Where Is the Semantic System? A Critical Review and Meta-Analysis of 120 Functional Neuroimaging Studies

Semantic memory refers to knowledge about people, objects, actions, relations, self, and culture acquired through experience. The neural systems that store and retrieve this information have been studied for many years, but a consensus regarding their identity has not been reached. Using strict inclusion criteria, we analyzed 120 functional neuroimaging studies focusing on semantic processing. Reliable areas of activation in these studies were identified using the activation likelihood estimate (ALE) technique. These activations formed a distinct, left-lateralized network comprised of 7 regions: posterior inferior parietal lobe, middle temporal gyrus, fusiform and parahippocampal gyri, dorsomedial prefrontal cortex, inferior frontal gyrus, ventromedial prefrontal cortex, and posterior cingulate gyrus. Secondary analyses showed specific subregions of this network associated with knowledge of actions, manipulable artifacts, abstract concepts, and concrete concepts. The cortical regions involved in semantic processing can be grouped into 3 broad categories: posterior multimodal and heteromodal association cortex, heteromodal prefrontal cortex, and medial limbic regions. The expansion of these regions in the human relative to the nonhuman primate brain may explain uniquely human capacities to use language productively, plan, solve problems, and create cultural and technological artifacts, all of which depend on the fluid and efficient retrieval and manipulation of semantic knowledge.

from Cerebral Cortex

Neural networks involved in voluntary and involuntary vocal pitch regulation in experienced singers

In an fMRI experiment, we tested experienced singers with singing tasks to investigate neural correlates of voluntary and involuntary vocal pitch regulation. We shifted the pitch of auditory feedback (±25 or 200 cents), and singers either: 1) ignored the shift and maintained their vocal pitch or 2) changed their vocal pitch to compensate for the shift. In our previous study, singers successfully ignored and compensated for 200-cent shifts; in the present experiment, we hypothesized that singers would be less able to ignore 25-cent shifts, due to a prepotent, corrective pitch-shift response. We expected that voluntary vocal regulation during compensate tasks would recruit the anterior portion of the rostral cingulate zone (RCZa) and posterior superior temporal sulcus (pSTS), as our earlier study reported; however, we predicted that a different network may be engaged during involuntary responses to 25-cent shifts. Singers were less able to ignore 25-cent shifts than 200-cent shifts, suggesting that pitch-shift responses to small shifts are under less voluntary control than responses to larger shifts. While we did not find neural activity specifically associated with involuntary pitch-shift responses, compensate tasks recruited a functionally connected network consisting of RCZa, pSTS, and anterior insula. Analyses of stimulus-modulated functional connectivity suggest that pSTS and intraparietal sulcus may monitor auditory feedback to extract pitch-shift direction in 200-cent tasks, but not in 25-cent tasks, which suggests that larger vocal corrections are under cortical control. During the compensate tasks, the pSTS may interact with the RCZa and anterior insula before voluntary vocal pitch correction occurs.

from Neuropsychologia

Language or music, mother or Mozart? Structural and environmental influences on infants’ language networks

Understanding how language emerged in our species calls for a detailed investigation of the initial specialization of the human brain for speech processing. Our earlier research demonstrated that an adult-like left-lateralized network of perisylvian areas is already active when infants listen to sentences in their native language, but did not address the issue of the specialization of this network for speech processing. Here we used fMRI to study the organization of brain activity in two-month-old infants when listening to speech or to music. We also explored how infants react to their mother’s voice relative to an unknown voice. The results indicate that the well-known structural asymmetry already present in the infants’ posterior temporal areas has a functional counterpart: there is a left-hemisphere advantage for speech relative to music at the level of the planum temporale. The posterior temporal regions are thus differently sensitive to the auditory environment very early on, channelling speech inputs preferentially to the left side. Furthermore, when listening to the mother’s voice, activation was modulated in several areas, including areas involved in emotional processing (amygdala, orbito-frontal cortex), but also, crucially, a large extent of the left posterior temporal lobe, suggesting that the mother’s voice plays a special role in the early shaping of posterior language areas. Both results underscore the joint contributions of genetic constraints and environmental inputs in the fast emergence of an efficient cortical network for language processing in humans.

