Developmental differences in white matter architecture between boys and girls

Previous studies have found developmental differences between males and females in brain structure. During childhood and adolescence, relative white matter volume increases faster in boys than in girls. Sex differences in the development of white matter microstructure were investigated in a cohort of normal children ages 5-18 in a cross-sectional diffusion tensor imaging (DTI) study. Greater fractional anisotropy (FA) in boys was shown in associative white matter regions (including the frontal lobes), while greater FA in girls was shown in the splenium of the corpus callosum. Greater mean diffusivity (MD) in boys was shown in the corticospinal tract and in frontal white matter in the right hemisphere; greater MD in girls was shown in occipito-parietal regions and the most superior aspect of the corticospinal tract in the right hemisphere. Significant sex-age interactions on FA and MD were also shown. Girls displayed a greater rate of fiber density increase with age when compared with boys in associative regions (reflected in MD values). However, girls displayed a trend toward increased organization with age (reflected in FA values) only in the right hemisphere, while boys displayed this trend only in the left hemisphere. These results indicate differing developmental trajectories in white matter for boys and girls and the importance of taking sex into account in developmental DTI studies. The results also may have implications for the study of the relationship of brain architecture with intelligence.     

Sex differences in finger tapping: a developmental study.

The development of serial order in a simple finger-tapping task was examined in normal school age boys and girls from 5 to 16 yr. Girls were consistently more accurate in adhering to the rate of an external entraining beat and in tapping to a steady rhythm. The right hand of all children was steadier than the left, and manual asymmetry for regularity of tapping was greater in girls than boys. The left hemisphere may be specialized for cortical functions controlling the serial organization of simple motor repetitions, and these functions mature earlier in girls than boys.     

Effect of alcohol and task on hemispheric asymmetry of visually evoked potentials in man.

The study examined the effect of a dose of alcohol producing a mean blood alcohol content of 90 mg% on components of the scalp-recorded visually evoked potential (VEP) both with and without a visual discrimination task to control the level of attention, and the interaction of amplitudinal hemispheric asymmetry of the VEP with alcohol treatment and the discrimination task. Ingestion of ethyl alcohol producing a mean blood alcohol content of 90 mg% affected VEPs recorded from the central scalp by attenuating the overall amplitude of the later VEP components (60-200 msec) and by significantly reducing hemispheric asymmetry in the amplitude of these VEP components. Alcohol attenuates VEP components P90-N120 and N120-P180, and the task of counting flashes and attending to discriminate double flashes increased amplitude of VEP components N60-P90 and P90-N120 in control and placebo conditions. Several studies have reported that the VEP recorded from the right hemisphere of human beings is larger than the VEP recorded from the homologous location in the left hemisphere. Evoked potentials recorded under control and placebo conditions in this study also demonstrated a hemispheric asymmetry with right larger than left for component P90-N120. We also found a reliable alcohol by hemispheric asymmetry interaction. Alcohol selectively depressed the amplitude of the right hemisphere VEP (P90-N120) component to a significantly greater extent than the left hemisphere VEP was affected.     

Effects of cortisol on the laterality of the neural correlates of episodic memory

Alterations in the laterality of cortical activity have been shown in depressive illnesses. One possible pathophysiological mechanism for this is an effect of corticosteroids. We have previously demonstrated that endogenous cortisol concentrations correlate with the asymmetry of cortical activity related to episodic memory in healthy subjects and depressed patients. To further-examine whether this is due to a causal effect of cortisol on the laterality of episodic memory, we studied the effect of exogenous administration of cortisol in healthy subjects. Twenty-three right-handed healthy male volunteers were tested in a double-blind cross-over study. Event-related potentials (ERPs) were recorded during an episodic memory task following a four-day course of 160mg/day cortisol or placebo. Low-resolution brain electromagnetic tomography (LORETA) was used to identify brain regions involved in the neurocognitive task. Cortisol levels were measured in saliva samples. ERP and LORETA analysis following placebo demonstrated significant left parahippocampal activation associated with successful retrieval. Cortisol led to a decrease in the mean early frontal ERP voltage and an increase in the late right ERP voltage. LORETA suggested this to be due to a significant increased late activation of the right superior frontal gyrus. There was no significant effect of cortisol on episodic memory performance. This study suggests that exogenous cortisol leads to more positive-going waveforms over the right than the left hemisphere, possibly due to increased monitoring of the products of retrieval. The results support the hypothesis of causal effects of cortisol on the laterality of cortical activity occurring during an episodic memory task.     

