Visual stimulation elicits locked and induced gamma oscillations in monkey intracortical- and EEG-potentials, but not in human EEG

Stimulus-related fast oscillations in the γ-range (30–100 Hz) were clearly demonstrated with microelectrode recordings in visual cortex of awake monkeys, and they were also reported for recordings of human electroencephalograms (EEG). However, the presence of stimulus-related γ-modulation in human EEG has repeatedly been disputed. To clarify this dispute, we recorded the scalp EEG of man and monkey as well as intracortical field potentials (LFP) from monkey primary visual cortex (V1) during identical visual stimulation (large-field sinusoidal gratings, which proved to induce the largest γ-amplitudes in monkey V1 and V2). We found a strong stimulus-related increase of γ-oscillations in monkey LFP and EEG, but no modulation of γ-activity in human EEG. In contrast to previous results, γ-oscillations in the monkey were strongly phase-locked to stimulus onsets in early response periods (80–160 ms) and became gradually independent in later periods. Our negative result on γ-modulation in human subjects contradicts several published findings. We conclude from our results that visually evoked γ-modulations in humans EEG are not as accessible as in the monkey. Received: 31 August 1998 / Accepted: 29 June 1999     

Cortical processing of tactile stimuli applied in quick succession across the fingertips: temporal evolution of dipole sources revealed by magnetoencephalography

We used magnetoencephalography (MEG) in 10 healthy human subjects to study cortical responses to tactile stimuli applied to the fingertips of digits 2–5 of the right hand. Each stimulus lasted 50 ms and was produced by air-driven elastic membranes. Four-hundred stimuli were delivered on each finger in three temporal patterns (conditions). In the “Discrete” condition, stimuli were applied to each finger repetitively with an interstimulus interval (ISI) of 1–2 s. In the “Continuous” condition, stimuli were applied to the fingers sequentially as four-stimulus trains with zero ISI and 1–2 s intervening between trains. Finally, in the “Gap” condition, stimuli were applied as in the Continuous condition but with an ISI of 50 ms. A sensation of tactile motion across fingers (digit 2 → digit 5) was reported by all subjects in the Continuous and Gap conditions. Cortical responses were extracted as single equivalent current dipoles over a period of 1 s following stimulus onset. In all three conditions, initial responses in left primary somatosensory cortex (SI) were observed ~20 to 50 ms after stimulus onset and were followed by additional left SI responses and bilateral responses in the secondary somatosensory cortex (SII). In addition, in the Continuous and Gap conditions, there was an activation of the precentral gyrus, the temporal aspects of which depended on the temporal relation of the administered stimuli, as follows. An ISI of 0 ms led to activation of the precentral gyrus shortly after the second stimulation, whereas an ISI of 50 ms led to activation of the precentral gyrus after the third stimulation. The current findings support results from previous studies on temporal activity patterns in SI and SII, verify the participation of the precentral gyrus during tactile motion perception and, in addition, reveal aspects of integration of sequential sensory stimulations over nonadjacent areas as well as temporal activity patterns in the postcentral and precentral gyri.     

Neuronal responses from beyond the classic receptive field in V1 of alert monkeys

Responses of primary visual cortex (V1) neurons to stimuli inside the classic receptive field (CRF) can be modulated by stimuli outside the CRF. We recently reported that responses of most V1 neurons to a line in the CRF center are inhibited by large surround-stimuli and that this modulation is stimulus selective. Here we report that a significant proportion of V1 neurons in alert monkeys respond directly to stimuli outside the CRF with very long latency and much reduced selectivity. When surround stimuli are presented alone, three response patterns can be distinguished in 153 single- or multiunits tested: (1) 31.4% have no significant response; (2) 50.3% show excitatory responses that are significantly higher than spontaneous activity. The average latency of these responses is about 145 ms, 2–3 times longer than center responses; (3) 18.3% show suppressed spontaneous activity after stimulus onset. The direct surround responses are found to be only weakly selective for the orientation of contextual lines, and not selective for other contextual patterns tested. While the outburst of responses to stimuli within the CRF is not affected by reducing stimulus duration from 500 ms to 50 ms, late excitatory surround responses are virtually eliminated. We propose that the late excitatory surround responses to extra-CRF stimulation alone are the reflection of feedback from higher cortical areas and may contribute to reduced contextual inhibition of cells in V1. This could play a role in figure-ground segregation. Electronic Publication     

