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What is Social Neuroscience?

Ian Morton's picture

Social Neuroscience:
The Search for the Social Brain
Ian Morton, 2007


This past summer (June-August, 2007) I participated in Bryn Mawr's Summer Science Research Fellowship Program, working for ten weeks in Paul Grobstein's lab along with friends and colleagues Rebecca Woodruf, Heather Fetting and Ashley Dawkins. In addition to contributing to collaborative, inquiry-based discussions (see PGLab-Summer2007 and Brain Stories), I pursued independent research, seeking to get a foundation in the developing field of social neuroscience. Here I offer a summary of my findings with the intent of offering a generalized understanding of what social neuroscience entails. Additionally, I hope to expand upon this page as my research continues and encourage anyone to offer his/her own thoughts and insight into this topic.


What is "Social Neuroscience?"

Social Neuroscience is a term applied to an emerging field of study concerned with identifying the neural processes underlying social behavior/cognition. However, the field extends beyond a search for neurochemicals and brain regions that cause behavior; social neuroscience seeks to better understand the relationship between the brain and (social) behavior (Waldrop, 1993; Decety & Keenan). This relationship is reciprocal: the brain affects social cognition and behavior, and social cognition/behavior affects the brain (Insel & Dernald, 2004). The underlying question of social neuroscience, then, is how do the agents of this dyadic relationship influence one another? From this question a multitude of increasingly specified questions arise, but all of which seek to examine some aspect of the nature of the social brain-behavior dyad.

The brain is an immensely complex system and it is becoming increasingly evident that the brain does not operate completely via feed-forward, hierarchical processes, stemming from discrete brain “centers”(Adolphs, 2003). Rather, the brain operates on parallel networks of both feed-forward and feedback loops, distributed across the brain (Waldrop, 1993). As behavior is a result of the brain, it too is highly complex, influenced by numerous factors, both neurological and environmental in nature (Insel & Dernald, 2004). Due to the complexity of both brain and behavior, dogmatically adhering to classical scientific methods (rigidly controlled hypothesis-testing experiments) will not suffice. Controlling for the multitude of factors seems out of the question, thus making it virtually impossible to reach any absolute conclusions (Adolphs, 2003). It seems that the best hope for progress in social neuroscience will be the search for overarching patterns of brain-behavior through identifying converging results from diverse studies (Adolphs, 2003; Raichle, 2003; Insel & Dernald, 2004; Decety & Keenan). Seeking fully objective conclusions about the social brain solely within the laboratory may hinder the field more than it would benefit its progress.

Consequently, social neuroscience is best served through interdisciplinary research. Researchers from across fields such as cognitive neuroscience, social psychology, psychiatry, sociology, and philosophy, are all contributing to this ongoing examination of the brain and the social condition. Research within this field should include studies of brain function and organization (Young, 2001; Gallese, 2003; Insel & Dernald, 2004; Amodio & Frith, 2006), functional imaging studies (Raichle, 2003; Heatherton et al., 2004), case studies (A. Demasio, 1995; Dobbs, 2007), animal studies (Young, 2001; Insel & Dernald, 2004; Miller, 2007), lesion studies (A. Demasio, 1994), behavior studies (Miller, 2007), meta-analyses (Zhou et al., 2005; Amodio & Frith, 2006) and perhaps additional techniques not yet devised. Social neuroscience can therefore be understood as an interdisciplinary field that seeks to better understand the relationship between brains and social cognition and behavior through the use of various experimental techniques and converging results.

Some Key Terms/Theories:

To date, several provocative and influential theories have emerged from the study of social cognition and behavior. As a part of understanding social neuroscience, it is important to examine the current directions in which research is headed, the predominant theories that are in play, and state of our current understanding of the “social brain.” Not only should one understand these theories, but also one should also critically examine their implications, the validity of their foundations and assumptions, and the nature of how research is approaching these topics.


Imitation is precisely as it sounds; it is the duplication of an observed behavior. However, as simple as it is to define, the precise mechanisms of imitation remain contentious (Brass & Heyes, 2005). A predominant view describes imitation as a behavior whereby an individual translates an observed, external action of an Other into an internal pattern of neuromuscular activity that produces an equivalent action, thereby implying intentionality (Meltzoff & Decety, 2003; Brass & Heyes, 2005). Meltzoff believes that infants are born with an innate capacity to imitate, which supplies them with a powerful learning tool in the social environment (Melzoff & Decety, 2003). Further, the mechanisms of imitation may play a decisive role in the development of theory of mind (Meltzoff & Decety, 2003).

