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Mucus ( ) is a slippery aqueous secretion produced by, and covering,
mucous membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It i ...
s. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It is a viscous colloid containing inorganic salts,
antimicrobial An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
enzymes (such as
lysozyme Lysozyme (EC 3.2.1.17, muramidase, ''N''-acetylmuramide glycanhydrolase; systematic name peptidoglycan ''N''-acetylmuramoylhydrolase) is an antimicrobial enzyme produced by animals that forms part of the innate immune system. It is a glycoside ...
s), immunoglobulins (especially IgA), and glycoproteins such as
lactoferrin Lactoferrin (LF), also known as lactotransferrin (LTF), is a multifunctional protein of the transferrin family. Lactoferrin is a globular glycoprotein with a molecular mass of about 80 kDa that is widely represented in various secretory fluids, s ...
and
mucin Mucins () are a family of high molecular weight, heavily glycosylated proteins (glycoconjugates) produced by epithelial tissues in most animals. Mucins' key characteristic is their ability to form gels; therefore they are a key component in most ...
s, which are produced by goblet cells in the mucous membranes and
submucosal glands Submucosal glands can refer to various racemose exocrine glands of the mucous type. These glands secrete mucus to facilitate the movement of particles along the body's various tubes, such as the throat and intestines. The mucosa is the lining of th ...
. Mucus serves to protect epithelial cells in the linings of the
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gre ...
, digestive, and
urogenital system The genitourinary system, or urogenital system, are the organs of the reproductive system and the urinary system. These are grouped together because of their proximity to each other, their common embryological origin and the use of common pathw ...
s, and structures in the visual and auditory systems from
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
ic
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
,
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
and
viruses A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsky's ...
. Most of the mucus in the body is produced in the gastrointestinal tract. Amphibians,
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
, snails, slugs, and some other
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordat ...
also produce external mucus from their epidermis as protection against pathogens, and to help in movement and is also produced in fish to line their gills. Plants produce a similar substance called
mucilage Mucilage is a thick, gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion. The direction of their movement is always opposite to that of the secretion of m ...
that is also produced by some microorganisms.


