Biomedical subjects

J R Wolpaw

Publications and source records attributed to J R Wolpaw.

At least 19 recordsLinked to original sources

PubMed1998

Answering questions with an electroencephalogram-based brain-computer interface.

OBJECTIVE: To demonstrate that humans can learn to control selected electroencephalographic components and use that control to answer simple questions. METHODS: Four adults (one with amyotrophic lateral sclerosis) learned to use electroencephalogram (EEG) mu rhythm (8 to 12Hz) or beta rhythm (18 to 25Hz) activity over sensorimotor cortex to control vertical cursor movement to targets at the top or bottom edge of a video screen. In subsequent sessions, the targets were replaced with the words YES and NO, and individuals used the cursor to answer spoken YES/NO questions from single- or multiple-topic question sets. They confirmed their answers through the response verification (RV) procedure, in which the word positions were switched and the question was answered again. RESULTS: For 5 consecutive sessions after initial question training, individuals were asked an average of 4.0 to 4.6 questions per minute; 64% to 87% of their answers were confirmed by the RV procedure and 93% to 99% of these answers were correct. Performances for single- and multiple-topic question sets did not differ significantly. CONCLUSIONS: The results indicate that (1) EEG-based cursor control can be used to answer simple questions with a high degree of accuracy, (2) attention to auditory queries and formulation of answers does not interfere with EEG-based cursor control, (3) question complexity (at least as represented by single versus multiple-topic question sets) does not noticeably affect performance, and (4) the RV procedure improves accuracy as expected. Several options for increasing the speed of communication appear promising. An EEG-based brain-computer interface could provide a new communication and control modality for people with severe motor disabilities.

Adult↗

PubMed1998

EEG-based communication and control: short-term role of feedback.

When people learn to control the amplitudes of certain electroencephalogram (EEG) components (e.g., the 8-12 Hz mu-rhythm over sensorimotor cortex) and use them to move a cursor to a target on a video screen, feedback about performance is normally provided by cursor movement and by trial outcome (i.e., success or failure). We assessed the short-term effects of this feedback on EEG control. After subjects received initial training with feedback present, feedback was removed intermittently for periods of several minutes. Subjects still displayed EEG control when feedback was removed. Removal of cursor movement alone appeared to have effects comparable to removal of both cursor movement and trial outcome. These results show that, in the short-term at least, mu-rhythm control is not dependent on the sensory input provided by cursor movement. They also suggest that feedback can have inhibitory as well as facilitory effects on EEG control, and that these effects vary across subjects. This finding has implications for the design of training procedures.

Adult↗

PubMed1998

EEG-based communication: improved accuracy by response verification.

Humans can learn to control the amplitude of electroencephalographic (EEG) activity in specific frequency bands over sensorimotor cortex and use it to move a cursor to a target on a computer screen. EEG-based communication could provide a new augmentative communication channel for individuals with motor disabilities. In the present system, each dimension of cursor movement is controlled by a linear equation. While the intercept in the equation is continually updated, it does not perfectly eliminate the impact of spontaneous variations in EEG amplitude. This imperfection reduces the accuracy of cursor movement. We evaluated a response verification (RV) procedure in which each outcome is determined by two opposite trials (e.g., one top-target trial and one bottom-target trial). Success, or failure, on both is required for a definitive outcome. The RV procedure reduces errors due to imperfection in intercept selection. Accuracy for opposite-trial pairs exceeds that predicted from the accuracies of individual trials, and greatly exceeds that for same-trial pairs. The RV procedure should be particularly valuable when the first trial has >2 possible targets, because the second trial need only confirm or deny the outcome of the first, and it should be applicable to nonlinear as well as to linear algorithms.

Adult↗

PubMed1997

Spatial filter selection for EEG-based communication.

