Functional connectivity in an fMRI study of semantic and phonological processes and the effect of L-Dopa

We describe an fMRI experiment examining the functional connectivity (FC) between regions of the brain associated with semantic and phonological processing. We wished to explore whether L-Dopa administration affects the interaction between language network components in semantic and phonological categorization tasks, as revealed by FC. We hypothesized that L-Dopa would decrease FC due to restriction of the semantic network. During two test sessions (placebo and L-Dopa) each participant performed two fMRI runs, involving phonological and semantic processing. A number of brain regions commonly activated by the two tasks were chosen as regions if interest: left inferior frontal, left posterior temporal and left fusiform gyri, and left parietal cortex. FC was calculated and further analyzed for effects of either the drug or task. No main effect for drug was found. A significant main effect for task was found, with a greater average correlation for the phonological task than for the semantic task. These findings suggest that language areas are activated in a more synchronous manner for phonological than for semantic tasks. This may relate to the fact that phonological processes are mediated to a greater extent within language areas, whereas semantic tasks likely require greater interaction outside of the language areas. Alternatively, this may be due to differences in the attentional requirements of the two tasks.     

An investigation of immune system disorder as a "marker" for anomalous dominance

Geschwind and Galaburda (1987) proposed that immune disorder (ID) susceptibility, along with left handedness and familial sinistrality (FS), is a "marker" for anomalous dominance. The theory predicts lesser left lateralization for language processes, lessened left hemisphere abilities, and enhanced right hemisphere abilities. We assessed language laterality (dichotic consonant vowel task) and performances on spatial and verbal tasks. Subjects were 128 college students. The factors of handedness, sex, FS, and immune disorder history (negative or positive) were perfectly counterbalanced. Left-handers were significantly less lateralized for language and scored lower than right-handers on the spatial tasks. Females scored lower on mental rotation than males, but performed comparably to males on the spatial relations task. The only effect of ID was by way of interaction with FS on both spatial tasks--subjects who were either negative or positive on both FS and ID status factors scored significantly higher than subjects negative for one but positive for the other factor. A speculative explanatory model for this interaction was proposed. The model incorporates the notion that FS and ID factors are comparably correlated, but in opposite directions, with hormonal factors implicated by other research as relevant for spatial ability differences. Finally, no support for the "anomalous dominance" hypothesis predictions was found.     

Multitask investigation of individual differences in hemispheric asymmetry

Right-handed subjects (N = 120) participated in four different laterality tasks designed to measure aspects of cerebral hemisphere asymmetry: identification of dichotically presented consonant-vowel syllables (CVs), examination of the effects of concurrent repetition of CVs and concurrent anagram solution on finger-tapping by the right and left hands, lateralized identification of CVs presented tachistoscopically to the left and right visual fields, and left/right biases on a free-vision face task involving judgments of emotion. Ear differences in the dichotic listening task were related to the pattern of lateralized interference in the dual-task finger-tapping paradigm. There were no other significant relations between pairs of tasks, but when the present results are considered in the light of other recent experiments, there appears to be a relation between lateral bias on the free-vision face task and visual field differences in tachistoscopic identification. The pattern of results has implications for hypothesized individual differences among right-handers in cerebral dominance for verbal processes, input pathway dominance, and asymmetric arousal of the two cerebral hemispheres.     

An fMRI study of sex differences in regional activation to a verbal and a spatial task

Sex differences in cognitive performance have been documented, women performing better on some phonological tasks and men on spatial tasks. An earlier fMRI study suggested sex differences in distributed brain activation during phonological processing, with bilateral activation seen in women while men showed primarily left-lateralized activation. This blood oxygen level-dependent fMRI study examined sex differences (14 men, 13 women) in activation for a spatial task (judgment of line orientation) compared to a verbal-reasoning task (analogies) that does not typically show sex differences. Task difficulty was manipulated. Hypothesized ROI-based analysis documented the expected left-lateralized changes for the verbal task in the inferior parietal and planum temporal regions in both men and women, but only men showed right-lateralized increase for the spatial task in these regions. Image-based analysis revealed a distributed network of cortical regions activated by the tasks, which consisted of the lateral frontal, medial frontal, mid-temporal, occipitoparietal, and occipital regions. The activation was more left lateralized for the verbal and more right for the spatial tasks, but men also showed some left activation for the spatial task, which was not seen in women. Increased task difficulty produced more distributed activation for the verbal and more circumscribed activation for the spatial task. The results suggest that failure to activate the appropriate hemisphere in regions directly involved in task performance may explain certain sex differences in performance. They also extend, for a spatial task, the principle that bilateral activation in a distributed cognitive system underlies sex differences in performance.     

