Feedback control of human alpha rhythm from the central area.

Twenty subjects, aged 17 to 25, were given from 5 to 10 sessions of training in controlling alpha. They were divided into three groups, respectively reinforced for increasing alpha from the central area, reinforced for decreasing alpha from the central area, and given noncontingent reinforcement. Compared with the initial baseline, the alpha of the noncontingent subjects did not change, while those reinforced for increases were reliably higher and those reinforced for decreases reliably lower than the noncontingent group. A slight trend toward improvement during successive sessions was not reliable. Since the experiment was conducted in the Soviet Union, the subjects had no expectations of an "alpha experience." Although tests showed a slight elevation in mood at the end of the sessions, there were no differences among the groups. There was an increase in reports of fatigue after the training sessions. There were no reports of using visual or somatomotor maneuvers as a means of controlling alpha. Furthermore, alpha rhythm control was not found to be consistently correlated with changes in heart rate, respiration, or mood, as determined by cross-correlation analysis.     

Is the alpha rhythm a control parameter for brain responses?

 The main goal of the present study is to develop a conceptual analysis of alpha response in the brain based on single sweep evaluation. A new method was employed to estimate a set of single-sweep parameters and quantify the oscillatory behaviour of single, electroencephalograph (EEG) sweeps. It was aimed to demonstrate that brain alpha responses are governed by spontaneous alpha activity and to validate the principle of brain response excitability. Because the spontaneous alpha activity depends on both the topology of recording and the subject’s age, topology and age models were used. Spontaneous and evoked alpha activity were recorded at frontal and occipital sites in three groups of subjects: 3-year-old children, young adults and middle-aged subjects. Amplitude, enhancement and phase-locking of single alpha responses to visual stimuli were analysed. Major results showed that: (1) visual alpha responses could be recorded only if the alpha rhythm was developed in the spontaneous EEG independent of electrode location; (2) middle-aged adults showed more expressed frontal spontaneous alpha activity in comparison with young adults; (3) accordingly, alpha responses with higher amplitude and stronger phase-locking were produced over the frontal brain area in middleaged than young adults. These results validate the principle of brain response excitability and demonstrate that a shift towards frontal brain areas for both the spontaneous and evoked alpha activity occurs with increasing age in adults. The results are discussed in the context of the diffuse and distributed alpha system of the brain. Age-dependent changes in frontal alpha activity are suggested to be related to frontal brain functioning during aging. Received: 6 November 1995 / Accepted in revised form: 13 March 1997     

Trajectories of alpha rhythm dipoles shifting over the human brain cortex

Dynamic study of 3D localization of the equivalent current dipoles (ECD)--sources of the EEG alpha rhythm in the human brain was performed in seven subjects with closed eyes using a one-dipole model. An exact localization of ECDs was obtained by combination of EEG and MRI mapping that allowed tracing of ECD shifts over the cortex with 4 ms step. Our data confirmed localization of these ECDs mainly in the occipital cortex and revealed their successive shift over this area during generation of each alpha-wave. Typical trajectories of these shifts were revealed and quantitatively compared by the hierarchical cluster analysis. The data obtained directly proved periodical rhythmic alpha-wave spreading process in the human visual cortex and an external control of this process. The data are discussed in terms of the "scanning hypothesis" (Pitts W., McCulloch W.H. Bull. Math. Biophys. 1947. V. 9. P. 127) which predicted a certain functional meaning of the alpha activity for cortical processing of sensory information in the human brain.     

The lability of the human background alpha rhythm under functional loads

Some characteristics of electroencephalographic reaction (alpha frequency, whole alpha range power spectrum and alpha index) have been studied. In the first series the indifferent single, rhythmic and complex stimuli were presented to the test subjects, in another series a conditioned motor reflex was elaborated, in the third series the test subjects rhythmically reproduced fixed temporal interval. It has been shown that the functional loads lead to higher alpha frequency. The alpha rhythm frequency was higher in subjects verbally underestimating and reproducing intervals with some delay than in ones overestimating intervals and reproducing them in advance. It has been suggested that alpha rhythm instability is due to reorganization processes in neuronal complexes of different lability.     

