Differential sensitivity of human visual cortex to faces, letterstrings, and textures: a functional magnetic resonance imaging study

“Twelve normal right-handed subjects (Edinburgh Handedness Inventory range 70 –100) (Oldfield, 1970) without a previous neurological or psychiatric history participated in this study.”

“There are advocates of a unitary system (Damasio et al., 1982; Critchley, 1986; Biederman, 1987), two subsystems dealing with faces and words (Farah, 1990), two subsystems dealing with living and nonliving objects (Sheridan and Humphreys, 1993; Newcombe et al.”

“These results are consistent with the neuropsychology literature (for review, see Rhodes, 1993) demonstrating a right hemisphere advantage for face recognition and with previous PET (Horwitz et al., 1992; Haxby et al., 1995) and scalp-recorded evoked potential (Bentin et al., 1996) studies.”

“Semantic processing of auditorily presented words also activated regions of the temporal lobe distant from ventral cortex (Petersen et al., 1988; Wise et al., 1991; Démonet et al., 1992).”

“Malach et al. (1995) compared the fMRI activation produced by objects (including faces) with the activation produced by textures.”

“It is activated by colors, shapes, and scrambled faces (Corbetta et al., 1991; Clark et al., 1995; Puce et al., 1995a), but not specifically by faces (Table 1) (see also Clark et al., 1995; Puce et al., 1995a).”

“Little activation and no letterstring-specific activation of left superior and middle temporal cortex was seen, regions previously activated in PET studies involving phonological processing or word reading (Petersen et al., 1989; Démonet et al., 1992; Howard et al., 1992; Price et al., 1994).”

“Little activation and no letterstring-specific activation of left superior and middle temporal cortex was seen, regions previously activated in PET studies involving phonological processing or word reading (Petersen et al., 1989; Démonet et al., 1992; Howard et al., 1992; Price et al., 1994).”

“Previous PET studies have consistently shown face activation of this region (Sergent et al., 1992; Haxby et al., 1994, 1995).”

“Regions engaged in processing of color and form (for review, see Maunsell and Newsome, 1987), faces (Gross, 1992; Perrett et al., 1992), facial expression (for review, see Rolls, 1992), motion (for review, see Albright, 1993), and “biological motion” (Oram and Perrett, 1994) have been described.”

“Semantic processing of auditorily presented words also activated regions of the temporal lobe distant from ventral cortex (Petersen et al., 1988; Wise et al., 1991; Démonet et al., 1992).”

“The region activated by unpronounceable nonwords here partially overlapped, but was inferior and lateral to a medial extrastriate region activated by words and pronounceable pseudowords (but not by false fonts or unpronounceable nonwords ) in the study of Petersen et al. (1990).”

“, concrete nouns or unpronounceable nonwords), whereas longer-latency potentials recorded from anterior regions of the inferior temporal lobe were highly sensitive to word type (Nobre et al., 1994).”

“The t-maps computed for each stimulus alternation order were further analyzed using two methods: (1) A " split-t " map (Puce et al., 1995b) was generated for each subject in which only those voxels whose associated t values exceeded Ϯ 1.5 for both stimulus alternation orders were retained.”

“Little activation and no letterstring-specific activation of left superior and middle temporal cortex was seen, regions previously activated in PET studies involving phonological processing or word reading (Petersen et al., 1989; Démonet et al., 1992; Howard et al., 1992; Price et al., 1994).”

“We predicted from electrophysiological studies (Allison et al., 1994a; Nobre et al., 1994) that activation of ventral surface cortex by letterstrings would be medial to that for faces.”

“…with the present study, the lesion evidence suggests that prosopagnosia (for review, see De Renzi et al., 1994) and pure alexia (for review, see Damasio and Damasio, 1983) are forms of visual agnosia preferentially involving lesions of the right and left occipitotemporal regions, respectively.”

“This processing occurs within two separate but interconnected pathways, a ventral pathway dealing with object recognition, and a dorsal pathway dealing with motion, spatial relationships, and visually guided movement (Ungerleider and Mishkin, 1982; Goodale and Milner, 1992; Ungerleider, 1995) (however, see Merigan and…”

“The collateral sulcus contains portions of areas V2, ventral V3 (VP), and V4 (Clarke and Miklossy, 1990; Clarke et al., 1995; Shipp et al., 1995).”

“, 1992), facial expression (for review, see Rolls, 1992), motion (for review, see Albright, 1993), and “biological motion” (Oram and Perrett, 1994) have been described.”

