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Coherent high-frequency neuronal activity has been proposed as a physiological indicator of perceptual and higher cognitive processes. Some of these processes can only be investigated in humans and the use of non-invasive recording techniques appears to be a prerequisite for investigating their physiological substrate in the healthy human brain. After addressing methodological issues in the non-invasive recording of high-frequency responses, we summarize studies indicating co-occurrence of neuronal synchrony of single cells exhibiting rhythmic activity at high frequencies, oscillations in the local field potential and dynamics in high frequencies recorded using high-resolution electroencephalography (EEG) and magnetoencephalography (MEG). We then review EEG and MEG studies of attention, perception, and language processing in humans indicating that dynamics in the high-frequency range > 20 Hz reflect specific cognitive processes. Types of high-frequency (HF) activity can be distinguished according to their latency after stimulus onset, stimulus-locking, cortical topography and frequency. There appears to be a systematic relationship between specific cognitive processes and types of HF activity. The findings are related to recent theories about the generation of HF activity and their possible role in binding of stimulus features. Dynamics of HF cortical activity reflecting higher cognitive processes can be accounted for based on the assumption that the elements of cognitive processing, e.g. visual objects and words, are organized in the brain as distributed neuronal assemblies with defined cortical topographies generating well-timed spatio-temporal activity patterns.

The causes underlying phantom limb pain are still unknown. Recent studies on the consequences of nervous system damage in animals and humans reported substantial reorganization of primary somatosensory cortex subsequent to amputation, and one study showed that cortical reorganization is positively correlated with phantom limb pain. This paper examined the hypothesis of a functional relationship between cortical reorganization and phantom limb pain. Neuroelectric source imaging was used to determine changes in cortical reorganization in somatosensory cortex after anesthesia of an amputation stump produced by brachial plexus blockade in six phantom limb pain patients and four pain-free amputees. Three of six phantom limb subjects experienced a virtual elimination of current phantom pain attributable to anesthesia (mean change: 3.8 on an 11-point scale; Z = -1.83; p < 0.05) that was mirrored by a very rapid elimination of cortical reorganization in somatosensory cortex (change = 19.8 mm; t(2) = 5.60; p < 0.05). Cortical reorganization remained unchanged (mean change = 1.6 mm) in three phantom limb pain amputees whose pain was not reduced by brachial plexus blockade and in the phantom pain-free amputation controls. These findings suggest that cortical reorganization and phantom limb pain might have a causal relationship. Methods designed to alter cortical reorganization should be examined for their efficacy in the treatment of phantom limb pain.

We examined slow potentials, transient event-related potentials, and oscillatory-like responses in the electroencephalogram during aversive conditioning in humans, in order to determine what is happening in the neocortex when behavioral adaptations are learned. Pictures of an angry and a happy human face served as reinforced (CS+) and unreinforced (CS-) conditioned stimuli, respectively, in one group, and either the reversed condition or two discriminably different neutral faces in two other groups (total n = 48 subjects). The unconditioned stimulus (US) was intracutaneous shock delivered to the left hand 5 s after CS+ onset. The electroencephalographic (EEG) activity was recorded from Fz, Cz, Pz, C3, and C4, electromyographic (EMG) activity from bilateral forearm and corrugator muscles, and skin conductance from the right hand. During acquisition a negative slow potential developed after CS+ (not CS-), which was more pronounced when a neutral face served as CS+. Early (iCNV, initial contingent negative variation) and late (tCNV, terminal contingent negative variation) components of the slow-potential response were positively related to the magnitude of conditioned EMG responses. Differentiation of tCNV was larger when neutral faces signaled the US; iCNV persisted during extinction when a happy face served as CS+. Late-occurring event-related potentials (ERPs) elicited by the US diminished over conditioning, whereas short-latency US components and ERPs elicited by CS events did not. Fourier analysis revealed oscillatory ("gamma-band") activity between 30 and 40 Hz, which persisted up to 3 s after US delivery and diminished as conditioning progressed. Our findings indicate that learning is expressed in neocortical structures at the earliest stages of conditioning. The functional roles of the three types of EEG response in learning are discussed.

Thirteen subjects were extensively trained on nine multiplication problems varying in difficulty. Practice was associated with a reaction time speed up and an attenuation of the problem size effect. The introduction of previously unpracticed problems led to a performance rebound to pretraining levels, indicating practice specificity. The eventrelated potentials were characterized by a late positive complex, followed by a positive slow wave. Offset latency of positive slow wave and preresponse amplitude at parietal electrodes showed practice specificity effects that systematically changed with practice and problem size, indicating an association with the load imposed on working memory. The peak of the late positive complex probably reflects task learning or adaptation effects because it was attenuated by practice predominantly at frontal electrodes, showed no practice specificity, and was not affected by problem size.