Feedback equilibrium control during human standing

Equilibrium maintenance during standing in humans was investigated with a 3-joint (ankle, knee and hip) sagittal model of body movement. The experimental paradigm consisted of sudden perturbations of humans in quiet stance by backward displacements of the support platform. Data analysis was performed using eigenvectors of motion equation. The results supported three conclusions. First, independent feedback control of movements along eigenvectors (eigenmovements) can adequately describe human postural responses to stance perturbations. This conclusion is consistent with previous observations (Alexandrov et al., 2001b) that these same eigenmovements are also independently controlled in a feed-forward manner during voluntary upper-trunk bending. Second, independent feedback control of each eigenmovement is sufficient to provide its stability. Third, the feedback loop in each eigenmovement can be modeled as a linear visco-elastic spring with delay. Visco-elastic parameters and time-delay values result from the combined contribution of passive visco-elastic mechanisms and sensory systems of different modalities     

Feedback control and quantification of the response of EEG alpha to visual stimulation

The response of the posterior EEG alpha rhythms to visual stimulation is so variable that it is difficult to obtain reliable on-line measurement of it. Feedback between the EEG alpha and the visual stimulus (1) reduces random variation in the response and (2) facilitates on-line quantification. With feedback EEG, the response to visual stimulation is measured as a series of time durations of alpha and of no-alpha intervals in the EEG. This time series occurs in two stages: an initial disturbance followed by a recovery. The quantification of the series of time durations is achieved by fitting curves to the series of alpha time intervals and of no-alpha time intervals. These functions, computed in each trial of 30 stimulations, are an objective, quantitative definition of EEG response. The utility of the method was demonstrated by testing it with reference to well-known effects. Habituation to a repeated stimulus, dishabituation, habituation to a class of stimuli, dishabituation by changing the class of stimuli, and differences among brain-lesioned, psychiatric patients and normals were shown with a detailed quantification. It was concluded that biofeedback is the method of choice for quantitative research on the EEG component of the human orienting response.     

Feedback regulation of the alpha electroencephalogram activity through control of the internal and external parameters

Summary Experiments with feedback stimulation triggered from the subject's electroencephalogram result in changing the sequential time series of intervals of occipital alpha and intervals of little or no alpha EEG activity. The rate of recurrence of alpha and no-alpha EEG can be changed by regulating the external feedback stimuli or by asking the subject to change his internal state. Four different paradigms were investigated and the results interpreted in terms of the hypothesis that oculomotor functions regulate the occurrence and nonoccurrence of alpha.This work was supported by NIGMS Grants 5 PO1 GM 14940-04 and GM 15006-03.Most of this research was conducted at the Perception Laboratory, Veterans' Administration Hospital, Bedford, Massachusetts, with the cooperation of Dr. Thomas B. Mulholland.     

The effect of mobile phone electromagnetic fields on the alpha rhythm of human electroencephalogram

Mobile phones (MP) emit low-level electromagnetic fields that have been reported to affect neural function in humans; however, demonstrations of such effects have not been conclusive. The purpose of the present study was to test one of the strongest findings in the literature; that of increased "alpha" power in response to MP-type radiation. Healthy participants (N = 120) were tested using a double-blind counterbalanced crossover design, with each receiving a 30-min Active and a 30-min Sham Exposure 1 week apart, while electroencephalogram (EEG) data were recorded. Resting alpha power (8-12 Hz) was then derived as a function of time, for periods both during and following exposure. Non-parametric analyses were employed as data could not be normalized. Previous reports of an overall alpha power enhancement during the MP exposure were confirmed (relative to Sham), with this effect larger at ipsilateral than contralateral sites over posterior regions. No overall change to alpha power was observed following exposure cessation; however, there was less alpha power contralateral to the exposure source during this period (relative to ipsilateral). Employing a strong methodology, the current findings support previous research that has reported an effect of MP exposure on EEG alpha power.     

