Robust Selectivity for Faces in the Human Amygdala in the Absence of Expressions

There is a well-established posterior network of cortical regions that plays a central role in face processing and that has been investigated extensively. In contrast, although responsive to faces, the amygdala is not considered a core face-selective region, and its face selectivity has never been a topic of systematic research in human neuroimaging studies. Here, we conducted a large-scale group analysis of fMRI data from 215 participants. We replicated the posterior network observed in prior studies but found equally robust and reliable responses to faces in the amygdala. These responses were detectable in most individual participants, but they were also highly sensitive to the initial statistical threshold and habituated more rapidly than the responses in posterior face-selective regions. A multivariate analysis showed that the pattern of responses to faces across voxels in the amygdala had high reliability over time. Finally, functional connectivity analyses showed stronger coupling between the amygdala and posterior face-selective regions during the perception of faces than during the perception of control visual categories. These findings suggest that the amygdala should be considered a core face-selective region.

[1]  A. O'Toole,et al.  Prototype-referenced shape encoding revealed by high-level aftereffects , 2001, Nature Neuroscience.

[2]  Marco Tamietto,et al.  Emotion in the brain: of low roads, high roads and roads less travelled , 2011, Nature Reviews Neuroscience.

[3]  Alexander Todorov,et al.  Modeling Social Perception of Faces , 2011 .

[4]  Luiz Pessoa,et al.  Target visibility and visual awareness modulate amygdala responses to fearful faces. , 2006, Cerebral cortex.

[5]  A. Lüthi,et al.  Processing of Temporal Unpredictability in Human and Animal Amygdala , 2007, The Journal of Neuroscience.

[6]  Kalanit Grill-Spector,et al.  The improbable simplicity of the fusiform face area , 2012, Trends in Cognitive Sciences.

[7]  A. Mikami,et al.  Activity of single neurons in the monkey amygdala during performance of a visual discrimination task. , 1992, Journal of neurophysiology.

[8]  Kalanit Grill-Spector,et al.  Sparsely-distributed organization of face and limb activations in human ventral temporal cortex , 2010, NeuroImage.

[9]  P. Vuilleumier,et al.  How brains beware: neural mechanisms of emotional attention , 2005, Trends in Cognitive Sciences.

[10]  J. Tanaka,et al.  The NimStim set of facial expressions: Judgments from untrained research participants , 2009, Psychiatry Research.

[11]  Ingrid R. Olson,et al.  Beyond the FFA: The role of the ventral anterior temporal lobes in face processing , 2014, Neuropsychologia.

[12]  Leslie G. Ungerleider,et al.  Dissociation of object and spatial visual processing pathways in human extrastriate cortex. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[13]  M. Bar,et al.  Very first impressions. , 2006, Emotion.

[14]  Michael Davis,et al.  The amygdala: vigilance and emotion , 2001, Molecular Psychiatry.

[15]  A. Todorov,et al.  EvaluaTiNg faCES ON TruSTwOrThiNESS afTEr miNimal TimE ExpOSurE , 2009 .

[16]  T. Allison,et al.  Social perception from visual cues: role of the STS region , 2000, Trends in Cognitive Sciences.

[17]  Alexander Todorov,et al.  Brain systems for assessing the affective value of faces , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

[18]  J. Sergent,et al.  Functional neuroanatomy of face and object processing. A positron emission tomography study. , 1992, Brain : a journal of neurology.

[19]  Peter Mende-Siedlecki,et al.  Social judgments from faces , 2013, Current Opinion in Neurobiology.

[20]  A. O'Toole,et al.  Three-Dimensional Information in Face Representations Revealed by Identity Aftereffects , 2009, Psychological science.

[21]  C. Büchel,et al.  The robustness of perception , 2005, The European journal of neuroscience.

[22]  M M Mesulam,et al.  Impact of signal-to-noise on functional MRI of the human amygdala , 2001, Neuroreport.

[23]  Andrew D. Engell,et al.  Task-invariant Brain Responses to the Social Value of Faces , 2011, Journal of Cognitive Neuroscience.

[24]  A. Ohman,et al.  Masking the face: recognition of emotional facial expressions as a function of the parameters of backward masking. , 1993, Scandinavian journal of psychology.

