Mitral cell

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Mitral cell
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Coronal section of olfactory bulb.
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Plan of olfactory neurons.
Details
System Smell
Location Olfactory bulb of mammals
Identifiers
NeuroLex ID nifext_120
Anatomical terms of neuroanatomy

Mitral cells are neurons that are part of the olfactory system. They are located in the olfactory bulb in the mammalian central nervous system. They receive information from the axons of olfactory receptor neurons, forming synapses in neuropils called glomeruli. Axons of the mitral cells transfer information to a number of areas in the brain, including the piriform cortex, entorhinal cortex, and amygdala. Mitral cells receive excitatory input from olfactory sensory neurons and external tufted cells on their primary dendrites, whereas inhibitory input arises either from granule cells onto their lateral dendrites and soma or from periglomerular cells onto their dendritic tuft. Mitral cells together with tufted cells form an obligatory relay for all olfactory information entering from the olfactory nerve. Mitral cell output is not a passive reflection of their input from the olfactory nerve. In mice, each mitral cell sends a single primary dendrite into a glomerulus receiving input from a population of olfactory sensory neurons expressing identical olfactory receptor proteins, yet the odor responsiveness of the 20-40 mitral cells connected to a single glomerulus (called sister mitral cells) [1] is not identical to the tuning curve of the input cells, and also differs between sister mitral cells. [2] Odorant response properties of individual neurons in an olfactory glomerular module. The exact type of processing that mitral cells perform with their inputs is still a matter of controversy. One prominent hypothesis is that mitral cells encode the strength of an olfactory input into their firing phases relative to the sniff cycle. A second hypothesis is that the olfactory bulb network acts as a dynamical system that decorrelates to differentiate between representations of highly similar odorants over time. Support for the second hypothesis comes primarily from research in zebrafish (where mitral and tufted cells cannot be distinguished). [3]

Contents

Structure

Mitral cells are a neuronal cell type in the mammalian olfactory bulb, distinguished by the position of their somata located in an orderly row in the mitral cell layer of the bulb. [4] They typically have a single primary dendrite, which they project into a single glomerulus in the glomerular layer, and a few lateral dendrites that project laterally in the external plexiform layer. Mitral cells are closely related to the second type of projection neuron in the mammalian bulb, known as the tufted cell. In lower vertebrates, mitral cells cannot be morphologically distinguished from tufted cells, and both are substantially morphologically different from the mammalian mitral cells. The cells often have multiple primary dendrites innervating different glomeruli and they are sometimes called simply projection neurons, to indicate that they are the main neural element which project outside the olfactory bulb. The morphology of mitral cells was an advantage in early studies of synaptic processing, because the soma and the primary dendrite could be independently stimulated by appropriate positioning of stimulating electrodes in different layers of the olfactory bulb. [5]

Function

Synaptic processing

Mitral cells are a key part of the olfactory bulb microcircuit. Mitral cells receive input from at least four cell types: olfactory sensory neurons, periglomerular neurons, external tufted cells and granule cells. The synapses made by external tufted cells and olfactory sensory neurons are excitatory, whereas those of granule cells and periglomerular neurons are inhibitory. In addition, sister mitral cells are reciprocally connected by gap junctions. The mitral to granule and mitral to periglomerular cell synapse was the first description of the rather atypical reciprocal dendrodendritic synapses (in contrast to the more common axodendritic synapse). The action of the full glomerular microcircuit is a topic that is under intense scientific investigation. Certain principles are starting to emerge. One discovery points to the idea of the microcircuit between mitral, tufted and periglomerular cells in separating the output of mitral and tufted cells in time. [6] It appears that tufted cells receive strong olfactory nerve input, [7] fire close to inhalation onset and their firing phase is relatively concentration insensitive, whereas mitral cells receive relatively weak olfactory nerve input [8] and strong periglomerular inhibition, which delays their firing relative to the tufted cells. This escape from inhibition can be sped up by increasing the stimulating odorant concentration, and thus mitral cell firing phase acts as one possible way the olfactory system encodes concentration. The role of the mitral cell lateral dendrite and granule cell circuit is currently a bit more uncertain. One possible hypothesis implicates the system in forming sparse representation which enable more effective pattern separation. [9] The action of this circuit is heavily influenced by both short term and long term plasticity and ongoing granule cell neurogenesis. [10] The circuit requires the animal to be awake if it is to have full functionality.

Projection targets

Mitral and tufted cells project to various targets in the brain. Most importantly, projections target the olfactory cortex, where odor information can be integrated with input from other sensory modalities and used to drive behavior. Tufted cells project mainly to the anterior olfactory nucleus, a center that also performs comparison between left and right side olfactory input. Mitral cells project to the olfactory tubercle, where chemical information is integrated with auditory signals. Mitral cells carrying pheromonal inputs project to the amygdala and hypothalamus to drive instinctive behaviors. A major integrative center is the piriform cortex, where mitral cells make non-topographic projections to pyramidal cells which integrate information across glomeruli. Projections also go to the entorhinal cortex. Anatomical connectivity of a mitral cell axon can be quite different depending on the target structure. Whereas piriform cortex is innervated mostly randomly, projections to the anterior olfactory nucleus and amygdala retain some topographic order. Finally, mitral cell axons also make intrabulbar connections to granule cells and in the mouse olfactory system they project selectively to granule cells underlying the second ipsilateral homotypic (expressing the same olfactory receptor) glomerulus.

