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).”

“Activation of these posterior ventral occipitotemporal regions in facial perception tasks is frequently reported in recent functional brain imaging studies (Haxby et al., 1991, 1994; Kapur et al., 1995; Puce et al., 1996; Gorno Tempini et al., 1998; Nakamura et al., 2000).”

“…Dehaene, 1995; Helenuis et al., 1999; Liu and Perfetti, 2003; Martı́n-Loeches et al., 2004; Mondini et al., 2008; Moscoso del Prado Martı́n et al., 2006; Puce et al., 1996; Rossion et al., 2003; Salmelin et al., 1996; Schendan et al., 1998; Spironelli and Angrilli, 2007; Tarkiainen et al., 1999).”

“This region is more activated to faces compared with other objects or body parts (Kanwisher et al., 1997; Puce et al., 1996).”

“…activity in ventral visual cortex is consistent with reports showing robust and reproducible neural activation in left occipito-temporal cortex in response to words and letters (Polk et al., 2002; Cohen et al., 2000; Polk & Farah, 1998; Puce et al., 1996; Petersen, Fox, Snyder, & Raichle, 1990).”

“, 2002), face processing (Puce et al., 1996), pre-lexical processing of words (Nobre et al.”

“As shown in Figure 1B, ROIs defined in this study are likely to include previously described headless body-selective (Downing et al., 2001; Peelen and Downing, 2005) and face-selective (Puce et al., 1996; Kanwisher et al., 1997) areas.”

“OFAwas defined by the peak voxel of activity in the inferior occipital gyrus (Calder and Young, 2005; Gauthier et al., 2000; Haxby et al., 2000; Puce et al., 1996), inferior and more medially located than the activity for LO.”

“Cortical sites encompassing the current region of interest also correspond to reports of functional response to objects including chairs, faces and letter strings (Ishai et al., 2000; Puce et al., 1996 ).”

“On the other hand, the neural basis of the perception of printed letters has been frequently studied (Allison et al., 1994; Behrmann et al., 1998; Puce et al., 1996; Tarkiainen et al., 1999; Raij et al., 2000; Polk et al., 2002; Cohen et al., 2003), with results emphasizing the importance of the…”

“…to the perception of simple features, such as colour (Lueck et al., 1989) and motion (Watson et al, 1993; McCarthy et al., 1995; Tootell et al., 1995), and to the perception of complex objects and faces A. C. Nobre et al. (Malach et al., 1995; Puce et al., 1995, 1996; Kanwisher et al., 1997).”

“Last, the distinction between face-selective and word-selective sites has also been noted in studies using intracranial recordings used to monitor pharmacologically resistant epilepsy (Allison, Puce, Spencer, & McCarthy, 1999; McCarthy, Puce, Belger, & Allison, 1999; Puce et al., 1995, 1996).”

“…individuals (Gaz-zaniga et al. 1965; Newcombe et al. 1989; Rizzolatti et al. 1971; Farah 1990; early neuroimaging evidence: Petersen et al. 1988; Puce et al. 1996; EEG: Salmelin et al. 1996; Tarkiainen et al. 1999; Rossion et al. 2003; lesion studies: Déjerine 1891; Hillis et al. 2001; Hillis…”

“2), a region of the visual ventral stream specific to face processing [16,21,23,25,40].”

“It has been consistently shown that letter string presentation elicits greater activity in the fusiform gyri and inferior occipital cortices compared to nonletter stimuli (Petersen et al., 1990; Puce et al., 1996; Uchida et al., 1999).”

“The fusiform gyrus contains both a brain region critical for face processing (the fusiform face area [FFA] [24, 25]) and an abutting region involved in body processing (the fusiform body area [FBA] [26, 27]).”

“Critchley et al, 2000 ; Schultz et al, 2000 ) that is specialized for face perception (Kanwisher, McDermott, & Chun, 1997 ; Puce, Allison, Asgari, Gore, & McCarthy, 1996 ), has been observed in participants with autism in comparison to typically developing participants.”

“Given the well-established laterality of face processing to the right hemisphere (Puce et al., 1996; De Renzi, 1997; Kanwisher et al., 1997) and also to maximize comparability between the PPA and FFA, we first focused our analyses to the right hemisphere ROIs (see Epstein et al., 2003).”

