The Paradigmatic Body - — Open MIND

41
The Paradigmatic Body Embodied Simulation, Intersubjectivity, the Bodily Self, and Language Vittorio Gallese & Valentina Cuccio In this paper we propose a way in which cognitive neuroscience could provide new insights on three aspects of social cognition: intersubjectivity, the human self, and language. We emphasize the crucial role of the body, conceived as the con- stitutive source of pre-reflective consciousness of the self and of the other. We provide a critical view of contemporary social cognitive neuroscience, arguing that the brain level of description is a necessary but not sufficient condition for study- ing intersubjectivity, the human self, and language; which are only properly vis- ible if coupled with a full appreciation of their intertwined relationship with the body. We introduce mirror mechanisms and embodied simulation and discuss their relevance to a new account of intersubjectivity and the human self. In this context, we focus on a specifically human modality of intersubjectivity: language. Aspects of social cognition related to language are discussed in terms of embodiment, while emphasizing the progress and limitations of this approach. We argue that a key aspect of human language consists in its decoupling from its usual denotative role, hence manifesting its power of abstraction. We discuss these features of hu- man language as instantiations of the Greek notion of paradeigma, originally ex- plored by Aristotle to refer to a typical form of rhetorical reasoning and relate it to embodied simulation. Paradigmatic knowledge connects the particular with the particular, moving from the contingent particular situation to an exemplary case. Similarly, embodied simulation is the suspension of the “concrete” application of a process: reuse of motor knowledge in the absence of the movement it realizes is an example of “paradigmatic knowledge.” This new epistemological approach to intersubjectivity generates predictions about the intrinsic functional nature of our social cognitive operations, cutting across, and not subordinated to, a specific on- tology of mind. Keywords Cognitive neuroscience | Embodied simulation | Intersubjectivity | Language | Mirror neurons | Paradigm | Social cognition Authors Vittorio Gallese vittorio.gallese @ unipr.it Università degli Studi di Parma Parma, Italy Valentina Cuccio valentina.cuccio @ unipa.it Università degli Studi di Palermo Palermo, Italy Commentator Christian Pfeiffer christian.pfeiffer @ epfl.ch Ecole Polytechnique Fédérale Lausanne, Switzerland Editors Thomas Metzinger metzinger @ uni-mainz.de Johannes Gutenberg-Universität Mainz, Germany Jennifer M. Windt jennifer.windt @ monash.edu Monash University Melbourne, Australia 1 Introduction The last decades of the twentieth century were marked by great progress in cognitive neuros- cience, made possible by recently-developed brain imaging technologies such as functional magnetic resonance imaging (fMRI)—which al- lowed for the first time non-invasive study of the human brain. But what is cognitive neuroscience? We think it is fair to say that it is above all a meth- odological approach whose results are strongly influenced by which questions are being asked and how. Studying single neurons and/or the brain does not necessarily predetermine the questions to be asked that will help us under- stand how and how much our human nature de- pends upon our brains. Even less so the an- swers. Our purpose here is twofold. On the one hand, we aim to provide a brief overview of cur- rent cognitive neuroscience and its methods. We first present the limitations displayed by most Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 1 | 22

Transcript of The Paradigmatic Body - — Open MIND

The Paradigmatic BodyEmbodied Simulation, Intersubjectivity, the Bodily Self, and Language

Vittorio Gallese & Valentina Cuccio

In this paper we propose a way in which cognitive neuroscience could providenew insights on three aspects of social cognition: intersubjectivity, the human self,and language. We emphasize the crucial role of the body, conceived as the con-stitutive source of pre-reflective consciousness of the self and of the other. Weprovide a critical view of contemporary social cognitive neuroscience, arguing thatthe brain level of description is a necessary but not sufficient condition for study-ing intersubjectivity, the human self, and language; which are only properly vis-ible if coupled with a full appreciation of their intertwined relationship with thebody. We introduce mirror mechanisms and embodied simulation and discuss theirrelevance to a new account of intersubjectivity and the human self. In this context,we focus on a specifically human modality of intersubjectivity: language. Aspectsof social cognition related to language are discussed in terms of embodiment,while emphasizing the progress and limitations of this approach. We argue that akey aspect of human language consists in its decoupling from its usual denotativerole, hence manifesting its power of abstraction. We discuss these features of hu-man language as instantiations of the Greek notion of paradeigma, originally ex-plored by Aristotle to refer to a typical form of rhetorical reasoning and relate itto embodied simulation. Paradigmatic knowledge connects the particular with theparticular, moving from the contingent particular situation to an exemplary case.Similarly, embodied simulation is the suspension of the “concrete” application of aprocess: reuse of motor knowledge in the absence of the movement it realizes isan example of “paradigmatic knowledge.” This new epistemological approach tointersubjectivity generates predictions about the intrinsic functional nature of oursocial cognitive operations, cutting across, and not subordinated to, a specific on-tology of mind.

KeywordsCognitive neuroscience | Embodied simulation | Intersubjectivity | Language |Mirror neurons | Paradigm | Social cognition

Authors

Vittorio [email protected]   Università degli Studi di ParmaParma, Italy

Valentina [email protected]   Università degli Studi di PalermoPalermo, Italy

Commentator

Christian [email protected]   Ecole Polytechnique FédéraleLausanne, Switzerland

Editors

Thomas [email protected]   Johannes Gutenberg-UniversitätMainz, Germany

Jennifer M. [email protected]   Monash UniversityMelbourne, Australia

1 Introduction

The last decades of the twentieth century weremarked by great progress in cognitive neuros-cience, made possible by recently-developedbrain imaging technologies such as functionalmagnetic resonance imaging (fMRI)—which al-lowed for the first time non-invasive study ofthe human brain.

But what is cognitive neuroscience? Wethink it is fair to say that it is above all a meth-odological approach whose results are strongly

influenced by which questions are being askedand how. Studying single neurons and/or thebrain does not necessarily predetermine thequestions to be asked that will help us under-stand how and how much our human nature de-pends upon our brains. Even less so the an-swers. Our purpose here is twofold. On the onehand, we aim to provide a brief overview of cur-rent cognitive neuroscience and its methods. Wefirst present the limitations displayed by most

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 1 | 22

www.open-mind.net

current mainstream cognitive neuroscience, fol-lowed by a proposal for an alternative approach,both in terms of the employed methodology andof its main goals. In short, in contrast withwhat many normally take for granted1, we as-sume the brain level of description to be a ne-cessary but not sufficient condition for studyingintersubjectivity, language, and the human self,which are only properly visible if coupled with afull appreciation of their intertwined relation-ship with the body. This overview has been spe-cifically designed to provide a useful tool for re-searchers working in the humanities. Section 2is entirely devoted to this goal.

On the other hand, the authors of this es-say are a cognitive neuroscientist and a philo-sopher of language, and as such our second pur-pose is to propose how cognitive neurosciencecould provide new insights on specific aspects ofhuman cognition. In section 3 we introduce mir-ror mechanisms and embodied simulation and,in the following sections, we discuss their relev-ance for a new account of intersubjectivity, thehuman self, and language—which privileges thebody as the transcendental foundation of each.

We emphasize the crucial role of the body,conceived as the constitutive source of pre-re-flective consciousness of the self and of theother and as the ground upon which linguisticmeaning is also based. The body we talk aboutin this paper manifests itself in two different,complementary, and closely intertwined ways:2it is a Leib, a lived body entertaining experi-ences of self and others, and a Körper, the so-matic object, of which the brain is a con-1 “Still other accounts of grounded cognition focus on situated action,

social interaction, and the environment (e.g., Barsalou 2003, Bars-alou et al. 2007a, Glenberg 1997, W. Prinz 1997, Rizzolatti &Craighero 2004, Robbins & Aydede 2007, E. Smith & Semin 2004,Yeh & Barsalou 2006). From this perspective, the cognitive systemevolved to support action in specific situations, including social inter-action. These accounts stress interactions between perception, action,the body, the environment, and other agents, typically during goalachievement. It is important to note that the phrase ‘embodied cog-nition’ is often used when referring to this collection of literatures.Problematically, however, ‘embodied cognition’ produces the mis-taken assumption that all researchers in this community believe thatbodily states are necessary for cognition and that these researchersfocus exclusively on bodily states in their investigations. Clearly,however, cognition often proceeds independently of the body, andmany researchers address other forms of grounding.” (Barsalou 2008,p. 619)

2 We find it useful to employ here the distinction originally proposedby Edmund Husserl in Husserl (1973, p. 119).

stitutive part. We posit that this dual nature ofour experienced body can be fully understood—and its genesis revealed—by investigating itsmotor neurophysiological underpinnings at thesub-personal level. The naturalization of inter-subjectivity, the self, and language implies afirst attempt to isolate the constituent compon-ents of the concepts we use to refer to these as-pects of human social cognition by literally in-vestigating what they are made of at the levelof description of the brain–body system. Thisattempt in relation to the notions of intersub-jectivity, the self, and language is intended hereto form an identification of their constitutivemechanisms. We believe that this investigationbecomes really effective only when it is framedwithin both comparative and developmentalperspectives.3

The comparative perspective not only al-lows us to frame human social cognition withinan evolutionary picture, thus providing accessto its phylogenetic antecedents.4 It also greatlyreduces the risk of the empirical investigation ofthe human brain being subordinated to a spe-cific human ontology of mind. A further reasonfor privileging the comparative perspectiveresides in the fact that it also brings us themost finely grained approach to date for study-ing the brain, and the possibility of correlatingsingle neurons’ activity with behaviour and cog-nition—as when studying single neurons’ activ-ity in non-human primates, like macaque mon-keys.

The immanent transcendence5 of thebody’s corporeality can be revealed, we contend,by bringing the analysis back to the level of thebrain–body system; that is, to the level of theKörper. We show that this particular neurocog-3 One of the major contributions to our understanding of human social

cognition is provided by developmental psychology. In this paper, forsake of concision we don’t focus on developmental aspects, in spite ofthe crucial importance we attribute to them to thoroughly addressthe issues we want to address here.

4 It is interesting to note that this comparative perspective seems tosupport the hypothesis of an evolutionary continuity between humanand non-human primates in relation to the emergence of languagefrom our sensori-motor abilities. For a discussion of this topic, seeGlenberg & Gallese (2012).

5 The expression immanent transcendence is meant to signify herethe fact that the body, by means of a biological mechanism (im-manent), can transcend his usual function (for example, motion)to become expression (a model or paradeigma) of this bodilyknowledge.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 2 | 22

www.open-mind.net

nitive approach is beginning to reveal the tightrelationship between a core notion of the bodilyself, its potentiality for action, and motor simu-lation at the level of the cortical motor system.Cognitive neuroscience can enable the analysisof several concepts and notions we normallyrefer to when describing ourselves and our socialcognitive lives. In the present paper we applythis method to the notions of intersubjectivity,the self, and language.

To fully account for the specific quality ofhuman social cognition one cannot undervaluethe linguistic dimension. For this reason, we in-troduce aspects of social cognition related tolanguage and discuss them in terms of embodi-ment, emphasizing the progress and limitationsof this approach. Traditionally, the linguisticand corporeal sensorimotor dimensions of socialcognition have been considered entirely unre-lated. We posit that embodied simulation, con-ceived of as a model for important aspects ofour relation to the world, might help in over-coming this apparently unsolvable dichotomy.We argue that a key aspect defining the uniquespecificity of human language consists in its de-coupling from its usual denotative role. Thismeans that language allows us to talk aboutgeneral concepts such as beauty or mankind,without denoting any particular instance ofthese concepts. In so doing, human languagemanifests its power for abstraction. We discussthese features of human language as instanti-ations of the Greek notion of paradeigma, ori-ginally explored by Aristotle, and relate it toembodied simulation. When a word or syntagm,like the Latin word Rosa, is decoupled from itsusual denotative role, it can function as a gen-eral rule of knowledge, e.g., as a paradigm forthe female nominative case of Latin nouns be-longing to the first declension. The notion ofparadeigma does not establish a connectionbetween a universal principle and its contingentaspects, as in deduction, it rather exemplifies aparticular case of induction: specifically thetransposition of inductive reasoning in in thefield of studies of persuasive communication,known as rethorics, where contingent particularcases lead to general rules describing them.Paradigmatic knowledge, however, differently

from standard cases of induction, connects theparticular with the particular, moving from thecontingent particular situation to an exemplarycase. We propose that embodied simulationcould instantiate such a notion of paradigmaticknowledge, hence enabling its naturalizationand helping us overcome the apparent gapbetween the linguistic and corporeal dimensions.

We conclude by emphasizing how the spe-cific use of cognitive neuroscience here proposedcan lead to a new take on social cognition. Thisnew take brings about a demonstration on em-pirical grounds of the constitutive role played infoundational aspects of social cognition by thehuman body, when conceived of in terms of itsmotor potentialities; hence its transcendentalquality.6 Of course this only covers a partial as-pect of social cognition. However, we think thisapproach has the merit of providing an epistem-ological model, which is also potentially usefulfor empirical investigation into the more cognit-ively sophisticated aspects of human social cog-nition.

2 The cognitive neuroscience of what?

It is a fact and an undisputable truth that therecannot be any mental life without the brain.More controversial is whether the level of de-scription offered by the brain is also sufficientfor providing a thorough and biologically-plaus-ible account of social cognition. We think itisn’t. We would like to ground this assertion ontwo arguments: the first deals with the oftenoverlooked intrinsic limitations of the approachadopting the brain level of description, particu-larly when the brain is considered in isolationand its intimate relation with the body is neg-lected; the second deals with social cognitiveneuroscience’s current prevalent explanatory ob-jectives and contents.

The contemporary emphasis divulgated bythe popular media, namely the supposedly re-volutionary heuristic value of cognitive neuros-cience, mostly rests upon the results of brainimaging techniques, and in particular on fMRI.

6 The “transcendental quality” attributed to the body is intended tomean that the body is considered as the a priori, non-further redu-cible condition of the possibility of experience.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 3 | 22

www.open-mind.net

fMRI is often presented as the ultimate methodof investigation of the human mind. It shouldbe pointed out, though, that fMRI studies donot constitute the whole story in cognitive neur-oscience. Cognitive neuroscience can indeedcarry out its investigation at a more drasticsub-personal level, such as at the level of singleneurons (see below), both in macaque monkeysand, although much more rarely, even in hu-mans. These alternative approaches notwith-standing, the main thrust of cognitive neuros-cience in studying human brain function is, aswe speak, mostly confined to fMRI.7

Unfortunately fMRI only indirectly “sees”the workings of the brain, by measuring neur-ons’ oxygen consumption. Such a measure isalso indirect, as it depends on the local differ-ence between oxygenated and deoxygenatedhemoglobin—the iron-rich molecule housed byred blood cells, which carriess oxygen to allbodily organs and tissues. Oxygenated anddeoxygenated hemoglobin have different para-magnetic behaviours in relation to the strongmagnetic field that is created by a big coil, in-side which the head is placed. The measure ofthis functional parameter allows scientists to es-timate local neural activity in terms of differentMRI (magnetic resonance imaging) signals. Theindirect quality of this kind of estimation ofbrain activity, which is based on local hemody-namic brain responses, inevitably introducesdistortions and noise. Indeed, when studyingany sensorimotor, perceptual, or cognitive func-tion, in order to maximize the so-called “signal-to-noise ratio”, several repetitions of the sametask in many individuals are required.

This means that fMRI allows us to indir-ectly assess the average brain-activation levelinduced by any given task across a populationof no less than twelve to fifteen different indi-viduals. Within each studied individual brainthe spatial resolution of fMRI is within the or-der of few millimetres. This implies that we areable to measure at best the potentially coherentactivation pattern of several hundred thousandneighbouring neurons, possibly also differing

7 For an intriguing discussion of the historical antecedents of brain imagingtechniques and a passionate criticism of the limitations of current use ofbrain imaging by cognitive neuroscience, see Legrenzi & Umiltà (2011).

among one another in terms of their excitatoryor inhibitory role.

Temporal resolution is even worse, since itis in the order of a few seconds. One shouldconsider that action potentials, or “spikes” asneurophysiologists like to call them—the electriccode employed by neurons to “communicate”with each other, and ultimately the true essenceof neurons’ activity—last less than one milli-second. fMRI cannot match such temporal res-olution because it measures the delayed (ofabout two seconds) and prolonged (for aboutfive seconds) local hemodynamic responseproviding neurons with all the oxygen theirelectric activity requires.

As we have previously argued, fMRI is notthe only available experimental methodology forstudying the brain. Many different techniquesare available nowadays (e.g., PET (positronemission tomography), NIRS (near-infraredspectroscopy), Tdcs (transcranial direct-currentstimulation) or TMS (transcranial magneticstimulation)). Particularly, since the revolution-ary introduction in 1927 by the Nobel Prizelaureate Edgar Adrian (Adrian & Matthews1927a, 1927b) of the extracellular microelec-trode, which allows the recording of action po-tentials discharged by single neurons, neuro-physiology has made enormous progress in re-vealing the brain’s physiological mechanisms.Such neurophysiological investigation startedwith the study of the neural circuits thatpreside over elementary sensorimotor beha-viours, like spinal reflexes, finally moving all theway up to the investigation of action and per-ception, reward and emotions, spatial mappingand navigation, working memory, decision-mak-ing, etc., in behaving animals like macaquemonkeys. Unfortunately, such a finely grainedlevel of description—both in terms of spatialand temporal resolution, is most of the timeprecluded in humans.

We posit that the scientific study of inter-subjectivity and the human self requires a com-parative approach, and that this is the only onecapable of connecting the distinctive traits ofhuman nature to their likely phylogenetic pre-cursors. In so doing, and by making use of thesingle-neuron recording approach, neuro-

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 4 | 22

www.open-mind.net

physiological mechanisms and the cortico-cor-tical networks expressing them can be relatedwith several aspects of primates’ social cognitivebehaviour and thus be thoroughly investigated.Conceptual notions like intersubjectivity andthe self should be analyzed in order to betterunderstand their nature, structure, and proper-ties. Such an analysis, which provides us a de-flationary notion of the same concepts, intendedas the identification of their minimal componentand the detailed study of their origin, will bemost successful if driven by a meticulous invest-igation of the underpinning neurophysiologicalmechanisms—which most of the time are avail-able only from the study of non human prim-ates’ brains, hence the necessity of a comparat-ive perspective.

Human brain imaging, because of the in-trinsic limitations we briefly outlined above, canonly provide correlations between particularbrain patterns of activation and particular be-haviours or mental states. This implies that thecorrelation between a particular brain state anda particular phenomenal mental state of a givenindividual human being8 is most informativewhen the specificity and uniqueness of such acorrelation can be firmly established. Unfortu-nately, this is not always the case with fMRIstudies. Very telling is the supposed mindread-ing specificity of some cortical circuits compris-ing the ventral portion of the mesial frontal cor-tex and the TPJ (temporo-parietal junction;e.g., Leslie 2005; Saxe 2006). Such specificity isnot only so far unproven, but is actually con-futed by accumulated evidence (for a lengthierdiscussion of this point and for arguments andexperimental evidence against such specificitysee Ammaniti & Gallese 2014; Gallese 2014).

