Overview
Neural oscillations are observed throughout the central nervous system and at all levels, e.g., spike trains, local field potentials and large-scale oscillations which can be measured by electroencephalography. In general, oscillations can be characterized by their frequency, amplitude and phase. These signal properties can be extracted from neural recordings using time-frequency analysis. In large-scale oscillations, amplitude changes are considered to result from changes in synchronization within a neural ensemble, also referred to as local synchronization. In addition to local synchronization, oscillatory activity of distant neural structures (single neurons or neural ensembles) can synchronize. Neural oscillations and synchronization have been linked to many cognitive functions such as information transfer, perception, motor control and memory.
Neural oscillations have been most widely studied in neural activity generated by large groups of neurons. Large-scale activity can be measured by techniques such as electroencephalography (EEG). In general, EEG signals have a broad spectral content similar to pink noise, but also reveal oscillatory activity in specific frequency bands. The first discovered and best-known frequency band is alpha activity (8–12 Hz) that can be detected from the occipital lobe during relaxed wakefulness and increases when the eyes are closed. Other frequency bands are: delta (1–4 Hz), theta (4–8 Hz), beta (13–30 Hz) and gamma (30–70 Hz) frequency band, where faster rhythms such as gamma activity have been linked to cognitive processing. Indeed, EEG signals change dramatically during sleep and show a transition from faster frequencies to increasingly slower frequencies such as alpha waves. In fact, different sleep stages are commonly characterized by their spectral content. Consequently, neural oscillations have been linked to cognitive states, such as awareness and consciousness.
Although neural oscillations in human brain activity are mostly investigated using EEG recordings, they are also observed using more invasive recording techniques such as single-unit recordings. Neurons can generate rhythmic patterns of action potentials or spikes. Some types of neurons have the tendency to fire at particular frequencies, so-called resonators. Bursting is another form of rhythmic spiking. Spiking patterns are considered fundamental for information coding in the brain. Oscillatory activity can also be observed in the form of subthreshold membrane potential oscillations (i.e. in the absence of action potentials). If numerous neuron spike in synchrony, they can give rise to oscillations in local field potentials (LFPs). Quantitative models can estimate the strength of neural oscillation in recorded data.
Neural oscillations are commonly studied from a mathematical framework and belong to the field of “neurodynamics”, an area of research in the cognitive sciences that places a strong focus upon the dynamic character of neural activity in describing brain function. It considers the brain a dynamical system and uses differential equations to describe how neural activity evolves over time. In particular, it aims to relate dynamic patterns of brain activity to cognitive functions such as perception and memory. In very abstract form, neural oscillations can be analyzed analytically. When studied in a more physiologically realistic setting, oscillatory activity is generally studied using computer simulations of a computational model.
The functions of neural oscillations are wide ranging and vary for different types of oscillatory activity. Examples are the generation of rhythmic activity such as a heartbeat and the neural binding of sensory features in perception, such as the shape and color of an object. Neural oscillations also play an important role in many neurological disorders, such as excessive synchronization during seizure activity in epilepsy or tremor in patients with Parkinson's disease. Oscillatory activity can also be used to control external devises in brain-computer interfaces, in which subjects can control an external device by changing the amplitude of particular brain rhythmics.
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