Showing posts with label self-referential. Show all posts
Showing posts with label self-referential. Show all posts

Thursday, January 02, 2014

Art Reaches Within: Aesthetic Experience, the Self, and the Default Mode Network


From the open access Frontiers in Neuroscience, this is an intriguing study that asked subjects to rate images of artworks while in an fMRI scanner. The researchers found that regions in the medial prefrontal cortex that are known to be part of the default mode network (DMN) were positively activated on the highest-rated trials (presumably, these same regions were not activated during the lowest-rated trials).

Typically, at least in our current understanding, the DMN is somewhat off-line when we are engaged in externally oriented activity; conversely, it is more active when we are not engaged in a mental activity, for example, when we are daydreaming. However, the DMN is activated when the activities involve self-referential tasks or if tasks involve self-relevant information.

It seems that rating an artwork as a 4 (1-4 scale) required subjects to be somewhat self-referential, perhaps requiring them to determine why they felt moved by that particular work of art. If this is true, then the study adds to the data around the DMN being active in self-referential tasks. The authors reach the same conclusion:
This account is consistent with the modern notion that individuals' taste in art is linked with their sense of identity, and suggests that DMN activity may serve to signal “self-relevance” in a broader sense than has been thought so far.
It's an interesting article - give it a read.

Full Citation: 
Vessel EA, Starr GG, and Rubin N. (2013, Dec 30). Art reaches within: aesthetic experience, the self and the default mode network. Frontiers in Neuroscience; 7:258. doi: 10.3389/fnins.2013.00258

Art reaches within: aesthetic experience, the self and the default mode network
 
Edward A. Vessel [1], G. Gabrielle Starr [2], and Nava Rubin [3,4]
1. Center for Brain Imaging, New York University, New York, NY, USA
2. Department of English, New York University, New York, NY, USA
3. Center for Neural Science, New York University, New York, NY, USA
4. ICREA and DTIC, Universitat Pompeu Fabra, Barcelona, Spain

Abstract


In a task of rating images of artworks in an fMRI scanner, regions in the medial prefrontal cortex that are known to be part of the default mode network (DMN) were positively activated on the highest-rated trials. This is surprising given the DMN's original characterization as the set of brain regions that show greater fMRI activity during rest periods than during performance of tasks requiring focus on external stimuli. But further research showed that DMN regions could be positively activated also in structured tasks, if those tasks involved self-referential thought or self-relevant information. How may our findings be understood in this context? Although our task had no explicit self-referential aspect and the stimuli had no a priori self-relevance to the observers, the experimental design we employed emphasized the personal aspects of aesthetic experience. Observers were told that we were interested in their individual tastes, and asked to base their ratings on how much each artwork “moved” them. Moreover, we used little-known artworks that covered a wide range of styles, which led to high individual variability: each artwork was rated highly by some observers and poorly by others. This means that rating-specific neural responses cannot be attributed to the features of any particular artworks, but rather to the aesthetic experience itself. The DMN activity therefore suggests that certain artworks, albeit unfamiliar, may be so well-matched to an individual's unique makeup that they obtain access to the neural substrates concerned with the self—access which other external stimuli normally do not get. This mediates a sense of being “moved,” or “touched from within.” This account is consistent with the modern notion that individuals' taste in art is linked with their sense of identity, and suggests that DMN activity may serve to signal “self-relevance” in a broader sense than has been thought so far.


Introduction


The burgeoning field of neuroaesthetics attempts to address the mysteries of the human preoccupation with art by studying the underlying brain mechanisms. And, while understanding the artistic creative process itself is certainly a formidable challenge, many of the open questions concern the response to works of art by their viewers, listeners, and readers. What makes us so drawn to certain artistic creations, so influenced and moved by them? In recent years, we have learned a considerable amount from brain imaging studies about the neural correlates of aesthetic experience and how they relate to sensory, reward, and emotion neural processes (for reviews see Di Dio and Gallese, 2009; Brown et al., 2011; Chatterjee, 2011; Nadal and Pearce, 2011). One aspect that has so far received little investigation is that of individual differences: although it is widely recognized that individuals can differ markedly in their aesthetic response, previous research in neuroaesthetics tended to utilize art pieces that were manipulated in a manner intended to have a consistent effect on observers' preferences or that were generally highly regarded and often, widely known (e.g., the Mona Lisa). It seems reasonable to expect that studying widely admired artwork can help uncover the universal aspects of aesthetic experience. But studying artworks that generate a diversity of responses can also be valuable. Brain imaging can, in principle, be used to probe the neural correlates of an experience in a manner dissociable from the external stimuli that gave rise to this experience. In particular, it is possible to capitalize on the differences in individual's responses to artworks to search for commonalities in brain activity associated with the aesthetic experience itself, irrespective of the stimulus properties of specific works of art that gave rise to it. We have used this strategy in a recent study (Vessel et al., 2012) and the results underscore its power and promise, by confirming known results while at the same time revealing new and hitherto unsuspected findings.

KEY CONCEPT 1. Neuroaesthetics
A multi-disciplinary field aimed at understanding the neural basis of aesthetic experience and behavior. This includes interactions with art-objects as well as aesthetic modes of interaction with non-art objects, such as faces, natural objects, and scenes.

KEY CONCEPT 2. Aesthetic experience
Aesthetics is a discipline concerned with the perception, appreciation, and production of art. Aesthetic experiences, such as looking at paintings, listening to music or reading poems, are linked to the perception of external objects, but not to any apparent functional use the objects might have. Aesthetic experience involves more than preference, encompassing a variety of emotional responses ranging from beauty to awe, sublimity, and a variety of other (often knowledge-based) emotions.

Highly Individualized Responses to Visual Art


As in much previous work in neuroaesthetics, we wished to compare fMRI brain activity during observation of visual art that elicited a high level of aesthetic appreciation with responses to unappreciated artworks. But there was an important difference: a primary goal of our study was to move away from the scenario whereby different observers tend to respond similarly to the art presented to them. (The rationale for this goal is explained below, section Neural Correlates of Aesthetic Appreciation: Two Distinct Activity Patterns). To achieve this, we collated a set of images of two-dimensional visual artwork spanning a wide variety of periods, regions, styles and genres (fifteenth to twentieth century, Western and Eastern works, including a range of representational and abstract genres). Importantly, although the images were taken from museum collections, the artworks were not commonly reproduced and were therefore novel to our observers. Moreover, the instructions to the participants emphasized that we were interested in their own, individual response (rather than in what may be the “normative” assessment of each artwork), and that aesthetic experiences may come in a variety of forms: “The paintings may cover the entire range from ‘beautiful’ to ‘strange’ or even ‘ugly.’ Respond on the basis of how much this image moves you.” Each observer (N = 16) was shown the same series of 109 color artworks (in randomized order) while being scanned using fMRI, and was asked to rate each artwork on a 4-point scale according to these instructions. For a list of artworks and other experimental details, see Vessel et al. (2012), Materials and Methods and List of Artworks.

Analysis of the behavioral responses revealed that responses were indeed highly individual: there was little agreement between observers regarding how moving each painting was (0.13 average correlation between the ratings of pairs of observers, computed over the entire set of images; SD = 0.17). This means that, on average, each image was rated as highly moving by one subset of observers and rated poorly by another subset of observers (Figure 1). These results stand in contrast with the rather high agreement obtained when observers make preference judgments for real-world scenes [e.g., 0.46 between-observer correlation in Vessel and Rubin (2010)] or attractiveness judgments for faces [0.41 correlation between pairs of strangers in Bronstad and Russell (2007); 0.40 in Honeköpp (2006)]. As we shall see below, the low agreement between individuals in terms of their aesthetic response is what allowed us to disentangle the external attributes of specific stimuli from the internal (neural) states to which they gave rise.

FIGURE 1

http://c431376.r76.cf2.rackcdn.com/52124/fnins-07-00258-HTML/image_m/fnins-07-00258-g001.jpg

Figure 1. Aesthetic appreciation of visual art is highly individual. (A) Two sample images from the set observers were shown. Images were reproductions of museum artworks that are not commonly reproduced (see Acknowledgments for image credits). Observers rated each image for how much the artwork “moved” them on a scale of 1 (lowest) to 4 (highest). (B) Ratings of all 16 observers for the two images in (A). As was typical for the artworks used in the experiment, observers differed widely in their response to the pair of images. In particular, some observers rated the top image (blue bars) to be highly moving, while others rated the bottom image (red bars) to be highly moving. (For this bar plot, observers were first sorted by their rating to the top image, then by their rating to the bottom image).
Another finding from the behavioral data that will play a role in interpreting the brain imaging results is that, on average, observers used the highest (“4”) rating significantly less than 25% of the time (mean: 16.7%; SD = 11.6%; 4 of 16 observers gave more than 25% “4” responses). This is interesting given that there was no special mention of the highest rating in our instructions, and that in rating sensory/perceptual attributes (e.g., perceived brightness) observers tend to distribute their responses across all available options. That the observers in our experiments behaved differently, and did not calibrate their responses so as to give a rating of “4” to roughly a quarter of the stimuli, suggests that they reserved this response for images which met a certain internal (and generally high) criterion.
 

Neural Correlates of Aesthetic Appreciation: Two Distinct Activity Patterns


The fMRI data were analyzed to compare responses during trials in which the artworks were highly-rated with trials of low-rated artwork. Contrasting brain activity between conditions that differ by the observers' own responses, or performance, has been used successfully in many domains of cognitive neuroscience (e.g., studying neural correlates of memory encoding by contrasting activity in subsequently-remembered and forgotten trials; Brewer et al., 1998; Wagner et al., 1998). But in the context of neuroaesthetics, extra care must be taken to dissociate neural correlates of the aesthetic experience itself from other aspects of brain activity elicited by the stimuli. As a simple example, suppose observers are presented with a set of paintings comprised mainly of portraits and landscapes, and suppose further that most of them happen to appreciate portraiture more than landscapes. Face-selective brain regions would then likely show up in a contrast between highly-rated and low-rated trials, but is it warranted to interpret their activity as pertaining to aesthetic experience? In this case, the (conjured) agreement in aesthetic preference is simple enough, and our knowledge of face-selectivity in the brain sound enough, to easily discern that the activity can be explained by other aspects of the stimuli (the types of objects depicted). But in fact, such potential confounds are present whenever there is high agreement between observers about the art: the highly-rated and low-rated trials in such cases correspond to different sets of (artwork) stimuli, which may well result in some differential activation unrelated to the aesthetic experience they produce. Conversely, high variability between different observers' aesthetic judgments alleviates the potential confound: in the limit of completely uncorrelated ratings, the highly-rated trials and the low-rated trials contain identical sets of stimuli (each contributed by a different observer to each set). This was therefore our motivation in creating a stimulus set that generated highly individualized responses: rating-specific neural responses would then not be attributable to the features of any particular artworks, thus allowing us to isolate neural correlates of the aesthetic experience itself.