from Brain and Language

The functional organization of trial-related activity in lexical processing after early left hemispheric brain lesions: An event-related fMRI study

Abstract
Children with congenital left hemisphere damage due to perinatal stroke are capable of acquiring relatively normal language functions despite experiencing a cortical insult that in adults often leads to devastating lifetime disabilities. Although this observed phenomenon is accepted, its neurobiological mechanisms are not well characterized. In this paper we examined the functional neuroanatomy of lexical processing in 13 children/adolescents with perinatal left hemispheric damage. In contrast to many previous perinatal infarct fMRI studies, we used an event-related design, which allowed us to isolate trial-related activity and examine correct and error trials separately. Using both group and single subject analysis techniques we attempt to address several methodological factors that may contribute to some discrepancies in the perinatal lesion literature. These methodological factors include making direct statistical comparisons, using common stereotactic space, using both single subject and group analyses, and accounting for performance differences. Our group analysis, investigating correct trial-related activity (separately from error trials), showed very few statistical differences in the non-involved right hemisphere between patients and performance matched controls. The single subject analysis revealed atypical regional activation patterns in several patients; however, the location of these regions identified in individual patients often varied across subjects. These results are consistent with the idea that alternative functional organization of trial-related activity after left hemisphere lesions is in large part unique to the individual. In addition, reported differences between results obtained with event-related designs and blocked designs may suggest diverging organizing principles for sustained and trial-related activity after early childhood brain injuries.

from Brain and Language

The functional organization of trial-related activity in lexical processing after early left hemispheric brain lesions: An event-related fMRI study

Children with congenital left hemisphere damage due to perinatal stroke are capable of acquiring relatively normal language functions despite experiencing a cortical insult that in adults often leads to devastating lifetime disabilities. Although this observed phenomenon is accepted, its neurobiological mechanisms are not well characterized. In this paper we examined the functional neuroanatomy of lexical processing in 13 children/adolescents with perinatal left hemispheric damage. In contrast to many previous perinatal infarct fMRI studies, we used an event-related design, which allowed us to isolate trial-related activity and examine correct and error trials separately. Using both group and single subject analysis techniques we attempt to address several methodological factors that may contribute to some discrepancies in the perinatal lesion literature. These methodological factors include making direct statistical comparisons, using common stereotactic space, using both single subject and group analyses, and accounting for performance differences. Our group analysis, investigating correct trial-related activity (separately from error trials), showed very few statistical differences in the non-involved right hemisphere between patients and performance matched controls. The single subject analysis revealed atypical regional activation patterns in several patients; however, the location of these regions identified in individual patients often varied across subjects. These results are consistent with the idea that alternative functional organization of trial-related activity after left hemisphere lesions is in large part unique to the individual. In addition, reported differences between results obtained with event-related designs and blocked designs may suggest diverging organizing principles for sustained and trial-related activity after early childhood brain injuries.

from Brain and Language

Dyslexic Children Show Short-Term Memory Deficits in Phonological Storage and Serial Rehearsal: An fMRI Study

Dyslexia is primarily associated with a phonological processing deficit. However, the clinical manifestation also includes a reduced verbal working memory (WM) span. It is unclear whether this WM impairment is caused by the phonological deficit or a distinct WM deficit. The main aim of this study was to investigate neuronal activation related to phonological storage and rehearsal of serial order in WM in a sample of 13-year-old dyslexic children compared with age-matched nondyslexic children. A sequential verbal WM task with two tasks was used. In the Letter Probe task, the probe consisted of a single letter and the judgment was for the presence or absence of that letter in the prior sequence of six letters. In the Sequence Probe (SP) task, the probe consisted of all six letters and the judgment was for a match of their serial order with the temporal order in the prior sequence. Group analyses as well as single-subject analysis were performed with the statistical parametric mapping software SPM2. In the Letter Probe task, the dyslexic readers showed reduced activation in the left precentral gyrus (BA6) compared to control group. In the Sequence Probe task, the dyslexic readers showed reduced activation in the prefrontal cortex and the superior parietal cortex (BA7) compared to the control subjects. Our findings suggest that a verbal WM impairment in dyslexia involves an extended neural network including the prefrontal cortex and the superior parietal cortex. Reduced activation in the left BA6 in both the Letter Probe and Sequence Probe tasks may be caused by a deficit in phonological processing. However, reduced bilateral activation in the BA7 in the Sequence Probe task only could indicate a distinct working memory deficit in dyslexia associated with temporal order processing.