[25]  M. Mesulam,et al.  Dissociation of Neural Representation of Intensity and Affective Valuation in Human Gustation , 2003, Neuron.

[26]  T. Allison,et al.  Face-sensitive regions in human extrastriate cortex studied by functional MRI. , 1995, Journal of neurophysiology.

[27]  G. Glover,et al.  Dissociated neural representations of intensity and valence in human olfaction , 2003, Nature Neuroscience.

[28]  R. Tootell,et al.  An anterior temporal face patch in human cortex, predicted by macaque maps , 2009, Proceedings of the National Academy of Sciences.

[29]  David G. Amaral,et al.  Neuroanatomy of the primate amygdala. , 2009 .

[30]  Ralph Adolphs,et al.  Fear, faces, and the human amygdala , 2008, Current Opinion in Neurobiology.

[31]  E. Rolls,et al.  Visual responses of neurons in the dorsolateral amygdala of the alert monkey , 1979, Experimental Neurology.

[32]  D. Perrett,et al.  Visual neurones responsive to faces in the monkey temporal cortex , 2004, Experimental Brain Research.

[33]  A. David,et al.  Predictors of amygdala activation during the processing of emotional stimuli: A meta-analysis of 385 PET and fMRI studies , 2008, Brain Research Reviews.

[34]  K. Zilles,et al.  ALE meta-analysis on facial judgments of trustworthiness and attractiveness , 2010, Brain Structure and Function.

[35]  Mark H. Johnson,et al.  Newborns' preference for face-relevant stimuli: effects of contrast polarity. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[36]  C. Koch,et al.  Category-specific visual responses of single neurons in the human medial temporal lobe , 2000, Nature Neuroscience.

[37]  Paul J. Whalen,et al.  Amygdaloid contributions to conditioned arousal and sensory information processing. , 1992 .

[38]  E. Rolls,et al.  The effects of stimulus novelty and familiarity on neuronal activity in the amygdala of monkeys performing recognition memory tasks , 2004, Experimental Brain Research.

[39]  D. Perrett,et al.  A differential neural response in the human amygdala to fearful and happy facial expressions , 1996, Nature.

[40]  T. Allison,et al.  Face recognition in human extrastriate cortex. , 1994, Journal of neurophysiology.

[41]  Christopher P. Said,et al.  The social evaluation of faces: a meta-analysis of functional neuroimaging studies. , 2013, Social cognitive and affective neuroscience.

[42]  K. Zilles,et al.  An investigation of the structural, connectional, and functional subspecialization in the human amygdala , 2012, Human brain mapping.

[43]  I. Gauthier,et al.  Expertise for cars and birds recruits brain areas involved in face recognition , 2000, Nature Neuroscience.

[44]  N. Kanwisher Functional specificity in the human brain: A window into the functional architecture of the mind , 2010, Proceedings of the National Academy of Sciences.

[45]  Charles L. Wilson,et al.  Single Neuron Activity in Human Hippocampus and Amygdala during Recognition of Faces and Objects , 1997, Neuron.

[46]  Dimitri Van De Ville,et al.  White-Matter Connectivity between Face-Responsive Regions in the Human Brain , 2012 .

[47]  J. Haxby,et al.  The distributed human neural system for face perception , 2000, Trends in Cognitive Sciences.

[48]  D. B. Bender,et al.  Visual properties of neurons in inferotemporal cortex of the Macaque. , 1972, Journal of neurophysiology.

[49]  John Patrick Aggleton,et al.  The Amygdala : a functional analysis , 2000 .

[50]  Derek K. Jones,et al.  Occipito-temporal connections in the human brain. , 2003, Brain : a journal of neurology.

[51]  B. Rossion,et al.  Defining face perception areas in the human brain: A large-scale factorial fMRI face localizer analysis , 2012, Brain and Cognition.

[52]  Alexander Todorov,et al.  The amygdala and FFA track both social and non-social face dimensions , 2010, Neuropsychologia.

[53]  Gilles Pourtois,et al.  Temporal precedence of emotion over attention modulations in the lateral amygdala: Intracranial ERP evidence from a patient with temporal lobe epilepsy , 2010, Cognitive, affective & behavioral neuroscience.

[54]  K. Nakayama,et al.  RESPONSE PROPERTIES OF THE HUMAN FUSIFORM FACE AREA , 2000, Cognitive neuropsychology.