“…those associated with perceptual processing and sensory recapitulation (e.g., occipital cortex) (e.g., Allison et al., 1994; Prince et al., 2009; Puce et al., 1996; Vaidya et al., 2002), as well as memory-related regions within the bilateral hippocampus and PHG and decision-making regions…”

“, 2012) of the left hemisphere [average MNI coordinates ( SE): 45(1), 66(2), 14(2)] was determined as responding more intensely to RLs than to FFs and Fourier noise images (Puce et al., 1996; Thesen et al., 2012).”

“The cortical network underlying face perception is well known and includes the fusiform face area (FFA; Kanwisher et al., 1997), the occipital face area (Puce et al., 1996; Gauthier et al., 2000), the superior temporal sulcus (STS) and the amygdala (AMG; Haxby et al., 2000).”

“These include general language processing [17,18], semantic processing [7–9,19,20,34], phonological retrieval [4,25,36], face processing [6,10,14,15,26] and object recognition [22,24,27,28,37].”

“The involvement of lingual gyrus in face perception has been previously reported (Puce et al., 1996).”

“Left inferior parietal activation is often reported in the same studies that report left fusiform/extrastriate activation for letters (James et al., 2005; Jessen et al., 1999; Longcamp et al., 2003; Pernet et al., 2004; Price et al., 1996; Puce et al., 1996; Sergent et al., 1992; Tagamets et al., 2000).”

“…“letterbox area” of the brain (Cohen and Dehaene, 2004; Dehaene, 2009; Dehaene and Cohen, 2011), is one of the subdivisions in the ventral occipitotemporal region involved in object recognition (Puce et al., 1996; Kanwisher et al., 1997; Ishai et al., 1999; Haxby et al., 2001; Malach et al., 2002).”

“An additional right middle temporal activation for faces has been observed also in the intracranial investigations mentioned above [Puce et al., 1996] although at a later period in time.”

“First, letters and words are processed in distinct areas from faces and visual textures; specifically, words and letter strings lead to more activation in the postlateral fusiform gyrus, left lateralized (Herbster, Mintun, Nebes, & Becker, 1997; Puce et al., 1996).”

“Written characters were compared with faces and line drawings to reveal category selectivity for visual words in the VWFA [6,12,33,34].”

“For example, in patients who hallucinated in colour, activity was recorded in the fusiform gyrus, an area in V4 corresponding to the colour centre, whereas in the patient who hallucinated in black and white, the activity was outside this region (Puce et al 1996).”

“Although these regions are typically activated in both hemispheres, the right lateralization of the face perception network is well established (De Renzi, 1986; Kanwisher et al., 1997; Puce et al., 1996; Wada and Yamamoto, 2001).”

“Such studies identified the FFA (Kanwisher et al. 1997; McCarthy et al. 1997; Tong et al. 2000), the OFA (Gauthier et al. 2000; Puce et al. 1996), and face selective regions in the pSTS (Allison et al. 2000; Puce et al. 1996).”

“Within these brain areas, different regions are found to respond preferentially to different object categories, including faces (Kanwisher, McDermott, & Chun, 1997; Puce, Allison, Asgari, Gore, & McCarthy, 1996 ), words (Cohen et al., 2000), letters (James, James, Jobard, Wong, & Gauthier, 2005; Polk et al., 2002), bo…”

“[8–10] and anomalous face recognition [11].”

“the Bilingual Brain”

“These findings suggest that the function of the posterior STS is related to perception of motion of the hands.”

“Hemodynamic [functional magnetic resonance imaging (fMRI) and positron emission tomography] studies of word and letter recognition have also indicated involvement of the left occipitotemporal cortex in orthographic analysis (Petersen et al., 1990; Puce et al., 1996; Pugh et al., 1996; Paulesu et al., 2000; Polk et al.…”

“Several imaging (Hasson et al., 2002; Puce et al.,1996; Sergent et al., 1992), ERP (Kovács et al., 2005; Rossion et al., 2003) and lesion studies of acquired prosopagnosia (Sorger et al., 2007) suggest a right hemisphere advantage in face processing.”

