Bereitschaftspotential
Encyclopedia
In neurology
, the Bereitschaftspotential or BP (from German
, "readiness potential"), also called the pre-motor potential or readiness potential (RP), is a measure of activity in the motor cortex
of the brain leading up to voluntary muscle movement. The BP is a manifestation of cortical contribution to the pre-motor planning of volitional movement. It was first recorded and reported in 1964 by Hans Helmut Kornhuber and Lüder Deecke
at the University of Freiburg in Germany
. In 1965 the full publication appeared after many control experiments.
(his doctoral student) went for lunch to the 'Gasthaus zum Schwanen' at the foot of the Schlossberg hill in Freiburg. Sitting alone in the beautiful garden they discussed their frustration with the passive brain research prevailing worldwide and their desire to investigate self-initiated action of the brain and the will. Consequently they decided to look for cerebral potentials in man related to volitional acts and to take voluntary movement as their research paradigm.
The possibility to do research on electrical brain potentials preceding voluntary movements came with the advent of the 'computer of average transients,' the first, primitive instrument available at that time in the Freiburg laboratory. In the electroencephalogram (EEG) little is to be seen preceding actions, except of an inconstant diminution of the α- (or μ-) rhythm. The young researchers stored the electroencephalogram and electromyogram of self-initiated movements (fast finger flexions) on tape and analyzed the cerebral potentials preceding movements time-reversed with the start of the movement as the trigger, literally turning the tape over for analysis since they had no reversal playback or programmable computer. A potential preceding human voluntary movement was discovered and published in the same year. After detailed investigation and control experiments such as passive finger movements the Citation Classic with the term Bereitschaftspotential was published.
The BP is ten to hundred times smaller than the α-rhythm
of the EEG; only by averaging, relating the electrical potentials to the onset of the movement it becomes apparent. Figure shows the typical slow shifts of the cortical DC potential, called Bereitschaftspotential, preceding volitional, rapid flexion
s of the right index finger. The vertical line indicates the instant of triggering t = 0 (first activity in the EMG of the agonist muscle). Recording positions are left precentral (L prec, C3), right precentral (R prec, C4), mid-parietal (Pz); these are unipolar recordings with linked ears as reference. The difference between the BP in C3 and in C4 is displayed in the lowest graph (L/R prec). Superimposed are the results of eight experiments as obtained in the same subject (B.L.) on different days [citation needed].
Note that the BP has two components, the early one (BP1) lasting from about −1.2 to −0.5; the late component (BP2) from −0.5 to shortly before 0 sec. The pre-motion positivity is even smaller, and the motor-potential which starts about fifty to sixty milliseconds before the onset of movement and has its maximum over the contralateral precentral hand area is still smaller. Thus, it takes great care to see these potentials: exact triggering by the real onset of movement is important, which is especially difficult preceding speech movements. Furthermore artifacts due to head-, eye-, lid-, mouth-movements and respiration have to be eliminated before averaging because such artifacts may be of a magnitude which makes it difficult to render them negligible even after hundreds of sweeps. In the case of eye movements eye muscle potentials have to be distinguished from cerebral potentials. In some cases animal experiments were necessary to clarify the origin of potentials such as the R-wave. Therefore, it took many years until some of the other laboratories were able to confirm the details of Kornhuber & Deecke's results. In addition to the finger or eye movements as mentioned above, the BP has been recorded accompanying willful movements of the wrist, arm, shoulder, hip, knee, foot and toes. It was also recorded prior to speaking, writing and also swallowing.
The magnetoencephalographic (MEG) equivalent of the Bereitschaftspotential (BP), 'Bereitschafts(magnetic)field' (BF), or readiness field (RF) was first recorded in Hal Weinberg's laboratory at Simon Fraser University Burnaby B.C. Canada in 1982. It was confirmed that the early component, BP 1 or BF1, respectively was generated by the supplementary motor area (SMA), including the pre-SMA, while the late component, BP2 or BF2, was generated by the primary motor area, MI.
