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Table of contents
Preface
Chapter I – Introduction
I) Neurons as individual, electrically excitable, nerve cells, communicating through synapses
Historical overview on the mode of communication between neurons
II) Electrical synaptic communication
Historical experiment revealing electrical coupling
Slow acceptance of electrical coupling by the scientific community
Electrical synapses as the molecular basis of electrical coupling
Definition of electrical synapses
Molecular composition of electrical synapses, and plasticity
Molecular composition of gap junctions
Plasticity of electrical synapses
Structural and functional differences between electrical and chemical synapses
Physiological roles of gap junctions and electrical synapses
Physiological roles of gap junctions
Physiological role of electrical synapses
III) Dendritic and synaptic integration
Passive properties and cable filtering
Active conductances
Non-linear dendritic integration
Active conductances can counteract electrical filtering
Active conductances in non-linear dendritic integration
Role of neuronal morphology in non-linear dendritic integration
Integration of excitatory and inhibitory inputs
Subcellular compartment targeting, and input/output relationship
Temporal integration of excitatory and inhibitory inputs: example of feed-forward inhibition
Electrical synapses in the context of dendritic integration
IV) The cerebellum as a canonical microcircuit
The cerebellar cortex
Cell-type diversity and cytoarchitecture
Differences between stellate cells and basket cells
Synaptic plasticity in the cerebellar cortex
Cerebellum-like structures
Role of the cerebellum in physiology
Early models of pattern separation in the cerebellar cortex
Eye-blink conditioning – a classical example of cerebellar temporal learning
Implication of the cerebellum in sensory motor control
Implication of the cerebellum in non-motor tasks
Proposed unifying theories of the role of cerebellum and cerebellar-like structures
Temporal specific learning
Multisensory integration
Multisensory integration and temporal specific learning in cerebellum-like structures
Long- and short-term plasticity as bases for temporal learning
Inhibitory interneurons and time processing
Chapter II – Materials and Methods
Electrophysiology
Transmitted light and fluorescence imaging
Image analysis
Pharmacological agents
Parallel fibre-mediated responses
Detecting electrical and/or chemical synapses in paired recordings
Data analysis and statistics
Chapter III: Electrical synapses within a feed-forward electrical circuit generate temporal contrast enhancement
I) Abstract
II) Introduction
III) Results
PF-evoked and direct recruitment of MLI’s spikelets
Modulation of spikelet polarity by presynaptic membrane potential
Electrical synapses form the majority of inhibitory connections between BCs in adult animals
Feed-forward recruitment of spikelets narrows single EPSP time-window, and dampens temporal summation
Spikelet signalling enables temporal contrast enhancement of temporally coded excitation
IV) Discussion
Electrical connectivity of cerebellar cortex is tuned for rapid output synchrony
Electrical synapses reinforce coincidence detection in electrically connected interneurons
Implications of eFFM in fine-tuning cerebellar-dependent motor behaviours
V) Supplementary figures
Chapter IV – Integration and modulation of electrical synapses-mediated inputs in cerebellar basket cells
I) Evidence for transmission of subthreshold EPSCs across electrical synapses
Discussion
II) Role of HCN channels in shaping AP waveform, and transmitted spikelets
Discussion
III) Frequency-dependent temporal summation of spikelets
Discussion
IV) Conclusion
Chapter V – Evidence for differential dendritic integration properties between SCs and BCs
I) Differences in dendritic morphology between stellate and basket cells
BCs have larger dendrites than SCs
Differences of cable filtering influence between theoretical predications and in vivo data
II) Impact of cable filtering on PF-mediated EPSCs in BCs and SCs
Synaptic currents and potentials are unevenly affected by electrical filtering
Indirect evidence of the passive role of electrical synapses in sharpening EPSP kinetics
III) Discussion
Differential dendritic integration behaviour of EPSCs in MLIs
Passive effect of electrical synapses
Chapter VI – Discussion, and perspectives
I) Electrical connectivity and spikelet transmission in BCs
Spikelets can be used to retrieve the resting membrane potential of unperturbed, electrically-connected cells
Technical considerations on the new method
Electrical synapses are more frequent than chemical synapses in adult mice
Spikelet transmission displays characteristic features of FFI
Narrowing EPSP time window
Dampening temporal summation
Frequency-dependent inhibitory action of spikelets
Spikelets mediate temporal contrast enhancement and coincidence detection
Transient excitation followed by long-lasting inhibition
Comparison with previous studies of spikelet transmission in cerebellar basket cells
Implication for cerebellar processing
II) Addressing the dendritic integration properties of cerebellar basket cells
III) Large scale implications
Implication for cerebellar computation
Conclusion
References
Appendices
Evidence for indirect coupling in paired recordings
Simultaneous recruitment of EPSP and spikelets narrows half-width of EPSPs
Impact of stimulation intensity of PFs on the temporal summation of EPSPs
Testing the hypothesis that HCN channels can shape transmitted spikelets: inconclusive but insightful experiments
Testing the hypothesis that spikelets influence EPSPs’ kinetics – inconclusive but insightful experiments
Supplementary methods
Retrieving the resting membrane potential of electrically connected cells
