Manuo-ocular coordination in target tracking. I. A model simulating human performance

During eye tracking of a self-moved target, human subjects' performance differs from eye-alone tracking of an external target. Typical latency between target and eye motion onsets is shorter, ocular smooth pursuit (SP) saturation velocity increases and the maximum target motion frequency at which the SP system functions correctly is higher. Based on a previous qualitative model, a quantitative model of the coordination control between the arm motor system and the SP system is presented and evaluated here. The model structure maintains a high level of parallelism with the physiological system. It contains three main parts: the eye motor control (containing a SP branch and a saccadic branch), the arm motor control and the coordination control. The coordination control is achieved via an exchange of information between the arm and the eye sensorimotor systems, mediated by sensory signals (vision, proprioception) and motor command copy. This cross-talk results in improved SP system performance. The model has been computer simulated and the results have been compared with human subjects' behavior observed during previous experiments. The model performance is seen to quantitatively fit data on human subjects. Received: 6 March 1997 / Accepted in revised form: 15 July 1997     

Relative phase dynamics in perturbed interlimb coordination: the effects of frequency and amplitude

Sch?ner [Sch?ner G (1995) Ecol Psychol 7: 291-314] argued that the relative phase dynamics of rhythmic interlimb coordination may be attributed to the timing level in that the stability properties of the relative phase are largely independent of dynamical principles operating at the goal level, such as those related to the maintenance of a particular amplitude or target position. Yet, according to the coupling functions in the coupled oscillator model proposed by Haken et al. [Haken H, Kelso JAS, Bunz H (1985) Biol Cybern 51: 347-356], the effect of frequency on the stability properties of relative phase is either wholly or partially mediated by frequency-induced changes in amplitude, implying that the relative phase dynamics strongly depends on spatial factors. In order to distinguish between these contrasting interpretations of the organizational principles underwriting the phase dynamics of interlimb coordination, an experiment was conducted in which the effects of frequency and amplitude on the stability of relative phase were separated. Six subjects performed both in-phase and antiphase coordination patterns at seven different frequencies and three different amplitudes. Two measures of pattern stability were used, the standard deviation of relative phase and the exponent of the relaxation process following phasic perturbations of relative phase. According to both measures, pattern stability decreased with increasing frequency, whereas the amplitude manipulation only had a significant effect on the standard deviation of relative phase. This result was interpreted to imply that the organizational principles at the (relative) timing level are affected only moderately by task constraints pertaining to the goal level, and that models of interlimb coordination in which amplitude coupling plays a partial or subordinate role should be preferred above models relying solely on amplitude coupling.     

Manuo-ocular coordination in target tracking. II. Comparing the model with human behavior

Several studies have shown that humans track a moving visual target with their eyes better if the movement of this target is directly controlled by the observer's hand. The improvement in performance has been attributed to coordination control between the arm motor system and the smooth pursuit (SP) system. In such a task, the SP system shows characteristics that differ from those observed during eye-alone tracking: latency (between the target-arm and the eye motion onsets) is shorter, maximum SP velocity is higher and the maximum target motion frequency at which the SP can function effectively is also higher. The aim of this article is to qualitatively evaluate the behavior of a dynamical model simulating the oculomotor system and the arm motor system when both are involved in tracking visual targets. The evaluation is essentially based on a comparison of the behavior of the model with the behavior of human subjects tracking visual targets under different conditions. The model has been introduced and quantitatively evaluated in a companion paper. The model is based on an exchange of internal information between the two sensorimotor systems, mediated by sensory signals (vision, arm muscle proprioception) and motor signals (arm motor command copy). The exchange is achieved by a specialized structure of the central nervous system, previously identified as a part of the cerebellum. Computer simulation of the model yielded results that fit the behavior of human subjects observed during previously reported experiments, both qualitatively and quantitatively. The parallelism between physiology and human behavior on the one hand, and structure and simulation of the model on the other hand, is discussed. Received: 6 March 1997 / Accepted in revised form: 15 July 1997     