Another ERP component which was discovered in 1964 is the contingent negative variation
, or CNV. The CNV also composes two waves; the initial wave (i.e., O wave) and the terminal wave (i.e., E wave). The terminal CNV has similar characteristics as the BP and many researchers have claimed that the BP and the terminal CNV are the same component. Other researchers have shown that they can be disassociated.
who investigated the laws of motion of the universe with metal balls on an inclined plane”. The interest was even greater in psychology and philosophy because volition is traditionally associated with human freedom (cf. Kornhuber 1984). The spirit of the time, however, was hostile to freedom in those years; it was believed that freedom is an illusion. The tradition of behaviourism and Freudism was deterministic. While will and volition were frequently leading concepts in psychological research papers before and after the first world war and even during the second war, after the end of the second world war this declined, and by the mid-sixties these key words completely disappeared and were abolished in the thesaurus of the American Psychological Association. The BP is an electrical sign of participation of the supplementary motor area (SMA) prior to volitional movement, which starts activity prior to the primary motor area. The BP has preciptated a worldwide discussion about free will
(cf. the closing chapter in the recent book "The Bereitschaftspotential").
EEG
s and EMG
s are used in combination with Bayesian inference
to construct Bayesian network
s which attempt to predict general patterns of Motor Intent Neuron Action Potential
s firing. Researchers attempting to develop non-intrusive brain-machine interfaces are interested in this, as are system analysis, operations research, and epistemology (e.g. the Smith predictor has been suggested in the discussion). (explanation needed!)
studied the relationship between conscious experience of volition
and the BP e.g. and found that the BP started about 0.35 sec earlier than the subject's reported conscious awareness that 'now he or she feels the desire to make a movement.' Libet concludes that we have no free will
in the initiation of our movements; though, since subjects were able to prevent intended movement at the last moment, we do have a veto.
These studies have provoked widespread debate. See also the recent extensive critique by Klemm, 2010).
(BCI) applications; this signal feature can be identified from scalp recording (even from single-trial measurements) and interpreted for various uses, for example control of computer displays or control of peripheral motor units in spinal cord injuries.
Neurology
Neurology is a medical specialty dealing with disorders of the nervous system. Specifically, it deals with the diagnosis and treatment of all categories of disease involving the central, peripheral, and autonomic nervous systems, including their coverings, blood vessels, and all effector tissue,...
, the Bereitschaftspotential or BP (from German
German language
German is a West Germanic language, related to and classified alongside English and Dutch. With an estimated 90 – 98 million native speakers, German is one of the world's major languages and is the most widely-spoken first language in the European Union....
, "readiness potential"), also called the pre-motor potential or readiness potential (RP), is a measure of activity in the motor cortex
Motor cortex
Motor cortex is a term that describes regions of the cerebral cortex involved in the planning, control, and execution of voluntary motor functions.-Anatomy of the motor cortex :The motor cortex can be divided into four main parts:...
of the brain leading up to voluntary muscle movement. The BP is a manifestation of cortical contribution to the pre-motor planning of volitional movement. It was first recorded and reported in 1964 by Hans Helmut Kornhuber and Lüder Deecke
Lüder Deecke
Dr. Lüder Deecke is a neurologist, neuroscientist, teacher and physician whose scientific discoveries have influenced brain research and the treatment and rehabilitation of neurological disorders....
at the University of Freiburg in Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...
. In 1965 the full publication appeared after many control experiments.
Discovery
In the spring of 1964 Hans Helmut Kornhuber (then docent and chief physician at the department of neurology, head Professor Richard Jung, university hospital Freiburg im Breisgau) and Lüder DeeckeLüder Deecke
Dr. Lüder Deecke is a neurologist, neuroscientist, teacher and physician whose scientific discoveries have influenced brain research and the treatment and rehabilitation of neurological disorders....
(his doctoral student) went for lunch to the 'Gasthaus zum Schwanen' at the foot of the Schlossberg hill in Freiburg. Sitting alone in the beautiful garden they discussed their frustration with the passive brain research prevailing worldwide and their desire to investigate self-initiated action of the brain and the will. Consequently they decided to look for cerebral potentials in man related to volitional acts and to take voluntary movement as their research paradigm.
The possibility to do research on electrical brain potentials preceding voluntary movements came with the advent of the 'computer of average transients,' the first, primitive instrument available at that time in the Freiburg laboratory. In the electroencephalogram (EEG) little is to be seen preceding actions, except of an inconstant diminution of the α- (or μ-) rhythm. The young researchers stored the electroencephalogram and electromyogram of self-initiated movements (fast finger flexions) on tape and analyzed the cerebral potentials preceding movements time-reversed with the start of the movement as the trigger, literally turning the tape over for analysis since they had no reversal playback or programmable computer. A potential preceding human voluntary movement was discovered and published in the same year. After detailed investigation and control experiments such as passive finger movements the Citation Classic with the term Bereitschaftspotential was published.
