ADP-ribosylation raises a handful of sensible questions. This page answers them in order, starting with the fundamentals and moving to applications.
This page was last updated on 2025-09-07 and is reviewed periodically as new material appears.
Related compounds include NADH, the reduced form, and NADP+, which carries an additional phosphate group. NADP+ and NADPH often serve in biosynthetic and antioxidant reactions, while NAD+ and NADH are more associated with energy-yielding catabolism. Nicotinamide, nicotinic acid, and nicotinamide riboside are precursors that can enter salvage pathways. The exact contribution of dietary precursors to tissue NAD+ pools is an area of active investigation. Some studies measure labeled precursors to trace those routes.
NAD+ is the oxidized form of nicotinamide adenine dinucleotide, a coenzyme built from two nucleotides joined by a phosphate linkage. One nucleotide carries adenine, and the other carries nicotinamide; the plus sign denotes a formal positive charge on the nicotinamide ring, not a free proton. In cells, NAD+ and its reduced partner NADH form a reversible redox pair. That pair participates in electron transfer reactions throughout metabolism. The abbreviation NAD+ is common in biochemistry, while NAD(H) sometimes denotes the combined pool.
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 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.
| Property | Value | Notes |
|---|---|---|
| Chemical name | Nicotinamide adenine dinucleotide (oxidized form) | NAD+ denotes the oxidized redox state |
| Common synonyms | Diphosphopyridine nucleotide; coenzyme I | Older names appear in historical literature |
| Molar mass | About 663.43 g/mol | Free acid value; salts and hydrates differ |
| Appearance | White to off-white powder | The purified solid is white; solutions are clear |
| Solubility | Highly soluble in water | Aqueous buffers are common laboratory solvents |
Quality control for NAD+ materials typically combines identity, purity, and water content checks. Identity may be confirmed by ultraviolet spectrum, retention time in chromatography, or mass accuracy, while purity is assessed by HPLC peak area or quantitative nuclear magnetic resonance. Residual water and solvents can affect molar calculations and enzyme assays, so Karl Fischer titration or thermogravimetric analysis may be used. Commercial materials vary in grade and counterion form, and published methods should specify the exact salt or hydrate when reporting concentrations. Regulatory status depends on intended use, with research reagents, dietary ingredients, and clinical products treated under different frameworks.
Quantification of NAD+ in biological samples usually relies on separation techniques coupled to sensitive detection. High-performance liquid chromatography with ultraviolet detection can measure the oxidized form by its absorbance near 260 nm, while mass spectrometry provides greater specificity and can distinguish NAD+ from close analogs. Enzymatic cycling assays use coupled dehydrogenase reactions to amplify signal and estimate NAD+ concentrations in cell or tissue extracts. Because NAD+ and NADH interconvert rapidly, sample preparation must quench metabolism quickly and preserve the redox state before analysis.
Beyond redox chemistry, NAD+ acts as a substrate for several enzyme families. ADP-ribosyltransferases, sirtuins, and CD38 ectoenzymes cleave the molecule into nicotinamide and ADP-ribose or related products. These reactions connect NAD+ availability to processes such as DNA repair, chromatin modification, and calcium signaling. Because the coenzyme is used in both electron transfer and signaling, cells maintain separate pools in compartments including the cytosol, mitochondria, and nucleus. The relative sizes and regulation of those pools remain active areas of study.
Cells produce NAD+ through several biosynthetic routes. The salvage pathway recycles nicotinamide, while the Preiss-Handler pathway uses nicotinic acid, and a de novo route can start from tryptophan in some organisms. In mammals, the salvage pathway is generally considered the main source under ordinary conditions. Tissue concentrations vary widely by cell type and compartment, and measured declines with age have been reported in some studies. Whether such changes drive aging or mainly accompany it remains an open question.
In humans, NAD+ can be synthesized from nicotinic acid, nicotinamide, nicotinamide riboside, and tryptophan through overlapping pathways. The salvage pathway recycles nicotinamide back to NAD+ and is often considered a major route in many tissues. Dietary precursors and intracellular recycling both contribute to the pool, but the quantitative importance of each source remains an active research question. NAD+ levels are not uniform across organs or cell compartments. Measurements in blood do not necessarily reflect concentrations inside tissues.