Grade-related changes in event-related potentials (ERPs) in primary school children: differences between two reading tasks.

Event-related potentials (ERPs) were recorded from the left and right temporal and parietal sites during word reading in a group of children who were followed over three consecutive years, starting at Grade 1. The first task required the child to read repeatedly presented words; the second task consisted of a series of different words, with each word being presented only once. All ERP components showed changes in amplitude as a function of grade: P240, N530, the parietal N150, and SW all decreased, whereas the temporal N360 and SW both increased. In addition, SW changed from no asymmetry to larger positivity over the right site, whereas N360 changed from symmetric amplitudes to larger amplitudes over the left hemisphere with grade. Single word presentations were associated with smaller SW activity and larger N530 and N360 amplitudes than were repeated word presentations. It was also found that proficient readers showed larger SW and N360 asymmetries and shorter vocal response times than did less proficient readers. In addition, a positive relationship was found between reading performance and ERP amplitudes over the left temporal hemisphere at Grades 2 and 3, particularly in the single word reading task. The results are discussed in terms of age-related changes in right and left hemisphere functions involved in learning to read.     

Abnormal asymmetry in language association cortex in autism

Autism is a neurodevelopmental disorder affecting cognitive, language, and social functioning. Although language and social communication abnormalities are characteristic, prior structural imaging studies have not examined language-related cortex in autistic and control subjects. Subjects included 16 boys with autism (aged 7-11 years), with nonverbal IQ greater than 80, and 15 age- and handedness-matched controls. Magnetic resonance brain images were segmented into gray and white matter; cerebral cortex was parcellated into 48 gyral-based divisions per hemisphere. Asymmetry was assessed a priori in language-related inferior lateral frontal and posterior superior temporal regions and assessed post hoc in all regions to determine specificity of asymmetry abnormalities. Boys with autism had significant asymmetry reversal in frontal language-related cortex: 27% larger on the right in autism and 17% larger on the left in controls. Only one additional region had significant asymmetry differences on post hoc analysis: posterior temporal fusiform gyrus (more left-sided in autism), whereas adjacent fusiform gyrus and temporooccipital inferior temporal gyrus both approached significance (more right-sided in autism). These inferior temporal regions are involved in visual face processing. In boys with autism, language and social/face processing-related regions displayed abnormal asymmetry. These structural abnormalities may relate to language and social disturbances observed in autism.     

Grade-related changes in event-related potentials (ERPs) in primary school children: Differences between two reading tasks

Abstract

Event-related potentials (ERPs) were recorded from the left and right temporal and parietal sites during word reading in a group of children who were followed over three consecutive years, starting at Grade 1. The first task required the child to read repeatedly presented words; the second task consisted of a series of different words, with each word being presented only once. All ERP components showed changes in amplitude as a function of grade: P240, N530, the parietal N150, and SW all decreased, whereas the temporal N360 and SW both increased. In addition, SW changed from no asymmetry to larger positivity over the right site, whereas N360 changed from symmetric amplitudes to larger amplitudes over the left hemisphere with grade. Single word presentations were associated with smaller SW activity and larger N530 and N360 amplitudes than were repeated word presentations. It was also found that proficient readers showed larger SW and N360 asymmetries and shorter vocal response times than did less proficient readers. In addition, a positive relationship was found between reading performance and ERP amplitudes over the left temporal hemisphere at Grades 2 and 3, particularly in the single word reading task. The results are discussed in terms of age-related changes in right and left hemisphere functions involved in learning to read.     

