Functional coupling of parietal α rhythms is enhanced in athletes before visuomotor performance: a coherence electroencephalographic study.

It has been shown that elite pistol shooters are characterized by a power increase of wide cortical electroencephalographic (EEG) alpha (about 8-12 Hz) and beta (about 14-35 Hz) rhythms during the preparation of air pistol shots, possibly related to selective attentional and "neural efficiency" processes [Del Percio C, Babiloni C, Bertollo M, Marzano N, Iacoboni M, Infarinato F, Lizio R, Stocchi M, Robazza C, Cibelli G, Comani S, Eusebi F (2009a) Hum Brain Mapp 30(11):3527-3540; Del Percio C, Babiloni C, Marzano N, Iacoboni M, Infarinato F, Vecchio F, Lizio R, Aschieri P, Fiore A, Toràn G, Gallamini M, Baratto M, Eusebi F (2009b) Brain Res Bull 79(3-4):193-200]. Here, we tested the hypothesis that such processes are associated with an enhanced functional coupling of posterior cortical regions involved in task-relevant attentional processes and visuo-motor transformations. To this aim, between-electrodes spectral coherence was computed from spatially enhanced EEG data collected during a previous study (i.e. right handed 18 elite air pistol shooters and 10 matched non-athletes; augmented 10-20 system; surface Laplacian estimation). Theta (about 4-6 Hz), low-frequency alpha (about 8-10 Hz), high-frequency alpha (about 10-12 Hz), low-frequency beta (14-22 Hz), high-frequency beta (23-35 Hz), and gamma (36-44 Hz) bands were considered. Statistical results showed that intra-hemispheric low-frequency alpha (parietal-temporal and parietal-occipital regions), high-frequency alpha (parietal-temporal and parietal-occipital regions), high-frequency beta, and gamma (parietal-temporal regions) coherence values were stable in amplitude in the elite athletes but not in the non-athletes during the preparation of pistol shots. The same applies to inter-hemispheric low-frequency alpha (parietal regions), high-frequency alpha (parietal regions), high-frequency beta and gamma coherence values. These findings suggest that under the present experimental conditions, elite athletes are characterized by the stabilization of functional coupling of preparatory EEG rhythms between "visuo-spatial" parietal area and other posterior cortical areas.

Effects of tiredness on visuo-spatial attention processes in elite karate athletes and non-athletes.

"Attentional" adaptations are fundamental effects for sport performance. We tested the hypothesis that tiredness and muscular fatigue poorly affect visuo-spatial attentional processes in elite karate athletes. To this aim, 14 elite karate athletes and 11 non-athletes were involved in an isometric contraction exercise protocol up to muscular fatigue. Blood lactate and attention measurements were taken. Posner's test probed "endogenous" (i.e., internally planned allocation of spatial attention) and "reflexive" (i.e., brisk variation of endogenous spatial attention due to unexpected external stimuli) attention. Lactate and attentional measurements were performed before (Block 1, B1) and after the fatiguing exercise (B2) and at the end of a recovery period (B3). Compared to the non-athletes, the athletes showed a better performance in the fatigue protocol, confirmed by the higher absolute lactate values in B2. The correct responses in the "valid trials" probing "endogenous" attention were 92.4% (B1), 93.9% (B2), and 95.8% (B3) in the non-athletes, and 98.5%, 96.4%, 95.5% in the elite karate athletes. The correct responses in the "invalid trials" probing "reflexive" attention were 95.4%, 89.7%, 93.2% in the non-athletes, and 96.4%, 97.3%, 98.5% in the elite karate athletes. The percentage of correct responses in the "invalid" trials significantly decreased from B1 to B2 in the non-athletes but not in the elite karate athletes. In conclusion, tiredness and muscular fatigue do not affect "reflexive" attentional processes of elite karate athletes, which is crucial to contrast attacks coming from an unexpected spatial region.

Neural basis of maternal communication and emotional expression processing during infant preverbal stage.