NAD+ is a dinucleotide composed of nicotinamide, ribose, and adenine linked by phosphate groups. Its full name is nicotinamide adenine dinucleotide, with "+" denoting the oxidized form. The molecule acts as a coenzyme in redox reactions, cycling between NAD+ and NADH. In cells, it participates in electron transfer during glycolysis, the citric acid cycle, and oxidative phosphorylation. It is distinct from NADP+, which carries an additional phosphate group and supports different biosynthetic reactions.
Glucose is converted to glucose-6-phosphate catalyzed by the enzyme hexokinase. Fructose-6-phosphate is converted to fructose 1,6-bisphosphate. This reaction is catalyzed by phosphofructokinase. Glyceraldehyde 3-phosphate is again phosphorylated to give 1,3-bisphosphoglycerate. This reaction is catalyzed by glyceraldehyde-3-phosphate dehydrogenase (GAPDH).
=== Affinity chromatography === Affinity chromatography is a method of separating biochemical mixtures, based on a highly specific biologic interaction such as that between antigen and antibody, enzyme and substrate, or receptor and ligand. The method was subsequently adopted for a variety of other techniques. Specific uses of affinity chromatography include antibody affinity, Immobilized metal ion affinity chromatography and purification of recombinant proteins - possibly the most common use of the method. To purify, proteins are tagged e.g. using His-tags or GST (glutathione-S-transferase) tags, which can be recognized by a metal ion ligand, such as imidazole. In 1971, Wilchek and colleagues applied this method to show that protein kinase is composed of regulatory and catalytic subunits. In 1972, Wilchek showed that the method can be used to remove toxic compounds from blood, as exemplified by the removal of heme peptides from blood using immobilized human serum albumin, thus laying the grounds for modern hemoperfusion
=== Pregnancy and breastfeeding === Hypothyroidism is common among pregnant women. A nationwide cohort study showed that 1.39% of all pregnant women in 2010 in Denmark received a prescription of levothyroxine during pregnancy. According to the U.S. Food and Drug Administration pregnancy categories, levothyroxine has been assigned category A. Given that no increased risk of congenital abnormalities has been demonstrated in pregnant women taking levothyroxine, therapy should be continued during pregnancy. Furthermore, treatment should be immediately administered to women diagnosed with hypothyroidism during pregnancy, as hypothyroidism is associated with a higher rate of complications, such as spontaneous abortion, preeclampsia, and premature birth. Thyroid hormone requirements increase during and last throughout pregnancy. As such, pregnant women are recommended to increase to nine doses of levothyroxine each week, rather than the usual seven, as soon as their pregnancy is confirmed. Repeat thyroid function tests should be done five weeks after the dosage is increased. While a minimal amount of thyroid hormones is found in breast milk, the amount does not influence infant plasma thyroid levels. Furthermore, levothyroxine was not found to cause any adverse events to the infant or mother during breastfeeding. Since adequate thyroid hormone concentrations are required to maintain normal lactation, appropriate levothyroxine doses should be administered during breastfeeding.
One rationale was to amputate as close to the damage as possible to save as much of the remaining limb as possible. Others argued to amputate in a location that would fit a prosthetic the best. This was especially debated for lower leg amputation because a prosthetic would fit better if the leg was amputated closer to the knee. Paré is the first known surgeon to decide the location of an amputation based on how a prosthetic would fit on it in 1564.
Sources: en.wikipedia.org
Some stegosaurs such as Kentrosaurus do not have a distinct "thagomizer" consisting of two pairs of spikes, as in these species there is not clear differentiation between the last two pairs on the tail and the other tail osteoderms, which are also spike-like, unlike the broad plate-like osteoderms on the back of Stegosaurus. There has been debate about whether the thagomizer was used simply for display, as posited by Gilmore in 1914, or used as a defensive weapon. Robert Bakker noted that it is likely that the stegosaur tail was much more flexible than those of other ornithischian dinosaurs because it lacked ossified tendons, thus lending credence to the idea of the thagomizer being a weapon. He also observed that Stegosaurus could have maneuvered its rear easily by keeping its large hindlimbs stationary and pushing off with its very powerfully muscled but short forelimbs, allowing it to swivel deftly to deal with attack. In 2010, analysis of a digitized model of Kentrosaurus aethiopicus showed that the tail could bring the thagomizer around to the sides of the dinosaur, possibly striking an attacker beside it.