Respiratory system

In the human
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies ...
, mucus is part of the airway surface liquid (ASL), also known as epithelial lining fluid (ELF), that lines most of the respiratory tract. The airway surface liquid consists of a ''sol'' layer termed the periciliary liquid layer and an overlying ''gel'' layer termed the mucus layer. The periciliary liquid layer is so named as it surrounds the cilia and lies on top of the surface epithelium. The periciliary liquid layer surrounding the cilia consists of a gel meshwork of cell-tethered mucins and polysaccharides. The mucus blanket aids in the protection of the lungs by trapping foreign particles before they enter them, in particular through the nose during normal breathing. Mucus is made up of a fluid component of around 95% water, the
mucin Mucins () are a family of high molecular weight, heavily glycosylated proteins (glycoconjugates) produced by epithelial tissues in most animals. Mucins' key characteristic is their ability to form gels; therefore they are a key component in most ...
secretions from the goblet cells, and the submucosal glands (2–3% glycoproteins), proteoglycans (0.1–0.5%), lipids (0.3–0.5%), proteins, and DNA. The major mucins secreted –
MUC5AC Mucin-5AC (MUC-5AC) is a protein that in humans is encoded by the ''MUC5AC'' gene. MUC-5AC is a large gel-forming glycoprotein. In the respiratory tract it protects against infection by binding to inhaled pathogens that are subsequently removed ...
and
MUC5B Mucin-5B (MUC-5B) is a protein that in humans is encoded by the ''MUC5B'' gene and by ''Muc5b'' gene in mouse. It is one of the five gel-forming mucins. MUC-5B can be found in whole saliva, normal lung mucus, and cervical mucus. In some disease ...
- are large polymers that give the mucus its rheologic or
viscoelastic In materials science and continuum mechanics, viscoelasticity is the property of materials that exhibit both viscous and elastic characteristics when undergoing deformation. Viscous materials, like water, resist shear flow and strain linearly ...
properties. MUC5AC is the main gel-forming mucin secreted by goblet cells, in the form of threads and thin sheets. MUC5B is a polymeric protein secreted from submucosal glands and some goblet cells, and this is in the form of strands. In the airways—the trachea,
bronchi A bronchus is a passage or airway in the lower respiratory tract that conducts air into the lungs. The first or primary bronchi pronounced (BRAN-KAI) to branch from the trachea at the carina are the right main bronchus and the left main bronchus. ...
, and
bronchiole The bronchioles or bronchioli (pronounced ''bron-kee-oh-lee'') are the smaller branches of the bronchial airways in the lower respiratory tract. They include the terminal bronchioles, and finally the respiratory bronchioles that mark the start ...
s—the lining of mucus is produced by specialized airway epithelial cells called goblet cells, and
submucosal glands Submucosal glands can refer to various racemose exocrine glands of the mucous type. These glands secrete mucus to facilitate the movement of particles along the body's various tubes, such as the throat and intestines. The mucosa is the lining of th ...
. Small particles such as dust, particulate
pollutant A pollutant or novel entity is a substance or energy introduced into the environment that has undesired effects, or adversely affects the usefulness of a resource. These can be both naturally forming (i.e. minerals or extracted compounds like o ...
s, and allergens, as well as infectious agents and
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
are caught in the viscous nasal or airway mucus and prevented from entering the system. This process, together with the continual movement of the cilia on the respiratory epithelium toward the
oropharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
(
mucociliary clearance Mucociliary clearance (MCC), mucociliary transport, or the mucociliary escalator, describes the self-clearing mechanism of the airways in the respiratory system. It is one of the two protective processes for the lungs in removing inhaled particl ...
), helps prevent foreign objects from entering the lungs during breathing. This explains why coughing often occurs in those who smoke cigarettes. The body's natural reaction is to increase mucus production. In addition, mucus aids in moisturizing the inhaled air and prevents tissues such as the nasal and airway epithelia from drying out. Mucus is produced continuously in the respiratory tract. Mucociliary action carries it down from the nasal passages and up from the rest of the tract to the pharynx, with most of it being swallowed subconsciously. Sometimes in times of respiratory illness or inflammation, mucus can become thickened with cell debris, bacteria, and inflammatory cells. It is then known as phlegm which may be
cough A cough is a sudden expulsion of air through the large breathing passages that can help clear them of fluids, irritants, foreign particles and microbes. As a protective reflex, coughing can be repetitive with the cough reflex following three ph ...
ed up as sputum to clear the airway.