from Neuroscience Research

Aging Influences the Neural Correlates of Lexical Decision but Not Automatic Semantic Priming

Human behavioral data indicate that older adults are slower to perform lexical decisions (LDs) than young adults but show similar reaction time gains when these decisions are primed semantically. The present study explored the functional neuroanatomic bases of these frequently observed behavioral findings. Young and older groups completed unprimed and primed LD tasks while functional magnetic resonance imaging (fMRI) was recorded, using a fully randomized trial design paralleling those used in behavioral research. Results from the unprimed task found that age-related slowing of LD was associated with decreased activation in perceptual extrastriate regions and increased activation in regions associated with higher level linguistic processes, including prefrontal cortex. In contrast to these age-related changes in brain activation, the older group showed a preserved pattern of fMRI decreases in inferior temporal cortex when LD was primed semantically. These findings provide evidence that older adults’ LD abilities benefit from contexts that reduce the need for frontally mediated strategic processes and capitalize on the continued sensitivity of inferior temporal cortex to automatic semantic processes in aging.

from Cerebral Cortex

The Influence of Multiple Primes on Bottom-Up and Top-Down Regulation during Meaning Retrieval: Evidence for 2 Distinct Neural Networks

Meaning retrieval of a word can proceed fast and effortlessly or can be characterized by a controlled search for candidate lexical items and a subsequent selection process. In the current study, we facilitated meaning retrieval by increasing the number of words that were related to the final target word in a triplet (e.g., lion–stripes–tiger). To induce higher search and selection demands, we presented ambiguous words as targets (i.e., homonyms like ball) in half of the trials. Hereby, the dominant (game), low-frequent (dance), or both meanings of the homonym were primed. Participants performed a relatedness judgment during functional magnetic resonance imaging. Activation in a bilateral network (angular gyrus, rostromedial prefrontal cortex) increased linearly with multiple related primes, whereas the posterior left inferior prefrontal cortex (pLIPC) showed the reverse activation pattern for unambiguous trials. When homonyms served as targets, pLIPC responded strongest when both meanings or low-frequent concepts were addressed. Additional anterior left inferior prefrontal cortex activation was observed for the latter trials only. The data support an interaction between 2 distinct cerebral networks that can be linked to automatic bottom-up support and top-down control during meaning retrieval. They further imply a functional specialization of the LIPC along an anterior–posterior dimension.

from Cerebral Cortex

Regional changes in word-production laterality after a naming treatment designed to produce a rightward shift in frontal activity

Five nonfluent aphasia patients participated in a picture-naming treatment that used an intention manipulation (opening a box and pressing a button on a device in the box with the left hand) to initiate naming trials and was designed to re-lateralize word production mechanisms from the left to the right frontal lobe. To test the underlying assumption regarding re-lateralization, patients participated in fMRI of category-member generation before and after treatment. Generally, the four patients who improved during treatment showed reduced frontal activity from pre- to post-treatment fMRI with increasing concentration of activity in the right posterior frontal lobe (motor/premotor cortex, pars opercularis), demonstrating a significant shift in lateraliity toward the right lateral frontal lobe, as predicted. Three of these four patients showed no left frontal activity by completion of treatment, indicating that right posterior lateral frontal activity supported category-member generation. Patients who improved in treatment showed no difference in lateralization of lateral frontal activity from normal controls pre-treatment, but post-treatment, their lateral frontal activity during category-member generation was significantly more right lateralized than that of controls. Patterns of activity pre- and post-treatment suggested increasing efficiency of cortical processing as a result of treatment in the four patients who improved. The one patient who did not improve during treatment showed a leftward shift in lateral frontal lateralization that was significantly different from the four patients who did improve. Neither medial frontal nor posterior perisylvian re-lateralization from immediately pre- to immediately post-treatment images was a necessary condition for significant treatment gains or shift in lateral frontal lateralization. Of the three patients who improved and in whom posterior perisylvian activity could be measured at post-treatment fMRI, all maintained equal or greater amounts of left-hemisphere perisylvian activity as compared to right. This finding is consistent with reviews suggesting both hemispheres are involved in recovery of language in aphasia patients.