Individuals can learn to control the amplitude of mu-rhythm activity in the EEG recorded over sensorimotor cortex and use it to move a cursor to a target on a video screen. The speed and accuracy of cursor movement depend on the consistency of the control signal and on the signal-to-noise ratio achieved by the spatial and temporal filtering methods that extract the activity prior to its translation into cursor movement. The present study compared alternative spatial filtering methods. Sixty-four channel EEG data collected while well-trained subjects were moving the cursor to targets at the top or bottom edge of a video screen were analyzed offline by four different spatial filters, namely a standard ear-reference, a common average reference (CAR), a small Laplacian (3 cm to set of surrounding electrodes) and a large Laplacian (6 cm to set of surrounding electrodes). The CAR and large Laplacian methods proved best able to distinguish between top and bottom targets. They were significantly superior to the ear-reference method. The difference in performance between the large Laplacian and small Laplacian methods presumably indicated that the former was better matched to the topographical extent of the EEG control signal. The results as a whole demonstrate the importance of proper spatial filter selection for maximizing the signal-to-noise ratio and thereby improving the speed and accuracy of EEG-based communication.

Adult↗

PubMed1997

The complex structure of a simple memory.

Operant conditioning of the vertebrate H-reflex, which appears to be closely related to learning that occurs in real life, is accompanied by plasticity at multiple sites. Change occurs in the firing threshold and conduction velocity of the motoneuron, in several different synaptic terminal populations on the motoneuron, and probably in interneurons as well. Change also occurs contralaterally. The corticospinal tract probably has an essential role in producing this plasticity. While certain of these changes, such as that in the firing threshold, are likely to contribute to the rewarded behavior (primary plasticity), others might preserve previously learned behaviors (compensatory plasticity), or are simply activity-driven products of change elsewhere (reactive plasticity). As these data and those from other simple vertebrate and invertebrate models indicate, a complex pattern of plasticity appears to be the necessary and inevitable outcome of even the simplest learning.

Animals↗

PubMed1997

Timing of EEG-based cursor control.

Recent studies show that humans can learn to control the amplitude of electroencephalography (EEG) activity in specific frequency bands over sensorimotor cortex and use it to move a cursor to a target on a computer screen. EEG-based communication could be a valuable new communication and control option for those with severe motor disabilities. Realization of this potential requires detailed knowledge of the characteristic features of EEG control. This study examined the course of EEG control after presentation of a target. At the beginning of each trial, a target appeared at the top or bottom edge of the subject's video screen and 1 sec later a cursor began to move vertically as a function of EEG amplitude in a specific frequency band. In well-trained subjects, this amplitude was high at the time the target appeared and then either remained high (i.e., for a top target) or fell rapidly (i.e., for a bottom target). Target-specific EEG amplitude control began 0.5 sec after the target appeared and appeared to wax and wane with a period of approximately 1 sec until the cursor reached the target (i.e., a hit) or the opposite edge of the screen (i.e., a miss). Accuracy was 90% or greater for each subject. Top-target errors usually occurred later in the trial because of failure to reach and/or maintain sufficiently high amplitude, whereas bottom-target errors usually occurred immediately because of failure to reduce an initially high amplitude quickly enough. The results suggest modifications that could improve performance. These include lengthening the intertrial period, shortening the delay between target appearance and cursor movement, and including time within the trial as a variable in the equation that translates EEG into cursor movement.

Adult↗

PubMed1997

Dorsal column but not lateral column transection prevents down-conditioning of H reflex in rats.