Respiratory tract

Increased mucus production in the
upper respiratory tract The respiratory tract is the subdivision of the respiratory system involved with the process of respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed in through the nose to t ...
is a symptom of many common ailments, such as the common cold, and influenza. Nasal mucus may be removed by blowing the nose or by using nasal irrigation. Excess nasal mucus, as with a cold or
allergies Allergies, also known as allergic diseases, refer a number of conditions caused by the hypersensitivity of the immune system to typically harmless substances in the environment. These diseases include hay fever, food allergies, atopic dermat ...
, due to vascular engorgement associated with vasodilation and increased capillary permeability caused by histamines, may be treated cautiously with
decongestant A decongestant, or nasal decongestant, is a type of pharmaceutical drug that is used to relieve nasal congestion in the upper respiratory tract. The active ingredient in most decongestants is either pseudoephedrine or phenylephrine (the latter ...
medications. Thickening of mucus as a "rebound" effect following overuse of decongestants may produce nasal or sinus drainage problems and circumstances that promote infection. During cold, dry seasons, the mucus lining nasal passages tends to dry out, meaning that mucous membranes must work harder, producing more mucus to keep the cavity lined. As a result, the nasal cavity can fill up with mucus. At the same time, when air is exhaled, water vapor in breath condenses as the warm air meets the colder outside temperature near the nostrils. This causes an excess amount of water to build up inside nasal cavities. In these cases, the excess fluid usually spills out externally through the nostrils. In the
lower respiratory tract The respiratory tract is the subdivision of the respiratory system involved with the process of Respiration (physiology), respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed ...
impaired
mucociliary clearance Mucociliary clearance (MCC), mucociliary transport, or the mucociliary escalator, describes the self-clearing mechanism of the airways in the respiratory system. It is one of the two protective processes for the lungs in removing inhaled particl ...
due to conditions such as
primary ciliary dyskinesia Primary ciliary dyskinesia (PCD) is a rare, autosomal recessive genetic ciliopathy, that causes defects in the action of cilia lining the upper and lower respiratory tract, sinuses, Eustachian tube, middle ear, Fallopian tube, and flagella of sper ...
may result in mucus accumulation in the bronchi. The dysregulation of mucus homeostasis is the fundamental characteristic of cystic fibrosis, an inherited disease caused by mutations in the ''
CFTR Cystic fibrosis transmembrane conductance regulator (CFTR) is a membrane protein and anion channel in vertebrates that is encoded by the ''CFTR'' gene. Geneticist Lap-Chee Tsui and his team identified the CFTR gene in 1989 as the gene linked wi ...
'' gene, which encodes a
chloride channel Chloride channels are a superfamily of poorly understood ion channels specific for chloride. These channels may conduct many different ions, but are named for chloride because its concentration ''in vivo'' is much higher than other anions. Several ...
. This defect leads to the altered electrolyte composition of mucus, which triggers its hyperabsorption and dehydration. Such low-volume, viscous, acidic mucus has a reduced antimicrobial function, which facilitates bacterial colonisation. The thinning of the mucus layer ultimately affects the periciliary liquid layer, which becomes dehydrated, compromising ciliary function, and impairing mucociliary clearance. A respiratory therapist can recommend airway clearance therapy which uses a number of clearance techniques to help with the clearance of mucus.


Mucus hypersecretion

In the
lower respiratory tract The respiratory tract is the subdivision of the respiratory system involved with the process of Respiration (physiology), respiration in mammals. The respiratory tract is lined with respiratory epithelium as respiratory mucosa. Air is breathed ...
excessive mucus production in the bronchi and bronchioles is known as mucus hypersecretion. Chronic mucus hypersecretion results in the chronic productive cough of
chronic bronchitis Bronchitis is inflammation of the bronchi (large and medium-sized airways) in the lungs that causes coughing. Bronchitis usually begins as an infection in the nose, ears, throat, or sinuses. The infection then makes its way down to the bronchi. ...
, and is generally synonymous with this. Excessive mucus can narrow the airways, limit airflow, and accelerate a decline in lung function.


Digestive system

In the
human digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compo ...
, mucus is used as a lubricant for materials that must pass over membranes, e.g., food passing down the
esophagus The esophagus ( American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to ...
. Mucus is extremely important in the gastrointestinal tract. It forms an essential layer in the colon and in the small intestine that helps reduce intestinal inflammation by decreasing bacterial interaction with intestinal epithelial cells. The layer of mucus of the
gastric mucosa The gastric mucosa is the mucous membrane layer of the stomach, which contains the glands and the gastric pits. In humans, it is about 1 mm thick, and its surface is smooth, soft, and velvety. It consists of simple columnar epithelium, lamina ...
lining the
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
is vital to protect the stomach lining from the highly acidic environment within it.


Reproductive system

In the human female reproductive system, cervical mucus prevents infection and provides lubrication during sexual intercourse. The consistency of cervical mucus varies depending on the stage of a woman's menstrual cycle. At
ovulation Ovulation is the release of eggs from the ovaries. In women, this event occurs when the ovarian follicles rupture and release the secondary oocyte ovarian cells. After ovulation, during the luteal phase, the egg will be available to be fertilize ...
cervical mucus is clear, runny, and conducive to sperm; post-ovulation, mucus becomes thicker and is more likely to block sperm. Several Fertility Awareness methods rely on observation of cervical mucus, as one of three primary fertility signs, to identify a woman's fertile time at the mid-point of the cycle. Awareness of the woman's fertile time allows a couple to time intercourse to improve the odds of pregnancy. It is also proposed as a method to avoid pregnancy.