from the Journal of Voice

Category-specific neural processing for naming pictures of animals and naming pictures of tools: An ALE meta-analysis

Using activation-likelihood estimation (ALE) meta-analysis, we identified brain areas that are invoked when people name pictures of animals and pictures of tools. We found that naming animals and naming tools invoked separate distributed networks in the brain. Specifically, we found that naming animals invoked greater responses than naming tools in frontal lobe structures that are typically modulated by emotional content and task demands, and in a number of visual areas in the ventral stream. In contrast, naming tools invoked greater responses in a different set of areas in the ventral stream than those invoked by naming animals. Naming tools also invoked greater responses than naming animals in motor areas in the frontal lobe as well as in sensory areas in the parietal lobe. The only overlapping sites of activation that we found for naming these two categories of objects were in the left pars triangularis, the left inferior temporal gyrus, and the left parahippocampal gyrus. Taken together, our meta-analysis reveals that animals and tools are categorically represented in visual areas but show convergence in higher-order associative areas in the temporal and frontal lobes in regions that are typically regarded as being involved in memory and/or semantic processing. Our results also reveal that naming tools not only engages visual areas in the ventral stream but also a fronto-parietal network associated with tool use. Whether or not this network associated with tool use contributes directly to recognition will require further investigation.

from Neuropsychologia

Differences in task demands influence the hemispheric lateralization and neural correlates of metaphor

This study investigated metaphor comprehension in the broader context of task-difference effects and manipulation of processing difficulty. We predicted that right hemisphere recruitment would show greater specificity to processing difficulty rather than metaphor comprehension. Previous metaphor processing studies have established that the left inferior frontal gyrus strongly correlates with metaphor comprehension but there has been controversy about whether right hemisphere (RH) involvement is specific for metaphor comprehension. Functional MRI data were recorded from healthy subjects who read novel metaphors, conventional metaphors, definition-like sentences, or literal sentences. We investigated metaphor processing in contexts where semantic judgment or imagery modulates linguistic judgment. Our findings support the position that the type of task rather than figurative language processing per se modulates the left inferior gyrus (LIFG). RH involvement was more influenced by processing difficulty and less by the novelty or figurativity of linguistic expressions. Our results suggest that figurative language processing depends upon the effects of task-type and processing difficulty on imaging results.

from Brain and Language

Neural development of networks for audiovisual speech comprehension

Everyday conversation is both an auditory and a visual phenomenon. While visual speech information enhances comprehension for the listener, evidence suggests that the ability to benefit from this information improves with development. A number of brain regions have been implicated in audiovisual speech comprehension, but the extent to which the neurobiological substrate in the child compares to the adult is unknown. In particular, developmental differences in the network for audiovisual speech comprehension could manifest through the incorporation of additional brain regions, or through different patterns of effective connectivity. In the present study we used functional magnetic resonance imaging and structural equation modeling (SEM) to characterize the developmental changes in network interactions for audiovisual speech comprehension. The brain response was recorded while children 8- to 11-years-old and adults passively listened to stories under audiovisual (AV) and auditory-only (A) conditions. Results showed that in children and adults, AV comprehension activated the same fronto-temporo-parietal network of regions known for their contribution to speech production and perception. However, the SEM network analysis revealed age-related differences in the functional interactions among these regions. In particular, the influence of the posterior inferior frontal gyrus/ventral premotor cortex on supramarginal gyrus differed across age groups during AV, but not A speech. This functional pathway might be important for relating motor and sensory information used by the listener to identify speech sounds. Further, its development might reflect changes in the mechanisms that relate visual speech information to articulatory speech representations through experience producing and perceiving speech.

from Brain and Language

Primary and multisensory cortical activity is correlated with audiovisual percepts

