Rostral and caudal prefrontal contribution to creativity: a meta-analysis of functional imaging data.

Creativity is of central importance for human civilization, yet its neurocognitive bases are poorly understood. The aim of the present study was to integrate existing functional imaging data by using the meta-analysis approach. We reviewed 34 functional imaging studies that reported activation foci during tasks assumed to engage creative thinking in healthy adults. A coordinate-based meta-analysis using Activation Likelihood Estimation (ALE) first showed a set of predominantly left-hemispheric regions shared by the various creativity tasks examined. These regions included the caudal lateral prefrontal cortex (PFC), the medial and lateral rostral PFC, and the inferior parietal and posterior temporal cortices. Further analyses showed that tasks involving the combination of remote information (combination tasks) activated more anterior areas of the lateral PFC than tasks involving the free generation of unusual responses (unusual generation tasks), although both types of tasks shared caudal prefrontal areas. In addition, verbal and non-verbal tasks involved the same regions in the left caudal prefrontal, temporal, and parietal areas, but also distinct domain-oriented areas. Taken together, these findings suggest that several frontal and parieto-temporal regions may support cognitive processes shared by diverse creativity tasks, and that some regions may be specialized for distinct types of processes. In particular, the lateral PFC appeared to be organized along a rostro-caudal axis, with rostral regions involved in combining ideas creatively and more posterior regions involved in freely generating novel ideas.

The influence of reading ability on subsequent changes in verbal IQ in the teenage years.

Intelligence Quotient (IQ) is regularly used in both education and employment as a measure of cognitive ability. Although an individual's IQ is generally assumed to stay constant across the lifespan, a few studies have suggested that there may be substantial variation at the individual level. Motivated by previous reports that reading quality/quantity has a positive influence on vocabulary acquisition, we hypothesised that reading ability in the early teenage years might contribute to changes in verbal IQ (VIQ) over the next few years. We found that good readers were more likely to experience relative improvements in VIQ over time, with the reverse true for poor readers. These effects were largest when there was a discrepancy between Time 1 reading ability and Time 1 VIQ. In other words, VIQ increases tended to be greatest when reading ability was high relative to VIQ. Additional analyses supported these findings by showing that variance in VIQ change associated with Time 1 behaviour was also associated with independent measurements of brain structure. Our finding that reading in the early teenage years can predict a significant proportion of the variance in subsequent VIQ change has implications for targeted education in both home and school environments.

Automated identification of brain tumors from single MR images based on segmentation with refined patient-specific priors.

Brain tumors can have different shapes or locations, making their identification very challenging. In functional MRI, it is not unusual that patients have only one anatomical image due to time and financial constraints. Here, we provide a modified automatic lesion identification (ALI) procedure which enables brain tumor identification from single MR images. Our method rests on (A) a modified segmentation-normalization procedure with an explicit "extra prior" for the tumor and (B) an outlier detection procedure for abnormal voxel (i.e., tumor) classification. To minimize tissue misclassification, the segmentation-normalization procedure requires prior information of the tumor location and extent. We therefore propose that ALI is run iteratively so that the output of Step B is used as a patient-specific prior in Step A. We test this procedure on real T1-weighted images from 18 patients, and the results were validated in comparison to two independent observers' manual tracings. The automated procedure identified the tumors successfully with an excellent agreement with the manual segmentation (area under the ROC curve = 0.97 ± 0.03). The proposed procedure increases the flexibility and robustness of the ALI tool and will be particularly useful for lesion-behavior mapping studies, or when lesion identification and/or spatial normalization are problematic.

The brain's dorsal route for speech represents word meaning: evidence from gesture.

The dual-route model of speech processing includes a dorsal stream that maps auditory to motor features at the sublexical level rather than at the lexico-semantic level. However, the literature on gesture is an invitation to revise this model because it suggests that the premotor cortex of the dorsal route is a major site of lexico-semantic interaction. Here we investigated lexico-semantic mapping using word-gesture pairs that were either congruent or incongruent. Using fMRI-adaptation in 28 subjects, we found that temporo-parietal and premotor activity during auditory processing of single action words was modulated by the prior audiovisual context in which the words had been repeated. The BOLD signal was suppressed following repetition of the auditory word alone, and further suppressed following repetition of the word accompanied by a congruent gesture (e.g. ["grasp" + grasping gesture]). Conversely, repetition suppression was not observed when the same action word was accompanied by an incongruent gesture (e.g. ["grasp" + sprinkle]). We propose a simple model to explain these results: auditory and visual information converge onto premotor cortex where it is represented in a comparable format to determine (in)congruence between speech and gesture. This ability of the dorsal route to detect audiovisual semantic (in)congruence suggests that its function is not restricted to the sublexical level.

