If you have been reading about thiol and want a single page that covers the useful parts, this is it: definitions, context, how it is studied, and the questions that come up repeatedly.
Last reviewed on 2025-09-22. Where a claim depends on a specific study, the study is described rather than over-claimed.
In cells, glutathione helps maintain the reducing environment of the cytosol and supports enzymes that counteract reactive oxygen species. It acts as a cofactor for glutathione peroxidases, which reduce hydrogen peroxide and lipid peroxides, and for glutathione S-transferases, which conjugate electrophiles. The ratio of GSH to GSSG is often used as an indicator of oxidative stress, although the ratio can vary by compartment and cell type. Glutathione also stores cysteine, an amino acid that can be limiting for protein synthesis and antioxidant defense.
Synthesis occurs in two ATP-dependent steps. The enzyme glutamate-cysteine ligase joins glutamate and cysteine to form gamma-glutamylcysteine, and glutathione synthetase adds glycine. The first step is rate-limiting and is influenced by cysteine availability and feedback inhibition by GSH. Breakdown involves gamma-glutamyl transferase and subsequent peptidases, which release constituent amino acids for reuse. Because turnover differs among tissues, measurements from blood, plasma, and tissues are not directly interchangeable. Research continues to clarify how compartment-specific pools are regulated in health and disease.
Glutathione is a tripeptide composed of glutamate, cysteine, and glycine. It contains an unusual gamma-glutamyl bond between glutamate and cysteine, which resists cleavage by many peptidases. The reduced form, GSH, carries a thiol group on cysteine and is the dominant intracellular form in most cells. Its structure allows it to participate in redox reactions and to serve as a sulfur donor. The oxidized form, GSSG, consists of two GSH molecules joined by a disulfide bond.
Because GSH is central to redox balance, its status is studied in aging, liver disease, neurodegenerative conditions, and metabolic disorders. Observational studies often report lower GSH or higher GSSG in affected tissues, but such associations do not establish that raising glutathione changes disease outcomes. Oral glutathione is digested into amino acids, and whether intact absorption occurs remains debated; precursors such as N-acetylcysteine and cysteine donors are also investigated. Regulatory agencies generally treat glutathione as a dietary supplement, not an approved drug, and clinical claims require evidence from controlled trials.
Glutathione is a small tripeptide composed of glutamate, cysteine, and glycine, with the unusual gamma-glutamyl linkage between glutamate and cysteine. Its cysteine thiol group makes it a major non-enzymatic antioxidant in cells. The reduced form, GSH, predominates in most intracellular compartments, while the oxidized disulfide form, GSSG, is produced when GSH reduces reactive oxygen species. Intracellular concentrations often reach millimolar levels, whereas plasma concentrations are much lower, typically in the low micromolar range. This gradient reflects active synthesis, transport, and consumption rather than passive distribution.
Synthesis occurs in two ATP-dependent steps: glutamate-cysteine ligase joins glutamate and cysteine to form gamma-glutamylcysteine, and glutathione synthetase adds glycine to complete the tripeptide. The pathway is feedback-inhibited by GSH and limited by cysteine availability, so cysteine supply often constrains production. Once formed, GSH participates in redox buffering, xenobiotic conjugation, and protein glutathionylation. Glutathione peroxidase uses GSH to reduce hydrogen peroxide and lipid peroxides, yielding GSSG, while glutathione reductase regenerates GSH using NADPH. Glutathione S-transferases conjugate electrophiles to GSH, supporting detoxification and excretion.
| Property | Value | Notes |
|---|---|---|
| Chemical formula | C10H17N3O6S | For reduced glutathione; the oxidized dimer has two sulfur atoms. |
| Molar mass | 307.32 g/mol | Calculated for the reduced form. |
| Appearance | White to off-white crystalline powder | Typical for solid reagent; solutions are usually colorless. |
| Solubility | Freely soluble in water | Poorly soluble in nonpolar organic solvents. |
| Typical storage | -20 °C, desiccated, protected from light | Limits oxidation, moisture uptake, and degradation. |
In cells, glutathione exists mainly in a reduced form called GSH. When two GSH molecules react, they form oxidized glutathione, or GSSG, which contains a disulfide bond. The ratio of GSH to GSSG is often used as an indicator of oxidative stress. Enzymes such as glutathione peroxidase and glutathione reductase help cycle the molecule between these two states. This cycling supports antioxidant defense, detoxification of reactive molecules, and regulation of certain signaling pathways.
Glutathione is present in most tissues, with especially high concentrations in the liver. It also serves as a cofactor for some enzymes and helps transport amino acids across cell membranes. In plants and microorganisms, glutathione contributes to stress responses and metal handling. The molecule is synthesized in two ATP-dependent steps, first producing gamma-glutamylcysteine and then adding glycine. Because cysteine availability often limits synthesis, dietary and metabolic factors can influence glutathione levels. Research continues to examine how these levels relate to health and disease.
Functionally, glutathione supports redox balance by donating electrons and becoming oxidized. It also serves as a cofactor for enzymes such as glutathione peroxidases and glutathione S-transferases. These enzymes participate in peroxide reduction and in conjugation reactions that help process reactive molecules. Separate from antioxidant roles, glutathione can modify protein cysteines through S-glutathionylation, influencing enzyme activity and signaling. Research continues to examine how these chemical roles translate into whole-organism effects.
