F O C U S O N soc i a l n e u r osc i e nc e

review

The animal and human neuroendocrinology of social

cognition, motivation and behavior

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2012 Nature America, Inc. All rights reserved.

Cade McCall & Tania Singer

Extensive animal and recent human research have helped inform neuroendocrinological models of social cognition, motivation and
behavior. In this review, we first summarize important findings regarding oxytocin, arginine vasopressin and testosterone in the
domains of affiliation, social cognition, aggression and stress/anxiety. We then suggest ways in which human research can continue to
profit from animal research, particularly by exploring the interactive nature of neuromodulatory effects at neurochemical, organismic
and contextual levels. We further propose methods inspired by the animal literature for the ecologically valid assessment of affiliative
behavior in humans. We conclude with suggestions for how human research could advance by directly assessing specific social
cognitive and motivational mechanisms as intermediate variables. We advocate a more comprehensive look at the distinct networks
identified by social neuroscience and the importance of a motivational state, in addition to approach and avoidance, associated with
quiescence and homeostatic regulation.
Decades of animal research have provided a rich account of neuroendocrinological influences on the social behavior of non-human animals.
Inspired by these findings, human social neuroendocrinological
research has flourished in recent years. Nevertheless, seemingly
contradictory data, multi-leveled interactions and methodological
hurdles have raised serious challenges. Here we summarize important
findings in animal and human social neuroendocrinology, focusing
on how roots in animal research have sprung new branches of human
research. We then outline ways in which human research can continue to profit from the methods and discoveries of animal research
and ways in which human research offers unique opportunities for
increasing our understanding of the social cognitive, affective and
motivational processes underlying social neuroendocrinology.
The neuroendocrinology of animal social behavior
Neuropeptides (that is, oxytocin and arginine vasopressin, AVP) and
steroid hormones (that is, testosterone and estradiol) have a central
role in the social lives of animals. As comprehensive reviews on
animal social neuroendocrinology have appeared recently13, we will
only briefly touch on themes in this literature that have provided the
roots for research in humans (Fig. 1). With that in mind, we focus on
oxytocin, AVP and testosterone and their influences on four general
categories: affiliation, social cognition, aggression and anxiety/stress
responses. Although other hormones (such as estradiol) are important
for animal social behavior, their effects on human social behavior are
not well explored and will therefore not be reviewed here.
Oxytocin and AVP perform social functions in a wide variety of
species2 and show remarkable evolutionary preservation of structure and function4. Oxytocin has proven to be particularly critical
for the expression of affiliative behaviors. Its involvement in maternal behavior is well established2. It also determines whether or not
Max Planck Institute for Human Cognitive and Brain Sciences, Department of
Social Neuroscience, Leipzig, Germany. Correspondence should be addressed
to T.S. (singer@cbs.mpg.de).
Published online 15 April 2012; doi:10.1038/nn.3084

nature neuroscience VOLUME 15 | NUMBER 5 | MAY 2012

individuals of various rodent species engage in alloparental behavior5, the parenting of non-offspring. Oxytocin is also important
for pair bonding and partner preference in a number of species.
In prairie voles, a rodent that exhibits unusually monogamous patterns of behavior, oxytocin increases females preference for specific partners and oxytocin antagonists decrease that preference6.
Accordingly, extracellular concentrations of oxytocin increase (in
the nucleus accumbens, NAcc) during mating in females and during
parturition7. It is important to note that much of oxytocin function,
particularly in regards to affiliative behaviors, is regulated by estrogens influence on the expression of its receptors8.
Mother-offspring bonding, pair bonding and partner preferences
all rely on the ability of the animal to form a memory of the offspring
or partner. Oxytocin is important for social memory in a variety of
species9, determining scent-based memory in animals that rely on
scents to identify conspecifics. It appears to be particularly important
for the acquisition of social memories. Rats with a null mutation in
their oxytocin gene, for example, regain social memory if they receive
oxytocin centrally before their initial encounter with a target animal10.
It may also facilitate maternal memory consolidation11.
Oxytocin is not usually associated with aggression per se, although
it appears to be involved in maternal aggression12. Finally, oxytocin
moderates physiological stress responses and anxious behavior. In hamsters, for example, high cortisol levels inhibit wound healing such that
wound size is a gauge of the stress response and its effect on the body;
oxytocin administration buffers the stress response such that wound
size is decreased for isolated animals, whereas a centrally delivered
oxytocin antagonist impairs wound healing in paired animals13. Data
also demonstrate its anxiolytic qualities. Mice lacking the oxytocin gene
exhibit more anxious behavior in their exploration of mazes14.
AVP also affects affiliative behaviors15. In rats, AVP is implicated in
paternal behaviors, such as grooming, crouching over and contacting
pups16. AVP is also important for partner preference and pair bonding, particularly for males in a variety of species. Central administration of AVP in prairie vole males who have not mated triggers a
selective preference for mates15. The distribution of AVP receptors in
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Basic findings in animals