Clinical significance

In general, nasal mucus is clear and thin, serving to filter air during inhalation. During times of infection, mucus can change color to yellow or green either as a result of trapped
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
or due to the body's reaction to viral infection. The green color of mucus comes from the heme group in the iron-containing enzyme
myeloperoxidase Myeloperoxidase (MPO) is a peroxidase enzyme that in humans is encoded by the ''MPO'' gene on chromosome 17. MPO is most abundantly expressed in neutrophil granulocytes (a subtype of white blood cells), and produces hypohalous acids to carry ...
secreted by white blood cells as a cytotoxic defense during a
respiratory burst Respiratory burst (or oxidative burst) is the rapid release of the reactive oxygen species (ROS), superoxide anion () and hydrogen peroxide (), from different cell types. This is usually utilised for mammalian immunological defence, but also play ...
. In the case of bacterial infection, the bacterium becomes trapped in already-clogged sinuses, breeding in the moist, nutrient-rich environment.
Sinusitis Sinusitis, also known as rhinosinusitis, is inflammation of the mucous membranes that line the sinuses resulting in symptoms that may include thick nasal mucus, a plugged nose, and facial pain. Other signs and symptoms may include fever, head ...
is an uncomfortable condition which may include congestion of mucus. A bacterial infection in sinusitis will cause discolored mucus and would respond to antibiotic treatment; viral infections typically resolve without treatment. Almost all sinusitis infections are viral and antibiotics are ineffective and not recommended for treating typical cases. In the case of a viral infection such as
cold Cold is the presence of low temperature, especially in the atmosphere. In common usage, cold is often a subjective perception. A lower bound to temperature is absolute zero, defined as 0.00K on the Kelvin scale, an absolute thermodynamic ...
or flu, the first stage and also the last stage of the infection cause the production of a clear, thin mucus in the nose or back of the throat. As the body begins to react to the virus (generally one to three days), mucus thickens and may turn yellow or green. Viral infections cannot be treated with antibiotics, and are a major avenue for their misuse. Treatment is generally symptom-based; often it is sufficient to allow the immune system to fight off the virus over time.
Obstructive lung disease Obstructive lung disease is a category of respiratory disease characterized by airway obstruction. Many obstructive diseases of the lung result from narrowing (obstruction) of the smaller bronchi and larger bronchioles, often because of excessiv ...
s often result from impaired mucociliary clearance that can be associated with mucus hypersecretion, and these are sometimes referred to as ''mucoobstructive lung diseases''. Techniques of airway clearance therapy can help to clear secretions, maintain respiratory health, and prevent inflammation in the airways. A unique umbilical cord lining epithelial stem cell expresses MUC1, termed (CLEC-muc). This has been shown to have good potential in the regeneration of the cornea.


Properties of mucus


Tunable swelling capacity

Mucus is able to absorb water or dehydrate through pH variations. The swelling capacity of mucus stems from the bottlebrush structure of
mucin Mucins () are a family of high molecular weight, heavily glycosylated proteins (glycoconjugates) produced by epithelial tissues in most animals. Mucins' key characteristic is their ability to form gels; therefore they are a key component in most ...
within which hydrophilic segments provide a large surface area for water absorption. Moreover, the tunability of swelling effect is controlled by polyelectrolyte effect.


Polyelectrolyte effect in mucus

Polymers with charged molecules are called
polyelectrolyte Polyelectrolytes are polymers whose repeating units bear an electrolyte group. Polycations and polyanions are polyelectrolytes. These groups dissociate in aqueous solutions (water), making the polymers charged. Polyelectrolyte properties are ...
s. Mucins, a kind of polyelectrolyte proteoglycans, are the main component of mucus, which provides the polyelectrolyte effect in mucus. The process of inducing this effect comprises two steps: attraction of counter-ions and water compensation. When exposed in physiological ionic solution, the charged groups in the polyelectrolytes attract counter-ions with opposite charges, thereby leading to a solute concentration gradient. An osmotic pressure is introduced to equalize the concentration of solute throughout the system by driving water to flow from the low concentration areas to the high concentration areas. In short, the influx and outflux of water within mucus, managed by the polyelectrolyte effect, contribute to mucus' tunable swelling capacity.