The discontinuous nature of motor execution

Human movement control requires adequate coordination of different movements, which is particularly important when different motor tasks are simultaneously executed by the same effector(s) (e.g. a muscle or a joint). The process of movement execution involves a series of highly nonlinear elements; for instance, a motor unit of a muscle produces force only in the direction of muscle shortening, thus representing a threshold operator that transforms the bipolar (i.e. excitatory or inhibitory) information at its spinal input into a purely unipolar signal (i.e. muscle force). This tripartite research report addresses the contribution of the nonlinearity of neuromuscular elements to the coordination of different motor tasks simultaneously executed by the same limb. In this first part of the series, a new hypothesis for such a single-muscle multiple-task coordination is presented which suggests an essentially threshold-linear coordination mechanism. Control signals generated by the central nervous system for each individual movement independently and feedback information from peripheral receptors are linearly superimposed. This compound control/feedback signal is processed by a nonlinear limiter element reflecting the discontinuous properties of the muscle and its reflex circuitry. It is shown that threshold-linear interaction of descending commands and afferent feedback information can lead to complex interdependent patterns of compound motor action. This includes the possibility of gating (i.e. the ability of one movement pattern to constrain or even impede the execution of another pattern) and of delayed response initiation when simultaneously performing more than one voluntary motor task. A theoretical analysis of the threshold-linear coordination mechanism and an extensive experimental validation of the model is provided in part II and part III of the report. Received: 6 October 1998 / Accepted in revised form: 2 June 1999     

A Direct Brain-to-Brain Interface in Humans

We describe the first direct brain-to-brain interface in humans and present results from experiments involving six different subjects. Our non-invasive interface, demonstrated originally in August 2013, combines electroencephalography (EEG) for recording brain signals with transcranial magnetic stimulation (TMS) for delivering information to the brain. We illustrate our method using a visuomotor task in which two humans must cooperate through direct brain-to-brain communication to achieve a desired goal in a computer game. The brain-to-brain interface detects motor imagery in EEG signals recorded from one subject (the “sender”) and transmits this information over the internet to the motor cortex region of a second subject (the “receiver”). This allows the sender to cause a desired motor response in the receiver (a press on a touchpad) via TMS. We quantify the performance of the brain-to-brain interface in terms of the amount of information transmitted as well as the accuracies attained in (1) decoding the sender’s signals, (2) generating a motor response from the receiver upon stimulation, and (3) achieving the overall goal in the cooperative visuomotor task. Our results provide evidence for a rudimentary form of direct information transmission from one human brain to another using non-invasive means.     

Artificial feedback for invasive brain–computer interfaces

During the last two decades, considerable progress has been made in the studies of brain–computer interfaces (BCIs)—devices in which motor signals from the brain are registered by multi-electrode arrays and transformed into commands for artificial actuators such as cursors and robotic devices. This review is focused on one problem. Voluntary motor control is based on neurophysiological processes, which strongly depend on the afferent innervation of skin, muscles, and joints. Thus, invasive BCI has to be based on a bidirectional system in which motor control signals are registered by multichannel microelectrodes implanted in motor areas, whereas tactile, proprioceptive, and other useful signals are transported back to the brain through spatiotemporal patterns of intracortical microstimulation (ICMS) delivered to sensory areas. In general, the studies of invasive BCIs have advanced in several directions. The progress of BCIs with artificial sensory feedback will not only help patients, but will also expand base knowledge in the field of human cortical functions.     

Toward electrocorticographic control of a dexterous upper limb prosthesis: building brain-machine interfaces