Theory of Mind (ToM):

(Also referred to as mentalization and mind reading)

Theory of Mind refers to the cognitive ability to attribute states of mind such as desire, intention and emotion to other people (and to oneself) as a means for predicting their behavior (Apperly, I.A., 2007). The cognitive ability to recognize and understand that others behave in a goal-oriented manner is a valuable tool for surviving in a social environment, as it allows individuals to predict the actions of others and therefore to manipulate and influence their behavior (Gallese, 2003). As ToM involves the recognition that others have mental states and perceptions that are distinct and independent of one’s own, of which one cannot have any direct knowledge, one can only have a “theory” of the other’s mind. ToM is an essential cognitive ability for successful social existence and has thus been a major focus of cognitive social neuroscience research. From the research with ToM, two major theoretical mechanisms have been proposed: Simulation Theory and Theory Theory (Vogeley K et al., 2001).

Simulation Theory (ST):

Simulation Theory is one of two leading theories as to what serves as the basis for ToM capacity. According to ST, the capacity for ToM is based in one’s ability to take on another’s perspective via simulation, the cognitive act of placing oneself in another’s mental “shoes” (Gallese, 2003; Vogeley K et al., 2001). According to Gallese, simulation is embodied in the sense that the process of mentally modeling the behavior of other draws upon the very cognitive modeling used to shape one’s own behavior (Gallese, 2003). Similarly, Vogeley et al. argue that in process of simulation, the mechanisms used for one’s own motivations, emotions and rationality are employed to imagine the motivations and emotions of the Other (Vogeley K et al., 2001). Thus in ST, one not only takes the perspective of the Other, but also projects one’s own subjectivity onto the Other.

Theory Theory (TT):

TT proposes that one’s ability to understand and predict the behavior of others is dependent on the employment of a social knowledge base, often called folk psychology, to infer the mental states of others (Stich & Nichols, 1993; Gallese, 2003). Thus, one’s social understanding of another is accomplished entirely and solely through mental metarepresentations, contrary to the notion of embodied simulation inherent to ST (Gallese, 2003).

Folk Psychology:

Folk psychology refers to a theoretical framework about the social world (Stich & Nichols, 1993). It is a body of theories or principles, constructed automatically and unconsciously throughout one’s ontogenic development; one continually creates, tests, and revises theories about the social world (Gopnik & Meltzoff, 1997). As folk psychology refers to a body of theories which serves as the foundation for TT, it is occasionally used synonymously with theory of mind.


Note: a DOI (Digital Object Identifier) is an identifier for a body information located on a digital network. That is, DOIs refer to the body of information itself, not just a URL where the body of information can be found. DOIs remain constant for a specific intellectual property, such as an article, and can be resolved using Resolving a DOI will supply information on the body of information such as its title and where it can be found digitally (a URL). For further information on the DOI system, visit


Meeting of minds: the medial frontal cortex and social cognition

David M. Amodio and Chris D. Frith

Amodio, David M., and Chris D. Frith. "Meeting of Minds: the Medial Frontal Cortex and Social Cognition." Nature Reviews Neuroscience 7 (2006): 268-277.

In their paper, David Amodio and Chris Frith incorporate a wide range of meta-analysis research across fields associated with social cognition along with anatomical characteristics of the medial frontal cortex (MFC) and neighboring regions to draft proposal for the various roles of the MFC in social cognition. While the anatomy is somewhat specific and the number of study results covered vast, giving the paper an overwhelming effect, upon careful reading the layman can extract the major proposals of this paper. The authors present a theoretical model of the MFC that seems well founded in the current research, but which remains to be confirmed. However, their proposal could offer a valuable foundation for drafting future research on the role of the MFC in social cognition. Further, such a detailed analysis of the MFC and its subdivisions is beneficial when juxtaposed with other theoretical works concerning social cognition that may or may not implicate the involvement of MFC activity.

According to Amodio and Frith’s model, the MFC is important for guiding behavior based on anticipated value within a social context. The posterior rostral region (prMFC), the most caudal of the regions of concern, is associated with a process of internal monitoring of response selection, thus regulating behavior in terms of the predicted value of possible actions. The orbital region (oMFC) guides behavior in terms of the value of possible future outcomes. Finally, the anterior region (arMFC), lying between the two and most caudally positioned, has been associated with self-knowledge, person knowledge and mentalizing. The authors go on to argue that within the overall architecture of the MFC, processes become more abstract and complex in more rostral/anterior regions. The activity of posterior MFC regions appears to be associated with given information such as a felt sense of pain while anterior regions are concerned with more abstract representations of experience such as sensing that a loved-one is in pain (empathy), a process independent of sensory input. The authors propose that the arMFC is responsible for meta-cognitive processes of reflecting on feelings, intentions and values linked to outcomes and actions, thereby implicating its crucial role in empathy, morality and reputation, all key aspects of social cognition.