Haptic memory and handedness in 2-month-old infants

This study examined the robustness of infant haptic memory, asymmetry between hands, and sex differences in haptic memory in infancy. A total of 96 2-month-old infants (half males, half females) were habituated haptically to an object with their right and their left hand, out of the field of view. Haptic memory was then tested under three conditions: after haptic interference, after a 30-second delay, or after no delay. The results show that haptic habituation occurred for both hands. The girls needed more time to habituate with their left hand than with their right hand, and they habituated more slowly than the boys did. Discrimination was also found in both hands and in both sexes. Haptic delayed recognition memory was only found in young boys mainly after a short delay and under certain conditions after interference. In young girls, recognition memory was found after interference only with the left hand. This result seemed to depend on the information processing speed. Thus, for memory performance, a sex difference was clearly observed. Moreover, the infants' left hand retained better information on object shape than did the right hand for both the sexes. Asymmetries in infancy are discussed in connection with the difference in brain maturation rate.     

Haptic Memory and Handedness in 2-month-old Infants

This study examined the robustness of infant haptic memory, asymmetry between hands, and sex differences in haptic memory in infancy. A total of 96 2-month-old infants (half males, half females) were habituated haptically to an object with their right and their left hand, out of the field of view. Haptic memory was then tested under three conditions: after haptic interference, after a 30-second delay, or after no delay. The results show that haptic habituation occurred for both hands. The girls needed more time to habituate with their left hand than with their right hand, and they habituated more slowly than the boys did. Discrimination was also found in both hands and in both sexes. Haptic delayed recognition memory was only found in young boys mainly after a short delay and under certain conditions after interference. In young girls, recognition memory was found after interference only with the left hand. This result seemed to depend on the information processing speed. Thus, for memory performance, a sex difference was clearly observed. Moreover, the infants' left hand retained better information on object shape than did the right hand for both the sexes. Asymmetries in infancy are discussed in connection with the difference in brain maturation rate.     

Significant relation between MR measures of planum temporale area and dichotic processing of syllables in dyslexic children

In the present study, we investigated differences between dyslexic and normal reading children in asymmetry of the planum temporale area in the upper posterior part of the temporal lobe and dichotic listening performance to consonant-vowel syllables. The current study was an extension of previous studies in our laboratory on the same participants, now including also girls and left-handers. There were 20 boys and 3 girls in the dyslexic group and 19 boys and 4 girls in the normal reading group. The age of the participants was 10-12 years for both groups. The participants were screened from a population of 950 students in the fourth school grade in the greater Bergen district. The planum temporale area was measured in sagittal magnetic resonance (MR) images. Mean left and right area and asymmetry index were compared between the groups. Dichotic presentations of consonant-vowel syllables made it possible to separately probe left and right hemisphere phonological function, and to correlate this with planum temporale area. The results showed a significantly larger left than right planum temporale area for both groups. However, while the right planum temporale area was similar for the dyslexic and control groups, the left planum temporale was significantly (one-tailed t-test) smaller in the dyslexic group. Both groups also showed a significant right ear advantage to the consonant-vowel syllables in the dichotic listening test. The relation between planum temporale and dichotic listening asymmetry showed a significant correlation for the dyslexic group only, indicating a positive relation between brain structure and function in dyslexic children. The results are discussed in terms of important subject characteristics with regard to brain markers of dyslexia.     

Mouth asymmetry during spontaneous speech

A high proportion of normal subjects have speech expression controlled predominantly by the left hemisphere. Since the left hemisphere also has stronger control of the right side of the lower face, it might be expected that normal subjects would show a right-sided asymmetry in mouth opening during speech. This hypothesis was tested by measuring lip opening in 196 subjects. Of these, 150 (76%) showed greater right-side opening. This tendency was found for males, females, left- and right-handers in four experiments using two different techniques. Mouth asymmetry during speech may provide an indication of which hemisphere is dominant for expressive speech.     

Effects of anticaricaturing vs. caricaturing and their neural correlates elucidate a role of shape for face learning

To assess the role of shape information for unfamiliar face learning, we investigated effects of photorealistic spatial anticaricaturing and caricaturing on later face recognition. We assessed behavioural performance and event-related brain potential (ERP) correlates of recognition, using different images of anticaricatures, veridical faces, or caricatures at learning and test. Relative to veridical faces, recognition performance improved for caricatures, with performance decrements for anticaricatures in response times. During learning, an amplitude pattern with caricatures>veridicals=anticaricatures was seen for N170, left-hemispheric ERP negativity during the P200 and N250 time segments (200-380ms), and for a late positive component (LPC, 430-830ms), whereas P200 and N250 responses exhibited an additional difference between veridicals and anticaricatures over the right hemisphere. During recognition, larger amplitudes for caricatures again started in the N170, whereas the P200 and the right-hemispheric N250 exhibited a more graded pattern of amplitude effects (caricatures>veridicals>anticaricatures), a result which was specific to learned but not novel faces in the N250. Together, the results (i) emphasise the role of facial shape for visual encoding in the learning of previously unfamiliar faces and (ii) provide important information about the neuronal timing of the encoding advantage enjoyed by faces with distinctive shape.     