Brain-speech alignment enhances auditory cortical responses and speech perception.

Asymmetry in auditory cortical oscillations could play a role in speech perception by fostering hemispheric triage of information across the two hemispheres. Due to this asymmetry, fast speech temporal modulations relevant for phonemic analysis could be best perceived by the left auditory cortex, while slower modulations conveying vocal and paralinguistic information would be better captured by the right one. It is unclear, however, whether and how early oscillation-based selection influences speech perception. Using a dichotic listening paradigm in human participants, where we provided different parts of the speech envelope to each ear, we show that word recognition is facilitated when the temporal properties of speech match the rhythmic properties of auditory cortices. We further show that the interaction between speech envelope and auditory cortices rhythms translates in their level of neural activity (as measured with fMRI). In the left auditory cortex, the neural activity level related to stimulus-brain rhythm interaction predicts speech perception facilitation. These data demonstrate that speech interacts with auditory cortical rhythms differently in right and left auditory cortex, and that in the latter, the interaction directly impacts speech perception performance.

Verbal and non-verbal intelligence changes in the teenage brain.

Intelligence quotient (IQ) is a standardized measure of human intellectual capacity that takes into account a wide range of cognitive skills. IQ is generally considered to be stable across the lifespan, with scores at one time point used to predict educational achievement and employment prospects in later years. Neuroimaging allows us to test whether unexpected longitudinal fluctuations in measured IQ are related to brain development. Here we show that verbal and non-verbal IQ can rise or fall in the teenage years, with these changes in performance validated by their close correlation with changes in local brain structure. A combination of structural and functional imaging showed that verbal IQ changed with grey matter in a region that was activated by speech, whereas non-verbal IQ changed with grey matter in a region that was activated by finger movements. By using longitudinal assessments of the same individuals, we obviated the many sources of variation in brain structure that confound cross-sectional studies. This allowed us to dissociate neural markers for the two types of IQ and to show that general verbal and non-verbal abilities are closely linked to the sensorimotor skills involved in learning. More generally, our results emphasize the possibility that an individual's intellectual capacity relative to their peers can decrease or increase in the teenage years. This would be encouraging to those whose intellectual potential may improve, and would be a warning that early achievers may not maintain their potential.

Regional and hemispheric determinants of language laterality: implications for preoperative fMRI.

Language is typically a function of the left hemisphere but the right hemisphere is also essential in some healthy individuals and patients. This inter-subject variability necessitates the localization of language function, at the individual level, prior to neurosurgical intervention. Such assessments are typically made by comparing left and right hemisphere language function to determine "language lateralization" using clinical tests or fMRI. Here, we show that language function needs to be assessed at the region and hemisphere specific level, because laterality measures can be misleading. Using fMRI data from 82 healthy participants, we investigated the degree to which activation for a semantic word matching task was lateralized in 50 different brain regions and across the entire cortex. This revealed two novel findings. First, the degree to which language is lateralized across brain regions and between subjects was primarily driven by differences in right hemisphere activation rather than differences in left hemisphere activation. Second, we found that healthy subjects who have relatively high left lateralization in the angular gyrus also have relatively low left lateralization in the ventral precentral gyrus. These findings illustrate spatial heterogeneity in language lateralization that is lost when global laterality measures are considered. It is likely that the complex spatial variability we observed in healthy controls is more exaggerated in patients with brain damage. We therefore highlight the importance of investigating within hemisphere regional variations in fMRI activation, prior to neuro-surgical intervention, to determine how each hemisphere and each region contributes to language processing.

Automatic top-down processing explains common left occipito-temporal responses to visual words and objects.

Previous studies have demonstrated that a region in the left ventral occipito-temporal (LvOT) cortex is highly selective to the visual forms of written words and objects relative to closely matched visual stimuli. Here, we investigated why LvOT activation is not higher for reading than picture naming even though written words and pictures of objects have grossly different visual forms. To compare neuronal responses for words and pictures within the same LvOT area, we used functional magnetic resonance imaging adaptation and instructed participants to name target stimuli that followed briefly presented masked primes that were either presented in the same stimulus type as the target (word-word, picture-picture) or a different stimulus type (picture-word, word-picture). We found that activation throughout posterior and anterior parts of LvOT was reduced when the prime had the same name/response as the target irrespective of whether the prime-target relationship was within or between stimulus type. As posterior LvOT is a visual form processing area, and there was no visual form similarity between different stimulus types, we suggest that our results indicate automatic top-down influences from pictures to words and words to pictures. This novel perspective motivates further investigation of the functional properties of this intriguing region.