Discrete hierarchical organization of social group sizes

W.-X.Zhou, D. Sornette, R. A. Hill and R. I. M. Dunbar

Zhou, W. -X., D. Sornette, R. A. Hill, and R. I. M. Dunbar. "Discrete Hierarchical Organization of Social Group Sizes." Proc. R. Soc. B 272 (2005): 439-444. PubMed. Haverford, Haverford.

The “social brain hypothesis” suggests that group sizing is somewhat limited by cognitive factors such as the volume of neural material available for processing complex social information and dynamics. When groups attempt to extend beyond their cognitive boundaries, social stability suffers and groups fall apart. In this paper the authors make their own contribution to the ongoing discussion of grouping patterns in humans, identifying a hierarchy of social group sizes.

Typically humans have a small core group of friends (the ‘support clique’). This core group is part of a larger sympathy group of friends, which in turn is part of the larger band. Bands, as termed in reference to hunter-gatherer societies, consist of a dynamic group of individuals, but which are all pulled from a common pool of people, the clan. The clan in turn is part of the megaband, which is a subgroup of the overall tribe. Using quantitative analysis of social networks from census data from various countries, the authors found that between these successive groupings there was a constant scaling ratio of about 3. That is they found that support cliques had a mean size of 4.6, sympathy groups a mean size of 14.3, bands 42.6 etc. From these results the authors propose that humans form groups according to a distinct hierarchy with a preference for a scaling ratio of about 3.

To support the quantitative analysis the authors also did a systematic analysis that took into account all the data, not just the means, as well as analysis of a third data set. In each case, they observed a scaling ratio of about 3. The data presented here is interesting as it shows a common scaling ratio for hierarchical group organizations from a diverse set of data, thereby suggesting that this is more than coincidental. Similar organization is observed in other facets of human life such as the military and stock market. One is left to wonder if this form of structuring is innate to human cognition, or perhaps the limits of human cognition. While this paper is not necessarily neural in nature, it suggests a neural basis and has implications for how we understand the neural processes of social understanding and categorization. How might this discrete hierarchical processing of social networking relate to the evolution of the social brain?


How The Brain Processes Social Information: Searching for the Social Brain

Thomas R. Insel and Russel D. Dernald. Annural Review of Neuroscience. 2004;27:697-722.

DOI: 10.1146/annurev.neuro.27.070203.144148

In this paper, Insel and Dernald describe some of the recent studies related to the processing of social information with the intent of highlighting some of the neural mechanisms that appear to be specialized for social cognition. While the bulk of research covered in this paper does not pertain directly to humans, the correlations between specific neural mechanisms and particular aspects of social behavior observed in model organisms can offer some general insight into how the nervous system shapes behavior and vice versa. With these general concepts in mind, future research can be shaped to more precisely understand the neural correlates of social behavior and could shed new light on the role of the human nervous system in social cognition thereby offering potential benefits for better understanding neurological social disorders. The authors also critically consider the research they present in their paper and offer questions and ideas to help shape future experiments.

Insel and Dernald call upon research that illustrate both ways in which social behavior can be influenced by physiological processes and ways that social behavior and perception can influence physiological processes. This is an important concept to highlight, that it is the interplay between one’s physiology and one’s environment that shapes behavior, not one or the other. With this in mind, the authors recognize that studying social neuroscience is difficult, as results can be skewed/misleading depending on the context (environment, social) of the experiments. However, Insel and Dernald believe that a blend of multiple studies will allow us to begin defining the circuitry of the “social brain” and could thus help us to better understand how the brain operates within the social world. Additionally, in their paper the authors present a simplified and theoretical model for mammalian social processing.


The Neurobiology of Social Recognition, Approach, and Avoidance

Larry J. Young

The neurobiology of social recognition, approach, and avoidance. Biological Psychiatry, Volume 51, Issue 1, 2001: Pages 18-26.

In this paper, Young discusses three main rodent models that are valuable for understanding the basic neural mechanisms involved with social processing, approach and avoidance. First, studies with oxytocin knockout (OTKO) mice suggest that the amygdala, rich in OT receptors, differentially processes social versus nonsocial information, but requires the presence of OT in order to do so. Young’s research with voles was the first to illustrate the direct correlation of the regional distribution and density of neuropeptide receptors with unique sets of social behavior. Young observed a high concentration of OT and vasopressin (AVP) receptors in regions associated with the dopamine reward system in highly social voles, but found low concentrations in the same regions of nonsocial voles. Young infers that OT and AVP promote social behavior in social voles through activation of the neural circuitry of reward thus reinforcing highly social behavior, while nonsocial voles, not expressing significant receptor distribution in regions connected to reward, do not receive the same motivation for social behavior. Finally, studies with hamsters suggest that corticotropin releasing factor (CRF) is involved in the expression of social avoidance and submissive behavior.