The first documented use of electrical stimulation for pain relief dates back to 46 AD when Scribonius Largus of the ancient Roman Empire used the electric properties of torpedo fish to relieve headaches. Scientific studies of neuromodulation began in 1745, when German physician De Haen published "a number of cases of spasmodic, paralytic and other nervous affections cured by electricity". The first implementation of electrocutical apparatus in hospital medical treatment recorded in Middlesex Hospital of London in 1767. In 1780, Italian physicist and biologist Luigi Galvani investigated how electrical signals in nerves could control muscle movement, further evidence of the electrical nature of the neuronal activity.
== Further reading == Breker, Arno (1970). Patis, Hitler et moi. Paris: Presses de la Cité. Giesler, Hermann (1977). Ein anderer Hitler: Bericht seines Architekten: Erlebnisse, Gespräche, Reflexionen (2nd ed.). Leoni am Starnberger See: Druffel. ISBN 978-3-8061-0820-0. Larsson, Lars Olof (1998). Albert Speer: Plan de Berlin, 1937-1943. [S.I.]: Aam. ISBN 978-2-87143-034-6. O'Donnell, James (1978). The Bunker. New York: Da Capo Press. ISBN 978-0-306-80958-3. Scobie, Alexander (1990). Hitler's State Architecture: The Impact of Classical Antiquity. University Park: Pennsylvania State University Press. ISBN 978-0-271-00691-8. Speer, Albert (1996). Erinnerungen. Frankfurt am Main: Ullstein. ISBN 978-3-550-07616-9. Speer, Albert (1970). Inside The Third Reich. New York: Macmillan. ISBN 978-0-380-00071-5.
A newer process, solid oxide membrane technology, involves the electrolytic reduction of MgO. At the cathode, Mg2+ ion is reduced by two electrons to magnesium metal. The electrolyte is yttria-stabilized zirconia (YSZ). The anode is a liquid metal. At the YSZ/liquid metal anode O2− is oxidized. A layer of graphite borders the liquid metal anode, and at this interface carbon and oxygen react to form carbon monoxide. When silver is used as the liquid metal anode, there is no reductant carbon or hydrogen needed, and only oxygen gas is evolved at the anode. It was reported in 2011 that this method provides a 40% reduction in cost per pound over the electrolytic reduction method.
=== Human civilization: a model for extrapolation === Kardashev poses the following question: "Is it possible to describe the development of a civilization in general terms over large cosmological periods?" Now many of the fundamental parameters that characterize the development of civilization on Earth are growing exponentially. In the field of energy, astronomer Don Goldsmith estimated that the Earth receives about one billionth of the Sun's energy, and that humans use about one millionth of it. So we consume about one millionth of a billionth of the Sun's total energy. Since human expansion is exponential, we can determine how long it will take for humanity to go from Type II to Type III according to Michio Kaku. Thus, the rate of development of our own world remains the only criterion for extrapolating the state of civilizations older than humanity. The same is true for social values and basic needs according to Ashkenazi. Therefore, the time to double technical knowledge is about 10 years, and to double energy output, available reserves, and population is about 25 years. Two scenarios are then possible: spatial expansion or energy stagnation, the latter being possible only for 125 years, according to Kardashev, using the following relationship
Sources: en.wikipedia.org
Portugal has a literary tradition that predates the Portuguese language going back into the early 13th century. Portuguese literature developed through song as well as the written page known as cantigas. The cantigas drew practitioners from all social ranging from King Denis I to Martin Codax who was a minstrel. The earliest known work of literature produced by a Portuguese is the Ora faz ost'o senhor de Navarra, a cantiga de escárnio e maldizer written in Galician–Portuguese by João Soares de Paiva at around the year 1200. Portuguese literature developed under the influence of both European geopolitical developments and broader European literary traditions. The Hundred Years' War helped foster the development of Portuguese chronicles by Fernão Lopes, which constitute a valuable record of some of Europe's early encounters with peoples beyond the continent. European medieval chivalric literature, together with didactic religious literature transmitted through adaptations and partial translations, contributed to the development of Portuguese poetry in the translated works of Norman French Arturian narratives. Portuguese literature flourished during the Age of Discovery with writers such as Luís Vaz de Camões and António Ferreira. Modern Portuguese literature took shape through the work of Almeida Garrett, one of the founders of Portuguese Romanticism. Portugal has one Nobel Prize–winning author—José Saramago (1998).
