This is a working overview of ADP-ribosylation, written for readers who want more than a one-paragraph summary but less than a textbook.
Reviewed 2025-08-15. Anything still debated is marked as such rather than presented as settled.
Beyond electron transfer, NAD+ serves as a substrate for enzymes that cleave the molecule and transfer ADP-ribose units. Sirtuins, poly(ADP-ribose) polymerases, and CD38 consume NAD+ in regulatory reactions. These activities link NAD+ availability to DNA repair, chromatin modification, calcium signaling, and metabolic stress responses. Because consumption can exceed biosynthesis under some conditions, cellular NAD+ levels are dynamic rather than fixed. Enzyme affinity and local synthesis also influence how much NAD+ is available for signaling.
NAD+ is a dinucleotide composed of adenine, ribose, and nicotinamide moieties linked by phosphate groups. Its oxidized form carries a positive charge on the nicotinamide ring, which enables reversible hydride transfer. The molecule functions as a coenzyme in oxidoreductase reactions rather than as a dietary vitamin in its intact form. Cells maintain separate pools in cytoplasm, mitochondria, and nucleus. This compartmentalization allows distinct redox environments while preserving a shared chemical identity.
In glycolysis, NAD+ accepts electrons during the oxidation of glyceraldehyde-3-phosphate, forming NADH. The tricarboxylic acid cycle and fatty acid oxidation also generate NADH, which donates electrons to the mitochondrial electron transport chain. This flow supports ATP synthesis and helps maintain the redox balance of the cell. Other dehydrogenases use NAD+ as a cofactor for biosynthetic reductions and detoxification reactions. NADH is later reoxidized to sustain continued flux through these pathways.
Beyond electron transfer, NAD+ serves as a substrate for enzymes that cleave it and attach its ADP-ribose portion to other molecules. This group includes poly(ADP-ribose) polymerases, CD38, and sirtuins. Such reactions consume NAD+ and can influence its availability for metabolism. Cells replenish NAD+ through a salvage pathway that recycles nicotinamide and through routes starting from tryptophan or vitamin B3 forms. How these synthesis and consumption routes are coordinated across tissues remains an active area of study, and compartment-specific concentrations are difficult to measure directly.
Nicotinamide adenine dinucleotide, abbreviated NAD+, is a dinucleotide built from adenine, nicotinamide, two ribose sugars, and two phosphate groups. The oxidized form carries a positive charge on the nicotinamide ring, while the reduced form, NADH, is neutral. This pair acts as a reversible electron carrier in cells. NAD+ is present in bacteria, plants, animals, and fungi. Its structure allows it to accept and donate electrons without being consumed in the reactions it supports.
| Property | Value | Notes |
|---|---|---|
| Chemical name | Nicotinamide adenine dinucleotide | Oxidized form abbreviated NAD+ |
| Molecular formula | C21H27N7O14P2 | Free acid form |
| Molar mass | 663.43 g/mol | Calculated for free acid |
| CAS Registry Number | 53-84-9 | Common entry for beta-NAD+ |
| Appearance | White to off-white powder | Hygroscopic solid |
In cells, NAD+ functions primarily as an electron carrier. Dehydrogenase enzymes in glycolysis and the citric acid cycle transfer hydride from substrates to NAD+, producing NADH. NADH then delivers electrons to the mitochondrial respiratory chain, supporting ATP synthesis. In fermentation, NADH is reoxidized to NAD+ so that glycolysis can continue. The balance between NAD+ and NADH helps set metabolic flux. Beyond redox, NAD+ serves as a substrate for enzymes that cleave it, including sirtuins, poly(ADP-ribose) polymerases, and CD38. These reactions consume NAD+ and release nicotinamide and ADP-ribose products.
Biosynthesis occurs through salvage, Preiss-Handler, and de novo pathways. In mammals, the salvage pathway from nicotinamide predominates, and NAMPT is often described as rate-limiting. Nicotinamide riboside and nicotinic acid enter related routes that converge on NAD+ production. Tissue NAD+ concentrations vary widely and are maintained by a balance of synthesis and consumption. Some studies report age-related declines in certain tissues, but whether these changes cause disease or can be reversed to improve human health remains an open question.
NAD+ stands for nicotinamide adenine dinucleotide, the oxidized form of a coenzyme found in all living cells. The molecule consists of two nucleotides, adenine and nicotinamide ribose, joined through phosphate groups. Its chemical formula is C21H27N7O14P2, and the free acid has a molar mass near 663.43 grams per mole. In redox reactions, NAD+ accepts a hydride ion and becomes NADH. The pair NAD+ and NADH participates in hundreds of metabolic reactions, including steps in glycolysis, the citric acid cycle, and oxidative phosphorylation.