[55]  K. Grill-Spector,et al.  Neural representations of faces and limbs neighbor in human high-level visual cortex: evidence for a new organization principle , 2011, Psychological Research.

[56]  Marta I. Garrido,et al.  Functional Evidence for a Dual Route to Amygdala , 2012, Current Biology.

[57]  N. Kanwisher,et al.  The fusiform face area: a cortical region specialized for the perception of faces , 2006, Philosophical Transactions of the Royal Society B: Biological Sciences.

[58]  R. Adolphs,et al.  Single-Unit Responses Selective for Whole Faces in the Human Amygdala , 2011, Current Biology.

[59]  Juraj Kukolja,et al.  Selective processing of social stimuli in the superficial amygdala , 2009, Human brain mapping.

[60]  R. Desimone,et al.  Neural mechanisms of selective visual attention. , 1995, Annual review of neuroscience.

[61]  Nicholas B. Turk-Browne,et al.  Representations of Facial Identity in the Left Hemisphere Require Right Hemisphere Processing , 2012, Journal of Cognitive Neuroscience.

[62]  S. Rauch,et al.  Masked Presentations of Emotional Facial Expressions Modulate Amygdala Activity without Explicit Knowledge , 1998, The Journal of Neuroscience.

[63]  Marlene Behrmann,et al.  Monocular Advantage for Face Perception Implicates Subcortical Mechanisms in Adult Humans , 2014, Journal of Cognitive Neuroscience.

[64]  Leslie G. Ungerleider,et al.  Selective dissociation between core and extended regions of the face processing network in congenital prosopagnosia. , 2014, Cerebral cortex.

[65]  D. Sparks,et al.  Population coding of saccadic eye movements by neurons in the superior colliculus , 1988, Nature.

[66]  P. Whalen Fear, Vigilance, and Ambiguity , 1998 .

[67]  Nicholas B. Turk-Browne,et al.  Face-Specific Resting Functional Connectivity between the Fusiform Gyrus and Posterior Superior Temporal Sulcus , 2010, Front. Hum. Neurosci..

[68]  E. Rolls,et al.  Neurons in the amygdala of the monkey with responses selective for faces , 1985, Behavioural Brain Research.

[69]  Andreas Meyer-Lindenberg,et al.  Evidence for a general face salience signal in human amygdala , 2011, NeuroImage.

[70]  Werner Strik,et al.  The effect of appraisal level on processing of emotional prosody in meaningless speech , 2008, NeuroImage.

[71]  R. Desimone Face-Selective Cells in the Temporal Cortex of Monkeys , 1991, Journal of Cognitive Neuroscience.

[72]  Karl J. Friston,et al.  Psychophysiological and Modulatory Interactions in Neuroimaging , 1997, NeuroImage.

[73]  K M Gothard,et al.  Neural responses to facial expression and face identity in the monkey amygdala. , 2007, Journal of neurophysiology.

[74]  D. Zald The human amygdala and the emotional evaluation of sensory stimuli , 2003, Brain Research Reviews.

[75]  M. Giese,et al.  Norm-based face encoding by single neurons in the monkey inferotemporal cortex , 2006, Nature.

[76]  D. Amaral,et al.  The organization of projections from the amygdala to visual cortical areas TE and V1 in the macaque monkey , 2005, The Journal of comparative neurology.

[77]  G. Glover,et al.  Self‐navigated spiral fMRI: Interleaved versus single‐shot , 1998, Magnetic resonance in medicine.

[78]  D. Schacter,et al.  Prefrontal Contributions to Executive Control: fMRI Evidence for Functional Distinctions within Lateral Prefrontal Cortex , 2001, NeuroImage.

[79]  Declan G. M. Murphy,et al.  The anatomy of extended limbic pathways in Asperger syndrome: A preliminary diffusion tensor imaging tractography study , 2009, NeuroImage.

[80]  Russell A. Poldrack,et al.  Large-scale automated synthesis of human functional neuroimaging data , 2011, Nature Methods.

[81]  A. Ishai,et al.  Effective connectivity within the distributed cortical network for face perception. , 2007, Cerebral cortex.