Transient visual field defects induced by transcranial magnetic stimulation over human occipital pole

Transient visual field defects (VFDs) and phosphenes were induced in normal volunteers by means of transcranial magnetic stimulation (TMS) using a circular magnetic coil of 12.5 cm diameter placed with its lower rim 2–4 cm above the inion in the midline. Subjects had to detect small, bright dots presented randomly for 14 ms in one of 60 locations on a computer screen resulting in a plot of the central 9° of the visual field. In 8 of 17 subjects, transient VFDs were inducible at peak magnetic field strenghts of 1.1–1.4 T. In the central 1–3°, detection of targets was impaired in both the upper and lower visual field, whereas at 4–9° large parts of only the lower visual field were affected with a sharp cut-off along the horizontal meridian. Targets at 1° in the lower field were affected with lower TMS intensities than corresponding locations in the upper or peripheral locations in the lower field. Detection of central targets was affected at more caudal stimulation sites than detection of peripheral targets. Phosphenes were elicitable in 14 of 17 subjects at clearly lower field strengths of 0.6–1.0 T. Many subjects perceived chromatophosphenes. From a discussion of the literature on patients with VFDs and the known topography of the human visual system, it is concluded that the transient VFDs at 1–3° are probably due to stimulation of both striate cortex (V1) and extrastriate areas (V2/V3), while VFDs in the lower visual field at eccentricities 4–9° are due to stimulation of V2/V3 but not V1. Received: 14 January 1997 / Accepted: 2 June 1997     

Microstimulation of V1 delays visually guided saccades: a parametric evaluation of delay fields

Electrical microstimulation of macaque striate cortex (area V1) delays the execution of saccadic eye movements made to a visual target placed in the receptive field of the stimulated neurons. The region of visual space within which saccades are delayed is called a delay field. We examined the effects of changing the parameters of stimulation and target size on the size of a delay field. Rhesus monkeys were required to generate a saccadic eye movement to a punctate and white visual target presented within or outside the receptive field of the neurons under study. On 50% of trials, a train of stimulation consisting of 0.2-ms anode-first pulses was delivered to the neurons before the onset of the visual target. Stimulations were performed in the operculum at 0.9–2.0 mm below the cortical surface. It was found that increases in current (50–100 μA), pulse frequency (100–300 Hz), or train duration (75–300 ms) increased the size of a delay field and increases in target size (0.1°–0.2° of visual angle) decreased the size of a delay field. Delay fields varied in size between 0.1 and 0.6° of visual angle. These results are related to the properties of phosphenes induced by electrical stimulation of V1 in humans and compared to the interference effects observed following transcranial magnetic stimulation of human V1.     

Limited plasticity of difference neurons in the visual cortex and hippocampus in rabbits during the oddball (random substitutions) test

The activity of 41 visual cortex neurons and 20 hippocampal field CA1 neurons was studied in rabbits during application of the oddball stimulation paradigm using color stimuli of different intensities. Among these cells, about one third were plastic cells (34% of cortical cells and 37% of hippocampal cells). These neurons showed significant increases in late responses, at times 200–500 and 200–1000 msec for visual cortex neurons and 300–550 msec for hippocampal neurons, to rare deviant stimuli of lesser intensity as compared with responses to the frequent standard stimuli of greater intensity. The initial peak of the response (40–120 msec), the “difference discharge,” remained stable in responses to deviant and standard stimuli throughout the experiment. It is suggested that the strengthening of the late components of neuron responses to rare deviant stimuli (limited plasticity) reflects inclusion of the mechanisms of the orientational reflex. __________ Translated from Zhurnal Vysshei Nervnoi Deyatel’nosti imeni I. P. Pavlova, Vol. 55, No. 3, pp. 360–367, May–June, 2005.     