Recent discoveries in humans

Affiliation
Increases trust, generosity and cooperation in
economic games (in men)

Maternal behavior
Alloparenting in females
Partner preference in females
Pair bond formation in females

Improves paternal play and partner

communication

OT

Enhances ingroup preferences (in men)

Paternal behavior
Alloparenting in males
Partner preference in males
Pair bond formation in males
Species-dependent moderation
of paternal behavior

AVP
Decreases trust in women who are
dispositionally high in trust
T
Increases womens, and lowers mens, offers
in the Ultimatum Game

Social cognition

2012 Nature America, Inc. All rights reserved.

Improves face recognition

Improves facial affect recognition
Social memory in females

OT
Enhances attention toward the eyes (in men)
Improves detection of social words and
biological motion

Social memory in males

Improves encoding of happy and angry faces

(in men)

AVP

Improves detection of social words (in men)

npg

Aggression
Maternal aggression

OT

Maternal and paternal aggression

Post-mating aggression
Intermale aggression

AVP

Social aggression
Mate guarding
Dominance motivation

Increases hostile facial responses in men, but

not women

Increases preparedness for action (in women)

Increases reward sensitivity (in women)

Anxiety/stress
Reduces cortisol response to social stress
Inhibition of HPA response
Anxiolysis

OT

Extinguishes fear conditioning (in men)

Reduces amygdala response to threatening
stimuli

Regulation of HPA axis

Anxiogenesis

AVP

Inhibition of HPA axis

Anxiolysis

Increases cortisol response to social stressors

(in men)

Reduces startle response (in women)

Reduces central response to aversive images
(in women)

Figure 1 Social neuroendocrinological findings from animal and human research. Column 1 lists well-established patterns regarding the influence of
oxytocin (OT), AVP and testosterone (T) on affiliative behavior, social cognition, aggression and anxiety/stress in non-human animals. Column 2 lists
initial, parallel findings from acute administration studies in humans. Where noted, human research on a given finding was limited to one gender.

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review
the ventral pallidum in monogamous versus non-monogamous male
voles (prairie voles versus meadow and montane voles) determines the
degree of pair bonding in that species4,17. AVP also influences social
memory, most likely through effects on memory consolidation10,17.
AVP is also associated with intermale aggression18 and maternal
aggression19. For example, direct central manipulation of AVP in the
amygdala20 of castrated male rats facilitates attack behavior, and AVP
receptor (Avpr1b) knockout mice show normal predatory behavior,
but show reduced attacks against conspecifics21. A substantial body of
evidence suggests that AVP regulates systemic responses to stressors
via its influence on adrenocorticotropic hormone secretion and, in
turn, the hypothalamic-pituitary-adrenal (HPA) axis18,22. Data also
suggest that AVP has anxiogenic qualities22.
Testosterone is related to affiliative behaviors, particularly through
its relationship to other systems. Early exposure to testosterone is
necessary for AVP-triggered partner preference behavior in male
voles23. Similarly, paternal behaviors in the California mouse, such
as pup grooming and huddling, are testosterone dependent, but
this effect is mediated by conversion of testosterone to estradiol24.
Testosterone is associated with social aggression in a wide range of
species, affecting such behaviors as mate and territorial guarding 25.
According to the influential challenge hypothesis26, testosterone is
released in response to social challenges and facilitates contextually
appropriate responses to them. For example, male chimpanzees show
a substantial increase in testosterone and aggression (that is, chase
and attacks and charging displays) when females are in estrus27. In
terms of the stress response, testosterone is traditionally associated
with inhibiting HPA function28 and anxiolytic properties that correspond with a cognitive performance29.
The neuroendocrinology of human social behavior
The wealth of discoveries in animals has naturally inspired research
into human social neuroendocrinology. We are only beginning to find
the human analogs of well-established patterns from animal research
(Fig. 1). Although major findings in animals have been investigated
across sexes, species and a variety of procedures, human research has
limited data points that are often specific to one gender or one protocol (Fig. 1). We briefly review some important findings in humans,
focusing on oxytocin, AVP and testosterone, and highlight current
methodological challenges. Because we are primarily interested
in experimental work that identifies causal relationships between
hormone release and its consequences, we focus on studies in which
hormones are acutely administered (for more exhaustive reviews of
human research, see refs. 1,2,3034).
Effects on affiliation, cognition, aggression and stress. One methodological hurdle to human neuroendocrinology is the fact that one
cannot directly manipulate or measure the concentration of a given
hormone in specific regions of the brain. As such, nasal administrations of oxytocin and AVP and sublingual administrations of testosterone have been important developments for research in this domain35.
With these methods, researchers aim to manipulate central levels of
these neuromodulators and measure the effects on social behavior
and social cognition. Here we examine important findings, highlighting evidence for neuroendocrinological effects on affiliation, social
cognition, aggression and anxiety/stress.
The human analogs of oxytocins effects on affiliative behavior in
animals have mostly been studied using monetary social exchange protocols borrowed from experimental micro-economics and game theory.
These studies reveal that it increases mens trust36, generosity37,
and willingness to cooperate38. Evidence from non-economic
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protocols also supports an effect of oxytocin on human trust; oxytocin