Lateralization is predicted by reduced coupling from the left to right prefrontal cortex during semantic decisions on written words.

Over 90% of people activate the left hemisphere more than the right hemisphere for language processing. Here, we show that the degree to which language is left lateralized is inversely related to the degree to which left frontal regions drive activity in homotopic right frontal regions. Lateralization was assessed in 60 subjects using functional magnetic resonance imaging (fMRI) activation for semantic decisions on verbal (written words) and nonverbal (pictures of objects) stimuli. Regional interactions between left and right ventral and dorsal frontal regions were assessed using dynamic causal modeling (DCM), random-effects Bayesian model selection at the family level, and Bayesian model averaging at the connection level. We found that 1) semantic decisions on words and pictures modulated interhemispheric coupling between the left and right dorsal frontal regions, 2) activation was more left lateralized for words than pictures, and 3) for words only, left lateralization was greater when the coupling from the left to right dorsal frontal cortex was reduced. These results have theoretical implications for understanding how left and right hemispheres communicate with one another during the processing of lateralized functions.

Left hemisphere regions are critical for language in the face of early left focal brain injury.

A predominant theory regarding early stroke and its effect on language development, is that early left hemisphere lesions trigger compensatory processes that allow the right hemisphere to assume dominant language functions, and this is thought to underlie the near normal language development observed after early stroke. To test this theory, we used functional magnetic resonance imaging to examine brain activity during category fluency in participants who had sustained pre- or perinatal left hemisphere stroke (n = 25) and in neurologically normal siblings (n = 27). In typically developing children, performance of a category fluency task elicits strong involvement of left frontal and lateral temporal regions and a lesser involvement of right hemisphere structures. In our cohort of atypically developing participants with early stroke, expressive and receptive language skills correlated with activity in the same left inferior frontal regions that support language processing in neurologically normal children. This was true independent of either the amount of brain injury or the extent that the injury was located in classical cortical language processing areas. Participants with bilateral activation in left and right superior temporal-inferior parietal regions had better language function than those with either predominantly left- or right-sided unilateral activation. The advantage conferred by left inferior frontal and bilateral temporal involvement demonstrated in our study supports a strong predisposition for typical neural language organization, despite an intervening injury, and argues against models suggesting that the right hemisphere fully accommodates language function following early injury.

Predicting language lateralization from gray matter.

It has long been predicted that the degree to which language is lateralized to the left or right hemisphere might be reflected in the underlying brain anatomy. We investigated this relationship on a voxel-by-voxel basis across the whole brain using structural and functional magnetic resonance images from 86 healthy participants. Structural images were converted to gray matter probability images, and language activation was assessed during naming and semantic decision. All images were spatially normalized to the same symmetrical template, and lateralization images were generated by subtracting right from left hemisphere signal at each voxel. We show that the degree to which language was left or right lateralized was positively correlated with the degree to which gray matter density was lateralized. Post hoc analyses revealed a general relationship between gray matter probability and blood oxygenation level-dependent signal. This is the first demonstration that structural brain scans can be used to predict language lateralization on a voxel-by-voxel basis in the normal healthy brain.

The main sources of intersubject variability in neuronal activation for reading aloud.

The aim of this study was to find the most prominent source of intersubject variability in neuronal activation for reading familiar words aloud. To this end, we collected functional imaging data from a large sample of subjects (n = 76) with different demographic characteristics such as handedness, sex, and age, while reading. The subject-by-subject error variance was estimated from a one-sample t test (on all 76 subjects) and was reduced to a lower dimension using principal components decomposition. A Gaussian Mixture Model was then applied to dissociate different subgroups of subjects that explained the main sources of variability in the data. This resulted in the identification of four different subject groups. The comparison of these subgroups to the subjects' demographic details showed that age had a significant effect on the subject partitioning. In addition, a region-by-group dissociation in the dorsal and the ventral inferior frontal cortex was consistent with previously reported dissociations in semantic and nonsemantic reading strategies. In contrast to these significant findings, the groupings did not differentiate subjects on the basis of either sex or handedness, nor did they segregate the subjects with right- versus left-lateralized reading activation. We therefore conclude that, of the variables tested, age and reading strategy were the most prominent source of variability in activation for reading familiar words aloud.

Explaining function with anatomy: language lateralization and corpus callosum size.