Measurement of spontaneous rotational movement (circling) in normal children

An asymmetry of basal ganglia dopaminergic function has been demonstrated in rats and related to both spontaneous and drug-induced rotation. An electronic device that measures the same kind of rotational movements in humans has been developed, and we have utilized this "rotometer" to study spontaneous rotational movement in prepubertal children. There was no significant difference between boys and girls in their average rate of rotation; however, left hemisphere-dominant boys were stronger rotators than left hemisphere-dominant girls. Both boys and girls made significantly more full turns to the left than to the right. These findings did not vary with age. Our observations are strikingly different from those obtained in previous studies of normal adults, in which women were stronger rotators than men, left hemisphere-dominant women turned to the left, and left hemisphere-dominant men rotated to the right. This study suggests that maturational changes in rotational behavior must occur, perhaps progressing to the adult pattern during puberty. The rotometer used in this study may provide useful information regarding the status of the basal ganglia in children with specific neurobehavioral conditions such as attention deficit disorder and Tourette's syndrome.     

Pattern reversal evoked potentials: age, sex and hemispheric asymmetry

Binocular pattern reversal evoked potentials (PREPs) and EEG power were recorded bilaterally from occipital scalp of 50 boys and 50 girls aged 5-14. PREP latency was unaffected by either age or gender. Amplitudes, however, were consistently larger for girls than boys, with differences diminishing by adolescence. Significant electrophysiological asymmetries were also found; larger PREP amplitudes were measured at the right occipital electrode than the left for the two waves investigated. These asymmetries were apparent in both girls and boys across all ages, although they tended to diminish with maturation. EEG alpha asymmetry did not correlate with PREP amplitude asymmetry. PREPs elicited by half-field stimulation of 8 girls aged 9-10 years determined that 'paradoxical lateralization' of the half-field response was the same for both left and right half-fields, thus failing to account for the asymmetry of the full-field binocular response.     

The development of lateral event-related potentials (ERPs) related to word naming: a four year longitudinal study

Event-related potentials (ERPs) were recorded from the left and right temporal and parietal sites during a word naming task. Subjects were a group of children that were followed over four consecutive years starting at Kindergarten. ERP waveforms contained a sequence of positive and negative components (N150, P240, N360, N530 and SW). All components, except N150, showed changes in amplitude as a function of age, whereas SW, N360 and N150 also changed in hemispheric distribution. In addition, a relationship was found between reading performance and ERP amplitudes over the right parietal hemisphere in young children, and over the left temporal hemisphere in older children. Proficient readers showed larger (more negative) parietal N530 amplitudes than less proficient readers, especially when stimuli were degraded words. The results are discussed in terms of age-related changes in right and left hemisphere functions involved in early and advanced stages of reading, that might possibly be related to visual word recognition.     

The face-selective N170 component is modulated by facial color

Faces play an important role in social interaction by conveying information and emotion. Of the various components of the face, color particularly provides important clues with regard to perception of age, sex, health status, and attractiveness. In event-related potential (ERP) studies, the N170 component has been identified as face-selective. To determine the effect of color on face processing, we investigated the modulation of N170 by facial color. We recorded ERPs while subjects viewed facial color stimuli at 8 hue angles, which were generated by rotating the original facial color distribution around the white point by 45° for each human face. Responses to facial color were localized to the left, but not to the right hemisphere. N170 amplitudes gradually increased in proportion to the increase in hue angle from the natural-colored face. This suggests that N170 amplitude in the left hemisphere reflects processing of facial color information.     