[82]  D. Amaral,et al.  Topographic organization of projections from the amygdala to the visual cortex in the macaque monkey , 2003, Neuroscience.

[83]  Peter J. Lang,et al.  Parallel amygdala and inferotemporal activation reflect emotional intensity and fear relevance , 2005, NeuroImage.

[84]  Pawan Sinha,et al.  Role of color in face recognition , 2010 .

[85]  N. Kanwisher,et al.  PSYCHOLOGICAL SCIENCE Research Article Visual Recognition As Soon as You Know It Is There, You Know What It Is , 2022 .

[86]  Bruce Fischl,et al.  Connectivity-based segmentation of human amygdala nuclei using probabilistic tractography , 2011, NeuroImage.

[87]  P. Holland,et al.  Amygdala circuitry in attentional and representational processes , 1999, Trends in Cognitive Sciences.

[88]  R. Matthew Hutchison,et al.  Resting-state fMRI reveals functional connectivity between face-selective perirhinal cortex and the fusiform face area related to face inversion , 2014, NeuroImage.

[89]  Beatriz Luna,et al.  Visual category-selectivity for faces, places and objects emerges along different developmental trajectories. , 2007, Developmental science.

[90]  Alexander Todorov,et al.  Nonlinear Amygdala Response to Face Trustworthiness: Contributions of High and Low Spatial Frequency Information , 2009, Journal of Cognitive Neuroscience.

[91]  Per B Sederberg,et al.  Scene Representations in Parahippocampal Cortex Depend on Temporal Context , 2012, The Journal of Neuroscience.

[92]  Cindy M. Bukach,et al.  Beyond faces and modularity: the power of an expertise framework , 2006, Trends in Cognitive Sciences.

[93]  T. Allison,et al.  Human extrastriate visual cortex and the perception of faces, words, numbers, and colors. , 1994, Cerebral cortex.

[94]  Mark H. Johnson,et al.  Oxford Handbook of Face Perception , 2011 .

[95]  G. Rhodes,et al.  The dynamics of visual adaptation to faces. , 2005, Proceedings. Biological sciences.

[96]  Edward E. Smith,et al.  Temporal dynamics of brain activation during a working memory task , 1997, Nature.

[97]  Paul J. Whalen,et al.  The uncertainty of it all , 2007, Trends in Cognitive Sciences.

[98]  Doris Y. Tsao,et al.  What's so special about the average face? , 2006, Trends in Cognitive Sciences.

[99]  Stephan Eliez,et al.  Amygdalar activation associated with positive and negative facial expressions , 2002, Neuroreport.

[100]  Marlene Behrmann,et al.  The nature of face representations in subcortical regions , 2014, Neuropsychologia.

[101]  Keiko Usui,et al.  Temporal Profile of Amygdala Gamma Oscillations in Response to Faces , 2012, Journal of Cognitive Neuroscience.

[102]  J. Haxby,et al.  Distributed Neural Systems for Face Perception , 2011 .

[103]  James M. Kilner,et al.  Brain systems for assessing facial attractiveness , 2007, Neuropsychologia.

[104]  A Belger,et al.  Abnormal Neural Activation to Faces in the Parents of Children with Autism. , 2015, Cerebral cortex.

[105]  G. Rhodes,et al.  Adaptive norm-based coding of facial identity , 2006, Vision Research.

[106]  E. Lauterbach The Amygdala: Neurobiological Aspects of Emotion, Memory, and Mental Dysfunction , 1993 .

[107]  Nicolas Davidenko,et al.  Face‐likeness and image variability drive responses in human face‐selective ventral regions , 2012, Human brain mapping.

[108]  L. Goldstein The Amygdala: Neurobiological Aspects of Emotion, Memory, and Mental Dysfunction , 1992, The Yale Journal of Biology and Medicine.

[109]  Jasmin Cloutier,et al.  The perceptual determinants of person construal: reopening the social-cognitive toolbox. , 2005, Journal of personality and social psychology.

[110]  Rodrigo Quian Quiroga,et al.  Human medial temporal lobe neurons respond preferentially to personally relevant images , 2009, Proceedings of the National Academy of Sciences.

[111]  E. Rolls Neurons in the cortex of the temporal lobe and in the amygdala of the monkey with responses selective for faces. , 1984, Human neurobiology.