One of the most exciting and compelling areas of research and development is building brain machine interfaces (BMIs) for controlling prosthetic limbs. Prosthetic limb technology is advancing rapidly, and the modular prosthetic limb (MPL) of the Johns Hopkins University/ Applied Physics Laboratory (JHU/APL) permits actuation with 17 degrees of freedom in 26 articulating joints. There are many signals from the brain that can be leveraged, including the spiking rates of neurons in the cortex, electrocorticographic (ECoG) signals from the surface of the cortex, and electroencephalographic (EEG) signals from the scalp. Unlike microelectrodes that record spikes, ECoG does not penetrate the cortex and has a higher spatial specificity, signal-to-noise ratio, and bandwidth than EEG signals. We have implemented an ECoG-based system for controlling the MPL in the Johns Hopkins Hospital Epilepsy Monitoring Unit, where patients are implanted with ECoG electrode grids for clinical seizure mapping and asked to perform various recorded finger or grasp movements. We have shown that low-frequency local motor potentials (LMPs) and ECoG power in the high gamma frequency (70,150 Hz) range correlate well with grasping parameters, and they stand out as good candidate features for closed-loop control of the MPL.     

Improving the separability of multiple EEG features for a BCI by neural-time-series-prediction-preprocessing

A recently proposed method for EEG preprocessing is extended and analyzed in this work via a range of different tests in combination with various other BCI components. Neural-time-series-prediction-processing (NTSPP) is a predictive approach to EEG preprocessing where prediction models (PMs) are trained to perform one-step-ahead prediction of the EEG times-series which reflect motor imagery induced alterations in neuronal activity. Due to the specialization of distinct PMs, the predicted signals (Ys) and error signals (Es) are distinctly different from the original (Os) signals. The PMs map the Os signals to a higher dimension which, in the majority of cases, produces features that are more separable than those produced by the Os signals. Four feature extraction procedures, ranging in complexity and in terms of the information which is extracted i.e., time domain, frequency domain and time–frequency (t–f) domain, are used to determine the separability enhancements which are verified by comparative statistical tests and brain–computer interface (BCI) tests on six subjects. It is shown that, in the majority of the tests, features extracted from the NTSPP signals are more separable than those extracted from the Os signals, in terms of increased Euclidean distance between class means, reduced inter-class correlations and intra-class variance, and higher classification accuracy (CA), information transfer (IT) rate and mutual information (MI).     

Effect of alpha-ketoglutarate on neurobehavioral, neurochemical and oxidative changes caused by sub-chronic cyanide poisoning in rats

Recent studies revealed that alpha-ketoglutarate (A-KG) alone or with sodium thiosulfate (STS) provide significant protection against acute and sub-acute cyanide poisoning in rodents. This study addresses the protective effect of A-KG and/or STS in sub-chronic (90 days) cyanide poisoning. Wistar rats were divided into seven groups (n = 10): Control animals, potassium cyanide (KCN) A-KG, STS, KCN + A-KG, KCN + STS and KCN + A-KG + STS. Spontaneous motor activity and motor coordination were recorded every 15th day. Lipid peroxidation (LPO), reduced glutathione (GSH), glutathione peroxidase (GPx), superoxide dismutase (SOD) and catalase (CAT) in blood, brain, liver and kidney, and glutamate, aspartate and dopamine in discrete regions of brain were measured following 90 days exposure. Cyanide significantly decreased motor coordination, accompanied by increase in LPO (blood, brain and liver) and dopamine (corpus striatum and cerebral cortex) levels, and depletion in GSH (blood, brain and liver), GPx (brain and liver), SOD (brain and liver), and CAT (blood and brain) levels. Although treatment of A-KG and STS alone significantly blunted the toxicity of KCN, concomitant use of both afforded the maximum protection. This study shows a promising role of A-KG and STS as treatment regime for long term cyanide exposure.     

fMRI signal changes in response to forced expiratory loading in congenital central hypoventilation syndrome.