While the studies presented by Young do not directly pertain to human systems, they provide valuable correlations between basic neural mechanisms and social behavior that may help to elucidate the role of the brain in the social behavior of humans. Young clearly presents the observations from rodent systems and does well at relating the observations to human systems and conditions. In particular, this paper could offer new and better ways to understand the human condition of social anxiety. Young’s work with voles implicates the importance of brain organization, a result of genetic sequence, for shaping behavior. It therefore seems plausible that individual variation in organization of the dopamine system would result in different predispositions for social behavior, including prosocial, affiliative behavior or nonsocial, anxious behavior. What roles might OT, CRFs and the amygdala play in prosocial versus socially anxious behaviors? While this paper can be useful for better understanding the neural machinery behind social approach and avoidance, one should not be misled to believe that brain organization solely determines human behavior. It should rather be understood that the brain creates strong dispositions for certain sets of behavior, which get reinforced or inhibited with experience in the world.


Investigating the cognitive neuroscience of social behavior

Ralph Adolphs. Neuropsychologia. 2003;41(2):119-26.

DOI: 10.1016/S0028-3932(02)00142-2

Ralph Adolph wrote his paper to complement several featured papers from this issue of Neuropsychologia, which cover some recent findings pertaining to social knowledge and behavior. Adolph did not intend to review the neurobiology of these articles, but rather to outline and discuss key issues provoked by these papers that pertain to the field of social neurobiology as a whole. This paper begins with a brief presentation of the featured articles, in which Adolph discusses the benefits offered by different techniques utilized in each, such as functional imaging, while also illustrating that results from a single study are incomplete and inconclusive. Adolphs’ intent is to show that social neuroscience differs from other “strict” scientific fields in both theory and methodology. Individuals results from hypothesis-testing studies are inconclusive on their own, however the accrual of results from multiple studies, using diverse techniques, could outline a more reliable brain-cognition-behavior relationship. Social neuroscience will thereby benefit from a collaborative, interdisciplinary, meta-analysis approach.

Social cognition and behavior involves a wealth of non-controllable factors, thereby complicating the approach to its study. Further, processes in the brain run in parallel and utilize both feed-forward and feedback information. It is clear that social neuroscience will prove a challenging field for researchers, and in order to best approach research within this field, there are some foundational issues that should be addressed. The paper goes on to discuss some of these key issues, as Adolphs poses five questions to theory and five to the methodology involved in social neuroscience and offers his own thoughts on these topics. Adolphs considers topics such as the very language utilized by this field, how social behavior is to be qualified and quantified, how data is presented and interpreted, whether or not social cognitive processes are reducible to non-social processes, and the goal of drafting neurological explanations for social behavior. Many of the questions and concerns raised by Adolphs in this paper are crucial for consideration within this field, and consequently anyone interested in social neuroscience would be best served to read Adolphs’ paper and consider the questions posed from one’s own stance.


What the Social Brain Sciences Can Tell Us About the Self

Todd F. Heatherton, C. Neil Macrae, and William M. Kelley

Current Directions in Psychological Science, Volume 13, Number 5, October 2004, pp. 190-193(4).

In this paper the authors set out to show how recent advances in brain imaging, in particular PET and fMRI, can help to better understand the “self.” The authors address two topics concerning the self. First, they discuss the self-referential effect in memory, this referring the apparent augmentation of memory when events or information are associated with the self. There has been debate over whether or not the enhanced memory effect is a result of neural self-reference processes, a debate that was hard to resolve using behavioral observations. With the use of fMRI, the authors observed that memory involving self-reference differed from other memory in that it selectively activated regions of the medial prefrontal cortex (MPFC). Further testing with fMRI showed that the greater the MPFC activation observed during self-referential judgments the more likely an item was to be remembered in later testing. Trough the use of fMRI the authors were able to show that MPFC activation correlates with self-referential processing and also plays a role in forming memories pertinent to the self. The authors were also able to use fMRI with split-brain patients to show a correlation between the left-brain hemisphere and self-recognition, while general recognition of others correlates to the right hemisphere.

This paper offers some interesting findings related to the self that researchers were able to infer with the aid of brain imaging techniques. The results reported could prove valuable for understanding social cognition and also illustrate the benefits of brain imaging. However, the authors did not offer a comprehensive discussion of the merits of cognitive neuroscience techniques. This paper could have benefited from a more critical analysis of the techniques, recognizing both the pros and cons of brain imaging, as well as describing what these processes actually are and how it is they offer a unique way to understand the self. The authors say that fMRI was able to show activation of the MPFC during self-reference, but they don’t explain what that means or how it’s quantified. This paper reports some interesting findings within the field of social-neuroscience, but could do better in its consideration of the techniques utilized in social neurosciences.