During the first year of life, exchanges and communication between a mother and her infant are exclusively preverbal and are based on the mother's ability to understand her infant's needs and feelings (i.e., empathy) and on imitation of the infant's facial expressions; this promotes a social dialog that influences the development of the infant self. Sixteen mothers underwent functional magnetic resonance imaging while observing and imitating faces of their own child and those of someone else's child. We found that the mirror neuron system, the insula and amygdala were more active during emotional expressions, that this circuit is engaged to a greater extent when interacting with one's own child, and that it is correlated with maternal reflective function (a measure of empathy). We also found, by comparing single emotions with each other, that joy expressions evoked a response mainly in right limbic and paralimbic areas; by contrast, ambiguous expressions elicited a response in left high order cognitive and motor areas, which might reflect cognitive effort.

Reafferent copies of imitated actions in the right superior temporal cortex.

Imitation is a complex phenomenon, the neural mechanisms of which are still largely unknown. When individuals imitate an action that already is present in their motor repertoire, a mechanism matching the observed action onto an internal motor representation of that action should suffice for the purpose. When one has to copy a new action, however, or to adjust an action present in one's motor repertoire to a different observed action, an additional mechanism is needed that allows the observer to compare the action made by another individual with the sensory consequences of the same action made by himself. Previous experiments have shown that a mechanism that directly matches observed actions on their motor counterparts exists in the premotor cortex of monkeys and humans. Here we report the results of functional magnetic resonance experiments, suggesting that in the superior temporal sulcus, a higher order visual region, there is a sector that becomes active both during hand action observation and during imitation even in the absence of direct vision of the imitator's hand. The motor-related activity is greater during imitation than during control motor tasks. This newly identified region has all the requisites for being the region at which the observed actions, and the reafferent motor-related copies of actions made by the imitator, interact.

A probabilistic atlas and reference system for the human brain: International Consortium for Brain Mapping (ICBM).

Motivated by the vast amount of information that is rapidly accumulating about the human brain in digital form, we embarked upon a program in 1992 to develop a four-dimensional probabilistic atlas and reference system for the human brain. Through an International Consortium for Brain Mapping (ICBM) a dataset is being collected that includes 7000 subjects between the ages of eighteen and ninety years and including 342 mono- and dizygotic twins. Data on each subject includes detailed demographic, clinical, behavioural and imaging information. DNA has been collected for genotyping from 5800 subjects. A component of the programme uses post-mortem tissue to determine the probabilistic distribution of microscopic cyto- and chemoarchitectural regions in the human brain. This, combined with macroscopic information about structure and function derived from subjects in vivo, provides the first large scale opportunity to gain meaningful insights into the concordance or discordance in micro- and macroscopic structure and function. The philosophy, strategy, algorithm development, data acquisition techniques and validation methods are described in this report along with database structures. Examples of results are described for the normal adult human brain as well as examples in patients with Alzheimer's disease and multiple sclerosis. The ability to quantify the variance of the human brain as a function of age in a large population of subjects for whom data is also available about their genetic composition and behaviour will allow for the first assessment of cerebral genotype-phenotype-behavioural correlations in humans to take place in a population this large. This approach and its application should provide new insights and opportunities for investigators interested in basic neuroscience, clinical diagnostics and the evaluation of neuropsychiatric disorders in patients.

Experience-dependent modulation of motor corticospinal excitability during action observation.

The primate premotor cortex is endowed with an "action observation/execution matching system", that is, the same premotor neurons discharge when actions are performed and when actions are observed. Hence, this system predicts a strong visual input to the motor system. Whether this input is dependent on visual experience or not has not been previously investigated. To address this issue we compared corticospinal excitability while subjects viewed frequently observed and less frequently observed hand actions of others and of themselves. Motor corticospinal excitability was larger when the action orientations were as they are frequently observed (Self-away, subject's own hand facing out from the subject, or Other-toward, an unknown hand facing toward the subject) compared with less frequently observed actions (Self-toward, subject's own hand facing "toward" the subject, or Other-away, an unknown hand facing out from the subject). This finding suggests that the modulation of motor corticospinal excitability during action observation and hence the "action observation/execution matching system" is largely dependent upon visual experience.

