributions of each.
However, additional findings support the involvement of different types of gene-environment
interplay in empathy. A molecular genetics GxE is exemplified by the interaction between the DRD4
genotype and maternal negativity to predict 3.5 year olds’ response to a pain simulation paradigm
(Knafo & Uzefovsky, 2013). Another type of GxE is the changing effect of genes throughout
development. A meta-analysis of the available twin studies of empathy, in both adults and in
children, may begin to answer this question with the finding that heritability of empathy increases
with age (Knafo & Uzefovsky, 2013). This is a common finding in behaviour genetics, often explained
by gene environment correlations (rGE), i.e., the increasing ability of children, as they grow into
adulthood, to choose environments that best suit their needs and allow for manifestation of genetic
potential. Additional processes might be at play as well, such as that genes exert greater influence
on empathy with age.
Importantly, different genes come into play at different times throughout development, and may
have different effects throughout development. A possible example of such an effect comes from a
study of the DRD4 gene in relation to affective knowledge in young children (Ben-Israel, Uzefovsky,
Ebstein, & Knafo-Noam, 2015). In this study, affective knowledge was measured using illustrated
emotional situations. For each of four situations describing four basic emotions (sadness, fear,
anger, and disgust) children were asked what the main characters feels and to choose the
corresponding illustrated facial expression. Findings showed that, as in the study with adults, DRD4
genotype interacted with sex to predict levels of affective knowledge. Therefore these studies
further substantiate the role of the dopaminergic system in cognitive empathy. However, the
direction of the effect was opposite to that found in adults, i.e., males carriers of the 7-repeat allele
scored higher than female carriers. Although these findings need to be further substantiated by

26
replication, they are important in exemplifying that in order to understand the mechanism of
empathy it must be done while taking developmental and age effects into account. A finding of an
effect in one age group does not necessitate the same effect in another age group. Even though our
genetic makeup is, for the most part, determined at conception, the effect of each gene is
dependent upon other genes and the ever changing biological environment in which they operate.
Conclusions
Empathy has been a focus of human thought for generations, and that is not surprising, as it
provides a basis for human communication and relationships. Maybe because of that, there are
many definitions of empathy and research on empathy have diverged to cover different
methodologies and disciplines, from developmental and social psychology to neurobiology and
molecular genetics; each contributing specific insights to our understanding of empathy. We can
learn from each field and methodology, yet it is also important to begin combining our knowledge
and clearing up the differences in definitions and measurement so that we can hope to truly
understand the nature of empathy.
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Table 1. Summary of empathy relevant concepts.
Notes: *Sympathy/empathic concern is thought to motivate prosocial behaviour. That is, experiencing
sympathy towards the other means wanting to act prosocially towards them. **Personal distress and
emotional contagion would motivate prosocial behaviour only in cases where acting prosocially would
decrease the actor’s distress.
Definition
Self-Other
Differentiation
Motivates
Prosocial behavior
Towards
distress only
Emotional
involvement
Emotional Empathy
The ability to share the
emotions of another, while
maintaining a self-other
distinction (Davis, 1983;
Zahn-Waxler, Radke-
Yarrow, Wagner &
Chapman, 1992)
Yes
can
no
yes
Cognitive Empathy/
Affective ToM/
Perspective taking
The ability to infer affective
mental states (Shamay-
Tsoory & Aharon-Peretz,
2007).
Yes
can
no
no
Sympathy/ Empathic
concern
An other-regarding
emotion. Feelings of
concern for the well-being
of the other (Decety, 2010;
Eisenberg et al., 1989).
Yes
yes*
yes
yes
Personal Distress
A self-regarding emotion.
Perceiving the distress of
the other causes feelings of
distress in the observer,
motivating withdrawal
from the situation (Decety,
2010)
No
situation-
dependent**
yes
yes
Emotional Contagion
The process of being
affected by another’s
emotional or arousal state
(de Waal, 2007; Hoffman,
1975).
No
situation-
dependent**
no
yes

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Figure 1. A schematic representation of the brain regions involved in emotional and cognitive
empathy.
The emotional empathy tract may develop faster than the cognitive empathy tract.
IFG –Inferior Frontal Gyrus, STS - Superior Temporal Sulcus, IPL - Inferior Parietal Lobule, ACC – Anterior
Cingulate Cortex, TPJ – Temporoparietal Junction, vMPFC – Ventromedial Prefrontal Cortex.