Congenital central hypoventilation syndrome (CCHS) patients show impaired ventilatory responses to CO2 and hypoxia and reduced drive to breathe during sleep but retain appropriate breathing patterns in response to volition or increased exercise. Breath-by-breath influences on heart rate are also deficient. Using functional magnetic resonance imaging techniques, we examined responses over the brain to voluntary forced expiratory loading, a task that CCHS patients can perform but that results in impaired rapid heart rate variation patterns normally associated with the loading challenge. Increased signals emerged in control (n = 14) over CCHS (n = 13; ventilator dependent during sleep but not waking) subjects in the cingulate and right parietal cortex, cerebellar cortex and fastigial nucleus, and basal ganglia, whereas anterior cerebellar cortical sites and deep nuclei, dorsal midbrain, and dorsal pons showed increased signals in the patient group. The dorsal and ventral medulla showed delayed responses in CCHS patients. Primary motor and sensory areas bordering the central sulcus showed comparable responses in both groups. The delayed responses in medullary sensory and output regions and the aberrant reactions in cerebellar and pontine sensorimotor coordination areas suggest that rapid cardiorespiratory integration deficits in CCHS may stem from defects in these sites. Additional autonomic and perceptual motor deficits may derive from cingulate and parietal cortex aberrations.     

Schema-based learning of adaptable and flexible prey-catching in anurans I. The basic architecture

A motor action often involves the coordination of several motor synergies and requires flexible adjustment of the ongoing execution based on feedback signals. To elucidate the neural mechanisms underlying the construction and selection of motor synergies, we study prey-capture in anurans. Experimental data demonstrate the intricate interaction between different motor synergies, including the interplay of their afferent feedback signals (Weerasuriya 1991; Anderson and Nishikawa 1996). Such data provide insights for the general issues concerning two-way information flow between sensory centers, motor circuits and periphery in motor coordination. We show how different afferent feedback signals about the status of the different components of the motor apparatus play a critical role in motor control as well as in learning. This paper, along with its companion paper, extend the model by Liaw et al. (1994) by integrating a number of different motor pattern generators, different types of afferent feedback, as well as the corresponding control structure within an adaptive framework we call Schema-Based Learning. We develop a model of the different MPGs involved in prey-catching as a vehicle to investigate the following questions: What are the characteristic features of the activity of a single muscle? How can these features be controlled by the premotor circuit? What are the strategies employed to generate and synchronize motor synergies? What is the role of afferent feedback in shaping the activity of a MPG? How can several MPGs share the same underlying circuitry and yet give rise to different motor patterns under different input conditions? In the companion paper we also extend the model by incorporating learning components that give rise to more flexible, adaptable and robust behaviors. To show these aspects we incorporate studies on experiments on lesions and the learning processes that allow the animal to recover its proper functioning     

A review on directional information in neural signals for brain-machine interfaces

Brain-machine interfaces (BMIs) can be characterized by the technique used to measure brain activity and by the way different brain signals are translated into commands that control an effector. We give an overview of different approaches and focus on a particular BMI approach: the movement of an artificial effector (e.g. arm prosthesis to the right) by those motor cortical signals that control the equivalent movement of a corresponding body part (e.g. arm movement to the right). This approach has been successfully applied in monkeys and humans by accurately extracting parameters of movements from the spiking activity of multiple single-units. Here, we review recent findings showing that analog neuronal population signals, ranging from intracortical local field potentials over epicortical ECoG to non-invasive EEG and MEG, can also be used to decode movement direction and continuous movement trajectories. Therefore, these signals might provide additional or alternative control for this BMI approach, with possible advantages due to reduced invasiveness.     

Methamphetamine Exposure During Development Causes Lasting Changes to Mesolimbic Dopamine Signaling in Mice

Methamphetamine (MA) abuse remains a public health issue. Prenatal MA exposure (PME) poses a significant health problem, as we know very little about the drug's long-term physiological impact on the developing human brain. We investigated the long-term consequences of early MA exposure using a mouse model that targets the brain growth spurt, which occurs during human third-trimester. Adult mice previously subjected to acute MA during post-natal days 4–9 exhibited hyperactivity during the Open-Field Test, while exhibiting no motor coordination changes during the Rotarod Test. Neonatal MA exposure reduced basal dopamine (DA) uptake rates in adult nucleus accumbens slices compared with saline-injected controls. Although slices from neonatal MA-exposed mice showed no change in evoked DA signals in the presence of MA, they exhibited potentiated non-evoked DA release through DA efflux in response to MA. These data suggest that developmental MA exposure alters brain development to produce long-lasting physiological changes to the adult mesolimbic DA system, as well as altering responses to acute MA exposure in adulthood. This study provides new insights into an important, under-investigated area in drugs of abuse research.