enhances the degree to which participants rate faces as trustworthy39
and the degree to which they are willing to trust others with confidential information40.
A handful of studies used more naturalistic settings to further
demonstrate an affiliative role for oxytocin in everyday life. For example, acute administration improves both communication between
couples discussing a conflict in their relationship41 and fathers behavior toward their children during a short interaction42. It is important to note that oxytocins affiliative effects are not always positively
valenced; it appears to increase gloating and envy43, and the relative
preference for ingroup members44. When identifying the boundaries
of oxytocins prosocial effects, however, it is critical to distinguish
between preferential treatment of some individuals and derogation
of others33,45.
Given the critical role of oxytocin in social recognition in animals,
researchers have also explored its influence on human social memory
and, more broadly, social cognition. A preponderance of evidence
regarding the cognitive effects of oxytocin deals with the memory and
perception of faces (but see refs. 46,47). It appears to improve memory
for specific faces as compared with nonsocial objects or landscapes48,
but the effect is moderated by expressions of affect on those faces 49.
Data also suggest that oxytocin facilitates recognition of facial expressions (independent of face recognition itself)33, although this effect is
again moderated by the specific expression. The precise mechanism of
oxytocins facilitative effects on facial identity and affect recognition
are not yet clear, but intriguing findings demonstrate that oxytocin
increases gaze toward the eyes50,51. Indeed, the eyes are likely critical
for making inferences about the face-based stimuli in many of these
studies and may also be critical for the encoding of facial identity.
Another stream of research on oxytocin and social cognition has dealt
more directly with its possibly facilitative role in making inferences
about the mental states of others52. Again, these studies have a bias
toward using facial stimuli and, as a consequence, suggest that oxytocin enhances peoples ability to read thoughts and emotions from
the face.
In contrast with data relating to social cognition, the pattern of
findings regarding the effects of oxytocin on anxiety, stress and fear
are clearer. As in animals, oxytocin reduces the stress response in
humans. Participants who were given an acute administration along
with the social support of a friend reported less anxiety and showed
a suppressed cortisol response after the Trier stress test53. Similarly,
couples showed a reduced cortisol response after the discussion of a
conflict when oxytocin was administered beforehand41. In terms of
fear, acute administration has been shown to abolish the effects of fear
conditioning via attenuation of amygdala responses54. Oxytocin also
attenuates amygdala responses to fearful stimuli, apparently reducing the coupling of activity in the amygdala and brainstem regions
involved in autonomic responses to fear55.
Animal research on AVP has inspired work in humans as well,
although far less so than oxytocin. For instance, the affiliative effects
of AVP that have been demonstrated in animals have so far not been
studied in humans, but the effects of AVP on social cognition have.
At least in men, AVP appears to facilitate the encoding of facial identity (although this effect is moderated by the target faces affect)56. It
also enhances detection of sexual words in men57. Facial EMG data
suggest that AVP may, as with animals, have gender-specific effects
on aggression58. AVP is also related to the stress response in humans,
increasing mens cortisol response to social stressors 59. The neural
mechanisms underlying AVPs effects in humans are not yet understood, although AVP may modulate activity in the subgenual cingulate
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cortex and, in turn, the emotion regulatory processes of the medial