Social Neuroscience: A Role for Brain Imaging

Marcus E. Raichle

Political Psychology, Vol. 24, No. 4, Special Issue: Neuroscientific Contributions to Political Psychology (Dec., 2003), pp. 759-764.

Social neuroscience has benefited greatly through its adoption of techniques utilized in cognitive neuroscience research, namely functional imaging. These techniques allow researchers a noninvasive, “ethical” glimpse into the activity of living brains in human subjects. Raichle stresses that if functional imaging is to optimally benefit this research, social and cognitive neurologists must work closely together in drafting questions and research methods, thereby making this an interdisciplinary process. As functional imaging has proven key to the development of social neurobiology as a field, Raichle set out to describe the current state of functional imaging as well as its development.

There is a tendency when interpreting functional imaging data to describe cognition in terms of localized domains. This is partially due to the presentation of imaging results in which emphasis is placed on specific brain regions while neglecting to include information from the entire data set. Raichle stresses that such interpretation of data is incomplete and that complex mental functions are the result of a wide distribution of activity. Therefore the role of functional imaging should not be to find domains of social cognition, but rather to identify the multiple regions activated during cognition and behavior, and their temporal relationships. How then should such data be utilized to address the questions facing social neuroscience? Raichle believes that eventually, through many well-designed experiments, converging results from various laboratories could reveal the nature of the neural processes underlying certain tasks or behaviors. With these results, social neurologists should be better prepared to move forward and address behaviors in more life-like environments.

As an interdisciplinary field it is indispensable that scientists across fields communicate not only results, but understand what aspects of brain activity each are measuring. For instance, neurophysiologists tend to focus on changes in neuron spiking or firing rates, which reflect output activity, however, functional imaging used by cognitive neurologists reflects changes in input activity. The nature of the interdisciplinary dialogue is thereby crucial and scientists should remain informed on the evolution of this dialogue. Raichle goes on to address additional issues facing cognitive neuroscience and brain imaging such as individual differences between test subjects, the current lack of information on brain maturation on a systems level, and the tendency to overlook baseline activity. Raichle does well to discuss the important applications of functional imaging within the field of social neuroscience and addresses some of the key issues concerning this developing process.


Cognitive Neuroscience: A World With a Future

M. Mitchell Waldrop

Science. 1993 Sep 24;261(5129):1805-7. Cognitive neuroscience: a world with a future MM Waldrop (24 September 1993).

DOI: 10.1126/science.8378783

This article, written for Science in 1993, describes cognitive neuroscience as a fast-emerging field. With the development of new scanning techniques such as functional imaging, researchers are no longer limited to studying behavior from an external standpoint nor to studying the brain through invasive techniques or through removing the brain itself. With this new technology, cognitive neuroscience has emerged as a field in which researchers can pursue an understanding of high-level brain functions and ultimate behavior through examining the activity of widespread, distributed and parallel neural networks. Additionally, computer modeling has also proven to be a valuable tool for cognitive neuroscience, allowing researchers greater into the operations of neural networking. Requiring backgrounds in functional imaging, experimental skills, computational skills and knowledge of brain structure and organization, the demand this field places on researchers is high, but with the demand for new skill sets comes plentiful job opportunities. This article is a good source for a brief review of the emergence of cognitive neuroscience as its own field and its promise of many years of future research to come. Understanding how the mind arises from activity of the brain is becoming a more realistic possibility as cognitive neuroscience programs continue to grow, new methods emerge and new technology develops. These are exciting times for this young field and its budding divisions including social neuroscience.


What Imitation Tells Us about Social Cognition: A Rapprochement between Developmental Psychology and Cognitive Neuroscience

Andrew N. Meltzoff; Jean Decety

Philosophical Transactions: Biological Sciences, Vol. 358, No. 1431, Decoding, Imitating and Influencing the Actions of Others: The Mechanisms of Social Interaction. (Mar. 29, 2003), pp. 491-500.

This paper investigates the innate capacity of humans for imitative behavior with the presumption that understanding the mechanisms underlying imitative behavior can provide a foundation for understanding the greater faculty of social cognition. The authors propose that imitation, an innate human function, developmentally and evolutionarily precedes Theory of Mind (ToM) and further, that ToM develops from the neural machinery underlying imitation. With the observation that imitation is beyond mere visual-motor resonance, involving the use of memory, the authors are lead to believe that imitation involves more than the activity of mirror matching neural systems and that consequently, mirror neurons alone are insufficient to account for the development of ToM. The authors argue that motor imitation serves as the crucial link between internal representations and the ‘like me’ analogy of interpersonal relations.

