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WIREs Cogn Sci
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Functional development of the brain's face‐processing system

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In the first 20 years of life, the human brain undergoes tremendous growth in size, weight, and synaptic connectedness. Over the same time period, a person achieves remarkable transformations in perception, thought, and behavior. One important area of development is face processing ability, or the ability to quickly and accurately extract extensive information about a person's identity, emotional state, attractiveness, intention, and numerous other types of information that are crucial to everyday social interaction and communication. Associating particular brain changes with specific behavioral and intellectual developments has historically been a serious challenge for researchers. Fortunately, modern neuroimaging is dramatically advancing our ability to make associations between morphological and behavioral developments. In this article, we demonstrate how neuroimaging has revolutionized our understanding of the development of face processing ability to show that this essential perceptual and cognitive skill matures consistently yet slowly over the first two decades of life. In this manner, face processing is a model system of many areas of complex cognitive development. WIREs Cogn Sci 2017, 8:e1423. doi: 10.1002/wcs.1423 This article is categorized under: Psychology > Development and Aging Neuroscience > Cognition
Description of commonly used noninvasive imaging techniques in terms of their spatial (in mm) and temporal (in seconds) resolution. Electroencephalographic and the signal‐averaged EEG measure of event‐related potentials measures, and the closely related magnetoencephalography (MEG) measures have very good temporal resolution, but limited spatial resolution. That means they can distinguish different brain events with millisecond accuracy, but are relatively limited in defining the precise location of the activity in the brain. Functional magnetic resonance imaging and a technique called functional near‐infrared spectroscopy, a technique that uses near‐infrared light to observe underlying brain activity, provide exceptional spatial resolution but are less precise in defining the timing of brain events. Thus, to understand functional brain development with good spatial and temporal resolution, multiple techniques must be used in combination. Although not directly measuring brain function, traditional magnetic resonance imaging measures of brain development such as brain volume and cortical thickness, and diffusion tensor imaging that can measure white matter changes, may be correlated with behavioral measures (e.g., IQ, face processing accuracy) to make structure‐functional associations. Two highly invasive techniques, the study of permanent brain lesions acquired in development and positron emission tomography, a functional brain imaging technique that requires the injection of radioactive substances to obtain its measures are shown to illustrate the functional imaging space they cover. Note that fine resolution imaging using noninvasive techniques is currently not possible for developmental populations. (Adapted with permission from Ref . Copyright 1988 Science)
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Examples of FFA functional connectivity and activation of the extended face network across development. (a) Findings from a study of functional connectivity within the core face network for children and adults. Here, functional connectivity is measured by a specialized statistical technique that examines how information flows from one region to another. The abbreviations stand for: FG = fusiform gyrus in the region of the fusiform face area (FFA); IOG = inferior occipital gyrus in the region of the occipital face area (OFA); STG = superior temporal gyrus. The researchers tested participants in each group with tasks that asked them to identify an individual, to judge the emotional expression in a face, and to judge the direction of eye gaze. They calculated the strength of the connection between the core network regions as an estimate of how much each region depends on the information from other regions feeding into it. In this example, the FG region is highly dependent on information from the IOG in all three groups (solid lines). However, as signified by the values next to the lines, which is a measure of the strength of the connection between IOG and FG, both child groups displayed weaker connectivity compared to adults. In the emotion task, the connection between IOG and STG in children was not reliable (ns = not significant), whereas this connection was significant in adults (solid line). Overall, the findings suggest that certain aspects of the core network are integrated in childhood, but not with the strength seen in adults, and other aspects of the network are not integrated in childhood. Thus, functional connectivity in the core network has a protracted developmental trajectory that extends beyond the 10‐ to 11‐year‐old range that was tested. (b) Findings from face‐specific processing regions across the whole brain when children, adolescents, and adults viewed face passively. Regions in blue indicate activity that was greater in children than adults. Children showed greater activation than adults in many extended face processing regions. Adults are expected to activate regions in the extended network to match task demands. In this passive viewing task, there were no explicit demands that should invoke such processing. The fact children activate these regions suggests children do not modulate extended face network activity as efficiently as adults, consistent with the proposal that such modulation is the result of face processing expertise that takes an extended period to develop. (Reprinted with permission from Ref . Copyright 2013 Elsevier)
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Functional magnetic resonance imaging (FMRI) evidence of development of the fusiform face area (FFA) across development. (a) Changes in the size of the right hemisphere FFA, the dominant brain hemisphere for face processing, across development measured as the volume of the fusiform gyrus that was more active to face than objects. From age 6 years through adulthood, the FFA gradually increases in size. The correlation between age and FFA size is shown in the regression line. Open circles indicate participants (N = 71) that showed a detectable FFA and closed circles show the participants that did not produce a reliable FFA. (b) Developmental changes in the location of the FFA. Regions in warm colors indicated the region in the fusiform gyrus that became more consistently associated with face processing in adults relative to children. No regions were observed as consistently active in children but not adults. This suggests that the FFA location in children is more variable than in adults. (Reprinted with permission from Ref . Copyright 2013 Elsevier)
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Electroencephalographic and event‐related potentials in face processing studies of development. (a) Example of electrode placement in the scalp for typical EEG and ERP studies (source: https://commons.wikimedia.org/wiki/File:EEG_cap.jpg). (b) Findings from developmental studies of a face‐specific ERP component sensitive to processing faces, the N170, a negative voltage wave with a peak in adults at about 170 ms after the presentation of a face. The N170 shows that face processing expertise develops gradually across development not reaching mature levels until early adulthood. (Reprinted with permission from Ref . Copyright 2004 MIT Press)
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Description of the core and extended face networks. (a) Schematic of various regions associated with the core (cooler colors) and extended (warmer colors) face networks. Core face network regions, particularly the fusiform face area and occipital face area, are activated automatically in response to viewing a face. Extended network regions tend to be activated on a task‐specific basis. For example, the amygdala and limbic regions may be activated during tasks that require analysis of face emotion. (b) Diagram of information flow between regions in the core and extended regions and brief descriptions of the role of the different regions in face processing. (Adapted with permission from Ref . Copyright 2000 Cell)
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