prefrontal cortexamygdala circuitry60.
Testosterone has also been shown to exhibit social effects in
humans. As in animals, the effects suggest that testosterone does not
simply promote aggression34. In a particularly interesting demonstration of that fact, researchers analyzed the effects of testosterone during
an ultimatum game while accounting for whether or not participants
believed that they had received the hormone. Testosterone increased
strategically fair offers, but the belief that they had been given the
hormone increased their unfair behavior61. Other data further suggest
an increase in strategic behavior. Sublingual administration to female
participants led to decreased trust levels of highly trusting individuals62. Testosterone might act, at least in part, by increasing reward
sensitivity63,64. It also affects anxiety and fear responses, reducing
the startle response65 and the central stress response to aversive pictures66. The mechanisms here are unclear. On a neuroscientific level,
testosterone appears to enhance amygdala and orbitofrontal activity
in response to threatening stimuli67 and to enhance NAcc responses
to reward anticipation64.
Methodological challenges for human research. The data from acute
administration studies provide a promising, if incomplete, picture of
the central hormonal influence on human social life. Nevertheless,
acute administration methodologies have their limits. For one, it is
not clear what proportion of nasally administered oxytocin or AVP, or
sublingually administered testosterone enters the brain30. Moreover,
it is not clear how the distribution of a given neuropeptide or
hormone compares to its endogenous release when delivered by these
means. Indeed, the endogenous central release of neuropeptides and
hormones is a complex process that we are only beginning to understand68. Peripheral measurement of peptides and steroids may provide a gauge of the endogenous central release of neuropeptides and
hormones, but the relationship between those measures and central
levels is, again, not clear31.
As others have pointed out2, the use of radioactive labeling of neuro
peptides and positron emission tomography (PET) scanning will provide a better understanding of the density and location of oxytocin
and AVP receptors in the brain. These developments will be a major
breakthrough in our understanding of the critical regions involved in
neuromodulation. They will, furthermore, improve our understanding
of nasal, intravenous and sublingual manipulations. The development
of oxytocin and AVP antagonists and agonists for humans will also help
the study of causal relations between neuropeptides and social behavior
through direct manipulation of central neuropeptide functioning2,31.
How animal research can continue to inspire human research
Animal research laid the groundwork for human social neuroendocrinology and it will continue to be important for resolving the critical
questions driving current work in humans30. Here we address how
animal research could inspire future directions in human research,
focusing on two broad themes. First, animal research has exposed the
multidimensional nature of these neuromodulators, highlighting the
importance of taking interactions between biochemical, contextual
and organismic features into account. Second, animal research provides methods for assessing implicit affiliative behavior in ecologically valid ways that could inspire analogous methods for measuring
affiliative behavior in humans.
The multidimensional nature of neuroendocrinology. The influence
of neuropeptides and steroid hormones on behavior is modulated by
neurochemical interactions, intraspecies variation in receptor density,
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individual differences such as the sex of the animal, and the specific environmental context and task that the animal is confronting. As human
research progresses and its methodologies improve, it will ideally move
to a multidimensional approach that accounts for these factors30,32.
Research on rats and other animals has shown that the neuromodulatory capabilities of neuropeptides rely on their influence on different neurotransmitter systems. For example, oxytocin modulation
of dopamine activity in the NAcc is critical for bond formation in
prairie voles6. Oxytocin also interacts with the opioid system by, for
instance, increasing opioid release in the rat hypothalamus69. In turn,
opioids modulate the activity of both oxytocin- and AVP-secreting
cells70. Interactions with these systems are critical in the context of
social affiliation given that the dopaminergic system is implicated in
the drive to affiliate30 and the consolidation of social memories71 and
that opioids come into play in relation to the reward derived from
affiliation71,72. Along these lines, researchers have used a variety of
methods to identify the specific pathways that are responsible for
various behaviors in animals, including blocking dopamine or opioid
receptors with antagonists. Human imaging research on oxytocin and
AVP has not directly tested the specific interactions between these
molecules and their associated neurotransmitter systems. This may
have led to an overgeneralization of the role of oxytocin for affiliative
and social behavior in humans while ignoring the important and welldocumented role of classical neurotransmitter systems such as the
dopamine and opioid systems for social and affiliative behaviors 71,72.
Future human research could adapt psychopharmacological intervention strategies from animal research by giving humans D2 antagonists
such as haloperidol or opioid antagonists such as naltrexone and using
different tasks to test the specific role of dopamine and opioids in
observed anxiolytic and affiliative effects of oxytocin and AVP in
different stages of the human affiliative process.
Animal research has also found that neuropeptide influences on