“Consistent with the present study, the lesion evidence suggests that prosopagnosia (for review, see De Renzi et al., 1994) and pure alexia (for review, see Damasio and Damasio, 1983) are forms of visual agnosia preferentially involving lesions of the right and left occipitotemporal regions,…”

“Intracranial recordings also have revealed a left dominant asymmetry (our unpublished observations), and evoked potential studies have shown that a letterstring N180 is larger over the left than the right temporal scalp (Nobre and McCarthy, 1994), complementary to the distribution obtained by Bentin et al.”

“The t-maps computed for each stimulus alternation order were further analyzed using two methods: (1) A " split-t " map (Puce et al., 1995b) was generated for each subject in which only those voxels whose associated t values exceeded Ϯ 1.5 for both stimulus alternation orders were retained.”

“Previous PET studies have consistently shown face activation of this region (Sergent et al., 1992; Haxby et al., 1994, 1995).”

“Previous PET studies have consistently shown face activation of this region (Sergent et al., 1992; Haxby et al., 1994, 1995).”

“Previous PET studies have consistently shown face activation of this region (Sergent et al., 1992; Haxby et al., 1994, 1995).”

“These results are consistent with the neuropsychology literature (for review, see Rhodes, 1993) demonstrating a right hemisphere advantage for face recognition and with previous PET (Horwitz et al., 1992; Haxby et al., 1995) and scalp-recorded evoked potential (Bentin et al., 1996) studies.”

“Visual cortex located posterio-medial to the face, letterstring, and numberstring areas was preferentially activated by colored checkerboards (Allison et al., 1993).”

“The frequency analysis has been described in detail (McCarthy et al., 1995; Puce et al., 1995a).”

“It has been argued that faces are an important category of objects for which rapid recognition requires specialized processing (Young and Bruce, 1991; Newcombe et al., 1994).”

“We suspect that our face activation is somewhat more lateralized to the right hemisphere than that seen in Courtney et al. (1997) and Puce et al. (1995, 1996), because our use of objects as comparison stimuli allowed us to effectively subtract out the contribution of general object processing to…”

“For example, Puce et al. (1996) found bilateral fusiform gyrus activation for faces, but with more extensive activation in the right hemisphere.”

“…in patients with the syndrome of prosopagnosia indicate that the homologous areas specialized for face and object identification are located predominantly within the mid-portion of the fusiform gyrus (BA 37 and BA 20) (Damasio, 1985; Sergent et al., 1992; Allison et al., 1994; Puce et al., 1996).”

“Excluded also were studies employing emotional words or pictures of facial expressions, as these stimuli involve highly specific neurophysiological processing (Britton et al., 2006; Horovitz et al., 2004; Puce et al., 1996), stimulus assessment, and specific procedures not employed in most affective ERP studies (Batty…”

“This area has been found to activate more for letter strings than face”

“…faces per se began in the mid-1990s, with demonstrations of fusiform regions that responded more strongly to faces than to letter strings and textures (Puce et al. 1996), flowers (McCarthy et al. 1997a), and other stimuli, including mixed everyday objects, houses, and hands (Kanwisher et al. 1997).”

“…science model’’ shared by many anthropologists, sociologists, and some psychologists (see Barkow et al., 1992; and chapter 2 in Pinker, 2002), but also some neuroscientists who view the cortical surface as ‘‘largely equipotential and free of domain-specific structure’’ (Quartz and Sejnowski, 1997).”

“…(either pronounceable pseudowords or consonant strings) relative to non-alphabetic visual stimuli such as false fonts or faces (Price et al., 1996a; Puce et al., 1996), or by similar contrasts using real words (Beauregard et al., 1997; Wagner et al., 1998; Fiez et al., 1999; Kiehl et al., 1999;…”

“…strong activations to specific object categories such as faces (Allison et al. 1994, Ishai et al. 1999, Kanwisher et al. 1997, Levy et al. 2001, Puce et al. 1995), animals (Chao et al. 1999, Martin et al. 1996), body parts (Downing et al. 2001, Grossman & Blake 2002), tools (Beauchamp et al.…”

“Each consisted of two subtasks (A and B) that alternated throughout each imaging run as described previously (Puce et al., 1995, 1996).”

“A region of ventral occipito-temporal cortex called the fusiform face area, or FFA (Kanwisher, McDermott, & Chun, 1997), responds strongly when subjects view photographs of faces, but only weakly when they view other classes of stimuli such as familiar objects or complex scenes (McCarthy, Puce, Gore, & Allison, 1997;…”

“, but activated at least twice as strongly for faces as for a wide variety of nonface stimuli, including letter string”

“The fusiform gyrus was selected as the primary cortical ROI due to reported consistent involvement of this structure during face processing tasks (Allison et al., 1994; Farah, 1996; Haxby et al., 1996; Puce et al., 1996; Kanwisher et al., 1999).”