We performed several different analyses, using both statistical activation maps and regions of interest (ROIs) generated from the same data set or from separate “localizer” runs. We first created whole-brain activation maps by contrasting the group-level brain response to the most moving trials (rated as “4”) with the responses to the least moving trials (rated as “1”). This “4-vs.-1” analysis revealed a network of regions distributed across posterior, anterior, and subcortical structures (Figure 2A; note that, in addition, extensive portions of visual sensory cortex were strongly activated by all stimuli, but the magnitude of response did not differ by rating; Figure 2B). This is consistent with conclusions from previous research using a variety of stimuli that multiple brain regions are engaged during aesthetic appreciation (Aharon et al., 2001; Blood and Zatorre, 2001; Cela-Conde et al., 2004; Kawabata and Zeki, 2004; Vartanian and Goel, 2004; Jacobsen et al., 2006; Koelsch et al., 2006; Di Dio et al., 2007; Kim et al., 2007; Yue et al., 2007; Calvo-Merino et al., 2008; Fairhall and Ishai, 2008; Cupchik et al., 2009; Ishizu and Zeki, 2011; Lacey et al., 2011; Salimpoor et al., 2011; Jacobs et al., 2012; Kuhn and Gallinat, 2012). Note that the large inter-observer variability in behavioral responses to our stimulus set means that the common (group-level) activation in the 4-vs.-1 contrast must reflect effects of the aesthetic experience itself, i.e., it could not be due to any attributes of particular art stimuli that gave rise to this experience. This is because, at the group level, the set of highly rated trials consisted mostly of the same images as the poorly rated trials (recall that for every image rated as high by one observer there was, on average, another observer that rated it as low). This also means, however, that our approach is more restrictive than that in some other studies, which could give rise to differences in the activations observed. We will not go here into details of comparing and contrasting the loci of activation with those previously reported in the literature (see Vessel et al., 2012). Instead, we focus below on those aspects most relevant for a novel and intriguing finding: the activation by highly moving stimuli of the default mode network (DMN).

FIGURE 2

http://c431376.r76.cf2.rackcdn.com/52124/fnins-07-00258-HTML/image_m/fnins-07-00258-g002.jpg

Figure 2. Distinct patterns of response to artworks as a function of their ratings in a distributed network of brain regions. (A) Center panel: a whole-brain analysis contrasting trials on which observers rated artworks as highly moving (4) vs. trials where artworks were given the lowest rating (1), showing a lateral (top) and ventral (middle) view of an inflated left hemisphere, and a coronal section (bottom) through the striatum (data thresholded at a False Discovery Rate of q < 0.05 in volumetric space and projected on a hemisphere of a single observer for visualization). Right-side panels: a linear increase with rating was observed for the activation loci in occipitotemporal cortex and some subcortical loci (shown here: left inferior temporal sulcus, lITS; left parahippocampal cortex, lPHC; left striatum, lSTR; see (Vessel et al., 2012) for additional ROIs and further detail). Left-side panels: a nonlinear, “step”-like response pattern was observed in the anterior activation loci; responses did not differ for images rated 1, 2, or 3, but were significantly elevated for images rated 4 (shown here: left inferior frontal gyrus par triangularis, lIFGt; left lateral orbitofrontal cortex, lLOFC). (B) Extensive portions of early visual cortex were strongly activated by all paintings, but the magnitude of fMRI response did not differ by rating.
The bar graphs surrounding the activation map in Figure 2A show fMRI response magnitude as a function of observers' ratings for select ROIs, revealing that different ROIs exhibited distinct response patterns. Moreover, ROIs could be grouped in two main categories: for one set of ROIs, response magnitudes varied linearly with rating (right-side panels: lITS, lPHC, and lSTR). The linear response pattern was observed in different variations in terms of its relation to the baseline (“rest”) level: in occipitotemporal cortex, higher ratings were accompanied by linearly changing BOLD signals that either increased well above a resting baseline (lITS, and lPHC) or, in one case, decreased well below it (rSTG, not shown). In subcortical regions, fMRI activity was suppressed below its resting level for low-rated stimuli and rose progressively to above-rest for highly rated stimuli [lSTR, bottom right panel; PRF, not shown; see Vessel et al. (2012) for ROIs not shown here and further details]. Since the 4-vs.-1 contrast selects for regions that responded differently to trials rated “4” compared with trials rated “1,” the pattern of response for the intermediate ratings of 2 or 3 in these regions is a priori unknown. It is therefore noteworthy that responses in these ROIs followed a linear trend so closely. Moreover, regions whose response patterns were significantly non-linear all showed the same distinct pattern, as follows.

A second category of regions revealed by the 4-vs.-1 contrast were characterized by a distinct “step” pattern: fMRI responses in those regions did not differ significantly for images rated 1, 2, or 3; only for the highest (4) rating was there a significant difference in response magnitude, and it was marked and dramatic (Figure 2A, left-side panels; see Vessel et al., 2012 for other examples; see also below, Figure 3). We performed several additional analyses in order to examine more closely the nature and spatial distribution of these nonlinear “step” responses. A whole-brain analysis contrasting the highest-rated trials with an average of all other trials (4-vs.-321; Vessel et al., 2012) gave us more power to detect regions that may not have reached the significance threshold in the 4-vs.-1 contrast due to the lower number of trials. A conjunction was subsequently computed to specifically capture the regions that, while showing a differential response to the highest-rated stimuli (“4”), showed no significant differences in responses within the lower ratings (1, 2, and 3). The resulting statistical map contained large swaths of highly significant differences in several regions known to be part of the DMN, and further examination indicated that the pattern of responses in those regions consisted of a strong deactivation in trials rated 1, 2, or 3 (with no significant differences in magnitude), which was greatly alleviated or even eliminated in the highest-rated trials [“4”; see Vessel et al. (2012), Figure 6]. To better underscore the commonalities and differences from what is currently known about the DMN, below we represent our results in a different format than before, which is modeled after that used in the DMN literature.

FIGURE 3

http://c431376.r76.cf2.rackcdn.com/52124/fnins-07-00258-HTML/image_m/fnins-07-00258-g003.jpg

Figure 3. The default mode network (DMN) deactivation during task performance is alleviated when viewing highly moving artworks. (A) Lateral (left) and medial (right) views of an inflated cortical surface are overlaid with statistical maps comparing fMRI responses during task (viewing and rating of artworks) vs. “rest” periods. Maps were computed separately for trials from each of the four possible ratings, 1 (top) to 4 (bottom). The warm colors indicate greater fMRI response during task; the cool colors indicate greater response during rest (“deactivation”; data were thresholded at a False Discovery Rate of q < 0.05 before projection onto one observer's inflated cortex). In trials rated 1, 2, or 3 (top three panels) there were deactivations in medial prefrontal cortex (MPFC), posterior cingulate cortex (PCC), lateral temporal cortex (LTC), temporoparietal junction (TPJ), and superior frontal gyrus (SFG). The suppression was greatly reduced for the highest-rated trials (4; bottom panel). (B) The spatial pattern of deactivation during the lower-rated trials (1–3) closely resembles that of the default mode network [DMN; image adapted with permission from Fox et al. (2005) Copyright 2005 National Academy of Sciences, U.S.A.]. (C) Average fMRI response in the MPFC region of interest (ROI) was markedly and uniformly below rest for trials rated 1, 2, or 3, but was not different from rest for the highest-rated trials (4). (D) fMRI signal timecourse in the MPFC for the lower-rated trials (cyan) and the highest-rated trials (magenta). Note that activity initially fell below its level during rest also for the highest rated trials, yet it rapidly recovered and then proceeded to increase above rest level. The fMRI response used for both C and D was estimated from an ROI defined via a contrast of the response on “4” trials vs. the other trials (4-vs.-321), conjoined with a map of regions showing no difference in the low-rated trials. The timecourses for each rating level were extracted by modeling the average timecourse from this ROI as a set of four finite impulse response functions (Dale and Buckner, 1997).
Figure 3A shows statistical activation maps contrasting the task-induced fMRI responses with “Rest”—intervals interspersed between the trials when only a blank screen was shown—overlaid on the inflated surface of the left hemisphere. The maps were generated separately for each of the four sets of trials corresponding to the four possible ratings (from top to bottom: 1-vs.-Rest to 4-vs.-Rest). Large regions in occipital cortex, as well as portions of parietal and frontal cortex, showed activation above rest for all four rating levels (warm colors, red-yellow). The cool colors (blue–green) denote regions that showed a reduced fMRI signal during viewing and rating of the artworks, compared to during rest. For the sets of trials rated 1, 2, or 3 (top three panels), extensive regions of reduced activity can be seen; their anatomical loci and spatial distribution closely resembles that observed in studies that contrasted activity during a wide range of cognitive and perceptual tasks with periods of rest (Shulman et al., 1997; Simpson et al., 2001), shown in Figure 3B (adapted from Fox et al., 2005). Specifically, reduced activity was observed in the medial prefrontal cortex (MPFC), posterior cingulate cortex (PCC), precuneus (PCu), temporo-parietal junction (TPJ), lateral temporal cortex (LTC) and superior frontal gyrus (SFG). Studies of blood flow and oxygen utilization indicate that the baseline level of these regions—that measured during rest—corresponds not to a lack of activity, but rather to activity associated with an ongoing, organized “default mode” of brain processing, which is suspended during performance of tasks that require externally directed attention (Gusnard and Raichle, 2001; Raichle et al., 2001). The reduced fMRI response in regions of this default mode network (DMN) during task performance is therefore widely referred to as deactivation (although the mechanisms giving rise to it are not fully understood).

KEY CONCEPT 3. Default mode network
A network of brain regions typically found to be suppressed when observers engage in externally oriented tasks, which includes the medial prefrontal cortex (MPFC), posterior cingulate cortex (PCC), temporo-parietal junction (TPJ), lateral temporal cortex (LTC), superior frontal gyrus (SFG) and the hippocampus. Patterns of spatial correlation measured in the absence of directed tasks (resting state fMRI) support this network structure and suggest that the DMN is composed of midline hub regions (MPFC, PCC) and two subsystems.
In contrast with the pattern observed for trials rated 1, 2, and 3, DMN regions showed markedly less deactivation during the highest-rated trials (“4”; bottom panel in Figure 3A). Indeed, in some portions of the DMN—most notably, in the MPFC—the deactivation seems all but gone. ROI analysis confirmed that the MPFC was strongly and uniformly deactivated for lower-rated trials (1–3), but not at all during those trials when the artworks were given the highest rating (4), resulting in a step-like response pattern [Figure 3C; for plots of several other DMN components, see Vessel et al. (2012)].
 

The Default Mode Network and Self-Referential Mental Processing


A defining characteristic of the DMN—indeed, how it was discovered—is that it is suppressed when observers are engaged in demanding tasks that require them to focus on external stimuli, compared with its level of activity during passive viewing or periods of rest between the tasks (Shulman et al., 1997; Buckner et al., 2008). The ubiquity of DMN deactivation during many different cognitive tasks with a variety of stimuli and response demands, along with studies of functional connectivity during rest, have led to the view that the DMN represents a “task-negative” network of brain regions that normally functions in an anti-correlated manner from “task-positive” networks such as sensory-semantic pathways and the dorsal attention network (Ingvar, 1979; Corbetta and Shulman, 2002; Fox et al., 2005; Buckner and Carroll, 2007). The finding that, in our own task, the cortical regions that overlap with previously identified components of the DMN (MPFC, PCC, TPJ, LTC) showed significant deactivation below their baseline (rest) level during a majority of the trials, those rated 1–3 (Figure 3A, top three panels) is therefore consistent with what is known about the DMN. From this same perspective, the dramatic reduction of deactivation in the trials rated “4” (Figure 3A, bottom panel) and its complete absence in the MPFC (Figure 3C) therefore seems puzzling. But consideration of additional findings about the DMN offers a potential explanation.

