The importance of neural aromatization in the acquisition, recall, and integration of song and spatial memories in passerines

In addition to their well-studied and crucial effects on brain development and aging, an increasing number of investigations across vertebrate species indicate that estrogens like 17β-estradiol (E2) have pronounced and rapid effects on cognitive function. The incidence and regulation of the E2-synthesizing enzyme aromatase at the synapse in regions of the brain responsible for learning, memory, social communication and other complex cognitive processes suggest that local E2 production and action affect the acute and chronic activity of individual neurons and circuits. Songbirds in particular are excellent models for the study of this “synaptocrine” hormone provision given that aromatase is abundantly expressed in neuronal soma, dendrites, and at the synapse across many brain regions in both sexes. Additionally, songbirds readily acquire and recall memories in laboratory settings, and their stereotyped behaviors may be manipulated and measured with relative ease. This leads to a rather unparalleled advantage in the use of these animals in studies of the role of neural aromatization in cognition. In this review we describe the results of a number of experiments in songbird species with a focus on the influence of synaptic E2 provision on two cognitive processes: auditory discrimination reliant on the caudomedial nidopallium (NCM), a telencephalic region likely homologous to the auditory cortex in mammals, and spatial memory dependent on the hippocampus. Data from these studies are providing evidence that the local and acute provision of E2 modulates the hormonal, electrical, and cognitive outputs of the vertebrate brain and aids in memory acquisition, retention, and perhaps the confluence of memory systems.