A four-dimensional probabilistic atlas of the human brain.

The authors describe the development of a four-dimensional atlas and reference system that includes both macroscopic and microscopic information on structure and function of the human brain in persons between the ages of 18 and 90 years. Given the presumed large but previously unquantified degree of structural and functional variance among normal persons in the human population, the basis for this atlas and reference system is probabilistic. Through the efforts of the International Consortium for Brain Mapping (ICBM), 7,000 subjects will be included in the initial phase of database and atlas development. For each subject, detailed demographic, clinical, behavioral, and imaging information is being collected. In addition, 5,800 subjects will contribute DNA for the purpose of determining genotype- phenotype-behavioral correlations. The process of developing the strategies, algorithms, data collection methods, validation approaches, database structures, and distribution of results is described in this report. Examples of applications of the approach are described for the normal brain in both adults and children as well as in patients with schizophrenia. This project should provide new insights into the relationship between microscopic and macroscopic structure and function in the human brain and should have important implications in basic neuroscience, clinical diagnostics, and cerebral disorders.

Parallel visuomotor processing in the split brain: cortico-subcortical interactions.

We tested nine patients with callosal pathology in a simple reaction time task with and without redundant targets in the same or opposite visual hemifield. Four patients showed large facilitation (redundancy gain) in the presence of a redundant target, exceeding probability summation models (neural summation). Five patients showed redundancy gain not exceeding probability models. Violation of probability models was not associated with a specific type of callosal lesion. Neural summation, which probably occurs at collicular level, may be modulated by cortical activity. To test this hypothesis, we used functional MRI. During detection of redundant simultaneous targets, activations in the extrastriate cortex were observed in a patient with callosal agenesis and redundancy gain violating probability models, but not in a patient with callosal agenesis and redundancy gain not exceeding probability models. We conclude that cortical activity in the extrastriate cortex may be a modulating factor in the magnitude of the redundancy gain during parallel visuomotor transforms.

Crossed-uncrossed difference in simple reaction times to lateralized flashes: between- and within-subjects variability.

In unimanual reaction times (RT) to lateralized flashes, contralateral responses tend to be slower than ipsilateral responses. This has been called Crossed-Uncrossed Difference (CUD). The CUD tends to show variability across subjects and across studies, but until now the stability of the CUD in an individual subject has not been investigated. To address the role of inter- and intra-subject variability in the CUD, three normal right handers were tested over 50 experimental sessions of 800 trials each, for a total of 40,000 trials of simple reaction times to lateralized flashes. In each subject, CUDs were computed for each session, over two, three, or more sessions, and over the entire dataset. These CUDs were then compared to the CUDs obtained in a group of 15 normal right handers, each tested once in a single session. Results show that: (i) CUD variability across several sessions in a single subject mimics the variability observed in a sample of subjects tested in a single session; (ii) this variability is considerably reduced when the CUD is computed over at least 2400 trials per subject; (iii) CUDs computed over 2400 and up to 12,000 of trials tend to be extremely similar ( approximately 2 ms) across the three subjects tested here; (iv) when reaction times are ordered from the fastest to the slowest and divided into bins, the CUD is remarkably stable over the entire reaction time distribution; and (v) in contrast to the variability of the CUD, the variability for crossed and uncrossed responses across several sessions in a single subject is small and does not mimic the variability observed in a sample of subjects tested in a single session. Taken together, these data suggest that the intersubject variability in the CUD observed in single experimental sessions does not represent a reliable intersubject difference and that the CUD computed over thousands of trials reflects hard-wired mechanisms of callosal transmission.

Synthetic brain imaging: grasping, mirror neurons and imitation.