Mechanism
Alpha wave
Alpha waves are neural oscillations in the frequency range of 8–12 Hz arising from synchronous and coherent electrical activity of thalamic pacemaker cells in humans...
of the EEG; only by averaging, relating the electrical potentials to the onset of the movement it becomes apparent. Figure shows the typical slow shifts of the cortical DC potential, called Bereitschaftspotential, preceding volitional, rapid flexion
Flexion
In anatomy, flexion is a position that is made possible by the joint angle decreasing. The skeletal and muscular systems work together to move the joint into a "flexed" position. For example the elbow is flexed when the hand is brought closer to the shoulder...
s of the right index finger. The vertical line indicates the instant of triggering t = 0 (first activity in the EMG of the agonist muscle). Recording positions are left precentral (L prec, C3), right precentral (R prec, C4), mid-parietal (Pz); these are unipolar recordings with linked ears as reference. The difference between the BP in C3 and in C4 is displayed in the lowest graph (L/R prec). Superimposed are the results of eight experiments as obtained in the same subject (B.L.) on different days [citation needed].
Note that the BP has two components, the early one (BP1) lasting from about −1.2 to −0.5; the late component (BP2) from −0.5 to shortly before 0 sec. The pre-motion positivity is even smaller, and the motor-potential which starts about fifty to sixty milliseconds before the onset of movement and has its maximum over the contralateral precentral hand area is still smaller. Thus, it takes great care to see these potentials: exact triggering by the real onset of movement is important, which is especially difficult preceding speech movements. Furthermore artifacts due to head-, eye-, lid-, mouth-movements and respiration have to be eliminated before averaging because such artifacts may be of a magnitude which makes it difficult to render them negligible even after hundreds of sweeps. In the case of eye movements eye muscle potentials have to be distinguished from cerebral potentials. In some cases animal experiments were necessary to clarify the origin of potentials such as the R-wave. Therefore, it took many years until some of the other laboratories were able to confirm the details of Kornhuber & Deecke's results. In addition to the finger or eye movements as mentioned above, the BP has been recorded accompanying willful movements of the wrist, arm, shoulder, hip, knee, foot and toes. It was also recorded prior to speaking, writing and also swallowing.
The magnetoencephalographic (MEG) equivalent of the Bereitschaftspotential (BP), 'Bereitschafts(magnetic)field' (BF), or readiness field (RF) was first recorded in Hal Weinberg's laboratory at Simon Fraser University Burnaby B.C. Canada in 1982. It was confirmed that the early component, BP 1 or BF1, respectively was generated by the supplementary motor area (SMA), including the pre-SMA, while the late component, BP2 or BF2, was generated by the primary motor area, MI.
Another ERP component which was discovered in 1964 is the contingent negative variation
Contingent negative variation
The contingent negative variation was one of the first event-related potential components to be described. The CNV component was first described by Dr. W. Grey Walter and colleagues in an article published in Nature in 1964...
, or CNV. The CNV also composes two waves; the initial wave (i.e., O wave) and the terminal wave (i.e., E wave). The terminal CNV has similar characteristics as the BP and many researchers have claimed that the BP and the terminal CNV are the same component. Other researchers have shown that they can be disassociated.
Outcomes
The Bereitschaftspotential was received with great interest by the scientific community, as reflected by Sir John Eccles's comment: “There is a delightful parallel between these impressively simple experiments and the experiments of Galileo GalileiGalileo Galilei
Galileo Galilei , was an Italian physicist, mathematician, astronomer, and philosopher who played a major role in the Scientific Revolution. His achievements include improvements to the telescope and consequent astronomical observations and support for Copernicanism...
who investigated the laws of motion of the universe with metal balls on an inclined plane”. The interest was even greater in psychology and philosophy because volition is traditionally associated with human freedom (cf. Kornhuber 1984). The spirit of the time, however, was hostile to freedom in those years; it was believed that freedom is an illusion. The tradition of behaviourism and Freudism was deterministic. While will and volition were frequently leading concepts in psychological research papers before and after the first world war and even during the second war, after the end of the second world war this declined, and by the mid-sixties these key words completely disappeared and were abolished in the thesaurus of the American Psychological Association. The BP is an electrical sign of participation of the supplementary motor area (SMA) prior to volitional movement, which starts activity prior to the primary motor area. The BP has preciptated a worldwide discussion about free will
Free will
"To make my own decisions whether I am successful or not due to uncontrollable forces" -Troy MorrisonA pragmatic definition of free willFree will is the ability of agents to make choices free from certain kinds of constraints. The existence of free will and its exact nature and definition have long...
