Spike sorting is a class of techniques used in the analysis of electrophysiological data. Spike sorting algorithms use the shape(s) of waveforms collected with one or more electrodes in the brain to distinguish the activity of one or more neurons from background electrical noise.
Neurons produce action potentials that are referred to as 'spikes' in laboratory jargon. Frequently this term is used for electrical signals recorded in the vicinity of individual neurons with a microelectrode (exception: 'spikes' in EEG recordings). In these recordings action potentials appear as sharp spikes (deviations from the baseline). These extracellular electrodes pick up all the components constituting the field at the point of its contact. This includes the component due to the synaptic currents and the action potentials. The synaptic currents have slower time course and the spikes have faster time course. They are thus easily separated by filtering: highpass for spikes and low pass for the synaptic mechanisms. The synaptic current contributes to the global local field potential (LFP) [1] . Spike sorting refers to the process of assigning spikes to different neurons. The background to this is that the exact time course of a spike event as recorded by the electrode depends on the size and shape of the neuron, the position of the recording electrode relative to the neuron, etc. These electrodes, positioned outside of the cells in the tissue, however, often 'see' the spikes generated by several neurons in their vicinity. Since the spike shapes are unique and quite reproducible for each neuron they can be used to distinguish spikes produced by different neurons, i.e. to separate the activity produced by each.
Technically this is often achieved based on different sizes of the spikes (simple but inaccurate version) or more sophisticated analyses which make use of the entire waveform of the spikes. The techniques often use tools such as principal components or wavelet analysis.
Multiple electrodes record different waveforms for each individual spike elicited by the neurons in the vicinity of the electrodes. The geometric configuration of the electrodes can then be used to define additional dimensions to analyze which spikes originated from which individual cell in the recorded population of cells. Thus the spike sorting using multiple electrodes is better than sorting based simply on waveform shape. The most popular setup involves the use of four micro electrodes, called 'tetrodes' (different from the vacuum tube Tetrode), though more electrodes may be used. Recording electrodes can be metal wires or fine print on a PCB with gold or platinum plated at their exposed tips to ensure good contact and prevent change in resistance while the experiment is going on.
Principal Component weights of spikes from two different neurons
Spike shapes colored according to their assignment to different neurons. The blue trace could not be assigned unequivocally.
An axon or nerve fiber is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action potentials away from the nerve cell body. The function of the axon is to transmit information to different neurons, muscles, and glands. In certain sensory neurons, such as those for touch and warmth, the axons are called afferent nerve fibers and the electrical impulse travels along these from the periphery to the cell body and from the cell body to the spinal cord along another branch of the same axon. Axon dysfunction can be the cause of many inherited and acquired neurological disorders that affect both the peripheral and central neurons. Nerve fibers are classed into three types – group A nerve fibers, group B nerve fibers, and group C nerve fibers. Groups A and B are myelinated, and group C are unmyelinated. These groups include both sensory fibers and motor fibers. Another classification groups only the sensory fibers as Type I, Type II, Type III, and Type IV.
A dendrite or dendron is a branched protoplasmic extension of a nerve cell that propagates the electrochemical stimulation received from other neural cells to the cell body, or soma, of the neuron from which the dendrites project. Electrical stimulation is transmitted onto dendrites by upstream neurons via synapses which are located at various points throughout the dendritic tree.
Within a nervous system, a neuron, neurone, or nerve cell is an electrically excitable cell that fires electric signals called action potentials across a neural network. Neurons communicate with other cells via synapses, which are specialized connections that commonly use minute amounts of chemical neurotransmitters to pass the electric signal from the presynaptic neuron to the target cell through the synaptic gap.
Electrophysiology is the branch of physiology that studies the electrical properties of biological cells and tissues. It involves measurements of voltage changes or electric current or manipulations on a wide variety of scales from single ion channel proteins to whole organs like the heart. In neuroscience, it includes measurements of the electrical activity of neurons, and, in particular, action potential activity. Recordings of large-scale electric signals from the nervous system, such as electroencephalography, may also be referred to as electrophysiological recordings. They are useful for electrodiagnosis and monitoring.