In spite of all these limitations, thisneuroimaging approach turned out to be veryproductive, enabling us to study for the firsttime in parallel brains, behaviour and cognition,shedding new light not only on human brainstructure, but also on its wiring pattern of con-

8 For sake of concision and focus we do not discuss here the implicatedtopic of the apparently absent synchronicity between brain statesand phenomenal consciousness (remember that fMRI does notprovide a good temporal resolution to firmly match brain states andphenomenal consciousness). We simply want to stress the parallel ex-istence of particular experiences and particular brain states.

nectivity and many of its functions. If we putthe newly-acquired knowledge on brain functionprovided by cognitive neuroscience under scru-tiny, we can make very interesting discoveries.For example, we discovered that in many areasof investigation brain imaging replicates andvalidates at a different scale what had been pre-viously discovered at the single neuron level inanimals like macaques.9

The prominent discoveries, among others,of David Hubel, Torsten Wiesel, and Semir Zekion the functional organization of primates’ cor-tical visual system, like the orientation-, shape-,motion- and colour-selectivity of visual neurons,were made by correlating the discharge activityof single neurons in macaques’ visual corticeswith different parameters of the visual stimulimacaques were looking at (for a comprehensivereview of this literature, see Zeki 1993). Theseresults later promoted a similar investigationcarried out on the human brain by means offMRI. Remarkably enough, a similar functionalarchitecture was detected in the human visualbrain, in spite of the species difference and,most importantly, the different scale at whichthese investigations were carried out: a few hun-dreds recorded neurons at best, in the case ofmacaques’ brains, versus hundreds of thousandsif not millions of activated neurons detected bya local increase in blood flow in the case of thehuman brain.

Face-selective neurons, first described inthe early nineteen-seventies by Charles Grossand colleagues in macaques’ temporal cortex(Gross et al. 1972), and immediately ridiculedas “grand-mother cells”, offer another verytelling example.10 Face-selective brain circuitsappear to be strikingly similar in macaques andhumans (see Ku et al. 2011). Furthermore, evenin human brains single neurons were detectedthat selectively respond to a single face—such

9 The comparative approach is primarily intended here as a compar-ison between humans and other animals. As a consequence, it alsoimplies a comparison between different experimental methods.

10 The notion of the grandmother cell was originally introduced by theneuroscientist Jerry Lettvin to refer to neurons with high integrativepower, which are able to map concepts or objects. The term hassince then mostly been employed with a negative connotation bysupporters of a more distributed population-coding of objects, per-cepts, and memories. For an historical account of the notion of thegrandmother cell, see Gross (2002).

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 5 | 22

www.open-mind.net

as the so-called Jennifer Aniston’s selectiveneurons (see Quiroga et al. 2005). These neur-ons respond to multiple representations of aparticular individual, regardless of the specificvisual features of the picture used. Indeed, theseneurons respond similarly to different picturesof the same person and even to his or her writ-ten or spoken name. The authors of this studyclaimed that their evidence supported the no-tion that single neurons within the human me-dial temporal lobe cortex instantiate the ab-stract representation of the identity of a singleindividual.

Such examples seem to suggest that inspite of the big scale magnification impliedwhen confronting single-neuron data frommacaques and fMRI results from humans, someimportant functional features are neverthelessmanifest across these different levels of descrip-tion (i.e., single neurons vs. brain areas). Onecan study canonical or mirror neurons (see be-low) by recording the activity of a few hundredspiking neurons from a behaving macaque mon-key during object and action observation, re-spectively. The same results can be replicatedby detecting, by means of fMRI, and during ob-ject and action observation, the simultaneousactivation of hundreds of thousands of humanneurons within analogous cortical areas of thehuman brain.

This remarkable but often neglected factcannot be the result of a pure coincidence. Thisevidence should thus invite us to resist and argueagainst those who downplay the heuristic powerof single-neuron recording. Their thesis is that be-cause of a supposedly incommensurable gapbetween single neurons and the incredible com-plexity of the human brain, where informationwould be exclusively mapped at the level of largepopulations of poorly selective neurons, it doesn’tmake any sense to study the brain by recordingsingle neurons. The fact however is that in spiteof the almost astronomic figures characterizingthe human brain (about 100 billion neurons, eachof which connects with thousands of other neur-ons), its complexity does not parallel such astro-nomic figures, or at least not in such a way as todeny any heuristic value to the single-neuron re-cording approach. Let’s see why.

As argued by Chittka & Niven (2009),brain size may have less of a relationship withbehavioural repertoire and cognitive capacitythan generally assumed. According to the sameauthors, larger brains are, in part at least, aconsequence of larger neurons that are neces-sary in large animals due to basic biophysicalconstraints. Larger brains also contain greaterreplication of neuronal circuits, adding precisionto sensory processes, detail to perception, moreparallel processing, enlarged storage capacity,and greater plasticity. These advantages, main-tain Chittka & Niven (2009), are unlikely toproduce the qualitative shifts in behaviour thatare often assumed to accompany increased brainsize, or at least not in a one-to-one manner.

The evidence so far briefly reviewed andthat we will present in the next sections suggestthat some functional properties of the brain ex-hibit a sort of “fractal quality”, such that theycan be appreciated at different scales and levelsof investigation. For these reasons fMRI cannotbe the sole neurocognitive approach to humansocial cognition,11 but it must be complementedby other approaches compensating for some ofits deficiencies: like TMS, EEG (electroenceph-alography), and the comparative functionalstudy of non-human primates by means of brainimaging and single-neuron recordings.

After having clarified what we take to bethe often-neglected limitations of cognitiveneuroscience that may hinder its potential heur-istic power (namely, that fMRI offers only anindirect estimation of brain activity, inferred bymeasuring neurons’ oxygen consumption, andthis inevitably leads to distortions and noise,and that it does not provide good temporal res-olution because it measures the delayed andprolonged hemodynamic responses due to neur-ons oxygenation), let us now move to thesecond argument against the sufficiency satis-faction condition of the current approach of cog-nitive neuroscience to the study of human socialcognition. This argument concerns the explanat-ory goals and contents of contemporary main-11 Maybe no one explicitly claims this. However, it is very common that

researchers neglect many (or all) of the complementary techniquesthat have been here suggested to be necessary and draw inferencesabout human social cognition and its neural implementation. As anexample, see papers by Ian Apperly.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 6 | 22

www.open-mind.net

stream cognitive neuroscience. Vast quarterswithin cognitive neuroscience are still todaystrongly influenced by classical cognitivism, onone side, and by evolutionary psychology on theother. Classical cognitive science is the bearer ofa solipsistic vision of the mind, according towhich focusing on the mind of the single indi-vidual is all that is required in order to definewhat a mind is and how it works. The image ofthe mind that classical cognitive science gives usis that of a functional system whose processesare described in terms of manipulations of in-formational symbols in accordance with a seriesof formal syntactic rules.

According to evolutionary psychology, bycontrast, the human mind is a set of cognitivemodules, each of which has been selected duringevolution for its adaptive value. Major figures ofthis current, such as John Tooby and Leda Cos-mides, have gone as far as maintaining that thebrain is a physical system that works like acomputer (Cosmides & Tooby 1997). Accordingto Steven Pinker (1994, 1997), our cognitive lifecan be referred to in terms of the function of aseries of modules like the linguistic module, themodule for the Theory of the Mind, etc.

Based on this theoretical framework, inthe last twenty years cognitive neurosciencewhen investigating human social cognition hasmainly tried to locate—as mentioned above—the cognitive modules in the human brain. Suchan approach suffers from ontological reduction-ism, because it reifies human subjectivity andintersubjectivity within a mass of neurons vari-ously distributed in the brain. This ontologicalreductionism chooses as a level of descriptionthe activation of segregated cerebral areas or, atbest, the activation of circuits that connect dif-ferent areas and regions of the brain. However,if brain imaging is not backed up by a detailedphenomenological analysis of the perceptual,motor, and cognitive processes that it aims tostudy and—even more importantly—if the res-ults are not interpreted, as previously argued,on the basis of the study of the activity ofsingle neurons in animal models, and the studyof clinical patients, then cognitive neuroscience,when exclusively consisting in brain imaging,loses much of its heuristic power. Without the

demonstration of the specific correlationbetween brain states and mind states and theexplanation of such correlation, much of thecontemporary brain imaging approach to socialcognition looks like a sort of high-tech versionof phrenology.

For this reason a “phenomenologization”of cognitive neuroscience is desirable, as Gallesehas proposed before (see Gallese 2007, 2009,2011, 2014). In Gallese’s view, to “phenomeno-logize” cognitive neuroscience means to startneuroscientific research from the analysis ofsubjective experience and of the role that theliving body plays in the constitution of our ex-perience of material objects and of other livingindividuals. In this way, the empirical study ofthe genetic aspects of subjectivity and intersub-jectivity can be pursued on new bases—if com-pared to those thus far adopted by classical cog-nitivism. Francisco Varela a few years ago real-ized a similar possibility and set out on a path-way of analysis in this direction (Varela &Shear 1999).

Times change, however. We are insistingon nothing less than a change of paradigm. Anew neuroscientific approach to the study of thehuman mind is gaining momentum. It capital-izes upon the study of the bodily dimension ofknowledge: the so-called “embodied cognition”approach. In the next section we introduce mir-ror neurons and embodied simulation. Our pur-pose is to show that, starting from a sub-per-sonal neuroscientific description of the prag-matic relationship with the world, a pathwaycan be traced to define the forms of subjectivityand intersubjectivity that distinguish humannature, rooted in the bodily interacting natureof human beings.

3 Mirroring mechanisms and embodied simulation

The discovery in the early 1990s of mirror neur-ons in the brain of macaques (Gallese et al.1996; Rizzolatti et al. 1996), and the subsequentdiscovery of mirror mechanisms in the humanbrain (see Gallese et al. 2004; Rizzolatti & Sini-gaglia 2010) suggest that there exists a directmodality of access to the meaning of other

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 7 | 22

www.open-mind.net

people’s behaviours—a modality that can be setaside from the explicit attribution of proposi-tional attitudes. Mirror neurons are motor neur-ons, originally discovered in macaques’ ventralpremotor cortex area F5, later on also found inthe reciprocally-connected posterior parietalareas AIP and PFG. Mirror neurons not onlyrespond to the execution of movements and ac-tions, but also respond to the perception of ac-tions executed by others. Mirror neurons mapthe action of others on the observers’ motorrepresentation of the same action. Further re-search also demonstrated in the human brainthe existence of a mechanism directly mappingaction perception and execution, defined as theMirror Mechanism (MM; for a recent review,see Ammaniti & Gallese 2014; Gallese 2014). Inaddition, in humans the motor brain is mul-timodal. Thus, it doesn’t matter whether we seeor hear the noise made by someone crackingpeanuts, or locking a door. Different—visualand auditory—sensory accounts of the samemotor behaviour activate the very motor neur-ons that normally enable it. The brain circuitsshowing evidence of the MM, connecting frontaland posterior parietal multimodal motor neur-ons, most likely analogous to macaques’ mirrorneurons, map a given motor content like “reachout” or “grasp” not only during their perform-ance, but also when perceiving the same motorbehaviour performed by someone else, when im-itating it, or when imagining performing itwhile remaining perfectly still. The relationalcharacter of behaviour as mapped by the cor-tical motor system enables the appreciation ofpurpose without relying on explicit proposi-tional inference.

Altogether, these findings led to the for-mulation of the “Motor Cognition” hypothesisas a crucial element in the emergence of socialcognition (Gallese 2009). According to this hy-pothesis, cognitive abilities like the hierarchicalrepresentation of action with respect to a distalgoal, the detection of motor goals in others’ be-haviour, and action anticipation are possible be-cause of the peculiar functional architecture ofthe motor system, organized in terms of goal-directed motor acts. Traditionally, the relationbetween actions and their outcomes is assumed

to be largely independent of the motor pro-cesses and representations underpinning actionexecution. Such processes and representationsallegedly concern elementary motor featuressuch as joint displacements or muscle contrac-tions only. However, solid empirical evidencechallenged this view. Motor processes may in-volve motor representations of action goals (e.g.,to grasp, to place, etc.), and not only kinematicor dynamic components of actions. This sug-gests that beliefs, desires, and intentions areneither primitive nor the only bearers of inten-tionality in action. We do not necessarily needto metarepresent in propositional format the in-tentions of others to understand them. Motoroutcomes and motor intentions are part of the“vocabulary” that is spoken by the motor sys-tem. On occasion we do not explicitly ascribeintentions to others; we simply detect them. In-deed, we posit that motor representation isenough to ground the directedness of an actionto its outcome (Gallese 2000, 2003b; Butterfill& Sinigaglia 2014; compare also Gallagher’s2005 notion of direct perception).

One of the consequences of the discoveryof mirror neurons was the possibility of derivingsubjectivity from intersubjectivity at the sub-personal level of description. The sense of self isprecociously developed, beginning from a selfthat is first of all physical and bodily, andwhich is constituted precisely by the possibilityof interacting and acting with the other. Em-bodied simulation can provide the neurobiolo-gical basis for early forms of intersubjectivity,from which the sense of the self is built. Thediscovery of mirror neurons and the simulationmechanism would therefore seem to furtherstress that being a self also implies being withthe other. The model of intersubjectivity sug-gested by mirror mechanisms and embodiedsimulation correlatively sheds new light on thesubjective dimension of existence. Let us seefirst what type of intersubjectivity mirror neur-ons seem to suggest.

The discovery of mirror neurons gives us anew empirically-grounded notion of intersub-jectivity connoted first and foremost as intercor-poreality—the mutual resonance of intentionallymeaningful sensorimotor behaviours. The ability

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 8 | 22

www.open-mind.net

to understand others as intentional agents doesnot exclusively depend on propositional compet-ence, but it is highly dependent on the rela-tional nature of action. According to this hypo-thesis, it is possible to directly understand themeaning of other people’s basic actions thanksto a motor equivalence between what others doand what the observer can do. Intercorporealitythus becomes the main source of knowledgethat we have of others. The motor simulationinstantiated by neurons endowed with “mirrorproperties” is probably the neural correlate ofthis human faculty, describable in functionalterms such as “embodied simulation” (Gallese2003a, 2005, 2011; Gallese & Sinigaglia 2011b).

Action constitutes only one dimension ofthe rich baggage of experiences involved in in-terpersonal relations. Every interpersonal rela-tion implies the sharing of a multiplicity ofstates like, for instance, the experience of emo-tions and sensations. Today we know that thevery nervous structures involved in the subject-ive experience of emotions and sensations arealso active when such emotions and sensationsare recognized in others. A multiplicity of “mir-roring” mechanisms is present in our brain. Itwas proposed that these mechanisms, thanks tothe “intentional attunement” they generate(Gallese 2006), allow us to recognize others asour fellows, likely making intersubjective com-munication and mutual implicit understandingpossible. The functional architecture of embod-ied simulation seems to constitute a basic char-acteristic of our brain, making possible our richand diversified intersubjective experiences, andlying at the basis of our capacity to empathizewith others.

4 Body and self

After having delineated a deflationary neurobio-logically-grounded account of basic aspects ofintersubjectivity, namely an account focused onthe minimal core mechanisms of intersubjectiv-ity, let us now address the relationship betweenbody and self. A minimal manifestation of thesense of self can already be identified in ourfirst bodily experiences, and this highlights thepotential contribution of bodily experiences to

its constitution. Some aspects of the minimalself proposed by contemporary philosophicaland empirical research are the notion of first-person perspective, the “mineness” of the phe-nomenal field (Meinigkeit), embodiment ofpoint of view, and issues of agency and bodyownership (Cermolacce et al. 2007).12 On thephilosophical side, phenomenology emphasizesthe necessity of embodiment of the self for allthe above-cited aspects of self experience. As ar-gued by Cermolacce et al. (2007, p. 704, foot-note 3), in phenomenology

the field of experience is not yet consideredto be subjective because this predicatealready implies that there is a subject. Forphenomenology, the very idea of the subjectarticulates itself in experience. In this sense,the manifestation and appearing of experi-ence are the conditions for the experience ofthe subject in question.

This philosophical standpoint has important im-plications for the empirical investigation of theneural correlates of the self.13 Rather than em-pirically addressing the self by starting with asearch for the neural correlates of a pre-defined,explicit, and reflective self-consciousness, we be-lieve it to be more productive to investigatewhat what set of constitutive conditions allowsan implicit and pre-reflective sense of self toemerge, and how this is effected. The interestingquestions to be first answered are: “What en-ables the basic experience of ourselves as bodilyselves? What enables us to implicitly distin-guish ourselves, as bodily selves, from other hu-man bodily selves?” In the following we reviewand discuss recent empirical evidence providingpreliminary answers to these questions.

12 Again, for sake of concision, we do not deal here with the relation-ship between the notion of a core, minimal self as a bodily self andagency and body ownership. On this topic, see Gallese & Sinigaglia(2010, 2011a). Moreover, it is worth noting that the arguments pro-posed in this section in relation to the notion of a minimal bodilyself could be applied to other non-human animals. The possibilitythat high-level self-awareness can emerge from a primitive and non-conceptual form of self-awareness, and that it is possible that weshare this basic level of the sense of self with other non-human anim-als, has already been discussed. For a discussion of this and other re-lated topics see Bermúdez (2003).

13 See Vogeley et al. (2003) and Vogeley et al. (2004) for an investiga-tion of the neural correlates of the first-person perspective.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 9 | 22

www.open-mind.net

The relationship between the minimalsense of self and the cortical motor system wasrecently revealed. The motor experience of one’sown body, even at a covert level, allows an im-plicit and pre-reflective bodily self-knowledge toemerge, leading to a self/other distinction. In-deed it was recently shown that in a task inwhich differently rotated static pictures of rightand left human hands were presented, parti-cipants who had to determine whether each ob-served hand was the right or the left one pro-duced faster responses when observing the pic-tures of their dominant hand with respect toothers’ hands (Ferri et al. 2011). However, whenparticipants were asked to explicitly discrimin-ate between their hands and the hands of oth-ers, the self-advantage disappeared. Implicit andexplicit recognition of the bodily self dissoci-ated: only implicit recognition of the bodily self,mapped in motor terms, facilitated implicitbodily self-processing.

A subsequent fMRI study by Ferri et al.(2012), using a similar hand mental rotationtask, demonstrated that a bilateral cortical net-work formed by the supplementary and pre-sup-plementary motor areas, the anterior insula,and the occipital cortex was activated duringprocessing of participants’ own hands. Further-more, the contralateral ventral premotor cortexwas uniquely and specifically activated duringmental rotation of the participants’ own domin-ant hands. The ventral premotor cortex mightrepresent one of the essential anatomo-func-tional bases for the motor aspect of bodily self-hood, also in light of its role in integrating self-related multisensory information. This hypo-thesis is corroborated by clinical and functionalevidence showing its systematic involvementwith body awareness (Ehrsson et al. 2004; Bertiet al. 2005; Arzy et al. 2006). This evidencedemonstrates a tight relationship between thebodily self-related multimodal integration car-ried out by the cortical motor areas, specifyingthe motor potentialities of one’s body and guid-ing its motor behaviour, and the implicit aware-ness one entertains of one’s body as one’s ownbody and of one’s behaviour as one’s own beha-viour. Because the ventral premotor cortex isanatomically connected to visual and somato-

sensory areas in the posterior parietal cortexand to frontal motor areas we hypothesize thatpremotor cortex activity, by underpinning thedetection of congruent multisensory signals fromone’s own body, could be at the origin of theexperience of owning one’s own body parts.

This minimal notion of the self, namelythe bodily self as power-for-action (see Gallese& Sinigaglia 2010, 2011a), tacitly presupposesownership of an action-capable agentive entity;hence it primarily rests upon the functionalityof the motor system. As we just saw, empiricalevidence supports the neural realization of thisimplicit aspect of selfhood in the brain’s motorcortex. Since the minimal bodily self rests neur-ally on the motor system, it logically followsthat characteristics of the latter are defining forthe former. This implies that one could attrib-ute to the minimal bodily self known features ofthe motor system, including its capacities andlimitations. The motor aspects of the bodily selfprovide the means to integrate self-related mul-timodal sensory information about the bodyand the world with which it interacts. This isalso important from a theoretical point of view,because it opens the possibility of linking theopenness of the self to the world to the motorpotentialities its bodily nature entails.