(cf. the closing chapter in the recent book "The Bereitschaftspotential").
EEG
Electroencephalography
Electroencephalography is the recording of electrical activity along the scalp. EEG measures voltage fluctuations resulting from ionic current flows within the neurons of the brain...
s and EMG
Electromyography
Electromyography is a technique for evaluating and recording the electrical activity produced by skeletal muscles. EMG is performed using an instrument called an electromyograph, to produce a record called an electromyogram. An electromyograph detects the electrical potential generated by muscle...
s are used in combination with Bayesian inference
Bayesian inference
In statistics, Bayesian inference is a method of statistical inference. It is often used in science and engineering to determine model parameters, make predictions about unknown variables, and to perform model selection...
to construct Bayesian network
Bayesian network
A Bayesian network, Bayes network, belief network or directed acyclic graphical model is a probabilistic graphical model that represents a set of random variables and their conditional dependencies via a directed acyclic graph . For example, a Bayesian network could represent the probabilistic...
s which attempt to predict general patterns of Motor Intent Neuron Action Potential
Action potential
In physiology, an action potential is a short-lasting event in which the electrical membrane potential of a cell rapidly rises and falls, following a consistent trajectory. Action potentials occur in several types of animal cells, called excitable cells, which include neurons, muscle cells, and...
s firing. Researchers attempting to develop non-intrusive brain-machine interfaces are interested in this, as are system analysis, operations research, and epistemology (e.g. the Smith predictor has been suggested in the discussion). (explanation needed!)
BP and free will
In a series of experiments in the 1980s, Benjamin LibetBenjamin Libet
Benjamin Libet April 12, 1916, Chicago, Illinois - July 23, 2007, Davis, California) was a pioneering scientist in the field of human consciousness. Libet was a researcher in the physiology department of the University of California, San Francisco...
studied the relationship between conscious experience of volition
Volition
Volition may refer to:*Volition , the cognitive process by which an individual decides on and commits to a particular course of action...
and the BP e.g. and found that the BP started about 0.35 sec earlier than the subject's reported conscious awareness that 'now he or she feels the desire to make a movement.' Libet concludes that we have no free will
Free will
"To make my own decisions whether I am successful or not due to uncontrollable forces" -Troy MorrisonA pragmatic definition of free willFree will is the ability of agents to make choices free from certain kinds of constraints. The existence of free will and its exact nature and definition have long...
in the initiation of our movements; though, since subjects were able to prevent intended movement at the last moment, we do have a veto.
These studies have provoked widespread debate. See also the recent extensive critique by Klemm, 2010).
Applications
An interesting use of the Bereitschaftspotential is in Brain-Computer InterfaceBrain-computer interface
A brain–computer interface , sometimes called a direct neural interface or a brain–machine interface , is a direct communication pathway between the brain and an external device...
(BCI) applications; this signal feature can be identified from scalp recording (even from single-trial measurements) and interpreted for various uses, for example control of computer displays or control of peripheral motor units in spinal cord injuries.
Further reading
- Deecke, L.; Kornhuber, H.H. (2003). Human freedom, reasoned will, and the brain. The Bereitschaftspotential story. In: M Jahanshahi, M Hallett (Eds.) The Bereitschaftspotential, movement-related cortical potentials. Kluwer Academic / Plenum Publishers [ISBN 0-306-47407-7] pp. 283–320.
- Kornhuber HH; Deecke L (2007) Wille und Gehirn [will and the brain]. Edition Sirius. Bielefeld/Locarno. 149 pp [ISBN 987-3-89528-628-5]
External links
- http://www.cmds.canterbury.ac.nz/documents/huckabee_swallowing.pdf
- http://www.cs.washington.edu/homes/rao/shenoy_rao05.pdf
See also
- EpiphenomenalismEpiphenomenalismIn philosophy of mind, epiphenomenalism, also known as Type-E Dualism, is a view that "mental" states do not have any influence on "physical" states.-Background:...