This study investigated age-related changes in the human brain function using both traditional EEG analysis (power spectra) and the correlational dimension, a measure reflecting the complexity of EEG dynamics and, probably, the complexity of neurophysiological processes generating the EEG. Assuming that the accumulation of individual experience is determined by the formation of functionally related groups of neurons showing a repetitive synchronous activation (cell assemblies), an increase in the number of such independently oscillating cortical cell assemblies can be expected, despite a decline of some metabolic and memory functions with normal ageing. Thus, the "wisdom of old age' may find its neurophysiological basis in greater complexity of brain dynamics compared to young ages. The experimental hypothesis was that EEG dimension steadily increases with age. In order to test this hypothesis the resting EEGs of 5 age groups from 7 to 60 were analysed. The results confirm the hypothesis: after a jump in the brain dynamics complexity during puberty a linear increase with age is observed. During maturation (7-25 years), the maximum gain in complexity occurs over the frontal associative cortex.

Patients with idiopathic Parkinson's syndrome, most of them in early stages of the disease, and matched healthy controls participated in a continuous performance task while their EEGs were recorded from 15 electrodes. During preparation of movements, a contingent negative variation (CNV) maximal at central and posterior sites was visible. This CNV was reduced in the patient population. A large P3-like positive deflection occurred after go and no-go stimuli that called for execution (go) or suppression (no-go) of a button press. Compared to healthy controls, the positive wave in Parkinson patients was significantly reduced after go stimuli and maximally attenuated when no-go stimuli had indicated to suppress the motor response. In contrast, P3 amplitudes after irrelevant "ignore' stimuli was not significantly reduced in the patients. These results are interpreted in the framework of a model of striatal function postulating (i) that populations of cortical and striatal neurons form distributed functional units (Hebbian cell assemblies), and (ii) that mutual inhibition between such cortico-striatal cell assemblies is mediated by the neostriatum, the forebrain structure primarily affected in Parkinson's disease.

Electrocortical activity was recorded from scalps of human subjects reading nouns and verbs. Current source density analysis of EEG signals and calculation of spectral responses revealed differences between word categories in the 30 Hz range. Verbs elicited stronger 30 Hz activity at recording sites over the motor cortices, while nouns elicited stronger responses at sites over visual cortices in the occipital lobes. Behavioural testing indicated that, at the cognitive level, this double dissociation corresponds to motor and visual associations prompted by verbal stimuli. These results suggest that local high-frequency brain responses can be indicators of conscious processing of motor and visual associations of verbal material. Furthermore, the results provide additional evidence that nouns and verbs have distinct neuronal generators in the intact human brain.

This study shows that divergent thinking, considered the general process underlying creative production, can be distinguished from convergent, analytical thought based on the dimensional complexity of ongoing electroencephalographic (EEG) activity. EEG complexity over the central and posterior cortex was higher while subjects solved tasks of divergent than convergent thinking, and also higher than during mental relaxation. Over the frontal cortex, EEG complexity was comparable during divergent thinking and mental relaxation, but reduced during convergent thinking. Results indicate that the basic process underlying the generation of novel ideas expresses itself in a strong increase in the EEG's complexity, reflecting higher degrees of freedom in the competitive interactions among cortical neuron assemblies. Frontocortical EEG complexity being comparable with that during mental relaxation, speaks for a loosened attentional control during creative thinking.

Meaningful words and matched pseudowords, such as moon vs. noom, are of equal perceptual complexity, but invoke different cognitive processes. To investigate high-frequency cortical responses to these stimuli, biomagnetic signals were recorded simultaneously over both hemispheres of right-handed individuals listening to words and pseudowords. Consistent with earlier EEG studies, evoked spectral responses recorded from the left hemisphere revealed depression of spectral power in the low gamma band (around 30 Hz) after pseudowords but not after words. Similar differences between stimulus categories were present in the beta range. These results indicate that distinct patterns of high-frequency cortical responses correspond to the different cognitive processes invoked by words and pseudowords. It is hypothesized that differential high-frequency cortical responses signal the activation or activation failure of distributed Hebbian cell assemblies representing words and other elements of cognitive processing.

Recording of slow brain potentials (SPs) and transcranial magnetic stimulation (TCMS) of the human motor cortex were combined to probe the relationship between SP level and excitability of cortical neurons. In experiment 1, TCMS was applied during and shortly after the warning interval in a forewarned reaction time task. Electromyographic (EMG) responses to TCMS increased only slightly during the warning interval and were significantly elevated 150 ms after the imperative stimulus. In experiment 2, TCMS was applied during biofeedback-induced cortical positivity and negativity. In this non-motor task a dependence of TCMS response on SP amplitude was not significant. Results indicate higher local excitability of motor cortex during cortical negativity when a preparatory motor task is required. TCMS may better be suited to probe processes involved in motor tasks rather than non-motor and cognitive conditions.