One could then posit that the minimalbodily self when conceived in terms of its motorpotentialities has a dual function. On the onehand, it constitutes important aspects of thebasic sense of self. On the other, it shapes ourperception and pre-reflective conception of oth-ers as other selves incarnated in a motorly-cap-able physical body with capacities and experi-ences similar to ours. Through mirror mechan-isms and embodied simulation, others appear tous as second selves, or second persons. We be-lieve that this perspective provides a more vividexperience of intersubjectivity, relative to thedetached, propositional deliberation on the ex-periences of others available in standard mindreading of others.

5 Body and language: Reflexiveness

According to the perspective so far delineated,body, actions, and feelings play a direct role in

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 10 | 22

www.open-mind.net

our knowledge of others. The question remainsopen as to whether our propositional represent-ations are totally separate from this bodily di-mension. Our hypothesis is that they are not.But it remains a fact that linguistic and bodilycognition afford us diversified modalities of epi-stemic access to the world, even though oftensuch modalities contaminate one another andare inevitably interwoven.

The mind, from the perspective delineatedhere, is therefore an embodied mind, though itwould be more correct to speak of a corporealmind. The concept of embodiment can induceone to think that a mind pre-existing the bodycan subsequently live in it, and use it. Thetruth is that mind and body are two levels ofdescription of the same reality, which manifestsdifferent properties according to the chosenlevel of description and the language employedto describe it. A thought is neither a muscle nora neuron. But its contents, the contents of ourmental representations, are inconceivablewithout our corporeity. Likewise it is difficult toimagine how the representational format of apropositional type can have developed withoutour corporeality. Language somehow allows us,as we will see, to transcend our corporeity; nev-ertheless, we posit that the bond with the bodyis always present.

A few years ago, Gallese (2000) proposedthat we look at the evolution of human lan-guage as an exaptation14 of functional sensor-imotor processes, which put them into the ser-vice of human linguistic competence. The hypo-thesis of exaptation was then developed in sub-sequent papers and later elaborated in terms of“neural exploitation” (Gallese & Lakoff 2005;Gallese 2008), or “neural reuse” (Gallese 2014).“Neural exploitation” consists in the reuse ofneural resources, originally evolved to guide ourinteractions with the world, to serve the morerecently evolved linguistic competence. This no-tion of reuse implies a functional uncoupling ofthe sensorimotor system from muscular output,to guide the generative-syntactic aspects of lan-guage by functionally connecting it to the pre-

14 Exaptation refers to the shift in the course of evolution of a giventrait or mechanism, which is later on reused to serve new purposesand functions (see Gould & Lewontin 1979).

frontal and, more generally, non-sensorimotorcircuits. According to this view, intentionality,the aboutness of our representations, is—in thefirst place—an exapted property of the actionmodels instantiated by the cortical motor sys-tem (Gallese 2000, p. 34). The sensorimotor sys-tem, when uncoupled from muscular output,makes available to us a model, or paradigm, ofour motor knowledge. As such, not only ithouses causative properties but also contentproperties. And this relation to a content, oraboutness, is a primitive expression of intention-ality, then exploited by other forms of repres-entations. This perspective on reuse is acquiringmore and more supporters (see Dehaene 2005;Anderson 2010 ).15

Compelling evidence shows that humans,when processing language, activate the motorsystem both at the phono-articulatory and atthe semantic level. When listening to spokenwords or looking at someone speaking to us, ourmotor system simulates the phono-articulatorygestures employed to produce those very samewords. Furthermore, processing action-relatedlinguistic expressions activates regions of themotor system congruent in somatotopic fashionwith the processed semantic content. Reading orlistening to a sentence describing a hand actionactivates the motor representation of the sameaction (for a review, see Gallese 2008; Glenberg& Gallese 2012). Interestingly, somatotopic mo-tor activation has also been observed during thecomprehension of abstract and figurative use oflanguage such as metaphors and idioms (e.g.,Guan et al. 2013; Boulenger 2012; see alsoGallese & Lakoff 2005 on the bodily foundationof concepts). However, it is important to notethat embodied simulation is not always involvedin language comprehension, and that there is nocontradiction in saying this. There are cases inwhich language, at least at the content level, isnot tied to any form of bodily knowledge. Insuch cases (e.g., when we talk about the notionssuch as moral judgement or intelligence) no em-bodied simulation is likely to be at play.

Nevertheless, the problems that languageraises for the embodied perspective on human15 For a discussion of different views on the notion of reuse, see Gallese

(2014).

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 11 | 22

www.open-mind.net

social cognition are still enormous. As clearlyunderlined, among others, by the Italian philo-sopher Paolo Virno (2003, 2011), the commonlinguistic space shared by a community ofspeakers proves to be incommensurably differ-ent from the pre-linguistic one. The linguisticdimension is based on a distinction between lin-guistic utterances and facts about the world, bethey referable to physical or psychologicalevents. We can say that “today the sun is shin-ing” and be understood, even if outside the win-dow snow is falling. Or we can maintain that“all Italians want to pay taxes”, again being un-derstood and simultaneously contradicted bythe factual truth of the enormous tax evasion inour country.

According to Virno, the gap betweenmeaning and denotation (what he calls theneutrality of meaning, namely the fact that themeaning of a word such as, for example, “man”,can be understood apart from any reference toan instance of man) is referable to linguistic re-flexiveness, i.e., to the fact that language refersto itself and that with words we can talk aboutother words. It seems to us that the reflexive-ness of language is a product of the symbolicnature of linguistic representations. The sym-bolic nature of such representations is what al-lows language to break away from the “here andnow”; it is what allows the neutrality of mean-ing.

In order for a sign to be symbolic it neces-sarily has to be reflexive. What makes a sign asymbol is its being part of a system in whicheach term is correlatively defined in relation tothe other terms within the system and in rela-tion to the renegotiation of this relationshipconstantly taking place within the system itself.It is the use of a symbol within a given contextthat each time redefines relationships inside thelanguage system.

Thus, symbolic relationships are by defini-tion characterized by reflexiveness. Symbols aredefined through other symbols. This level of re-flexiveness is pre-theoretical; it emerges in thelinguistic activity of each speaker and leads to aform of linguistic awareness of a practical char-acter. This practical linguistic awareness hasbeen called the epilinguistic quality and thus it

has been distinguished from the theoreticalquality that is expressed in the metalanguage oflinguistics (Culioli 1968; Lo Piparo 2003). Theconcept of epilinguistic quality refers to the nat-ural tendency of speakers to reflect on their ownlanguage—a tendency made possible by the dis-tinctive quality of language being able to speakof itself.

The uniqueness of human language is alsomaintained by classical cognitivism and by cog-nitive linguistics, but for very different reasons.The otherness of human language in comparisonwith other systems of communication known inthe animal world derives from its linguistic re-cursive quality. In an often-quoted article writ-ten some years ago (Hauser et al. 2002) definedthe faculty of language in a narrow sense (FLN)as being expressed by recursivity. Nevertheless,this perspective, in addition to suffering fromthe usual cognitivist solipsism, is exposed tocomparative verification in the animal world. Ifthe FLN marks human linguistic uniqueness interms of syntactic recursivity, the latter must beentirely absent in the extra-human animal king-dom.

Actually, the facts tell us exactly the op-posite. Recent studies (Gentner et al. 2006; Abe& Watanabe 2011; see also Margoliash & Nus-baum 2009; Bloomfield et al. 2011) have shownthat singing species of birds like starlings orfinches demonstrate, both in the production andin the reception of conspecifics’ vocalizations,the ability to produce and to extract recursivesyntactic characteristics. The study by Abe andWatanabe also shows that the development ofthis competence is dependent on social encoun-ters with the vocalizations of other conspecificindividuals. Finally, these authors have shownthat lesion of the lateral magnocellular nucleusof the anterior nidopallium, a motor structurecomparable to the basal ganglia of primates, in-volved both in the production and the percep-tion of song, prevents finches from discriminat-ing the syntactic-recursive characteristics of thesong they hear.

These results show that the best strategyfor studying some of the most relevant aspectsof human social cognition, even demonstratingthe bases of their uniqueness, consists in a pre-

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 12 | 22

www.open-mind.net

liminary recognition of the mechanisms and fac-ulties that we share with the rest of the animalworld. As maintained in the past (Gallese2003b, 2008), the difference between human andnonhuman nature could originally have been ofa quantitative rather than a qualitativenature.16

6 Body and language: Facts and challenges

One of the key challenges for the embodied ap-proach to human social cognition consists intrying to understand whether and how our bod-ily nature determines some of our linguisticactivities, such as denying, asking, or doubting,that seem to be exclusively human. Are lin-guistic activities as those ones anchored to bod-ily mechanisms? The question is open and em-pirical research must address this challenge inthe coming years. In the meantime, at least at apurely speculative level, let us try to delineate apossible point of contact between the anthropo-genic power of language and embodied simula-tion.17

There is indeed a way to connect the com-mon pre-linguistic sphere to the linguistic one(Gallese 2003b, 2007, 2008; Gallese & Lakoff2005; Glenberg & Gallese 2012). This consists inshowing that language, when it refers to the bodyin action, brings into play the neural resourcesnormally used to move that very same body. See-ing someone performing an action, like grabbingan object, and listening to or reading the lin-guistic description of that action lead to a similarmotor simulation that activates some of the sameregions of our cortical motor system, includingthose with mirror properties, normally activatedwhen we actually perform that action.

These data on the role of simulation in un-derstanding language (see Pulvermüller 2013 fora review of this topic) broadly confirm a thesisalready discussed in the history of philosophy(for instance, by Epicurus, Campanella, Vico,

16 According to this perspective, linguistic syntax could originate andbe modelled upon syntactic motor competence, the latter being ex-apted and put at the service of the new linguistic competence (seeGallese 2007, 2008).

17 With the expression “anthropogenic power of language” we meanthat the human nature, as we know it, depends on language.

see Usener 1887 and Firpo 1940 or Condillaco2001). The thesis in question claims for thebodily, sensory, and motor dimensions a con-stitutive role in language production and under-standing. However, it seems that the relation-ship between language and body does not movein a single direction. The fact is that languageis without doubts constitutive of human natureand, as such, it seems to offer us wholly humanmodalities of experiencing our corporeity.

In this sense, neuroscientific data on therole of simulation during understanding of lan-guage also lend themselves to a mirror and com-plementary reading with respect to that previ-ously proposed. On the one hand it is plausiblethat embodied simulation might play a crucialrole in understanding language. Indeed, if onereversibly interferes with this process, for in-stance by means of TMS stimulation, under-standing of language is jeopardized. On theother hand, language allows us—and in this weare unique among all living species—to fix andrelive specific aspects of our bodily experience.Through language we can crystallize and relivefragments of experiences that are not topical,that is to say are not my experiences now, butbecome a paradigm, a model, for understandingourselves and others.

In the following section we discuss the roleof embodied simulation seen as a paradigm ormodel in the light of the Aristotelian notion ofparadeigma.18 For the time being it suffices tostress that the possibility of hypostatizing andthen segments of our experiences independentlyof our immediate physical context, or independ-ently of specific physical stimuli, is a possibilitythat only the possession of language allows usto experience. The faculty of language is there-fore, on one side, rooted in corporeality but, inturn, changes and moulds our way of livingbodily experiences.

7 Body and language: Embodied simulation as a paradigm?

The relation between body and language was toa great extent underestimated in the last cen-18 For an earlier formulation of this hypothesis, see Gallese

(2013).

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 13 | 22

www.open-mind.net

tury, thanks, above all, to Chomsky’s major in-fluence. In 1966 Chomsky published a book sig-nificantly entitled Cartesian Linguistics.Descartes is the originator of the idea that lan-guage has little to do with the body.19 TheCartesian thesis on the relationship betweenlanguage and body implies, on one side, thatthe body is not a substratum and material oflanguage and, on the other, that language is ex-clusively a tool that expresses a thought formedindependently of language itself. According toDescartes (1642) and the Cartesian tradition inwhich Chomsky stands, language is a toolthrough which we manifest an autonomousthought that precedes language—a thoughtstructured by logic but certainly not by lan-guage, whose role is circumscribed and downs-ized to that of being a mere label of thoughts(cf. Hinzen & Sheehan 2013 for a critical discus-sion of the issue).

The theses informing the Cartesian idea oflanguage are challengeable nowadays. Languagemakes meaning general, releasing it from thecontext, that is, from the dimensions of who,what, how, where, and when. Language, inother words, provides us with a unique modalityof reference to the world, allowing us at thesame time to transcend contingent determina-tions and to define them at a different level,thanks to the use of concepts like subject, ob-ject, time, space, universal, etc. It is perhapsnot trivial to notice that such concepts corres-pond to precise grammatical structures andthat, most likely, the use of a grammatically-structured language contributed, by co-evolu-tionary dynamics, to the structuring of rationalthought characterized by such features (Hinzen& Sheehan 2013).

Hence, thanks to language we can speakof humankind without referring in particularto any of the single individuals sharing theproperty of belonging to the human species.We can speak of a subject aside from the indi-vidual embodiments of this attribute, etc.Language, as stressed by Virno, furnishes uswith general meanings, that is, meanings valid

19 It is worth noting that Descartes also defended the related thesisthat animals don’t have soul exactly because they do not have lan-guage. Cf. Descartes (1637).

for everybody but, at the same time, mean-ings that do not necessarily denote a particu-lar instantiation.

Interestingly enough, according to GiorgioAgamben (2008) what holds “for everybody andnobody” is referable to the Greek notion ofparadeigma, originally explored by Aristotle.The paradeigma is a typical form of rhetoricalreasoning that moves between individual andindividual according to a form of bipolar analo-gical knowledge. Agamben (2008, pp. 23-24),radicalizing Aristotle’s theses, maintains thatthe paradigm can only be conceived of byabandoning the dichotomy between individualand universal: the rule does not exist before thesingle cases to which it is applied. The rule isnothing but its own exhibition in the singlecases themselves, which thus it renders intelli-gible.

By applying the notion of paradigm to thegrammatical “rules” of language, Agambentouches upon a central point: the so-called lin-guistic rule derives from the suspension of theconcrete denotative application:

[t]hat is to say, in order to be able to serveas an example, the syntagm must be sus-pended from its normal function, and, nev-ertheless, it is precisely through this non-operation and this suspension that it canshow how the syntagm works, can allowthe formulation of the rule. (Agamben2008, p. 26)

To better explain the notions of rule and sus-pension of a denotative application, Agambenrefers to Latin declensions. When we want tolearn the first declension we inflect a noun, forexample “rosa”, “rosae”, etc… In so doing, weare suspending the usual denotative applicationof this noun and, by means of this suspension,we are showing how the declension works. Ac-cording to Agamben, “[…] in the paradigm, in-telligibility does not precede the phenomenon,but is, so to speak, ‘alongside’ it (parà)” (2008,p. 29). In other words “[…] in the paradigmthere is not an origin or an arché: every phe-nomenon is the origin, every image is archaic”(Agamben 2008, p. 33).

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 14 | 22

www.open-mind.net

On Agamben’s reading, the Aristotelianparadeigma is a good model for describing thecreation of linguistic rules. Starting from Agam-ben’s intuition and seeking to move one stepfurther, the hypothesis that we want to explorehere is that the notion of paradeigma is a goodmodel not only for the creation of linguisticrules but also for the definition of the embodiedsimulation mechanism. In this connection, simu-lation allows us, at a sensorimotor level, to hy-postatize and reuse what holds “for everybodyand nobody”.

To understand to what extent the analogybetween embodied simulation and paradeigmaworks it is necessary to go back to Aristotle.What is meant by paradeigma in Aristotelianthought and in what context does Aristotlemake use of this notion?

The paradeigma, as already anticipated, isa typical is a typical argument form used topersuade and devoted to the discussion of“things that can be otherwise.” Aristotle dis-cusses this argument form, which does not haveany demonstrative aim, both in Prior Analyticsand in Rhetoric. Argumentation based on theparadeigma, for example, consists in thepresentation by the orator of an exemplary case,based on a historical fact or a figment of theimagination, as in the case of fables. It is thejuxtaposition of the present situation and an ex-emplary one that guides, or should guide, theactions of the person to whom the argumenta-tion is addressed. Thus the paradeigma, amongrhetorical argumentations, is that which goesfrom the particular to the particular, from anexemplary case to the present situation. Argu-mentation based on the paradeigma does notmake a claim for universality. The orator is notbound to offer an exhaustive number of casesjustifying a universally valid conclusion. Onecase is sufficient, provided that it is particularlysuitable, and precisely exemplary, in relation tothe context in which the argumentative dis-course takes place.

For these reasons, though resembling in-ductive reasoning (epagoghé), which proceedsfrom the particular to the universal, and indeedconsidered by Aristotle himself as the transposi-tion of inductive reasoning to the rhetorical

sphere, the paradeigma conquers its ownautonomous space. To confirm this, one needonly to think that in Prior Analytics Aristotle(Ross 1978) devotes two separate chapters toparadigm and induction: respectively XXIV andXXI of Book II.

On the one side the paradeigma, whichproceeds “from the part to the part” (PriorAnalytics 69a, 15), are peculiar aspects distin-guishing it from the epagoghé; on the other, it isby all means a form of induction, as Aristotleexpressly affirms at the start of Chapter XX ofBook II of Rhetoric. According to Piazza (2008,p. 117) there are at least two reasons why theparadeigma can still be considered a form ofepagoghé, despite the peculiarities that charac-terize it. Both these reasons seem interesting,not only for the definition of paradigm that,starting from Aristotle, Agamben discusses inrelation to linguistic praxis, but also and aboveall in the framework of a reflection on the mir-ror mechanisms enacted in embodied simula-tion.

Following Piazza’s (2008, p. 117) readingof Aristotle, the first of the characteristics of in-ductive reasoning also found in the paradeigmaconsists in always proceeding from what is “bestknown and first for us” (Aristotle, Analyticaposteriora II.19), or from what is for us mostimmediate and most easily accessible, becausebeing part of our baggage of experiences andknowledge. The second characteristic is, instead,identifying similarities between particular cases.

At another level of analysis, both thesefeatures also characterize embodied simulation.One condition for the simulation mechanism’sbeing enacted is sharing a baggage of (motor)experiences and knowledge. Embodied simula-tion is enacted starting from what for us is“first,” i.e., what for us is known and easily ac-cessible in terms of motor potentialities and ex-periences. Sharing a repertoire of practices, ex-periences, and sensations is therefore an essen-tial condition, since only by starting from whatis well known to us it is possible to identify ana-logies between our actions and others’. We un-derstand the other starting from our own bodilyexperience, which is what is “best known andfirst for us”, again using Aristotle’s words. On

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 15 | 22

www.open-mind.net

the basis of this knowledge we identify similarelements in our experiences as well as in thoseof others.

Embodied simulation, when manifested inthe phenomenon of action, emotion, or sensa-tion mirroring always involves an original I-thourelationship in which the “thou” is the termwith respect to which the self is constituted. Onthe other hand, the “self” is the basis on whichimmediate and implicit understanding of the“thou” is possible.