Following its initial identification, further research showed that the DMN regions can maintain their baseline activity not only during periods of (waking) rest, but that they can escape deactivation, or even become activated above baseline, also during the performance of structured tasks. Ventral portions of the MPFC are involved in affective decision making processes, including (but not restricted to) encoding the subjective value of future rewards and assessing the emotional salience of stimuli (Bechara et al., 1999; Knutson et al., 2005; Kringelbach, 2005; Kable and Glimcher, 2007; Schmitz and Johnson, 2007; Levy and Glimcher, 2011). The anterior and dorsal portions of MPFC are active in tasks involving self-knowledge such as making judgments about oneself as well as about close others (family and friends), self-relevant moral decision-making (Reniers et al., 2012) and in “theory of mind” tasks that require gauging others' perspectives (Zysset et al., 2002; Ochsner et al., 2004, 2005; Amodio and Frith, 2006; Mitchell et al., 2006; Enzi et al., 2009; Andrews-Hanna et al., 2010; Whitfield-Gabrieli et al., 2011). The PCC and medial temporal lobe regions are active during tasks that involve retrieving autobiographical memories as well as planning or simulating the future (Buckner and Carroll, 2007; Buckner et al., 2008; Andrews-Hanna et al., 2010).

The DMN is thus emerging as a highly interconnected network of brain regions that support self-referential mental processing (Northoff et al., 2006). Such processing is, of course, ubiquitous in everyday life and is undoubtedly important for normal functioning. In experimental settings it can occur spontaneously (e.g., as “mind wandering” during periods of rest) but it can also be triggered in structured tasks, by external stimuli that cause observers to draw on self-referential information (intentionally or automatically), or to engage in inwardly focused attention. Could this have been the case with the images that our observers rated as “highly moving”? We propose that the answer is yes, as detailed in the account provided below.

KEY CONCEPT 4. DMN and self-referential mental processing
Structured tasks can activate the DMN if they require some self-referential processing (e.g., introspection, autobiographical memory recall). Similarly, it is presumed that the DMN is metabolically active during baseline non-task periods (e.g., fixation or “rest” conditions) because observers engage in such processes spontaneously.

Intense Aesthetic Experience: A (Non-Personal) External Stimulus Reaches the Self


Taste in art is highly individual and can be hard to predict by even the most well-informed bystander (e.g., Bell and Koren, 2007), yet it is strongly felt. Indeed, many individuals consider their artistic taste to be an important part of their identity, their sense of who they are. This is not limited to connoisseurs of “high art”: from teenagers whose tumultuous struggles for self-determination are conducted to the soundtrack of meticulously compiled music collections, to adults of all ages who repeatedly turn to their favorite genres of fiction or film to escape the tedium of their daily lives, our taste in art is intertwined with the choices we make about how to spend our time and with whom to spend it, and as such it is part of who we are. How does this come about? What gives certain artworks their mysterious “pull”? Our data say nothing about this in terms of the attributes of the artwork itself. (Whether this will remain a mystery forever or may yield to future research is an interesting question that will not be discussed here). But our results suggest that the strong effect of certain artworks can be understood in terms of the physiological state they generate and how this state is experienced, or interpreted, by the observer.

We propose that certain artworks can “resonate” with an individual's sense of self in a manner that has well-defined physiological correlates and consequences: the neural representations of those external stimuli obtain access to the neural substrates and processes concerned with the self—namely to regions of the DMN. This access, which other external stimuli normally do not obtain, allows the representation of the artwork to interact with the neural processes related to the self, affect them, and possibly even be incorporated into them (i.e., into the future, evolving representation of self). This hypothesis gains considerable support from the way that the fMRI responses evolved over time in the MPFC, the region most associated with evaluations of self-relevance. As can be seen from the time course plots in Figure 3D, immediately following stimulus presentation the fMRI signal in the MPFC fell below baseline for all images, i.e., also for those images that were (later) rated by the observer as highly moving (4). Thus, the initial predisposition of this DMN region was, for all external stimuli, to deactivate. But in contrast with the MPFC response to the artworks rated 1, 2, or 3, which was suppressed during image presentation and remained below baseline throughout the subsequent recovery (Figure 3D, cyan line), in the 4-rated trials activity started recovering soon after stimulus presentation and then continued to rise above baseline (magenta line). This is reminiscent of the MPFC recovery from deactivation observed when a highly self-relevant stimulus such as one's own name is presented in a stream of self-irrelevant stimulation, as in the “cocktail party effect” (e.g., Cherry, 1953; Bargh, 1982; Wood and Cowan, 1995; Perrin et al., 2005). But why should a hitherto unseen artwork, that has no a priori personal relevance for the observer, have this effect of engaging the DMN system? Again, we cannot say what attributes make specific artworks so exquisitely attuned to an individual's unique makeup. And yet this hypothesis provides a coherent explanation of our data in that it is consistent not only with what we know about the DMN, but also with what we know about art.

Great art is, almost by definition, universal: the wide appeal it commands comes from a connection with fundamental aspects of human nature and human cognition (Kant, 1790/1987). Yet, at its best, art in any of its forms—visual art, music, literature, etc.—can feel strikingly personal. Intense aesthetic experience often carries with it a sense of intimacy, “belonging,” and closeness with the artwork. It may be hard to imagine that the experiences of our observers, lying in an MRI scanner watching images of little-known artworks selected by an experimenter who knew nothing about them, reached the profound levels that give art its intense power. And yet the data are compellingly in line with the phenomenology of aesthetic experience: in the small subset of the trials that observers rated as “highly moving,” DMN regions and in particular the MPFC were released from deactivation and even activated above baseline, a hallmark of self-relevant neural processing. Perhaps the key to this was in our experimental design, which relied on a stimulus set that maximized individual differences in behavioral response. As already mentioned, the original motivation for this design was to measure neural correlates of aesthetic experience in the absence of potential confounds with effects of stimulus attributes. But the emphasis on a diversity of artistic styles and topics may have, serendipitously, also increased the chances that a few of the artworks resonate with each observer in a particularly powerful way.

Note that the “resonance” between certain artworks and observers' sense of self that, we propose, occurs during intense aesthetic experience, is different from explicitly self-referential emotions such as pride, shame, guilt and embarrassment, as these involve an appraisal of self-responsibility for an event (Silvia, 2012). It is also interesting to note in this context that intense aesthetic experience can sometimes be thrillingly bidirectional: not only does the perceiver feel as if they understand the artwork, but there is a sense that the artwork “understands” the perceiver, expressing one's own innermost thoughts, feelings, or values. The latter sense points to the possibility that it is the artist, not the artwork, who has understood something deep about the perceiver's experience; hence the intensely personal connection felt by many people toward favorite artists who are, after all, strangers to them. In some cases, this bidirectionality is accompanied by a perceived or real congruence with the intentions of the artist (Jucker and Barrett, 2011; Tinio, 2013). Thus, unlike in self-referential emotions, in aesthetic experience the relation to others is not focused on appraisal but on a sense of understanding, gained insight and meaning. The extraction of meaning has been suggested previously as a primary factor of aesthetic experience (Martindale, 1984; Leder et al., 2004). But, while those authors suggest that an appeal to self-related information is but one way in which viewers extract meaning from artwork, the release of the DMN from suppression on only the trials rated “4” suggests that, in fact, self-relevance is an integral aspect of intensely moving aesthetic experience.

What internal signal did the observers use to provide their responses? It is tempting to think that they were able to detect the unusual release from deactivation in the DMN when viewing artworks which they (later) rated “highly moving,” and that they based their responses on this internal signal. Indeed, the MPFC and PCC respond to self-relevant information even when there is no explicit requirement to evaluate self-relevance, and such information is in fact task-irrelevant (Moran et al., 2009; Reniers et al., 2012). Perhaps observers conferred the highest rating on those artworks that invoked in them a sense of self-relevance, even though they were not instructed to do so, and may well be unable to explicitly state this as their strategy. Yet given the poor temporal information provided by fMRI, it is too early to rule out the possibility that responses on the “4” trials arose from posterior regions whose activity grew linearly with rating or from other frontal regions that showed positive activation for only the “4” trials, and that the release from suppression in the DMN for highly moving artworks occurred subsequent to the evaluation. A recent MEG study of aesthetic appreciation reported coherence between frontal midline, posterior and temporal regions that was detectable 1 s after onset of images deemed “beautiful” (1000–1500 ms analysis window) but not in an earlier epoch (250–750 ms; Cela-Conde et al., 2013). This finding is consistent with our proposal that the release of the DMN from suppression for intensely moving artworks occurs subsequent to an initial perceptual and semantic analysis, and early enough to be a potential basis for response selection; however, it leaves open the question of how, in time, explicit evaluation relates to these dynamics.

A coactivation of the DMN and stimulus-driven sensory system as we have observed for strongly moving aesthetic experiences has so far not been reported in other contexts. Yet, if our self identity is to be influenced by the world we inhabit, it may be that similar moments should occur with greater frequency than would be expected based on the current conceptualization of the DMN as a network that is invariably suppressed during mental activity which is directed at the external world. It may be that our findings are just the “tip of the iceberg”—i.e., that instances of resonance between external stimuli and internal, self-related processing are more commonplace in daily life than what has so far been captured in fMRI experiments in the laboratory. By that view, much of our existence may be well-served by switching between periods of dominance of externally-directed (“task-positive”) brain networks over the DMN and vice versa, but those periods are punctuated by significant moments when our brains detect a certain “harmony” between the external world and our internal representation of the self—allowing the two systems to co-activate, interact, influence and reshape each other.


Conflict of Interest Statement


The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
 

Acknowledgments


This project was supported by an ADVANCE Research Challenge Grant funded by the NSF ADVANCE-PAID award # HRD-0820202 and by the Andrew W. Mellon Foundation (as a New Directions Fellowship). Cloud Study, c. 1822. John Constable. Oil on paper, 29.21 × 48.26 cm. The Frick Collection, Bequest of Henrietta E.S. Lockwood in memory of her father and mother, Ellery Sedgwick and Mabel Cabot Sedgewick, 2001.3.134. An Ecclesiastic, c. 1874. Mariano José Maria Bernardo Fortuny y Carbo. Oil on panel, 19 × 13 cm. The Walters Art Museum, Bequest of William T. Walters. 37.150.

References available at the Frontiers site.


Saturday, August 17, 2013

Anne Beate Reinertsen, PhD - Welcome to My Brain


Wow. This paper has, perhaps, the most convoluted and self-reflexive abstract I think I have ever seen. And yet I suspect that is the point - the author is using her own brain function as the research design for a qualitative study of the subject ("recursive, intrinsic, self-reflexive as de-and/or resubjective always evolving living research designs").

The author employs the Möbius Strip as an image for her ideas.

File:Möbius strip.jpg
A Möbius strip made with a piece of paper and tape. If an ant were to crawl along the length of this strip, it would return to its starting point having traversed the entire length of the strip (on both sides of the original paper) without ever crossing an edge.

Interesting . . . "auto- brain- biography - ethnomethodology attempt."