Computation of Color and Brightness Differences by Rabbit Visual Cortex Neurons

Extracellular recording of the activity of 54 neurons in the rabbit visual cortex in responses to substitutions of eight colored and eight monochromatic stimuli in pairs was studied. Stimuli were uniform flashes of light displayed on an SVGA monitor and illuminated the whole retina. The responses of phasic neurons showed an initial discharge (50–90 msec from the moment of the change in stimulus), associated with the brightness or color difference between the stimuli. These “discrimination discharges” were used to construct an 8 × 8 matrix for each neuron, showing the mean number of spikes per sec in responses to changes in different pairs of stimuli. Processing of the matrix by factor analysis identified the major factors determining the axes of the sensory space. A brightness space with only two dimensions, with darkness and brightness orthogonal axes, was seen for 30% of neurons. A four-dimensional color space was seen in 22% of neurons, with two color and two achromatic axes. The sensory space of these neurons was similar to the spaces obtained by analyzing the early components of visual evoked potentials in rabbits induced by changes in color stimuli and behavioral operant responses in conditioned reflex color differentiation. The fundamental coincidence of the sensory spaces obtained by different methods identifies the general nature of the principle of vector coding and the existence of special neuronal mechanisms for detection of color and brightness differences in the visual field. __________ Translated from Zhurnal Vysshei Nervnoi Deyatel'nosti imeni I. P. Pavlova, Vol. 55, No. 1, pp 60–70, January–February, 2005.     

Auditory evoked visual awareness following sudden ocular blindness: an EEG and TMS investigation

Neurophysiological and neuroanatomical studies have provoked controversy about whether the visual cortex may be more modifiable than previously believed. Auditory processing is enhanced in blind compared to sighted people, and the enhancement might reflect encroachment of auditory transmission onto visual cortex. To address this issue, we recorded the auditory event-related potentials (ERPs) correlated with auditory related paradoxical visual awareness in a subject with traumatic total late-onset blindness. We found that (1) there was auditory related brain activity over the occipital visual scalp regions starting from a very early stage (< 80 ms) and (2) this occipital activity was significantly different between “visually aware” and “visually unaware” responses in the P1 (40–80 ms) component following meaningful stimuli. There was also a significant difference between responses with and without visual awareness in the N1 (100–120 ms) component following either tones or meaningful stimuli. The phosphenes accompanying auditory stimuli in the ERP experiment were always perceived to be directly in front of the subject and this was reproduced by transcranial magnetic stimulation over the blind primary visual cortex and by sudden sounds delivered to the side or behind the subject. The TMS induced phosphenes were restricted to the central part of the space and were, at least qualitatively, the same as those induced by sounds. The results are clear evidence that human perceptual functions can be reorganized after sudden, late-onset, total ocular blindness.     

Early extrastriate activity without primary visual cortex in humans

Damage to the primary visual cortex (V1) destroys the major source of anatomical input to extrastriate cortical areas (V2, V3, V4 and V5) and produces cortical blindness--an absence of any sensation of light and colour--in the visual field contralateral to the side of the lesion. Neuroimaging studies, nevertheless, have recently demonstrated dorsal and ventral extrastriate activation for stationary stimuli presented to the blind visual field in the absence of V1 activity in human subjects. To clarify the moment in time that visual information reaches extrastriate areas, by means of event-related potentials (ERPs) we tracked the temporal course of responses to complex visual stimuli (faces) presented in the blind field of a hemianopic patient. Stimulation of the normal visual field elicited a positive occipital deflection (P1) at 140 ms. A P1 response was also observed with stimulation of the blind field, although slightly delayed (20 ms) and reduced. Its topography and timing demonstrate that early neural activity for stationary stimuli takes place within extrastriate regions despite V1 denervation.     

