Computational Models of Timing Mechanisms in the Cerebellar Granular Layer

A long-standing question in neuroscience is how the brain controls movement that requires precisely timed muscle activations. Studies using Pavlovian delay eyeblink conditioning provide good insight into this question. In delay eyeblink conditioning, which is believed to involve the cerebellum, a subject learns an interstimulus interval (ISI) between the onsets of a conditioned stimulus (CS) such as a tone and an unconditioned stimulus such as an airpuff to the eye. After a conditioning phase, the subject’s eyes automatically close or blink when the ISI time has passed after CS onset. This timing information is thought to be represented in some way in the cerebellum. Several computational models of the cerebellum have been proposed to explain the mechanisms of time representation, and they commonly point to the granular layer network. This article will review these computational models and discuss the possible computational power of the cerebellum.

• Publication year

  2009

• Venue

  Cerebellum

• Publication date

  2009-06-05

• Fields of study

  Medicine, Computer Science, Psychology

• Identifiers
  DOI 10.1007/s12311-009-0115-7PMID 19495900PMCID 2788136
• External record

  Open on Semantic Scholar

• Source metadata

  Semantic Scholar, PubMed

• No claims are published for this paper.

• No concepts are published for this paper.

• Timing and plasticity in the cerebellum: focus on the granular layer.

  2009cited by this paper
• Characteristics of Responses of Golgi Cells and Mossy Fibers to Eye Saccades and Saccadic Adaptation Recorded from the Posterior Vermis of the Cerebellum

  2009cited by this paper
• Interactions between prefrontal cortex and cerebellum revealed by trace eyelid conditioning.

  2009cited by this paper
• State-dependent computations: spatiotemporal processing in cortical networks

  2009cited by this paper
• Where is the trace in trace conditioning?

  2008cited by this paper
• Interaction between Purkinje Cells and Inhibitory Interneurons May Create Adjustable Output Waveforms to Generate Timed Cerebellar Output

  2008cited by this paper
• Functions of Interneurons in Mouse Cerebellum

  2008cited by this paper
• The Contribution of Single Synapses to Sensory Representation in Vivo

  2008cited by this paper
• A spiking network model for passage-of-time representation in the cerebellum

  2007cited by this paper
• High-fidelity transmission of sensory information by single cerebellar mossy fibre boutons

  2007cited by this paper
• 2007 Special Issue: The cerebellum as a liquid state machine

  2007cited by this paper
• Generative Modeling of Autonomous Robots and their Environments using Reservoir Computing

  2007cited by this paper
• Special issue on echo state networks and liquid state machines

  2007cited by this paper
• Acquisition, Extinction, and Reacquisition of a Cerebellar Cortical Memory Trace

  2007cited by this paper
• 2007 Special Issue: Automatic speech recognition using a predictive echo state network classifier

  2007cited by this paper
• Ketamine and xylazine depress sensory-evoked parallel fiber and climbing fiber responses.

  2007cited by this paper
• Properties of Somatosensory Synaptic Integration in Cerebellar Granule Cells In Vivo

  2006cited by this paper
• The Random Projection Method

  2005cited by this paper
• Neural Modeling of an Internal Clock

  2005cited by this paper
• Time and tide in cerebellar memory formation.

  2005cited by this paper
• A discrete approach for a model of temporal learning by the cerebellum: in silico classical conditioning of the eyeblink reflex.

  2004cited by this paper
• Harnessing Nonlinearity: Predicting Chaotic Systems and Saving Energy in Wireless Communication

  2004cited by this paper
• The neural representation of time.

  2004cited by this paper
• Model of granular layer encoding in the cerebellum

  2004cited by this paper
• A Biophysical Model of Synaptic Delay Learning and Temporal Pattern Recognition in a Cerebellar Purkinje Cell

  2004cited by this paper
• Integration of quanta in cerebellar granule cells during sensory processing

  2004cited by this paper
• Developmental changes in eyeblink conditioning and neuronal activity in the pontine nuclei.

  2003cited by this paper
• Neural substrates of eyeblink conditioning: acquisition and retention.

  2003cited by this paper
• Trace eyeblink conditioning in decerebrate guinea pigs

  2003cited by this paper
• The Functional Anatomy of Skeletal Conditioning

  2002cited by this paper
• A Neuroscientist's Guide to Classical Conditioning

  2002cited by this paper
• Real-Time Computing Without Stable States: A New Framework for Neural Computation Based on Perturbations

  2002cited by this paper
• Unsupervised learning of granule cell sparse codes enhances cerebellar adaptive control.