The spatial and temporal variability of the magnetoencephalogram alpha rhythm in the normal human subject

Using one-channel SQUID biomagnetometer alpha rhythm was recorded in five healthy men and eight healthy women. Maps of distribution of maximum values of mean MEG amplitudes were analyzed. Most frequently MEG maxima predominated in the parietal parts of the head. Less frequently they were found in the occipital parts. In single instances they could be found in the posterocentral sections and on the midline. In most cases an amplitude interhemispheric asymmetry of extrema was revealed. In men the mean amplitude of the alpha-rhythm predominated more frequently on the left side and in women--on the right one. The map of distribution of mean amplitudes was stable at least during two weeks. Narrow-band filtration allowed to reveal up to three discrete amplitude modes. At definite frequencies the MEG amplitude was continuously stable during several seconds.     

Circadian rhythm of proboscis extension responsiveness in the blowfly: central control of threshold changes

ABSTRACT. The proboscis extension response of the blowfly, Protophormia terraenovae (R.-D.), elicited by stimulation of the tarsi with sucrose, was studied behaviourally under LD 12:12. A diel rhythm in responsiveness was found that occurred at successively higher daily mean levels as the test flies became increasingly starved. This rhythm persisted for at least one cycle in constant light, demonstrating it to be truly endogenous and circadian. The tarsal chemo-receptors concerned in initiating the response were examined electrophysiologically in whole fly preparations. No daily change occurred in the response of these receptors to sucrose. Hence central control of the behavioural rhythm of response to sucrose is inferred, and a model is proposed which incorporates a circadian oscillator into the currently accepted model for the control of proboscis extension thresholds.     

Feedback control of the protein kinase TAK1 by SAPK2a/p38alpha

TAB1, a subunit of the kinase TAK1, was phosphorylated by SAPK2a/p38alpha at Ser423, Thr431 and Ser438 in vitro. TAB1 became phosphorylated at all three sites when cells were exposed to cellular stresses, or stimulated with tumour necrosis factor-alpha (TNF-alpha), interleukin-1 (IL-1) or lipopolysaccharide (LPS). The phosphorylation of Ser423 and Thr431 was prevented if cells were pre-incubated with SB 203580, while the phosphorylation of Ser438 was partially inhibited by PD 184352. Ser423 is the first residue phosphorylated by SAPK2a/p38alpha that is not followed by proline. The activation of TAK1 was enhanced by SB 203580 in LPS-stimulated macrophages, and by proinflammatory cytokines or osmotic shock in epithelial KB cells or embryonic fibroblasts. The activation of TAK1 by TNF-alpha, IL-1 or osmotic shock was also enhanced in embryonic fibroblasts from SAPK2a/p38alpha-deficient mice, while incubation of these cells with SB 203580 had no effect. Our results suggest that TAB1 participates in a SAPK2a/p38alpha-mediated feedback control of TAK1, which not only limits the activation of SAPK2a/p38alpha but synchronizes its activity with other signalling pathways that lie downstream of TAK1 (JNK and IKK).     

A model of the spatial-temporal characteristics of the alpha rhythm

A linear spatially distributed model of a chain of neurons and interneurons was investigated in relation to the generation of propagated alpha rhythmic activity. It was assumed that the elements of the chain were interconnected by means of recurrent collaterals and inhibitory fibres in such a way that the connectivity functions were assumed to be homogeneous and their strength was an exponentially decreasing function of distance. It was found that such a neuronal chain shows propagation properties for frequencies in the alpha band. The results obtained with the model are in agreement with the phase velocities encountered experimentally. In this way, it was possible to estimate the length of the neural fibres responsible for the phenomenon of propagated activity. The estimates obtained are in good agreement with recent quantitative neuroanatomical data on the circuitry of the neocortex.     

The spatial structure of the alpha rhythm in the healthy human being studied by EEG mapping]

EEG study was conducted in the state of relative rest in 15 healthy subjects with predominance of the alpha rhythm. Data processing was performed on neuromapper ("Neuroscience", Great Britain). Principal attention was paid to the dynamics of spatial-temporal relations of the alpha rhythm. Cyclic changes were shown of the alpha-rhythm amplitude, proceeding with change of high and low amplitudes during seconds, varying in time in various individuals. Three types of spatial distribution of the alpha rhythm over the cortex at the relative rest were obtained: 1) generalized distribution over the cortex with the wavy shift of the frequency fields; 2) formation of delimited local zones of the alpha rhythm, differing by the frequency from the rhythm in other cortical areas; 3) local zone of the low-frequency alpha rhythm in the visual projection zone (18 and 19 fields). Change of the distribution types of the alpha rhythm in the state of rest takes place in the seconds of time intervals, corresponding, according to the literature data, to the proceeding of elementary mental processes.     