- Somatosensory evoked potentialSomatosensory Evoked PotentialSomatosensory Evoked Potentials are a useful, noninvasive means of assessing somatosensory system functioning. By combining SEP recordings at different levels of the somatosensory pathways, it is possible to assess the transmission of the afferent volley from the periphery up to the cortex...
- C1 and P1C1 & P1 (Neuroscience)The C1 and P1 are two human scalp-recorded event-related brain potential components, collected by means of a technique called electroencephalography . The C1 is named so because it was the first component in a series of components found to respond to visual stimuli when it was first discovered...
- Visual N1Visual N1The Visual N1 is a visual evoked potential, a type of event-related electrical potential , that is produced in the brain and recorded on the scalp. The N1 is so named to reflect the polarity and typical timing of the component. The "N" indicates that the polarity of the component is negative with...
- Mismatch negativityMismatch negativityThe mismatch negativity or mismatch field is a component of the event-related potential to an odd stimulus in a sequence of stimuli. It arises from electrical activity in the brain and is studied within the field of cognitive neuroscience and psychology. It can occur in any sensory system, but...
- N100
- N200N200 (neuroscience)The N200, or N2, is an event-related potential component. An ERP can be monitored using a non-invasive electroencephalography cap that is fitted over the scalp on human subjects...
- N2pcN2pcN2pc refers to an ERP component linked to selective attention. The N2pc appears over visual cortex contralateral to the location in space to which subjects are attending; if subjects pay attention to the left side of the visual field, the N2pc appears in the right hemisphere of the brain, and...
- N170N170The N170 is a component of the event-related potential that reflects the neural processing of faces.When potentials evoked by images of faces are compared to those elicited by other visual stimuli, the former show increased negativity 130-200 ms after stimulus presentation...
- P200P200In neuroscience, the visual P200 or P2 is a waveform component or feature of the event-related potential measured at the human scalp. Like other potential changes measurable from the scalp, this effect is believed to reflect the post-synaptic activity of a specific neural process...
- N400
- P300 (neuroscience)P300 (neuroscience)The P300 wave is an event related potential elicited by infrequent, task-relevant stimuli. It is considered to be an endogenous potential as its occurrence links not to the physical attributes of a stimulus but to a person's reaction to the stimulus. More specifically, the P300 is thought to...
- P3aP3aThe P3a, or novelty P3, is a component of time-locked signals known as event-related potentials . The P3a is a positive-going scalp-recorded brain potential that has a maximum amplitude over frontal/central electrode sites with a peak latency falling in the range of 250-280 ms...
- P3bP3bThe P3b is a subcomponent of the P300, an event-related potential component that can be observed in human scalp recordings of brain electrical activity...
- Late Positive ComponentLate Positive ComponentThe LPC is a positive-going event-related brain potential component that has been important in studies of explicit recognition memory...
- Difference due to MemoryDifference due to MemoryDifference due to Memory indexes differences in neural activity during the study phase of an experiment for items that subsequently are remembered compared to items that are later forgotten...
- Contingent negative variationContingent negative variationThe contingent negative variation was one of the first event-related potential components to be described. The CNV component was first described by Dr. W. Grey Walter and colleagues in an article published in Nature in 1964...
- Error-related negativityError-related negativityError-related negativity , , is a component of an event-related potential . ERPs are electrical activity in the brain as measured through electroencephalography and time-locked to an external event...
- Lateralized readiness potentialLateralized readiness potentialIn neuroscience, the lateralized readiness potential is an event-related brain potential, or increase in electrical activity at the surface of the brain, that is thought to reflect the preparation of motor activity on a certain side of the body; in other words, it is a spike in the electrical...
- Early left anterior negativityEarly left anterior negativityThe early left anterior negativity is an event-related potential in electroencephalography , or component of brain activity that occurs in response to a certain kind of stimulus...
- P600P600The P600 is an event-related potential , or peak in electrical brain activity measured by electroencephalography . It is a language-relevant ERP and is thought to be elicited by hearing or reading grammatical errors and other syntactic anomalies...