“…also localized primarily to the lateral fusiform gyrus (Kanwisher et al., 1997; McCarthy et al., 1997; Halgren et al., 1999). fMRI studies suggest that cortex within the midfusiform and occipitotemporal sulci is also activated by faces (Clark et al., 1996; Puce et al., 1996; Halgren et al., 1999).”

“Puce et al. (1996) report single-subject activations during passive presentation of visual words, faces, and textures.”

“…faces and objects in the regions of the inferior occipital gyrus and middle fusiform gyrus (e.g., Kanwisher et al., 1997; Sergent et al., 1992; Puce et al., 1996; see Haxby et al., 2000), and in the left fusiform gyrus for letterstrings compared to numbers and shapes (the " letter area, "…”

“While the essential neural structures of face perception are not absolutely clear, the principal components of the human neural system for face perception include the inferior occipital gyrus (IOG), the superior temporal sulcus (STS), and the anterior fusiform gyrus or ‘fusiform face area’ (FFA; Haxby et al., 2002; Ka…”

“…fMRI revealed more of the specificity of these cortical regions for faces, with demonstrations of fusiform regions that responded more strongly to faces than to letter strings and textures (Puce et al 1996), flowers (McCarthy et al 1997), everyday objects, houses, and hands (Kanwisher et al 1997).”

“In literate people, the VWFA is always located next to a cortical region that is selective for faces: the left fusiform face area (left FFA) 62–64 .”

“For instance, distinct brain areas respond preferentially to letter strings, chairs, buildings or face”

“…of letter- and word-related regions averaged across six different studies (Fiez and Petersen, 1998; Mechelli et al., 2000; Moore and Price, 1999; Puce et al., 1996; Rumsey et al., 1997; Tagamets et al., 2000; Uchida et al., 1999). d Talairach coordinates of face-related regions obtained in…”

“These studies indicate that in human adults, particular areas of ventral and lateral occipital cortex are preferentially activated by faces (Kanwisher, McDermott, & Chun, 1997; Puce, Allison, Gore, & McCarthy, 1995; Puce, Allison, Asgari, Gore, & McCarthy, 1996 ) and that face processing is impaired if they are damage…”

“Previous work has demonstrated that within occipito-temporal regions, perceptual features of objects and faces (e.g., Puce et al., 1996) and of different categories of objects (e.g., Chao, Haxby, & Martin, 1999) are represented in distinguishable cortical areas.”

“An unexpected finding was the rise in fMRI signal before the onset of the conscious experience, the opposite of the normal delayed response to visual stimulation found in fMRI experiment”

“The OT-ECD is far inferior and posterior to a third face-selective area seen with fMRI in the region of the superior temporal sulcus ( Puce et al., 1995, 1996, 1998; Halgren et al., 1999).”

“…functional imaging studies have linked the fusiform and lingual gyri with perceptual, pre-semantic analysis of faces (Haxby et al., 1994; Grady et al., 1996; Puce et al., 1996; Dolan et al. 1997), even when they were compared with complex visual stimuli such as houses (Kanwisher et al., 1997).”

“10) and fMRI (Puce et al., 1996, 1997, 1998) studies demonstrate that the ventral and lateral face areas are separated by cortex (the superior portion of the inferior temporal gyrus) that is relatively uninvolved in face processing.”

“The lateral fusiform gyrus subserves selective activation to faces (110, 111), and because of this area’s specificity and the consistency with which it has been linked to facial recognition, it has been dubbed the “fusiform face area.”

“The mean PSC across the 1983; Puce et al. 1996).”

“sulcus dividing the fusiform into lateral and medial partitions first appeared in modern writing by Puce and colleagues (Puce et al., 1996) and was later named the MFS (Allison et al.”

“Thus, areas in the occipitotemporal region appear to be specialized for the processing of colour, texture, and form differences of objects (e.g. Puce et al., 1996; Price et al., 1996; Malach et al., 1995; Kanwisher et al., 1996).”

“[27,21,18,16,19]) have 200 ms (N200) recorded from ventral occipitotemporal located an area in the fusiform gyrus that is strongly cortex was unaffected by the familiarity of faces and activated when faces are presented, but not in response to showed no habituation, suggesting that this potential houses, scrambled fac…”

“, 2000, 2002), but also relative to other categories of visual objects such as faces or houses (Gauthier et al., 2000; Hasson et al., 2002; Puce et al., 1996).”

“In the left hemisphere the centroid of letter-stringspecific N200s is near the occipitotemporal sulcus, which Puce et al. (Puce et al., 1996) found to be activated more by letter-strings than by faces or textures.”

“, lateral to the fusiform face responses (as previously reported in [80]) and overlapping with the most anterior part of the lateral object-responsive cortex (Fig 2).”

