Spine Calcium Transients Induced by Synaptically-Evoked Action Potentials Can Predict Synapse Location and Establish Synaptic Democracy

CA1 pyramidal neurons receive hundreds of synaptic inputs at different distances from the soma. Distance-dependent synaptic scaling enables distal and proximal synapses to influence the somatic membrane equally, a phenomenon called “synaptic democracy”. How this is established is unclear. The backpropagating action potential (BAP) is hypothesised to provide distance-dependent information to synapses, allowing synaptic strengths to scale accordingly. Experimental measurements show that a BAP evoked by current injection at the soma causes calcium currents in the apical shaft whose amplitudes decay with distance from the soma. However, in vivo action potentials are not induced by somatic current injection but by synaptic inputs along the dendrites, which creates a different excitable state of the dendrites. Due to technical limitations, it is not possible to study experimentally whether distance information can also be provided by synaptically-evoked BAPs. Therefore we adapted a realistic morphological and electrophysiological model to measure BAP-induced voltage and calcium signals in spines after Schaffer collateral synapse stimulation. We show that peak calcium concentration is highly correlated with soma-synapse distance under a number of physiologically-realistic suprathreshold stimulation regimes and for a range of dendritic morphologies. Peak calcium levels also predicted the attenuation of the EPSP across the dendritic tree. Furthermore, we show that peak calcium can be used to set up a synaptic democracy in a homeostatic manner, whereby synapses regulate their synaptic strength on the basis of the difference between peak calcium and a uniform target value. We conclude that information derived from synaptically-generated BAPs can indicate synapse location and can subsequently be utilised to implement a synaptic democracy

The effect of noise in models of spiny dendrites

The dendritic tree provides the surface area for synaptic connections between the 100 billion neurons in the brain. 90% of excitatory synapses are made onto dendritic spines which are constantly changing shape and strength. This adaptation is believed to be an important factor in learning, memory and computations within the dendritic tree. The environment in which the neuron sits is inherently noisy due to the activity in nearby neurons and the stochastic nature of synaptic gating. Therefore the effects of noise is a very important aspect in any realistic model. This work provides a comprehensive study of two spiny dendrite models driven by different forms of noise in the spine dynamics or in the membrane voltage. We investigate the effect of the noise on signal propagation along the dendrite and how any correlation in the noise may affect this behaviour. We discover a difference in the results of the two models which suggests that the form of spine connectivity is important. We also show that both models have the capacity to act as a robust filter and that a branched structure can perform logic computations