The body tracks short intervals of time using neural oscillator networks
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Scientific literature demonstrates that recurrent neural networks and oscillatory models can successfully simulate interval timing and temporal processing in the brain.
Theta and gamma frequency oscillations occur in the same brain regions and interact with each other, a process called cross-frequency coupling. Here, we review evidence for the following hypothesis: that the dual oscillations form a code for representing multiple items in an ordered way. This form of coding has been most clearly demonstrated in the hippocampus, where different spatial information is represented in different gamma subcycles of a theta cycle. Other experiments have tested the functional importance of oscillations and their coupling. These involve correlation of oscillatory properties with memory states, correlation with memory performance, and effects of disrupting oscillations on memory. Recent work suggests that this coding scheme coordinates communication between brain regions and is involved in sensory as well as memory processes.
Evidence has amassed from both animal intracranial recordings and human electrophysiology that neural oscillatory mechanisms play a critical role in a number of cognitive functions such as learning, memory, feature binding and sensory gating. The wide availability of high-density electrical and magnetic recordings (64-256 channels) over the past two decades has allowed for renewed efforts in the characterization and localization of these rhythms. A variety of cognitive effects that are associated with specific brain oscillations have been reported, which range in spectral, temporal, and spatial characteristics depending on the context. Our laboratory has focused on investigating the role of alpha-band oscillatory activity (8-14 Hz) as a potential attentional suppression mechanism, and this particular oscillatory attention mechanism will be the focus of the current review. We discuss findings in the context of intersensory selective attention as well as intrasensory spatial and feature-based attention in the visual, auditory, and tactile domains. The weight of evidence suggests that alpha-band oscillations can be actively invoked within cortical regions across multiple sensory systems, particularly when these regions are involved in processing irrelevant or distracting information. That is, a central role for alpha seems to be as an attentional suppression mechanism when objects or features need to be specifically ignored or selected against.
Covert spatial attention is essential for humans’ ability to direct limited processing resources to the relevant aspects of visual scenes. A growing body of evidence suggests that rhythmic neural activity in the alpha frequency band (8–12 Hz) tracks the spatial locus of covert attention, which suggests that alpha activity is integral to spatial attention. However, extant work has not provided a compelling test of another key prediction: that alpha activity tracks the temporal dynamics of covert spatial orienting. In the current study, we examined the time course of spatially specific alpha activity after central cues and during visual search. Critically, the time course of this activity tracked trial-by-trial variations in orienting latency during visual search. These findings provide important new evidence for the link between rhythmic brain activity and covert spatial attention, and they highlight a powerful approach for tracking the spatial and temporal dynamics of this core cognitive process.
Extensive research has described two key features of interval timing. The bias property is associated with accuracy and implies that time is overestimated for short intervals and underestimated for long intervals. The scalar property is linked to precision and states that the variability of interval estimates increases as a function of interval duration. The neural mechanisms behind these properties are not well understood. Here we implemented a recurrent neural network that mimics a cortical ensemble and includes cells that show paired-pulse facilitation and slow inhibitory synaptic currents. The network produces interval selective responses and reproduces both bias and scalar properties when a Bayesian decoder reads its activity. Notably, the interval-selectivity, timing accuracy, and precision of the network showed complex changes as a function of the decay time constants of the modeled synaptic properties and the level of background activity of the cells. These findings suggest that physiological values of the time constants for paired-pulse facilitation and GABAb, as well as the internal state of the network, determine the bias and scalar properties of interval timing. SIGNIFICANCE STATEMENT Timing is a fundamental element of complex behavior, including music and language. Temporal processing in a wide variety of contexts shows two primary features: time estimates exhibit a shift toward the mean (the bias property) and are more variable for longer intervals (the scalar property). We implemented a recurrent neural network that includes long-lasting synaptic currents, which cannot only produce interval-selective responses but also follow the bias and scalar properties. Interestingly, only physiological values of the time constants for paired-pulse facilitation and GABAb, as well as intermediate background activity within the network can reproduce the two key features of interval timing.