An action potential occurs when the membrane potential of a specific cell rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, called excitable cells, which include neurons, muscle cells, and in some plant cells. Certain endocrine cells such as pancreatic beta cells, and certain cells of the anterior pituitary gland are also excitable cells.
In neuroscience, an excitatory postsynaptic potential (EPSP) is a postsynaptic potential that makes the postsynaptic neuron more likely to fire an action potential. This temporary depolarization of postsynaptic membrane potential, caused by the flow of positively charged ions into the postsynaptic cell, is a result of opening ligand-gated ion channels. These are the opposite of inhibitory postsynaptic potentials (IPSPs), which usually result from the flow of negative ions into the cell or positive ions out of the cell. EPSPs can also result from a decrease in outgoing positive charges, while IPSPs are sometimes caused by an increase in positive charge outflow. The flow of ions that causes an EPSP is an excitatory postsynaptic current (EPSC).
An apical dendrite is a dendrite that emerges from the apex of a pyramidal cell. Apical dendrites are one of two primary categories of dendrites, and they distinguish the pyramidal cells from spiny stellate cells in the cortices. Pyramidal cells are found in the prefrontal cortex, the hippocampus, the entorhinal cortex, the olfactory cortex, and other areas. Dendrite arbors formed by apical dendrites are the means by which synaptic inputs into a cell are integrated. The apical dendrites in these regions contribute significantly to memory, learning, and sensory associations by modulating the excitatory and inhibitory signals received by the pyramidal cells.
Neuronal noise or neural noise refers to the random intrinsic electrical fluctuations within neuronal networks. These fluctuations are not associated with encoding a response to internal or external stimuli and can be from one to two orders of magnitude. Most noise commonly occurs below a voltage-threshold that is needed for an action potential to occur, but sometimes it can be present in the form of an action potential; for example, stochastic oscillations in pacemaker neurons in suprachiasmatic nucleus are partially responsible for the organization of circadian rhythms.
In neuroscience, single-unit recordings provide a method of measuring the electro-physiological responses of a single neuron using a microelectrode system. When a neuron generates an action potential, the signal propagates down the neuron as a current which flows in and out of the cell through excitable membrane regions in the soma and axon. A microelectrode is inserted into the brain, where it can record the rate of change in voltage with respect to time. These microelectrodes must be fine-tipped, impedance matching; they are primarily glass micro-pipettes, metal microelectrodes made of platinum, tungsten, iridium or even iridium oxide. Microelectrodes can be carefully placed close to the cell membrane, allowing the ability to record extracellularly.
Local field potentials (LFP) are transient electrical signals generated in nerves and other tissues by the summed and synchronous electrical activity of the individual cells in that tissue. LFP are "extracellular" signals, meaning that they are generated by transient imbalances in ion concentrations in the spaces outside the cells, that result from cellular electrical activity. LFP are 'local' because they are recorded by an electrode placed nearby the generating cells. As a result of the Inverse-square law, such electrodes can only 'see' potentials in spatially limited radius. They are 'potentials' because they are generated by the voltage that results from charge separation in the extracellular space. They are 'field' because those extracellular charge separations essentially create a local electric field. LFP are typically recorded with a high-impedance microelectrode placed in the midst of the population of cells generating it. They can be recorded, for example, via a microelectrode placed in the brain of a human or animal subject, or in an in vitro brain thin slice.
In neuroscience, a population spike (PS) is the shift in electrical potential as a consequence of the movement of ions involved in the generation and propagation of action potentials. Population spikes often reflect synaptically induced firing and therefore, they can be classified as a type of field excitatory postsynaptic potentials.
In some areas of the brain, such as the hippocampus, neurons are arranged in such a way that they all receive synaptic inputs in the same area. Because these neurons are in the same orientation, the extracellular signals from the generation of action potentials don't cancel out, but rather add up to give a signal that can easily be recorded with a field electrode.
The PS is usually recorded with an extracellular electrode placed close to neural cell bodies or axons. The first interpretations of hippocampal field potentials were developed by Per Andersen.