Incongruent auditory and visual stimuli can elicit audiovisual illusions such as the McGurk effect where visual /ka/ and auditory /pa/ fuse into another percept such as/ta/. In the present study, human brain activity was measured with adaptation functional magnetic resonance imaging to investigate which brain areas support such audiovisual illusions. Subjects viewed trains of four movies beginning with three congruent /pa/ stimuli to induce adaptation. The fourth stimulus could be (i) another congruent /pa/, (ii) a congruent /ka/, (iii) an incongruent stimulus that evokes the McGurk effect in susceptible individuals (lips /ka/ voice /pa/), or (iv) the converse combination that does not cause the McGurk effect (lips /pa/ voice/ ka/). This paradigm was predicted to show increased release from adaptation (i.e. stronger brain activation) when the fourth movie and the related percept was increasingly different from the three previous movies. A stimulus change in either the auditory or the visual stimulus from /pa/ to /ka/ (iii, iv) produced within-modality and cross-modal responses in primary auditory and visual areas. A greater release from adaptation was observed for incongruent non-McGurk (iv) compared to incongruent McGurk (iii) trials. A network including the primary auditory and visual cortices, nonprimary auditory cortex, and several multisensory areas (superior temporal sulcus, intraparietal sulcus, insula, and pre-central cortex) showed a correlation between perceiving the McGurk effect and the fMRI signal, suggesting that these areas support the audiovisual illusion. Hum Brain Mapp, 2009. © 2009 Wiley-Liss, Inc.

from Human Brain Mapping

Sensory-motor brain network connectivity for speech comprehension

Abstract
The act of listening to speech activates a large network of brain areas. In the present work, a novel data-driven technique (the combination of independent component analysis and Granger causality) was used to extract brain network dynamics from an fMRI study of passive listening to Words, Pseudo-Words, and Reverse-played words. Using this method we show the functional connectivity modulations among classical language regions (Broca’s and Wernicke’s areas) and inferior parietal, somatosensory, and motor areas and right cerebellum. Word listening elicited a compact pattern of connectivity within a parieto-somato-motor network and between the superior temporal and inferior frontal gyri. Pseudo-Word stimuli induced activities similar to the Word condition, which were characterized by a highly recurrent connectivity pattern, mostly driven by the temporal lobe activity. Also the Reversed-Word condition revealed an important influence of temporal cortices, but no integrated activity of the parieto-somato-motor network. In parallel, the right cerebellum lost its functional connection with motor areas, present in both Word and Pseudo-Word listening. The inability of the participant to produce the Reversed-Word stimuli also evidenced two separate networks: the first was driven by frontal areas and the right cerebellum toward somatosensory cortices; the second was triggered by temporal and parietal sites towards motor areas. Summing up, our results suggest that semantic content modulates the general compactness of network dynamics as well as the balance between frontal and temporal language areas in driving those dynamics. The degree of reproducibility of auditory speech material modulates the connectivity pattern within and toward somatosensory and motor areas. Hum Brain Mapp, 2009. © 2009 Wiley-Liss, Inc.

from Human Brain Mapping

Age-Related Differences in Laterality of Cortical Activations in Swallowing

The present study examined age differences in neural lateralization patterns during swallowing and three related tasks, using functional magnetic resonance imaging (fMRI). Ten healthy right-handed young adults (mean age = 21.7 years, SD = 2.1 years) and nine healthy elders (mean age = 70.2 years, SD = 3.9 years) were scanned in a 3-T MRI head scanner. Participants were visually cued to “prepare to swallow,” “swallow,” “tap your tongue,” and “clear your throat” in randomized order. Laterality preference for each task was examined within and between groups using region-of-interest (ROI) analyses in seven areas of the left and right primary sensorimotor and premotor cortices. Results of the within-group comparisons verified a more active role of the left premotor cortex in motor-cognitive planning of deglutition in both young and older adults and a more active role of selected areas of the right hemisphere during swallowing in young adults. Greater variability was seen during tongue tapping and throat clearing in both groups. Finally, as people age the cortical hemispheric control of swallowing seems to start becoming more symmetrical/bilateral, which may indicate neural compensatory mechanisms of the aging brain commonly seen for other motor and cognitive functions.

from Dysphagia