Occipital network for figure/ground organization

To study the cortical mechanism of figure/ground categorization in the human brain, we employed fMRI and the temporal-asynchrony paradigm. This paradigm is able to eliminate any differential activation for local stimulus features, and thus to identify only global perceptual interactions. Strong segmentation of the image into different spatial configurations was generated solely from temporal asynchronies between zones of homogeneous dynamic noise. The figure/ground configuration was a single geometric figure enclosed in a larger surround region. In a control condition, the figure/ground organization was eliminated by segmenting the noise field into many identical temporal-asynchrony stripes. The manipulation of the type of perceptual organization triggered dramatic reorganization in the cortical activation pattern. The figure/ground configuration generated suppression of the ground representation (limited to early retinotopic visual cortex, V1 and V2) and strong activation in the motion complex hMT+/V5+; conversely, both responses were abolished when the figure/ground organization was eliminated. These results suggest that figure/ground processing is mediated by top-down suppression of the ground representation in the earliest visual areas V1/V2 through a signal arising in the motion complex. We propose a model of a recurrent cortical architecture incorporating suppressive feedback that operates in a topographic manner, forming a figure/ground categorization network distinct from that for “pure” scene segmentation and thus underlying the perceptual organization of dynamic scenes into cognitively relevant components. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect of stimulus probability on the somatosensory mismatch field

We investigated the effect of deviant stimulus probability on the somatosensory magnetic mismatch negativity (MMNm) using an electrical two-point stimulation. First, we determined the discrimination threshold (DT) of the two-point distance. We applied standard stimuli at a distance that subjects felt as one point and deviant stimuli at a distance that subjects definitely felt as two points. We used three deviant stimulus probabilities, 10, 30, and 50%. The components peaking around 30–70 ms (first component) and 150–250 ms (fourth component) following deviant stimuli were significantly larger than those following standard stimuli in 10% condition, but not in 30 or 50% condition. The equivalent current dipole (ECD) was located in the contralateral primary somatosensory cortex (cSI) for the first component, and in the cSI and in the contralateral secondary somatosensory cortex (cSII) for the fourth component. The peak amplitude of the MMNm decreased as the probability of the deviant stimulus increased. The Somatosensory MMNm was affected by deviant stimulus probability similar to an auditory mismatch negativity (MMN).     

Multiple temporal components of optic flow responses in MST neurons

 Neurons in monkey medial superior temporal cortex (MST) respond to optic flow stimuli with early phasic, tonic, and after-phasic response components. In these experiments we characterized each response component to compare its potential contributions to visual motion processing. The early responses begin 60–100 ms after stimulus onset and last between 100 and 250 ms, the tonic responses begin 100–300 ms after stimulus onset and last for as long as the evoking stimulus persists, and the after-responses begin about 60 ms after the stimulus goes off and last for 100–350 ms. A neuron’s tonic responses were evoked by specific optic flow stimuli: over two-thirds of the 264 neurons showed tonic responses evoked by two to five stimuli, whereas only 15% responded to either all or none of the stimuli. The tonic responses continued with stimulus presentations as long as 15 s, with their directional preferences being maintained throughout stimulation. However, the tonic response to a given stimulus was seen to change in amplitude when it was presented in random sequence with different sets of other stimuli. Thus, the tonic responses might convey substantial information about optic flow patterns, which continue with prolonged stimulation, but can be modified by the visual context created by other visual motion stimuli. Only about one-third of the 264 neurons had early responses that were selective for specific stimuli. In neurons yielding at least one early response, that neuron was most often activated by all the visual motion stimuli. After-rsponses occurred in only half the neurons, but they were more often specifically related to particular optic flow stimuli, regardless of whether those stimuli had evoked tonic excitatory or tonic inhibitory responses. The presence of early and after-responses complicates the interpretation of activity evoked when one stimulus immediately follows another. However, under those conditions, early responses and after-responses might contribute to signaling changes in the ongoing pattern of optic flow. We conclude that several components of MST responses should be recognized and that they potentially play different roles in the cortical analysis of optic flow. Tonic responses show the greatest specificity for particular optic flow stimuli, and possess characteristics which make them suitable neuronal participants in self-movement perception. Received: 1 April 1996 / Accepted: 1 October 1996     