It has recently been proposed that brain responses in the gamma-range (> 20 Hz) include information about specific cognitive processes in the human brain. Empirical data substantiating this assumption come from EEG and MEG recordings during visual and language processing. For example, 30 Hz activity has been found to be stronger during processing of words than during processing of meaningless but pronounceable pseudo-words. However, it has been reported that power changes in the gamma-band are sometimes artefacts of changes taking place in other frequencies, such as the alpha-band. In this study, we investigated power changes in frequency bands outside the gamma-range and found no evidence that such changes are related to the dynamics in gamma-power distinguishing between words and pseudowords. These results are consistent with the view that gamma-band responses are an indicator of lexical processing and other cognitive processes related to binding of stimulus features into a whole (Gestalt), as proposed by Singer.

Fractal dimension has been proposed as a useful measure for the characterization of electrophysiological time series. This paper investigates what the pointwise dimension of electroencephalographic (EEG) time series can reveal about underlying neuronal generators. The following theoretical assumptions concerning brain function were made (i) within the cortex, strongly coupled neural assemblies exist which oscillate at certain frequencies when they are active, (ii) several such assemblies can oscillate at a time, and (iii) activity flow between assemblies is minimal. If these assumptions are made, cortical activity can be considered as the weighted sum of a finite number of oscillations (plus noise). It is shown that the correlation dimension of finite time series generated by multiple oscillators increases monotonically with the number of oscillators. Furthermore, it is shown that a reliable estimate of the pointwise dimension of the raw EEG signal can be calculated from a time series as short as a few seconds. These results indicate that (i) The pointwise dimension of the EEG allows conclusions regarding the number of independently oscillating networks in the cortex, and (ii) a reliable estimate of the pointwise dimension of the EEG is possible on the basis of short raw signals.

Psycholinguistic theories propose that words of the 2 major vocabulary classes, content (open-class) and function (closed-class) words, are computationally distinct and have different neuronal generators. This predicts distinct EEG patterns elicited by words of these 2 classes. To test this prediction, content and function words, together with matched pseudowords, were presented in a lexical decision task (where subjects had to decide whether stimuli were meaningful words or not). Evoked potentials were recorded from 17 electrodes 12 of which were located in close vicinity of the perisylvian cortices. Already 160 msec post stimulus onset, substantial differences in activity patterns distinguish the 2 vocabulary classes. A hemisphere by word class interaction revealed interhemispheric differences for function words but not for content words. Potentials evoked by function words were more negative over the left hemisphere compared to the right. These results evidence that brain mechanisms underlying function and content word processing are different. The following explanation of the data is proposed: content words correspond to neuronal assemblies equally distributed over both hemispheres, while assemblies corresponding to function words are strongly lateralized to the left hemisphere and primarily located in the perisylvian region.

Differential hemispheric involvement in controlling simple and complex motor movements was investigated in humans using EEG spectral responses. Analysis of spectral power in the alpha band revealed the following. While during a simple motor task (tapping) signs of unilateral cortical activation were present, more complex sequential motor behaviour (Luria finger apposition task) led to symmetrical bihemispheric activation. It appears that unilateral cortical processing is present in the normal brain when very simple computations are performed. Bilateral hemispheric activity and interhemispheric interaction may be general features of more complex information processing in the cortex.

The present study was designed to examine brain activity underlying mental imagery is conceptualized as behavior guided by internal representation only, the activity of the prefrontal lobes was assumed to be a measure of differentiation of imagery from perception. Twenty-one subjects were requested to observe and imagine a swinging pendulum and to touch and imagine a coshball in separate trials. The EEG was recorded from 15 standard electrode sites and analyzed with (1) traditional alpha power and (2) an estimation of dimensional complexity (a measure derived from nonlinear dynamics). Both EEG measures revealed expected object-related differences during perception as well as during imagery. The visual pendulum showed relative to the tactile coshball increased dimensional complexity and less alpha power at parietal and frontal sites. However, only the EEG dimension supported the main hypothesis: Imagery resulted in increased prefrontal dimensional complexity in comparison to perception independent of the modality of the image. In contrast, for alpha power the difference between imagery and perception was due to stimulus modality.