The anatomy of the corpus callosum (CC) has been advocated as a potential marker for functional lateralization because its size is supposedly proportional to the number of fibers connecting the hemispheres. Previous morphometric studies of this relationship have compared CC size in groups of subjects who are more or less likely to show differences in their lateralization (e.g., left vs right handers). The findings, however, have been inconsistent, and to our knowledge, no previous study has directly compared CC size with lateralization assessed by functional imaging data. We therefore combined anatomical measurements of CC size with left versus right hemisphere language activation in 74 normal subjects. After controlling for perceptual and motor output effects, as well as for global white-matter volume, handedness, gender and age, we found that subjects who had a larger CC showed more left lateralization for language in posterior temporal and inferior frontal regions. Examination of these effects revealed that, as CC size increased, stronger lateralization resulted from more left hemisphere activation in both regions as well as reduced right hemisphere activation in the posterior temporal region. Our observations provide the first clear evidence in normal subjects that the midsagittal surface area of the CC contributes to the degree to which language is functionally lateralized. We discuss the complex interhemispheric processes that might underlie this effect.

Hemispheric specialization for language: Brain volume matters.

Increasing brain volume may impose constraints, through longer information transfer delays, on the distributed networks supporting language. Here, we assessed the relative effects of brain volume and other putative predictors of the functional variability of perisylvian language areas, as probed with PET, during both a language comprehension and a language production task. In the case of language comprehension (story listening), a linear combination of planum temporale surface, brain volume and handedness could explain almost 60% of the functional asymmetry observed in the perisylvian area. Without brain volume, the goodness of fit was significantly decreased (39%, P < 0.05), and furthermore, the effect of handedness was not detected anymore. This was due to the fact that in our sample, left-handers (n = 12) had a significantly larger brain volume as compared to right-handers (n = 8, P = 0.03). As for language production (verb generation), brain volume and the planum temporale also played a role. However, in this case, the main predictor of functional variability was handedness, where a greater degree of right-handedness was associated with larger activation of left inferior frontal regions. Depending on the language component of interest, these results support different (yet compatible) theories on hemispheric specialization. Left specialization for comprehension could be attributed to the constraints of processing speech stimuli, while a gestural origin of language is mostly supported by the relation we observed between left specialization for production and right-handedness.

Hemispheric specialization for language.

Hemispheric specialization for language is one of the most robust findings of cognitive neuroscience. In this review, we first present the main hypotheses about the origins of this important aspect of brain organization. These theories are based in part on the main approaches to hemispheric specialization: studies of aphasia, anatomical asymmetries and, nowadays, neuroimaging. All these approaches uncovered a large inter-individual variability which became the bulk of research on hemispheric specialization. This is why, in a second part of the review, we present the main facts about inter-individual variability, trying to relate findings to the theories presented in the first part. This review focuses on neuroimaging as it has recently given important results, thanks to investigations of both anatomical and functional asymmetries in healthy subjects. Such investigations have confirmed that left-handers, especially "familial left-handers", are more likely to have an atypical pattern of hemispheric specialization for language. Differences between men and women seem less evident although a less marked hemispheric specialization for language was depicted in women. As for the supposed relationship between anatomical and functional asymmetries, it has been shown that the size of the left (not the right) planum temporale could explain part of the variability of left hemispheric specialization for language comprehension. Taken as a whole, findings seem to vary with language tasks and brain regions, therefore showing that hemispheric specialization for language is multi-dimensional. This is not accounted for in the existing models of hemispheric specialization.

Left planum temporale: an anatomical marker of left hemispheric specialization for language comprehension.

We report on a study aimed at investigating the relationships between handedness, anatomical data and functional data related to speech processing. Twenty subjects with variable handedness (Edinburgh score ranging from -100 to 100) underwent both anatomical magnetic resonance imaging (MRI) and Positron Emission Tomography (PET) during story listening and rest. The surface areas of the left and right planum temporale (PT) were measured on each subject's MRI scan. A multiple regression analysis of PET data was conducted using these PT surface areas as well as handedness scores as predictors. The surface of the left PT explained a significant part of the functional variability. We observed that subjects who had the larger left PT were likely to show a larger leftward functional asymmetry of several perisylvian areas, namely the inferior parietal lobule outside the supra-marginal gyrus (the angular gyrus and the cortex above), Heschl's gyrus, the rolandic operculum, and the temporal pole. The size of the right PT explained only a little part of functional variability and we found no evidence that the anatomical asymmetry of the PT explained functional variability. In addition, we could not evidence any relationship between handedness and functional data. These results, which confirm previous work, argue for a perceptive origin of hemispheric specialization for language comprehension as has been suggested by others.