Parvocellular and Magnocellular Contributions to the Initial Generators of the Visual Evoked Potential: High-Density Electrical Mapping of the “C1” Component

The C1 component of the VEP is considered to index initial afference of retinotopic regions of human visual cortex (V1 and V2). C1 onsets over central parieto–occipital scalp between 45 and 60 ms, peaks between 70 and 100 ms, and then resolves into the following P1 component. By exploiting isoluminant and low-contrast luminance stimuli, we assessed the relative contributions of the Magnocellular (M) and Parvocellular (P) pathways to generation of C1. C1 was maximal at 88 ms in a 100% luminance contrast condition (which stimulates both P and M pathways) and at 115 ms in an isoluminant chromatic condition (which isolates contributions of the P pathway). However, in a 4% luminance contrast condition (which isolates the M pathway), where the stimuli were still clearly perceived, C1 was completely absent. Absence of C1 in this low contrast condition is unlikely to be attributable to lack of stimulus energy since a robust P1–N1 complex was evoked. These data therefore imply that C1 may be primarily parvocellular in origin. The data do not, however, rule out some contribution from the M system at higher contrast levels. Nonetheless, that the amplitude of C1 to P-isolating isoluminant chromatic stimuli is equivalent to that evoked by 100% contrast stimuli suggests that even at high contrast levels, the P system is the largest contributor. These data are related to intracranial recordings in macaque monkeys that have also suggested that the initial current sink in layer IV may not propagate effectively to the scalp surface when M-biased stimuli are used. We also discuss how this finding has implications for a long tradition of attention research that has␣used C1 as a metric of initial V1 afference in humans. C1 has been repeatedly interrogated for potential selective attentional modulations, particularly in spatial attentional designs, under the premise that modulation of this component, or lack thereof, would be evidence for or against selection at the initial inputs to visual cortex. Given the findings here, we would urge that in interpreting C1 effects, a consideration of the dominant cellular contributions will be necessary. For example, it is plausible that spatial attention mechanisms could operate primarily through the M system and that as such C1 may not always represent an adequate dependent measure in such studies.     

Neuronal activity of somatosensory cortex in a cross-modal (visuo-haptic) memory task

Studies have shown that in the monkey′s associative cerebral cortex, cells undergo sustained activation of discharge while the animal retains information for a subsequent action. Recent work has revealed the presence of such ″memory cells″ in the anterior parietal cortex (Brodmann′s areas 3a, 3b, 1, and 2) – the early stage of the cortical somatosensory system. Here we inferred that, in a cross-modal visuo-haptic short-term memory task, somatosensory cells would react to visual stimuli associated with tactile features. Single-unit discharge was recorded from the anterior parietal cortex – including areas of hand representation – of monkeys performing a visuo-haptic delayed matching-to-sample task. Units changed firing frequency during the presentation of a visual cue that the animal had to remember for making a correct tactile choice between two objects at the end of a delay (retention period). Some units showed sustained activation during the delay. In some of them that activation differed depending on the cue. These findings suggest that units in somatosensory cortex react to visual stimuli behaviorally associated with tactile information. Further, the results suggest that some of these neurons are involved in short-term active memory and may, therefore, be part of cross-modal memory networks. Received: 24 March 1997 / Accepted: 8 May 1997     

Analysis of Evoked EEG Synchronization and Desynchronization During Perception of Emotiogenic Stimuli: Association with Autonomic Activation Processes