Time is essential for understanding the brain. A temporal theory for realizing major brain functions (e.g., sensation, cognition, motivation, attention, memory, learning, and motor action) is proposed that uses temporal codes, time-domain neural networks, correlation-based binding processes and signal dynamics. It adopts a signal-centric perspective in which neural assemblies produce circulating and propagating characteristic temporally patterned signals for each attribute (feature). Temporal precision is essential for temporal coding and processing. The characteristic spike patterns that constitute the signals enable general-purpose, multimodal, multidimensional vectorial representations of objects, events, situations, and procedures. Signals are broadcast and interact with each other in spreading activation time-delay networks to mutually reinforce, compete, and create new composite patterns. Sequences of events are directly encoded in the relative timings of event onsets. New temporal patterns are created through nonlinear multiplicative and thresholding signal interactions, such as mixing operations found in radio communications systems and wave interference patterns. The newly created patterns then become markers for bindings of specific combinations of signals and attributes (e.g., perceptual symbols, semantic pointers, and tags for cognitive nodes). Correlation operations enable both bottom-up productions of new composite signals and top-down recovery of constituent signals. Memory operates using the same principles: nonlocal, distributed, temporally coded memory traces, signal interactions and amplifications, and content-addressable access and retrieval. A short-term temporary store is based on circulating temporal spike patterns in reverberatory, spike-timing-facilitated circuits. A long-term store is based on synaptic modifications and neural resonances that select specific delay-paths to produce temporally patterned signals. Holographic principles of nonlocal representation, storage, and retrieval can be applied to temporal patterns as well as spatial patterns. These can automatically generate pattern recognition (wavefront reconstruction) capabilities, ranging from objects to concepts, for distributed associative memory applications. The evolution of proposed neural implementations of holograph-like signal processing and associative content-addressable memory mechanisms is discussed. These can be based on temporal correlations, convolutions, simple linear and nonlinear operations, wave interference patterns, and oscillatory interactions. The proposed mechanisms preserve high resolution temporal, phase, and amplitude information. These are essential for establishing high phase coherency and determining phase relationships, for binding/coupling, synchronization, and other operations. Interacting waves can sum constructively for amplification, or destructively, for suppression, or partially. Temporal precision, phase-locking, phase-dependent coding, phase-coherence, synchrony are discussed within the context of wave interference patterns and oscillatory interactions. Sequences of mixed neural oscillations are compared with a cascade of sequential mixing stages in a single-sideband carrier suppressed (SSBCS) radio communications system model. This mechanism suggests a manner by which multiple neural oscillation bands could interact to produce new emergent information-bearing oscillation bands, as well as to abolish previously generated bands. A hypothetical example illustrates how a succession of different oscillation carriers (gamma, beta, alpha, theta, and delta) could communicate and propagate (broadcast) information sequentially through a neural hierarchy of speech and language processing stages. Based on standard signal mixing principles, each stage emergently generates the next. The sequence of oscillatory bands generated in the mixing cascade model is consistent with neurophysiological observations. This sequenc
A crucial mechanism for the brain to make sense of the auditory environment is the synchronization of neural responses to external temporal regularities, such as a musical beat. It is debated whether this synchronization and the resulting beat percept reflect phase alignment of endogenous neural oscillations to the external regularity ("entrainment"), or evoked responses to the rhythmic stimulus ("tracking"). Here, we use the tempo-dependent properties of beat processing to differentiate between the two accounts. Participants listened to a repeating rhythmic pattern at different speeds. Behaviorally, they consistently tapped at the preferred beat rate (around 2 Hz) across tempi, shifting to higher metrical levels as tempo increased. We found a similar shift in EEG data, where the metrical level at which neural synchronization was strongest depended on tempo. This selective enhancement is consistent with entrainment accounts and could indeed be mimicked by an oscillator model. However, importantly, the results were also captured by a model simulating evoked responses. Together, our findings demonstrate that while neural responses to rhythm are selectively enhanced at the beat rate, this enhancement need not be taken as evidence for entrainment, but can also be explained by successive evoked responses.
Ermentrout B (1994). "An introduction to neural oscillators". In F Ventriglia (ed.). Neural Modeling and Neural Networks. pp. 79–110. Breakspear M, Heitmann
Neural oscillations, or brainwaves, are rhythmic or repetitive patterns of neural activity in the central nervous system. Neural tissue can generate oscillatory activity in many ways, driven either by mechanisms within individual neurons or by interactions between neurons. In individual neurons, oscillations can appear either as oscillations in membrane potential or as rhythmic patterns of action
Oscillations can often be described and analyzed using mathematics. Mathematicians have identified several dynamical mechanisms that generate rhythmicity. Among the most important are harmonic (linear) oscillators, limit cycle oscillators, and delayed-feedback oscillators. Harmonic oscillations appear very frequently in nature—examples are sound waves, the motion of a pendulum, and vibrations of every sort. They generally arise when a physical system is perturbed by a small degree from a minimum-energy state, and are well understood mathematically.
Noise-driven harmonic oscillators realistically simulate alpha rhythm in the waking EEG as well as slow waves and spindles in the sleep EEG. Successful EEG analysis algorithms were based on such models. Several other EEG components are better described by limit-cycle or delayed-feedback oscillations.
Limit-cycle oscillations arise from physical systems that show large deviations from equilibrium, whereas delayed-feedback oscillations arise when components of a system affect each other after significant time delays. Limit-cycle oscillations can be complex but there are powerful mathematical tools for analyzing them; the mathematics of delayed-feedback oscillations is primitive in comparison. Linear oscillators and limit-cycle oscillators qualitatively differ in terms of how they respond to fluctuations in input. In a linear oscillator, the frequency is more or less constant but the amplitude can vary greatly. In a limit-cycle oscillator, the amplitude tends to be more or less constant but the frequency can vary greatly. A heartbeat is an example of a limit-cycle oscillation in that the frequency of beats varies widely, while each individual beat continues to pump about the same amount of blood.
Computational models adopt a variety of abstractions in order to describe complex oscillatory dynamics observed in brain activity. Many models are used in the field, each defined at a different level of abstraction and trying to model different aspects of neural systems. They range from models of the short-term behaviour of individual neurons, through models of how the dynamics of neural circuitry arise from interactions…
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