Full Citation:
Reinersten, AB. (2013, Jul 12). Welcome to My Brain. Qualitative Inquiry, XX(X); 1-12. doi: 10.1177/1077800413489534

Abstract 
This is about developing recursive, intrinsic, self-reflexive as de-and/or resubjective always evolving living research designs. It is about learning and memory cognition and experiment poetic/creative pedagogical science establishing a view of students ultimately me as subjects of will (not) gaining from disorder and noise: Antifragile and antifragility and pedagogy as movements in/through place/space. Further, it is about postconceptual hyperbolic word creation thus a view of using language for thinking not primarily for communication. It is brain research with a twist and becoming, ultimately valuation of knowledges processes: Becoming with data again and again and self-writing theory. I use knitting the Möbius strip and other art/math hyperbolic knitted and crocheted objects to illustrate nonbinary . . . perhaps. Generally; this is about asking how-questions more than what-questions.
The article is available to read or download at Scribd, but here is a taste of the first couple of pages:

Introduction and the Möbius Strip
“So freedom of thought exists when I can have all possible thoughts; but the thoughts become property only by not being able to become masters. In the time of freedom of thought, thoughts (ideas) rule; but, if I attain to property in thought, they stand as my creatures. 
 If the hierarchy had not so penetrated men to the innermost as to take from them all courage to pursue free thoughts, that is, thoughts perhaps displeasing to God, one would have to consider freedom of thought just as empty a word as, say, a freedom of digestion.  
According to the professionals’ opinion, the thought is given to me; according to the freethinkers’, I seek the thought. There the truth is already found and extant, only I must—receive it from its Giver by grace; here the truth is to be sought and is my goal, lying in the future, toward which I have to run.” (Stirner, 2012: Kindle locations 5514-5519).
Welcome to my brain. It is plastic, mentally creative, and physically adaptable just like yours if we want to. It is a chaotic noisy place wanting to produce results. I will not let one area dominate however. Also, there are places I do not want to go. Cortex- Hippocampus- learning and memory Cognition, emotional, sensory, bodily, social centers . . . —in the brain . . . So first; start with a long rectangle (ABCD) made of paper. Then give the rectangle a half twist. Third; join the ends so that A is matched with D and B is matched with C. Now you have created a continuous one-sided surface from this rectangular strip only by rotating one end 180° and attaching it to the other end. The brain is/has architecture (The Cortex) and neurons form networks forming larger networks processing information. Networks that for example allow us to recognize and code space, develop tools and navigate thus decide, make matter, and plan ahead. These are processes of orientation in/through space. Navigating and thinking about navigating simultaneously. Movements and moving and words as thinking tools: Complex processes of building complex representations of compass distances—learning and memory—measurements. And important: Thinking/planning also without direct sensory impulses: Thus having the ability to generate new ideas and ideas about the future too . . . in the brain. To live such complexity I turn to “knitting around” Möbius Band Scarves (Zimmerman, 1989) as you will see below. Very easy but not; I had to rehearse. Thus calming but not, product oriented nice and warm—comfortable—if I finish but also while doing.
As you now know, this nonorientable surface is called a Möbius Strip or Möbius Band, named after August F. Möbius, a 19th century German mathematician and astronomer, who was a pioneer in the field of topology. Möbius, along with his contemporaries, Riemann, Lobachevsky, and Bolyai, created a non-Euclidean revolution in geometry. Möbius strips have found a number of applications that exploit a remarkable property they possess: one-sidedness. Joining A to C and B to D (no half twist) would produce a simple belt-shaped loop with two sides and two edges—impossible to travel from one side to the other without crossing an edge. But, as a result of the half twist, the Möbius Strip has only one side and one edge, and I am the no-one-sided teacher/researcher/professor in education: a nonorientable surface with a boundary—architecture—Cortex—and you. Thus I/you/we must/can make choices and make knowledges matter. And sometimes I am forced or I force myself to go places. Qualia: the subjective experience of things, a property of something . . . its feel or appearance perhaps rather than the thing itself . . .: This is about finding the out there in the in here. 
Here is what another teacher/researcher/professor, and this time in brain research at Center for Biology of Memory, Trondheim, Norway (CBM): www.ntnu.no/cbm, says about what he does, knows and thinks: “The Hippocampus is a part of the brain we know is relevant for learning and memory. There are brain structures there that are involved. The brain research area has exploded the recent years but still we are in the beginning of discovering general rules about how the brain processes information. Experimental evidence based science does not—must not/cannot—move too fast. We do not know that much and other so quickly. At least if we speak about evidence. Grounded scientific research is a privilege” (Interview, October 28th, 2011). 
At CBM they started to study memory but ended up in studying sense of place/space: “Now we know that this sense is closely linked to memory. It is almost like a human GPS with grid cells in the part of the brain called the Entorhinal Cortex. Grid cells collaborate with other specialized nerve cells with complementary roles in the sense of place/space and direction. Together they build a map in the brain. And the brain uses this map to orient itself in both familiar and unfamiliar environments. Signals create a coordination system in which positions can be registered. They register our movements and are closely linked to our memory. The grid cells do not reflect signals that come in from the outside from any of our senses. The grid patterns are made by the brain itself. Therefore we can use the grid cells as a way of understanding how activity patterns emerge in the brain. The grid is opening up new possibilities to study how the brain stores information” (Moser & Moser, in Jacobsen, 2012). 
“It is very difficult and risky to deduct anything directly between brain research and pedagogy, but one thing that we might say is that if you want to learn something different and other, it might be sensible to shift between places: Learning different things in different places that is” (Interview, October 28th, 2011). 
Ethnomethodology is an approach to sociological inquiry as the study of the everyday methods that people use for the production of social order (Garfinkel, 1967). It is also called bottom-up microsociology and a member’s methods inquiring into common sense knowledge, self organizing systems, and situated natural language. The aim is to document methods and practices through which society’s members make sense of the world. It is in itself however not a method. It does not have a set of formal research methods or procedures. 
Knitting is a repetitive visual spatial task. These tasks also include e.g. running and folding origami and can put our brain into the state of Theta (Retrieved Dec.7th. 2012). States of Theta can increase creativity, lower stress/anxiety and increase objectivity in difficult situations. Theta is also that state between falling asleep and waking up when we seem to have all of our best ideas. Because the brain is focusing on (literally) the task at hand, it isn’t as judgmental and has lower standards/barriers. Theta is nonjudgmental, more observant . . . objective? No but I try. And as you will see below, I try to hold my brain in my hands.
I and My Research . . . Brain . . . Questions
“Some things benefit from shocks; they thrive and grow when exposed to volatility, randomness, disorder, and stressors and love adventure, risk, and uncertainty. Yet in spite of the ubiquity of the phenomenon, there is no word for the exact opposite of fragile. Let us call it antifragile. Antifragility is beyond resilience or robustness. The resilient resist shocks and stays the same; the antifragile gets better” (Taleb, 2012, p. 3). 
This article is therefore about developing recursive intrinsic self-reflexive as de- and/or resubjective always evolving living research designs. Inquiry perhaps full stop—me: An auto-brain—biography and/or a brain theorizing itself; me theorizing my brain. It is thus about theorizing bodily here brain and transcorporeal materialities, in ways that neither push us back into any traps of biological determinism or cultural essentialism, nor make us leave bodily matter and biologies behind. It is an attempt of seeing the real as/through/in its material-discursive coconstitutive complexity and produce research from within an ontology and epistemology where ‘matter and meaning are mutually articulated’ (Barad 2007, p. 152). It is about learning and memory cognition and experiment poetic and/or creative pedagogical science; learning ultimately pedagogy as movements in/through space. 
It is brain research with a twist and becoming, ultimately valuation of knowledges process; a personal antifragile will born from knowledge. I use knitting and other, as you will see, to illustrate or rather live nonbinary. First, I will write more about research designing, second, about knitting Möbius bands. Third, I will philosophize a bit with Socrates, Meno and Plato: “Meno’s Paradox,” or “The Paradox of Inquiry” (Meno 80d-e) and Max Stirner (1806-1856) over learning and memory, one-sidedness, antifragility research, pedagogy, and will only to end in wonder. Eventually, this is a philosophical brain journey in which the question “how do you know” is more difficult but vital to ask than the “what do you know” question we traditionally have asked both ourselves and our students through years. 
I treat theory, transcribed interview notes, pieces of art, creating knitting Möbius band scarves and myself as data (text). Data shapes and negotiate. Data are shaped and negotiated. There are data dilemmas—paradoxes. I am at hearing of the data; text and textuality. Thus I am (my own) data but as a “montage” in which “several different images are superimposed onto one another” (Denzin and Lincoln, 2003, p. 6). Several different texts: My brain. My knowledge of my ordinary affairs, of my own organized enterprises, as part of the same setting that makes it orderable. 
It is broad and multifaceted and with open-ended references to any kind of sense-making procedure, a domain of uncharted dimensions my auto- brain- biography - ethnomethodology attempt. 
I turn knitting, art into data and tool to see other and beget thinking; activate brain cells—circuits—inquiring minds; experiment—poetic—creative—pedagogical science—and language . . . making, de/re/constructing the world? I told you this was chaotic and noisy and my own moving sensations of sound touch taste and smell. And further, my amalgamations of images in order to make a very unique image of my own, and mine. Seeking to describe the common sense methods through which I produce myself as teacher/researcher. A member’s methods; my methods.

Thursday, August 08, 2013

Role of Mid-Line Brain Regions in Socio-Emotional Function and Self-Evaluation


In this new article from Frontiers in Human Neuroscience, Flagan and Beer look at the ways in which the mid-line regions of the brain function individual and as a network. The authors propose a new framework . . .
to account for the contribution of MPFC, VACC, and MOFC to social cognition. MPFC is broadly implicated in self-evaluation but may be characterized by a ventral to dorsal division when evaluating others based on their intimacy. Certainty about evaluation may better characterize the contribution of DMPFC to social cognition than the correction of self-projection. The association between VMPFC and self-relatedness will be clearer once future research disentangles shared emotional and cognitive properties of evaluation of self and close others. Further, previous research has failed to take into account the fundamental role that motivation has in self-evaluations. As a result, the role of VACC and MOFC in social cognition has been obscured until recently. VACC may mediate bottom-up sensitivity to information based on its potential for helping us evaluate ourselves and others the way we want. MOFC may mediate top-down motivational influences on self-evaluation. 
Here are the main brain regions involved and their roles (briefly):
  • Medial PreFrontal Cortex (MPFC): Broadly modulated by encoding and remembering information in relation to the self , but subdivisions within MPFC are differentially modulated by the evaluation of close others (ventral MPFC) and the evaluation of other social targets (dorsal MPFC
  • Ventral MPFC (VMPFC): Certainty in evaluation
  • Dorsal MPFC (DMPFC): Socioemotional significance or cognitive quality of information?
  • Ventral Anterior Cingulate Cortex (VACC): Sensitivity to information based on its potential to fulfill motivation
  • Medial OrbitoFrontal Cortex (MOFC): Shifts in evaluation standards under threat
Here is the article in full - it's very geeky, but it's also very interesting for anyone seeking a better understanding of how our brains create social values and relate social experience to the "self."


Three ways in which midline regions contribute to self-evaluation



Taru Flagan and Jennifer S. Beer
  • Department of Psychology, University of Texas at Austin, Austin, TX, USA
ABSTRACT
An integration of existing research and newly conducted psychophysiological interaction (PPI) connectivity analyses suggest a new framework for understanding the contribution of midline regions to social cognition. Recent meta-analyses suggest that there are no midline regions that are exclusively associated with self-processing. Whereas medial prefrontal cortex (MPFC) is broadly modulated by self-processing, subdivisions within MPFC are differentially modulated by the evaluation of close others (ventral MPFC: BA 10/32) and the evaluation of other social targets (dorsal MPFC: BA 9/32). The role of DMPFC in social cognition may also be less uniquely social than previously thought; it may be better characterized as a region that indexes certainty about evaluation rather than previously considered social mechanisms (i.e., correction of self-projection). VMPFC, a region often described as an important mediator of socioemotional significance, may instead perform a more cognitive role by reflecting the type of information brought to bear on evaluations of people we know well. Furthermore, the new framework moves beyond MPFC and hypothesizes that two other midline regions, ventral anterior cingulate cortex (VACC: BA 25) and medial orbitofrontal cortex (MOFC: BA 11), aid motivational influences on social cognition. Despite the central role of motivation in psychological models of self-perception, neural models have largely ignored the topic. Positive connectivity between VACC and MOFC may mediate bottom-up sensitivity to information based on its potential for helping us evaluate ourselves or others the way we want. As connectivity becomes more positive with striatum and less positive with middle frontal gyrus (BA 9/44), MOFC mediates top-down motivational influences by adjusting the standards we bring to bear on evaluations of ourselves and other people.

