Long-term potentiation is associated with new excitatory spine synapses on rat dentate granule cells

To investigate possible morphological correlates to long-term potentiation (LTP), three-dimensional reconstruction of serial electron micrographs was employed. LTP was induced in the perforant path/dentate granule cell synapse in two rats. The surgically isolated contralateral side served as control, along with two untreated animals. Longitudinally sectioned and transversally sectioned dendrites were sampled from the middle fifth of the molecular layer and all visibly connected spines were identified. A mixed, unbalanced, nested variance component model was used to make a valid statistical comparison between the LTP and control groups. The spine density was higher in the experimental than in the control groups. The changes were statistically significant in both the longitudinal and transverse sample. In addition, spines with a divided stem and two heads (bifurcating spines) were seen at a higher frequency in the LTP material compared with the contralateral material. From a subset of dendrites all connected spines were reconstructed and detailed measurements of head, neck, and PSD dimensions were made. We failed to find significant differences following LTP on either of the dimensions measured. The results suggest that new spine synapses are formed following LTP, including some of the bifurcating type.