Hemispheric specialization for reading

Behavioral laterality tasks with linguistic stimuli were used to assess the differential processing efficiencies of the cerebral hemispheres in right- and left-handed adults. Findings from a lateralized lexical decision task with concrete nouns supported Zaidel's (1983) "direct access" model of hemispheric functioning. A dual task consisting of oral and silent reading indicated that the right hand was significantly more disrupted than the left during unimanual finger tapping; however, some bilateral interference was observed. Taken together the findings suggest that although the left hemisphere was relatively more efficient, the right hemisphere was dynamically involved in the reading process.     

Sex task and processing strategy effects in hemispheric alpha asymmetries for the recall and recognition of arousal words: results from perceptual and motor tasks in males and females

Hemispheric alpha asymmetries of males and females were observed during perceptual and motor tasks requiring recall and recognition of words controlled for level of arousal (positive, negative, and neutral). Verbal reports of individual processing strategy were collected and analyzed relative to hemispheric alpha ratios. Results showed greater alpha suppression in the left relative to right hemisphere for recall as compared to recognition tasks and for word presentation when contrasted with motor conditions. High positive correlations were found between narrative report of processing strategy and hemispheric alpha data. A separate analysis revealed that seven subjects identified as highly analytic processors showed greater alpha suppression in the left relative to right hemisphere across tasks, conditions, and stimuli than did seven highly visual processors who, in contrast, demonstrated greater right hemispheric alpha suppression. Task difficulty and individual differences in processing style that modify cerebral laterality effects are discussed.     

The influence of gender, handedness, and performance level on specialized cognitive functioning

Individual differences in performance on neuropsychological tests were analyzed across age (prepubertal to adult), gender, and handedness groups and examined for performance level as a moderating variable. No differences were observed for the factor structure of these tests across ages and between genders, suggesting similar cognitive structures among these groups. Significant differences in performance were observed between males and females and, to a lesser extent, between right and left handers. Of interest, were significant Gender x Handedness and Gender x Handedness x Level (of performance) interactions seen especially in the older (postpubertal and adult) subjects. High performing, right handed males and left handed females performed better on visuospatial tasks while left handed males and right handed females performed better on verbosequential tasks.     

Priming vs. rhyming: orthographic and phonological representations in the left and right hemispheres

The right cerebral hemisphere has long been argued to lack phonological processing capacity. Recently, however, a sex difference in the cortical representation of phonology has been proposed, suggesting discrete left hemisphere lateralization in males and more distributed, bilateral representation of function in females. To evaluate this hypothesis and shed light on sex differences in the phonological processing capabilities of the left and right hemispheres, we conducted two experiments. Experiment 1 assessed phonological activation implicitly (masked homophone priming), testing 52 (M=25, F=27; mean age 19.23years, SD 1.64years) strongly right-handed participants. Experiment 2 subsequently assessed the explicit recruitment of phonology (rhyme judgement), testing 50 (M=25, F=25; mean age 19.67years, SD 2.05years) strongly right-handed participants. In both experiments the orthographic overlap between stimulus pairs was strictly controlled using DICE [Brew, C., & McKelvie, D. (1996). Word-pair extraction for lexicography. In K. Oflazer & H. Somers (Eds.), Proceedings of the second international conference on new methods in language processing (pp. 45-55). Ankara: VCH], such that pairs shared (a) high orthographic and phonological similarity (e.g., not-KNOT); (b) high orthographic and low phonological similarity (e.g., pint-HINT); (c) low orthographic and high phonological similarity (e.g., use-EWES); or (d) low orthographic and low phonological similarity (e.g., kind-DONE). As anticipated, high orthographic similarity facilitated both left and right hemisphere performance, whereas the left hemisphere showed greater facility when phonological similarity was high. This difference in hemispheric processing of phonological representations was especially pronounced in males, whereas female performance was far less sensitive to visual field of presentation across both implicit and explicit phonological tasks. As such, the findings offer behavioural evidence indicating that though both hemispheres are capable of orthographic analysis, phonological processing is discretely lateralised to the left hemisphere in males, but available in both the left and right hemisphere in females.     