The cortical apparatus involved in performing autonomic responses in conditions of emotional activation has received little study. The aim of the present work was to assess the dynamics of evoked EEG synchronization and desynchronization at different frequency ranges during the perception of emotiogenic visual stimuli depending on the extent of accompanying autonomic activation as measured by skin galvanic responses. Studies were performed on 33 students (all right-handed) aged 18–28 years. Difference between subjects with weak (SGR^-) and strong (SGR⁺) skin galvanic responses were seen only in the θ1 range (4–6 Hz). At the stage at which emotiogenic information was perceived (the first second after the start of stimulus presentation), both groups showed similar dynamics of increases in evoked synchronization in the parietal-temporal-occipital areas of the cortex, with greater involvement of the right hemisphere. From the second second to the end of presentation (2–6 sec), emotiogenic signals gave significantly greater levels of evoked synchronization in these cortical areas as compared with neutral stimuli, and only in the SGR⁺ group. These data provide evidence for the involvement of the posterior areas of the cortex of the right hemisphere in the mechanisms of motivational attention and sympathetic activation. __________ Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 90, No. 11, pp. 1314–1323, November, 2004.     

TMS disruption of V5/MT+ indicates a role for the dorsal stream in word recognition

Although word recognition is a skill commonly expected to rely more on ventral rather than dorsal stream processing, there is some evidence for a magnocellular/dorsal impairment in dyslexia. The early rapid feedforward/feedback loop through the dorsal stream seen in primate has been suggested to allow an initial global analysis, and in human early activation of parietal attention mechanisms for detecting salient stimuli, facilitating more local level detailed ventral stream processing. To test this model in humans, transcranial magnetic stimulation (TMS) was used to probe the role of early visual cortex (V1/V2) and V5/MT+ in single word identification. TMS over V1/V2 between word onset and 36 ms post word onset disrupted accurate word discrimination, with disruption also evident at approximately 99 ms. TMS over V5/MT+ also disrupted accuracy following stimulation at approximately the same time as word onset and again at 130 ms post word onset. Thus, a role for V5/MT+ in accurate single word identification is apparent suggesting rapid triggering of attention to salient exogenous stimuli may be required prior to processing in primary and temporal cortical regions.     

Asymmetry of the internal connections of the striate cortex of the cat in the projection zone of the center of the field of vision

Studies were carried out on the organization of the internal connections of the striate cortex in cats in the projection zone of the center (0–5°) of the field of vision by microintophoretic application of horseradish peroxidase to electrophysiologically identified orientational columns. The area containing neurons showing retrograde labeling in most cases extended in the mediolateral direction. Labeled cells were located in the upper (II, III) and lower (V, VI) layers of the cortex, and the shapes and orientations of the areas containing labeled neurons in these layers coincided. Spatial asymmetry was detected in the distribution of labeled neurons relative to the orientational column studied. Labeled cells were located predominantly medial to the columns, regardless of the distance from the projection of the area centralis. Considering the visuotopical map of field 17, the asymmetry detected here provides evidence that neurons in orientational columns have more extensive connections with neurons of the peripheral part of the cortex. An asymmetrical distribution of “silent” zones around the receptive fields of neurons in orientational columns is suggested, and that these appear to receive influences from the periphery of the visual field. Laboratory of Visual Physiology and Laboratory of Central Nervous System Morphology, I. P. Pavlov Institute of Physiology, Russian Academy of Sciences, 6 Makarov Bank, 199034 St. Petersburg, Russia. Translated from Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 82, No. 12, pp. 23–29, December, 1996.     