“This is also consistent with earlier fMRI studies using artificial abstract textures, which revealed texture selectivity in the medial ventral visual cortex (Puce et al., 1996; Peuskens et al., 2004).”

“The occipital face area (OFA) was localized based on the same contrast as FFA and restricted to face-selective voxels in the occipital lobe (Puce et al., 1996; Gauthier et al., 2000).”

“…while the pattern was similar for the poses used in the present study, other studies from our lab (Allison, McCarthy, Nobre, Puce, & Belger, 1994b; Puce et al., 1995, 1996) have found that scrambled faces and complex textures also activate medial VOTC regions while faces activated more lateral…”

“Second, the fusiform face area (FFA), a region preferentially activated by faces (Puce et al. 1996; Kanwisher et al. 1997; Yovel et al. 2008), is generally described as showing a preferential lateralization to the right hemisphere (McCarthy et al. 1997; Haxby et al. 1999; Dien 2009), which may…”

“... and pseudoword processing involves a left hemisphere (LH) posterior reading system (see Figure 1) consisting of both ventral and dorsal components.1 The ventral region includes lateral extrastriate areas and a left occipito-temporal area where functional imaging studies show robust activation in word and pseudowor…”

“Electrophysiological (Bentin et al., 1996) and fMRI studies of face processing have shown greater activation at right than at left hemisphere sites, as well as more extensive fusiform gyrus activation in the right hemisphere (Puce et al., 1996; Kanwisher et al., 1997).”

“In human neuroimaging studies, a number of studies have revealed that observation of faces (relative to the other categories of common objects) selectively activates regions in and around the superior temporal sulcus (e.g., Puce et al., 1996; Haxby et al., 1999).”

“A corresponding neural model was proposed by Haxby, Hoffman, and Gobbini (2000) based on a review of single unit and fMRI studies in both humans and monkeys (Hoffman & Haxby, 2000; Puce et al., 1996).”

“PET studies of reading revealed that the left FG and extrastriate cortex were involved in the processing of letter strings (Price et al., 1996; Puce et al., 1996).”

“…the superior temporal sulcus (STS), for face perception has been supported by previous studies in adults by using functional magnetic resonance imaging (fMRI) (Kanwisher et al. 1997; Puce et al. 1996) and in infants by using nearinfrared spectroscopy (NIRS) (Nakato et al. 2009; Otsuka et al. 2007).”

“110 Journal of Cognitive Neuroscience Volume 14, Number 1 ventral visual pathway that may cover a wide range of visual objects, both verbal and nonverbal ( Puce, Allison, Asgari, Gore, & McCarthy, 1996; Ungerleider & Haxby, 1994), it seems unlikely that the visual images of kana are stored exceptionally in a separate…”

“1-Frith et al. (1995); 2-Kapur et al. (1994); 3- Kuriki et al. (1998); 4-Menard et al. (1996); 5-Petersen et al. (1990): 6-Price, Moore, & Frackowiak (1996); 7-Puce et al. (1996); 8-Pugh et al. (1996).”

“Functional magnetic resonance imaging (MRI) has identified areas selective for both faces and scrambled faces in ventral occipito-temporal cortex, the middle occipital gyri, and the superior temporal or lateral occipital sulci (Puce et al. 1996; Clark et al. 1996).”

“…2003; Deen and McCarthy, 2010; Grossman et al ., 2010), the lateral portion of the fusiform gyrus to represent information about faces and bodies (Puce et al ., 1996; Kanwisher et al ., 1997; Schwarzlose et al ., 2005; Kanwisher and Yovel, 2006), the posterior cingulate and precuneus to…”

“, 1999), neuroimagery (Haxby, Ungerleider, Clark, Schouten, Hoffman and Martin, 1999; Ishay, Ungerleider, Martin, Schouten and Haxby, 1999; Puce et al., 1996; Sergent et al., 1992), and neuropsychological (De Renzi, 1986; Farah et al.”

“While it is known that these regions are involved in visual feature perception [45,54,61] and object identification [1,33,66] in young individuals, the pattern of change observed here suggests that age-related decline of right hemisphere regions may be counteracted by an increase in homologous regions of the left hemi…”