=== Taxonomic paradigm === The two species of the Cannabis genus that are most commonly grown are Cannabis indica and Cannabis sativa. A third species, Cannabis ruderalis, is very short and produces only trace amounts of tetrahydrocannabinol (THC), and thus is not commonly grown for industrial, recreational or medicinal use. However, because Cannabis ruderalis flowers independently of the photoperiod and according to age, it has been used to breed autoflowering strains. Pure sativas are relatively tall (reaching as high as 4.5 meters), with long internodes and branches, and large, narrow-bladed leaves. Pure indica varieties are shorter and bushier, with wider leaflets. They are often favored by indoor growers for their size. Sativas bloom later than indicas, often taking a month or two longer to mature. The subjective effects of sativas and indicas are said to differ, but the ratio of tetrahydrocannabinol (THC) to cannabidiol (CBD) in most named drug varieties of both types is similar (averaging about 200:1). Unlike most commercially developed strains, indica landraces exhibit plants with varying THC/CBD ratios. The informal designation sativa and indica may have various, controversial meanings. Morphologically, the name sativa designates tall plants with narrow leaves, while indica refers to short plants with wide leaves. Among the marijuana community however, sativa rather refers to equatorial varieties producing stimulating psychoactive effects, whereas indica-type plants from Central Asia are used for relaxing and sedative drugs.
Described retrospectively as an "outlier", the album had been made by a group in disagreement over method and taste. The more rhythm-and-blues-oriented Haskell and McCulloch both found the music difficult to relate to, and tedious and confusing to record. Collins disliked how his parts were composed, while both Fripp and Haskell detested Sinfield's lyrics. This line-up of the band did not survive much longer than the Lizard recording sessions. Haskell quit the band acrimoniously during initial tour rehearsals, after refusing to sing live with distortion and electronic effects on his voice, and McCulloch departed soon after. With Sinfield not being a musician and Fripp having seemingly given up on the band, Collins was left to search for new members.
=== French Mandate === After the fall of the Ottoman Empire in 1918, the Syrian National Congress was convened in May 1919 in Damascus. In September 1920, Henri Gouraud, High Commissioner of the Levant, formed a representative council, with two-thirds elected and one-third appointed by the French administration. On 28 June 1922, the Syrian Federation was established, creating a Federation Council of 15 members from various states. Due to the lack of elections, these members were appointed by the High Commissioner in 1923, and their terms were extended the following year. In 1925, after the formation of the State of Syria, President Ahmad Nami and High Commissioner Henri Ponsot agreed to hold elections for a constituent assembly to draft a constitution. This led to the first Syrian legislative elections in 1928, which elected 68 representatives but was later disbanded on 5 February 1929. Article 30 of the 1930 Syrian constitution established a legislative authority known as the Chamber of Deputies, with representatives elected for four-year terms. The first elections for the Chamber of Deputies were held in December 1931 and January 1932. The first council met in June 1932 and facilitated a compromise that led to Muhammad Ali Bey al-Abid's presidency. In the 1936 elections, the National Bloc won the majority of seats in the Chamber, and Hashim al-Atassi was elected president. Concurrently, negotiations with France led to the independence treaty, ratified by the Chamber of Deputies in December 1936.
Sources: en.wikipedia.org
It indicates a formal positive charge on the nicotinamide ring. The molecule is not simply a protonated acid, and the charge is part of its redox chemistry.
NAD+ is the oxidized form, while NADH is the reduced form carrying two additional electrons and a proton. The two forms interconvert in many metabolic reactions.
No. NADP+ contains an extra phosphate group on the adenine ribose. NADP+ and NADPH tend to participate in different biosynthetic and antioxidant pathways.
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.