Biosynthesis of NAD+ starts from nicotinamide, nicotinic acid, or nicotinamide riboside through salvage pathways. A rate-limiting enzyme, nicotinamide phosphoribosyltransferase, converts nicotinamide to nicotinamide mononucleotide. Further coupling with ATP yields NAD+. In mammals, the liver and muscle can synthesize NAD+ from dietary precursors, but tissue levels vary widely. Researchers study these pathways to understand age-related changes, metabolic disorders, and neurodegeneration. Direct causal links between NAD+ decline and disease remain an active area of investigation.
NAD+ is a dinucleotide composed of adenine, ribose, and nicotinamide groups joined by phosphate linkages. It serves as a coenzyme in oxidoreductase reactions, cycling between oxidized NAD+ and reduced NADH. The molecule is water-soluble and occurs in all living cells. Its nicotinamide ring accepts hydride ions during catabolic reactions, linking substrate oxidation to electron transport. This redox couple supports ATP production and helps maintain cytosolic and mitochondrial redox balance in many cell types.
A submarine can have a variety of sensors, depending on its missions. Modern military submarines rely almost entirely on a suite of passive and active sonars to locate targets. Active sonar relies on an audible "ping" to generate echoes to reveal objects around the submarine. Active systems are rarely used, as doing so reveals the sub's presence. Passive sonar is a set of sensitive hydrophones set into the hull or trailed in a towed array, normally trailing several hundred feet behind the sub. The towed array is the mainstay of NATO submarine detection systems, as it reduces the flow noise heard by operators. Hull mounted sonar is employed in addition to the towed array, as the towed array can not work in shallow depth and during maneuvering. In addition, sonar has a blind spot "through" the submarine, so a system on both the front and back works to eliminate that problem. As the towed array trails behind and below the submarine, it also allows the submarine to have a system both above and below the thermocline at the proper depth; sound passing through the thermocline is distorted resulting in a lower detection range. Global climate change and warmer oceans may complicate detecting submarines at depth in most places in the world. Submarines also carry radar equipment to detect surface ships and aircraft. Submarine captains are more likely to use radar detection gear than active radar to detect targets, as radar can be detected far beyond its own return range, revealing the submarine. Periscopes are rarely used, except for position fixes and to verify a contact's identity.
=== Xen Museum === The Crowbar Collective released an expansion in April 2021 called Xen Museum that presents a virtual museum that documents the team's past five years of effort in creating Black Mesa and mostly their work in recreating Xen from the original Half-Life.
3,4-Methylenedioxymethamphetamine (MDMA, ecstasy, or molly) is an entactogen, euphoriant, and psychostimulant of the amphetamine class. Briefly used by some psychotherapists as an adjunct to therapy, the drug became popular recreationally and, in the United States, the DEA listed MDMA as a Schedule I controlled substance, prohibiting most medical studies and applications. MDMA is known for its entactogenic properties. The stimulant effects of MDMA include hypertension, anorexia (appetite loss), euphoria, social disinhibition, insomnia (enhanced wakefulness/inability to sleep), improved energy, increased arousal, and increased perspiration, among others. Relative to catecholaminergic transmission, MDMA enhances serotonergic transmission significantly more, when compared to similar psychostimulants like amphetamine. Due to the relative safety of MDMA, some researchers such as David Nutt have criticized the scheduling level, writing a satirical article finding MDMA to be 28 times less dangerous than horseriding, a condition he termed "equasy" or "Equine Addiction Syndrome".
== Definition == A variety of definitions have been proposed throughout the years, but the simplest and most recent definition of fermentation proposed is "catabolism where organic compounds are both the electron donor and acceptor." This definition distinguishes fermentation from aerobic respiration (when oxygen is the acceptor) and types of anaerobic respiration (when an inorganic species is the acceptor). However, this definition does not encompass all forms of fermentation. For example, propionate fermentation which uses H2 as an electron donor, or the second step of butyrate fermentation where CO2 can act as an electron acceptor. Thus, it is simplest to use this definition while acknowledging that protons can also be used as electron donors and CO2 as acceptors. In 1876, before the discovery of anaerobic respiration, Louis Pasteur described it as "la vie sans air" (life without air). It was also common for fermentation to be defined based on how fermentation forms ATP which was catabolism that forms ATP through only substrate-level phosphorylation. Industrial fermentation is another type of fermentation that is defined loosely as a large-scale biological manufacturing process; however, this definition focuses on the process of manufacturing rather than metabolic details.
