Vomeronasal organ

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Vomeronasal organ
Gray51.png
Frontal section of nasal cavities of a human embryo 28 mm long (vomeronasal organ labeled at right)
Details
Precursor Nasal placode
Lymph Node
Identifiers
Latin organum vomeronasale
MeSH D019147
TA98 A06.1.02.008
TA2 3141
FMA 77280
Anatomical terminology

The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. [1] The name is derived from the fact that it lies adjacent to the unpaired vomer bone (from Latin vomer 'plowshare', for its shape) in the nasal septum. It is present and functional in all snakes and lizards, and in many mammals, including cats, dogs, cattle, pigs, and some primates. Some humans may have physical remnants of a VNO, but it is vestigial and non-functional.

Contents

The VNO contains the cell bodies of sensory neurons which have receptors that detect specific non-volatile (liquid) organic compounds which are conveyed to them from the environment. These compounds emanate from prey, predators, and the compounds called sex pheromones from potential mates. Activation of the VNO triggers an appropriate behavioral response to the presence of one of these three.

VNO neurons are activated by the binding of certain chemicals to their G protein-coupled receptors: they express receptors from three families, called V1R, [2] [3] [4] V2R, and FPR. [5] [6] The axons from these neurons, called cranial nerve zero (CN 0), project to the accessory olfactory bulb, which targets the amygdala and bed nucleus of the stria terminalis, which in turn project to the anterior hypothalamus. These structures constitute the accessory olfactory system.

The VNO triggers the flehmen response in some mammals, which helps direct liquid organic chemicals to the organ. The VNO was discovered by Frederik Ruysch prior to 1732, and later by Ludwig Jacobson in 1813. [7]

Structure

The organ

Placement of Jacobson's organ in a snake Jacobson's organ in a reptile.svg
Placement of Jacobson's organ in a snake

The VNO is found at the base of the nasal cavity. It is split into two, being divided by the nasal septum, with both sides possessing an elongated C-shaped, or crescent, lumen. It is encompassed inside a bony or cartilaginous capsule which opens into the base of the nasal cavity. [8]

The system

The vomeronasal receptor neurons possess axons which travel from the VNO to the accessory olfactory bulb (AOB), also known as the vomeronasal bulb. These sensory receptors are located on the medial concave surface of the crescent lumen. The lateral, convex surface of the lumen is covered with non-sensory ciliated cells, where the basal cells are also found. At the dorsal and ventral aspect of the lumen are vomeronasal glands, which fill the vomeronasal lumen with fluid. Sitting next to the lumen are blood vessels that dilate or constrict, forming a vascular pump that deliver stimuli to the lumen. A thin duct, which opens onto the floor of the nasal cavity inside the nostril, is the only way of access for stimulus chemicals.

During embryological development, the vomeronasal sensory neurons form from the nasal (olfactory) placode, at the anterior edge of the neural plate (cranial nerve zero).

Sensory epithelium and receptors

The VNO is a tubular crescent shape and split into two pairs, separated by the nasal septum. The medial, concave area of the lumen is lined with a pseudo stratified epithelium that has three main cell types: receptor cells, supporting cells, and basal cells. The supporting cells are located superficially on the membrane while the basal cells are found on the basement membrane near the non-sensory epithelium. The receptor neurons possess apical microvilli, to which the sensory receptors are localized. These are G-protein-coupled receptors, which are often referred to as pheromone receptors since vomeronasal receptors have been tied to detecting pheromones.

Three G-protein-coupled receptors have been identified in the VNO, each found in distinct regions: the V1Rs, V2Rs, and FPRs. V1Rs, V2Rs and FPRs are seven transmembrane receptors which are not closely related to odorant receptors expressed in the main olfactory neuroepithelium. [9]

The vomeronasal organ's sensory neurons act on a different signaling pathway than that of the main olfactory system's sensory neurons. Activation of the receptors stimulates phospholipase C, [11] which in turn opens the ion channel TRPC2. [12] [13] Upon stimulation activated by pheromones, IP3 production has been shown to increase in VNO membranes in many animals, while adenylyl cyclase and cyclic adenosine monophosphate (cAMP), the major signaling transduction molecules of the main olfactory system, remain unaltered. This trend has been shown in many animals, such as the hamster, the pig, the rat, and the garter snake upon introduction of vaginal or seminal secretions into the environment.

V1Rs and V2Rs are activated by distinct ligands or pheromones.

Many vomeronasal neurons are activated by chemicals in urine. Some of the active compounds are sulfated steroids. [17] Detecting the types and amounts of different sulfated steroids conveys information about the urine donor's physiological state, and may therefore serve as an honest signal.