The analogy with the cognitive mechanismsubtended by paradigmatic reasoning appearsevident. Indeed, in the case of Aristotle’sparadeigma, an example, a particular case, isunderstood because it is close to our feeling, ourexperiences, and our baggage of knowledge. Andnevertheless the process does not stop here.This form of understanding of a particular thatis not me will lead me to new conclusions andto a deeper understanding of myself, of my par-ticular case, and of my situation. Our experi-ences are therefore the measure from which weunderstand others and their experiences (i.e.,our previous actions, emotions, and so forth).And others’ experiences (i.e., their actions, emo-tions and so forth) are for us a condition fordeeper understanding of ourselves. Thus, theembodied simulation underpinning my presentexperience is also a paradeigma from which Ican understand what I observe in others anddraw inferences from it for others and for my-self.

The embodied simulation mechanism, thusdefined, is constitutive of the process of con-struction of meaning. In this connection, em-bodied simulation enacted while understandinglanguage is not my present experience but theparadeigma in relation to which some of our lin-guistic expressions acquire a meaning that isrooted in the body. When we read or listen tothe description of an action, the process of sim-ulation that takes place in us is not the enact-ment of the same action; we would be echop-ractic if we were unable to avoid imitating andreproducing all the actions that we see or whosedescription we listen to or read. According toour hypothesis, embodied simulation rathermakes available to us an exemplary case, a

model, in relation to which understanding oflanguage is also realized. If therefore it is truethat the symbolic dimension opens up somepossibilities for us and creates worlds for usthat only linguistic creatures can enter, it is alsotrue that language strongly exploits mechanismsrooted in our corporeality. Enactment of thesimulation process in understanding languageseems to suggest that the symbolic dimensionand the bodily dimension cohabit in linguisticpraxis.

Nevertheless, the nature of this relation-ship is still not entirely clear, nor is the confinebetween the bodily dimension and the typicallyor exclusively symbolic dimension. Can it be hy-pothesized that corporeal knowledge also playsa role in understanding logical operators suchas, for instance, negation or disjunction, or thatit plays a role in understanding the interrogat-ive form? The whole symbolic nature of theselinguistic structures appears in some respectsbeyond question. Research on these issues isnow open (Kaup et al. 2006, 2007; Tettamantiet al. 2005; Christensen 2009; Tomasino et al.2010; Liuzza et al. 2011; Kumar et al. 2013) andtoday many wonder about the possibility ofidentifying mechanisms that can anchor suchstructures to our bodily experience. We takethis to be the real challenge for the embodiedcognition approach to the role played by lan-guage in human social cognition.

Let us once more return to the Aris-totelian notion of paradeigma and appraiseother possible hints for substantiating the ana-logy with the embodied simulation mechanism.The understanding that the rhetor calls forthrough reasoning based on the paradeigmashould lead the citizen to choose what is bestfor him in various circumstances. The goal ofsuch reasoning is to determine understanding ofa present situation, by analogy with a historicalexample or a fable, and, on the basis of thismore informed knowledge, to guide the humanbeing’s choices. In other words, the rhetoricalexample or paradeigma is knowledge whosemain goal is practical and not theoretical.

A practical aim also characterizes embod-ied simulation. Embodied simulation is alwaysaimed at “navigating” in the world and, there-

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 16 | 22

www.open-mind.net

fore, eventually at acting. It was hypothesizedthat embodied simulation allows us a direct, ex-periential way of understanding other people’sactions and experiences and, on the basis of thisunderstanding, it allows us to regulate our ac-tions and our experiences. These goals are al-ways practical. In some respects, the process ofembodied simulation that is enacted, for in-stance, when reading a novel (see Wojciehowski& Gallese 2011), also has a practical aim. Liter-ature recreates a world of emotions and experi-ences: the emotions and the experiences of theliterary characters inhabit the fictional world ofthe novel. The simulation mechanism helps usto “navigate” that world, even if it is a fictitiousworld; it allows us to understand and, in part,to relive the emotions of the protagonists andtheir vicissitudes. The aim in this case is prac-tical insofar as the simulation mechanism allowsus to approach the fictitious other with asecond-person epistemic perspective (Gallese2014).20

Embodied simulation makes implicitknowledge about others immediately available,with the aim of regulating our interactions withthem. Our understanding of the literary other isalmost always second-person, based on the pos-sibility of perceiving analogies between our ownexperiences and others’ and made possiblethrough a hypostatization of our experiencesthat is achieved through the simulation mechan-ism (Wojciehowski & Gallese 2011).

In the end, what is embodied simulation ifnot a suspension of the application of a process?Let us think of when mirror neurons are activatedin observing actions performed by others; or ofwhen canonical neurons are activated while weare looking at the keyboard of a computer think-ing about what we want to write; or when cor-tical motor neurons are activated when we ima-gine ourselves writing on that keyboard. These re-sponses on the part of motor neurons manifestthe activation of implicit knowledge, that is, bod-ily motor knowledge expressing the motor poten-tialities of the bodily self mapped by the motorsystem in terms of their motor outcomes.

20 A second-person perspective is adopted in social contexts when, im-plicitly or explicitly, we re-use our own experiences to understandothers. On the notion of second-person perspective see Pauen (2012).

Reuse of motor knowledge, in the absence ofthe movement that realizes it, as exemplified byembodied simulation, is an example of “paradig-matic knowledge.” Thus, embodied simulation is acase of implicit paradigmatic knowledge. Accord-ing to our hypothesis, embodied simulation allowsus to naturalize the notion of paradigm, anchor-ing it at a level of sub-personal description, whoseneural correlates we can study.

Our openness to the world is constitutedand made possible by a motor system predispos-ing and allowing us to adapt our daily and con-tingent pragmatic relationships with the worldagainst the background of a prefigured but highlyflexible plan of motor intentionality. Such a plan,intended as the sum of our motor potentialities,provides its coordination to any single contingentmodality of relation with the world, that is, toany single action we perform, in which it contin-ues to actualize itself. This aspect seems import-ant to us because it shows how specific aspects ofhuman social cognition are made possible andscaffolded upon processes not necessarily specificto humans, like embodied simulation.

8 Body and soul

It is improper, or at least it seems so to us, to saythat the soul, the spirit, or intelligence are em-bodied. If it were so, we would thus return to adualistic conception of human nature. Such dual-ism is always present in the tradition of Westernthought, though it is often disguised in formsthat, rightly or wrongly, are deemed politicallymore correct. Today nearly all cognitive scientistsdeclare themselves to be monists and physicalists.Nevertheless, the conception still dominant todayabout the cognitive structure of humans and theirfunctions draws a clear-cut and apparently un-solvable division between linguistic-cognitive pro-cesses and sensory-motor processes. It matterslittle that everybody admits that both are insome way referable to the biological physicality ofthe brain. The brain, according to classical cog-nitivism, is divorced from the body and conceivedof as a box of algorithmic wonders.

Classical cognitivism sees the body as anappendix of little or no interest for decoding thesupposed algorithms reportedly presiding over

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 17 | 22

www.open-mind.net

our cognitive life. Such is a cognitive life withvery little vitality, totally divorced not onlyfrom effectual reality, as we try to show here,but also from our daily phenomenal experience.It is not by chance that the language usuallyused to describe cognitive processes is borrowedfrom artificial intelligence: algorithms, informa-tion processing, etc.

Humans, however, cannot be assimilatedto information-processing entities. Even less ac-ceptable is the thesis that the concept of mean-ing is wholly assimilable to the concept of in-formation. Classical cognitivism has maintainedfor decades that intelligence depends on the al-gorithms that substantiate it and not on thematerial substrata on which the algorithmsthemselves are believed to be implemented.This is the so-called principle of the multiplerealizability of cognitive processes. EmbodiedSimulation and its relation to language and cog-nition casts severe doubts on this principle andadds arguments in support of the thesis thathuman cognition is tightly and necessarily de-pendent on the kind of body we have. As such,the mechanism of Embodied Simulation and therole it plays in human cognition provide furtherarguments in support of the idea that the prin-ciple of multiple realizability is false. We arewho we are because we evolved by adapting to aphysical world that obeys a series of physicallaws, such as that of gravity.

As the art historian Heinrich Wölfflin wrotein his 1886 Prolegomena zu einer Psychologie derArchitektur, if we were exclusively opticalcreatures, aesthetic judgment of the physicalworld would be precluded to us. Are theamazement and sense of elevation transmitted tous by the contemplation of a Gothic cathedralconceivable in purely algorithmic terms? Is it con-ceivable to divorce aesthetic experience from ourdaily muscular, tactile, and viscero-motor experi-ence of reality? Wölfflin (and together with himmany others, among them Merleau-Ponty) main-tained it was not, and we think that he was right.

We believe that our “natural” propensity todualism is, on the one side, the product of our be-ing asymmetrically positioned between mind andbody, as Helmuth Plessner (2006) maintained.We are corporeal beings, but at the same time we

maintain that we have a body. On the other side,the account of the historical result of the pro-gressive de-centralization of the anthropologicaldimension leads us to be dualist. We are nolonger the living image of God, we are no longerat the centre of the universe, and perhaps in post-modern times we are no longer even subjects orselves. What are we left with but with the claimof the total otherness and discontinuity of ourcognitive social life and its underlying processes?Their immaterial nature, or more exactly theirtotal otherness in relation to a corporeity whoseanimal origin or essence is—evolutionarily speak-ing—pretty much clear, is perhaps the only wayof reaffirming our uniqueness. The dualismbetween mind and body become, thus, a mechan-ism of defence. The so-called mental Rubicon thatseparates us from other non-human living beingsis a very powerful anti-depressive argument for adisorientated humanity.

At this point, however, a clarification is re-quired in order to avoid unpleasant misunder-standings. It is beyond doubt that the least intel-ligent among humans is incommensurably differ-ent and other in relation to the most intelligentamong chimpanzees, despite their almost com-plete sharing of a genetic endowment. The pointis that this quantum leap can be explained, per-haps, by remaining within an evolutionary frame-work that does not look for discontinuities foun-ded on theories of “cognitive catastrophes,” ge-netic big bangs (as in the case of the so-called“grammar gene” invoked by Pinker), and so forth.The mysterious uniqueness—and loneliness—ofhumankind in the universe proves more compre-hensible, or at least more easily approachable, ifempirically investigated after having set aside theanti-continuist and self-consoling recipes of classiccognitive science. In our continuist approach, hu-mankind is not special, because his evolution fol-lows the same laws that regulate evolution of allother animals and is in continuity with evolution-ary paths of other animals. However, our peculiarevolutionary path leaded us to acquire species-specific characteristics that only human beingsshare.

Sigmund Freud realized long before othershow much the self is a bodily self (1923). Freudalso helped us to understand how little we know

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 18 | 22

www.open-mind.net

about who we are, particularly when aspiring toground this knowledge solely on self-questioningrationality. What are the drives of which Freudspoke but a further manifestation of the doublestatus of our flesh? We are Körper (objectualand represented body) and Leib (lived body), asEdmund Husserl maintained. Today cognitiveneuroscience can shed new light on the Leib byinvestigating the Körper. The point is not to re-duce the Leib to the Körper, but to understandthat the empirical investigation of the Körpercan tell us new things about the Leib.

9 Provisional conclusions

In this paper we addressed and discussed thenotions of intersubjectivity and of the self as in-dissolubly intertwined outcomes of the bodilyand symbolic dimensions. We proposed that em-bodied simulation seems to be able to naturalizethe notion of paradigm, thus naturalizing one ofthe processes that makes language reflexivenesspossible, and thus contributing to “creating”the human being. Being a subject perhapsmeans being a body that learns to express itselfand to express its world thanks to the paradigm—embodied simulation—that allows one to gobeyond the body while remaining anchored toit. A new understanding of intersubjectivity canbenefit from a bottom-up study and character-ization of the non-declarative and non-metarep-resentational aspects of social cognition (seeGallese 2003a, 2007).

One key issue of the new approach to in-tersubjectivity we proposed here is the investig-ation of the neural bases of our capacity to beattuned to the intentional relations of others.At a basic level, our interpersonal interactionsdo not make explicit use of propositional atti-tudes. This basic level consists of embodied sim-ulation processes that enable the constitution ofa shared meaningful interpersonal space. Theshared intersubjective space in which we livefrom birth constitutes a substantial part of oursemantic space. Self and other relate to eachother because are opposite extensions of thesame correlative and reversible we-centric space(Gallese 2003a). Observer and observed are partof a dynamic system governed by reversible

rules. By means of intentional attunement, “theother” is much more than a different representa-tional system; it becomes a bodily self, like us.

This new epistemological approach to inter-subjectivity has the merit of generating predic-tions about the intrinsic functional nature of oursocial cognitive operations, cutting across, andnot being subordinated to a specific ontology ofmind, like that purported by the classic cognitiv-ist approach. Open questions that need to be fur-ther investigated in the future concern the biolo-gical mechanisms underlying our species-specificforms of self-knowledge and intersubjectivity.Language will have a special role in this investiga-tion. To what extent and how are symbolic opera-tions constrained by biological mechanisms? Isthis connection between symbolic representationsand bodily mechanisms that has been responsiblefor our specificity? These and other questions willbe object of investigation in the next years

Acknowledgements

This work was supported by the EU Grant To-wards an Embodied Science of InterSubjectivity(TESIS, FP7-PEOPLE-2010-ITN, 264828) andby the KOSMOS Fellowship from HumboldtUniversity, Berlin to VG.

The authors wish to thank Anna Strasser,Michael Pauen and two anonymous reviewersfor their most useful comments and criticismson an earlier version of this article.

Although both authors discussed and de-signed the article together, sections 2, 3, 4, 5,and 8 were written by Vittorio Gallese, whilesections 6, and 7 were written by ValentinaCuccio. Sections 1 and 9 were written jointly byboth authors.

References

Abe, K. & Watanabe, D. (2011). Songbirds possess thespontaneous ability to discriminate syntactic rules.Nature Neuroscience, 14 (8), 1067-1074.10.1038/nn.2869

Adrian, E. D. & Matthews, R. (1927a). The action of lighton the eye: Part I. The discharge of impulses in the opticnerve and its relation to the electric changes in the retina.The Journal of Physiology, 63 (4), 378-414.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 19 | 22

www.open-mind.net

(1927b). The action of light on the eye: Part II.The processes involved in retinal excitation. TheJournal of Physiology, 64 (3), 279-301.

Agamben, G. (2008). Signatura Rerum. Sul Metodo.Torino, IT: Bollati-Boringhieri.

Ammaniti, M. & Gallese, V. (2014). The birth of inter-subjectivity: Psychodynamics, neurobiology, and theself. New York, NY: Norton.

Anderson, M. L. (2010). Neural reuse: A fundamental re-organizing principle of the brain. Behavioral Brain Sci-ences, 33 (4), 245-266. 10.1017/S0140525X10000853

Aristotle, (2012). The art of rhetoric. London, UK:Harper Press.

Arzy, S., Overney, L. S., Landis, T. & Blanke, O. (2006).Neural mechanisms of embodiment: Asomatognosia dueto premotor cortex damage. Archives of Neurology, 63(7), 1022-1025.10.1001/archneur.63.7.1022

Barsalou, L. W. (2008). Grounded cognition. Annual Re-view of Psychology (59), 617-645. 10.1146/annurev.psych.59.103006.093639

Bermúdez, J. L. (2003). Thinking without words. NewYork, NY: Oxford University Press.

Berti, A., Bottini, G., Gandola, M., Pia, L., Smania, N.,Stracciari, A., Castiglioni, I., Vallar, G. & Paulesu, E.(2005). Shared cortical anatomy for motor awarenessand motor control. Science, 309 (5733), 488-491.10.1126/science.1110625

Bloomfield, T. C., Gentner, T. Q. & Margoliash, D.(2011). What birds have to say about language. NatureNeuroscience, 14 (8), 947-948. 10.1038/nn.2884

Boulenger, V., Shtyrov, Y. & Pulvermüller, F. (2012).When do you grasp the idea? MEG evidence for in-stantaneous idiom understanding. NeuroImage, 59 (4),3502-3513. 10.1016/j.neuroimage

Butterfill, S. A. & Sinigaglia, C. (2014). Intention andmotor representation in purposive action. Philosophyand Phenomenological Research, 88 (1), 119-145.10.1111/j.1933-1592.2012.00604.x

Cermolacce, M., Naudin, J. & Parnas, J. (2007). The“minimal self” in psychopathology: Re-examining theself-disorders in the schizophrenia spectrum. Con-sciousness and Cognition, 16 (3), 703-714. 10.1016/j.concog.2007.05.013

Chittka, L. & Niven, J. (2009). Are bigger brains better? Cur-rent Biology, 19 (21), R995-R1008. 10.1016/j.cub.2009.08.023.

Chomsky, N. (1966). Cartesian linguistics: A chapter inthe history of rationalist thought. Lanham, MD: Uni-versity Press of America.

Christensen, K. R. (2009). Negative and affirmative sen-tences increase activation indifferent areas in the brain.Journal of Neurolinguistics, 22 (1), 1-17.10.1016/j.jneuroling.2008.05.001

Condillaco, E. B. (2001). Essay on the origin of humanknowledge. Cambridge, UK: Cambridge University Press.

Cosmides, L. & Tooby, J. (1997). The multimodularnature of human intelligence. In A. Schiebel & J. W.Schopf (Eds.) (p. Origin and Evolution of Intelligence).Toronto, Canada: Jones and Bartlett Publishers.

Culioli, A. (1968). La formalisation en linguistique.Cahiers Pour l’Analyse, 9, 106-117.

Dehaene, S. (2005). Evolution of human cortical circuitsfor reading and arithmetic: The neuronal recycling hy-pothesis. In S. Dehaene, J.-R. Duhamel, M. D. Hauser& G. Rizzolatti (Eds.) From Monkey Brain to HumanBrain. A Fyssen Foundation Symposium (pp. 133-157).Cambridge, MA: MIT Press.

Descartes, R. (1637). Discours de la methode pour bien con-duire sa raison, & chercher la verité dans les sciences:plus la dioptrique, les meteores, et la geometrie, qui sontdes essais de cette methode. Leiden: Jan Maire.

(1642). Meditationes de prima philosophia, inquibus Dei existentia & animae humanae a corpore dis-tinctio demonstrantur: his adjunctae sunt variae objec-tiones doctorum virorum in istas de Deo & animademonstrationes, cum responsionibus authoris. 2nd edi-tion. Amsterdam, NL: Elzevir.

Ehrsson, H. H., Spence, C. & Passingham, R. E. (2004).That’s my hand! Activity in premotor cortex reflectsfeeling of ownership of a limb. Science, 305 (5635), 875-877.10.1126/science.1097011

Ferri, F., Frassinetti, F., Costantini, M. & Gallese, V.(2011). Motor simulation and the bodily self. PLoSOne, 6 (3), e17927. 10.1371/journal.pone.0017927

Ferri, F., Frassinetti, F., Ardizzi, M., Costantini, M. &Gallese, V. (2012). A sensorimotor network for thebodily self. Journal of Cognitive Neuroscience, 24 (7),1584-1595. 10.1162/jocn_a_00230

Firpo, L. (1940). Bibliografia degli scritti di TommasoCampanella. Turin, I: V. Bona.

Freud, S. (1923). The ego and the I. London, UK: TheHogarth Press Ltd.

Gallagher, S. (2005). How the body shapes the mind. Ox-ford, UK: Clarendon Press.