Sex differences in semantic language processing: a functional MRI study

Predictions based on two models of sex differences in cerebral organization of language were compared by examining fMRI patterns of 10 females and 9 males during a semantic processing task. Both groups displayed activation of left inferior frontal gyrus (IFG), left superior temporal gyrus (STG), and cingulate. Females, but not males, showed bilateral IFG and STG activation. Further analyses revealed females had less diffuse left activation and greater right posterior temporal and insula region activation than males. Results support both an interhemispheric and an intrahemispheric model of sex differences in language, suggesting that the models may not be mutually exclusive.     

Visual field differences in verbal tasks: Effects of task familiarity and sex of subject

Four experiments using normal subjects investigated differences in magnitude of the right visual field (RVF) superiority as a function of word material (frequency and concreteness/imageability status), nonword letter strings (some of which were homophonic with nonpresented real words), and type of task (overt naming or lexical decision with discriminatory manual responses) as well as sex of the subject and the subject's familiarity with the material. Both latency and error measures showed that RVF superiority was more consistent when overt naming was required and with male subjects. For female subjects engaged in lexical decisions, a left visual field (LVF) superiority was often apparent, especially in the first half of an experimental sequence; when actually naming the items aloud, they showed field asymmetries similar to males. Except from an analysis of errors, there was little evidence to support differential right hemisphere mediation of high frequency concrete/imageable materials. It is suggested that in females, right hemisphere space normally reserved for visuospatial processing may have been invaded by secondary speech mechanisms. These mechanisms appear to operate at an essentially lexical level and may act in a supportive or auxiliary capacity for difficult or unfamiliar material; they seem to be equally concerned with both phonological and graphological processing and may account for the well-known female superiority in verbal tasks and inferiority in visuospatial tasks. Other findings are discussed such as the degree of consistency of the field differences, both for the same subjects and for the same stimulus materials under different task requirements and experimental conditions.     

Brain-behavior relationships in reading acquisition are modulated by socioeconomic factors

Functional neuroimaging may provide insights into the achievement gap in reading skill commonly observed across socioeconomic status (SES). Brain activation during reading tasks is known to be associated with individual differences in children's phonological language skills. By selecting children of equivalent phonological skill, yet diverse socioeconomic backgrounds, we use functional magnetic resonance imaging (fMRI) to demonstrate that a child's experience, as operationalized by SES, can systematically modulate the relationship between phonological language skills and reading-related brain activity in left fusiform and perisylvian regions. Specifically, at lower socioeconomic levels, individual differences in skill result in large differences in brain activation. In contrast, as SES increases, this relationship between phonological language skill and activation is attenuated. Socioeconomic background factors are thus found to modulate brain-behavior relationships in reading, indicating that cognitive, social, and neurobiological influences on reading development are fundamentally intertwined.     

Postural effect on manual laterality in different tasks in captive grey-cheeked mangabey (Lophocebus albigena)

The authors examined the effects of task complexity and posture on laterality and compared lateralization during different tasks in 9 captive grey-cheeked mangabeys (Lophocebus albigena) during spontaneous food processing and 3 experimental tasks. Comparisons with data of red-capped mangabeys (Cercocebus torquatus torquatus; semiterrestrial species) were used. Less than half the monkeys were lateralized for simple everyday activities, but 6 were lateralized for complex daily activities. Moreover, all the monkeys were lateralized when performing experimental tasks. Laterality at the group level was found for the bipedal task: Mangabeys were right-handed. Complexity of tasks increases laterality at the individual level. Significant differences between the 2 species of mangabeys were found, allowing us to confirm that postural constraints are a major factor in the emergence of group handedness.     