  2001cited by this paper
• Beyond parallel fiber LTD: the diversity of synaptic and non-synaptic plasticity in the cerebellum

  2001cited by this paper
• Timing Mechanisms in the Cerebellum: Testing Predictions of a Large-Scale Computer Simulation

  2000cited by this paper
• Short‐lasting conditioned stimulus applied to the middle cerebellar peduncle elicits delayed conditioned eye blink responses in the decerebrate ferret

  1999cited by this paper
• Neural Network Model of the Cerebellum: Temporal Discrimination and the Timing of Motor Responses

  1999cited by this paper
• Classical conditioning

  1998cited by this paper
• Internal models in the cerebellum.

  1998cited by this paper
• Intracellular Correlates of Acquisition and Long-Term Memory of Classical Conditioning in Purkinje Cell Dendrites in Slices of Rabbit Cerebellar Lobule HVI

  1998cited by this paper
• Classical conditioning increases membrane‐bound protein kinase C in rabbit cerebellum

  1998cited by this paper
• Autonomous development of decorrelation filters in neural networks with recurrent inhibition.

  1998cited by this paper
• Pairing‐specific long‐term depression prevented by blockade of PKC or intracellular Ca2+

  1998cited by this paper
• Synchronization of golgi and granule cell firing in a detailed network model of the cerebellar granule cell layer.

  1998cited by this paper
• A model of Pavlovian eyelid conditioning based on the synaptic organization of the cerebellum.

  1997influential reference
• The detection and generation of sequences as a key to cerebellar function: Experiments and theory

  1997cited by this paper
• Metabotropic Glutamate Receptor Activation in Cerebellar Purkinje Cells as Substrate for Adaptive Timing of the Classically Conditioned Eye-Blink Response

  1996cited by this paper
• Frequency dependent activation of a slow N-methyl-D-aspartate-dependent excitatory postsynaptic potential in turtle cerebellum by mossy fibre afferents.

  1995cited by this paper
• A neural model of timed response learning in the cerebellum

  1994cited by this paper
• Signal transmission in the parallel fiber-Purkinje cell system visualized by high-resolution imaging.

  1994influential reference
• Morphology of parallel fibres in the cerebellar cortex of the rat: An experimental light and electron microscopic study with biocytin

  1994influential reference
• Cerebellar cortex: its simulation and the relevance of Marr's theory.

  1992cited by this paper
• A neural network model of the cerebellar cortex performing dynamic associations

  1991cited by this paper
• Adaptively timed conditioned responses and the cerebellum: A neural network approach

  1989cited by this paper
• Adaptive timing in neural networks: The conditioned response

  1988cited by this paper
• Adaptive filter model of the cerebellum

  1982cited by this paper
• Acoustic input to the lateral pontine nuclei.

  1978cited by this paper
• A Theory of Cerebellar Function

  1971cited by this paper
• Neurophysiological aspects of the cerebellar motor control system.

  1970cited by this paper
• A theory of cerebellar cortex

  1969cited by this paper

• Classical conditioning of Purkinje cell responses in vitro produces in vivo-like simple spike suppressions during the conditional stimulus

  2026cites this paper
• The human cerebellum encodes temporally sensitive reinforcement learning signals

  2025cites this paper
• Exploring the Mechanisms of Bidirectional Plasticity From Temporal Feature Stimuli in Cerebellar Motor Learning: A Ca2+-Mediated Simulation

  2025cites this paper
• Neural circuit mechanisms to transform cerebellar population dynamics for motor control in monkeys

  2025cites this paper
• Hippocampus shapes entorhinal cortical output through a direct feedback circuit

  2025cites this paper
• A novel fuzzy cerebellar adaptive neuro intelligent controller for high-performance brushless DC motor drives

  2025cites this paper
• A spiking network model of the cerebellum for predicting movements with diverse complex spikes

  2025cites this paper
• A cerebellar granule cell–climbing fiber computation to learn to track long time intervals

  2024cites this paper
• Cerebellar encoding of prior knowledge of temporal statistics

  2024cites this paper
• Cerebellum-Inspired Model Predictive Control for Redundant Manipulators With Unknown Structure Information