Operant conditioning of the H reflex, the electrical analogue of the spinal stretch reflex, in freely moving rats is a relatively simple model for studying long-term supraspinal control over spinal cord function. Motivated by food reward, rats can gradually increase or decrease the soleus H reflex. This study is the first effort to determine which spinal cord pathways convey the descending influence from supraspinal structures that changes the H reflex. In anesthetized Sprague-Dawley rats, the entire dorsal column (DC), which includes the main corticospinal tract, or the right lateral column (LC) was transected by electrocautery. Animals recovered quickly and the minimal transient effects of transection on the right soleus H reflex disappeared within 16 days. Beginning at least 18 days after transection, 12 rats were exposed to the HRdown-conditioning mode, in which reward was given when the H reflex of the right soleus muscle was below a criterion value. In seven LC rats exposed to the HRdown mode, the H reflex fell to 71 +/- 8% (mean +/- SE) of its initial value. In six of the seven, conditioning was successful (i.e., decrease to < or = 80%). These results were comparable with those previously obtained from normal rats. In contrast, in five DC rats exposed to the HRdown mode, the H reflex at the end of exposure was 106 +/- 12% of its initial value. In none of these rats was HRdown-conditioning successful. DC rats differed significantly from normal and LC rats in both final H reflex values and number successful. In five DC and three LC rats that continued under control conditions over 30-78 days, the H reflex at the end of the period was 98 +/- 4% and 100 +/- 8%, respectively, of its initial value, indicating that DC or LC transection itself did not lead to gradual increase or decrease in the H reflex. The results indicate that the DC, containing the main corticospinal tract, is essential for HRdown-conditioning, whereas the ipsilateral LC, containing the main rubrospinal, vestibulospinal, and reticulospinal tracts, is not essential. Combined with the known muscular specificity of conditioning, these results suggest that the main corticospinal tract is essential for HRdown-conditioning. The DC ascending tract might also be necessary. The respective roles of the DC descending and ascending tracts, and transection effects on HRup-conditioning and on the maintenance of both HRup- and HRdown-conditioning after they have occurred, remain to be defined.

Animals↗

PubMed1997

EEG-based communication: evaluation of alternative signal prediction methods.

Individuals can learn to control the amplitude of EEG activity in specific frequency bands over sensorimotor cortex and use it to move a cursor to a target on a computer screen. For one-dimensional (i.e., vertical) cursor movement, a linear equation translates the EEG activity into cursor movement. To translate an individual's EEG control into cursor control as effectively as possible, the intercept in this equation, which determines whether upward or downward movement occurs, should be set so that top and bottom targets are equally accessible. The present study compares alternative methods for using an individual's previous performance to select the intercept for subsequent trials. In offline analyses, five different intercept selection methods were applied to EEG data collected while trained subjects were moving the cursor to targets at the top or bottom edge of the screen. In the first two methods-moving average, and weighted sum-a single intercept was selected for the entire 1-2 sec period of each trial. In the other three methods-blocked moving average, blocked weighted sum, and blocked recursive sum (a variation of the weighted sum)-an intercept was selected for each 200-ms segment of the trial. The results from these methods were compared in regard to their balance between upward and downward movements and their consistency of performance across trials. For all subjects combined, the five methods performed similarly. However, performance across subjects was more consistent for the moving average, blocked moving average, and blocked recursive sum methods than for the weighted sum and blocked weighted sum methods. Due to its consistent performance and its computational simplicity, the moving average method, using the five most recent pairs of top and bottom trials, appears to be the method of choice.

Adult↗

PubMed1996

Operant conditioning of H-reflex changes synaptic terminals on primate motoneurons.

Operant conditioning of the primate triceps surae H-reflex, the electrical analog of the spinal stretch reflex, creates a memory trace that includes changes in the spinal cord. To define the morphological correlates of this plasticity, we analyzed the synaptic terminal coverage of triceps surae motoneurons from animals in which the triceps surae H-reflex in one leg had been increased (HRup mode) or decreased (HRdown mode) by conditioning and compared them to each other and to motoneurons from unconditioned animals. Motoneurons were labeled by intramuscular injection of cholera toxin-horseradish peroxidase. A total of 5055 terminals on the cell bodies and proximal dendrites of 114 motoneurons from 14 animals were studied by electron microscopy. Significant differences were found between HRup and HRdown animals and between HRup and naive (i.e., unconditioned) animals. F terminals (i.e., putative inhibitory terminals) were smaller and their active zone coverage on the cell body was lower on motoneurons from the conditioned side of HRup animals than on motoneurons from the conditioned side of HRdown animals. C terminals (i.e., terminals associated with postsynaptic cisterns and rough endoplasmic reticulum) were smaller and the number of C terminals in each C complex (i.e., a group of contiguous C terminals) was larger on motoneurons from the conditioned side of HRup animals than on motoneurons either from the conditioned side of HRdown animals or from naive animals. Because the treatment of HRup and HRdown animals differed only in the reward contingency, the results imply that the two contingencies had different effects on motoneuron synaptic terminals. In combination with other recent data, they show that H-reflex conditioning produces a complex pattern of spinal cord plasticity that includes changes in motoneuron physiological properties as well as in synaptic terminals. Further delineation of this pattern should reveal the contribution of the structural changes described here to the learned change in behavior.

