Quantal analysis and synaptic anatomy--integrating two views of hippocampal plasticity

The excitatory synapses onto CA1 pyramidal cells have become a model system for understanding the activity-dependent changes in synapses that underlie learning and memory. Here we examine physiological and anatomical results that are relevant to understanding the mechanisms of synaptic transmission and plasticity at these synapses. Three main points are discussed. First, quantal analysis indicates a large heterogeneity of postsynaptic efficacies for different synapses on the same cell. Reconstructions from electron microscopy show that synapse size is also highly heterogeneous. Reasons for suspecting a relationship between synaptic size and efficacy are discussed. Second, physiological evidence indicates that the changes during long-term potentiation are both pre- and postsynaptic. Similarly, several lines of anatomical evidence suggest that plasticity affects the structure of both the pre- and postsynaptic elements. The detailed registration of structures across the synapse and the physical linkage between pre- and postsynaptic elements suggest a 'structural unit hypothesis' for coordinating pre- and postsynaptic modifications. Third, quantal analysis indicates that stimulation of a single axon can release multiple quanta. Anatomical evidence shows that cell pairs can be connected by multiple synapses, suggesting that multiple quanta may be released at independent sites. These results raise the possibility that one component of synaptic plasticity is mediated by changes in the number of functional synaptic sites.