The article contributes to the quest to relate global data on brain and behavior (e.g. from PET, Positron Emission Tomography, and fMRI. functional Magnetic Resonance Imaging) to the underpinning neural networks. Models tied to human brain imaging data often focus on a few "boxes" based on brain regions associated with exceptionally high blood flow, rather than analyzing the cooperative computation of multiple brain regions. For analysis directly at the level of such data, a schema-based model may be most appropriate. To further address neurophysiological data, the Synthetic PET imaging method uses computational models of biological neural circuitry based on animal data to predict and analyze the results of human PET studies. This technique makes use of the hypothesis that rCBF (regional cerebral blood flow) is correlated with the integrated synaptic activity in a localized brain region. We also describe the possible extension of the Synthetic PET method to fMRI. The second half of the paper then exemplifies this general research program with two case studies, one on visuo-motor processing for control of grasping (Section 3 in which the focus is on Synthetic PET) and the imitation of motor skills (Sections 4 and 5, with a focus on Synthetic fMRI). Our discussion of imitation pays particular attention to data on the mirror system in monkey (neural circuitry which allows the brain to recognize actions as well as execute them). Finally, Section 6 outlines the immense challenges in integrating models of different portions of the nervous system which address detailed neurophysiological data from studies of primates and other species; summarizes key issues for developing the methodology of Synthetic Brain Imaging; and shows how comparative neuroscience and evolutionary arguments will allow us to extend Synthetic Brain Imaging even to language and other cognitive functions for which few or no animal data are available.

The crossed-uncrossed difference in simple reaction times to lateralized auditory stimuli is not a measure of interhemispheric transmission time: evidence from the split brain.

In a complete commissurotomy patient, the difference in simple (detection) reaction times between responses to contralateral and ipsilateral auditory stimuli was found to be small (less than 5 ms) and not reliable, whereas the difference between contralateral and ipsilateral responses to lateralized visual stimuli was found to be large (ranging from 25 ms to 45 ms in different previous studies) and always reliable. This suggests that the reaction times difference in detecting lateralized auditory stimuli is not a valid estimate of interhemispheric transmission time.

Emission tomography contribution to clinical neurology.

The role of functional neuroimaging techniques in furthering the understanding of pathophysiological mechanisms of neurological diseases and in the assessment of neurological patients is increasingly important. Here, we review data mainly from emission tomography techniques, namely positron emission tomography (PET) and single photon emission computerized tomography (SPECT), that have helped elucidate the pathophysiology of a number of neurological diseases and have suggested strategies in the treatment of neurological patients. We also suggest possible future developments of functional neuroimaging applied to clinical populations and briefly touch on the emerging role of functional magnetic resonance imaging (fMRI) in clinical neurology and neurosurgery.

Individual differences in the hemispheric specialization of dual route variables.

The dual route model suggests that reading of letter strings can occur through both a lexical and a nonlexical route. Hemispheric specialization of these routes has also been posited, suggesting that the left hemisphere has both lexical and nonlexical routes while the right hemisphere has only a lexical route. However, some recent data conflict with this hemispheric dual route model, suggesting that both hemispheres may have access to both routes. The purpose of the present study was to investigate individual differences in the hemispheric specialization of these routes and to determine whether these group differences in their specialization might explain conflicts in the literature. The effect of four individual difference factors was explored: handedness, biological sex, menstrual stage (i.e., fluctuations in estrogen), and self-rated degree of masculinity (i.e., sexual attribution). We looked at the interaction of these individual differences with the following dual route variables: (i) string length, (ii) word frequency, (iii) regularity of grapheme-phoneme correspondences of words, and (iv) the interaction of frequency and regularity using a bilateral lexical decision task. We observed that sex, menstrual stage, and masculinity each affected hemispheric specialization of the dual route variables, but did so in different ways. We posit that both hemispheres have orthographical (lexical) access as well as phonological (nonlexical) access to words. Further, we suggest that the presence of phonological processing in the right hemisphere depends on available resources and the strategies used, which are subject to individual differences.

Cortical mechanisms of human imitation.

