The recognition memory of imprinting: biochemistry and electrophysiology

A restricted part of the intermediate and medial part of the hyperstriatum ventrale (IMHV) of the domestic chick forebrain is pivotal to the learning process of imprinting and is probably the site at which information about an imprinting stimulus is stored. A range of learning-related changes occur in the IMHV between 1 and 24 h after training. The earliest change described is in Fos-like immunoreactivity. There follow changes in phosphorylation of the protein kinase C substrate MARCKS, morphological changes in axospinous synapses, an increase in NMDA receptor number and increases in amounts of the major isoforms of the neural cell adhesion molecule and clathrin heavy chain. All but the change in Fos-immunopositivity occurs in the left, but not the right, IMHV. Insufficient nitric oxide synthase is available in the IMHV to support the hypothesis that nitric oxide is a retrograde messenger contributing to the effect on Fos-like immunoreactivity. In chicks anaesthetised approximately 24 h after imprinting training, the spontaneous mean neuronal firing rate is related to a preference score (a measure of learning). In unanaesthetised chicks 24 h after training, the responsiveness of some IMHV neurons is biassed specifically towards the imprinting stimulus.The responses of other neurons in the IMHV generalise across some features of the training stimulus, such as form or colour. Some neurons in the IMHV of unanaesthetised chicks are responsive to the distance of an imprinting stimulus from the chick; distance-sensitive neurons can be distinguished from distance-insensitive neurones by the action potential shape.