Recent behavioral investigations indicate that the processes underlying mental arithmetic change systematically with practice from deliberate, conscious calculation to automatic, direct retrieval of answers from memory [Bourne, L.E.Jr. and Rickard, T.C., Mental calculation: The development of a cognitive skill, Paper presented at the Interamerican Congress of Psychology, San Jose, Costa Rica, 1991: Psychol. Rev., 95 (1988) 492-527]. Results reviewed by Moscovitch and Winocur [In: The handbook of aging and cognition, Erlbaum, Hillsdale, NJ, 1992, pp. 315-372] suggest that consciously controlled processes are more dependent on frontal lobe function than are automatic processes. It is appropriate, therefore to determine whether transitions in the locus of primary brain activity occur with practice on mental calculation. In this experiment, we examine the relationship between characteristics of event-related brain potentials (ERPs) and mental arithmetic. Single-digit mental multiplication problems varying in difficulty (problem size) were used, and subjects were trained on these problems for four sessions. Problem-size and practice effects were reliably found in behavioral measures (RT). The ERP was characterized by a pronounced late positivity after task presentation followed by a slow wave, and a negativity during response indication. These components responded differentially to the practice and problem-size manipulations. Practice mainly affected topography of the amplitude of positivity and offset latency of slow wave, and problem-size mainly offset latency of slow wave and pre-response negativity. Fronto-central positivity diminished from session to session, and the focus of positivity centered finally at centro-parietal regions.(ABSTRACT TRUNCATED AT 250 WORDS)

The negativity of slow cortical potentials (SCP) of the surface EEG is a measure of brain excitability, correlating with motor and cognitive preparation. Self-control of SCP positivity has been shown to reduce seizure activity. Following SCP biofeedback from a central EEG electrode position, subjects gained bidirectional control over their SCP. The current study used a modified feedback methodology, and found a positive relationship between negativity and magnitude of EMG startle response (a measure of cortical and subcortical arousal, particularly aversive response disposition). Greater success in SCP differentiation was associated with self-report of less relaxation during negativity training.

Cortical positivity as measured by slow event-related potentials is assumed to represent a decreased excitability of cortical networks and suppression of their behavioral-cognitive output. The blink reflex probe is a commonly used defensive electromyographic response whose amplitude was shown to be modulated by emotional and attentional orientation. It was used here as an indicator of cortico-subcortical excitation. In study 1, 33 healthy subjects took part in a continuous performance test (CPT). Event-related potentials were recorded from 15 standard scalp locations. Acoustic startling noise bursts were delivered during conditions that required either performance of prepared motor responses (Go), inhibition of prepared motor responses (NoGo), or had no motor significance (Irrelevant condition). During the NoGo condition, EEG surface potentials showed a widespread P300-like positivity with a central maximum. Startle responses were inhibited during the NoGo condition as compared to the Irrelevant condition. In study 2 (21 subjects) the same format was used, except that the startle reflex was elicited visually. Startle reflexes again showed smaller magnitude during the NoGo condition, which evoked larger positivity at central sites in comparison to the Irrelevant condition. The relationship between positivity in the EEG and inhibited startle responses is in line with the hypothesis that positive EEG shifts reflect a state of cortical disfacilitation.

Subjects viewed letters presented at 2 sec intervals and prepared a fast button press whenever an "O" appeared. If the next letter was an "X" the button press was executed (Go signal), but if the letter was a non-X character (T, H, Z) suppression of the response was required (No-Go cue). No-Go signals elicited a P300-like wave that was larger at central and frontal scalp sites contralateral to the prepared movement, compared to P300s elicited by Go cues which were symmetric about the sagittal midline and dominant at parietal sites. Subtraction of preparatory CNVs from the No-Go P300 did not remove differences in scalp topography, or reduce the amplitude of the No-Go P300 to that seen following control letters that required perceptual identification but did not call for suppression of prepared motor responses. Principal components analysis identified a middle positive wave following X-alone control stimuli whose topography resembled the No-Go P300. These findings suggest that the source of augmented No-Go P300s is a generator involved with sensorimotor inhibition. We discuss the mechanism of P300 waves and evidence linking these waves with inhibition in other task arrangements.

We explored classical conditioning in human subjects who had lesions in their cerebellar circuitry. Seven patients with damage to cerebellar structures and matched control subjects underwent simple delay tone-airpuff conditioning. Eyelid conditioned response (CR) acquisition was severely disrupted in the patient group, whereas autonomic CRs and slow cortical potentials developing between conditioned stimulus (CS) and the unconditioned stimulus (UCS) were unaffected. Results are consistent with animal studies and earlier case reports indicating that intact cerebellar structures are necessary for the acquisition of classically conditioned motor responses.

The present study aimed at investigating to what extent the regulation of excitability in cortical networks, as indicated by surface-negative slow cortical potentials (SCPs), is impaired in epileptic patients and to what extent training of SCP self-regulation by means of biofeedback and instrumental learning procedures might affect seizure frequency. Twenty-five patients suffering from drug-refractory epilepsies (complex focal, grand mal, and absence type of seizures) participated in 28 1-h sessions of feedback and instrumental conditioning of their SCPs. Subjects' EEGs were obtained from the vertex. Depending on discriminative stimuli DC shifts towards increased or suppressed negativity relative to the pre-trial baseline were demonstrated by on-line visual feedback during intervals of 8 s each; each session comprised 110 trials. While performance on the SCP self-regulation task was initially below normal (as compared to healthy subjects), significant increases in SCP control were achieved by the patients across the 28 training sessions. In 18 patients at least 1-year follow-up data are available. Changes in seizure frequency were related to transfer of SCP control with six of the patients becoming seizure-free. Age affected the ability to acquire SCP control and its impact on seizure frequency.