Related Research Articles

<span class="mw-page-title-main">Olfactory nerve</span> Cranial nerve I, for smelling

The olfactory nerve, also known as the first cranial nerve, cranial nerve I, or simply CN I, is a cranial nerve that contains sensory nerve fibers relating to the sense of smell.

<span class="mw-page-title-main">Olfactory bulb</span> Neural structure

The olfactory bulb is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It 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 bulb is divided into two distinct structures: the main olfactory bulb and the accessory olfactory bulb. The main olfactory bulb connects to the amygdala via the piriform cortex of the primary olfactory cortex and directly projects from the main olfactory bulb to specific amygdala areas. The accessory olfactory bulb resides on the dorsal-posterior region of the main olfactory bulb and forms a parallel pathway. Destruction of the olfactory bulb results in ipsilateral anosmia, while irritative lesions of the uncus can result in olfactory and gustatory hallucinations.

<span class="mw-page-title-main">Olfactory system</span> Sensory system used for smelling

The olfactory system or sense of smell is the sensory system used for smelling (olfaction). Olfaction is one of the special senses, that have directly associated specific organs. Most mammals and reptiles have a main olfactory system and an accessory olfactory system. The main olfactory system detects airborne substances, while the accessory system senses fluid-phase stimuli.

<span class="mw-page-title-main">Olfactory receptor neuron</span> Transduction nerve cell within the olfactory system

An olfactory receptor neuron (ORN), also called an olfactory sensory neuron (OSN), is a sensory neuron within the olfactory system.

<span class="mw-page-title-main">Olfactory epithelium</span> Specialised epithelial tissue in the nasal cavity that detects odours

The olfactory epithelium is a specialized epithelial tissue inside the nasal cavity that is involved in smell. In humans, it measures 5 cm2 (0.78 sq in) and lies on the roof of the nasal cavity about 7 cm (2.8 in) above and behind the nostrils. The olfactory epithelium is the part of the olfactory system directly responsible for detecting odors.

<span class="mw-page-title-main">Glomerulus (olfaction)</span>

The glomerulus is a spherical structure located in the olfactory bulb of the brain where synapses form between the terminals of the olfactory nerve and the dendrites of mitral, periglomerular and tufted cells. Each glomerulus is surrounded by a heterogeneous population of juxtaglomerular neurons and glial cells.

An apical dendrite is a dendrite that emerges from the apex of a pyramidal cell. Apical dendrites are one of two primary categories of dendrites, and they distinguish the pyramidal cells from spiny stellate cells in the cortices. Pyramidal cells are found in the prefrontal cortex, the hippocampus, the entorhinal cortex, the olfactory cortex, and other areas. Dendrite arbors formed by apical dendrites are the means by which synaptic inputs into a cell are integrated. The apical dendrites in these regions contribute significantly to memory, learning, and sensory associations by modulating the excitatory and inhibitory signals received by the pyramidal cells.

<span class="mw-page-title-main">Golgi cell</span>

In neuroscience, Golgi cells are the most abundant inhibitory interneurons found within the granular layer of the cerebellum. Golgi cells can be found in the granular layer at various layers. The Golgi cell is essential for controlling the activity of the granular layer. They were first identified as inhibitory in 1964. It was also the first example of an inhibitory feedback network in which the inhibitory interneuron was identified anatomically. Golgi cells produce a wide lateral inhibition that reaches beyond the afferent synaptic field and inhibit granule cells via feedforward and feedback inhibitory loops. These cells synapse onto the dendrite of granule cells and unipolar brush cells. They receive excitatory input from mossy fibres, also synapsing on granule cells, and parallel fibers, which are long granule cell axons. Thereby this circuitry allows for feed-forward and feed-back inhibition of granule cells.

The antennal lobe is the primary olfactory brain area in insects. The antennal lobe is a sphere-shaped deutocerebral neuropil in the brain that receives input from the olfactory sensory neurons in the antennae and mouthparts. Functionally, it shares some similarities with the olfactory bulb in vertebrates. The anatomy and physiology function of the insect brain can be studied by dissecting open the insect brain and imaging or carrying out in vivo electrophysiological recordings from it.

