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Emma Wilkinson sandbox

The Olfactory Bulb

Layers of Olfactory Bulb
The glomeruli layer of the olfactory bulb is the first level of synaptic processing. The glomeruli layer represents a spatial odor map organized by chemical structure of odorants like functional group and carbon chain length. This spatial map is divided into zones and clusters, which represent similar glomeruli and therefore similar odors. One cluster in particular is associated with rank, spoiled smells which are represented by certain chemical characteristics. This classification may be evolutionary to help identify food that is no longer good to eat. The spatial map of the glomeruli layer may be used for perception of odor in the olfactory cortex.

The next level of synaptic processing in the olfactory bulb occurs in the external plexiform layer. The external plexiform layer contains astrocytes, interneurons and some mitral cells. It does not contain many cell bodies, rather mostly dendrites of mitral cells and GABAergic granule cells. Interneurons in the external plexiform layer are responsive to pre-synaptic action potentials and exhibit both excitatory postsynaptic potentials and inhibitory postsynaptic potentials. Neural firing varies temporally, there are periods of fast, spontaneous firing and slow modulation of firing. These patterns may be related to sniffing or change in intensity and concentration of odorant. Temporal patterns may have effect in later processing of spatial awareness of odorant.

Lateral Inhibition of Mitral Cells in the External Plexiform Layer
The interneurons in the external plexiform layer perform feedback inhibition on the mitral cells to control back propagation. They also participate in lateral inhibition of the mitral cells. This inhibition is an important part of olfaction as it aids in odor discrimination by decreasing firing in response to background odors and differentiating the responses of olfactory nerve inputs in the mitral cell layer. . Inhibition of the mitral cell layer by the other layers contributes to odor discrimination and higher level processing by modulating the output from the olfactory bulb. These hyperpolarizations during odor stimulation shape the responses of the mitral cells to make them more specific to an odor. One layer deeper is the internal plexiform layer.

Lateral Inhibition of Mitral Cells in the Granule Cell Layer
The granule cell layer is the deepest layer in the olfactory bulb. It is made up of dendrodendritic granule cells that synapse to the mitral cell layer. The granule cell layer receives excitatory glutamate signals from the basal dendrites of the mitral and tufted cells. The granule cell in turn releases GABA to cause an inhibitory effect on the mitral cell. More neurotransmitter is released from the activated mitral cell to the connected dendrite of the granule cell, making the inhibitory effect from the granule cell to the activated mitral cell stronger than the surrounding mitral cells. These differentiated odor responses are later processed and perceived as distinct odors. . There is also evidence of cholinergic effects on granule cells that enhance depolarization of granule cells making them more excitable which in turn increases inhibition of mitral cells. This may contribute to a more specific output from the olfactory bulb that would closer resemble the glomerular odor map.

Further Olfactory Processing
The olfactory bulb sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex(OFC) and the hippocampus where it plays a role in emotion, memory and learning. The main olfactory bulb connects to the amygdala via the piriform cortex of the primary olfactory cortex and direct projections from the main olfactory bulb to specific amygdala areas. The amygdala passes olfactory information on to the hippocampus. The orbitofrontal cortex, amygdala, hippocampus, thalamus, and olfactory bulb have many interconnections directly and indirectly through the cortices of the primary olfactory cortex. These connections are indicative of the association between the olfactory bulb and higher areas of processing, specifically those related to emotion and memory. .

Amygdala
Associative learning takes place in the amygdala takes place between odors and behavioral responses. The odors serve as the reinforcers or the punishers during the associative learning process; odors that occur with positive states reinforce the behavior that resulted in the positive state while odors that occur with negative states do the opposite. Odor cues are coded by neurons in the amygdala with the behavioral effect or emotion that they produce. In this way odors reflect certain emotions or physiological states. Odors become associated with pleasant and unpleasant responses, and eventually the odor becomes a cue and can cause an emotional response. These odor associations contribute to emotional states such as fear. Brain imaging shows amygdala activation correlated with pleasant and unpleasant odors, reflecting the association between odors and emotions. .

Hippocampus
The hippocampus aids in olfactory memory and learning as well. Several olfaction-memory processes occur in the hippocampus. Similar to the process in the amygdala, an odor is associated with a particular reward, i.e. the smell of food with receiving sustenance. Odor in the hippocampus also contributes to the formation of episodic memory; the memories of events at a specific place or time. The time at which certain neurons fire in the hippocampus is associated by neurons with a stimulus such as an odor. Presentation of the odor at a different time may cause recall of the memory, therefore odor aids in recall of episodic memories. .

Olfactory Bulb and Depression Models
Further evidence of the link between the olfactory bulb and emotion and memory is shown through animal depression models. Olfactory bulb removal in rats effectively causes structural changes in the amygdala and hippocampus and behavioral changes similar to that of a person with depression. Researchers use rats with olfactory bulbectomies to research antidepressants. Research has shown that removal of the olfactory bulb in rats leads to dendrite reorganization, disrupted cell growth in the hippocampus, and decreased neuroplasticity in the hippocampus. These hippocampal changes due to olfactory bulb removal are associated with behavioral changes characteristic of depression, demonstrating the correlation between the olfactory bulb and emotion. The hippocampus and amygdala effect odor perception. During certain physiological states such as hunger a food odor may seem more pleasant and rewarding due to the associations in the amygdala and hippocampus of the food odor stimulus with the reward of eating. .

The Orbitofrontal Cortex
Olfactory information is sent to the primary olfactory cortex, where projections are sent to the orbitofrontal cortex. The OFC contributes to this odor-reward association as well as it assesses the value of a reward, i.e. the nutritional value of a food. The OFC receives projections from the piriform cortex, amygdala, and parahippocampal cortices. Neurons in the OFC that encode food reward information activate the reward system when stimulated, associating the act of eating with reward. The OFC further projects to the anterior cingulate cortex where it plays a role in appetite. The OFC also associates odors with other stimuli, such as taste. . Odor perception and discrimination also involve the OFC. The spatial odor map in the glomeruli layer of the olfactory bulb may contribute to these functions. The odor map begins processing of olfactory information by spatially organizing the glomeruli. This organizing aids the olfactory cortex in it's functions of perceiving and discriminating odors. .