[112]  R. J Dolan,et al.  Common and distinct neural responses during direct and incidental processing of multiple facial emotions , 2003, NeuroImage.

[113]  M. Posner,et al.  The attention system of the human brain. , 1990, Annual review of neuroscience.

[114]  Alexander Todorov,et al.  Modeling Social Perception of Faces [Social Sciences] , 2011, IEEE Signal Processing Magazine.

[115]  D. Amaral,et al.  Synaptic organization of projections from the amygdala to visual cortical areas TE and V1 in the macaque monkey , 2005, The Journal of comparative neurology.

[116]  J. Sprague,et al.  The role of the superior colliculus in facilitating visual attention and form perception. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[117]  D. Sparks The brainstem control of saccadic eye movements , 2002, Nature Reviews Neuroscience.

[118]  Alexander Todorov,et al.  The role of the amygdala in face perception and evaluation , 2011, Motivation and emotion.

[119]  Leslie G. Ungerleider,et al.  An area specialized for spatial working memory in human frontal cortex. , 1998, Science.

[120]  T. Allison,et al.  Differential Sensitivity of Human Visual Cortex to Faces, Letterstrings, and Textures: A Functional Magnetic Resonance Imaging Study , 1996, The Journal of Neuroscience.

[121]  Doris Y. Tsao,et al.  Taking Apart the Neural Machinery of Face Processing , 2011 .

[122]  T. Hendler,et al.  Feeling or Features Different Sensitivity to Emotion in High-Order Visual Cortex and Amygdala , 2001, Neuron.

[123]  C. Gross,et al.  Representations of faces and body parts in macaque temporal cortex: a functional MRI study. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[124]  Karine Sergerie,et al.  The role of the amygdala in emotional processing: A quantitative meta-analysis of functional neuroimaging studies , 2008, Neuroscience & Biobehavioral Reviews.

[125]  Nancy Kanwisher,et al.  The Functional Architecture of the Face System: integrating Evidence from fMRI and Patient Studies , 2011 .

[126]  N. Kanwisher,et al.  The Fusiform Face Area: A Module in Human Extrastriate Cortex Specialized for Face Perception , 1997, The Journal of Neuroscience.

[127]  R. Adolphs,et al.  Annals of the New York Academy of Sciences What Does the Amygdala Contribute to Social Cognition? , 2022 .

[128]  R. Desimone,et al.  Visual properties of neurons in a polysensory area in superior temporal sulcus of the macaque. , 1981, Journal of neurophysiology.

[129]  Nikos K. Logothetis,et al.  Facial-Expression and Gaze-Selective Responses in the Monkey Amygdala , 2007, Current Biology.

[130]  Harry Wechsler,et al.  The FERET database and evaluation procedure for face-recognition algorithms , 1998, Image Vis. Comput..

[131]  Krista A. Ehinger,et al.  SUN database: Large-scale scene recognition from abbey to zoo , 2010, 2010 IEEE Computer Society Conference on Computer Vision and Pattern Recognition.

[132]  Katsuki Nakamura,et al.  Responses of single neurons in monkey amygdala to facial and vocal emotions. , 2007, Journal of neurophysiology.

[133]  Francesco Fera,et al.  The Amygdala Response to Emotional Stimuli: A Comparison of Faces and Scenes , 2002, NeuroImage.

[134]  Timothy E. J. Behrens,et al.  Deep and Superficial Amygdala Nuclei Projections Revealed In Vivo by Probabilistic Tractography , 2011, The Journal of Neuroscience.

[135]  Thomas Vetter,et al.  Portraits made to measure: manipulating social judgments about individuals with a statistical face model. , 2009, Journal of vision.

[136]  A. Todorov,et al.  The functional basis of face evaluation , 2008, Proceedings of the National Academy of Sciences.

[137]  N. Logothetis,et al.  Functional imaging of the monkey brain , 1999, Nature Neuroscience.

[138]  T. Johnstone,et al.  Human amygdala responses during presentation of happy and neutral faces: correlations with state anxiety , 2004, Biological Psychiatry.

[139]  William A. Cunningham,et al.  PSYCHOLOGICAL SCIENCE Research Article Affective Flexibility Evaluative Processing Goals Shape Amygdala Activity , 2022 .