Sex differences in spatial task performance of patients with and without unilateral cerebral lesions

Two visuospatial tasks, the WAIS Block Design and the Street Gestalt Completion Test, were administered to men and women with and without unilateral cerebral lesions. These two tasks represent different categories of visuospatial functions. The Street test is a visual-perceptual gestalt task, requiring the closure of fragmented pictures, whereas Block Design is an analytical, manipulospatial task requiring rotation of spatial coordinates. For the non-brain-damaged group, the men showed a nonsignificant trend toward better Block Design performance relative to the women, whereas there was no sex-related difference in Street performance. For the brain-damaged groups, patients with right hemisphere lesions performed significantly worse than patients with left hemisphere lesions on both the Street test and Block Design, indicating that both tasks were more sensitive to right hemisphere functioning. There was, however, a significant sex X side of lesion interaction on Block Design only, with the men showing a more asymmetrical pattern of scores. These results suggest that sex differences in functional lateralization may underlie sex differences in visuospatial ability, and that sex differences in functional lateralization may be present for only certain visuospatial processes.     

Predicting spatial ability from hemispheric ‘non-verbal’ lateralisation: Sex,handedness and task differences implicate encoding strategy effects

Left- and right-handed males and females were given a divided visual field delayed form recognition task and three tests of visuospatial ability of varying degrees of complexity [(a) AH4 diagrammatic section, using 2-D items testing non-verbal reasoning; (b) Revised Minnesota Paper Form Board, using 2-D items testing simple spatial ability; (c) Differential Aptitude Test Space Relations, using 3-D items testing spatial manipulation]. Females of both handedness groups were left hemisphere dominant for the recognition task; males displayed no hemisphere difference. Only the most complex visuospatial test discriminated between groups, both left- right-handed males were superior to their female counterparts. There were no handedness effects for either laterality or spatial ability. Since it was also possible in some cases to predict spatial ability from laterality within sex/handedness groups, the findings were discussed in terms of the contribution of both structural and encoding factors to individual and group differences in laterality and cognitive abilities.     

What clocks tell us about the neural correlates of spatial imagery

We review a series of experimental studies aimed at answering some critical questions about the neural basis of spatial imagery. Our group used functional magnetic resonance imaging (fMRI) to explore the neural correlates of an online behaviourally controlled spatial imagery task without need for visual presentation—the mental clock task. Subjects are asked to imagine pairs of times that are presented acoustically and to judge at which of the two times the clock hands form the greater angle. The cortical activation elicited by this task was contrasted with that obtained during other visual, perceptual, verbal, and spatial imagery tasks in several block design studies. Moreover, our group performed an event‐related fMRI study on the clock task to investigate the representation of component cognitive processes in spatial imagery. The bulk of our findings demonstrates that cortical areas in the posterior parietal cortex (PPC), along the intraparietal sulcus, are robustly involved in spatial mental imagery and in other tasks requiring spatial transformations. PPC is bilaterally involved in different kinds of spatial judgement. Yet the degree to which right and left PPC are activated in different tasks is a function of task requirements. From event‐related fMRI data we obtained evidence that left and right PPC are activated asynchronously during the clock task and this could reflect their different functional role in subserving cognitive components of visuospatial imagery.     

Sex differences in visuospatial working memory: components of cognitive processing

Numerous studies have shown that sex differences in visuospatial tasks vary in size and direction depending on the nature of the task, with large differences favoring males on tasks that require transformations in visuospatial working memory. The cognitive processes underlying these differences were investigated using laboratory tasks developed by Dror and Kosslyn (1994). Four cognitive components of visuospatial working memory were assessed—image generation, maintenance, scanning, and transformation—in an attempt to identify the components that would show differential effects for females and males. The image generation task required retrieval of shape information from long-term memory, generation of a visual image in working memory, and utilization of the information about the shape in a decision task. The image maintenance task required only the latter two processes. The information processing demands required by scanning and rotation tasks came from the need to transform the visual image so that it could be used in decision making. Males responded more quickly on all four tasks (ds between .63 and .77), with no between-sex differences in accuracy. We concluded that speed of processing is central to understanding sex differences in visuospatial working memory. We discuss implications of these findings for performance on real-world visuospatial tasks.     