“Thus, the VWFA is activated by alphabetic strings relative to fixation but also relative to complex non-alphabetic stimuli such as false fonts, faces or geometrical patterns (e.g. Price et al., 1996b; Puce et al., 1996; Cohen et al., 2002).”

“…Stimulias well as other neuroimaging studies showing selective Objects-From-Motion (OFM) Experimentactivation to various object categories (Haxby et al., Epochs of visual stimulation 40 s long were alternated with 16 or 1996; Martin et al., 1996; Puce et al., 1996; Kanwisher et 20 s blank epochs.”

“Core brain regions of face processing are located in inferior occipital gyrus (occipital face area, OFA, Puce et al., 1996), lateral fusiform gyrus (fusiform face area, FFA, Kanwisher et al., 1997), and posterior superior temporal sulcus (pSTS, Hoffman and Haxby, 2000).”

“Pitcher et al. (2009) 45, i78, i6 Faces > Objects 15/15 Puce et al. (1996) 36, i66, i17 Faces > Letter strings Not reported”

“…gyrus, or it is possible that it could be replaced by external sources of motivation such as the kind of supervised learning that one can encounter for letters and words and which appears to lead to similar specialization for letters in other parts of the visual system (e.g., Puce et al., 1996).”

“Unlike the fusiform activation, this activation was strongly right-lateralized, consistent with previous studies (Puce et al., 1996).”

“…a cluster of brain regions in the ventral visual stream known as the lateral occipital complex (LOC; e.g. Kohler et al., 1995; Malach et al., 1995; Puce et al., 1996, 1999; Kanwisher et al., 1997; Allison et al., 1999; Haxby et al., 1999; Ishai et al., 1999; Doniger et al., 2000, 2001, 2002;…”

“As we were still puzzling over our initial results, others published reports on a specialized area in the fusiform gyrus that responded more during face perception tasks than during perception of other objects (Kanwisher et al., 1997; McCarthy et al., 1997; Puce et al., 2006).”

“These results are consistent with functional neuroimaging studies (Kanwisher et al., 1996, 1997; Puce et al., 1996; Hasson et al., 2002) and several ERP/MEG studies (Botzel et al., 1995; Bentin et al., 1996; Halgren et al., 2000; Liu et al., 2000; Rossion et al., 2003), where face-sensitive…”

“Several brain imaging and electrophysiological recording studies in humans have reported discrete cortical regions in the posterior ventral temporal cortex that respond preferentially to faces (Haxby et al., 1999; Kanwisher, McDermott, & Chun, 1997; McCarthy, Puce, Gore, & Allison, 1997; Clark et al., 1996; Puce, Alli…”

“Also consistent with this finding are the previous studies that have reported occipitotemporal and lateral occipital sensitivity to letter strings [Puce et al., 1996] and written words [Baker et al., 2007].”

“during viewing of faces than a variety of compar- 103 ison stimuli (Puce et al., 1996, 1995), collectively 104 suggest that there are cortical areas devoted pref- 105 erentially to the processing of faces.”

“…the fusiform gyrus are more active when adults passively view faces than when they view other objects such as hands or chairs (Puce et al 1995; Puce et al 1996; Kanwisher et al 1997) and the pattern of activation over the ventral temporal cortex differs for faces compared to other objects…”

“If it turns out that nonpreferred some classes of objects represented in localized regions responses carry as much information as preferred reof extrastriate cortex specialized for representing prisponses, that will call into question both the standard marily that class of object (Downing et al., 2001; Epstein techniq…”

“An expertise account would also have difficulty explaining why letters, which represent another highly overlearned visual category, activate different brain regions than faces (Puce et al., 1996).”

“Many previous studies have shown that face stimuli produce fusiform activity (Clark et al., 1998; Dolan et al., 1997; Grady et al., 1995; Halgren et al., 1999; Haxby et al., 1999, 1996, 1991; Kanwisher et al., 1997; McCarthy et al., 1997; Puce et al., 1996; Sergent et al., 1992).”

“Many regions important for face, facial feature and object perception, including the fusiform, superior and middle temporal gyri (Puce et al., 1996, 1998), were activated by covert and overt expressions of fear and disgust (Tables 1 – 4).”

“The sensitivity of these cortical areas to faces was confirmed by fMRI ( Puce, Allison, Asgari, Gore, & McCarthy, 1996 ), and these studies were the catalyst for a large number of subsequent functional neuroimaging studies on face processing.”

“1,A, B, andD), which is near regions observed in other studies using block letter stimuli (Puce et al. 1996).”

“A wide range of noninvasive techniques, including PET, fMRI, ERP, and MEG have shown that FFA responds selectively to images of faces, relative to a wide variety of control objects (1-7).”