[140]  Christopher J Rennie,et al.  Mode of Functional Connectivity in Amygdala Pathways Dissociates Level of Awareness for Signals of Fear , 2006, The Journal of Neuroscience.

[141]  J. Grafman,et al.  The Human Amygdala: An Evolved System for Relevance Detection , 2003, Reviews in the neurosciences.

[142]  Marlene Behrmann,et al.  Unraveling the distributed neural code of facial identity through spatiotemporal pattern analysis , 2011, Proceedings of the National Academy of Sciences.

[143]  P. Sinha,et al.  Contribution of Color to Face Recognition , 2002, Perception.

[144]  C. Koch,et al.  Invariant visual representation by single neurons in the human brain , 2005, Nature.

[145]  Andrew D. Engell,et al.  Probabilistic atlases for face and biological motion perception: An analysis of their reliability and overlap , 2013, NeuroImage.

[146]  N. Kanwisher,et al.  Domain specificity in visual cortex. , 2006, Cerebral cortex.

[147]  Rolls Et Neurons in the cortex of the temporal lobe and in the amygdala of the monkey with responses selective for faces. , 1984 .

[148]  O. Pascalis,et al.  The Origins of Face Processing in Humans: Phylogeny and Ontogeny , 2009, Perspectives on psychological science : a journal of the Association for Psychological Science.

[149]  Doris Y. Tsao,et al.  A Cortical Region Consisting Entirely of Face-Selective Cells , 2006, Science.

[150]  Malcolm J Avison,et al.  Regional brain response to faces of humans and dogs. , 2004, Brain research. Cognitive brain research.

[151]  C. Macrae,et al.  A face with a cue: exploring the inevitability of person categorization , 2007 .

[152]  Timothy J. Andrews,et al.  Response of face-selective brain regions to trustworthiness and gender of faces , 2012, Neuropsychologia.

[153]  C. Gross,et al.  Neural representations of faces and body parts in macaque and human cortex: a comparative FMRI study. , 2009, Journal of neurophysiology.

[154]  Lee Friedman,et al.  Report on a multicenter fMRI quality assurance protocol , 2006, Journal of magnetic resonance imaging : JMRI.

[155]  Andrew D. Engell,et al.  Facial expression and gaze-direction in human superior temporal sulcus , 2007, Neuropsychologia.

[156]  Marc G. Berman,et al.  Evaluating functional localizers: The case of the FFA , 2010, NeuroImage.

[157]  A. Ishai,et al.  Distributed and Overlapping Representations of Faces and Objects in Ventral Temporal Cortex , 2001, Science.

[158]  T. Allison,et al.  Face-Specific Processing in the Human Fusiform Gyrus , 1997, Journal of Cognitive Neuroscience.

[159]  P. Downing,et al.  Human Neuroscience , 2022 .

[160]  J. Moake,et al.  This article has been cited by other articles , 2003 .

[161]  Jerome Kagan,et al.  Differential amygdalar response to novel versus newly familiar neutral faces: a functional MRI probe developed for studying inhibited temperament , 2003, Biological Psychiatry.

[162]  Norihiro Sadato,et al.  Neural substrates participating in acquisition of facial familiarity: an fMRI study , 2003, NeuroImage.

[163]  Conny F. Schmidt,et al.  Face perception is mediated by a distributed cortical network , 2005, Brain Research Bulletin.

[164]  Paul J Whalen,et al.  Prior experience as a stimulus category confound: an example using facial expressions of emotion. , 2006, Social cognitive and affective neuroscience.

[165]  Hyeonjoon Moon,et al.  The FERET evaluation methodology for face-recognition algorithms , 1997, Proceedings of IEEE Computer Society Conference on Computer Vision and Pattern Recognition.

[166]  Mark H. Johnson Subcortical face processing , 2005, Nature Reviews Neuroscience.

[167]  H. Wilson,et al.  fMRI evidence for the neural representation of faces , 2005, Nature Neuroscience.

[168]  Hironobu Fujiwara,et al.  Neuroimaging studies of social cognition in schizophrenia , 2013, Psychiatry and clinical neurosciences.

[169]  Bryan R. Conroy,et al.  A Common, High-Dimensional Model of the Representational Space in Human Ventral Temporal Cortex , 2011, Neuron.