     

Altered Rolandic Gamma-Band Activation Associated with Motor Impairment and Ictal Network Desynchronization in Childhood Epilepsy

Epilepsy is associated with an abnormal expression of neural oscillations and their synchronization across brain regions. Oscillatory brain activation and synchronization also play an important role in cognition, perception and motor control. Childhood epilepsy is associated with a variety of cognitive and motor deficits, but the relationship between altered functional brain responses in various frequency ranges and functional impairment in these children remains poorly understood. We investigated functional magnetoencephalographic (MEG) responses from motor cortex in multiple functionally relevant frequency bands following median nerve stimulation in twelve children with epilepsy, including four children with motor impairments. We demonstrated that children with motor impairments exhibit an excessive gamma-band response from Rolandic cortex, and that the magnitude of this Rolandic gamma response is negatively associated with motor function. Abnormal responses from motor cortex were also associated with ictal desynchronization of oscillations within Rolandic cortex measured using intracranial EEG (iEEG). These results provide the evidence that ictal disruption of motor networks is associated with an altered functional response from motor cortex, which is in turn associated with motor impairment.     

The impact of basic motor abilities on the specific motoricity performance in elite karateka

The aim of the study was to identify motor structures that determine high performance in karate. The study included a group of 85 karateka aged 18-29 years, competing as senior category athletes within the Croatian Karate Society. A battery of 22 motor tests (9 basic motoricity tests and 13 specific motoricity tests) were used. Factor analysis of the basic motor variables pointed to the existence of three significant factors: coordination, explosive strength and movement frequency; whereas factor analysis of the specific motoricity area indicated two significant factors, i.e., factor of technical efficiency and factor of specific agility. Canonical correlation analysis showed the isolated set of basic motor factors to significantly determine both technical efficiency and specific agility-mobility of the karateka, with a predominance of the explosive strength (force) factor, followed by the speed and coordination factors.     

Contributions of Subsurface Cortical Modulations to Discrimination of Executed and Imagined Grasp Forces through Stereoelectroencephalography

Stereoelectroencephalographic (SEEG) depth electrodes have the potential to record neural activity from deep brain structures not easily reached with other intracranial recording technologies. SEEG electrodes were placed through deep cortical structures including central sulcus and insular cortex. In order to observe changes in frequency band modulation, participants performed force matching trials at three distinct force levels using two different grasp configurations: a power grasp and a lateral pinch. Signals from these deeper structures were found to contain information useful for distinguishing force from rest trials as well as different force levels in some participants. High frequency components along with alpha and beta bands recorded from electrodes located near the primary motor cortex wall of central sulcus and electrodes passing through sensory cortex were found to be the most useful for classification of force versus rest although one participant did have significant modulation in the insular cortex. This study electrophysiologically corroborates with previous imaging studies that show force-related modulation occurs inside of central sulcus and insular cortex. The results of this work suggest that depth electrodes could be useful tools for investigating the functions of deeper brain structures as well as showing that central sulcus and insular cortex may contain neural signals that could be used for control of a grasp force BMI.     