Recent studies proved a new family of formyl peptide receptor like proteins in VNO membranes of mice, which points to a close phylogenetic relation of signaling mechanisms used in olfaction and chemosensors. [5]

Sensory neurons

Vomeronasal sensory neurons are extremely sensitive and fire action potentials at currents as low as 1 pA. Many patch-clamp recordings have confirmed the sensitivity of the vomeronasal neurons. This sensitivity is tied to the fact that the resting potential of the vomeronasal neurons is relatively close to that of the firing threshold of these neurons. Vomeronasal sensory neurons also show remarkably slow adaptation and the firing rate increases with increasing current up to 10 pA. The main olfactory sensory neurons fire single burst action potentials and show a much quicker adaptation rate. Activating neurons that have V1 receptors, V1Rs, cause field potentials that have weak, fluctuating responses that are seen the anterior of the accessory olfactory bulb, AOB. Activation of neurons that contain V2 receptors, V2Rs, however, promote distinct oscillations in the posterior of the AOB. [18]

Function

In mammals, the sensory neurons of the vomeronasal organ detect non-volatile chemical cues, which requires direct physical contact with the source of odor. Notably, some scents act as chemical-communication signals (pheromones) from other individuals of the same species. Unlike the main olfactory bulb that sends neuronal signals to the olfactory cortex, the VNO sends neuronal signals to the accessory olfactory bulb and then to the amygdala, BNST, and ultimately hypothalamus. Since the hypothalamus is a major neuroendocrine center (affecting aspects of reproductive physiology and behavior as well as other functions such as body temperature), this may explain how scents influence aggressive and mating behavior. For example, in many vertebrates, nerve signals from the brain pass sensory information to the hypothalamus about seasonal changes and the availability of a mate. In turn, the hypothalamus regulates the release of reproductive hormones required for breeding. [19] Some pheromones are detected by the main olfactory system. [20]

In animals

The vomeronasal organ originated in tetrapods. The functional vomeronasal system is found in all snakes and lizards, [21] and many mammals.

Sagittal section of the vomeronasal organ of garter snake VO of garter snake sagittal section.jpg
Sagittal section of the vomeronasal organ of garter snake

In some other mammals the entire organ contracts or pumps in order to draw in the scents. [33]

Stallion exhibiting the flehmen response Flehmendes Pferd 32 c.jpg
Stallion exhibiting the flehmen response

Flehmen response

Some mammals, particularly felids (cats) and ungulates (which includes horses, cattle, and pigs among other species), use a distinctive facial movement called the flehmen response to direct inhaled compounds to the VNO. The animal lifts its head after finding the odorant, wrinkles its nose while lifting its lips, and ceases to breathe momentarily.

Flehmen behavior is associated with "anatomical specialization", and animals that present flehmen behavior have incisive papilla and ducts, which connect the oral cavity to the VNO, that are found behind their teeth. However, horses are the exception: they exhibit flehmen response but do not have an incisive duct communication between the nasal and the oral cavity because they do not breathe through their mouths; instead, the VNOs connect to the nasal passages by the nasopalatine duct. [34]

Cats use their vomeronasal organ when scent rubbing; they are able to discriminate between similar smelling substances using this organ, and then perform the rubbing behaviour. [35]

Evidence for existence in humans

Many studies have tried to determine whether there is a VNO in adult human beings. Trotier et al. [36] estimated that around 92% of their subjects that had no septal surgery had at least one intact VNO. Kjaer and Fisher Hansen, on the other hand, [37] stated that the VNO structure disappears during fetal development as it does for some primates. [38] However, Smith and Bhatnagar (2000) [39] asserted that Kjaer and Fisher Hansen simply missed the structure in older fetuses. Won (2000) found evidence of a VNO in 13 of his 22 cadavers (59.1%) and 22 of his 78 living patients (28.2%). [40] In a study using retrospective analysis of nearly one thousand outpatient nasal endoscopies, Stoyanov et al. (2016) found the organ to be present in 26.83% of the Bulgarian population. [41]

Given these findings, some scientists have argued that there is a VNO in adult human beings. [42] [43] However, most investigators have sought to identify the opening of the VNO in humans, rather than identify the tubular epithelial structure itself. [44] Thus it has been argued that such studies, employing macroscopic observational methods, have sometimes misidentified or even missed the vomeronasal organ. [44]

Among studies that use microanatomical methods, there is no reported evidence that human beings have active sensory neurons like those in working vomeronasal systems of other animals. [45] Furthermore, there is no evidence to date that suggests there are nerve and axon connections between any existing sensory receptor cells that may be in the adult human VNO and the brain. [46] Likewise, there is no evidence for any accessory olfactory bulb in adult human beings, [47] and the key genes involved in VNO function in other mammals have pseudogenized in human beings. Therefore, while many debate the structure's presence in adult human beings, a review of the scientific literature by Tristram Wyatt concluded that on current evidence, "most in the field... are skeptical about the likelihood of a functional VNO in adult human beings." [48]

History

The VNO was discovered by Frederik Ruysch prior to 1732, and later by Ludwig Jacobson in 1813. [7]

Related Research Articles

<span class="mw-page-title-main">Pheromone</span> Secreted or excreted chemical factor that triggers a social response in members of the same species

A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavior of the receiving individuals. There are alarm pheromones, food trail pheromones, sex pheromones, and many others that affect behavior or physiology. Pheromones are used by many organisms, from basic unicellular prokaryotes to complex multicellular eukaryotes. Their use among insects has been particularly well documented. In addition, some vertebrates, plants and ciliates communicate by using pheromones. The ecological functions and evolution of pheromones are a major topic of research in the field of chemical ecology.