Animals↗

PubMed1996

Reversal of H-reflex operant conditioning in the rat.

In response to an operant conditioning task, rats can gradually increase or decrease soleus H-reflex amplitude without change in background electromyographic activity or M response amplitude. Both increase (under the HRup mode) and decrease (under the HRdown mode) develop over weeks. The present study investigated reversal of conditioned H-reflex change. Following collection of control data, rats were exposed to one mode (HRup or HRdown) for 50 days, and then exposed to the opposite mode for up to 72 days. Rats responded to each mode exposure with gradual, mode-appropriate change in H-reflex amplitude. This finding is consistent with other evidence that H-reflex conditioning depends on spinal cord plasticity. The effects of exposure to the HRup (or HRdown) mode were not affected by whether exposure followed previous exposure to the HRdown (or HRup) mode. In accord with recent studies suggesting that HRup and HRdown conditioning have different spinal mechanisms, these results suggest that reversal of H-reflex change is due primarily to the superimposition of additional plasticity rather than to decay of the plasticity responsible for the initial change.

Animals↗

PubMed1996

Operant conditioning of H-reflex in spinal cord-injured rats.

Operant conditioning of the spinal stretch reflex or its electrical analog, the H-reflex, is a new model for exploring the mechanisms of supraspinal control over spinal cord function. Both rats and primates can gradually increase (HRup conditioning mode) or decrease (HRdown conditioning mode) soleus H-reflex magnitude when exposed to an operant conditioning task. This study used H-reflex operant conditioning to assess and modify spinal cord function after injury. Soleus H-reflexes were elicited and recorded with chronically implanted electrodes from rats that had been subjected to calibrated contusion injuries to the spinal cord at T8. From 18 to 140 days after injury, background EMG, M response amplitude, and initial H-reflex amplitude were not significantly different from those of normal rats. HRdown conditioning was successful in some, but not all, spinal cord-injured rats. The H-reflex decrease achieved by conditioning was inversely correlated with the severity of the injury as assessed histologically or by time to return of bladder function. It was not correlated with the length of time between injury and the beginning of conditioning. The results confirm the importance of descending control from supraspinal structures in mediating operantly conditioned change in H-reflex amplitude. In conjunction with recent human studies, they suggest that H-reflex conditioning could provide a sensitive new means for assessing spinal cord function after injury, and might also provide a method for initiating and guiding functional rehabilitation.

Animals↗

PubMed1996

Therapeutic neural effects of electrical stimulation.

The use of a functional neuromuscular stimulation (FNS) device can have therapeutic effects that persist when the device is not in use. Clinicians have reported changes in both voluntary and electrically assisted neuromuscular function and improvements in the condition of soft tissue. Motor recovery has been observed in people with incomplete spinal cord injury, stroke, or traumatic brain injury after the use of motor prostheses. Improvement in voluntary dorsiflexion and overall gait pattern has been reported both in the short term (several hours) and permanently. Electrical stimulation of skin over flexor muscles in the upper limb produced substantial reductions for up to 1 h in the severity of spasticity in brain-injured subjects, as measured by the change in torque generation during ramp-and-hold muscle stretch. There was typically an aggravation of the severity of spasticity when surface stimulation reached intensities sufficient to also excite muscle. Animals were trained to alter the size of the H-reflex to obtain a reward. The plasticity that underlies this operantly conditioned H-reflex change includes changes in the spinal cord itself. Comparable changes appear to occur with acquisition of certain motor skills. Current studies are exploring such changes in humans and animals with spinal cord injuries with the goal of using conditioning methods to assess function after injury and to promote and guide recovery of function. A better understanding of the mechanisms of neural plasticity, achieved through human and animal studies, may help us to design and implement FNS systems that have the potential to produce beneficial changes in the subject's central nervous systems.

Animals↗

PubMed1996

EEG-based communication: prospects and problems.