Using phase information to reveal the nature of event-related desynchronization

It is common knowledge among brain researchers that certain events, such as motor imagery, can elicit event-related synchronization/desynchronization (ERD/ERS) of neurons in the brain, and that these phenomena manifest themselves in terms of band power changes in brain signals. There are, however, also important phase related changes in the brain signals associated with ERD/ERS which we investigate here. We quantify for the first time the “delta-phase” component of the Complex Band Power (CBP) features introduced in [G. Townsend, B. Graimann, G. Pfurtscheller, A comparison of common spatial patterns with complex band power features in a four-class BCI Experiment, IEEE Trans. Biomed. Eng. 53 (2006) 642–651] and [G. Townsend, Brain Computer Interfaces: Phase Features and Post Processing Issues, Ph.D. Thesis, 2006, p. 97]. Although this component, which makes explicit use of phase, is known from these earlier studies to provide improved classification results in a brain–computer interface (BCI), this phenomenon has never been investigated purely in the context of a physiological phenomenon. The importance of this study is to fill this gap in our knowledge by providing an analysis and quantification of this new feature so that an improved understanding of its nature will allow further exploitation of this feature to further improve BCI systems. Given that the primary use of BCIs is as an assistive technology for the disabled, the importance of this work to society cannot be overstated. Changes in band power in the context of ERD/ERS have been investigated in the past using a visualization tool called an ERD/ERS map [B. Graimann, Movement-Related Patterns in ECoG and EEG: Visualization and Detection, Ph.D. Thesis, 2003, p. 107]. We present and discuss here for the first time, an equivalent tool called the delta-phase map used to visualize the occurrence of delta-phase in different frequency bands and over the time courses of motor imagery trials. In this study, the delta-phase feature is shown to be proportional to frequency and is quantified and presented in that context for clarity. It was found that the most prominent ERD activity occurring in the frequency bands studied here are correlated to the activity captured in the delta-phase maps. The underlying nature of ERD/ERS is explored based on evidence provided by the delta-phase feature.     

Encoding-based brain-computer interface controlled by non-motor area of rat brain

As the needs of disabled patients are increasingly recognized in society, researchers have begun to use single neuron activity to construct brain-computer interfaces (BCI), designed to facilitate the daily lives of individuals with physical disabilities. BCI systems typically allow users to control computer programs or external devices via signals produced in the motor or pre-motor areas of the brain, rather than producing actual motor movements. However, impairments in these brain areas can hinder the application of BCI. The current paper demonstrates the feasibility of a one-dimensional (1D) machine controlled by rat prefrontal cortex (PFC) neurons using an encoding method. In this novel system, rats are able to quench thirst by varying neuronal firing rate in the PFC to manipulate a water dish that can rotate in 1D. The results revealed that control commands generated by an appropriate firing frequency in rat PFC exhibited performance improvements with practice, indicated by increasing water-drinking duration and frequency. These results demonstrated that it is possible for rats to understand an encoding-based BCI system and control a 1D machine using PFC activity to obtain reward.     

The discontinuous nature of motor execution II. Merging discrete and rhythmic movements in a single-joint system -- the phase entrainment effect

 Initiation of rapid discrete flexion movements is significantly altered when a secondary rhythmic movement is performed simultaneously with the same limb; the onset of a stimulus-evoked discrete movement tends to occur time-locked to the oscillation: i.e., the rhythmic movement entrains the discrete response. This nonlinear interaction may reflect a specific principle of coordination of motor tasks which are simultaneously executed with the same effector. This part II of a tripartite research report on such single-muscle multiple-task coordination investigates the contribution of the dynamic properties of the muscle and its reflex circuitry to phase entrainment. Assuming a simple threshold-linear relationship between the control signals generated by the central nervous system and the observable kinematic and electromyographic signals, a secondary rhythmic movement will cause an additional phase-dependent delay between the central “go” command and the first observable change in actual kinematics of the compound movement. Several indicators for such threshold-linear interaction are derived and tested on real data obtained in psychophysical experiments. Four healthy subjects performed rapid lateral abductions of the index finger in response to a visual “go” signal. During a portion of the experiments, subjects produced additional low-amplitude oscillatory movements before stimulus presentation with either the same finger (one-handed task), or with the index finger of the other hand (two-handed task). Results showed phase entrainment and modulation of reaction times when the cyclic and the discrete movements were simultaneously executed by the same finger. But there was no entrainment in the bimanual execution of the tasks. The model was capable of reproducing the observed effects. It is concluded that coordination of voluntary movements which are concurrently performed by the same effector involves specific discontinuous operations, which represents an essential part of the mechanism of motor coordination. Phase entrainment reflects this characteristic discontinuous behavior of the lower stages of motor execution and does not necessarily require nonlinear interaction of motor commands at higher levels of motor processing. Received: 5 September 2001 / Accepted in revised form: 19 December 2001