Slow cortical potentials (SCPs) are considered to reflect the regulation of attention resources and cortical excitability in cortical neuronal networks. Impaired attentional functioning, as found in patients with schizophrenic disorders, may covary with impaired SCP regulation. This hypothesis was tested using a self-regulation paradigm. Twelve medicated male schizophrenic inpatients and 12 healthy male controls received continuous feedback of their SCPs, during intervals of 8 s each, by means of a visual stimulus (a stylized rocket) moving horizontally across a TV screen. The position of the feedback stimulus was a linear function of the integrated SCP at each point in time during the feedback interval. Subjects were required to increase or reduce negative SCPs (referred to pretrial baseline) depending on the presentation of a discriminative stimulus. The correct response was indicated by the amount of forward movement of the feedback stimulus and by monetary rewards. Schizophrenics participated in 20 sessions (each comprising 110 trials), while controls participated in 5 sessions. Compared with the healthy controls, schizophrenics showed no significant differentiation between negativity increase and negativity suppression during the first sessions. However, in the last 3 sessions, patients achieved differentiation similar to controls, demonstrating the acquisition of SCP control after extensive training.

Findings on depressive patients indicate that depressives have electrophysiological characteristics similar to those of schizophrenics, in that they exhibit reduced Contingent Negative Variation (CNV) amplitudes and more distinct Postimperative Negative Variations (PINVs) than normal controls. In a biofeedback experiment, 8 medicated male inpatients with the DSM III-R diagnosis of "Bipolar Disorder, Depressive," and "Major Depression" demonstrated no impairment in the self-regulation of Slow Cortical Potentials (SCP) in comparison to schizophrenics in terms of increasing and suppressing negativity. Continuous visual SCP feedback is presented to the patient as a horizontally moving rocket in a video game format. The direction changes of the rocket represented SCP changes at each point in time, recorded by the central EEG (based on the pretrial baseline). Depressives demonstrated SCP self-regulation across 20 sessions, although with many between-and-within variations. The 8 male controls were unable to regulate their SCPs across 5 sessions. This result contradicts other findings of our laboratory on normal controls. Motivational factors and insufficient operant reinforcement (financial reward) may have facilitated this effect.

The purpose of this study was the determination of the relationship between the dimensional complexity of the electroencephalogam (EEG) and the level of intelligence in humans. In two experiments 34 male subjects were divided into two groups, with high and low levels of intelligence (as measured by the intelligence quotient (IQ)). During a resting phase and various mental imagery conditions the EEG was recorded from several scalp sites. Nonlinear analysis, based on the theory of deterministic chaos, revealed that subjects with high IQs demonstrate higher dimensional complexity of the EEG attractors than subjects with low IQs only during resting conditions. During performance of the imagery tasks the less intelligent subjects increase the complexity of electrical brain dynamics such that IQ-dependency vanishes. The gross (mass) neuronal manifestation of general intelligence seems to depend on task conditions and may be related to the individual brain dynamics only when no specific task is present.

Exteroceptive feedback was given for negative and positive shifts in slow potentials (SPs) recorded from Fz, Cz, or Pz (between groups design). Slow potentials at the feedback site were referred to adjacent scalp and non-cephalic electrodes, so as to confine SP shifts to the feedback location. Area-specific regulation of SPs was obtained at each midsagittal site after 3 days of feedback training. Subjects reported sensorimotor and emotional arousal when negative SP shifts were trained frontally, but not when negative shifts were trained parietally (cognitive/attentional strategies reported after parietal feedback). Area-specific regulation of SPs was subsequently abolished when behavioral tasks were added to further probe frontal/parietal differences (dual-task procedure). These findings indicate that area-specific self-regulation of SPs is possible on the sagittal midline, and that self-regulated parietal SPs (in contrast to frontal ones) arise from non-motoric generators. The source of SP self-regulation was more readily probed by verbal reports of feedback strategy than by study of dual-task relations, because feedback control was disrupted by the dual-task requirement.

Symptom-specific psychophysiological responding was assessed in 20 chronic back pain patients, 20 patients suffering from temporomandibular pain and dysfunction, and 20 matched healthy controls. Surface EMG from the lower and upper back, the masseter, and the biceps muscles, and heart rate and skin conductance level were continuously recorded during adaptation, resting baseline, and stressful and neutral imagery phases. Univariate and multivariate analyses of variance were performed on raw data as well as data corrected for autocorrelation. The results showed significantly higher EMG reactivity which was lateralized to the left side at the patients' site of pain but not distal sites. This hyperreactivity was observed only during stressful imagery. The healthy controls displayed a significantly higher response in heart rate, but skin conductance level was not significantly different. The results are interpreted as indicative of idiosyncratic muscular response patterns to personally relevant situations at the site of pain in patients suffering from chronic muscular pain.