How does imitation occur? How can the motor plans necessary for imitating an action derive from the observation of that action? Imitation may be based on a mechanism directly matching the observed action onto an internal motor representation of that action ("direct matching hypothesis"). To test this hypothesis, normal human participants were asked to observe and imitate a finger movement and to perform the same movement after spatial or symbolic cues. Brain activity was measured with functional magnetic resonance imaging. If the direct matching hypothesis is correct, there should be areas that become active during finger movement, regardless of how it is evoked, and their activation should increase when the same movement is elicited by the observation of an identical movement made by another individual. Two areas with these properties were found in the left inferior frontal cortex (opercular region) and the rostral-most region of the right superior parietal lobule.

Bimodal (auditory and visual) left frontoparietal circuitry for sensorimotor integration and sensorimotor learning.

We used PET to test whether human premotor and posterior parietal areas can subserve basic sensorimotor integration and sensorimotor learning equivalently in response to auditory and visual stimuli, as has been shown in frontoparietal neurons in non-human primates. Normal subjects were studied while they performed a spatial compatibility task. They were instructed to respond to lateralized auditory and visual stimuli with the ipsilateral hand (compatible condition) or with the contralateral hand (incompatible condition). Reaction times were faster in the compatible than in the incompatible condition, for both auditory and visual stimuli. Left rostral dorsal premotor and posterior parietal blood-flow increases were observed in the incompatible condition, compared with the compatible condition, for both auditory and visual modalities. Blood-flow increases, which were correlated with the reaction-time learning curves, were observed in both auditory and visual modalities in the left caudal dorsal premotor cortex. These data suggest that, as in non-human primates, human frontoparietal areas can subserve basic sensorimotor transformations equivalently in the auditory and visual modality. Further, they reveal a functional rostrocaudal fractionation of human dorsal premotor cortex that resembles the rostrocaudal anatomical and physiological fractionation observed in non-human primates.

Merging of oculomotor and somatomotor space coding in the human right precentral gyrus.

Using PET and H215O, we investigated the cortical areas that merge two different ways of coding space in the cerebral cortex, those concerned with the oculomotor and the somatomotor space. Normal subjects performed a visuomotor task that required the spatial coding of visual stimuli in oculomotor space and of motor responses in somatomotor space. We manipulated the mapping of oculomotor and somatomotor space by instructing subjects to respond in half of the PET scans with uncrossed hands, i.e. each hand was in the homonymous hemispace (standard oculomotor-somatomotor mapping), and in the other half with crossed hands, i.e. with the left hand in the right hemispace and the right hand in the left hemispace (nonstandard oculomotor-somatomotor mapping). Reaction times were slower for crossed hands than uncrossed hands. Crossed hands produced increases in blood flow in the precentral and postcentral gyri of the right hemisphere. Increases in blood flow in the precentral gyrus were correlated with increases in reaction time comparing the crossed-hand task with the uncrossed one, whereas the increases in blood flow in the postcentral gyrus were not. These findings suggest that the right precentral gyrus merges oculomotor and somatomotor space coding in the human brain.

Does the previous trial affect lateralized lexical decision?

We investigated the effect of previous trial variables on performance in the current trial in a lexical decision task with unilateral presentation of one letter string or bilateral simultaneous presentation of two different letter strings, one cued to be processed (target) and the other uncued, to be ignored (distractor). The variables included correctness of the previous trial, visual hemifield and wordness of the previous trial, and presentation mode of the previous trial (unilateral or bilateral). An incorrect response on the previous trial enhanced the accuracy in the current trial only in the left visual field (LVF). A previous LVF target produced faster correct responses to LVF targets in the current trial and LVF word recognition was more accurate when the previous LVF target was a word rather than a nonword. Target processing in the current trial was not inhibited or facilitated if the target belonged to the same response category as the unattended stimulus in the previous trial (absence of "negative priming'). Taken together, our data suggest that previous trial effects in lateralized lexical decision are stronger for word decisions in the LVF, and may account for the inconsistency of the wordness effect in the LVF across different lateralized lexical decision experiments. Our data also suggest that behavioral laterality experiments are well advised to use random sequences that change across subjects in order to minimize previous trial effects.