Current rehabilitation engineering combines new prosthetic methods with recent developments in personal computers to provide alternative communication and control channels to individuals with motor impairments. Despite these advances, all commercially available systems still require some measure of voluntary motor control. Thus, these systems are not useful for individuals who are totally paralyzed. Electroencephalographic (EEG) activity may provide the basis for a system that would completely bypass normal motor output. EEG-based communication technology might provide assistive devices for individuals who have little or no reliable motor function. This paper reviews the prospects for and problems of EEG-based communication. It summarizes current approaches to development of this new technology, describes the major problems that must be resolved, and focuses on issues critical for its use by those with severe motor disabilities.

Brain↗

PubMed1995

Operantly conditioned plasticity and circadian rhythm in rat H-reflex are independent phenomena.

Recent studies indicate that rats can increase or decrease H-reflex amplitude in response to an operant conditioning paradigm. In addition, rats also display a circadian rhythm in H-reflex amplitude. As part of the development of H-reflex conditioning in the rat as a new model for defining the plasticity underlying a simple form of learning, this study examined the relationship in the rat between operantly conditioned H-reflex change and the H-reflex circadian rhythm. When H-reflex amplitude increased or decreased in response to the operant conditioning program, its circadian rhythm showed no changes in phase and minimal change in amplitude. Furthermore, animals did not alter daily performance schedule so as to use the rhythm to increase reward probability. Thus, in the rat, H-reflex operant conditioning and the H-reflex circadian rhythm appear to be independent phenomena. The circadian rhythm should not be a significant complicating factor in studies of operantly conditioned H-reflex change.

Animals↗

PubMed1995

Operant conditioning of H-reflex in freely moving rats.

1. Primates can increase or decrease the spinal stretch reflex and its electrical analogue, the H-reflex (HR), in response to an operant conditioning task. This conditioning changes the spinal cord itself and thereby provides an experimental model for defining the processes and substrates of a learned change in behavior. Because the phenomenon has been demonstrated only in primates, its generality and theoretical implications remain unclear, and its experimental use is restricted by the difficulties of primate research. In response to these issues, the present study explored operant conditioning of the H-reflex in the rat. 2. Seventeen Sprague-Dawley rats implanted with chronic electromyographic (EMG) recording electrodes in one soleus muscle and nerve cuff stimulating electrodes on the posterior tibial nerve were rewarded (either with medial forebrain bundle stimulation or food) for increasing (HRup conditioning mode) or decreasing (HRdown conditioning mode) soleus H-reflex amplitude without change in background EMG or M response (direct muscle response) amplitude. 3. H-reflex amplitude changed appropriately over 3-4 wk. Under the HRup mode, it rose to an average of 158 +/- 54% (mean +/- SD) of initial value, whereas under the HRdown mode it fell to an average of 67 +/- 11% of initial value. Background EMG and M response amplitude did not change. 4. Operant conditioning of the H-reflex in the rat appears similar in rate and final magnitude of change to that observed in the monkey.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1995

Operantly conditioned motoneuron plasticity: possible role of sodium channels.

1. Learning is traditionally thought to depend on synaptic plasticity. However, recent work shows that operantly conditioned decrease in the primate H reflex is associated with an increase in the depolarization needed to fire the spinal motoneuron (VDEP) and a decrease in its conduction velocity (CV). Furthermore, the increase in VDEP appears to be largely responsible for the H-reflex decrease. The conjunction of these changes in VDEP and CV suggests that an alteration in Na+ channel properties throughout the soma and axon could be responsible. 2. A mathematical model of the mammalian myelinated axon was used to test whether a positive shift in the voltage dependence of Na+ channel activation, a decrease in Na+ channel peak permeability, or changes in other fiber properties could have accounted for the experimental findings. 3. A positive shift of 2.2 mV in Na+ channel activation reproduced the experimentally observed changes in VDEP and CV, whereas a reduction in Na+ channel permeability or changes in other fiber properties did not. 4. These results are consistent with the hypothesis that operantly conditioned decrease in the primate H reflex is largely due to a positive shift in the voltage dependence of Na+ channel activation. Recent studies suggest that change in activation of protein kinase C may mediate this effect.