behavior are critically moderated by receptor densities in specific
regions of the brain4,9. For example, research on voles suggests that
oxytocin and AVP receptor density determines pair bonding and
parenting behaviors, regardless of variance in neuropeptide concentration, and that densities vary markedly across individuals and across
subspecies. Research has further identified genetic variants associated with receptor density that predict differences in social behavior4.
Following this logic, individual differences in human social behavior
and the abnormal social behaviors that are characteristic of specific
clinical populations may be a consequence of genetic variation determining reception density in critical areas 2,4,31. The development of
appropriate ligands for oxytocin and AVP research using PET methodology in humans in combination with human genetic neuroimaging
research could help to clarify the role of receptor density differences in
specific areas for different neuropeptides in human social and affiliative behavior2.
Furthermore, the animal literature has long described sex-specific
effects of steroids and neuropeptides 4. Oxytocin, for example, is
important for partner preference formation in female, but not male,
prairie voles2. AVP, on the other hand, is more critical for partner
preference in males15. For pragmatic reasons, human research has
tended to use one gender for research on a given neuromodulator
(Fig. 1). Men have been used predominantly for research on AVP and
oxytocin, whereas women have been used predominantly for research
on testosterone. Nevertheless, recent studies in humans have found
significant gender interactions in both AVP58 and oxytocin73. Clearly,
studies with combined samples of both men and women are necessary to achieve a better understanding of the gender-specific effects
of oxytocin, AVP and testosterone30,31.
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Evidence from animal research also demonstrates that neuropeptide
influences on behavior are shaped by early life experience. Research
in both voles74 and macaques75 suggests that rearing conditions alter
oxytocin synthesis and oxytocin receptor expression. Preliminary correlational work using cerebral spinal fluid concentrations in women
who experienced early childhood abuse suggests that the same may
be true for humans76. Again, future work relating information about
specific receptor densities using PET or genetic imaging approaches
to information about human life course and specific life events will
provide more definitive answers regarding the differential expression
of neuropeptides, such as oxytocin and AVP, in populations with different early life experiences and attachment styles. The apparent plasticity of neuropeptide function in early life further raises the question
of plasticity across the life course, including in adults and the elderly.
Future training and intervention research in humans should pursue
the possibility that behavioral treatment strategies such as mental
and affective training may afford beneficial changes in neuropeptide
functioning beyond early childhood.
Finally, the animal literature has repeatedly demonstrated that
a given hormones influence is highly specific to the environmental and social context. For example, testosterone function is just as
context-specific in humans as it is in animals. Despite laypeoples
beliefs regarding the link between testosterone and aggression61,
the acute administration of testosterone can both decrease trust62
and increase fairness61, depending on the circumstance. Similarly, as
human research on the influences of oxytocin on social interactions
between ingroup and outgroup members demonstrates 77, the same
neuropeptide may act very differently depending on the identity of
the interactants. Accordingly, human social endocrinological research
should always consider the complex nature of the interaction between
neuropeptides, social context and the targeted behavior30,32.
Measuring affiliative behavior in vivo. In bridging the gap between
animal and human research, it might also prove fruitful to borrow
from animal researchs strategies for measuring affiliative behaviors
in ecologically valid ways. The coding of social behaviors such as
physical proximity, grooming and huddling have been critical to the
study of hormonal influences on nurturance and attachment1. Along
these lines, the partner preference test78 measures pair bonding by
placing an experimental animal in an apparatus with an unfamiliar
animal tethered to one end of the apparatus and a familiar partner
animal tethered to the other. Partner preference is determined by the
experimental animals tendency to stay close to the partner animal.
Although we wouldnt recommend measuring a humans tendency
to approach a tethered spouse versus a tethered stranger, developments in the measurement of human nonverbal behavior provide new
techniques for measuring social approach and avoidance behaviors
reflected in proxemics. Digital body tracking allows researchers to
precisely measure interpersonal distance and gaze while the participants are not necessarily aware that their movements are being
tracked. As a consequence, participants approach or avoidance of
others can be measured during an actual social interaction with, for
example, a close relationship partner or an outgroup member79. These
measures would mimic the partner preference test and have already
been proven to be powerful indicators of social responses that are
relatively immune to deliberative control. As such, the digital measurement of proxemics would provide a naturalistic and ecologically
valid complement to the measurement of symbolic and communicative acts used in the recent wave of research in which explicit ratings
of memory and affect or economic and computer games measure
socio-affective responses.
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Social neuroendocrinology and the black box