The present study investigated the effects of biofeedback of arterial blood pressure on cortical, peripheral, and psychological measures and the dependence of these effects on nicotine. Four groups of subjects, nonsmokers, and habitual smokers who smoked cigarettes during the experimental sessions containing 0.3, 0.8, or 1.5 mg nicotine, respectively, participated in a feedback paradigm in which continuous feedback of mean blood pressure was provided for intervals of 8 s each. While tonic blood pressure did not differ between the groups, the ability to modulate blood pressure (under feedback conditions) was restricted in smokers as compared to nonsmoking subjects; increasing nicotine dosage was accompanied by poorer performance. Independently of habitual smoking and nicotine doses, heart rate increased during feedback and under conditions of blood pressure increase. In smokers, activity in the alpha band was reduced in a dose-dependent manner. Slow cortical potentials (SCPs) during the feedback interval varied with self-induced blood pressure changes in nonsmokers (blood pressure increase was accompanied by reduced surface-negative potential shifts and vice versa), while SCP variations during feedback conditions were small in smokers, more so under the influence of 0.3 and 0.8-mg nicotine, less so under 1.5 mg. Verbal reports suggest that awareness of performance strategies may not be a necessary variable for performance on the blood pressure regulation task.

To study physiological and psychological effects of baroreceptor activity, the cervical neck cuff technique has been frequently used to stimulate the carotid baroreceptors mechanically. Using this technique, no satisfying control conditions to date have been available. Because the carotid stretch receptors are sensitive not only to the pressure level, but also to the rate of change, it is possible to manipulate the receptor firing through changes in carotid pulse amplitude. The device described here relies on the application of short changes in cuff pressure tied to different phases within the cardiac cycle (phase related external suction (PRES)). A brief external suction during systole has potent stimulatory effects on baroreceptors whereas the application of the very same pressure pulse during diastole inhibits the firing burst associated with the pulse wave. To allow an ongoing period of stimulation, a sequence of alternating negative/positive pressure pulses is applied. In the stimulation condition, the R-wave of the electrocardiogram triggers a negative pulse which is followed by a positive one during diastole. In the control condition this relationship is reversed. Two experiments are reported confirming different baroreceptor effects of the two conditions. PRES allows for blind or double-blind experiments to investigate effects of baroreceptor activity on physiology and behavior.

We assessed whether instrumentally-learned pressor responses inhibit electrocortical activity, as predicted by learning theories of idiopathic hypertension. Subjects received beat-by-beat feedback for increases and decreases in mean arterial pressure measured from the finger (Peñáz method). Slow potentials were recorded from the midsagittal line during the final training session. Also recorded at this time were heart rate, eye movements, respiration, and post-session verbal reports of the subject's control strategies. Thirteen of 14 subjects differentiated blood pressure increases and decreases at p less than .05 or better during the final session (within-subject discriminative operant procedure). Slow potentials were less negative on blood pressure increase compared to decrease trials at all midsagittal sites (p less than .02), indicating relative cortical inhibition by pressor responses. This effect occurred even though subjects reported tensing of muscles on increase trials (p less than .01), a behavioral activity previously associated with augmented rather than diminished cortical negativity. On increase trials slow potentials shifted toward positivity just prior to heart rate deceleration (the latter effect confirming activation of the baroreceptors).

The present study employed event-related potentials (ERPs) of the brain to improve the understanding of temporal processing. A reproduction paradigm was realized by presenting a visual stimulus (illuminated screen) for intervals of varying length. A few seconds after presentation of such standard intervals the visual stimulus was switched on again and subjects were asked to reproduce the duration of the standard interval by turning off the illumination after a corresponding interval had elapsed. The length of standard intervals varied randomly with each of the following lengths being presented 20 times: 1, 2, 3, 4, 6, and 8 s. Reproduction was accurate for standard intervals up to 3 s but deteriorated with increasing interval length. Brain potentials during reproduction intervals of 1-3 s differed from those recorded during the longer intervals. A CNV-like slow negative shift developed during the shorter reproduction intervals. Negatively was reduced or even absent, when subjects had to reproduce standard intervals of 4 s or longer. The ERP results suggest that intervals shorter than 3-4 s may evoke a processing mode that is qualitatively different from the one dominating when periods in the range of several seconds have to be processed.