Action Potentials↗

PubMed1995

Motoneuron properties after operantly conditioned increase in primate H-reflex.

1. Monkeys can increase (HRup conditioning mode) or decrease (HRdown conditioning mode) the triceps surae (TS) H-reflex in response to an operant conditioning task. This conditioning modifies the spinal cord. To define this spinal cord plasticity and its role in the behavioral change (H-reflex increase or decrease), we have recorded intracellularly from TS motoneurons in conditioned animals. The present report describes data from HRup animals and compares them with data from previously studied naive (NV; i.e., unconditioned) animals. 2. Thirteen monkeys (Macaca nemestrina, male, 3.8-7.1 kg) were exposed to the HRup conditioning mode, in which reward occurred when H-reflex size in one leg (i.e., the trained leg) was above a criterion value. Conditioning was successful (i.e., increase of > or = 20%) in 12 of the 13 animals. At the end of conditioning, H-reflex size in the trained leg averaged 188% of its initial value, whereas size in the control leg averaged 134% of its initial value. 3. Intracellular recordings were obtained from 136 TS motoneurons on trained (UT + motoneurons) and control (UC + motoneurons) sides of the successful animals. Measurements included axonal conduction velocity, input resistance, time constant, electrotonic length, rheobase, firing threshold to current injection, afterhyperpolarization duration and amplitude, and composite homonymous and heteronymous excitatory postsynaptic potential (EPSP) size and shape. Results were compared with intracellular data from NV animals.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1994

Synaptic terminal coverage of primate triceps surae motoneurons.

This study examined the synaptic terminal coverage of primate triceps surae (TS) motoneurons at the electron microscopic level. In three male pigtail macaques, motoneurons were labeled by retrograde transport of cholera toxin-horseradish peroxidase that was injected into TS muscles bilaterally and visualized with tetramethylbenzidine stabilized with diaminobenzidine. Somatic, proximal dendritic, and distal dendritic synaptic terminals were classified by standard criteria and measured. Overall and type-specific synaptic terminal coverages and frequencies were determined. Labeled cells were located in caudal L5 to rostral S1 ventral horn and ranged from 40 to 74 microns in diameter (average, 54 microns). The range and unimodal distribution of diameters, the label used, and the presence of C terminals on almost all cells indicated that the 15 cell bodies and associated proximal dendrites analyzed here probably belonged to alpha-motoneurons. Synaptic terminals covered 39% of the cell body membrane, 60% of the proximal dendritic membrane, and 40% of the distal dendritic membrane. At each of these three sites, F terminals (flattened or pleomorphic vesicles, usually symmetric active zones, average contact length 1.6 microns) were most common, averaging 52%, 56%, and 58% of total coverage and 56%, 57%, and 58% of total number of cell bodies, proximal dendrites, and distal dendrites respectively. S terminals (round vesicles, usually asymmetric active zones, average contact length 1.3 microns) averaged 24%, 29%, and 33% of coverage and 33%, 35%, and 36% of number at these three sites, respectively. Thus, S terminals were slightly more prominent relative to F terminals on distal dendrites than on cell bodies. C terminals (spherical vesicles, subsynaptic cisterns associated with rough endoplasmic reticulum, average contact length 3.5 microns) constituted 24% and 11% of total terminal coverage on cell bodies and proximal dendrites, respectively, and averaged 11% and 6% of terminal number at these two locations. M terminals (spherical vesicles, postsynaptic Taxi bodies, some with presynaptic terminals, average contact length 2.7 microns) were absent on cell bodies and averaged 3% and 7% of total coverage and 2% and 5% of terminals on proximal and distal dendrites, respectively. Except for M terminals, which tended to be smaller distally, terminal contact length was not correlated with location. Total and type-specific coverages and frequencies were not correlated with cell body diameter. Primate TS motoneurons are similar to cat TS motoneurons in synaptic terminal morphology, frequency, and distribution. However, primate terminals appear to be smaller, so that the fraction of membrane covered by them is lower.

Animals↗