Human research provides the opportunity to push social neuroendocrinology beyond what is possible in animal research. Specifically, we
can tap into the black box by directly measuring neuroendocrinological effects on social cognition, social emotions and motivation with
human subjective experience as an indicator (Fig. 2). Furthermore,
we can use human imaging techniques to more precisely identify the
differential effects of hormones and neuropeptides on the different
neutral networks underlying human social cognition. We provide suggestions for a closer integration of human social neuroendocrinology
with current social neuroscientific models and describe ways in which
human social endocrinology might more precisely address the full
range of human affect and motivation.
The distinct routes underlying human social cognition. In recent
years, social neuroscience has identified a relatively refined framework of pathways involved in representing different categories of
social cognition. Although these networks certainly interact in
healthy adult brains, they are structurally and functionally distinct
and can be dissociated in their developmental trajectories during
ontogeny and in psychopathology. Human neuroendocrinology
would benefit from integrating its findings into this framework.
Specifically, different neuronal routes underlie our capacity to
understand others, be it their facial expressions, actions, thoughts or
emotions. Inspired by the discovery of the so-called mirror neurons
in monkeys80,81, a multitude of functional magnetic resonance imaging (fMRI) studies in humans have identified a network underlying
the representation of others motor actions and intentions comprised
of the inferior parietal lobule, the ventral premotor cortex and the
caudal part of the inferior frontal gyrus. A distinctly different network underlies empathy, the sharing of feelings. This vicarious experience of sensations, such as pain, disgust or touch, mostly engages
somatosensory cortices, as well as the insula and anterior cingulate
cortex (for reviews, see refs. 82,83). In addition to this affective route
to the understanding of others, researchers have distinguished a
cognitive route to understanding the beliefs, thoughts and desires
of others8486. This mentalizing or theory of mind network typically
comprises areas in the medial prefrontal cortex, precuneus, superior
temporal sulcus and right temporo-parietal junction87,88. Finally,
recognition of emotional facial expressions is again associated
Cognition
Neuropeptides
and hormones