  2024influential citation
• Discrimination and learning of temporal input sequences in a cerebellar Purkinje cell model

  2023cites this paper
• Heterogeneous encoding of temporal stimuli in the cerebellar cortex

  2023cites this paper
• Cognitive-Affective Functions of the Cerebellum

  2023cites this paper
• A computational passage-of-time model of the cerebellar Purkinje cell in eyeblink conditioning

  2023cites this paper
• Position-Based Visual Servo Control of Dual Robotic Arms With Unknown Kinematic Models: A Cerebellum- Inspired Approach

  2023cites this paper
• Electrical coupling regulated by GABAergic nucleo-olivary afferent fibres facilitates cerebellar sensory-motor adaptation

  2022cites this paper
• Quantitative properties of the creation and activation of a cell-intrinsic duration-encoding engram

  2022cites this paper
• Comparing the Representation of a Simple Visual Stimulus across the Cerebellar Network

  2022cites this paper
• Simulation of the effect of the randomness of climbing fiber input on the relationship between motor learning and Focal Task-Specific Dystonia

  2022cites this paper
• Computational Structure of the Cerebellar Molecular Layer

  2021cites this paper
• Computational epidemiology study of homeostatic compensation during sensorimotor aging

  2021cites this paper
• Timing in Purkinje Cells and a Novel Learning Mechanism

  2021cites this paper
• Evolution of the Marr-Albus-Ito Model

  2021cites this paper
• A cerebellar-based solution to the nondeterministic time delay problem in robotic control

  2021cites this paper
• Network model of predictive coding based on reservoir computing for multi-modal processing of visual and auditory signals

  2021cites this paper
• Homeostatic cerebellar compensation of age-related changes of vestibulo-ocular reflex adaptation: a computational epidemiology study

  2020cites this paper
• A computational model of homeostatic cerebellar compensation of ageing in vestibulo-ocular reflex adaptation

  2020cites this paper
• Kir3 channel blockade in the cerebellar cortex suppresses performance of classically conditioned Purkinje cell responses

  2020cites this paper
• A Cerebellar Computational Mechanism for Delay Conditioning at Precise Time Intervals

  2020cites this paper
• Quantitative Properties of the Creation and Activation of a Cell-Intrinsic Engram

  2020cites this paper
• An Internal Clock Based Space-time Neural Network for Motion Speed Recognition

  2020cites this paper
• On Robot Compliance: A Cerebellar Control Approach

  2020cites this paper
• VOR Adaptation on a Humanoid iCub Robot Using a Spiking Cerebellar Model

  2020cites this paper
• Revisiting a theory of cerebellar cortex.

  2019cites this paper
• Modeling the Encoding of Saccade Kinematic Metrics in the Purkinje Cell Layer of the Cerebellar Vermis

  2019cites this paper
• Cerebellum-inspired approach for adaptive kinematic control of soft robots

  2019cites this paper
• New Advances Regarding Adaptation of the Vestibulo-Ocular Reflex.

  2019cites this paper
• Intrinsic memory of temporal intervals in cerebellar Purkinje cells.

  2019cites this paper
• Purkinje Neurons During Eye Blink Conditioning and New Mechanisms of Cerebellar Learning and Timing

  2019cites this paper
• Absence of associative motor learning and impaired time perception in a rare case of complete cerebellar agenesis

  2018cites this paper
• Absence of Parallel Fibre to Purkinje Cell LTD During Eyeblink Conditioning

  2018cites this paper
• Spike burst-pause dynamics of Purkinje cells regulate sensorimotor adaptation

  2018cites this paper
• The Neurobiological Bases for the Computational Theory of Mind

  2017cites this paper
• Learned response sequences in cerebellar Purkinje cells

  2017cites this paper
• A cerebellar mechanism for learning prior distributions of time intervals

  2017cites this paper
• Non-synaptic signaling from cerebellar climbing fibers modulates Golgi cell activity

  2017cites this paper
• Theoretical implications of quantitative properties of interval timing and probability estimation in mouse and rat.