<span class="mw-page-title-main">Olfactory tubercle</span> Area at the bottom of the forebrain

The olfactory tubercle (OT), also known as the tuberculum olfactorium, is a multi-sensory processing center that is contained within the olfactory cortex and ventral striatum and plays a role in reward cognition. The OT has also been shown to play a role in locomotor and attentional behaviors, particularly in relation to social and sensory responsiveness, and it may be necessary for behavioral flexibility. The OT is interconnected with numerous brain regions, especially the sensory, arousal, and reward centers, thus making it a potentially critical interface between processing of sensory information and the subsequent behavioral responses.

A topographic map is the ordered projection of a sensory surface, like the retina or the skin, or an effector system, like the musculature, to one or more structures of the central nervous system. Topographic maps can be found in all sensory systems and in many motor systems.

Dysosmia is a disorder described as any qualitative alteration or distortion of the perception of smell. Qualitative alterations differ from quantitative alterations, which include anosmia and hyposmia. Dysosmia can be classified as either parosmia or phantosmia. Parosmia is a distortion in the perception of an odorant. Odorants smell different from what one remembers. Phantosmia is the perception of an odor when no odorant is present. The cause of dysosmia still remains a theory. It is typically considered a neurological disorder and clinical associations with the disorder have been made. Most cases are described as idiopathic and the main antecedents related to parosmia are URTIs, head trauma, and nasal and paranasal sinus disease. Dysosmia tends to go away on its own but there are options for treatment for patients that want immediate relief.

<span class="mw-page-title-main">Sense of smell</span> Sense that detects smells

The sense of smell, or olfaction, is the special sense through which smells are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste.

Gordon Murray Shepherd was an American neuroscientist who carried out basic experimental and computational research on how neurons are organized into microcircuits to carry out the functional operations of the nervous system. Using the olfactory system as a model that spans multiple levels of space, time and disciplines, his studies ranged from molecular to behavioral, recognized by an annual lecture at Yale University on "integrative neuroscience". At the time of his death, he was professor of neuroscience emeritus at the Yale School of Medicine. He graduated from Iowa State University with a BA, Harvard Medical School with an MD, and the University of Oxford with a DPhill.

<span class="mw-page-title-main">Granule cell</span> Type of neuron with a very small cell body

The name granule cell has been used for a number of different types of neurons whose only common feature is that they all have very small cell bodies. Granule cells are found within the granular layer of the cerebellum, the dentate gyrus of the hippocampus, the superficial layer of the dorsal cochlear nucleus, the olfactory bulb, and the cerebral cortex.

Tufted cells are found within the olfactory glomeruli. They receive input from the receptor cells of the olfactory epithelium found in areas of the nose able to sense smell. Both tufted cells and mitral cells are projection neurons. Projection neurons send the signals from the glomeruli deeper into the brain. The actual signal sent through these projection cells has been sharpened or filtered by a process called lateral inhibition. Both the periglomerular cells and the granule cells contribute to lateral inhibition. Projection neurons therefore transmit a sharpened olfactory signal to the deeper parts of the brain. Tufted cells project onto the anterior piriform cortex.

Dendrodendritic synapses are connections between the dendrites of two different neurons. This is in contrast to the more common axodendritic synapse (chemical synapse) where the axon sends signals and the dendrite receives them. Dendrodendritic synapses are activated in a similar fashion to axodendritic synapses in respects to using a chemical synapse. An incoming action potential permits the release of neurotransmitters to propagate the signal to the post synaptic cell. There is evidence that these synapses are bi-directional, in that either dendrite can signal at that synapse. Ordinarily, one of the dendrites will display inhibitory effects while the other will display excitatory effects. The actual signaling mechanism utilizes Na+ and Ca2+ pumps in a similar manner to those found in axodendritic synapses.

<span class="mw-page-title-main">Insect olfaction</span> Function of chemical receptors

Insect olfaction refers to the function of chemical receptors that enable insects to detect and identify volatile compounds for foraging, predator avoidance, finding mating partners and locating oviposition habitats. Thus, it is the most important sensation for insects. Most important insect behaviors must be timed perfectly which is dependent on what they smell and when they smell it. For example, olfaction is essential for locating host plants and hunting prey in many species of insects, such as the moth Deilephila elpenor and the wasp Polybia sericea, respectively.

Retronasal smell, retronasal olfaction, is the ability to perceive flavor dimensions of foods and drinks. Retronasal smell is a sensory modality that produces flavor. It is best described as a combination of traditional smell and taste modalities. Retronasal smell creates flavor from smell molecules in foods or drinks shunting up through the nasal passages as one is chewing. When people use the term "smell", they are usually referring to "orthonasal smell", or the perception of smell molecules that enter directly through the nose and up the nasal passages. Retronasal smell is critical for experiencing the flavor of foods and drinks. Flavor should be contrasted with taste, which refers to five specific dimensions: (1) sweet, (2) salty, (3) bitter, (4) sour, and (5) umami. Perceiving anything beyond these five dimensions, such as distinguishing the flavor of an apple from a pear for example, requires the sense of retronasal smell.

An axo-axonic synapse is a type of synapse, formed by one neuron projecting its axon terminals onto another neuron's axon.

References

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