Mechanism of pH-tunable swelling

The ionic charges of mucin are mainly provided by acidic amino acids including aspartic acid ( pKa=3.9) and glutamic acid (pKa=4.2). The charges of acidic amino acids will change with environmental pH value due to acid dissociation and association. Aspartic acid, for example, has a negative side chain when the pH value is above 3.9, while a neutrally charged side chain will be introduced as pH value drops below 3.9. Thus, the number of negative charges in mucus is influenced by the pH value of surrounding environment. That is, the polyelectrolyte effect of mucus is largely affected by the pH value of solution due to the charge variation of acidic amino acid residues on the mucin backbone. For instance, the charged residue on mucin is protonated at a normal pH value of the stomach, approximately pH 2. In this case, there is scarcely polyelectrolyte effect, thereby causing compact mucus with little swelling capacity. However, a kind of bacteria, ''
Helicobacter pylori ''Helicobacter pylori'', previously known as ''Campylobacter pylori'', is a gram-negative, microaerophilic, spiral (helical) bacterium usually found in the stomach. Its helical shape (from which the genus name, helicobacter, derives) is thoug ...
'', is prone to producing base to elevate the pH value in stomach, leading to the deprotonation of aspartic acids and glutamic acids, i.e., from neutral to negative-charged. The negative charges in the mucus greatly increase, thus inducing the polyelectrolyte effect and the swelling of the mucus. This swelling effect increases the pore size of the mucus and decreases mucus' viscosity, which allows bacteria to penetrate and migrate into the mucus and cause disease.


Charge selectivity

The high selective permeability of mucus plays a crucial role in the healthy state of human beings by limiting the penetration of molecules, nutrients, pathogens, and drugs. The charge distribution within mucus serves as a charge selective diffusion barrier, thus significantly affecting the transportation of agents. Among particles with various surface
zeta potential Zeta potential is the electrical potential at the slipping plane. This plane is the interface which separates mobile fluid from fluid that remains attached to the surface. Zeta potential is a scientific term for electrokinetic potential in coll ...
s, cationic particles tend to have a low depth of penetration, neutral ones possess medium penetration, and anionic ones have the largest penetration depth. Furthermore, the effect of charge selectivity changes when the status of the mucus varies, i.e., native mucus has a threefold higher potential to limit agent penetration than purified mucus.


Other animals

Mucus is also produced by a number of other animals. All
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
are covered in mucus secreted from glands all over their bodies. Invertebrates such as snails and slugs secrete mucus called snail slime to enable movement, and to prevent their bodies from drying out. Their reproductive systems also make use of mucus for example in the covering of their eggs. In the unique mating ritual of '' Limax maximus'' the mating slugs lower themselves from elevated locations by a mucus thread. Mucus is an essential constituent of
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, ...
slime used to deter predators. Mucus is produced by the endostyle in some tunicates and larval
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s to help in filter feeding.


See also

*
Alkaline mucus Alkaline mucus is a thick fluid produced by animals which confers tissue protection in an acidic environment, such as in the stomach. Properties Mucus that serves a protective function against acidic environments generally has a high viscosity, ...
*
Empty nose syndrome Empty may refer to: ‍ Music Albums * ''Empty'' (God Lives Underwater album) or the title song, 1995 * ''Empty'' (Nils Frahm album), 2020 * ''Empty'' (Tait album) or the title song, 2001 Songs * "Empty" (The Click Five song), 2007 * ...
*
Mucoadhesion Mucoadhesion describes the attractive forces between a biological material and mucus or mucous membrane. Mucous membranes adhere to epithelial surfaces such as the gastrointestinal tract (GI-tract), the vagina, the lung, the eye, etc. They are gener ...
* Mucophagy *
Sniffle A sniffle is the instinctive action of inhaling quickly in order to prevent mucus from flowing from one's nasal cavity, as an alternative to blowing the nose. Physiology For a fraction of a second, the performer inhales strongly, pulling mucus ...
* Spinnbarkeit * Feces


Notes


References

{{Authority control Body fluids Exocrine system Symptoms and signs: Respiratory system Excretion