Meltzoff and Decety’s paper is a valuable contribution to the ongoing discussion on imitation, mirror neurons and interpersonal relations, highlighting research observations that challenge proposals implicating mirror neurons as the foundation for social cognition. The authors propose that mirror neurons may not be an innate feature of the human brain, but rather develop from experience. What is innate to humans is the propensity for imitation, the recognition of coherence between perceived and performed actions, which may be the foundation for mentalization. Through experience and the functioning of proprioception, infants construct a link between their inner feelings and outward behavior, a link which they can then project onto the Other upon observing the Other’s behavior. Thus ToM could be a developmental feature of the mind stemming from the neural machinery allowing for imitation. The authors call on persuasive research to support their hypotheses and to highlight the possible neural mechanisms underlying imitation. Specifically, they implicate the inferior parietal cortex in mediating between self-other, which recognizes both similarities and differences between the behavior and the underlying mental states of the Other and oneself.


Imitation: is cognitive neuroscience solving the correspondence problem?

Marcel Brass and Cecilia Heyes

Marcel Brass and Cecilia Heyes, Imitation: is cognitive neuroscience solving the correspondence problem?, Trends in Cognitive Sciences, Volume 9, Issue 10, October 2005, Pages 489-495.

The authors of this article address what is called the “correspondence problem,” the question of how an observer is able to initiate the proper pattern of motor activations to imitate the externally observed actions of another. Two categories of theories have arisen to address the correspondence problem, specialist theories and generalist theories. Specialist theories argue that imitation is controlled by a set of dedicated neural mechanisms, while generalist theories argue that imitative ability arises in development through the integration of associative learning and action control processes. The authors examine recent findings from cognitive neuroscience research that suggest a generalist theory of imitation is more likely the case.

The authors do well to offer a general description of prominent generalist and specialist theories, specifically the ideomotor theory (IM), the associative sequence learning model (ASL), and the active intramodel matching model (AIM). Of these theories, ASL seems to be the most supported by recent observations. The authors present the research supporting ASL clearly and go on to offer a discussion of the role of learning in imitation. Learning, an understudied aspect of imitation refers to the process by which visual representations (action observation) become linked with motor representations (somatosensory, proprioceptive information) throughout one’s ontogenic development. Finally, the authors round out the discussion of imitation by presenting findings that implicate mechanisms involved in distinguishing self from other in the process of inhibiting imitative behavior. This evidence further supports the theory that action observation activates equivocal motor representations that can then be used to guide imitative behavior. Overall this paper is a decent source on some of the recent findings from cognitive neuroscience research that support generalist theories of imitation.


The Manifold Nature of Interpersonal Relations: The Quest for a Common Mechanism

Vittorio Gallese

Philosophical Transactions: Biological Sciences, Vol. 358, No. 1431, Decoding, Imitating and Influencing the Actions of Others: The Mechanisms of Social Interaction. (Mar. 29, 2003), pp. 517-528.

Vittorio Gallese, Associate Professor of Human Physiology at the University of Parma, Italy, researching with Giacomo Rizzolatti and Leonardo Fogassi, first discovered “mirror neurons” in the brains of macaque monkeys in the early 1990s. Since his initial discovery, Gallese has continued to research and publish extensively on mirror neurons in both monkeys and humans. In this paper, Gallese draws on developmental psychology research to suggest that a common neural format between observing the actions of an external subject and an internal neural representation (simulation) utilizing the somatosensory system, which models what it would be like for the perceiver to perform the observed actions, could be the foundation for one’s sense of existing within an intersubjective space, a space in which one has a sense of self/other equivalence (‘like me’). Gallese argues that this sense of intersubjectivity is a crucial precursor for social cognition, including functions such as imitation, empathy and “mind reading” (e.g. Theory of Mind). Applying his own research with mirror neurons, Gallese proposes that the functional mechanism underlying “embodied simulation” relies on the activity of a mirror-matching neural system, which occurs continually, automatically and unconsciously.

This paper serves as good introduction to Gallese’s perspective on Theory of Mind (ToM). ToM is essentially described as a rational function involving intentionality. Stemming from Simulation Theory (one approach to ToM), Gallese argues that ToM results from mentally modeling the behavior of others, but diverges from ST in his belief that simulations are not the result of a rational and intentional act, but rather occur automatically and unconsciously, thereby describing social cognition in terms that are not limited to the rational mind. Contrary to views of ToM that favor a domain-specific function, the Gallese proposes that ToM is built from a basic and distributed functional mechanism, the primary mechanism with which we model reality, embodied simulation. Further, Gallese argues that unlike TT, which implicates ToM as a distinct cognitive division between humans and nonhuman primates, ST allows for an evolutionary continuity between primates and humans, between behavior/mind reading and ToM. This paper is therefore valuable in that it offers a unique and parsimonious theory of social cognition with a practical evolutionary path.


Mind Reading: Neural Mechanisms of Theory of Mind and Self-Perspective

Vogeley K, Bussfeld P, Newen A, Herrmann S, Happé F, Falkai P, Maier W, Shah NJ, Fink GR, Zilles K.