Gallese, V. (2000). The inner sense of action: Agency andmotor representations. Journal of Consciousness Stud-ies, 7 (10), 23-40.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 20 | 22

www.open-mind.net

(2003a). The manifold nature of interpersonal re-lations: The quest for a common mechanism. Philo-sophical Transactions of the Royal Society of LondonB, 358 (1431), 517-528. 10.1098/rstb.2002.1234

(2003b). A neuroscientific grasp of concepts: Fromcontrol to representation. Philophical Transaction ofthe Royal Society of London B, 358 (1435), 1231-1240.10.1098/rstb.2003.1315

(2005). Embodied simulation: From neurons tophenomenal experience. Phenomenology and the Cog-nitive Sciences, 4 (1), 23-48. 10.1007/s11097-005-4737-z

(2006). Intentional attunement: A neurophysiolo-gical perspective on social cognition and its disruptionin autism. Cognitive Brain Research, 1079 (1), 15-24.10.1016/j.brainres.2006.01.054

(2007). Before and below theory of mind: Embod-ied simulation and the neural correlates of social cogni-tion. Philosophical Transactions of the Royal Society ofLondon B, 362 (1480), 659-669. 10.1098/rstb.2011.0029

(2008). Mirror neurons and the social nature oflanguage: The neural exploitation hypothesis. SocialNeuroscience, 3 (3-4), 317-333.10.1080/17470910701563608

(2009). Motor abstraction: A neuroscientific ac-count of how action goals and intentions are mappedand understood. Psychological Research, 73 (4), 486-498. 10.1007/s00426-009-0232-4

(2011). Neuroscience and phenomenology. Phe-nomenology & Mind, 1, 33-48.

(2013). Corpo non mente. Le neuroscienze cognit-ive e la genesi di soggettività ed intersoggettività. Edu-cazione Sentimentale, 20, 8-24. 10.3280/EDS2013-020002

(2014). Bodily selves in relation: Embodied simu-lation as second-person perspective on intersubjectivity.Philosophical Transaction of the Royal Society B, 369(1644), 20130177-20130177. 10.1098/rstb.2013.0177

Gallese, V., Fadiga, L., Fogassi, L. & Rizzolatti, G.(1996). Action recognition in the premotor cortex.Brain, 119 (2), 593-609. 10.1093/brain/119.2.593

Gallese, V., Keysers, C. & Rizzolatti, G. (2004). A unify-ing view of the basis of social cognition. Trends inCognitive Sciences, 8 (9), 396-403.10.1016/j.tics.2004.07.002

Gallese, V. & Lakoff, G. (2005). The brain’s concepts:The role of the sensory-motor system in reason andlanguage. Cognitive Neuropsychology, 22 (3), 455-479.10.1080/02643290442000310

Gallese, V. & Sinigaglia, C. (2010). The bodily self aspower for action. Neuropsychologia, 48 (3), 746-755.10.1016/j.neuropsychologia.2009.09.038

(2011a). How the body in action shapes the self.Journal of Consciousness Studies, 18 (7-8), 117-143.

(2011b). What is so special with embodied simula-tion. Trends in Cognitive Sciences, 15 (11), 512-519.10.1016/j.tics.2011.09.003

Gentner, T. Q., Fenn, K. M., Margoliash, D. & Nusbaum,H. C. (2006). Recursive synctactic pattern learning bysongbirds. Nature, 440, 1204-1207.10.1038/nature04675

Glenberg, A. M. & Gallese, V. (2012). Action-based lan-guage: A theory of language acquisition, comprehen-sion, and production. Cortex, 48 (7), 905-922.10.1016/j.cortex.2011.04.010

Gould, S. J. & Lewontin, R. C. (1979). The spandrels ofSan Marco and the Panglossian paradigm. A critique ofthe adoptionist programme. Proceedings of the RoyalSociety of London, 205 (1161), 281-288.10.1098/rspb.1979.0086

Gross, C. G. (2002). Genealogy of the ‘GrandmotherCell’. Neuroscientist, 8 (5), 512-518.10.1177/107385802237175

Gross, C. G., Rocha-Miranda, C. E. & Bender, D. B. (1972).Visual properties of neurons in inferotemporal cortex of themacaque. Journal of Neurophysiology, 35 (1), 96-111.

Guan, C. Q., Meng, W., Yao, R. & Glenberg, A. M.(2013). The motor system contributes to comprehen-sion of abstract language. PLoS One, 8 (9), e75183.10.1371/journal.pone.0075183

Hauser, M. D., Chomsky, N. & Fitch, W. T. (2002). Thefaculty of language: What is it, who has it, and howdid it evolve? Science, 298 (5598), 1569-1579.10.1126/science.298.5598.1569

Hinzen, W. & Sheehan, M. (2013). The philosophy of uni-versal grammar. Oxford, UK: Oxford University Press.

Husserl, E. (1973). Cartesianische Meditationen und Par-iser Vorträge. In S. Strasser (Ed.) Den Haag, NL: Mar-tinus Nijhoff.

Kaup, B., Yaxley, R. H., Madden, C. J., Zwaan, R. A. &Lüdtke, J. (2006). Experiential simulations of negatedtext information. Journal of Experimental Psychology,60 (7), 976-990. 10.1080/17470210600823512

Kaup, B., Lüdtke, J. & Zwaan, R. A. (2007). Processingnegated sentences with contradictory predicates: Is adoor that is not open mentally closed? Journal ofPragmatics, 38, 1033-1050.10.1016/j.pragma.2005.09.012

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 21 | 22

www.open-mind.net

Ku, S. P., Logothetis, N. K., Tolias, A. S. & Goense, J.(2011). fMRI of the face-processing network in theventral temporal lobe of awake and anesthetizedmacaques. Neuron, 70 (2), 352-362.10.1016/j.neuron.2011.02.048.

Kumar, U., Padakannaya, P., Mishra, R. K. & Khetrapal,C. L. (2013). Distinctive neural signatures for negativesentences in Hindi: An fMRI study. Brain Imaging andBehaviour, 7 (2), 91-101. 10.1007/s11682-012-9198-8

Legrenzi, P. & Umiltà, C. A. (2011). Neuromania: On the lim-its of brain science. Oxford, UK: Oxford University Press.

Leslie, A. M. (2005). Developmental parallels in under-standing minds and bodies. Trends in Cognitive Sci-ence, 9 (10), 459-462. 10.1016/j.tics.2005.08.002

Liuzza, M. T., Candidi, M. & Aglioti, S. M. (2011). Do notresonate with actions: Sentence polarity modulates cortico-spinal excitability during action-related sentence reading.PLoSONE, 6, e16855. 10.1371/journal.pone.0016855

Lo Piparo, F. (2003). Aristotele e il linguaggio. Roma, IT:Laterza.

Margoliash, D. & Nusbaum, H. C. (2009). Language. Theperspective from organismal biology. Trends in Cognit-ive Sciences, 13 (12), 505-510.10.1016/j.tics.2009.10.003

Pauen, M. (2012). The second-person perspective. In-quiry, 55 (1), 33-49. 10.1080/0020174X.2012.643623

Piazza, F. (2008). La retorica di Aristotele. Introduzionealla lettura. Roma, IT: Carocci.

Pinker, S. (1994). The language instinct. New York, NY:Harper Collins.

(1997). How the mind works. New York, NY:Norton.

Plessner, H. (2006). I gradi dell’organico e l’uomo.Torino, IT: Bollati-Boringhieri.

Pulvermüller, F. (2013). How neurons make meaning: Brainmechanisms for embodied and abstract-symbolic se-mantics. Trends in Cognitive Science, 458 (470), 1-17.10.1016/j.tics.2013.06.004

Quiroga, R., Reddy, L., Kreiman, G., Koch, C. & Fried, I.(2005). Invariant visual representation by single neur-ons in the human brain. Nature, 435, 1102-1107.10.1038/nature03687

Rizzolatti, G., Fadiga, L., Gallese, V. & Fogassi, L.(1996). Premotor cortex and the recognition of motoractions. Cognitive Brain Research, 3 (2), 131-141.

Rizzolatti, G. & Sinigaglia, C. (2010). The functional roleof the parieto-frontal mirror circuit: Interpretationsand misinterpretations. Nature Reviews Neuroscience,11 (4), 264-274. 10.1038/nrn2805

Ross, W. D. (Ed.) (1978). Aristotles prior and posterioranalytics. Oxford, UK: Clarendon Press.

Saxe, R. (2006). Uniquely human social cognition. Cur-rent Opinion in Neurobiology, 16 (2), 235-249.10.1016/j.conb.2006.03.001

Tettamanti, M., Buccino, G., Saccuman, M. C., Gallese,V., Danna, M., Scifo, P., Fazio, F., Rizzolatti, G.,Cappa, S. F. & Perani, D. (2005). Listening to action-related sentences activates fronto-parietal motor cir-cuits. Journal of Cognitive Neuroscience, 7 (2), 273-281. 10.1162/0898929053124965

Tomasino, B., Weiss, P. H. & Fink, G. R. (2010). Tomove or not to move: Imperatives modulate action-re-lated verb processing in the motor system. Neuros-cience, 168, 246-258.10.1016/j.neuroscience.2010.04.039

Usener, H. (1887). Epicurea. Leipzig: Teubner. Italiantranslation by Ilaria Ramelli, Epicurea: Testi diEpicuro e testimonianze epicuree nell’edizione di Her-mann Usener. Milan, IT: Bompiani, 2002.

Varela, F. J. & Shear, J. (Eds.) (1999). The view fromwithin: First-person approaches to the study of con-sciousness. Bowling Green, OH: Imprint Academic.

Virno, P. (2003). Quando il verbo si fa carne. Linguaggioe natura umana. Torino, IT: Bollati Boringhieri.

(2011). E così via all’infinito. Logica e antropolo-gia. Torino, IT: Bollati Boringhieri.

Vogeley, K., May, M., Ritzl, A., Falkai, P., Zilles, K. &Fink, G. R. (2004). Neural correlates of first-personperspective as one constituent of human self-conscious-ness. Journal of Cognitive Neuroscience, 16 (5), 817-827. 10.1162/089892904970799

Vogeley, K. & Fink, G. R. (2003). Neural correlates of thefirst-person-perspective. Trends in Cognitive Sciences,7 (1), 38-42.

Wojciehowski, H. C. & Gallese, V. (2011). How storiesmake us feel. Toward an embodied narratology. Cali-fornia Italian Studies, 2 (1).

Wölfflin, H. (1886). Prolegomena zu einer Psychologie derArchitektur. München, GER: University of Munich.

Zeki, S. (1993). A vision of the brain. London, UK:Wiley-Blackwell.

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body - Embodied Simulation, Intersubjectivity, the Bodily Self, and Language.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(T). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570269 22 | 22

Multisensory Spatial Mechanisms of the Bodily Self and Social CognitionA Commentary on Vittorio Gallese & Valentina Cuccio

Christian Pfeiffer

This commentary aims to find the right description of the pre-reflective brainmechanisms underlying our phenomenal experience of being a subject bound to aphysical body (bodily self) and basic cognitive, perceptual, and subjective aspectsrelated to interaction with other individuals (social cognition). I will focus on theproposal by Gallese and Cuccio that embodied simulation, in terms of motor res-onance, is the primary brain mechanism underlying the pre-reflective aspects ofsocial cognition and the bodily self. I will argue that this proposal is too narrowto serve a unified theory of the neurobiological mechanisms of both target phe-nomena. I support this criticism with theoretical considerations and empirical evid-ence suggesting that multisensory spatial processing, which is distinct from but apre-requisite of motor resonance, substantially contributes to the bodily self andsocial cognition.

My commentary is structured in three sections. The first section addressessocial cognition and compares embodied simulation to an alternative account,namely the attention schema theory. According to this theory we pre-reflectivelyempathize with others by predicting their current state of attention which involvespredicting the spatial focus of attention. Thereby we derive a representationalmodel of their state of mind. On this account, spatial coding of attention, ratherthan motor resonance, is the primary mechanism underlying social cognition. Itake this as a theoretical alternative complementing motor resonance mechanisms.

The second section focuses on the bodily self. Comparison of the brain net-works of the bodily self and social cognition reveals strong overlap, suggestingthat both phenomena depend on shared multisensory and sensorimotor mechan-isms. I will review recent empirical data about altered states of the bodily self interms of self-location and the first-person perspective. These spatial aspects ofthe bodily self are encoded in brain regions distinct from the brain network of em-bodied simulation. I argue that while motor resonance might contribute to bodyownership and agency, it does not account for spatial aspects of the bodily self.Thus, embodied simulation appears to be a necessary but insufficiently “primary”brain mechanism of the bodily self and social cognition.

The third section discusses the contributions of the vestibular system, i.e.,the sensory system encoding head motion and gravity, to the bodily self and so-cial cognition. Vestibular cortical processing seems relevant to both processes, be-cause it directly encodes the world-centered direction of gravity and allows us todistinguish between motions of the own body and motions of other individuals andthe external world. Furthermore, the vestibular cortical network largely overlapswith those neural networks relevant to the bodily self and social cognition. Thus,the vestibular system may play a crucial role in multisensory spatial coding relat-ing the bodily self to other individuals in the external world.

KeywordsAttention schema | Bodily self | Embodied simulation | First-person perspective |Mirror neurons | Self-location | Social cognition | Vestibular system

Commentator

Christian [email protected]   Ecole Polytechnique FédéraleLausanne, Switzerland

Target Authors

Vittorio [email protected]   Università degli Studi di ParmaParma, Italy

Valentina [email protected]   Università degli Studi di PalermoPalermo, Italy

Editors

Thomas [email protected]   Johannes Gutenberg-UniversitätMainz, Germany

Jennifer M. [email protected]   Monash UniversityMelbourne, Australia

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese & Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 1 | 14

www.open-mind.net

1 Introduction

The paper by Gallese and Cuccio provides anintegrated theoretical framework explaining howthe brain and body relate to social cognition,the human self, and language. The authors re-view empirical evidence from electrophysiolo-gical and neuroimaging studies supporting em-bodied simulation (ES) theory (Gallese & Cuc-cio this collection, p. 8). According to ES, thebrain covertly simulates the bodily actions, per-ceptions, and emotions observed in other indi-viduals by using parts of our neural architectureinvolved in acting, sensing, and feeling emo-tions. Thereby, we infer the goals, intentions,and states of mind of others in a pre-reflectiveand non-conceptual fashion. But the authorstake this a step further and propose that ES isthe key mechanism underlying, and hence unify-ing, both social cognition, the human self, andlanguage. Throughout the paper, the authorsemphasize the tight functional coupling betweenthe body and the brain, which when taken intoaccount bears the potential to significantly ad-vance the scientific study of the hard problem ofconsciousness (Chalmers 1996).

This commentary on Gallese and Cuccioaims to find the right description of the brainmechanisms underlying pre-reflective aspects ofboth the bodily self and social cognition. Spe-cifically, I will focus on Gallese and Cuccio’scentral claim that ES, based on motor reson-ance and neural processing in the motor system,is the primary brain mechanism underlying pre-reflective representations of the bodily self andsocial cognition (Gallese & Cuccio this collec-tion, pp. 8–14). I ask the following questions:Could there be an alternative theory or empir-ical evidence countering the claim of a primacyof motor resonance underlying social cognitionand the bodily self? Which brain mechanisms inaddition to motor resonance might contribute topre-reflective aspects of social cognition and thebodily self? I will defend the following threetheses:

(1) Social cognition and the bodily self de-pend on multisensory spatial coding,which is distinct from motor resonance.

Thus, motor resonance may be a necessarybut insufficiently “primary” brain mechan-ism of social cognition and the bodily self(cf. section 1, 2).

(2) The brain networks underlying socialcognition and the bodily self largely over-lap. Specific functional associations exist(a) between motor resonance and bodyownership/agency and (b) between multis-ensory spatial coding and self-location/thefirst-person perspective (cf. section 2).

(3) The vestibular system, i.e., the sensorysystem encoding head motion and gravity,might provide unique information used formultisensory spatial coding that relatesthe bodily self to other individuals and theexternal world. This is further suggestedby the large overlap existing between thehuman vestibular cortex and the brainnetworks underlying the bodily self andsocial cognition (cf. section 3).

My commentary is structured in three sec-tions. In the first section I shall compare ES toan alternative theory of social cognition that as-signs priority to spatial coding of attention,rather than to motor resonance. I shall showthat both theories bear the potential that theirproposed brain mechanisms cooperatively worktogether in order to support social cognition.The second section addresses the bodily self. Ishall review data from neurological patients andfull-body illusion experiments, which highlightthe importance of two spatial aspects of thebodily self not mentioned by Gallese and Cuc-cio, i.e., self-location and the first-person per-spective. These spatial aspects of the bodily selfdepend primarily on multisensory integrationand on cortical processing outside regions in-volved in ES. Furthermore, comparisonsbetween the brain networks encoding the bodilyself and social cognition show large overlaps,suggesting shared functional mechanisms. In thethird section I propose that because multisens-ory spatial processing appears to be critical for

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 2 | 14

www.open-mind.net

the bodily self and social cognition, importantcontributions may come from the vestibular sys-tem (Lenggenhager & Lopez this collection). Ishall show that the vestibular cortical networklargely overlaps with the brain networks under-lying the bodily self and social cognition. I shalldiscuss potential contributions of vestibular cor-tical processing to these target phenomena andsuggest directions for future research.

2 Is social cognition based on motor resonance or attention tracking?

Social cognition refers to cognitive processes,perceptions, and subjective experiences relatedto interaction with conspecifics. This sectionasks: Which are the brain mechanisms underly-ing pre-reflective aspects of social cognition?Could there be alternative theories and empir-ical evidence countering the primary role of mo-tor resonance?

Gallese and Cuccio propose that socialcognition mainly depends on ES based on motorresonance and processing of mirror neurons (seecitations in Gallese & Cuccio this collection).Mirror neurons were initially discovered infronto-parietal networks of the macaque monkeybrain. They are a specific type of canonicalneuron involved in planning and executing handactions and were found to be activated bothwhen the monkey executed a specific graspingor reaching action and when the monkey pass-ively observed somebody performing similar ac-tions (Gallese et al. 1996; Rizzolatti et al. 1996).Neuroimaging studies in humans also showedmirror neuron-like activation patterns at thelevel of populations of neurons in distinct brainregions—mainly the ventral premotor cortex(vPM), the intraparietal sulcus (IPS), but alsothe insula cortex and the secondary somato-sensory cortex (Rizzolatti & Sinigaglia 2010; seealso figure 1a gray dots). ES proposes thatbased on mirror neurons the brain maps ob-served actions into an action space, into motorpotentialities, within our hierarchically-organ-ized motor system, and thereby infers and pre-dicts the action goals of the individual. In thisway it penetrates the state of mind of the other,and thus links self and other in a pre-reflective

empathical fashion (Gallese & Cuccio this col-lection, p. 7).

I would like to point out that motor reson-ance, i.e., the mapping of observed actions intomotor potentialities, necessarily depends onmultisensory spatial coding. I argue that this isthe case because of five points: First, the brainhas access to the physical world only throughthe different sensory receptors of the body thatbombard it with exteroceptive (e.g., vision, au-dition), proprioceptive (somatosensory, vestibu-lar), and interoceptive (somatosensory, visceral)signals. Second, these multisensory signals mustbe integrated according to their spatial andtemporal parameters (Stein & Stanford 2008) toinform neural representations of the states ofthe body and of the world around us—includingthe agents whose actions are subject to motorresonance. Third, the observed movements ofthese agents are coded in coordinates distinctfrom the egocentric spatial frame of referenceupon which our motor system operates. Fourth,the brain must necessarily perform spatialtransformations of the observed movements bythe other agent into the egocentric frame of ref-erence, upon which motor resonance can oper-ate. In sum, multisensory spatial coding is apre-requisite of motor resonance.