The last RAF Mustang Mk I and Mustang Mk II aircraft were struck off charge in 1945. Army Co-operation Command used the Mustang's superior speed and long range to conduct low-altitude "Rhubarb" raids over continental Europe, sometimes penetrating German airspace. The V-1710 engine ran smoothly at 1,100 rpm, versus 1,600 for the Merlin, enabling long flights over water at 50 ft (15 m) altitude before approaching the enemy coastline. Over land, these flights followed a zig-zag course, turning every six minutes to foil enemy attempts at plotting an interception. During the first 18 months of Rhubarb raids, RAF Mustang Mk.Is and Mk.Ias destroyed or heavily damaged 200 locomotives, over 200 canal barges, and an unknown number of enemy aircraft parked on the ground, for a loss of eight Mustangs. At sea level, the Mustangs were able to outrun all enemy aircraft encountered. The RAF gained a significant performance enhancement at low altitude by removing or resetting the engine's manifold pressure regulator to allow overboosting, raising output as high as 1,780 horsepower at 70 in Hg. In December 1942, Allison approved only 1,570 horsepower at 60 in Hg manifold pressure for the V-1710-39. The RAF later operated 308 P-51Bs and 636 P-51Cs, which were known in RAF service as Mustang Mk IIIs; the first units converted to the type in late 1943 and early 1944. Mustang Mk III units were operational until the end of World War II, though many units had already converted to the Mustang Mk IV (P-51D) and Mk IVa (P-51K) (828 in total, comprising 282 Mk IV and 600 Mk IVa).
Sources: en.wikipedia.org
== Potentiality for use == Since the very first studies that gave insights into the properties of hydrophobins, these small proteins have been regarded as great candidates for technological use. The detailed understanding of the molecular mechanisms underlying hydrophobin self-assembly into amphipathic monolayer in hydrophobic:hydrophilic interfaces is of great academic interest but mainly of commercial interest. This is because a deep understanding of the elements driving these mechanisms would allow engineering of hydrophobins (or other biomolecules) for nano and biotechnological applications. An example is that the hydrophobin-coating of carbon nanotubes was found to increase their solubility and reduce their toxicity, a finding that increases the prospects of carbon nanotubes to be used as vehicles for drug delivery. Other areas of potential use of hydrophobins include:
== History == Biocon was formed in 1978 as Biocon India Private Limited, an India-based partner to the existing Irish multinational brewing enzyme producer Biocon Biochemicals Ltd., with US$10,000 as startup capital from Mazumdar-Shaw. The subsidiary initially produced enzymes for the brewing market such as the papaya enzyme papain and the fish collagen extract Iisinglass, a clarifying agent used for some beers and wines. In 1979, Biocon became the first Indian company to manufacture and export enzymes to the US and Europe, and spent the 1980s producing an increasing share of pharmaceuticals for the domestic market.
=== Coagulation factor disorders === The best-known coagulation factor disorders are the hemophilias. The three main forms are hemophilia A (factor VIII deficiency), hemophilia B (factor IX deficiency or "Christmas disease") and hemophilia C (factor XI deficiency, mild bleeding tendency). Von Willebrand disease (which behaves more like a platelet disorder except in severe cases), is the most common hereditary bleeding disorder and is characterized as being inherited autosomal recessive or dominant. In this disease, there is a defect in von Willebrand factor (vWF), which mediates the binding of glycoprotein Ib (GPIb) to collagen. This binding helps mediate the activation of platelets and formation of primary hemostasis. In acute or chronic liver failure, there is insufficient production of coagulation factors, possibly increasing risk of bleeding during surgery. Thrombosis is the pathological development of blood clots. These clots may break free and become mobile, forming an embolus or grow to such a size that occludes the vessel in which it developed. An embolism is said to occur when the thrombus (blood clot) becomes a mobile embolus and migrates to another part of the body, interfering with blood circulation and hence impairing organ function downstream of the occlusion. This causes ischemia and often leads to ischemic necrosis of tissue. Most cases of venous thrombosis are due to acquired states (older age, surgery, cancer, immobility).
== Principles and ethical foundations == Across its applications, harm reduction prioritizes reducing adverse consequences without requiring elimination of the underlying behavior. In drug policy, this orientation is commonly described as pragmatic: it begins from the continued existence of drug use and regards reductions in harm as worthwhile outcomes even when abstinence is not achieved. Ethical defenses have also drawn on consequentialist and rights-based reasoning. Consequentialist arguments emphasize reductions in illness, death, and social costs, while rights-based arguments appeal to autonomy and opposition to paternalism. The harm can be distinguished in physical harms, dependence and social harms including health care costs and community outcomes. Social justice approaches broaden the analysis of harm beyond individual behavior to include poverty, homelessness, criminalization, violence, stigma, and barriers to health care. From this perspective, harm reduction may respond to structural inequities as well as immediate individual risk. Relational and participatory approaches emphasize that people who use drugs possess knowledge relevant to effective services and should participate in their design and governance. User-led organizations and peer practices have been important within this tradition. Ethical analysis of harm-reduction research has also used communitarian approaches, emphasizing social relationships and responsibilities alongside individual rights.