Emotion

Behavior

Motivation
Behavior
Appetitive
motivation

Approach
(for example, fight)

Aversive
motivation

Avoidance
(for example, flight)

Quiescence

Neither approach
nor avoidance
(for example, repose,
restoration,
homeostasis)

Figure 2 Overview of neuroendocrinological influences on human social

cognition, emotion, and motivation as potential (and measureable)
mediators of behavior. Models of motivation should account for
quiescence as well as for appetitive and aversive motivational states.

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with another set of brain structures, which include the amygdala,
fusiform face area and somatosensory cortices89,90.
Although recent neuroendocrinological studies using fMRI have
suggested a generalized enhancement of social cognitive capacities
through the acute administration of steroid hormones and neuropeptides, these studies have not specified which networks are affected
and, as reviewed above, are biased toward the use of facial stimuli. For
example, one study52 suggested that oxytocin increases mind reading
in humans on the basis of an increase in scores on the Reading the
Mind in the Eyes test (RMET91). Although a superficial interpretation
of these results may suggest that oxytocin affects the theory of mind
or mentalizing network, a more careful inspection suggests that the
effect may rather have been mediated by the amygdala, which is associated with face processing. The RMET tests a very specific facet of
mind reading: the inference of others internal state from subtle cues
in the eye region of human faces, a capacity that is associated with
amygdala functions and is deficient in people with autism92. Similarly,
a number of fMRI studies using nasal oxytocin in the social domain
have focused on emotional face processing, which is known to stimulate amygdala, and found a modulation of amygdala activation51,55,73.
A recent behavioral study93 found an increase of empathy and social
learning as a result of oxytocin, an effect that was not observed in
two patients with amygdala dysfunctions. Given that the stimuli in
these studies again relied heavily on faces, responses may have again
depended entirely on oxytocins effects on the amygdala.
A systematic investigation of the effects of oxytocin should identify the specific social cognition pathways involved by comparing
modulation of brain activation in classical theory of mind or empathy
paradigms that do not typically involve amygdala activation during
processing of social stimuli82,83,88,94. Accordingly, one empathy study
using a classical empathy for pain paradigm95 failed to show an oxytocin effect on empathic brain responses to the others pain typically
associated with activation in anterior insula and anterior cingulate
cortex, but not the amygdala82. In contrast, however, the same subjects
showed an oxytocin-related reduction of amygdala activation during
the anticipation of nociceptive stimulation in a non-social condition
(see ref. 96 for oxytocin decreasing amygdala activation, but simultaneously increasing insula activation elicited by the sounds of infants
crying). These findings suggest that the effects of oxytocin may not
generalize to all social cognitive tasks equally and that, in contrast
with widely held beliefs, it may not increase empathy or mentalizing
in general, but only in social tasks that elicit amygdala activation.
In general, future human social neuroendocrinological research
must tease apart the specific effects of the given neuromodulators
on different brain regions involved in different social affective and
cognitive processes to prevent oversimplifications of their functions,
such as oxytocin being the social brain hormone that increases social
cognition or empathy in general.
Toward a more comprehensive model of motivation. In addition to
using a more precise approach to identify the influences of specific
hormones and neuropeptides on particular cognitive and neural
mechanisms, neuroendocrinology should also work toward a more
precise conceptualization of motivation as an important precursor of
social behavior. Along these lines, approach and avoidance motivation
represent a dominant framework for understanding affect, cognition
and behavior across a wide variety of domains. Not surprisingly, these
two motivational states factor importantly into various models of
neuroendocrine influences on social behavior3,30,97. Organisms are,
however, capable of more states than just approach and avoidance.
In fact, the calm, digestive, reconstitutive state associated with goal
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achievement is essential to homeostasis in animals, including humans

(Fig. 2)98. This state is central to social affiliation, which proceeds
from an approach-motivated appetitive state, to consummation, to
this quiescent state71. In the existing human social neuroendocrinological literature, it is difficult to distinguish between effects driven
by approach versus quiescence. Here we describe ways in which these
two states should differ in their neurochemical, subjective, autonomic
and behavioral qualities.
At the level of neurotransmitter systems, approach is likely associated with dopaminergic systems and quiescence with opioids71,98.
Oxytocin, for example, modulates both systems. Knowing which
combination of these systems is active in humans in a given context would allow for better predictions concerning subjective affect,
social cognition and behavior. Again, pharmacological manipulations
using dopaminergic and opioid antagonists would allow experimental
research.
At the level of subjectivity, approach and quiescent motivational
states likely differ in terms of their affective qualities. Quiescence
should be associated with positive feelings of warmth and calm.
Approach, on the other hand, may be negatively valenced, as in the
case of anger, or positively valenced, as in the case of excitement.
Although some researchers have measured subjective reports of affective states during the course of experiments (for example, see ref. 50),
future studies could provide a more comprehensive picture of the
affective experience of acute administration with particular focus on
warmth (associated with quiescence) and arousal (associated with
approach). Furthermore, quiescence and approach motivations should
be particularly distinguishable by their autonomic responses, with the
quiescent state being associated with parasympathetic activity and the
appetitive state being associated with increased sympathetic activity.
More comprehensive assessments of autonomic states during acute
administration might help to identify which motivational system is
active during a given task under the influence of a given hormone.
At the behavioral level, approach clearly motivates action or
readiness for action. Quiescence, on the other hand, motivates the
absence of any action. Currently the literature confounds these two.
For example, oxytocin may facilitate prosocial behavior by facilitating
approach, affiliative goals, or it may do so by reducing social avoidance (that is, via quiescence)97. Despite its prediction of inaction, the
quiescent state should be measureable given its reduction of stressrelated actions and associated subjective stress experience, as well as
in health-related indicators such as cortisol levels.
In addition to providing us with a more nuanced understanding
of human social endocrinology, acknowledgment of the quiescent
state will help to translate basic findings in this domain into clinical
applications. Indeed, compassion-based therapy and compassionbased stress reduction strategies are based on the activation of the
quiescence and demonstrate the profound benefits of this state to
long-term physical and mental health99,100.
COMPETING FINANCIAL INTERESTS
The authors declare no competing financial interests.
Published online at http://www.nature.com/natureneuroscience/.
Reprints and permissions information is available online at http://www.nature.com/
reprints/index.html.
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