Olfactory sensory organs
- Olfactory glands (Bowman glands) secrete a fluid that bathes the cilia of the receptors and acts as a solvent for odorant molecules.
Olfactory tracts
- First-order sensory neurons of the olfactory system
- Reside in the nasal cavity.
- Olfactory receptor cells (first-order neurons) are stimulated by the binding of odor molecules to their cilia
- G protein activation and activation of adenylyl cyclase → a rise in intracellular cAMP → causes opening of a cyclic-nucleotide gated ion channel → influx of Na+ and Ca2 + causing neuronal depolarization.
- The axons of the olfactory receptor cells form CN I (olfactory nerve); these project through the cribriform plate at the base of the cranium to synapse with the mitral cells of the olfactory bulb in olfactory glomeruli.
- The map of glomerular activation patterns within the olfactory bulb are thought to represent the quality of the odor being detected.
- Second order neurons (mitral cells)
- The mitral cells of the olfactory bulb are excitatory
- The output axons of the mitral cells form the olfactory tract + lateral olfactory stria, both of which project to the primary olfactory cortex (prefrontal cortex) and the amygdala.
- Because the olfactory bulb is composed of secondary, rather than primary, sensory neurons, it is, strictly speaking, a central nervous system (CNS) tract and not a cranial nerve.
Primary Olfactory Cortex
- Cortical brain regions which receive the mitral and tufted cell axon projections.
- Consist of
- Anterior olfactory nucleus
- A predominantly two-layered cortical-like structure with connections to the olfactory tract.
- Has an
- Anterior segment located posterior to the olfactory bulb
- Posterior segment within the anterior temporal lobe.
- Function
- Reciprocal transfer of information from the bulb to the piriform cortex
- Relay of information between (via the anterior commissure)
- Left and right olfactory bulbs,
- Left and right piriform cortices
- Piriform cortex
- Large three-layered cortex
- Pear-like shape
- Has
- Frontal (‘prepiriform’) components
- Posterior (temporal) components
- Function
- Reciprocally connects with a number of brain regions involved with behaviour and emotion and is critical for processing basic olfactory information.
- Learning and memory of odors
- Encodes representations of odor quality, identity, familiarity, and hedonics (plesure/displeasure).
- Multisensory integration
- For example, piriform activity becomes elicited by visual stimuli after they are paired with a pleasant food odor.
- When food associated with the odor is eaten to satiety, this response is attenuated.
- Red is temporal piriformis cortex;
- Blue is frontal piriformis cortex
- Regions of the amygdala and periamygdaloid complex
- Amygdala is positioned deep within the medial temporal lobe.
- It is closely and reciprocally related to the hypothalamus, and is intimately associated with sympathetic nervous system activity and emotion, such as fear.
- Its connections with the ventral tegmental area, locus coeruleus, and laterodorsal tegmental nucleus influence the release of such neurotransmitters as dopamine, norepinephrine, and epinephrine.
- Recent neuroimaging studies suggest that the amygdala responds to the intensity of emotionally significant, that is, pleasant or unpleasant, odors.
- Rostral entorhinal cortex
- The most caudal temporal lobe region that receives axonal projections from the olfactory bulb is the lateral entorhinal cortex.
- This six-layered cortex is a transitional cortex between the three-layered allocortex and the six-layered neocortex.
- This brain region has strong reciprocal connections with the piriform cortex.
- It preprocesses information entering the hippocampus and is intimately involved in learning and memory.
- In animals, conditioned odor-aversion learning is disrupted by lesions of the medial or lateral entorhinal cortex.
Secondary olfactory cortex (orbitofrontal cortex)
- Definition: Cortical regions receiving projections from the primary olfactory cortex.
- The structure most commonly termed secondary olfactory cortex in humans is the Orbitofrontal cortex.
- Located in the posterior ventral region of the frontal cortex.
- 5-layered agranular neocortex
- Receives projections from, and sends projections to, all primary olfactory regions,
- In addition to these direct connections, the orbitofrontal cortex also reciprocally connects to a number of these structures via the dorsomedial nucleus of the thalamus.
- This cortical region has extensive connections with brain regions associated with vision, touch, taste, and visceral sensations, providing cross-modal integration and associative learning.
- Function
- In addition to integrating food-related and odor-guided behaviours, the orbitofrontal cortex is intimately associated with judgments of odour familiarity, intensity, hedonicity, and quality, and facilitates the perception of flavour sensations