Full Citation:
Flagan T, and Beer JS. (2013, Aug 2). Three ways in which midline regions contribute to self-evaluation. Frontiers in Human Neuroscience, 7:450. doi: 10.3389/fnhum.2013.00450

Introduction

The speculation that some midline regions contribute to self-processing stems from research conducted over a decade ago (Beer et al., 2006b). What have we learned since those initial studies found that medial prefrontal cortex (MPFC) is modulated by encoding and remembering information in relation to the self (e.g., Kelley et al., 2002; Macrae et al., 2004; Ochsner et al., 2005)? This article draws on existing research and newly conducted psychophysiological interaction (PPI) analyses to describe a new framework for the contribution of how a subportion of midline regions to social cognition (see Figure 1). The new framework builds on previous discussions by (a) positing a new role for the MPFC in social cognition and (b) moving past the MPFC to consider the importance of ventral anterior cingulate (VACC: BA 25) and medial orbitofrontal cortex (MOFC: BA 11) in aiding motivational influences on social cognition. Recent meta-analyses suggests that there are no midline regions that are exclusively associated with self-processing. For example, meta-analyses of studies of social evaluation (i.e., traits, personal abilities, etc.) find that the MPFC likely mediates psychological processes that are brought to bear on self-evaluation but also evaluations of other kinds of people (e.g., close others vs. non-close-others: Ochsner et al., 2005; Qin and Northoff, 2011;Murray et al., 2012; Roy et al., 2012). Whereas it was once thought that regions within VMPFC (BA 10/32) and DMPFC (BA 9/32) mediated person evaluation through the correction of self-projection, there are a number of issues that must be addressed before strong conclusions can be drawn. For example, current research provides more consistent evidence for the role DMPFC in certainty about self-evaluation even during tasks that require evaluations of other people. Furthermore, recent research suggests that VACC and MOFC are just as importantly involved in social cognition as the MPFC. Neural models of social cognition have not incorporated motivated processing which is a fundamental element of psychological models of the self (Beer, 2007). A growing body of research suggests that motivational influences on self- and other-evaluation are mediated by VACC and MOFC. VACC may mediate bottom-up sensitivity to information based on its potential for helping us evaluate ourselves or others the way we want (Beer, 2012a). MOFC may mediate top-down motivational influences on self-evaluation. Taken together, the new framework highlights the progress that has been made over the past decade: MPFC is involved in social cognition but does not mediate “self-specific” processes and two additional regions, VACC and MOFC, play an important role in motivated self- and other-evaluation.
FIGURE 1 