According to Gallese and Cuccio, the out-comes of such multisensory spatial coding arereadily available to the brain network of ESthrough anatomical connections to the vPMthat are “anatomically connected to visual andsomatosensory areas in the posterior parietalcortex and to frontal motor areas” (Gallese &Cuccio this collection, p. 10). However, it seemsthat the multisensory spatial coding requiredfor a precise description of complex motor actsmight be computationally costly. Might there bea computationally more effective alternative bywhich multisensory spatial coding is used to de-code the intentions of observed agents?

The attention schema (AS) theory ofawareness (Graziano 2013; Graziano & Kastner2011) proposes that brain mechanisms relatedto attention and spatial coding, which are dis-tinct from neural processing relevant to ES,primarily underlie pre-reflective aspects of socialcognition. Graziano and Kastner define atten-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 3 | 14

www.open-mind.net

tion as an information-handling mechanism ofthe brain that serves to give priority to some in-formation (e.g., representational features) out ofseveral equally probable alternatives that are inconstant competition for awareness. Further-more, awareness is defined as the process ofconsciously experiencing something, it is theprocess of relating the subject (i.e., a phenom-enal self, see also Metzinger 2003) to the ob-ject/content of experience. Graziano and Kast-ner summarize AS as follows:

[Awareness is information and] dependson some system in the brain that musthave computed [it] […]; otherwise, the in-formation would be unavailable for re-port. […] People routinely compute thestate of awareness of other people [and]the awareness we attribute to anotherperson is our reconstruction of that per-son’s attention. […] The same machinerythat computes socially relevant informa-tion […] also computes […] informationabout our own awareness. […] Awarenessis […] a perceptual model […] a rich in-formational model that includes, amongother computed properties, a spatialstructure. […] Through the use of the so-cial perceptual machinery, we assign theproperty of awareness to a locationwithin ourselves. (Graziano & Kastner2011, pp. 98–99)

Related to social cognition, AS proposes that byusing a schematic representation of the state ofattention of other individuals—including a pre-diction of the spatial location of their focus ofattention—we predict the current state ofawareness of the individual, which is informat-ive about their intentions and potential futureactions. In short: Awareness of others is an at-tention schema. As compared to ES, AS is a rel-atively recent theory that requires extensive em-pirical studies. Yet the evidence so far showsthat indeed the brain has a neural circuitry formonitoring the spatial configuration of one’sown attention independent of the sensory mod-ality (Downar et al. 2000), including the direc-tion of gaze (Beck & Kastner 2009; Desimone &

Duncan 1995). These structures are the pro-posed neural expert system upon which AS isbased and consist of the right-hemispheric tem-poro-parietal junction (TPJ) and superior tem-poral sulcus (STS) (see figure 1a in black). Not-ably, this expert system relevant to AS showslittle anatomical overlap with the neural struc-tures relevant to ES (figure 1a compare blackwith gray).

Because the AS relies on coding of thespatial relationship between the location ofthe observed individual and the likely spatiallocation of this individual’s attention (i.e., in-dependent of a particular sensory modality),the required spatial computations seem simpleand straightforward. They require two points,i.e., the individual as a reference point andthe potential spatial location of the attentionof that individual. According to AS, usingsuch spatial labeling the brain is able to sim-ultaneously track the aware and attendingminds of several individuals simultaneously.Thus, spatial coding in the context of AS ap-pears to be less complex and less computa-tionally demanding than spatial transforma-tions underlying ES (see above).

Which of these seemingly distinct brainmechanisms proposed by AS and ES moreplausibly underlies social cognition: the neuralexpert system decoding the state of attentionaccording to AS or the mirror mechanism sys-tem decoding observed motor plans accordingto ES? It has been proposed that AS and ESmay in principle work together. Graziano andKastner propose that the expert system of ASmay take a leading role by formulating a hy-pothesis about the state of awareness of an in-dividual that is likely to drive further beha-vior and therefore provide a set of predictionsbased upon which motor resonance could moreefficiently perform simulations (Graziano &Kastner 2011). Motor resonance would thusadd richer detail to the state-of-attention hy-pothesis made by the expert system.

This combined mechanism is compatiblewith the predictive processing principle (Clarkthis collection; Hohwy 2013, this collection),which has been proposed relevant to the bod-ily self (Apps & Tsakiris 2013; Limanowski &

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 4 | 14

www.open-mind.net

Blankenburg 2013; Seth this collection). Ac-cording to predictive processing the brain con-stantly predicts the potential causes of sens-ory input by minimizing prediction errors viaupdate of the predicted causes or by actionthat changes sensory input (Friston 2005).Applying the predictive processing principle toGraziano and Kastner’s proposal that AS is ahypothesis-generating tool to which ES addsfurther detail, one could conceive of bothmechanisms as different predictive processingmodules aimed at anticipating the state ofawareness and of intentional actions observedin others. Although no empirical study so farhas addressed this specific hypothesis, a re-cent functional magnetic resonance imagingstudy found that predictive processing prin-ciples accounted for the blood oxygen-level de-pendent activity related to the perception offaces, which is an important perceptual func-tion for social cognition in the human species(Apps & Tsakiris 2013).

These common and distinct predictionsbased on ES, AS, and predictive processing

call for empirical research aimed at providingevidence to further refine, integrate, or rejectthem.

3 Multisensory and motor mechanisms ofthe multifaceted bodily self

The bodily self refers to the phenomenal experi-ence of being an experiencing subject (i.e., aphenomenal self) bound to a physical body,which gives rise to the dual nature of the body(Husserl 1950; Gallese & Cuccio this collection,p. 2). The unified experience of being a bodilyself can be decomposed into different aspects,including the experience that we identify with aparticular body (self-identification or body own-ership), the experience that the self is situatedin a specific spatial location (self-location), thatwe take a specific experiential perspective at theworld (first-person perspective), and that we arethe authors of our actions, including havingcontrol of attentional focus (agency; (Blanke2012; Ehrsson 2012; Jeannerod 2003; Metzinger2003).

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 5 | 14

Figure 1: Summary of cortical brain regions involved in social cognition, the bodily self, and vestibular processing. (a)Whereas for social cognition there is little overlap between the brain regions proposed relevant for the attention schema(in black) and embodied simulation (in gray), both sets of brain regions overlap with (b) the brain network of the bod-ily self as identified by full-body illusion experiments manipulating self-location and first-person perspective (in black)and the body-swap illusion manipulating mainly body ownership (in gray). (c) The human vestibular cortical regions(in black) are widely distributed and overlap with several regions relevant to both the bodily self and social cognition.(The images are derived from images by NASA, licensed under creative commons.)

www.open-mind.net

In their paper, Gallese & Cuccio highlightthe relevance of mirror mechanisms, in particu-lar related to processing in the cortical motorsystem, to the sense of body ownership and thesense of agency, in particular in the context ofaction and action observation:

This minimal notion of the self, namelythe bodily self as power-for-action […], ta-citly presupposes ownership of an action-capable agentive entity; hence, it primarilyrests upon the functionality of the motorsystem. (this collection, p. 10)

However, recent philosophy of mind and cognit-ive neuroscience research reveals the crucial roleof spatial aspects of the bodily self, consisting ofa first-person perspective and self-location. Inthis section I shall compare the brain networkcontributing to spatial aspects of the bodily selfwith the brain network underlying body owner-ship and ask: Do these neuroimaging resultssupport the proposal that motor resonance is aprimary mechanism underlying all aspects ofthe bodily self? What is the relationshipbetween the neural networks of the bodily selfand social cognition? Which functional associ-ations can be derived from this?

3.1 Brain mechanisms of spatial aspects of the bodily self

The phenomenal experience of being a subjectis associated with a spatial location, which typ-ically is the space of the physical body (see alsoAlsmith & Longo 2014; Limanowski & Hecht2011). However, there are exceptions to theseprototypical states of the bodily self in neurolo-gical disorders and experimental illusions point-ing to a specific set of brain regions involved inspatially linking the phenomenal self to thephysical body.

Which brain mechanisms link the phenom-enal self to the physical body to give rise to thedual nature of the body as lived body and asphysical object? Research in neurological pa-tients who have had out-of-body experiences(OBE) shows that damage or interference withthe right TPJ can lead to dissociations between

the bodily self and physical body (Blanke et al.2004; Blanke et al. 2002; De Ridder et al. 2007;Ionta et al. 2011). During an OBE, patientstypically experience a disembodied self-locationin elevation above their physical body, and analtered first-person perspective that originatesfrom an elevated location in the room and isdirected downwards at the physical body(Blanke et al. 2004; Metzinger 2009). These pa-tients do not identify with their physical bodybut with an illusory double outside of the bor-ders of the physical body. At the phenomenolo-gical level, self-location and the first-person per-spective are often experienced as having theirspatial origin in the same position. However,during OBE there are instances where self-loca-tion can be dissociated from the first-personperspective in different sensory modalities (DeRidder et al. 2007). Further evidence from aso-matic OBEs and bodiless dreams suggests thata phenomenal first-person perspective may bereducible to a single point in space (Windt2010). In fact, vestibular hallucinations system-atically preceded OBEs in patients with sleepparalysis, i.e., a motor paralysis characterisedby the transient inability to execute bodily ac-tions when waking up from sleep (Cheyne &Girard 2009), showing further dissociations ofthe spatial location of the bodily self and thephysical body and links to sensory processing.These studies seem to suggest that the first-per-son perspective and self-location may depend ondifferent neural mechanisms (Blanke 2012).

OBE in epileptic patients can be inducedby subcortical electrical stimulation of a specificintensity at the TPJ. However, stimulating thesame brain region with either lower or higherstimulation intensity induces bodily sensations(including vestibular, visual, somatosensory,kinesthetic sensations) without inducing anOBE (Blanke et al. 2002). These observationsgave rise to the idea that the spatial aspects ofthe bodily self are based on the accurate integ-ration of multisensory signals (i.e., which wasperturbed by electrical stimulation in the pa-tient in Blanke et al. 2002, which are sensorysignals from personal space to sensory signalsfrom the external environment Blanke et al.2004).

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 6 | 14

www.open-mind.net

These clinical observations in patientswere corroborated by different full-body illusionexperiments in healthy subjects, such as the so-called “body-swap illusion” (Petkova & Ehrsson2008; Petkova et al. 2011; van der Hoort et al.2011), the “full-body illusion” (Ionta et al. 2011;Lenggenhager et al. 2009; Lenggenhager 2007;Pfeiffer et al. 2013; Pfeiffer, Schmutz & Blanke2014), and the “out-of-body illusion” (Ehrsson2007; Guterstam & Ehrsson 2012). In these ex-periments, healthy subjects receive conflictingsignals about the spatial location of their bodyand of the temporal synchrony of exteroceptiveand interoceptive signals, including somato-sensory, cardiac, and vestibular signals that atthe same time are applied to a virtual or fakebody seen by the subject (Aspell et al. 2013;Ionta et al. 2011; Pfeiffer et al. 2013; Pfeiffer etal. 2014). For example, in the full-body illusion,synchronous stroking of a virtual or fake bodyseen from a distance can induce the feeling inparticipants that they are more closely locatedto the position of the virtual or fake body, andthat they experience and increase of ownershipfor the seen body. The brain regions involved inthese spatial experimental manipulations of theexperienced bodily self most consistently involvethe right TPJ region, but also draw on somato-sensory and visual regions that process the sens-ory inputs (Blanke 2012; Ionta et al. 2011; fig-ure 1b in black). Recently, several studies havemanipulated visual and vestibular signals aboutthe direction of gravity, affecting self-locationand perspective and thus showing that thosevisual spatial cues affect our subjective experi-ence of the first-person perspective (Ionta et al.2011; Pfeiffer et al. 2013). These authors presen-ted images on virtual-reality goggles showingvisual gravitational cues, similar to the visualperspective during an OBE showing a scenefrom an elevated spatial location and a visualviewpoint directed downwards into the room.At the same time the somatosensory and thevestibular signals received by the participant,who was lying on the back, suggested that thephysical body was oriented upwards with re-spect to veridical gravity. Thus the visual grav-ity cues (i.e., downwards) and the vestibulargravity cues (i.e., upwards) were in directional

conflict. When the full-body illusion was in-duced under these conflicting conditions, parti-cipants reported subjective changes in their ex-perienced direction of the first-person perspect-ive (upward or downward) in line with experi-mentally-induced multisensory conflict (Ionta etal. 2011; Pfeiffer et al. 2013).

3.2 Brain mechanisms of body ownership

A different brain network encodes experimentalmanipulations of another aspect of the bodilyself: body ownership. This was shown by thebody-swap illusion (Petkova & Ehrsson 2008;Petkova et al. 2011), during which the parti-cipant views from a first-person visual view-point the body of a mannequin or another per-son. Thus no conflict between the visual spatialcoordinates of the participant’s physical bodyand the visually-perceived location of the man-nequin is presented. However, conflicting sens-ory information about the shape, gender, size,or overall spatial context surrounding the vir-tual body were presented that typically preven-ted feeling ownership of the virtual body. If un-der these conditions visuo-tactile stroking onthe abdomen of the participant and the virtualbody was synchronously administered, an illu-sion of ownership for the body emerged, reflec-ted in increased responses to threatening themannequin. In different variants of the body-swap illusion subjects reported experiencing andadopting different sizes of both the virtual bodyand the contextual environment (Petkova &Ehrsson 2008; Petkova et al. 2011; van derHoort et al. 2011). Neuroimaging experiments ofthe body-swap illusion show activation of thevPM and IPS regions, notably without in-volving actions made by subjects or performedby the virtual body (Petkova et al. 2011). Thesebrain regions are key nodes of the mirror mech-anism network of ES (see Serino et al. 2013).For a recent review see figure 1b.

3.3 A shared brain network of bodily self and social cognition

Although the neuroimaging evidence so far sug-gests that distinct brain regions encode the spa-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 7 | 14

www.open-mind.net

tial aspects of the bodily self and body owner-ship (Blanke 2012; Serino et al. 2013), the en-semble of those bodily self-encoding regionsclosely matches the brain regions relevant forsocial cognition (compare in figure 1a with fig-ure 1b). These empirical data indeed suggestthat the bodily self and social cognition are en-coded by at least overlapping neural circuitssupporting the proposal of ES that neural capa-cities to control and monitor the own body areused in understanding others.

These neuroimaging data suggest particu-lar functional associations between different as-pects of social cognition and the bodily self. Inparticular, the brain network of ES anatomic-ally overlaps with regions encoding experiment-ally-induced changes in body ownership duringthe body-swap illusion (figure 1a‒b in gray),which involves spatial congruence of the obser-vational viewpoint and position of the fakebody and the participant’s body. A second asso-ciation can be observed between the brain net-work of AS and the brain regions encoding spa-tial aspects of the bodily self, as manipulatedduring the full-body illusion (figure 1a‒b inblack). During the latter, the position and ob-servational viewpoints of the virtual body andthe participant’s body are in spatial conflict,and thus closely resemble social interaction set-tings.

Based on these functional and neuroana-tomical observations, I propose that ES seemsto contribute to the bodily self and social cogni-tion in a way primarily related to the sense ofbody ownership and agency. However, ES doesnot account for multisensory spatial representa-tions that relate the physical body to the bodilyself in space. These spatial aspects of the bodilyself are encoded by brain regions outside of thebrain network of ES, and rather resemble thosebrain regions relevant for coding the spatialconfiguration of attention (or awareness, accord-ing to AS).

Because two crucial aspects of the bodilyself, i.e., self-location and the first-person per-spective, are encoded in the TPJ region, andfull-body illusions show that they can be ma-nipulated without action or motor manipula-tions, it seems implausible that ES as based on

motor resonance is the primary brain mechan-ism underlying the bodily self. Instead, thebrain networks coding self-location and thefirst-person perspective, which overlap withbrain regions proposed to encode spatial aspectsof an attention schema (see figure 1), seem tocontribute to at least an equal degree to boththe bodily self and social cognition. Thus, ESseems to be a necessary but insufficiently“primary” brain mechanism underlying the bod-ily self and social cognition.

I do not mean to imply that these are in-dependent processes, because it is possible thatthey cooperatively work together (Graziano &Kastner 2011). However, I think that Galleseand Cuccio’s claim of a primacy of motor reson-ance underlying the multifaceted aspects of thebodily self and social cognition is questionableon empirical and theoretical grounds.

4 Vestibular contributions to the bodily self and social cognition

In the previous sections I have provided theoret-ical considerations and empirical evidence as-signing a critical role to multisensory spatialprocessing in the neural computations underly-ing representations of the bodily self and socialcognition. This section will further examine themultisensory mechanisms relating the space ofthe bodily self to other individuals and the ex-ternal world. I propose that important contribu-tions to the brain’s multisensory spatial codingmight come from a particular sensory system,i.e., the vestibular system, which has often beenneglected in studies of higher brain functions re-lated to subjectivity and intersubjectivity. I willask: What might be the functional contributionof the vestibular system to pre-reflective repres-entations of the bodily self and social cognition?How does the human vestibular cortex relate tothe neural networks of the bodily self and socialcognition?

The vestibular system consists of sensoryorgans in the inner ear that sense accelerationsof the head in space, including rotational andlinear movement of the head and whole bodyand the constant acceleration of gravity onearth (Day & Fitzpatrick 2005). Vestibular sig-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 8 | 14

www.open-mind.net

nals are processed by subcortical and corticalstructures (Angelaki & Cullen 2008; Cullen2012; Lopez & Blanke 2011). Research initiallyfocused on subcortical processing as related togaze control, postural stabilization, and neuralcomputations of head motion directions(Fernandez & Goldberg 1971; Goldberg &Fernandez 1971). More recently, studies have re-vealed the contribution of vestibular corticalprocessing to spatial cognition, body percep-tion, and the bodily self (see Lenggenhager &Lopez this collection; Lopez & Blanke 2011;Pfeiffer et al. 2014 for reviews). These studiesshow that vestibular cortical processing is basedon a neural network of distinct, distributed, andmultisensory cortical regions. In distinctionfrom any other sensory modality, there is noprimary vestibular cortex that processes purelyvestibular signals. Instead, a core vestibular cor-tical input region, the human parieto-insularvestibular cortex (PIVC; Lopez et al. 2012; zuEulenburg et al. 2012), processes vestibular, so-matosensory, and visual signals and is connec-ted to a number of multisensory brain regions inthe parietal, temporal, cingulate, and frontal re-gions (figure 1c).

The vestibular system contributes to spa-tial aspects of the bodily self. For instance,OBEs were associated with vestibular sensation,such as floating in elevation (Blanke et al. 2004;Blanke & Mohr 2005; Blanke et al. 2002), andvestibular sensations preceded OBEs in personswith sleep paralysis (Cheyne & Girard 2009).Other studies presented conflicting visual andvestibular signals about earth gravity duringthe full-body illusion and induced changes inthe subjectively-experienced spatial direction ofthe first-person perspective and self-location(Ionta et al. 2011; Pfeiffer et al. 2013). Thus, ithas been argued that vestibular cortical pro-cessing does not merely signal the motions ofthe own body and the external world, but isalso constitutive of spatial aspects of the bodilyself (Lopez et al. 2008; Pfeiffer et al. 2014).

Previously, Lopez et al. (2013), Deroualle& Lopez (2014), and Lenggenhager & Lopez(this collection) have argued that the vestibu-lar system probably contributes to social cog-nition. I will briefly summarize their main ar-

guments and complement them with ownpoints:

First, because the human species evolvedunder the steady influence of the earth’s gravit-ational field, adaptation to gravity also framedand affected action, perception, and social inter-action. More recently, research has shown thatthe brain hosts internal models of gravity, rep-resenting the effects of gravity on the motion ofobjects under the influence of gravity, of self-motion, of bodily actions, and of the directionof the gravitational acceleration. Those internalmodels of gravity strongly overlap with the ves-tibular cortex (Indovina et al. 2005; Indovina etal. 2013; McIntyre et al. 2001; Sciutti et al.2012). More evidence for a vestibular contribu-tion to social perception comes from studiesshowing the effects of gravitational signals onthe perception of emotional faces (Thompson1980) and the perception of the spatial orienta-tion of bodies (Lopez et al. 2009).