The effects of the benzodiazepine clonazepam (a drug used as anticonvulsant) on event-related brain potentials were investigated in healthy human subjects. Thirty-six male student volunteers (mean age 30 years) received clonazepam or a placebo in a double-blind setting. VEPs (visual evoked potentials) were obtained from the standard checkerboard reversal procedure; AEPs (auditory evoked potentials) and slow cortical potentials (CNV) were measured during a 2-stimulus reaction time paradigm, in which the quality of the acoustic S1 signalled whether the acoustic S2 would follow after 2 sec or after 6 sec. Each S2 requested a speeded button press. Compared to placebo, clonazepam significantly reduced P100 amplitude of the VEP and the amplitudes of the AEP components N1 and P3. On the other hand, clonazepam boosted the development of a distinct N2 which was not apparent in placebo subjects. The CNV was significantly reduced and reaction time increased under clonazepam compared to placebo. Specific versus non-specific damping effects of the benzodiazepine are discussed, comparing the present result with the pattern of ERP effects of the anticonvulsant carbamazepine that had been obtained using the same experimental paradigms.

Two studies served to examine behavioural effects of slow cortical potentials (SPs). SPs were manipulated by means of a biofeedback procedure. The ability of human subjects to alter SPs differentially between the two hemispheres--specifically over the lateral aspects of the central sulcus--was tested by providing feedback of the SP difference between C3 and C4. In Expt. I, 21 of the 45 subjects produced hemispheric asymmetries of more than 2 microV between C3 and C4 on an average after 80 trials of analogue, continuous and immediate feedback. In Expt. II, SP changes were fed back digitally at the end of each trial. Within 120 trials, 20 of the 48 subjects reached the criterion of a minimum 2-microV difference in SPs between C3 and C4 on the average. Average differentiation remained significantly below the SP differentiations achieved for continuous feedback. Trials with feedback were followed by 'task' trials without feedback, during which subjects were still requested to produce SP changes but also had to complete a task: Either sensorimotor tasks (Expt. I) or forced choice handedness tasks (Expt. II) were presented to evaluate behavioural consequences of hemispheric SP differences. In subjects achieving the required SP differentiation it affected the behavioural output in agreement with the known functions of the respective cortical area.

Twenty subjects learned to control slow potential (SP) shifts of the brain by means of a biofeedback procedure. Depending upon the pitch of a signal tone, negative SP shifts had to be increased or reduced during intervals of 6 sec each. Visual feedback of the actual SP shift was given. Blocks of training trials alternated with blocks of test trials without any feedback of the SPs. At the end of every test trial a simple arithmetic problem had to be solved by the subjects. Subjects performed the computation in a shorter time interval if an increased negativity preceded task onset as compared to slower response times during suppression of negativity. Results suggest that cortical negativity reflects unspecific preparation for cerebral performance.

Following one base-line session, 20 normal subjects received four half hour sessions consisting of simultaneous feedback of heart rate and frontalis muscle (pretraining). Ten subjects received contingent (CF), the other ten noncontingent feedback (NCF). Subjects were asked to lower heart rate and frontal muscle tension (EMG). Heart rate within sessions decreased up to 19 bpm, with a mean of 4 bpm for the CF group. There was only a week decrease over sessions, however, because of the strong habituation effect. The following events accompanied the heart rate decrease: (1) an increase of the variability of the heart rate, (2) a decrease of the variance of the EMG, (3) an increased correlation between heart rate slowing and EMG decrease, and (4) an increasing subjective experience of control of heart rate and EMG. After pretraining, subjects received eight sessions of auditory feedback of their frontal EEG theta activity (four sessions with CF and four sessions with NCF in balanced order). There was a weak increase of theta for the CF condition over sessions, but a decrease within the sessions. Pretraining on heart rate and frontal EMG control had no influence on the performance during theta training. It was hypothesized that control of heart rate slowing and theta control involve different mechanisms.

Low-level direct currents (DC) are thought to polarize brain tissue and to affect brain processes. The present experiments investigated the effects of low-level transcortical DC-currents on response speed and physiological variables in humans. The DC-currents were applied during the warning interval within a constant foreperiod reaction time paradigm. Currents of less than 0.3 mA were applied between a vertex electrode and a noncephalic reference. The polarity of the electrical source varied randomly across trials within subjects. The first two studies showed that subjects respond fastest when the positive pole is applied to the vertex. Under the same conditions higher skin conductance responses were observed as compared to vertex negative conditions. The dynamics of the observed responses suggest that the brain "learns" to respond differentially to the different current polarities. This was confirmed by the third study, which documented different conditioned electroencephalic responses subsequent to current application.

Seventeen pairs of monozygotic twins, females and males, were tested in a dichotic listening task, containing several types of pips: standard and deviating target pips, which the subject either attended to, or not. Averaged auditory evoked potentials (AEPs) to the pips provided measures of different attentional processes. Furthermore, EEG power spectra, heart rate and blood pressure and behavioral performance were measured. Subjects received treatments (20 I.U. lysine-vasopressin vs. placebo) intranasally 48, 24, and 1 hour prior to the experimental sessions according to a co-twin control design. Whereas measures of voluntary selective attention remained unchanged by lysine-vasopressin (LVP) the peptide primarily affected an attentional mechanism responding in an automatic fashion to stimulus deviance. This effect was indicated by a substantial negative shift of the AEP amplitudes following deviating stimuli within the latency range of the N2/P2 components (about 200 msec post-stimulus). The effect seemed to be unrelated to modulations of cortical arousal after LVP.