Figure 1. A framework of cortical midline structures implicated in self-evaluation. DMPFC, dorsomedial prefrontal cortex (BA 9, 32); VMPFC, ventromedial prefrontal cortex (BA 10, 32); VACC, ventral anterior cingulate cortex (BA 25); MOFC, medial orbitofrontal cortex (BA 11, 12).
A New Conceptualization of the Role of MPFC in Social Cognition: Certainty in Evaluation? 
While much has been learned about the role of MPFC in social cognition in the past decade, much more remains to be known. A series of studies in the early 2000s found that MPFC (BA 9/10/32) was modulated by both self-evaluation and evaluations of a political figure but modulation was greatest for self-evaluation (e.g., Kelley et al., 2002; Macrae et al., 2004; Ochsner et al., 2005). This research sparked interest in testing the possibility that functional MPFC subdivisions distinguished self-processing from the evaluation of other people. In contrast to this possibility, recent meta-analyses have shown that MPFC (BA 9/10/32) modulation is not exclusive to self; this region is modulated by both self-processing and processing about other people (Ochsner et al., 2005; Qin and Northoff, 2011; Murray et al., 2012; Roy et al., 2012). Instead of a self vs. other distinction, meta-analyses suggest a close other vs. non-close-other distinction. A ventral subdivision of this MPFC region (BA: 10/32, see Figure 1) is associated with self-processing and evaluations of close others (Ochsner et al., 2005; Qin and Northoff, 2011; Murray et al., 2012). A more dorsal subdivision (BA 9/32, see Figure 1) is associated with self-processing and evaluations of non-close-others (Ochsner et al., 2005; Qin and Northoff, 2011; Murray et al., 2012). Therefore, the next step toward understanding the contribution of MPFC to social cognition should be focused on understanding the psychological significance of MPFC’s broad association with self-processing in combination with the ventral to dorsal differentiation of processing about other people. 
DMPFC: Correcting Self-Projection or Certainty in (Social) Judgment? 
The correction of self-projection was one of the first psychological processes hypothesized to explain DMPFC’s association with both self-processing and processing of non-close-others. Psychological models suggest that one way we evaluate a new person is through the correction of self-projection, that is, by drawing on self-representation to the extent it is perceived as applicable to the new person (i.e., assumed similarity: Nickerson, 1999;Epley et al., 2004; Srivastava et al., 2010). Does DMPFC modulation reflect the corrective adjustment processes that are engaged to the extent that a new person is evaluated as dissimilar to the self? A different mechanism is suggested by an examination of the broader role of DMPFC in evaluation (including non-social evaluation) and psychological models of the interrelation between self-evaluation and evaluation of non-similar others. Specifically, DMPFC is associated with evaluation outside the social domain (Krain et al., 2006) and has been implicated in greater certainty about an evaluation (e.g., Krain et al., 2006; Bhanji et al., 2010). In contrast to the correction of self-projection hypothesis, it may be that certainty about self-evaluation explains why MPFC is modulated by the degree to which novel others are evaluated as different than the self. 
Dissimilarity between the self and a novel person positively modulates DMPFC activation 
Studies have consistently found that DMPFC (BA 9/32) activation parametrically increases to the extent that a novel person is evaluated as dissimilar to the self (Mitchell et al., 2005; Tamir and Mitchell, 2010). For example, these studies ask participants to report their own preferences (e.g., “how much do you look forward to going home for Thanksgiving?”) and to evaluate the preferences of strangers. The strangers are often manipulated to vary in their dissimilarity to the participant (e.g., have a different or similar political orientation, gender, or race). When neural activation is measured during the evaluation of strangers’ preferences, DMPFC (BA 9/32) is parametrically modulated by the dissimilarity between the participant’s own preferences and the preferences they assign to the strangers (Mitchell et al., 2005; Tamir and Mitchell, 2010). In other words, the more participants evaluate the strangers as dissimilar to themselves, the more DMPFC activation increases when participants are evaluating the stranger’s preferences. 
The robust association between DMPFC modulation and dissimilarity between self and others has been theorized to reflect the role of DMPFC (BA 9/32) in correcting, that is, adjusting one’s own self-representation to estimate the experience of a stranger. This explanation stems from psychological models of person evaluation which suggest that people use themselves as a starting point and correct as needed to evaluate unknown others (Nickerson, 1999; Epley et al., 2004; Srivastava et al., 2010). So if you encounter someone who shares your political orientation and you have to evaluate their position on a particular issue, you are likely to use your own experience to evaluate the person’s position. However, if a new person does not share your political orientation, then you cannot simply use your own experience and your evaluation will likely correct for the extent to which the person differs in political orientation (e.g., a liberal may feel that a self-representation might partially apply to a stranger who is a moderate but not apply at all to someone who is conservative). 
Does MPFC modulation reflect a correction of self-projection in social evaluation or is there another explanation that warrants examination before a strong conclusion can be drawn? There are a number of findings which raise the possibility that DMPFC modulation may instead reflect greater certainty in evaluation rather than correction of self-projection. For example, the previous studies have looked at DMPFC modulation during the evaluation of others. How does this compare to DMPFC modulation during self-evaluation and is it consistent with a correction of self-projection explanation? Meta-analyses find that DMPFC (DMPFC is a label that is used in various ways in previous literature; the present article draws on published meta-analyses and uses the term DMPFC as label for relevant portions of BA 9/32. Within BA 9, z ranges from 20 to 42 in MNI coordinates, see Figure 1) is activated by both self-processing and evaluations of other people not personally known by the participant. In direction comparisons, some meta-analyses find that this activation is relatively greater for unknown others while other meta-analyses find no difference in this region’s activation for self-processing and evaluation of unknown others (Self vs. Other comparison: Ochsner et al., 2005; Qin and Northoff, 2011; Roy et al., 2012). If the DMPFC reflects a correction of self-projection that occurs while evaluating another person, then it is puzzling why DMPFC is modulated by self-evaluation to the same degree as the evaluation of a person who is not personally known but assumed to be dissimilar to the self. If this region of DMPFC indexes correction away from a self-representation, why would this correction be engaged when evaluating oneself? 
An alternate conceptualization of why DMPFC is modulated by self-other dissimilarity: certainty about the evaluation 
Although it has not received much empirical attention, there is an alternate mechanism which could explain the pattern of DMPFC modulation found in these studies of self-evaluation and evaluation of strangers. Research on evaluation in non-social domains finds an association between DMPFC activation and greater certainty in evaluation (e.g., Krain et al., 2006;Bhanji et al., 2010; Eldaief et al., 2012). An integration of the psychological research on the interplay between self- and other-evaluation with the established association between DMPFC and evaluation certainty suggests that the DMPFC modulation found in paradigms involving self-evaluation and evaluation of strangers is tracking certainty in self-evaluation. 
It has already been shown that certainty about one’s self-evaluation modulates DMPFC activation (D’Argembeau et al., 2012). Studies on self-referent processing ask participants to rate the self-descriptiveness of personality traits and find that MPFC activation (extending into the DMPFC) is increased to the extent the traits are evaluated as self-descriptive (e.g., Moran et al., 2006; D’Argembeau et al., 2012). A personality trait may be evaluated as self-descriptive because people are certain about their association with the trait or they may be motivated to see themselves as characterized by that trait. One study delved further into these underlying reasons and found that a region within DMPFC was modulated by degree of certainty that the trait applied to self (D’Argembeau et al., 2012). 
DMPFC is associated with certainty about evaluation both for non-social tasks and self-evaluation; but how can an increase in certainty explain why DMPFC activation increases to the extent we evaluate people who are dissimilar to the self? Wouldn’t we be feeling uncertain when evaluating people we presume do not share our own qualities? Psychological research finds that evaluations of dissimilar others elicits a spontaneous self-evaluation and ironically solidifies our certainty about our own opinions and attitudes. In fact, certainty about our own preferences increases to the extent that the we perceive the target of our evaluation to be dissimilar to ourselves (Holtz and Miller, 2001; Holtz and Nihiser, 2008). In other words, this research suggests that rather than using the self as an anchor for evaluating other people (i.e., self-projection), the evaluation of other people triggers a spontaneous self-evaluation. And the more we evaluate someone to be different from us, the more we feel certain about where we stand on that attribute. Therefore, the increasing DMPFC activation found during a task that requires the evaluations of others could also be indexing an aspect of concomitant, spontaneous self-evaluations. Specifically, DMPFC activation may be modulated by increased certainty about the self to the extent that the target of evaluation is perceived as dissimilar to the self.\ 
Implications of an association between DMPFC and certainty about self-evaluation 
If DMPFC modulation does reflect certainty in self-evaluation, then a reconceptualization of self-evaluation localizer tasks may be warranted. Some studies have used a self-referent processing task (i.e., asking participants to rate their own personality traits compared to rating the personality traits of a political figure) as a way of localizing neural regions associated with self-processing for subsequent tasks. Future research is needed to understand whether this task identifies regions within DMPFC that index the intended rich psychological aspects of self or simply certainty in evaluation (i.e., on average, we are likely more certain about self-evaluation than evaluation of a political figure only seen in the news).
VMPFC: Socioemotional Connection or Firsthand Experience? 
The correction of self-projection or certainty might explain the contribution of DMPFC to social cognition but what about the more ventral subdivision of MPFC (VMPFC) that is associated with evaluations of self and intimate others (VMPFC is a label that is used in various ways in previous literature; the present article draws on published meta-analyses and uses the term VMPFC as label for relevant portions of BA 10/32; z range −2 to 8, see Figure 1, Ochsner et al., 2005; Qin and Northoff, 2011; Murray et al., 2012;Roy et al., 2012)? Two different hypotheses have been proposed: the correction of self-projection (Mitchell et al., 2005; Tamir and Mitchell, 2010) and self-relatedness (Northoff et al., 2006; Krienen et al., 2010;Murray et al., 2012; Roy et al., 2012). Currently, there is only mixed support for the correction of self-projection perspective. The hypothesis that VMPFC may mediate self-relatedness is more consistent with the available data but more research is needed to unpack the psychological meaning of self-relatedness. 
VMPFC modulation and the correction of self-projection? Current studies find inconsistent associations 
Unlike the DMPFC, VMPFC modulation has not shown a consistent pattern of association with evaluations of others as function of self-other dissimilarity (Mitchell et al., 2005; Krienen et al., 2010; Tamir and Mitchell, 2010). For example, an initial study asked participants to rate the preferences of unknown others (i.e., pleasure at having their photograph taken: Mitchell et al., 2005). VMPFC activation during the preference-evaluation task decreased to the extent that the unknown others were evaluated as dissimilar to the self in a post-scan procedure. Yet a follow-up analysis found a different pattern: VMPFC activation did not show a parametric association and it showed a positive (i.e., opposite) association to dissimilarity. VMPFC showed little change (in relation to baseline) during evaluations of other people who were evaluated (to any degree) as dissimilar from the self and a significant deactivation (in relation to baseline) when evaluating similar others (Tamir and Mitchell, 2011). It has been suggested that the different findings might indicate the existence of different neural mediation for computing global vs. specific dissimilarity. Dissimilarity was operationalized as a person’s global political affiliation on the one hand (Mitchell et al., 2006) and trial-by-trial specific preferences on the other (i.e., Tamir and Mitchell, 2010). However, another series of studies found no association at all between VMPFC (BA 10) modulation and similarity between close others or strangers (Krienen et al., 2010). This research instead found that VMPFC shows increased activation for self-evaluations and evaluations of close others (regardless of similarity) and less activation for unknown others. Even if future research were to flesh out a robust association between VMPFC modulation and evaluations of dissimilar others, the correction of self-projection explanation still suffers from a parallel set of problems mentioned above in relation to DMPFC. Meta-analyses find that VMPFC activation is modulated by both self-evaluation and evaluation of intimate others (Ochsner et al., 2005; Qin and Northoff, 2011; Murray et al., 2012). It is unclear why people would need to correct the use of their self-representation when evaluating themselves or why they would use a self-projection process to evaluate someone they know well. 
VMPFC modulation and self-relatedness of social evaluation: A socioemotional or cognitive mechanism? 
Another predominant hypothesis arising from current social evaluation research is that VMPFC marks “self-relatedness,” that is, the socioemotional connection between the self and the person being evaluated (Northoff et al., 2006; Krienen et al., 2010; Murray et al., 2012; Roy et al., 2012). Self-relatedness is a socioemotional variable reflecting the extent to which the evaluation process draws on affectively rich, self-representations. Psychological models of social evaluation suggest that self-representations may be activated by evaluations of close others but not for the same reason as for unknown others. “Close others” are often defined by the extent to which representations of those people are associated with self-representations (Aron et al., 1992). It is not the case that the self-representation is theorized to serve as a starting point for evaluating the close other (i.e., a self-projection-like process which is then subject to correction). Instead, the evaluation of a close other draws on a representation of the close other that is emotionally charged because of its association with the self-representation. From this perspective, VMPFC is modulated by self-evaluation and evaluation of close others because those evaluations have a unique affective or socioemotional significance. 
However, it may not be that VMPFC marks whether social evaluations are “self-like” in a socioemotional sense. In the existing research, socioemotional relation between the self and another person has always been confounded with the quality of information (e.g., cognitive representation) used to make an evaluation. We simply have a different class of information to draw on when we evaluate ourselves and people we actually know (e.g., greater complexity, abstraction, actual experience) compared to unknown others. A novel person and a romantic partner elicit different emotional reactions but they also elicit different cognitive representations. For both the self and romantic partners, there is a long history of storing person information which creates a more elaborated representation that includes both abstract and biographical information when compared to representations that could be used to evaluate someone who is relatively unknown (e.g., Sherman and Klein, 1994; Kihlstrom et al., 2003). A brain region that indexes one or more cognitive qualities that are emphasized in the representations of people we know well (i.e., self, close other) would also behave like the VMPFC across these social evaluation tasks as reviewed above (i.e., similar modulation across self-evaluation and evaluation of close others but less modulation for unknown others: Ochsner et al., 2005; Krienen et al., 2010; Qin and Northoff, 2011; Murray et al., 2012; Roy et al., 2012). This raises the possibility that the contribution of VMPFC to social cognition is more a cognitive (rather than affective) “self-relatedness.” From this perspective, VMPFC may mediate a quality of the kind of information that feeds into self-evaluations that is also available for evaluations of people we actually know (but not as much for unknown others). 
Expanding Beyond MPFC (BA 9/10/32): VACC (BA 25) and MOFC (BA 11) Mediate Motivational Influences on Bottom-Up and Top-Down Processing of Social Targets 
Despite the heavy focus on MPFC (BA 9/10/32), an emerging body of literature suggests that at least two other midline regions are just as important for social cognitive processing: VACC (BA 25) and MOFC (BA 11) (see Figure 1). VACC and MOFC mediate motivational aspects of self-processing. Motivation has been ascribed a central role in psychological models of self-processing (Kunda, 1990; Robins and John, 1997). For example, self-evaluations tend to be positively tinged (also described as “self-serving,” “the above average effect,” “self-flattering,” “self-enhanced” “optimistic bias”: Alicke, 1985; Taylor and Brown, 1988; Dunning et al., 1989; Chambers and Windschitl, 2004). Self-evaluations are described as positively tinged to the extent that they are more positive than warranted by some other criterion and this positive slant may even be pre-potent, that is, the default mode of self-evaluation (Beer, 2007). Cognitive load makes self-evaluation even more positively tinged (Paulhus et al., 1989;Kruger, 1999; Koole and Dijksterhuis, 2001; Lench and Ditto, 2008; Beer and Hughes, 2010; Beer et al., 2013). Furthermore, the positive tinge of self-evaluation is not circumscribed to the lab (Dunning et al., 2004). People will wager money that their positively tinged views are accurate (Williams and Gilovich, 2008), expect that other people will share their positively tinged views (Hepper et al., 2011), and experience different life trajectories based on the extent of their positive slant (Robins and Beer, 2001). A positive tinge also pervades evaluations of close others but is less evident in evaluations of unknown others (Suls et al., 2002). A positive tinge may arise because people use incomplete information when making a social evaluation (e.g., using the first thing that comes to mind which happens to be positive: Chambers and Windschitl, 2004). However, a positive tinge can also arise from the motivation to cast oneself or a close other in a positive light (i.e., self-flattery: Taylor and Brown, 1988;Sedikides and Gregg, 2008). 
Despite the central role of motivation in psychological models of self- and person evaluation, neural models of self-processing have paid little attention to motivation (Beer, 2007, 2012a). Recent research that addresses this gap suggests that (a) VACC may be modulated by opportunities that have the potential to accomplish a motivated self-evaluation (i.e., motivational influences on bottom-up processing) and (b) MOFC may be modulated by the extent to which the motivation to cast oneself in a positive light requires the adjustment of evaluation thresholds across contexts (i.e., top-down processing). 
VACC: Motivational Influences on Bottom-Up Processing 
VACC may mediate bottom-up sensitivity to opportunities that have the potential to affirm the way someone wants to evaluate themselves; however, it does not predict whether the opportunity will successfully lead to motivated self-evaluation (Moran et al., 2006; Sharot et al., 2007; Beer and Hughes, 2010; Hughes and Beer, 2012a). Social psychological theories of self- and other-evaluation often characterize these evaluations in terms of the contribution of bottom-up and top-down processes (e.g., Duncan, 1976; Shavelson and Bolus, 1982; Devine, 1989; Fiske and Neuberg, 1990;Brown et al., 2001; Beer, 2012b). “Top-down” and “bottom-up” are terms that are used widely, but somewhat differently across fields. In the case of motivated self-evaluation, these terms can be used to distinguish between subjective and objective construals of information. People may be motivated to see themselves in a particular way and, therefore, interpret information in a top-down, subjective manner that ensures the information can be used to accomplish a motivated self-evaluation. Or the motivation may affect the kind of information that is distinguished from other kinds of information (i.e., the influence of the motivation on relatively bottom-up processing: e.g., Brown et al., 2001). 
The influence of motivation on relatively bottom-up processing of information can be illustrated by the example of people filling out an online dating profile who want to portray themselves as especially athletic compared to other people. If people scan the activities checklist with the goal of portraying themselves as especially athletic, then we predict that VACC will be modulated by activities on the checklist that objectively involve sports vs. activities that reflect poorly on athleticism (e.g., watching television). Similarly, someone with the goal of portraying themselves as artistic would show greater VACC activation when reading checklist activities that objectively involve artistic pursuits vs. all of the other options. In this way, VACC activation is implicated in the influence of motivation on bottom-up processing of the checklist because VACC modulation distinguishes between opportunities that are objectively consistent vs. inconsistent with the activated motivation. VACC is not implicated in purely top-down processing because research suggests that it would not predict the extent to which someone claims to be especially involved in each sport compared to other people (i.e., the success of the top-down goal of portraying oneself as particularly athletic). In other words, VACC modulation does not predict the extent to which the meaning or interpretation of the checklist activities are subjectively construed to fit with the activated motivation nor does it predict reported self-evaluation on those checklist activities. Instead, we hypothesize that VACC is modulated by a preliminary and relatively more bottom-up step of motivated evaluation: delineating the existence of opportunities that objectively have the potential to cast yourself in particular light. 
VACC activation differentiates positive valence from negative valence, especially for social targets we want to see in a positive light 
The distinction between opportunity for motivated evaluation and success in motivated evaluation is important because they have been conflated in the current literature (Beer, 2007, 2012a). It is inappropriate to use the term “bias” (e.g., positively tinged) to label a self-endorsement of a positive trait or likelihood of a positive future event. There is no way to know whether someone has successfully achieved a positively tinged evaluation simply because they are rating positive traits or future events as particularly self-descriptive. The person may truly possess high levels of that trait and be predisposed to a positive future or they may not. A response can be characterized as “biased” or positively tinged (rather than merely positive) when it is more positive than warranted by a benchmark criterion (Beer, 2007, 2012b). 
For example, one way that positively tinged self-evaluation has been operationalized is the extent to which people inflate their own standing when comparing themselves to other people (Taylor and Brown, 1988;Chambers and Windschitl, 2004). This line of research often asks participants to make social-comparative judgments. That is, participants are asked to evaluate how much they possess personality traits in comparison to their average peer (i.e., much less, about the same or much more than someone of the same, age, community, education level, etc.). When participants’ social-comparison evaluations are averaged across hundreds of personality traits, their average evaluation, by definition, should be somewhere near the average peer benchmark. However, the majority of people report having significantly higher levels of positive personality traits and significantly lower levels of negative traits than their average peer (Taylor and Brown, 1988; Chambers and Windschitl, 2004). In this social-comparison task, VACC is modulated by the condition that includes positive personality traits (compared to negative personality traits) but it does not predict the extent to which someone reports an overall significantly more desirable personality in comparison to their average peer (Beer and Hughes, 2010; Hughes and Beer, 2012a). 
VACC has been implicated in the detection of emotionally significant, that is, valenced information in a variety of tasks (compared to non-valenced information: Bush et al., 2000). However, research on social cognition has shown that VACC modulation may differentiate between particular classes of valence depending on motivational state. When people evaluate well-liked social targets (e.g., the self, romantic partner, well-liked roommate), VACC activation differentiates trials where endorsement would portray the target in a positive light (i.e., desirable personality traits, likelihood of a positive future) from trials where endorsement would portray the target in a negative light (i.e., undesirable personality traits, likelihood of a negative future: Moran et al., 2006; Sharot et al., 2007; Beer and Hughes, 2010;Hughes and Beer, 2012a). However, when there is reduced motivation to portray the target in a positive light (i.e., personality traits that are not considered central to one’s self-view: Sedikides and Gregg, 2008; a non-close other: Suls et al., 2002), VACC activation is less likely to differentiate trials on the basis of how endorsement would portray the target (i.e., the self: Moran et al., 2006; an assigned college roommate: Hughes and Beer, 2012a). This research suggests that VACC is important for identifying opportunities to portray someone in a particular light but it does not predict whether the opportunity actually leads to successful motivated evaluation. 
Bottom-up sensitivity to information based on its potential to affirm motivated self-evaluations: connectivity between VACC and MOFC 
Psychophysiological interaction connectivity analyses (Friston et al., 1997) conducted on previously published results (Beer and Hughes, 2010) further supports the hypothesis that VACC (BA 25) mediates a preliminary step but not the ultimate success of motivated evaluation (for the full set of results, see Figure 2; Table 1).
FIGURE 2 
Figure 2. PPI connectivity analyses for the VACC seed associated with social comparisons about Positive (i.e., desirable) vs. Negative (i.e., undesirable) personality traits. (A) Each participant’s time series was extracted from the VACC seed (5 mm radius sphere around group peak: 14, 38, −4 from the Positive vs. Negative contrast, Beer and Hughes, 2010). (B) The VACC seed shows relatively more positive covariation with an MOFC region. This MOFC region overlaps with the MOFC region that regulates the extent to which social comparisons are positively tinged (red: MOFC region found in PPI analyses; blue: MOFC region found in Beer and Hughes, 2010; purple: overlap between MOFC region in connectivity and primary analyses).
TABLE 1 
 