Event related potentials (ERP) and behavioral measurements to verbal stimulation of visual fields

Event related potentials (ERP) to visually presented linguistic stimuli were examined using a lexical-decision task and an oddball paradigm. Stimuli were presented to the central, right or left visual fields (CVF, RVF and LVF) and generated ERP with very clear N100-P300 components. The question addressed was whether there is ERP evidence for left hemisphere (LH) superiority in linguistic discrimination as reported behaviorally. Nineteen young, right-handed male subjects participated. The main factor influencing the latency and amplitude of N100 was that of contralateral versus ipsilateral stimulation. Shorter N100 latency and larger amplitude were recorded over the hemisphere contralateral to the visual field stimulated. In contrast, the factors influencing the P300 parameters were the visual field stimulated and the hemisphere over which the ERP was recorded. P300 amplitude was significantly larger and P300 latency significantly shorter over the LH than over the RH. Significantly shorter P300 latency and larger peak amplitude were found for RVF than for LVF stimulation.     

Subcortical volumes in girls with tourette syndrome: support for a gender effect

OBJECTIVE: To test whether girls with Tourette syndrome (TS) show subcortical morphology that differentiates them from control subjects. METHODS: MRI-based subcortical assessment was completed on 19 girls with TS age 7 to 15 years, 11 with TS only, and 8 with TS plus attention deficit hyperactivity disorder (TS + ADHD), and on 21 age- and sex-matched controls. The structures measured were the caudate, putamen, globus pallidus, and lateral ventricle volumes. Whole-brain-corrected volumes and asymmetry indices were compared using two- and three-group designs (i.e., TS versus control; TS-only versus TS + ADHD versus control). RESULTS: Two-group comparisons demonstrated no robust significant differences between girls with TS and gender-matched controls. Three-group comparisons demonstrated that TS-only subjects had significantly small lateral ventricles compared with TS + ADHD and control subjects. Because the two-group comparisons of the current study differed from previous reports of putamen asymmetry index as a marker for TS, retrospective comparisons with data from boys were performed. These additional comparisons showed that girls with TS had putamen asymmetry indices similar to those of boys with TS; however, control girls also showed those same patterns. CONCLUSIONS: Basal ganglia volume and asymmetry differences do not distinguish the girls with TS from matched controls. Gender differences confound the association between putamen asymmetry and TS. Although the numbers are small and the clinical significance is unclear, this study further indicates that girls with TS-only have smaller lateral ventricular volumes than control subjects and those with TS + ADHD.     

Asymmetry of flight and escape turning responses in horses

We investigated whether horses display greater reactivity to a novel stimulus presented in the left compared to the right monocular visual field, and whether a population bias exists for escape turning when the same stimulus was presented binocularly. Domestic horses (N=30) were tested on three occasions by a person opening an umbrella five metres away and then approaching. The distance each horse moved away before stopping was measured. Distance was greatest for approach on the left side, indicating right hemisphere control of flight behaviour, and thus followed the same pattern found previously in other species. When order of monocular presentation was considered, an asymmetry was detected. Horses tested initially on the left side exhibited greater reactivity for left approach, whereas horses tested on the right side first displayed no side difference in reactivity. Perhaps left hemisphere inhibition of flight response allowed horses to learn that the stimulus posed no threat and this information was transferred to the right hemisphere. No population bias existed for the direction of escape turning, but horses that turned to the right when approached from the front were found to exhibit longer flight distances than those that turned to the left.     

Hemispheric asymmetry and aging: right hemisphere decline or asymmetry reduction

We review evidence for two models of hemispheric asymmetry and aging: the right hemi-aging model, which proposes that the right hemisphere shows greater age-related decline than the left hemisphere, and the hemispheric asymmetry reduction in old adults (HAROLD) model, which proposes that frontal activity during cognitive performance tends to be less lateralized in older than in younger adults. The right hemi-aging model is supported by behavioral studies in the domains of cognitive, affective, and sensorimotor processing, but the evidence has been mixed. In contrast, available evidence is generally consistent with the HAROLD model, which is supported primarily by functional neuroimaging evidence in the domains of episodic memory encoding and retrieval, semantic memory retrieval, working memory, perception, and inhibitory control. Age-related asymmetry reductions may reflect functional compensation or dedifferentiation, and the evidence, although scarce, tends to support the compensation hypothesis. The right hemi-aging and the HAROLD models are not incompatible. For example, the latter may apply to prefrontal regions and the former to other brain regions.