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), which is 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. [29]

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. [30]

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. [31]

Evidence for existence in humans

Many studies have tried to determine whether there is a VNO in adult human beings. Trotier et al. [32] 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, [33] stated that the VNO structure disappears during fetal development as it does for some primates. [34] However, Smith and Bhatnagar (2000) [35] 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%). [36] 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. [37]

Given these findings, some scientists have argued that there is a VNO in adult human beings. [38] [39] However, most investigators have sought to identify the opening of the VNO in humans, rather than identify the tubular epithelial structure itself. [40] Thus it has been argued that such studies, employing macroscopic observational methods, have sometimes misidentified or even missed the vomeronasal organ. [40]

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. [41] 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. [42] Likewise, there is no evidence for any accessory olfactory bulb in adult human beings, [43] 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." [44]

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. 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.

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

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.

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

The rhinarium is the furless skin surface surrounding the external openings of the nostrils in many mammals. Commonly it is referred to as the tip of the snout, and breeders of cats and dogs sometimes use the term nose leather. Informally, it may be called a "truffle", "wet snout," or "wet nose” because its surface is moist in some species: for example, healthy dogs and cats.

<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.

<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.

Olfactory memory refers to the recollection of odors. Studies have found various characteristics of common memories of odor memory including persistence and high resistance to interference. Explicit memory is typically the form focused on in the studies of olfactory memory, though implicit forms of memory certainly supply distinct contributions to the understanding of odors and memories of them. Research has demonstrated that the changes to the olfactory bulb and main olfactory system following birth are extremely important and influential for maternal behavior. Mammalian olfactory cues play an important role in the coordination of the mother infant bond, and the following normal development of the offspring. Maternal breast odors are individually distinctive, and provide a basis for recognition of the mother by her offspring.

A sense is a biological system used by an organism for sensation, the process of gathering information about the surroundings through the detection of stimuli. Although, in some cultures, five human senses were traditionally identified as such, many more are now recognized. Senses used by non-human organisms are even greater in variety and number. During sensation, sense organs collect various stimuli for transduction, meaning transformation into a form that can be understood by the brain. Sensation and perception are fundamental to nearly every aspect of an organism's cognition, behavior and thought.

<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.

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.

Pherines, also known as vomeropherines, are odorless synthetic neuroactive steroids that engage nasal chemosensory receptors and induce dose-dependent and reversible pharmacological and behavioral effects. Pherines target human chemosensory receptors and possibly other receptors such as the GABAA receptor and influence central nervous system activity.

<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.

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

In zoology, copulation is animal sexual behavior in which a male introduces sperm into the female's body, especially directly into her reproductive tract. This is an aspect of mating. Many animals that live in water use external fertilization, whereas internal fertilization may have developed from a need to maintain gametes in a liquid medium in the Late Ordovician epoch. Internal fertilization with many vertebrates occurs via cloacal copulation, known as cloacal kiss, while most mammals copulate vaginally, and many basal vertebrates reproduce sexually with external fertilization.

References

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