Table 1. PPI connectivity analyses with VACC seed from data published in Beer and Hughes (2010).
Methods. Whole-brain PPI analyses were conducted in order to investigate the functional connectivity of the VACC region that differentiated social-comparative evaluations made in the Positive condition from the social-comparative evaluations made in the Negative condition (Beer and Hughes, 2010). Specifically, participants rated how much they had desirable (Positive condition) and undesirable (Negative condition) personality traits in comparison to their average peer. Imaging data were preprocessed using the FSL software toolbox [Oxford Center for Functional Magnetic Resonance Imaging (FMRIB); Smith et al., 2004]. Functional images were motion corrected using MCFLRT (Jenkinson et al., 2002) and non-brain structures were stripped from functional and structural volumes using the Brain Extraction Tool (BET; Smith, 2002). Images were then smoothed (8 mm full-width half-maximum) and normalized to MNI-152 space during preprocessing. Parameters for normalization into a standard space were obtained by multiplying the transformation matrices across a two-step process in which the functional images were registered to the MP-RAGE (6 DOF affine transformation), and the MP-RAGE was registered to the MNI-152 template (12 DOF affine transformation). 
Functional Magnetic Resonance Imaging analysis was performed using FSL’s FEAT (FMRI Expert Analysis Tool version 5.98). A fixed-effects analysis modeled event related responses for each participant. Responses made in the Positive and Negative conditions were modeled as events using a canonical hemodynamic response function with a temporal derivative. Motion regressors were modeled as regressors of no interest. Each participant’s time series was extracted from the VACC seed found in the group analyses of the Positive vs. Negative condition (5 mm radius sphere around group peak: 14, 38, −4 from the Positive vs. Negative contrast, Beer and Hughes, 2010). Two PPI regressors were created: the interaction of the time series of the VACC seed with (a) the time series of the Positive condition regressor and (b) the time series of the Negative condition regressor. 
A subsequent fixed-effects analysis was conducted modeling the following regressors: (a) Positive condition regressor, (b) Negative condition regressor, (c) temporally filtered activity across the time course from the VACC seed region, (d) PPI regressor for the Positive condition, and (e) PPI regressor for the negative condition. The PPI regressors were contrasted in a GLM. A second-level analysis created contrast estimates for each participant by collapsing across the two runs, treating runs as a fixed effect. FEAT’s FLAME module (FMRIB’s Local Analysis of Mixed Effects; Smith et al., 2004) was used to preformed mixed effects analysis which created group average maps for contrasts of interest (p < 0.005, uncorrected). The significance threshold was chosen because it is the recommended threshold for striking an optimal balance between Type I and Type II error when reporting analyses of brain activation in relation to complex psychological processes; simulation studies show that other significance thresholds raise the possibility of Type II error beyond acceptable limits (Lieberman and Cunningham, 2009). As the first report of functional connectivity in relation to motivated self-evaluation, the goal was to be as inclusive as reasonably possible to avoid missing true effects.
Results. When people make social comparisons about desirable traits, VACC shows relatively more positive covariation with a portion of MOFC (BA 11) that was found to regulate the extent to which social-comparative evaluations are positively tinged in the primary analyses. Although directionality cannot be determined from PPI analyses, it is possible that VACC is involved in analyzing the opportunities afforded by the content of an evaluation (i.e., a desirable trait vs. an undesirable trait). That information may then be processed upstream by the MOFC before an evaluation is expressed. 
MOFC: Motivational Influences on Top-Down Processing 
As mentioned above, the MOFC is implicated in self-evaluation. How should we conceptualize its role? Take the example mentioned earlier: people who view themselves as particularly athletic complete an activity checklist on an online dating profile. Their expectation may be met when they are able to endorse participation in numerous sports on the checklist. But if they find themselves able to only endorse involvement in just one or two of the numerous sports possibilities, they may have one of two possible reactions. If their self-esteem is not staked on their athletic ability, they might realize that they are not so different from other people in this regard. However, if the procedure threatens their self-esteem, they may react defensively by changing their evaluation threshold in such a way that they can evaluate themselves as having even more superior athleticism. For example, they may evaluate degree of athleticism based on the intensity of involvement in a particular sport, rather than on the number of sports activities they can endorse on the checklist. In one case, the initial expectation of portraying oneself as athletic is dismissed during activity endorsement (i.e., an initial top-down influence is controlled). In the case where self-esteem is threatened, motivation to portray oneself in a particular way exhibits a top-down influence on activity endorsement by biasing the standards with which the evaluation is made. MOFC modulation has been associated with both of the examples above: realizing the self is not as special as expected and defensive reactions when self-esteem is threatened. Connectivity analyses suggest that MOFC modulation likely reflects different psychological processes across these circumstances. In particular, a network involving MOFC and (a) relatively more positive covariation with striatum and (b) relatively less positive covariation with middle frontal gyrus may aid self-evaluations that protect the self in the face of self-esteem threat. However, MOFC activation found in association with dismissing the influence of a self-evaluation motivation does not show such connectivity. In this way, MOFC may mediate top-down motivational influences on social evaluation by supporting changes in evaluation standards to either facilitate or control an activated motivational state. 
When self-esteem is not at stake: OFC function is negatively associated with positively tinged social evaluations 
Both neuroimaging and lesion studies have shown that reduced MOFC function is associated with positively tinged social evaluations (i.e., self and close others: Beer et al., 2003; Beer et al., 2006a, 2010; Beer and Hughes, 2010; Hughes and Beer, 2012a). This relation holds across various operationalizations of self-flattery: the difference in the way you see yourself compared to how others view you, self-evaluation of task achievement compared to actual task performance on an unimportant task, and base rates of social comparisons. 
A series of studies found that patients with OFC damage tend to view their social behavior in a positively tinged manner (Beer et al., 2003, 2006a). In one study, patients with OFC damage were socially disinhibited compared to healthy control participants yet they expressed greater pride in their social behavior (i.e., inappropriate teasing of strangers: Beer et al., 2003). Another study found that patients with OFC damage did not evaluate the appropriateness of their social behavior any differently than healthy control participants or participants with dorsolateral prefrontal cortex (DLPFC) damage. Yet outside observers, blind to participant status, rated the social behavior of the patients with OFC damage to be significantly more inappropriate than the other groups (i.e., too familiar for an interaction with a stranger: Beer et al., 2006a). 
Neuroimaging results complement the lesion studies: reduced OFC activation (BA 11) is associated with positively tinged evaluations of one’s task performance and personality (Beer and Hughes, 2010; Beer et al., 2010; Hughes and Beer, 2012a). In one study, participants estimated their confidence in their answers to a trivia task. Reduced OFC activation (BA 11) predicted the extent to which participants were overconfident about their incorrect trivia answers (Beer et al., 2010). Reduced OFC activation also predicts the extent to which people view themselves and their romantic partners to have significantly more desirable personalities than their peers. As mentioned above in the section on VACC function, these studies ask participants to compare themselves or their romantic partners to an average peer (i.e., a person who is the same gender, age, from the same community, university campus, etc.). When these social-comparative judgments of personality traits are averaged across hundreds of traits, each participant (or their romantic partner) should, by definition, be evaluated as comparable to their average peer. Whereas VACC activation showed no relation, reduced OFC activation is associated with the extent to which people evaluate themselves (Beer and Hughes, 2010) or their romantic partners (Hughes and Beer, 2012a) to have significantly more positive traits and significantly fewer negative traits than their average peer. Taken together, these studies provide robust evidence that reduced OFC activation predicts positively tinged evaluations on a trial-by-trial, condition, and individual difference basis. 
The case of self-esteem defense: A positive association between OFC activation and self-protection 
There is an exception to the findings described above: increased MOFC (BA 11) activation predicts self-evaluations in situations where self-esteem comes under attack (Hughes and Beer, 2013). Self-esteem is typically threatened when people receive negative feedback about their personality, academic abilities, or skills (Baumeister et al., 1993; Leary et al., 1998;vanDellen et al., 2011). People cope with self-esteem threat by inflating the positively tinged nature of their self-evaluation (including social comparisons: Beer et al., 2013 and see vanDellen et al., 2011 for review). The lesion and fMRI research reviewed above did not include any manipulations to threaten self-esteem. What happens to the underlying neural modulation when social-comparison judgments are used to cope with self-esteem attack? In other words, what neural regions mediate self-evaluations that are self-flattering (e.g., positively tinged with the purpose of protecting the self against a self-esteem threat)? One fMRI study addressed this question by using the very same social-comparison evaluation as a previous study (Beer and Hughes, 2010) but added in a self-esteem threat manipulation (Hughes and Beer, 2013). Participants learned that other students had found them either likable or unlikable and then evaluated how their personalities compared to their peers. Consistent with previous research, evaluations made after learning that others found them unlikable were even more self-flattering (compared to learning that others found them likable). The extent to which social comparisons became even more self-flattering as a function of self-esteem attack was positively associated with increased MOFC modulation (Hughes and Beer, 2013). Therefore, this study found that MOFC modulation predicted a change in self-evaluation but, in the case of self-esteem attack, it shows a positive association with self-protection. 
Although the studies on social comparison (Beer and Hughes, 2010; Hughes and Beer, 2013) provided a rigorous test of the association between MOFC modulation and self-evaluation as a function of self-esteem threat, they were not designed to pinpoint the underlying psychological process that explained the association. One study has begun to address this issue by using Signal Detection Theory to investigate the neural associations of self-evaluations used to protect one’s self-esteem (Hughes and Beer, 2012b). Just as people tend to inflate their social standing on personality traits, they tend to claim knowledge about concepts beyond what they actually know or could know (i.e., overclaim knowledge: Paulhus et al., 2003; Beer et al., 2010). However, when self-esteem is potentially at stake (i.e., their false claims could be discovered), people reduce the extent to which they overclaim knowledge (Paulhus et al., 2003) or inflate their social standing on personality traits (McKenna and Myers, 1997). In conditions where false claims would make them look foolish, people protect their self-esteem by adopting a different standard (i.e., decision threshold) for claiming knowledge which consequently reduces overclaiming. An fMRI study found that MOFC (BA 11) modulation was positively associated with the shift toward a more conservative standard in conditions where participants would look foolish if they were to make false claims of knowledge (i.e., they were warned that some concepts in the list did not exist: Hughes and Beer, 2012b). 
A top-down role of MOFC in social evaluation 
Consistent with the hypothesis that self-evaluations used to cope with self-esteem threat are distinct from self-evaluations made in the absence of threat, a relatively consistent pattern of functional connectivity emerged in the studies that investigated the impact of self-esteem threat on self-evaluations (Hughes and Beer, 2012b, 2013) and was distinct from the pattern found in a parallel social-comparison procedure that did not manipulate self-esteem threat (Beer and Hughes, 2010, see Table 2).
TABLE 2 