Short and long duration transcranial direct current stimulation (tDCS) over the human hand motor area

The aim of the present paper is to study effects of short and long duration transcranial direct current stimulation (tDCS) on the human motor cortex. In eight normal volunteers, motor evoked potentials (MEPs) induced by transcranial magnetic stimulation (TMS) were recorded from the right first dorsal interosseous muscle, and tDCS was given with electrodes over the left primary motor cortex (M1) and the contralateral orbit. We performed two experiments: one for short duration tDCS (100 ms, 1, 3 or 5 mA) and the other for long duration tDCS (10 min, 1 mA). The stimulus onset asynchrony (SOA) between the onset of tDCS and TMS were 1–7 and 10–120 ms for the former experiment. In the latter experiment, TMS was given 0–20 min after the end of 10 min tDCS. We evaluated the effect of tDCS on the motor cortex by comparing MEPs conditioned by tDCS with control MEPs. Cathodal short duration tDCS significantly reduced the size of responses to motor cortical stimulation at SOAs of 1–7 ms when the intensity was equal to or greater than 3 mA. Anodal short duration tDCS significantly increased MEPs when the intensity was 3 mA, but the enhancement did not occur when using 5 mA conditioning stimulus. Moreover, both anodal and cathodal short duration tDCS decreased responses to TMS significantly at SOAs of 20–50 ms and enhanced them at an SOA of 90 ms. Long duration cathodal tDCS decreased MEPs at 0 and 5 min after the offset of tDCS and anodal long duration tDCS increased them at 1 and 15 min. We conclude that the effect at SOAs less than 10 ms is mainly caused by acute changes in resting membrane potential induced by tDCS. The effect at SOAs of 20–100 ms is considered to be a nonspecific effect of a startle-like response produced by activation of skin sensation at the scalp. The effect provoked by long duration tDCS may be short-term potentiation or depression like effects.     

Automatic and imperative motor activations in stimulus–response compatibility: magnetoencephalographic analysis of upper and lower limbs

The stimulus–response (S–R) compatibility effect refers to the difference in performance due to the spatial S–R relationship in choice reaction time. We investigated the mechanism of neural activities in S–R compatibility at the level of the primary motor cortices for upper and lower limbs responses using magnetoencephalography (MEG). In the S–R compatible task, subjects were required to respond on the same side of the stimulus light using either an upper or lower limb. In the incompatible task, subjects were required to respond in the reverse manner. Premotor times of upper and lower limbs were faster for the compatible response than for the incompatible response. The neuromagnetic brain activities related to response execution were estimated using a multi-dipole model. Stimulus-locked MEG indicated that the current moments of motor dipoles for both effectors occurred bilaterally and reached the first peak at a constant delay irrespective of whether the task was compatible or incompatible. This indicates that the neural activation of the primary motor cortex is automatically synchronized with the stimulus onset. Response-locked MEG showed that the peak current moment of the motor dipole contralateral to the response was stronger for the compatible task than for the incompatible one regardless of whether the responses were made using the upper or lower limbs. The MEG results suggest that automatic motor activation facilitates imperative motor activation for a compatible response, whereas it is not sufficient to prime imperative motor activation for an incompatible response.     

Foveal attention modulates responses to peripheral stimuli

When attending to a visual object, peripheral stimuli must be monitored for appropriate redirection of attention and gaze. Earlier work has revealed precentral and posterior parietal activation when attention has been directed to peripheral vision. We wanted to find out whether similar cortical areas are active when stimuli are presented in nonattended regions of the visual field. The timing and distribution of neuromagnetic responses to a peripheral luminance stimulus were studied in human subjects with and without attention to fixation. Cortical current distribution was analyzed with a minimum L1-norm estimate. Attention enhanced responses 100-160 ms after the stimulus onset in the right precentral cortex, close to the known location of the right frontal eye field. In subjects whose right precentral region was not distinctly active before 160 ms, focused attention commonly enhanced right inferior parietal responses between 180 and 240 ms, whereas in the subjects with clear earlier precentral response no parietal enhancement was detected. In control studies both attended and nonattended stimuli in the peripheral visual field evoked the right precentral response, whereas during auditory attention the visual stimuli failed to evoke such response. These results show that during focused visual attention the right precentral cortex is sensitive to stimuli in all parts of the visual field. A rapid response suggests bypassing of elaborate analysis of stimulus features, possibly to encode target location for a saccade or redirection of attention. In addition, load for frontal and parietal nodi of the attentional network seem to vary between individuals.