Sex-related developmental differences in the lateralized activation of the prefrontal cortex and amygdala during perception of facial affect

The lateralization of cognitive abilities is influenced by a number of factors, including handedness, sex, and developmental maturation. To date, a small number of studies have examined sex differences in the lateralization of cognitive and affective functions, and in only few of these have the developmental trajectories of these lateralized differences been mapped from childhood through early adulthood. In the present study, a cross-sectional design was used with healthy children (n=7), adolescents (n= 12), and adults (n= 10) who underwent functional magnetic resonance imaging (fMRI) during a task that required perceiving fearful faces. Males and females differed in the asymmetry of activation of the amygdala and prefrontal cortex across the three age groups. For males, activation within the dorsolateral prefrontal cortex was bilateral in children, right lateralized in adolescents, and bilateral in adults, whereas females showed a monotonic relationship with age, with older females showing more bilateral activation than younger ones. In contrast, amygdala activation was similar for both sexes, with bilateral activation in children, right-lateralized activation in adolescents, and bilateral activation in adults. These results suggest that males and females show different patterns of lateralized cortical and subcortical brain activation across the period of development from childhood through early adulthood.     

An investigation of bilateral asymmetries in electrodermal activity

Research in cerebral laterality supports the idea that functional differences between the left and right hemispheres exist with respect to cognitive style and perceptual ability. Related research, which has examined autonomic nervous system (ANS) correlates of cerebral laterality, suggests that a component of the ANS, electrodermal activity (EDA), is also lateralized. Some findings in the literature report the occurrence of bilateral asymmetries in phasic and tonic EDA as a function of a left or right hemisphere preference for information processing. This experiment used normal male subjects who were either left or right movers in a test of Conjugate Lateral Eye Movement (CLEM). This selection procedure served to maximize a subject’s preference for either a right or left hemisphere mode of information processing. Bilateral EDA was recorded continuously while subjects performed a visual recognition task using word (left hemisphere) and shape (right hemisphere) stimuli. The data do not support the contention that performance on a procedure chosen to selectively activate a given hemisphere elicits asymmetric tonic and/or phasic EDA. The results show no significant difference in the frequency of elicited skin conductance responses under either of the experimental conditions. Bilateral tonic EDA rose continuously over time and did not vary in either hand as a function of task. The present results fail to offer support for either of the hypotheses which argue for I) increased contralateralexcitation with selective hemispheric activation, or 2) increased contralateralinhibition of the EDR with hemispheric arousal.     

Cross-modal correlation of auditory and visual language laterality tasks: a serendipitous finding

Two experiments were conducted to investigate whether auditory and visual language laterality tasks test the same brain processes for verbal functions. In the first experiment, 48 undergraduate students (24 males, 24 females) completed both an auditory monitoring task and a visual monitoring task, with the Waterloo Handedness Questionnaire administered between the two tasks. The visual task was an analogue of the dichotic listening task used. It was hypothesized that a significant cross-modal correlation would be found, indicating that the dichotic listening task and the visual analogue task do, in fact, test the same brain processes for verbal functions. Results revealed a right ear advantage in the auditory task, a left visual field advantage (LVFA) in the visual task, and a cross-modal correlation of asymmetries of -.09. The LVFA observed in the visual task was replicated in Experiment 2, thus establishing its legitimacy. Results are discussed in relation with the type of processing that might produce such an unexpected finding on the visual task.     

Word matching and Lexical Decisions: A Visual Half-Field study

In unilateral Visual Half-Field tasks visuospatial and linguistic processing were compared. In a Word Matching task subjects judged the physical identity of simultaneously presented pairs of three-letter words or legal nonwords. No mainfield effects were found, but word pairs were recognized better and faster as "same" than nonword pairs. Latencies and errors in "different" pairs increased monotonically with position of letter change in the left but not in the right visual field (RVF), suggesting a serial, letter-by-letter way of processing for the right hemisphere and a whole word approach for the left. At this perceptual level the ability to store lexical information from the icon is stressed as a hemisphere-specific factor. In a Lexical Decision task the same subjects judged the same items on the word/nonword dimension. A RVF advantage associated with words as compared to nonwords occurred, as expected. Additional analysis suggests that order and difficulty of tasks may influence females' laterality, as compared to that of males.