Glomerulus (olfaction)

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Glomerulus (olfaction)
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Coronal section of olfactory bulb.
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Plan of olfactory neurons.
Identifiers
NeuroLex ID nlx_anat_1005011
Anatomical terms of neuroanatomy

The glomerulus (pl.: glomeruli) 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 (that include periglomerular, short axon, and external tufted cells) and glial cells. [1] [2] [3]

Contents

All glomeruli are located near the surface of the olfactory bulb. The olfactory bulb also includes a portion of the anterior olfactory nucleus, the cells of which contribute fibers to the olfactory tract. [4] They are the initial sites for synaptic processing of odor information coming from the nose. A glomerulus is made up of a globular tangle of axons from the olfactory receptor neurons, and dendrites from the mitral and tufted cells, as well as, from cells that surround the glomerulus such as the external tufted cells, periglomerular cells, short axon cells, and astrocytes. In mammals, glomeruli typically range between 50-120 µm in diameter and number between 1100 and 2400 depending on the species, with roughly between 1100 and 1200 in humans. [1] [2] The number of glomeruli in a human decreases with age; in humans that are over 80 they are nearly absent. [5] Each glomerulus is composed of two compartments, the olfactory nerve zone and the non-olfactory nerve zone. The olfactory nerve zone is composed of preterminals and terminals of the olfactory nerve and is where the olfactory receptor cells make synapses on their targets. [2] The non-olfactory nerve zone is composed of the dendritic processes of intrinsic neurons and is where dendrodendritic interactions between intrinsic neurons occur. [2]

Structure

Glomeruli are important waystations in the pathway from the nose to the olfactory cortex and have been found to be critical for odorant signal transduction. The olfactory receptor neurons (ORN), which originate in the nasal epithelium express only one type of olfactory receptor (OR). These ORNs then project their axons to the olfactory bulb. In the olfactory bulb, the ORNs synapse with termination in the glomeruli. [6] Each glomerulus receives input from olfactory receptor neurons expressing only one type of olfactory receptor. The glomerular activation patterns within the olfactory bulb are thought to represent the quality of the odor being detected. These activation patterns of glomeruli can change due to changes in airflow rate and odor concentration in the mucus layer of the nasal cavity. [7] [8] A certain odorant can activate a glomeruli strongly whilst affecting others with less efficiency to very little at all. Linda Buck and Richard Axel were awarded a Nobel prize in 2004 for heavily influencing the working out of the genetic basis for this Olfactory coding.

The current dogma is that axons from all ORNs expressing the same receptor converge onto one or two glomeruli of a possible 1800 glomeruli in each olfactory bulb. [6] As the axons of the ORNs migrate towards their specific glomeruli they often overshoot into neighboring glomeruli. Thus, a glomerulus representing a specific OR develops slowly and involves considerable axonal reorganization in order to achieve the highly topographical projection observed in adult mice. [9]

Function

The glomerulus is the basic unit in the odor map of the olfactory bulb. Each odor activates a different pattern of glomeruli, such that, simply by analyzing the different sets of activated glomeruli, one could, in theory, decode the identity of the odor. This odor map, however, is modified by the circuitry within the olfactory bulb so that the spiking pattern of the second-order mitral cells is usually different from that of the olfactory sensory neurons. [10]

Other species

Glomerulus in dogs

The glomerulus process in dogs is divided in 3 parts: signal acquisition, signal transduction, and signal processing. Some dogs have as many as 100 times more ORNs than humans do, producing a correspondingly sharpened ability to detect and discriminate among millions of odors. [11] Characteristics of these stages include: the role of olfactory plume and sniffing, the continuous renewal of Glomerulus receptors throughout the life cycle, and the relationship between the olfactory neuron and the glomerulus, and finally, the synthetic nature of glomerulus coding. [12]

Glomerulus in fish

One of the most distinctive features of fish olfaction is that it takes place entirely in the aquatic environment. The carrier of stimulant is water and therefore the chemicals must be soluble in water. The olfactory epithelium of fish consists of three cell types, like other vertebrates. These three cell types are the receptor cells, supporting cells and basal cells. [13]

Fish glomerulus differs from the mammalian glomerulus in terms of the number of dendrites that it receives from the mitral cells. In a mammalian olfactory system, a single dendrite from a mitral cell enters a single glomerulus. However, in fish, one or more dendrites from mitral cells enter one or more glomerulus. [13]

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 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.

Synaptogenesis is the formation of synapses between neurons in the nervous system. Although it occurs throughout a healthy person's lifespan, an explosion of synapse formation occurs during early brain development, known as exuberant synaptogenesis. Synaptogenesis is particularly important during an individual's critical period, during which there is a certain degree of synaptic pruning due to competition for neural growth factors by neurons and synapses. Processes that are not used, or inhibited during their critical period will fail to develop normally later on in life.

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.

<span class="mw-page-title-main">Mitral cell</span> Neurons that are part of the olfactory system

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 is not identical to the tuning curve of the input cells, and also differs between sister mitral cells. 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.

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.

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.

<span class="mw-page-title-main">Anterior olfactory nucleus</span> Portion of the forebrain of vertebrates

The anterior olfactory nucleus is a portion of the forebrain of vertebrates.

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.

The Mauthner cells are a pair of big and easily identifiable neurons located in the rhombomere 4 of the hindbrain in fish and amphibians that are responsible for a very fast escape reflex. The cells are also notable for their unusual use of both chemical and electrical synapses.

<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.

The lateral horn is one of the two areas of the insect brain where projection neurons of the antennal lobe send their axons. The other area is the mushroom body. Several morphological classes of neurons in the lateral horn receive olfactory information through the projection neurons.

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.

References

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