The olfactory bulb Mitral cells are the principal neurons in the olfactory bulb.

. There are about 45,000 of these cells in each bulb in the rat and around 50,000 in the adult human. They have a primary apical dendrite which extends into a spherical bundle of neuropil called a glomerulus (see below) which receives the input from the olfactory receptor neurons. Their axons merge together to form the lateral olfactory tract. They possess colaterals, involved in negative feedback and positive feed-forward. Glomeruli are roughly spherical bundles of dendritic processes - some 25 mitral cells may send their primary dendrites to a single glomerulus - and it is here that they make contact with incoming olfactory nerves (in rodents the branches of 17,000-25,000 olfactory axons). In the rabbit there are about 2000 glomeruli per olfactory bulb. Periglomerular cells are involved in lateral inhibition at the level of the glomeruli Granule cells are inhibitory interneurones. They receive both contra- and ipsilateral input. The lateral olfactory tract terminates in the pyriform and prepyriform areas (primary olfactory cortex) from where the primary projection goes to the thalamus (medialis dorsalis). Axons project from here to the neocortex (orbito-frontal). In addition, primates have a pathway that runs via the limbic brain to the hypothalamus and is involved with mood (and memory) and neuroendocrine regulation. This latter pathway is responsible for the so-called "affective" component of smell. Centrifugal pathways have a "wipe clean" function to reset the system ready for the next input and also with dis-inhibition. When hungry smells have a greater effect! The architecture of the bulb results in 1:1000 convergence of olfactory receptor neurons to mitral cells. Thus a lot of information about individual receptors is thrown away but this increases sensitivity since contributions from many receptors are added together.

Central olfactory pathways Neurons from the lateral olfactory tract project to; (1) the amygdala, septal nuclei, prepyriform cortex, the entorhinal cortex, hippocampus and the subiculum. Many of these structures form the limbic system, an ancient region of the brain concerned with motivation, emotion and certain kinds of memory. The septal nuclei and amygdala contain regions known as the "pleasure centres". The hippocampus is concerned with motivational memory (the association of certain stimuli with food). (2) Projections are also sent to the thalamus and thence to the frontal cortex for recognition. There are many forward and backward connections between each of these brain centres.

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pt un desn--------->a | Olfactory sensory neurons that express the same odorant receptor project to distinct glomeruli in the main olfactory bulb (MOB). b | Vomeronasal sensory neurons that express the same V1R or V2R project to multiple, small glomeruli in the accessory olfactory bulb (AOB). The apical layer of the epithelium projects to the rostral half of the AOB, and the basal layer projects to the posterior half. c | The four zones in the main olfactory epithelium (MOE) correspond roughly to four domains in the MOB. The boundaries between the domains have not been delineated precisely because not enough odorant receptors have been mapped onto the MOB.