  2017cites this paper
• Real-Time Simulation of Passage-of-Time Encoding in Cerebellum Using a Scalable FPGA-Based System

  2016cites this paper
• Distributed Circuit Plasticity: New Clues for the Cerebellar Mechanisms of Learning

  2016cites this paper
• Single-Neuron and Network Computation in Realistic Models of the Cerebellar Cortex

  2016cites this paper
• Classical Conditioning of Timed Motor Responses

  2016cites this paper
• Timing Rules for Synaptic Plasticity Matched to Behavioral Function.

  2016cites this paper
• Mechanisms for motor timing in the cerebellar cortex.

  2016cites this paper
• A Passage-of-time Model of the Cerebellar Purkinje Cell

  2016cites this paper
• Distributed Cerebellar Motor Learning: A Spike-Timing-Dependent Plasticity Model

  2016influential citation
• Neural networks-on-chip for hybrid bio-electronic systems

  2016cites this paper
• At the Edge of Chaos: How Cerebellar Granular Layer Network Dynamics Can Provide the Basis for Temporal Filters

  2015cites this paper
• Activation of a Temporal Memory in Purkinje Cells by the mGluR7 Receptor.

  2015influential citation
• Digital neural circuits : from ions to networks

  2015cites this paper
• A Purkinje cell Timing Mechanism. On the Physical Basis of a Temporal Duration Memory.

  2015influential citation
• Bio-inspired robotic control schemes using biologically plausible neural structures

  2014cites this paper
• Bidirectional Plasticity of Purkinje Cells Matches Temporal Features of Learning

  2014cites this paper
• Memory trace and timing mechanism localized to cerebellar Purkinje cells

  2014cites this paper
• Theoretical considerations for understanding a Purkinje cell timing mechanism

  2014cites this paper
• Integrated neural and robotic simulations. Simulation of cerebellar neurobiological substrate for an object-oriented dynamic model abstraction process

  2014cites this paper
• Modeling possible effects of atypical cerebellar processing on eyeblink conditioning in autism

  2014cites this paper
• Long-term depression as a model of cerebellar plasticity.

  2014cites this paper
• Fast convergence of learning requires plasticity between inferior olive and deep cerebellar nuclei in a manipulation task: a closed-loop robotic simulation

  2014cites this paper
• Distributed cerebellar plasticity implements adaptable gain control in a manipulation task: a closed-loop robotic simulation

  2013cites this paper
• An integrator circuit in cerebellar cortex

  2013cites this paper
• Classical conditioning of motor responses: What is the learning mechanism?

  2013cites this paper
• ADHD as a Model of Brain-Behavior Relationships

  2013cites this paper
• An adaptive filter model of cerebellar zone C3 as a basis for safe limb control?

  2013cites this paper
• Realtime cerebellum: A large-scale spiking network model of the cerebellum that runs in realtime using a graphics processing unit

  2013cites this paper
• Simulation of biological neuronal structures. Design and functional study of the cerebellum

  2012cites this paper
• [Cerebellar internal clock: its neural mechanisms and functional roles].

  2012cites this paper
• Consensus Paper: Roles of the Cerebellum in Motor Control—The Diversity of Ideas on Cerebellar Involvement in Movement

  2011cites this paper
• Multiple extra‐synaptic spillover mechanisms regulate prolonged activity in cerebellar Golgi cell–granule cell loops

  2011cites this paper
• Learning Stimulus Intervals—Adaptive Timing of Conditioned Purkinje Cell Responses

  2011cites this paper
• Stimulus-Dependent State Transition between Synchronized Oscillation and Randomly Repetitive Burst in a Model Cerebellar Granular Layer

  2011cites this paper
• Time Course of Classically Conditioned Purkinje Cell Response Is Determined by Initial Part of Conditioned Stimulus

  2011influential citation
• The neuronal substrate underlying order and interval representations in sequential tasks: A biologically based robot study

  2010cites this paper
• Adaptive Processes in Speech Perception: Contributions from Cerebral and Cerebellar Cortices

  2009cites this paper
• This Work Is Licensed under a Creative Commons Attribution 3.0 Unported License Simulation of Passage-of-time Encoding in Cerebellum Using a Scalable Fpga-based System. Ieee Transactions on Biomedical Circuits and Real-time Simulation of Passage-of-time Encoding in Cerebellum Using a Scalable Fpga-b

  year unknowncites this paper
• Neuroscience and Biobehavioral Reviews the Contribution of Brain Sub-cortical Loops in the Expression and Acquisition of Action Understanding Abilities

  year unknowncites this paper