Table 2. PPI connectivity analyses with MOFC seed*.
 
Methods. Whole-brain PPI analyses were conducted in order to investigate the functional connectivity of MOFC during social comparisons in the presence and absence of self-esteem threat from three previously published datasets (Dataset 1: Hughes and Beer, 2013; Dataset 2: Hughes and Beer, 2012a; Dataset 3: Beer and Hughes, 2010). For all three datasets, the preprocessing steps were the same as described earlier for the PPI analyses of the VACC seed. PPI analyses were conducted as follows. In Dataset 1 (Hughes and Beer, 2013), participants made social-comparative evaluations of their personality traits while the presence of self-esteem threat was manipulated. In other words, participants evaluated how their personality traits compared to an average peer after just learning that a majority of peers found them unlikable (Threat condition) or a majority of peers found them likable (No Threat condition). As previously published, increased MOFC activity is associated with positively tinged evaluations of one’s personality in the Self-esteem Threat condition (both a main effect and individual differences in evaluations made in the Threat vs. No Threat condition: Hughes and Beer, 2013). Each participant’s time series was extracted from the MOFC seed (group peak: −12, 54, −14 from the Threat vs. No Threat contrast, Hughes and Beer, 2012a). Two PPI regressors were created: interaction of the time series of the MOFC seed with (i) the time series of the Threat condition regressor and (ii) the time series of the No Threat condition regressor. 
In Dataset 2 (Hughes and Beer, 2012b), participants evaluated their familiarity with blocks of information they believed would make them appear intelligent while their awareness of the exposure of fake claims was manipulated. Specifically, all blocks of information contained items that existed and items that do not exist but participants were only warned of the possibility of non-existent items in half of the blocks (Accountable condition vs. an Unaccountable condition where they were not warned that they might be claiming familiarity with something that does not exist). Increased MOFC activity was associated with the shift toward a more conservative standard for claiming knowledge in the Accountable condition. Each participant’s time series was extracted from the MOFC seed (5 mm radius sphere around group peak: −6, 58, −20 from the Accountable vs. Not Accountable contrast; Hughes and Beer, 2012b). Two PPI regressors were created: interaction of the time series of the MOFC seed with (i) the time series of the Accountable condition regressor and (ii) the time series of the Not Accountable condition regressor. 
In Dataset 3 (Beer and Hughes, 2010), participants made the same social-comparative evaluations of their personality traits as in Dataset 1 but self-esteem threat was not manipulated. Instead, the breadth of personality traits were manipulated such that they could either be broadly construed (i.e., Broad condition: trait has a wide variety of behavioral manifestations such as “capable”) or more specifically construed (i.e., Specific condition: trait has few behavioral manifestations such as “talkative”). Reduced MOFC activity was associated with viewing the self as having more positive and fewer negative traits in comparison to the average peer (i.e., positively tinged evaluations of one’s personality). Each participant’s time series was extracted from the MOFC seed (5 mm radius sphere around group peak: −4, 46, −10 from the Specific vs. Broad contrast; Beer and Hughes, 2010). Two PPI regressors were created: interaction of the time series of the MOFC seed with (i) the time series of the Specific condition regressor and (ii) the time series of the Broad condition regressor. 
After PPI regressors were created, all of the datasets were subjected to a subsequent fixed-effects analyses in the same manner as described earlier for the PPI analyses of the VACC seed. Specifically, the fixed-effects analyses to modeled condition of interest regressors (i.e., Dataset 1: Threat and No Threat conditions (Hughes and Beer, 2013; Dataset 2: Accountable and Unaccountable conditions (Hughes and Beer, 2012b; Dataset 3: Specific and Broad conditions (Beer and Hughes, 2010), a temporal filter of activity across the time course from the MOFC seed region, and the PPI regressors for conditions of interest. The PPI regressors were contrasted in a GLM. FEAT’s FLAME module (FMRIB’s Local Analysis of Mixed Effects; Smith et al., 2004) was used to preform mixed effects analyses for each dataset, which created group average maps for contrasts of interest (p < 0.005, uncorrected). 
Results. PPI connectivity analyses (Friston et al., 1997) conducted on previously published results (Beer and Hughes, 2010; Hughes and Beer, 2012b, 2013) suggest that functional connectivity between MOFC, the striatum, and the middle frontal gyrus (BA 9) may support self-evaluations used to protect self-esteem in the face of threat (see Figure 3; Table 2). When self-esteem is at stake, the region of MOFC that is associated with self-evaluation shows relatively less positive covariation with middle frontal gyrus (BA 9) and relatively greater positive covariation with striatum. It is possible that functional connectivity between MOFC and striatal subregions reflects whether a shift to more conservative or liberal evaluation standards will be most rewarding in the face of self-esteem threat. For example, greater positive covariation between MOFC and caudate was found when liberal thresholds were advantageous for protecting self-esteem (Hughes and Beer, 2013) and between MOFC and putamen when conservative thresholds were advantageous for protecting self-esteem (Hughes and Beer, 2012b). Taken together, this research suggests that MOFC aids top-down influences on social cognition by adjusting evaluation standards as function of motivational state.
FIGURE 3 
Figure 3. PPI connectivity analyses for the MOFC seed associated with self-protection in the face of self-esteem threat. (A) MOFC seed regions for connectivity analyses of previously published studies (5 mm radius spheres around group peak). Dark green seed: social comparisons while under Threat vs. No Threat contrast, group peak −12, 54, −14 (Dataset 1: Hughes and Beer, 2013). Light green seed: claims of knowledge while Accountable vs. Not Accountable contrast, group peak: −6, 58, −20 (Dataset 2: Hughes and Beer, 2012b). (B) When false claims of knowledge could be discovered, the MOFC seed associated with self-protection (i.e., less inflated claims) shows relatively more positive covariation with the right putamen (22, −10, 6) and less positive covariation with the left middle frontal gyrus (BA 9; −24, 52, 30). (C)When self-esteem was threatened, the MOFC seed associated with more self-flattering evaluations shows relatively more positive covariation with the right caudate (14, 16, −10) and less positive covariation with the middle frontal gyrus (BA 9). 
Conclusions and Future Considerations for Research on the Role of MPFC, VACC, and MOFC in Self-Evaluation 
While much progress has been made since the discovery in the early 2000s that MPFC is associated with self-evaluation, several hypotheses have been tested and eliminated. The new hypotheses described here will benefit from future research guided by a number of questions. For example, even though MPFC has received the bulk of attention, there are still many questions that remain. It would be extremely useful (and feasible) to conduct connectivity analyses on the large, existing body of studies that have measured MPFC modulation in relation to both self-evaluation and the evaluation of unknown others. One potential drawback of the “correction of self-projection” hypothesis for both VMPFC and DMPFC is that these regions are activated for evaluation of targets where correction of self-projection is unlikely (e.g., self and/or close others). If the functional connectivity of VMPFC and DMPFC is different during self-evaluation compared to evaluations of unknown others, those results would eliminate some concerns about the correction of self-projection hypothesis. Furthermore, more research is needed to decouple the affective vs. cognitive qualities shared by evaluation of the self and close others to more clearly delineate the role of VMPFC in social evaluation. If VMPFC is similarly modulated by the evaluation of another person where there is an emotional association with the self but no actual firsthand experience or basis for self-projection, then that would be strong evidence that VMPFC indexes the emotional aspect of self-relatedness when evaluating other people. 
Additionally, more research is needed to clarify the possibility that VACC is involved in detecting opportunities that might fulfill expectations about self-evaluation. Does VACC mediate sensitivity to motivationally consistent information or positively valenced information when it is motivationally consistent? This question is important because psychological models show that motivation to see oneself in a positive light is not the only motivation that impacts self-evaluation. For example, the relation between valence and motivation becomes uncoupled when self-verification, another motivation known to influence self-evaluation, is activated. People often want to feel that their self-evaluations are correct and are vigilant for opportunities that have the potential to verify their current self-evaluations. In fact, this research finds that people with negative self-evaluations desire chances to confirm these negative self-evaluations (Swann et al., 1989). In this situation, the evaluation of negative traits (rather than positive traits) would have the potential to affirm motivated self-evaluation. If VACC mediates sensitivity to motivationally consistent information, then it should be modulated by opportunities to affirm a negative self-evaluation for people who are motivated to confirm a negative self-view. Furthermore, more research is needed to replicate and understand the psychological significance of the connectivity between VACC, MOFC and the other regions found in the PPI analyses. 
Finally, more research is needed to replicate and elucidate the functional connectivity of MOFC in association with self-evaluations made in the presence and in the absence of self-esteem threat. While there is convergent evidence that more positive covariation with striatum and reduced covariation with middle frontal gyrus is associated with self-evaluation used to protect self-esteem, much less is understood about the significance of regions that covary with MOFC modulation associated with self-evaluations made in the absence of self-esteem threat. 
In conclusion, a new framework is proposed to account for the contribution of MPFC, VACC, and MOFC to social cognition. MPFC is broadly implicated in self-evaluation but may be characterized by a ventral to dorsal division when evaluating others based on their intimacy. Certainty about evaluation may better characterize the contribution of DMPFC to social cognition than the correction of self-projection. The association between VMPFC and self-relatedness will be clearer once future research disentangles shared emotional and cognitive properties of evaluation of self and close others. Further, previous research has failed to take into account the fundamental role that motivation has in self-evaluations. As a result, the role of VACC and MOFC in social cognition has been obscured until recently. VACC may mediate bottom-up sensitivity to information based on its potential for helping us evaluate ourselves and others the way we want. MOFC may mediate top-down motivational influences on self-evaluation. 
Conflict of Interest Statement 
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. 
Acknowledgments
Funding for the article was supported by NSF Grant (NSF-BCS-1147776) to Jennifer S. Beer.
References are available at the Frontiers site.