Neuroimage. 2001 Jul;14(1 Pt 1):170-81.


This paper follows the form of typical scientific writing and is a good example of how fMRI is being utilized to reach new understandings of brain activity during cognitive processes. The authors are concerned with two important cognitive capacities employed by humans, the ability to recognize oneself as the immediate subject of one’s experiences (SELF) and the ability to attribute mental states to an Other, or theory of mind (ToM). Both of these cognitive functions are crucial to successful social cognition and behavior and the two seem closely related. The question is, how closely related are they? To what extent is SELF required for ToM? In an attempt to address this question, two theories have arisen over the years. The first, simulation theory (ST), suggests that SELF is a subcomponent of the greater capacity for ToM, and as such, these two functions would likely utilize common neural mechanisms. The second theory, theory theory (TT), argues that the capacity for ToM arises from a distinct theoretical knowledge base that is gained through one’s ontogenic development. According to TT, acquiring the capacity for ToM is distinct from developing a sense of SELF.

To begin addressing this issue the authors utilized fMRI and a selection of carefully categorized stories to assess what regions of the brain are activated during tasks that require SELF, ToM, neither (baseline) and both. The authors observed that both common and distinct brain regions are associated with SELF and ToM. Specifically, the exercise of ToM is correlated with activity in the right anterior cingulate cortex, irrespective of whether or not SELF is also being utilized. SELF also shows a correlation with the right anterior cingulate cortex activity, but also seems to require activity in the right inferior temporoparietal cortex, irrespective of whether or not ToM is being utilized at the same time. These results conflict with both strict ST or TT accounts of ToM. If ST were exclusively valid, ToM activity should always involve the neural processes necessary for SELF, which is not observed in the results. The authors instead propose that it is a blend of both ST and TT that allows for ToM to arise. This is a valuable paper not only for its contributions to an ongoing understanding of ToM, but also for the creative implementation of fMRI to address questions that have been previously unanswerable.


Beyond Simulation–Theory and Theory–Theory: Why social cognitive neuroscience should use its own concepts to study “theory of mind”

Apperly, I. A., Beyond Simulation–Theory and Theory–Theory: ..., Cognition (2007), doi:10.1016/j.cognition.2007.07.019


Getting Vygotskian about theory of mind: Mediation, dialogue, and the development of social understanding

Fernyhough, C., Getting Vygotskian about theory of mind: ..., Developmental Review (2007), doi:10.1016/j.dr.2007.03.001


Folk Psychology: Simulation or Tacit Theory?

Stephen Stich; Shaun Nichols

Philosophical Issues, Vol. 3, Science and Knowledge. (1993), pp. 225-270.


Words, thoughts, and theories

Gopnik, A. & Meltzoff, A.N. 1997 Words, thoughts, and theories. Cambridge, MA: MIT Press.


Descartes' Error

Antonio Demasio

Demasio, A. (1994). Descartes' error: Emotion, reason, and the human brain. New York: G.P. Putnam's Sons.


All Together Now—Pull!

Miller, Greg.

Science. 2007 Sep 7:317(5843), pp. 1338-1340.

DOI: 10.1126/science.317.5843.1338

Greg Miller reports on current research being performed with apes at wildlife sanctuaries. Specifically, at the Ngamba Island Chimpanzee Sanctuary in Uganda, researchers are observing chimp cooperation and its limitations to make inferences about the evolution of sophisticated social behavior. Chimps are observed while performing tasks such as the rope-pull, in which two chimps must cooperate by simultaneously pulling on separate ropes in order to obtain a reward (food). Chimps failed at this task when one was preoccupied with asserting dominance or one was too fearful to fulfill his part. However, two chimps of a similar rank cooperated successfully to complete the challenge. From these and additional observations of ape cooperation, Brain Hare has come to believe that social tolerance is a phylogenetic prerequisite for sophisticated social behavior. In support of this hypothesis, bonobos, which are said to be the most tolerant of the great apes, show an even greater ability to cooperate and complete the rope pull task than chimps. While bonobos do not show cooperation in the wild—perhaps because cooperation isn’t needed due to an abundance of resources—this evidence suggests that they do possess an advanced cognitive capacity to do so relative to chimps and other taxa.

How bonobos developed the capacity for such sophistication of social behavior is unknown, and the paper goes on to discuss tentative theories, but insufficient evidence exists to support them. Hare, however, believes that there were selective pressures against aggression amongst bonobos, which in turn augmented social tolerance, therefore allowing for greater accommodation of cooperative behavior. While the observations and results reported are inconclusive, they make intriguing implications for both the evolution of sophisticated social behavior and for its applications in human society. In studying differences of social behavior between bonobos and chimps, Hare hopes to learn not only about how these ape species came to differ, but also how humans came to surpass apes in social behavior. Additionally, it seems to me that this research suggests that if humanity is to maximize its fitness benefits garnered through social existence, we must learn to be more tolerant of others so as to more flexibly cooperate. Stemming from this is the question, how important is tolerance and cooperation for fitness? This seems to be an even more challenging question when one considers the success of humans afforded through manipulation, deceit, and control.