Sources: en.wikipedia.org
== Reception and legacy == The series was well received by viewers. Within a week after uploading, episode one was featured by several well-established internet magazines and blogs such as Destructoid, Kotaku, and Boing Boing. Video game journalist Justin McElroy praised it as "an awesome piece of classic fan fiction written by a nine-year-old and then animated by a group of evil geniuses." Within a year it had received over 1 million views. The popularity of Half-Life: Full Life Consequences inspired other spinoff works, many adapting other Squirrelking fan fiction. Although the fan fiction author Squirrelking was often cited as being a young child, it was later revealed to be a hoax account with the goal of making something "so mind-numbingly bad that it stands the test of time as one of the worst things ever written." In 2019, the series was showcased during the 12 Days of Garry's Mod event on the official Garry's Mod website.
== Clinical significance == Levels of the hormone prolactin within the body can indicate various conditions. Normal prolactin levels support reproductive and metabolic functions, while elevated prolactin levels (hyperprolactinemia), and low prolactin levels (hypoprolactinemia) can indicate an underlying medical disorder. Levels of prolactin within the body can also be affected by factors such as pregnancy, stress, and certain medications. Due to the widespread distribution of prolactin receptors across the body’s organs and tissues, elevated levels of prolactin can simultaneously influence multiple organ systems of the body. While prolactin cell levels rise during pregnancy and lactation, they can also rise due to stress, pain, exercise, sexual intercourse, and food consumption. Hyperprolactinemia (elevated prolactin levels) can be caused by excessive thyrotropin-releasing hormone production or decreased dopamine levels. This may cause symptoms in both males and females such as infertility, erectile dysfunction, and irregular periods. Excessive release of prolactin can be due to a prolactinoma, a tumor of the pituitary gland. The tumor itself may not cause symptoms, but may cause an increased production of prolactin and decreased levels of estrogen and testosterone. Symptoms due to a prolactinoma may include infertility, a decrease in sexual desire, and osteoporosis. In men, symptoms may include erectile dysfunction, enlarged breast tissue, and decreased body hair. In women, symptoms may include breast discharge, irregular menstrual cycles, acne, and increased body hair.
The share of the world's lead production by the Eastern Bloc increased from 10% to 30%, from 1950 to 1990, with the Soviet Union being the world's largest producer during the mid-1970s and the 1980s, and China starting major lead production in the late 20th century. Unlike the European communist countries, China was largely unindustrialized by the mid-20th century; in 2004, China surpassed Australia as the largest producer of lead. As was the case during European industrialization, lead has had a negative effect on health in China.
Periodic acid-Schiff is a histology special stain used to mark carbohydrates (glycogen, glycoprotein, proteoglycans). PAS is commonly used on liver tissue where glycogen deposits are made which is done in efforts to distinguish different types of glycogen storage diseases. PAS is important because it can detect glycogen granules found in tumors of the ovaries and pancreas of the endocrine system, as well as in the bladder and kidneys of the renal system. Basement membranes can also show up in a PAS stain and can be important when diagnosing renal disease. Due to the high volume of carbohydrates within the cell wall of hyphae and yeast forms of fungi, the Periodic acid -Schiff stain can help locate these species inside tissue samples of the human body.
=== Endocrine === Dexamethasone is the treatment for the very rare disorder of glucocorticoid resistance. In adrenal insufficiency and Addison's disease, dexamethasone is prescribed when the patient does not respond well to prednisone or methylprednisolone. It can be used in congenital adrenal hyperplasia in older adolescents and adults to suppress adrenocorticotropic hormone (ACTH) production. It is typically given at night.
Sources: en.wikipedia.org
NAD+ is the oxidized form, while NADH is the reduced form carrying an additional hydride equivalent. The pair participates in reversible electron transfer reactions. Their ratio helps indicate the redox state of a compartment.
NAD+ itself is not classified as a vitamin, but its precursor niacin is an essential nutrient in humans. Cells synthesize NAD+ from niacin, nicotinamide, nicotinamide riboside, or tryptophan. The intact dinucleotide is not obtained directly from typical diets in meaningful amounts.
Age-related studies often examine whether NAD+ levels decline in tissues and whether that decline affects mitochondrial function or DNA repair. Interventions using precursor molecules raise open questions about cause and effect. Current evidence does not establish that changing NAD+ levels slows human aging.
It indicates the oxidized form, which has a positive charge on the nicotinamide nitrogen. The reduced partner NADH lacks that charge and carries added electrons. The plus sign is part of the standard abbreviation, not a separate ion.