<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">Sensory nervous system</span> Part of the nervous system

The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons, neural pathways, and parts of the brain involved in sensory perception and interoception. Commonly recognized sensory systems are those for vision, hearing, touch, taste, smell, balance and visceral sensation. Sense organs are transducers that convert data from the outer physical world to the realm of the mind where people interpret the information, creating their perception of the world around them.

<span class="mw-page-title-main">Flehmen response</span> Behavior in which an animal curls back its upper lip exposing its front teeth

The flehmen response, also called the flehmen position, flehmen reaction, flehmen grimace, flehming, or flehmening, is a behavior in which an animal curls back its upper lip exposing its front teeth, inhales with the nostrils usually closed, and then often holds this position for several seconds. It may be performed over a sight or substance of particular interest to the animal, or may be performed with the neck stretched and the head held high in the air.

<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.

A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance to generate a biological signal. This signal may be in the form of an action potential, if the chemoreceptor is a neuron, or in the form of a neurotransmitter that can activate a nerve fiber if the chemoreceptor is a specialized cell, such as taste receptors, or an internal peripheral chemoreceptor, such as the carotid bodies. In physiology, a chemoreceptor detects changes in the normal environment, such as an increase in blood levels of carbon dioxide (hypercapnia) or a decrease in blood levels of oxygen (hypoxia), and transmits that information to the central nervous system which engages body responses to restore homeostasis.

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

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

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

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<span class="mw-page-title-main">VN1R1</span> Protein-coding gene in the species Homo sapiens

Vomeronasal type-1 receptor 1 is a protein that in humans is encoded by the VN1R1 gene.

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<span class="mw-page-title-main">Vomeronasal receptor</span> Class of olfactory receptors

Vomeronasal receptors are a class of olfactory receptors that putatively function as receptors for pheromones. Pheromones have evolved in all animal phyla, to signal sex and dominance status, and are responsible for stereotypical social and sexual behaviour among members of the same species. In mammals, these chemical signals are believed to be detected primarily by the vomeronasal organ (VNO), a chemosensory organ located at the base of the nasal septum.

Odour is sensory stimulation of the olfactory membrane of the nose by a group of molecules. Certain body odours are connected to human sexual attraction. Humans can make use of body odour subconsciously to identify whether a potential mate will pass on favourable traits to their offspring. Body odour may provide significant cues about the genetic quality, health and reproductive success of a potential mate.

<span class="mw-page-title-main">Mammalian reproduction</span> Most mammals are viviparous, giving birth to live young

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Odor molecules are detected by the olfactory receptors in the olfactory epithelium of the nasal cavity. Each receptor type is expressed within a subset of neurons, from which they directly connect to the olfactory bulb in the brain. Olfaction is essential for survival in most vertebrates; however, the degree to which an animal depends on smell is highly varied. Great variation exists in the number of OR genes among vertebrate species, as shown through bioinformatic analyses. This diversity exists by virtue of the wide-ranging environments that they inhabit. For instance, dolphins that are secondarily adapted to an aquatic niche possess a considerably smaller subset of genes than most mammals. OR gene repertoires have also evolved in relation to other senses, as higher primates with well-developed vision systems tend to have a smaller number of OR genes. As such, investigating the evolutionary changes of OR genes can provide useful information on how genomes respond to environmental changes. Differences in smell sensitivity are also dependent on the anatomy of the olfactory apparatus, such as the size of the olfactory bulb and epithelium.

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

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<span class="mw-page-title-main">Copulation (zoology)</span> Animal sexual reproductive act in which a male introduces sperm into the females body

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<span class="mw-page-title-main">BPIFB9P</span> Pseudogene in the species Homo sapiens

Vomeromodulin is a non-human protein also known as BPI fold containing family B, member 9 (BPIFB9) in the rat encoded by the Bpifb9/RYF3 gene, and as BPI fold containing family B, member 9A (BPIFB9A) encoded by the Bpifb9a gene in the mouse. This protein has been characterized in mammals such as rodents, carnivores, even-toed ungulates, insectivores, bats, lagomorphs, and shrews but is apparently absent in primates and other vertebrates such as birds, reptiles, and amphibians. Its function is associated with detection of chemical odorant pheromone molecules.

References

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  41. ↑ Stoyanov G, Moneva K, Sapundzhiev N, Tonchev AB (April 2016). "The vomeronasal organ - incidence in a Bulgarian population". The Journal of Laryngology and Otology. 130 (4): 344–347. doi:10.1017/S0022215116000189. PMID   26831012. S2CID   1696242.
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  46. ↑ Wysocki CJ, Preti G (November 2004). "Facts, fallacies, fears, and frustrations with human pheromones". The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology. 281 (1): 1201–1211. doi: 10.1002/ar.a.20125 . PMID   15470677.
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  48. ↑ Wyatt, Tristram D. (2003). Pheromones and Animal Behaviour: Communication by Smell and Taste. Cambridge: Cambridge University Press. ISBN   0-521-48526-6. p295

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