Evolution in the Social Brain

R.I.M. Dunbar and Susanne Shultz

Science. 2007 Sep 7:317(5843), pp. 1344-1347.

DOI: 10.1126/science.1145463

In their paper, Dunbar and Shultz report on the evolution of larger brains in social species. While previous research has suggested that the evolution of larger brains can be attributed to ecological pressures such as the demands of foraging skills, the authors believe that evidence more favorably supports the demands of social living as the driving force behind brain evolution. In particular, Dunbar and Shultz outline results that point to the demands of advanced pair-bonding as the initiating factor. They argue that primate sociality differs from sociality in other taxa in that it is based on bonded relationships, which could account for why the social brain hypothesis (SBH) seems primarily viable for primates.

Dunbar and Shultz argue that energetic demands of the environment were not the selective pressure behind brain evolution, but rather acted only as constraints. That is to say, the technical demands of ecological success, such as efficiency of metabolism, only limited the degree to which brains could grow. While a more efficient metabolism may have allowed for larger brains to develop, this offers little evidence to believe that the forces driving the evolution of metabolic efficiency were themselves responsible for the growth of brains, especially when one considers the sheer cost of maintaining large brains. Instead, Dunbar and Shultz believe brain evolution was driven by the cognitive demands of group living, which offers valuable fitness benefits such as protections against predation.

In order to reconcile the SBH with ecological pressures, Dunbar and Shultz propose that individuals needed to develop flexibility (tolerance) in social situations to overcome conflicts arising from sharing resource space and potential mates. In other words, group members must be willing to sacrifice immediate, individual benefits in expectation of greater benefits for fitness afforded through cooperative group life. In primates, such flexibility is maintained through building strong social bonds. Evidence shows that in many species of birds, carnivores, bats, and ungulates, there is a qualitative relationship between brain size and mating strategy; those species that mate monogamously have relatively larger brains than those that are polygamous, in order to support the demands of mate selection stemming from the investment cost of selecting only one mate. This suggests that the demands of pair-bonding initiated the evolution of larger brains. Dunbar and Shultz believe that in early evolution, primates called upon the same cognitive skills necessary for the intensity of pair-bonding found in monogamous species to build strong bonds between social group members who were not even reproductive partners. It would therefore have been the cognitive demands of pair-bonding that initiated social brain evolution.


Social Components of Fitness in Primate Groups

Joan B. Silk

Science. 2007 Sep 7:317(5843), pp. 1347-1351.

DOI: 10.1126/science.1140734

In this paper, Silk discusses the fitness benefits allotted through social-group living.


Prospection: Experiencing the Future

Daniel T. Gilbert and Timothy D. Wilson

Science. 2007 Sep 7:317(5843), pp. 1351-1354.

DOI: 10.1126/science.1144161


Humans Have Evolved Specialized Skills of Social Cognition: The Cultural Intelligence Hypothesis

Esther Herrmann, Josep Call, María Victoria Hernández-Lloreda, Brian Hare, Michael Tomasello

Science. 2007 Sep 7:317(5843), pp. 1360-1366.

DOI: 10.1126/science.1146282


The New Synthesis in Moral Psychology

Jonathan Haidt

Science 18 May 2007 316: 998-1002

DOI: 10.1126/science.1137651


Social Neuroscience

Social Neuroscience -- A New Academic Journal

Decety & Keenan

Homepage for the Social Neuroscience journal. This site contains information on the journal, links to other related journals and sites, as well as the introduction from the first issue of Social Neuroscience. The introduction (Jean Decety, editor, and Julian Paul Keenan, deputy editor) outlines the field of social neuroscience, providing a summary of what "social neuroscience" is, as well as a general description of the methodologies and impacts of the field.


The Gregarious Brain

David Dobbs

New York Times Magazine, July 8, 2007.

See also: /exchange/brainstories/dobbs


Ron C. de Weijze's picture

Scanning for sociosis

Hello Ian, perhaps your research should be extended to psychiatry, for even though more or less forgotten, the work of Dutch psychiatrist J.H. van den Berg is about 'sociosis' which is a social desease expressing itself in communities having developed their own coping strategies with reality, which may not in all respects be valid in the long run. In the 50s there were no possibilies to scan brain activity when these socioses occurred and could only be treated on the psychiatrist's couch. Now it is a different story. I would very much welcome research of that kind!