Multimorphic spines and complex postsynaptic structures in the rat and human brains: A common finding with intriguing morphology and open functional questions

Dendritic spines are postsynaptic specializations that mainly contact excitatory inputs and modulate a wide range of processes involving synaptic transmission and plasticity. Based on morphological features, they were classified into stubby, wide, thin, mushroom, ramified, and double spines. However, spines display other than these "classical" shapes, which are morphologically more convoluted and were initially called "atypical" spines. They have been much less studied and, then, worthy of investigation. Here, atypical (or, rather, multimorphic) spines, as well as complex dendritic protrusions, were examined using the Golgi method and after 2D and 3D image reconstructions in dendrites, cell bodies, and axon hillocks of several neuron types from both rats and humans. A variety of morphological features of complex dendritic protrusions and multimorphic spines were characterized in basket cells, Purkinje cells, brush neurons and granule cells of the cerebellum, in multipolar neurons of the inferior olivary nucleus, in multipolar neurons of the posterodorsal medial amygdaloid nucleus, in short-shaft pyramidal neurons of the hippocampus, in layers V-VI pyramidal and polymorphic neurons of the prefrontal cortex, and layers II-VI neurons of the anterior cingulate, precuneus, temporal, and occipital cortex. We provide evidence for the usual occurrence of these multimorphic spines, characterized by their heterogeneity in shape and size, and discuss the likely functional implications for synaptic processing, intraspine microdomains, compartmentalization features, and plasticity. Given their presence in rodents and humans, we also discuss the possible implications of multimorphic spines for more complex synaptic transmission across evolved neural circuits, laying the groundwork for future research.