Second, the vestibular system might con-tribute to social cognition because it detectshead motions in space and hence directly en-ables us, when compared to other sensory sig-nals, to discern movements made by our ownbody from motions of other individuals and mo-tions of the external environment (Deroualle &Lopez 2014).

Third, mental spatial transformation ofthe own visual viewpoint to that of another per-son presents an important underlying cognitiveaspect of social cognition (Furlanetto 2013;Hamilton 2009; Newen & Vogeley 2003; alsocited by Gallese & Cuccio this collection, pp. 9–11). More direct evidence supporting this hypo-thesis comes from a recent study that showedthat physical whole-body rotations, which stim-ulate the vestibular sensory organs, affected theability of participants to perform mental spatialtransformations (van Elk & Blanke 2013).

Fourth, I have argued in previous sectionsof this commentary that multisensory spatialcoding is a critical prerequisite that underliespre-reflective brain mechanisms of the bodilyself and social cognition. Because the vestibularcortical processing has been strongly associatedwith multisensory integration (for review seeLopez & Blanke 2011), it is likely that vestibu-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 9 | 14

www.open-mind.net

lar signals shape multisensory spatial coding rel-evant to the bodily self and social cognition(Deroualle & Lopez 2014; Pfeiffer et al. 2014).

Fifth, the distributed multisensory vesti-bular cortical network clearly overlaps with theneural structures involved in social cognitionand the bodily self, which suggests that there isa functional contribution on the part of vestibu-lar processing to these phenomena (compare fig-ure 1c to 1a and 1b; compare also to Deroualle& Lopez 2014).

Together, these five points suggest thatthe vestibular system may be a promising can-didate for future studies of the sensorimotormechanisms of social cognition, which shouldmotivate research on the intersection of vestibu-lar cortical processing, mirror mechanisms, andintersubjectivity. These studies may, for in-stance, question how vestibular stimulation af-fects our ability to reconstruct the process of at-tention of another person, a function critical inthe AS framework. Although the vestibular sys-tem is related to reflexive motor control, it isnot clear whether it also affects motor reson-ance (see Deroualle & Lopez 2014 for a relatedproposal). One might ask whether vestibularprocessing facilitates or inhibits motor reson-ance and our understanding of intentional ac-tion observed in others. How about vestibularcontributions to theory of mind and reasoning?On the other hand, does social interaction mod-ulate vestibular functions, such as self-motionperception, postural stabilization, and gaze con-trol? These questions address the role of vesti-bular processing in functional mechanisms relev-ant to the AS and ES frameworks. Furthermore,empirical research addressing the causal rela-tionship between the AS and ES brain mechan-isms and the bodily self and social cognition areneeded, for instance by brain lesion analysis ordirect brain stimulation.

5 Conclusion

At the beginning of this paper I asked whichbrain mechanisms underlie pre-reflective repres-entations of the bodily self and social cognition.ES, based on motor resonance, substantiallycontributes to the representation of the bodily

self and social cognition. However, a unified the-ory of the neural basis of these target phenom-ena cannot assign a primary role to motor res-onance. I have argued that multisensory spatialcoding is at least of equal importance and prob-ably more basic than ES in contributing to sev-eral key aspects of the bodily self and socialcognition.

Specifically, I have argued that:

(1) Social cognition and the bodily self de-pend on multisensory spatial coding,which is distinct from motor resonance.Thus, motor resonance may be a necessarybut insufficiently “primary” brain mechan-ism of social cognition and the bodily self(cf. section 1, 2).

(2) The brain networks underlying socialcognition and the bodily self largely over-lap. Specific functional associations exist(a) between motor resonance and bodyownership/agency and (b) between multis-ensory spatial coding and self-location/thefirst-person perspective (cf. section 2).

(3) The vestibular system, i.e., the sensorysystem encoding head motion and gravity,might provide unique information used formultisensory spatial coding that relatesthe bodily self to other individuals and theexternal world. This is further suggestedby the large overlap existing between thehuman vestibular cortex and the brainnetworks underlying the bodily self andsocial cognition (cf. section 3).

A unifying theory of pre-reflective brainmechanisms of the bodily self and social cogni-tion must be able to account for the empiricalevidence reviewed here; and it seems that sucha theory cannot exclusively depend on motorresonance. Multisensory spatial coding, motormechanisms, but also representations of the pro-cess of attention appear highly relevant to bod-ily self and social cognition.

I agree with Gallese & Cuccio (this collec-tion, pp. 3–7) that cognitive neuroscience can-not fully explore these exciting topics by limit-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 10 | 14

www.open-mind.net

ing itself to a specific neuroimaging method,such as functional magnetic resonance imaging.Instead, we should exploit multi-method ap-proaches in search for correlative and causalevidence relating brain function and anatomy tothe phenomenology of the bodily self and socialcognition. The body, but also the spatial repres-entation of the world around us, are relevant tounderstanding brain function, and when takeninto account can lead to novel approaches tophenomenal analysis of subjective experience.But we should be careful in assigning priority toa single brain mechanism when aiming to ex-plain the human self and intersubjectivity. Scru-tiny and dialogue at the intersection of philo-sophy of mind and cognitive neuroscience arenecessary in order to advance our understandingof the nature of the human mind.

Acknowledgments

I thank Thomas Metzinger, Jennifer Windt, andtwo anonymous reviewers for constructive com-ments on an earlier version of this commentary.

References

Alsmith, A. J. & Longo, M. R. (2014). Where exactly amI? Self-location judgements distribute between headand torso. Consciousness and Cognition, 24, 70-74.10.1016/j.concog.2013.12.005

Angelaki, D. E. & Cullen, K. E. (2008). Vestibular sys-tem: the many facets of a multimodal sense. AnnualReviews in Neuroscience, 31, 125-150. 10.1146/annurev.neuro.31.060407.125555

Apps, M. A. & Tsakiris, M. (2013). Predictive codes offamiliarity and context during the perceptual learningof facial identities. Nature Communications, 4 (2698),2698-2698. 10.1038/ncomms3698

Aspell, J. E., Heydrich, L., Marillier, G., Lavanchy, T.,Herbelin, B. & Blanke, O. (2013). Turning body andself inside out: Visualized heartbeats alter bodily self-consciousness and tactile perception. Psychological Sci-ence, 24 (12). 10.1177/0956797613498395

Beck, D. M. & Kastner, S. (2009). Top-down and bottom-up mechanisms in biasing competition in the humanbrain. Vision Research, 49 (10), 1154-1165.10.1016/j.visres.2008.07.012

Blanke, O. (2012). Multisensory brain mechanisms ofbodily self-consciousness. Nature Reviews Neuroscience,13 (8), 556-571. 10.1038/nrn3292

Blanke, O., Ortigue, S., Landis, T. & Seeck, M. (2002).Stimulating illusory own-body perceptions. Nature, 419(6904), 269-270. 10.1038/419269a

Blanke, O., Landis, T., Spinelli, L. & Seeck, M. (2004).Out-of-body experience and autoscopy of neurologicalorigin. Brain, 127 (Pt 2), 243-258.10.1093/brain/awh040

Blanke, O. & Mohr, C. (2005). Out-of-body experience,heautoscopy, and autoscopic hallucination of neurolo-gical origin Implications for neurocognitive mechanismsof corporeal awareness and self-consciousness. BrainResearch Reviews, 50 (1), 184-199. 10.1016/j.brainresrev.2005.05.008

Chalmers, D. J. (1996). The conscious mind: In search ofa fundamental theory. Oxford, UK: Oxford UniversityPress.

Cheyne, J. A. & Girard, T. A. (2009). The body un-bound: Vestibular-motor hallucinations and out-of-body experiences. Cortex, 45 (2), 201-215. 10.1016/j.cortex.2007.05.002

Clark, A. (2015). Embodied prediction. In T. Metzinger& J. M. Windt (Eds.) Open MIND (pp. 1-21). Frank-furt a. M., GER: MIND Group.

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 11 | 14

www.open-mind.net

Cullen, K. E. (2012). The vestibular system: Multimodalintegration and encoding of self-motion for motor con-trol. Trends in Neurosciences, 35 (3), 185-196.10.1016/j.tins.2011.12.001

Day, B. L. & Fitzpatrick, R. C. (2005). The vestibularsystem. Current Biology, 15 (15), R583-R586.10.1016/j.cub.2005.07.053

De Ridder, D., Van Laere, K., Dupont, P., Menovsky, T.& Van de Heyning, P. (2007). Visualizing out-of-bodyexperience in the brain. The New England Journal ofMedicine, 357 (18), 1829-1833. 10.1056/NEJMoa070010

Deroualle, D. & Lopez, C. (2014). Toward a vestibularcontribution to social cognition. Frontiers in Integrat-ive Neuroscience, 8 (16). 10.3389/fnint.2014.00016

Desimone, R. & Duncan, J. (1995). Neural mechanisms ofselective visual attention. Annual Reviews in Neuros-cience, 18, 193-222.10.1146/annurev.ne.18.030195.001205

Downar, J., Crawley, A. P., Mikulis, D. J. & Davis, K. D.(2000). A multimodal cortical network for the detectionof changes in the sensory environment. Nature Neuros-cience, 3 (3), 277-283. 10.1038/72991

Ehrsson, H. H. (2007). The experimental induction ofout-of-body experiences. Science, 317 (5841).10.1126/science.1142175

(2012). The new handbook of multisensory pro-cesses. Cambridge, MA: MIT Press.

Fernandez, C. & Goldberg, J. M. (1971). Physiology ofperipheral neurons innervating semicircular canals ofthe squirrel monkey. II. Response to sinusoidal stimula-tion and dynamics of peripheral vestibular system.Journal of Neurophysiology, 34 (4), 661-675.

Friston, K. (2005). A theory of cortical responses. Philo-sophical Transactions of the Royal Society of London.Series B, Biological Sciences, 360 (2456), 815-836.10.1098/rstb.2005.1622

Furlanetto, T., Bertone, C. & Becchio, C. (2013). The bi-located mind: New perspectives on self-localization andself-identification. Frontiers in Human Neuroscience, 7(71). 10.3389/fnhum.2013.00071

Gallese, V. & Cuccio, V. (2015). The paradigmatic body.Embodied simulation, intersubjectivity and the bodilyself. In T. Metzinger & J. M. Windt (Eds.) OpenMIND. Frankfurt a. M., GER: MIND Group.

Gallese, V., Fadiga, L., Fogassi, L. & Rizzolatti, G.(1996). Action recognition in the premotor cortex.Brain, 119 (Pt 2), 593-609. 10.1093/brain/119.2.593

Goldberg, J. M. & Fernandez, C. (1971). Physiology ofperipheral neurons innervating semicircular canals of

the squirrel monkey. I. Resting discharge and responseto constant angular accelerations. Journal of Neuro-physiology, 34 (4), 635-660.

Graziano, M. S. (2013). Consciousness and the socialbrain. New York, NY: Oxford University Press.

Graziano, M. S. & Kastner, S. (2011). Human conscious-ness and its relationship to social neuroscience: A novelhypothesis. Cognitive neuroscience, 2 (2), 98-113.10.1080/17588928.2011.565121

Guterstam, A. & Ehrsson, H. H. (2012). Disowning one’sseen real body during an out-of-body illusion. Con-sciousness and Cognition, 21 (2), 1037-1042.10.1016/j.concog.2012.01.018

Hamilton, A. F., Brindley, R. & Frith, U. (2009). Visualperspective taking impairment in children with autisticspectrum disorder. Cognition, 113 (1), 37-44.10.1016/j.cognition.2009.07.007

Hohwy, J. (2013). The predictive mind. Oxford, UK: Ox-ford University Press.

(2015). The neural organ explains the mind. In T.Metzinger & J. M. Windt (Eds.) Open MIND (pp. 1-23). Frankfurt a. M., GER: MIND Group.

Husserl, E. (1950). Cartesianische Meditationen und Par-iser Vorträge. The Hague, NLD: Martinus Nijhoff Pub-lishers.

Indovina, I., Maffei, V., Bosco, G., Zago, M., Macaluso,E. & Lacquaniti, F. (2005). Representation of visualgravitational motion in the human vestibular cortex.Science, 308 (5720), 416-419. 10.1126/science.1107961

Indovina, I., Maffei, V., Pauwels, K., Macaluso, E.,Orban, G. A. & Lacquaniti, F. (2013). Simulated self-motion in a visual gravity field: Sensitivity to verticaland horizontal heading in the human brain. NeuroIm-age, 71, 114-124. 10.1016/j.neuroimage.2013.01.005

Ionta, S., Heydrich, L., Lenggenhager, B., Mouthon, M.,Fornari, E., Chapuis, D., Gassert, R. & Blanke, O.(2011). Multisensory mechanisms in temporo-parietalcortex support self-location and first-person perspect-ive. Neuron, 70 (2), 363-374.10.1016/j.neuron.2011.03.009

Jeannerod, M. (2003). The mechanism of self-recognitionin humans. Behavioural Brain Research, 142 (1-2), 1-15. 10.1016/S0166-4328(02)00384-4

Lenggenhager, B., Tadi, T., Metzinger, T. & Blanke, O.(2007). Video ergo sum: Manipulating bodily self-con-sciousness. Science, 317 (5841), 1096-1099. 10.1126/science.1143439

Lenggenhager, B., Mouthon, M. & Blanke, O. (2009).Spatial aspects of bodily self-consciousness. Conscious-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 12 | 14

www.open-mind.net

ness and Cognition, 18 (1), 110-117.10.1016/j.concog.2008.11.003

Lenggenhager, B. & Lopez, C. (2015). Vestibular contri-butions to the sense of body, self and others. In T.Metzinger & J. M. Windt (Eds.) Open MIND. Frank-furt a. M., GER: MIND Group.

Limanowski, J. & Blankenburg, F. (2013). Minimal self-models and the free energy principle. Frontiers in Hu-man Neuroscience, 7 (547). 10.3389/fnhum.2013.00547

Limanowski, J. & Hecht, H. (2011). Where do we standon locating the self? Psychology, 2 (4), 312-317.10.4236/psych.2011.24049

Lopez, C. & Blanke, O. (2011). The thalamocortical ves-tibular system in animals and humans. Brain ResearchReviews, 67 (1-2), 119-146.10.1016/j.brainresrev.2010.12.002

Lopez, C., Halje, P. & Blanke, O. (2008). Body ownershipand embodiment: vestibular and multisensory mechan-isms. Clinical Neurophysiology, 38 (3), 149-161.10.1016/j.neucli.2007.12.006

Lopez, C., Bachofner, C., Mercier, M. & Blanke, O.(2009). Gravity and observer’s body orientation influ-ence the visual perception of human body postures.Journal of Vision, 9 (5), 11-14. 10.1167/9.5.1

Lopez, C., Blanke, O. & Mast, F. W. (2012). The humanvestibular cortex revealed by coordinate-based activa-tion likelihood estimation meta-analysis. Neuroscience,212, 159-179. 10.1016/j.neuroscience.2012.03.028

Lopez, C., Falconer, C. J. & Mast, F. W. (2013). Beingmoved by the self and others: Influence of empathy onself-motion perception. PloS one, 8 (1), e48293-e48293.10.1371/journal.pone.0048293

McIntyre, J., Zago, M., Berthoz, A. & Lacquaniti, F.(2001). Does the brain model Newton’s laws? NatureNeuroscience, 4 (7), 693-694. 10.1038/89477

Metzinger, T. (2003). Being no one. Boston: MIT Press. (2009). Why are out-of-body experiences interest-

ing for philosophers? The theoretical relevance of OBEresearch. Cortex, 45 (2), 256-258.10.1016/j.cortex.2008.09.004

Newen, A. & Vogeley, K. (2003). Self-representation:Searching for a neural signature of self-consciousness.Consciousness and Cognition, 12 (4), 529-543.10.1016/S1053-8100(03)00080-1

Petkova, V. I. & Ehrsson, H. H. (2008). If I were you:perceptual illusion of body swapping. PLOS ONE, 3(8), e3832-e3832. 10.1371/journal.pone.0003832

Petkova, V. I., Bjornsdotter, M., Gentile, G., Jonsson, T.,Li, T. Q. & Ehrsson, H. H. (2011). From part- to

whole-body ownership in the multisensory brain. Cur-rent Biology, 21 (13), 1118-1122.10.1016/j.cub.2011.05.022

Petkova, V. I., Khoshnevis, M. & Ehrsson, H. H. (2011).The perspective matters! Multisensory integration inego-centric reference frames determines full-body own-ership. Frontiers in Psychology, 2 (35).10.3389/fpsyg.2011.00035

Pfeiffer, C., Lopez, C., Schmutz, V., Duenas, J. A., Mar-tuzzi, R. & Blanke, O. (2013). Multisensory origin ofthe subjective first-person perspective: Visual, tactile,and vestibular mechanisms. PloS one, 8 (4).10.1371/journal.pone.0061751

Pfeiffer, C., Schmutz, V. & Blanke, O. (2014). Visuospa-tial viewpoint manipulation during full-body illusionmodulates subjective first-person perspective. Experi-mental Brain Research. 10.1007/s00221-014-4080-0

Pfeiffer, C., Serino, A. & Blanke, O. (2014). The vestibu-lar system: A spatial reference for bodily self-concious-ness. Frontiers in Integrative Neuroscience, 8 (31).10.3389/fnint.2014.00031

Rizzolatti, G., Fadiga, L., Gallese, V. & Fogassi, L.(1996). Premotor cortex and the recognition of motoractions. Cognitive Brain Research, 3 (2), 131-141.

Rizzolatti, G. & Sinigaglia, C. (2010). The functional roleof the parieto-frontal mirror circuit: Interpretationsand misinterpretations. Nature Reviews Neuroscience,11 (4), 264-274. 10.1038/nrn2805

Sciutti, A., Demougeot, L., Berret, B., Toma, S., Sandini,G., Papaxanthis, C. & Pozzo, T. (2012). Visual gravityinfluences arm movement planning. Journal of neuro-physiology, 107 (12), 3433-3445. 10.1152/jn.00420.2011

Serino, A., Alsmith, A., Costantini, M., Mandrigin, A.,Tajadura-Jimenez, A. & Lopez, C. (2013). Bodily own-ership and self-location: Components of bodily self-con-sciousness. Consciousness and Cognition, 22 (4), 1239-1252. 10.1016/j.concog.2013.08.013

Seth, A. (2014). The cybernetic Bayesian brain: From in-teroceptive inference to sensorimotor contingencies. InT. Metzinger & J. W. Windt (Eds.) Open MIND.Frankfurt am Main: MIND Group.