A tetrode is a type of electrode used in neuroscience for electrophysiological recordings. They are generally used to record the extracellular field potentials from nervous tissue, e.g. the brain. Tetrodes are constructed by bundling together four very small electrodes; each wire is generally less than 30 μm in diameter. Tetrodes are used to classify extra-cellular action potentials into sets generated by the individual neurons, as each channel of the tetrode is usually close enough to a cell such that action potentials emitted by that cell are detected on each of the four channels, but because of the spatial distribution of the individual channels, the amplitude of the signal varies across the four channels. Tetrodes can be built into an implantable device called micro-drive that moves them up and down with precision in the brain of experimental animals
The Mauthner cells are a pair of big and easily identifiable neurons located in the rhombomere 4 of the hindbrain in fish and amphibians that are responsible for a very fast escape reflex. The cells are also notable for their unusual use of both chemical and electrical synapses.
Microelectrode arrays (MEAs) are devices that contain multiple microelectrodes through which neural signals are obtained or delivered, essentially serving as neural interfaces that connect neurons to electronic circuitry. There are two general classes of MEAs: implantable MEAs, used in vivo, and non-implantable MEAs, used in vitro.
In neurophysiology, a dendritic spike refers to an action potential generated in the dendrite of a neuron. Dendrites are branched extensions of a neuron. They receive electrical signals emitted from projecting neurons and transfer these signals to the cell body, or soma. Dendritic signaling has traditionally been viewed as a passive mode of electrical signaling. Unlike its axon counterpart which can generate signals through action potentials, dendrites were believed to only have the ability to propagate electrical signals by physical means: changes in conductance, length, cross sectional area, etc. However, the existence of dendritic spikes was proposed and demonstrated by W. Alden Spencer, Eric Kandel, Rodolfo Llinás and coworkers in the 1960s and a large body of evidence now makes it clear that dendrites are active neuronal structures. Dendrites contain voltage-gated ion channels giving them the ability to generate action potentials. Dendritic spikes have been recorded in numerous types of neurons in the brain and are thought to have great implications in neuronal communication, memory, and learning. They are one of the major factors in long-term potentiation.
Electroencephalography (EEG) is a method to record an electrogram of the spontaneous electrical activity of the brain. The biosignals detected by EEG have been shown to represent the postsynaptic potentials of pyramidal neurons in the neocortex and allocortex. It is typically non-invasive, with the EEG electrodes placed along the scalp using the International 10–20 system, or variations of it. Electrocorticography, involving surgical placement of electrodes, is sometimes called "intracranial EEG". Clinical interpretation of EEG recordings is most often performed by visual inspection of the tracing or quantitative EEG analysis.
Nonsynaptic plasticity is a form of neuroplasticity that involves modification of ion channel function in the axon, dendrites, and cell body that results in specific changes in the integration of excitatory postsynaptic potentials and inhibitory postsynaptic potentials. Nonsynaptic plasticity is a modification of the intrinsic excitability of the neuron. It interacts with synaptic plasticity, but it is considered a separate entity from synaptic plasticity. Intrinsic modification of the electrical properties of neurons plays a role in many aspects of plasticity from homeostatic plasticity to learning and memory itself. Nonsynaptic plasticity affects synaptic integration, subthreshold propagation, spike generation, and other fundamental mechanisms of neurons at the cellular level. These individual neuronal alterations can result in changes in higher brain function, especially learning and memory. However, as an emerging field in neuroscience, much of the knowledge about nonsynaptic plasticity is uncertain and still requires further investigation to better define its role in brain function and behavior.
Ephaptic coupling is a form of communication within the nervous system and is distinct from direct communication systems like electrical synapses and chemical synapses. It may refer to the coupling of adjacent (touching) nerve fibers caused by the exchange of ions between the cells, or it may refer to coupling of nerve fibers as a result of local electric fields. In either case ephaptic coupling can influence the synchronization and timing of action potential firing in neurons. Myelination is thought to inhibit ephaptic interactions.
Electrocochleography is a technique of recording electrical potentials generated in the inner ear and auditory nerve in response to sound stimulation, using an electrode placed in the ear canal or tympanic membrane. The test is performed by an otologist or audiologist with specialized training, and is used for detection of elevated inner ear pressure or for the testing and monitoring of inner ear and auditory nerve function during surgery.
A chronic electrode implant is an electronic device implanted chronically into the brain or other electrically excitable tissue. It may record electrical impulses in the brain or may stimulate neurons with electrical impulses from an external source.