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Brain–Computer Interfaces - Index of

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210 D.M. Taylor and M.E. Stetner<br />

designed analog circuits that transformed the voltage spikes detected by the electrodes<br />

into the movement <strong>of</strong> a needle on a voltmeter. The circuits were designed so<br />

that the needle would move when the monkey increased the firing rate <strong>of</strong> individually<br />

recorded neurons sometimes while decreasing activity in associated muscles.<br />

Most importantly, the animal was allowed to watch the needle’s movements and<br />

was rewarded when he moved the needle past a certain mark. The animal quickly<br />

learned to control the firing activity <strong>of</strong> the neurons as assigned in order to move the<br />

voltmeter needle and get a reward. These early studies show the power <strong>of</strong> visual<br />

feedback, and they demonstrated that monkeys (and presumably humans) have the<br />

ability to quickly learn to willfully modulate the firing patterns <strong>of</strong> individual neurons<br />

for the purpose <strong>of</strong> controlling external devices.<br />

Along side this initial BCI work, other researchers were implanting intracortical<br />

microelectrodes in brains <strong>of</strong> monkeys simply to try to understand how our<br />

nervous system controls movement. These early studies have laid the foundation for<br />

many BCI systems that decode one’s intended arm and hand movement in real time<br />

and use that desired movement command to control the movements <strong>of</strong> a computer<br />

cursor, a robotic arm, or even one’s own paralyzed arm activated via implanted stimulators.<br />

One seminal study was done in the 1980s by Dr. Apostolos Georgopoulos<br />

and his colleagues who showed that the different neurons in the arm areas <strong>of</strong> the<br />

brain are “directionally tuned” [3]. Each neuron has what’s called a “preferred direction”.<br />

Each neuron will fire at its maximum rate as you are about to move your arm<br />

in that neuron’s preferred direction. The neuron’s firing rate will decrease as the arm<br />

movements start deviating from that cell’s preferred direction. Finally, the neuron<br />

will fire at its minimum rate as you move your arm opposite the neuron’s preferred<br />

direction. Different neurons have different preferred directions. By looking at the<br />

firing rates <strong>of</strong> many different directionally-tuned neurons, we can deduce how the<br />

arm is moving fairly reliably [4].<br />

Unfortunately, each neuron’s firing patterns are not perfectly linked to one’s<br />

movement intent. Neurons have a substantial amount <strong>of</strong> variability in their firing patterns<br />

and each one only provides a noisy estimate <strong>of</strong> intended movement. In 1988,<br />

Georgopoulos and Massey conducted a study to determine how accurately movement<br />

direction could be predicted using different numbers <strong>of</strong> motor cortex neurons<br />

[5]. They recorded individual neurons in monkeys from the area <strong>of</strong> the brain that<br />

controls arm movements. They recorded these neurons while the monkeys made a<br />

sequence <strong>of</strong> “center-out” movements with a manipulandum, which is a device that<br />

tracks hand position as one moves a handle along a tabletop. In these experiments,<br />

the table top had a ring <strong>of</strong> equally-spaced target lights and one light in the center<br />

<strong>of</strong> this ring that indicated the starting hand position. The monkey would start by<br />

holding the manipulandum over the center light. When one <strong>of</strong> the radial target lights<br />

would come on, the animal would move the manipulandum outward as fast as it<br />

could to the radial target to get a reward 1 . The researchers recorded many different<br />

1 Because the researchers imposed time limits on hitting the targets, the subjects tried to move<br />

quickly to the targets and did not always hit the target accurately.

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