Stein, B. E. & Stanford, T. R. (2008). Multisensory integ-ration: current issues from the perspective of the singleneuron. Nature Reviews Neuroscience, 9 (4), 255-266.10.1038/nrn2331

Thompson, P. (1980). Margaret Thatcher: A new illusion.Perception, 9 (4), 483-484.

van der Hoort, B., Guterstam, A. & Ehrsson, H. H.(2011). Being Barbie: The size of one’s own body de-

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 13 | 14

www.open-mind.net

termines the perceived size of the world. PLOS ONE, 6(5), e20195-e20195. 10.1371/journal.pone.0020195

van Elk, M. & Blanke, O. (2013). Imagined own-bodytransformations during passive self-motion. Psycholo-gical Research, 78 (1). 10.1007/s00426-013-0486-8

Windt, J. M. (2010). The immersive spatiotemporal hal-lucination model of dreaming. Phenomenology and theCognitive Sciences, 9 (2), 295-316. 10.1007/s11097-010-9163-1

zu Eulenburg, P., Caspers, S., Roski, C. & Eickhoff, S. B.(2012). Meta-analytical definition and functional con-nectivity of the human vestibular cortex. NeuroImage,60 (1), 162-169. 10.1016/j.neuroimage.2011.12.032

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms of the Bodily Self and Social Cognition - A Commentary on Vittorio Gallese &Valentina Cuccio. In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(C). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570450 14 | 14

Embodied Simulation: A Paradigm for the Constitution of Self and OthersA Reply to Christian Pfeiffer

Vittorio Gallese & Valentina Cuccio

The main criticism Pfeiffer advances in his commentary is that our proposal is toonarrow. Embodied simulation (ES), in his view equated to motor resonance, is nota sufficiently primary mechanism on which we can base a unified neurobiologicaltheory of the earliest sense of self and others. According to Pfeiffer, motor reson-ance needs to be complemented by other more basic and primary mechanisms.Hence, as an alternative to our proposal, he suggests that multisensory spatialprocessing can play this role, primarily contributing to the earliest foundation ofthe sense of self and others. In our reply we stress on the one hand that identify-ing ES only with motor resonance is a partial view that may give rise to falla-cious arguments, since ES also deals with emotions and sensations. We alsoshow, on the other hand, that ES and multisensory integration should not be seenas alternative solutions to the problem of the neural bases of the bodily self, be-cause multimodal integration carried out by the cortical motor system is an in-stantiation of ES. We conclude by stressing the role ES might have played in thetransition from bodily experience to symbolic expression.

KeywordsAttention schema theory | Bodily self | Embodied simulation | Language | Motorresonance | Multimodal integration | Paradigm | Peri-personal space | Social cog-nition

Authors

Vittorio [email protected]   Università degli Studi di ParmaParma, Italy

Valentina CuccioUniversità degli Studi di PalermoPalermo, Italy

Commentator

Christian [email protected]   Ecole Polytechnique FédéraleLausanne, Switzerland

Editors

Thomas [email protected]   Johannes Gutenberg-UniversitätMainz, Germany

Jennifer M. [email protected]   Monash UniversityMelbourne, Australia

1 An overview of Pfeiffer’s criticisms

We would like to thank Christian Pfeiffer for hisvery well-articulated commentary on our paper“The paradigmatic body: Embodied Simulation,Intersubjectivity, the Bodily Self, and Lan-guage” (Gallese & Cuccio this collection). Hiscomments and criticisms offered us the oppor-tunity to further reflect on some of the ideas

proposed in our piece. The aim of our paperwas to discuss the role of the body in the con-stitution of the earliest and primary sense of selfand others and, also, to emphasize the con-stitutive role of the body in a specifically hu-man modality of intersubjectivity: language. Tobe more precise, we identified a biological mech-

Gallese, V. & Cuccio, V. (2015). Embodied Simulation: A Paradigm for the Constitution of Self and Others - A Reply to Christian Pfeiffer.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(R). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570962 1 | 5

www.open-mind.net

anism, embodied simulation (ES), as a primarysource of intersubjectivity, the sense of self, andlanguage. The mechanism of ES is widely de-scribed in the paper and its role in human cog-nition is explained by also resorting to the Aris-totelian notion of paradeigma.

The commentary offered by ChristianPfeiffer is focused on a partial aspect of ourmuch wider proposal. In fact, the author onlydiscusses the constitutive role motor resonancehas for the sense of self and for social cognition.However, motor resonance is just one dimensionof the mechanism of ES. As argued in our paperand elsewhere (see Gallese & Sinigaglia 2011a;Gallese 2014) the mechanism of simulation iswidespread in the brain and it also characterizesthe nervous structures involved in the experi-ence of emotions and sensations. All these di-mensions of ES should be taken into account.To identify ES only with motor resonance is apartial view that may give rise to fallacious ar-guments.

The main criticism Pfeiffer advances in hiscommentary is that our proposal for the con-stitutive role of motor resonance is too narrow.ES, in his view equated to motor resonance,cannot be the primary neurobiological mechan-ism at the basis of both the sense of self andothers. According to Pfeiffer, motor resonanceneeds to be complemented by other more basicand primary mechanisms. Hence, as an alternat-ive to our proposal, he suggests that multisens-ory spatial processing can play this role, primar-ily contributing to the earliest foundation of thesense of self and others. To support this claim,he provides theoretical arguments and presentsempirical data structured in three different sec-tions. Each of these sections supposedlyprovides evidence of the role of multisensoryspatial processing in the foundation of a bodilysense of self and others.

In the first section Pfeiffer addresses theissue of intersubjectivity and presents the At-tention schema theory (AS). In his proposal,our ability to understand others is primarilybased on a mechanism more primitive than ES-as-motor-resonance: spatial coding of attention.AS predicts that we understand the currentstate of awareness of our conspecifics by means

of schematic representations of their states ofattention (Pfeiffer this collection, p. 4). In otherwords, according to AS, by using a representa-tion of the spatial relationship between the indi-vidual we are observing and the spatial focus ofher/his attention we can likely predict his inten-tions and, as a consequence, his actions. Pfeiffer(this collection, p. 4) also discusses recent em-pirical findings on the neural structures under-lying the AS. It seems that the neural struc-tures for the spatial coding of attention arebased in the right temporo-parietal junctio(TPJ) and in the superior temporal sulcus(STS). These neural structures do not overlapwith the neural circuits involved in ES.

In the second section Pfeiffer addresses theissue of the bodily foundation of the sense ofself. The experience of being a bodily self canbe decomposed into four different aspects (Pfeif-fer this collection, p. 5): body ownership, self-location, first-person perspective, and agency.According to Pfeiffer, motor resonance can ac-count only for body ownership and agency, dir-ectly contributing to these (non-spatial) aspectsof the bodily self. However, for the two spatialcomponents of the bodily self we need a differ-ent account. In fact, according to Pfeiffer, em-pirical evidence suggests that these spatial as-pects of the bodily self, which imply multisens-ory spatial representations, are encoded in abrain region, the TPJ, not characterized by mo-tor resonance. Hence, motor resonance, whilebeing still necessary for the bodily foundation ofsome basic aspects of the self, is not a suffi-ciently primary mechanism, since differentneural structures are also needed for the bodilyfoundation of the self. In support of this claim,Pfeiffer discusses data from neurological pa-tients with out-of-body experiences and otherkinds of altered states.

Finally, in the third section the con-stitutive role of the vestibular system to thebodily foundation of both the consciousness ofself and others is discussed. It is proposed thatthis system, which encodes gravity and headmotion and is associated with multisensory spa-tial processing, significantly and primarily con-tributes to our ability to distinguish betweenmotions of our own body and motions of other

Gallese, V. & Cuccio, V. (2015). Embodied Simulation: A Paradigm for the Constitution of Self and Others - A Reply to Christian Pfeiffer.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(R). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570962 2 | 5

www.open-mind.net

people’s bodies, in this way contributing toboth the foundation of the sense of self and so-cial cognition. Empirical studies are reported tosupport these claims. In addition, empiricaldata showing that the vestibular cortical net-work overlaps with neural structures underlyingthe bodily foundation of both the sense of selfand others, as discussed in the two previous sec-tions, are presented.

In the light of the empirical evidence dis-cussed in his commentary, Christian Pfeifferconcludes that ES-as-motor-resonance is not asufficiently primary mechanism on which we canbase a unified neurobiological theory of theearliest sense of self and others. In the next sec-tion we answer these criticisms.

2 Responses

First, we would like to point out that ES is notconfined to motor resonance of others’ actions,like that instantiated by macaques’ mirror neur-ons, as in humans ES also encompasses the ac-tivation of somatosensory areas during the ob-servation of others’ tactile experiences, the ac-tivation of pain-related areas like the anteriorinsula and the anterior cingulate cortex duringthe observation of others’ pain, and the activa-tion of the anterior insula and limbic structureslike the amygdala during the observation of oth-ers’ emotions like disgust and fear (see our pa-per, p. 9 and Gallese & Sinigaglia 2011a). Thus,motor resonance only describes one partial as-pect of ES.

Two distinct arguments can be used to ex-plain why we do not think that AS constitutesa valid alternative to ES, as argued by Pfeiffer.We certainly agree with Pfeiffer that shared at-tention, that is, the capacity to direct the gazeto an object gazed by someone else, is a basicingredient of social cognition. Indeed, as main-tained by Colwyn Trevarthen (1977), shared at-tention marks in human infants around 9months of age the transition from primary tosecondary intersubjectivity. However, shared at-tention constitutes only one aspect of intersub-jectivity and social cognition, thus AS at bestonly covers a partial aspect of social cognitionand therefore appears to be more limited than

ES in this respect. Furthermore, and most im-portantly, shared attention can be linked to mo-tor resonance. Shepherd, Klein, Deaner, andPlatt 2009) discovered in macaques a class ofmirror neurons in the lateral intraparietal (LIP)area involved in oculomotor control, signalingboth when the monkey looked at a given direc-tion in space and when it observed anothermonkey looking in the same direction. Theseauthors suggested that LIP mirror neurons forgaze might contribute to the sharing of ob-served attention. This evidence shows thatshared attention is not divorced from motor res-onance, but actually requires it.

A further argument in our opinion demon-strates that ES and AS should not be seen asalternative solutions to the problem of socialcognition. Multisensory integration is a pervas-ive feature of parieto-frontal centers involved insensory-motor planning and control. Indeed aninfluential theory about attention, the “Premo-tor Theory of Attention” (see Rizzolatti et al.1987; Rizzolatti et al. 1994) states that spatialattention results from the activation of the same“pragmatic” circuits that program oculomotorbehavior and other motor activities, even if suchactivation does not produce any overt motor be-havior, thus qualifying as motor simulation.

We would like to emphasize even morestrongly than we did in the paper that a crucialrole of the cortical motor system is preciselythat of integrating multiple sources of body-re-lated sensory signals, like tactile, visual andauditory stimuli (see our paper, pp. 10–11; seealso Gallese & Sinigaglia 2010, 2011b; Gallese2014). The ventral premotor cortex (vPMC)might represent one of the essential anatomo-functional bases for the motor aspect of bodilyselfhood, specifically because of its role in integ-rating self-related multisensory information.This hypothesis is corroborated by clinical andfunctional evidence showing the systematic in-volvement of vPMC with body awareness(Ehrsson et al. 2004; Berti et al. 2005; Arzy etal. 2006). This evidence demonstrates a tight re-lationship between the bodily self-related mul-timodal integration carried out by the corticalmotor areas specifying the motor potentialitiesof one’s body and guiding its motor behavior

Gallese, V. & Cuccio, V. (2015). Embodied Simulation: A Paradigm for the Constitution of Self and Others - A Reply to Christian Pfeiffer.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(R). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570962 3 | 5

www.open-mind.net

and the implicit awareness one entertains ofone’s body as one’s own body and of one’s be-havior as one’s own behavior.

The vPMC is anatomically connected tovisual and somatosensory areas in the posteriorparietal cortex and to frontal motor areas andfor this reason it is plausible to assume thatvPMC activity reflects the detection of congru-ent multisensory signals related to one’s ownbody parts: this mechanism could be respons-ible for the feeling of body ownership. The mo-tor aspects of the bodily self-enable the integra-tion of self-related multimodal sensory informa-tion about the body and about the world withwhich the body interacts, as epitomized by theproperties of macaques’ premotor neurons inarea F4 (see Fogassi et al. 1996; Rizzolatti et al.1997) and the analogous functional propertiesdisplayed by the human homologue of area F4(see Bremmer et al. 2001). The same neuronscontrolling the movement in space of the heador of the upper limb also respond to tactile,visual, and auditory stimuli, provided they areapplied to the same body part, like tactile stim-uli, or they occur in the body-part-centeredperi-personal space, like visual and auditorystimuli. Thus, we think that ES and multisens-ory integration should not be seen as alternativesolutions to the problem of the neural bases ofthe bodily self, because multimodal integrationcarried out by vPMC is an instantiation of ES.We agree with Pfeiffer, however, that otherbrain areas, like TPJ, might contribute to a co-herent sense of one’s own body. It must be ad-ded that TPJ is part of a network (includingthe posterior parietal cortex, and the premotorcortex) implicated in multisensory integrationduring self-related and other-related events andexperiences. Indeed, as shown by Ebisch et al.(2011), the observation of others’ affective tact-ile experiences leads to the activation of observ-ers’ vPMC and second somatosensory area andto the inactivation of observers’ posterior insula.Functional connectivity revealed a significant in-teraction between the posterior insula, rightTPJ, left pre-central gyrus, and right posteriorparietal cortex during the observation of other’saffective touch. These data suggest that TPJmight be involved in mapping the self–other dif-

ferentiation, by means of lower-level computa-tional mechanisms for generating, testing, andcorrecting internal predictions about externalsensory events.

Last, we agree with Pfeiffer that the vesti-bular system might contribute to the bodilyfoundation of both the consciousness of self andothers and we thank him for having pointedthis out, thus integrating our perspective.

3 Conclusions

It seems that the data discussed in the previoussection allow us to come to the conclusion thatES is the primary and earliest mechanism con-tributing to the foundation of the sense of selfand others. That said, in conclusion, we wouldlike to stress again the issue of the cognitiverole ES has in relation to language. Though theaspect of the relation between ES and languagewas not addressed in Pfeiffer’s commentary, thiswas a central point of our proposal. The rela-tion between ES and language is two-sided. Onthe one hand, empirical evidence has shown therole ES plays in language comprehension. Thesedata (for an overview see Gallese & Cuccio thiscollection, p. 13) suggest that the bodily, sens-ory, and motor dimensions play a constitutiverole in language, both ontogenetically andphylogenetically. On the other hand, being lin-guistic creatures, we humans are the only livingspecies able to fix and relive specific aspects ofour bodily experiences by means of symbols.Words or other forms of symbolic representa-tions such as art, for example, allow us to activ-ate and relive our bodily experiences. In thisway, by means of symbolic representations, wecan share our bodily experiences, enacted byES, even with people far away from us in timeand space. As argued in our paper, ES is amodel of our own experiences and its definingfeatures are best explained by resorting to theAristotelian notion of paradeigma. ES-as-paradeigma (and not just as motor resonance)provides a neurobiologically-based new per-spective on human social cognition and ulti-mately on the very definition of human nature.

Gallese, V. & Cuccio, V. (2015). Embodied Simulation: A Paradigm for the Constitution of Self and Others - A Reply to Christian Pfeiffer.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(R). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570962 4 | 5

www.open-mind.net

Acknowledgments

This work was supported by the EU Grant To-wards an Embodied Science of InterSubjectivity(TESIS, FP7-PEOPLE-2010-ITN, 264828) andby the KOSMOS Fellowship from HumboldtUniversity, Berlin to VG.

References

Arzy, S., Overney, L. S., Landis, T. & Blanke, O. (2006).Neural mechanisms of embodiment: Asomatognosia dueto premotor cortex damage. Archives of Neurology, 63(7), 1022-1025. Neural mechanisms of embodiment:Asomatognosia due to premotor cortex damage

Berti, A., Bottini, G., Gandola, M., Pia, L., Smania, N.,Stracciari, A., Castiglioni, I., Vallar, G. & Paulesu, E.(2005). Shared cortical anatomy for motor awarenessand motor control. Science, 309 (5733), 488-491.10.1126/science.1110625

Bremmer, F., Schlack, A., Shah, N. J., Zafiris, O., Kubis-chik, M., Hoffmann, K., Zilles, K. & Fink, G. R.(2001). Polymodal motion processing in posterior pari-etal and premotor cortex: a human fMRI studystrongly implies equivalencies between humans andmonkeys. Neuron, 29 (1), 287-296. 10.1016/S0896-6273(01)00198-2

Ebisch, S. J. H., Ferri, F., Salone, A., d’Amico, L.,Perucci, M. G., Ferro, F. M., Romani, G. L. & Gallese,V. (2011). Differential involvement of somatosensoryand interoceptive cortices during the observation of af-fective touch. Journal of Cognitive Neurosciences, 23(7), 1808-1822. 10.1162/jocn.2010.21551

Ehrsson, H. H., Spence, C. & Passingham, R. E. (2004).That’s my hand! Activity in premotor cortex reflectsfeeling of ownership of a limb. Science, 305 (5685), 875-877. 10.1126/science.1097011

Fogassi, L., Gallese, V., Fadiga, L., Luppino, G., Matelli,M. & Rizzolatti, G. (1996). Coding of peripersonalspace in inferior premotor cortex (area F4). Journal ofNeurophysiology, 76 (1), 141-157.

Gallese, V. (2014). Bodily Selves in Relation: Embodiedsimulation as second-person perspective on intersub-jectivity. Philosophical Transactions of the Royal Soci-ety, London. Series B Biological Sciences, 369 (1644),20130177-20130177. 10.1098/rstb.2013.0177

Gallese, V. & Cuccio, V. (2015). The Paradigmatic Body.In T. Metzinger & J. M. Windt (Eds.) Open MIND(pp. 1-23). Frankfurt a. M., GER: MIND Group.

Gallese, V. & Sinigaglia, C. (2010). The bodily self aspower for action. Neuropsychologia, 48 (3), 746-755.10.1016/j.neuropsychologia.2009.09.038

(2011a). What is so special with Embodied Simu-lation. Trends in Cognitive Sciences, 15 (11), 512-519.10.1016/j.tics.2011.09.003

(2011b). How the body in action shapes the self.Journal of Consciousness Studies, 18 (7-8), 117-143.

Pfeiffer, C. (2015). Multisensory Spatial Mechanisms ofthe Bodily Self and Social Cognition. In T. Metzinger& J. M. Windt (Eds.) Open MIND (pp. 1-14). Frank-furt a. M., GER: MIND Group.

Rizzolatti, G., Riggio, L., Dascola, I. & Umiltá, C.(1987). Reorienting attention across the horizontal andvertical meridians: evidence in favor of a premotor the-ory of attention. Neuropsychologia, 25 (1a), 31-40.

Rizzolatti, G., Riggio, L. & Sheliga, B. M. (1994). Spaceand selective attention. In C. Umiltá & M. Moscovitch(Eds.) Attention and performance XV (pp. 231-265).Cambridge, MA: MIT Press.

Rizzolatti, G., Fadiga, L., Fogassi, L. & Gallese, V.(1997). The space around us. Science, 277 (5323), 190-191. 10.1126/science.277.5323.190

Shepherd, S. V., Klein, J. T., Deaner, R. O. & Platt, M.L. (2009). Mirroring of attention by neurons inmacaque parietal cortex. Proceedings of The NationalAcademy of Sciences USA, 106 (23), 9489-9494.10.1073/pnas.0900419106

Trevarthen, C. (1977). Descriptive analyses of infant com-municative behavior. In H. R. Schaffer (Ed.) Studies inmother-infant interaction (pp. 227-270). London, UK:Academic Press.

Gallese, V. & Cuccio, V. (2015). Embodied Simulation: A Paradigm for the Constitution of Self and Others - A Reply to Christian Pfeiffer.In T. Metzinger & J. M. Windt (Eds). Open MIND: 14(R). Frankfurt am Main: MIND Group. doi: 10.15502/9783958570962 5 | 5