Phase Precession
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Phase precession is a
neurophysiological Neurophysiology is a branch of physiology and neuroscience that studies nervous system function rather than nervous system architecture. This area aids in the diagnosis and monitoring of neurological diseases. Historically, it has been dominated b ...
process in which the time of firing of
action potential An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s by individual
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
s occurs progressively earlier in relation to the
phase Phase or phases may refer to: Science *State of matter, or phase, one of the distinct forms in which matter can exist *Phase (matter), a region of space throughout which all physical properties are essentially uniform * Phase space, a mathematic ...
of the
local field potential Local field potentials (LFP) are transient electrical signals generated in nervous and other tissues by the summed and synchronous electrical activity of the individual cells (e.g. neurons) in that tissue. LFP are "extracellular" signals, meaning ...
oscillation Oscillation is the repetitive or periodic variation, typically in time, of some measure about a central value (often a point of equilibrium) or between two or more different states. Familiar examples of oscillation include a swinging pendulum ...
with each successive cycle. In place cells, a type of neuron found in the
hippocampal The hippocampus (via Latin from Greek , 'seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, an ...
region of the
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a ve ...
, phase
precession Precession is a change in the orientation of the rotational axis of a rotating body. In an appropriate reference frame it can be defined as a change in the first Euler angle, whereas the third Euler angle defines the rotation itself. In oth ...
is believed to play a major role in the
neural coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activit ...
of information. John O'Keefe, who later shared the 2014
Nobel Prize in Physiology or Medicine The Nobel Prize in Physiology or Medicine is awarded yearly by the Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or medicine. The Nobel Prize is not a single prize, but five separate prizes that, accord ...
for his discovery that place cells help form a "map" of the body's position in space, co-discovered phase precession with Michael Recce in 1993.


Place cells

Pyramidal cells in the hippocampus called place cells play a significant role in self-location during movement over short distances. As a rat moves along a path, individual place cells fire action potentials at an increased rate at particular positions along the path, termed "place fields". Each place cell's maximum firing ratewith action potentials occurring in rapid burstsoccurs at the position encoded by that cell; and that cell fires only occasionally when the animal is at other locations. Within a relatively small path, the same cells are repeatedly activated as the animal returns to the same position. Although simple
rate coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activity ...
(the coding of information based on whether neurons fire more rapidly or more slowly) resulting from these changes in firing rates may account for some of the neural coding of position, there is also a prominent role for the timing of the action potentials of a single place cell, relative to the firing of nearby cells in the local population. As the larger population of cells fire occasionally when the rat is outside of the cells' individual place fields, the firing patterns are organized to occur synchronously, forming wavelike
voltage Voltage, also known as electric pressure, electric tension, or (electric) potential difference, is the difference in electric potential between two points. In a static electric field, it corresponds to the work needed per unit of charge to ...
oscillations. These oscillations are measurable in local field potentials and electroencephalography (EEG). In the CA1 region of the hippocampus, where the place cells are located, these firing patterns give rise to
theta wave Theta waves generate the theta rhythm, a neural oscillation in the brain that underlies various aspects of cognition and behavior, including learning, memory, and spatial navigation in many animals. It can be recorded using various electrophysi ...
s. Theta oscillations have classically been described in rats, but evidence is emerging that they also occur in humans. In 1993, O'Keefe and Recce discovered a relationship between the theta wave and the firing patterns of individual place cells. Although the occasional action potentials of cells when rats were outside of the place fields occurred in phase with (at the peaks of) the theta waves, the bursts of more rapid spikes elicited when the rats reached the place fields were out of synchrony with the oscillation. As a rat approached the place field, the corresponding place cell would fire slightly in advance of the theta wave peak. As the rat moved closer and closer, each successive action potential occurred earlier and earlier within the wave cycle. At the center of the place field, when the cell would fire at its maximal rate, the firing had been advanced sufficiently to be anti-phase to the theta potential (at the bottom, rather than at the peak, of the theta waveform). Then, as the rat continued to move on past the place field and the cell firing slowed, the action potentials continued to occur progressively earlier relative to the theta wave, until they again became synchronous with the wave, aligned now with one wave peak earlier than before. O'Keefe and Recce termed this advancement relative to the wave phase "phase precession". Subsequent studies showed that each time a rat entered a completely different area and the place fields would be remapped, place cells would again become phase-locked to the theta rhythm. It is now widely accepted that the anti-phase cell firing that results from phase precession is an important component of information coding about place.


Other systems

There have been conflicting theories of how neurons in and around the hippocampus give rise to theta waves and consequently give rise to phase precession. As these mechanisms became better understood, the existence of phase precession was increasingly accepted by researchers. This, in turn, gave rise to the question of whether phase precession could be observed in any other regions of the brain, with other kinds of cell circuits, or whether phase precession was a peculiar property of hippocampal tissue. The finding that theta wave phase precession is also a property of grid cells in the
entorhinal cortex The entorhinal cortex (EC) is an area of the brain's allocortex, located in the medial temporal lobe, whose functions include being a widespread network hub for memory, navigation, and the perception of time.Integrating time from experience in the ...
demonstrated that the phenomenon exists in other parts of the brain that also mediate information about movement. Theta wave phase precession in the hippocampus also plays a role in some brain functions that are unrelated to spatial location. When rats were trained to jump up to the rim of a box, place cells displayed phase precession much as they do during movement along a path, but a subset of the place cells showed phase precession that was related to initiating the jump, independently of spatial location, and not related to the position during the jump. Phase precession in the entorhinal cortex has been hypothesized to arise from an attractor network process, so that two sequential neural representations within a single cycle of the theta oscillation can be temporally linked to each other downstream in the hippocampus, as
episodic memories Episodic memory is the memory of everyday events (such as times, location geography, associated emotions, and other contextual information) that can be explicitly stated or conjured. It is the collection of past personal experiences that occurred ...
.


References

{{reflist, 32em Neural coding Neural circuits Hippocampus (brain) Animal locomotion Electrophysiology