Subjects received exteroceptive feedback for bidirectional changes in slow cortical potentials or alpha power measured from the vertex. The slow potential group succeeded in shifting slow potentials toward negativity and positivity on feedback and transfer trials requiring these changes, after two sessions of training. Differentiation of negativity and positivity was accompanied by verbal reports of somatomotor activation that occurred on trials on which negative slow potentials were required (p less than .01). Vertical and lateral eye movements, chin and frontalis electromyogram, and heart rate did not differentiate between negativity and positivity trials in the slow potential negativity during feedback. Although the alpha power group did not succeed at controlling changes in alpha, evidence of a training effect appeared in verbal reports of emotional arousal (p less than .05) and focused vision (p less than .08) on alpha suppression trials in this group. We discuss the findings from the viewpoint that biofeedback tasks involving electrocortical responses are problems in the organization of action that subjects seek to solve.

An experiment was performed to investigate the self-regulation of slow cortical potentials (SCP) found in a previous study (Elbert et al. 1979). Seventeen subjects received continuous visual feedback of their actual cortical shift perceptible as a rocket moving across a TV-screen during intervals of 6 sec; subjects had to direct the rocket into one of two goals representing more or less cortical negativity, depending on the pitch of two signal tones. Within two identical experimental sessions feedback trials alternated with test trials without feedback. Highly significant differences of SCP between the two required polarities were demonstrated. The most pronounced differences were observed during test trials without feedback of the second session in which a positive shift below baseline level occurred when positivity (or less negativity) was required.

On the basis of Lang's (1979) theory of emotional imagery three experiments were conducted to investigate the relationship between slow cortical potentials (SP) and emotional imagery. According to the assumptions of Lang's theory and our model (Rockstroh et al., 1982) of SP-function imagery ability should be related to a person's capacity to generate and suppress preparatory activity in cortical networks "on demand." In order to test this hypothesis subjects in Experiment I were trained to regulate right- versus left-hemispheric SP-differentiation within an instrumental learning paradigm. Thirty-four subjects were reinforced for achieving maximal SP-differences between electrode locations C3-C4 over a 6 s interval across 120 trials. Success in the SP-regulation task correlated significantly (r = .37) with the capacity for vivid imagery as measured with the Questionnaire for Mental Imagery (QMI). In Experiment II instructions to imagine right- versus left-hand movements were introduced successively over 5 sessions of SP-self-regulation. Imagery clearly modified right- versus left-hand EMG-differentiation but had no influence on cortical SP-differentiation. Experiment III tested the influence of already achieved SP-regulation at the vertex on the perceived vividness of emotional images introduced after the SP-biofeedback training. Again, clear effects of imagery content on autonomic variables (HR, SCR) were found. However, SP-amplitude and SP-polarity had no effect on perceived vividness, arousal or emotional content. It may be concluded from the results of Experiment II and III that SPs either are not the crucial parameter to represent the cortical efferent outflow component of imagery, or that the dual task of SP-self-regulation and imagery prevented covariations to show up. Experiment I, on the other hand, points toward a positive relation of imagery-ability as a trait-variable and brain-self-regulation abilities.

The present study investigates the hemispheric specificity of event-related (ERP) and slow potentials (SP) of the brain during preparation and performance of sensorimotor (haptic) tasks. Tasks were selected which have been assumed to be processed primarily by only one brain hemisphere. ERPs, SPs and spontaneous EEGs were obtained from both hemispheres (C3-C4, P3-P4) and from Fz and Cz in a constant foreperiod (6 s) paradigm. Warning and task stimuli comprised up to four tactile stimuli, applied to index and middle finger of each hand. To maximize lateralized processing, the right-hand task required counting, the left hand task pattern matching. Components of the ERP (N1, P3, SW) and of SPs show task-dependent lateral asymmetry with the larger amplitudes contralateral to the preparing and responding hand. Results confirm hemisphere-specific processing of the tactile tasks but quantification as indicated by ERP amplitudes show considerable activity in both hemispheres. The SP-distribution rather reflects task-specific preparation than general alertness or orienting.

The comparison of principal component analyses between seven experimental studies demonstrates a remarkable similarity of the extracted components. Slow scalp-recorded potentials of the brain (SPs) during a constant 6 sec foreperiod can be described by an early frontal, and a late preparatory component. Furthermore, an additional intermediate component may be retained by the PCA. This component seems to reduce the between-subject variance and often describes processes dependent on stimulus repetition. There is evidence favoring the varimaxed solution of the PCA for the parametrization of most of the experimental data.