Feedback control of human metabolic work rate during robotics-assisted treadmill exercise

A novel approach is presented which suggests the use of human metabolic work rate to define and regulate exercise intensity during robotics-assisted treadmill training. The work describes the design and technical validation of the new method.A feedback structure is proposed which provides automatic regulation of metabolic work rate, in conjunction with an embedded feedback loop for volitional control of mechanical work rate. Human metabolic work rate was derived in real time from breath-by-breath measurements of oxygen uptake and carbon dioxide output.The results show that the feedback method provides close to nominal performance for square-wave and ramp reference tracking tasks and that good disturbance rejection properties are obtained. A collateral finding of this work is an estimate of 14.5% of the metabolic efficiency of robotics-assisted treadmill exercise.The use of feedback control of human metabolic work rate provides a direct measure of exercise intensity as perceived by the exercising human as it directly reflects the energy requirements of the working muscles. This complements previous approaches to guiding robotics-assisted treadmill training based on mechanical work rate, heart rate or oxygen uptake. The new approach based on metabolic work rate may have advantages in populations with compromised and widely varying exercise responses. This provides a new approach for driving and controlling active patient participation during robotics-assisted treadmill exercise.     

Redox control of cardiac rhythm

The rhythm of cardiac beats is generated by pacemaker cells differing from other cardiomyocytes by the presence of slow diastolic depolarization. Consistently activated transmembrane ionic currents provide cyclic excitation of pacemakers, forming the original “membrane clocks”. A new concept has been forwarded in the last decade according to which periodic fluctuations in myoplasmic Ca²⁺ level (“calcium clocks”) not only influence a course of “membrane clocks”, but they also can serve as independent generators of the rhythm. Transport of Ca²⁺ in cells is under constant influence of active forms of oxygen and nitrogen. Both superoxide and NO in moderate doses facilitate Ca²⁺ output from the sarcoplasmic reticulum, accelerating the course of “calcium clocks”, but in higher doses they have opposite effect that may be neutralized mainly by reduced glutathione. The control of cardiac rhythm by active forms of oxygen and nitrogen represents a feedback mechanism by which mitochondria and NO-synthases support Ca²⁺ homeostasis in cells that can be temporarily disturbed under mechanical loads or hypoxia.     

Feedback control of the Arabidopsis hypersensitive response

The plant hypersensitive response (HR) to avirulent bacterial pathogens results from programmed cell death of plant cells in the infected region. Ion leakage and changes in signaling components associated with HR progression were measured. These studies compared Arabidopsis mutants affecting feedback loops with wild-type plants, with timepoints taken hourly. In response to Pseudomonas syringae pv. tomato DC3000 x avrB, npr1-2 mutant plants showed increased ion leakage relative to wild-type plants. Hydrogen peroxide accumulation was similar to that in wild type, but salicylic acid accumulation was reduced at some timepoints. With DC3000 x avrRpt2, similar trends were seen. In response to DC3000 x avrB, ndr1-1 mutant plants showed more ion leakage than wild-type or npr1-2 plants. Hydrogen peroxide accumulation was delayed by approximately 1 h and reached half the level seen with wild-type plants. Salicylic acid accumulation was similar to npr1-2 mutant plants. With DC3000 x avrRpt2, ndr1-1 mutant plants showed no ion leakage, no hydrogen peroxide accumulation, and minimal salicylic acid accumulation. Results with a ndr1-1 and npr1-2 double mutant were similar to ndr1-1. A model consistent with these data is presented, in which one positive and two negative regulatory circuits control HR progression. Understanding this circuitry will facilitate HR manipulation for enhanced disease resistance.