Glutathione is a tripeptide composed of glutamate, cysteine, and glycine. Its glutamate-cysteine linkage uses the gamma-carboxyl group of glutamate, a feature that resists standard peptidases. The cysteine residue provides a thiol group, which gives the molecule its reducing character. In cells, glutathione is often the most abundant small-molecule thiol, with concentrations varying widely by tissue and compartment. It exists mainly in a reduced form called GSH, while oxidation produces a disulfide-linked dimer called GSSG.
Interpreting glutathione measurements requires attention to pre-analytical variables. The GSSG concentration in a sample can rise artificially during storage or processing, making the GSH/GSSG ratio unreliable if not controlled. Reference ranges vary by specimen type, assay, and population, so comparisons across studies are difficult. Plasma glutathione is low and sensitive to hemolysis, while whole blood reflects primarily erythrocyte content. Many studies measure total glutathione rather than the reduced and oxidized forms separately, which limits conclusions about redox status.
Accurate measurement of glutathione begins with careful sample handling. Because GSH oxidizes rapidly to GSSG, samples must be processed quickly or frozen immediately. Acid precipitation with metaphosphoric acid or perchloric acid is common; it lowers pH, precipitates proteins, and helps preserve the reduced form. Chelating agents such as EDTA can limit metal-catalyzed oxidation. For whole blood, hemolysis releases glutathione from erythrocytes, so plasma and serum values differ substantially from whole blood values.
Gary Keith Ackers (1939–2011) was Emeritus Professor of Biochemistry and Molecular Biophysics of Washington University School of Medicine. His research focused on thermodynamic linkage analysis of biological macromolecules, addressing the molecular mechanism of cooperative O2 binding to human hemoglobin since the early 1970s. He was a Fellow of the Biophysical Society and one of the founders of the annual Gibbs Conference. Ackers invented agarose gel chromatography when he was a teenager. He went on the develop analytical gel chromatography methods for determinations of many important characteristics of water-soluble proteins; diffusion coefficient, molecular size, thermodynamics of protein-protein interactions including important changes due to single amino acid substitutions.
He also played a significant role in the winning elimination final against Western Bulldogs, but an injury sustained in the game ended his season. It was announced in early March 2021 that Ryder would temporarily step away from the club to spend time with his family and elders. Ryder received the full support of the club. Ryder returned to the club in April and played his first game for the season in the round seven win over Hawthorn in which he gathered five tackles, three clearances and 36 hitouts (significantly above the AFL average). Ryder had a purple patch in rounds 10-12, collecting 41, 34 and 34 hitouts in three games. Ryder also had a solid game in round 16 against Collingwood and champion Brodie Grundy, collecting 23 hitouts and kicking two goals. Ryder's last game for 2021 was Round 19, after which he battled an achilles injury which saw him miss the remainder of the season. After playing 12 of a possible 22 games and averaging over 30 hitouts a game, Ryder re-signed with the Saints for another season. After another injury-interrupted year for St Kilda during the side's 2022 season, Ryder announced his retirement on 20 August, ending a career of 281 games across three clubs.
In particular, decreased access to public health services such as syringe exchange programs and confiscation of syringes can precipitate a cascade of health harms. Geographic diffusion of epidemics from the northern border states elsewhere is also possible with the rotation of police and military personnel stationed in drug conflict areas with high infection prevalence. With increased drug use, there has been a parallel rise in demand for drug user treatment in Mexico.
== N == NEMS-MS – Nanoelectromechanical systems mass spectrometry NETD – Negative electron-transfer dissociation NICI – Negative ion chemical ionization NRMS – Neutralization reionization mass spectrometry
== History == The company (the Ranpak name standing for Random Packaging) traces its origins to a machine patented by George R. Johnson in 1970 that converted kraft paper into shock absorbing crinkled paper for use in cushioning auto-parts to prevent such items being damaged in shipment. Johnson, working with entrepreneur and engineer, Raymond Q. Armington, an inventor with several packing- and storage related patents to his name, incorporated the company in 1972. Armington became its chairman, a position he held until his death in 1993. In early 2002, David M. Gabrielsen was appointed president and CEO of Ranpak replacing Steven E. Armington, son of the co-founder. In December 1991 Ranpak opened its first foreign manufacturing center in Heerlen, Holland. The plant produced packing material and packing machines for the European market. Ranpak's acquisitions include:
Sources: en.wikipedia.org
==== S ==== Salami – cured sausage, fermented and air-dried meat, originating from one or a variety of animals. Historically, salami was popular among Southern European peasants because it can be stored at room temperature for periods of up to 30–40 days once cut, supplementing a possibly meager or inconsistent supply of fresh meat. Varieties of salami are traditionally made across Europe. Salumi – Italian cured meat products that are predominantly made from pork. It comes from the Italian word salume, pl. salumi "salted meat", derived from Latin sal "salt". The term salumi also encompasses bresaola, which is made from beef, and also cooked products such as mortadella and prosciutto cotto. Secca de bœuf – a type of dried salted beef made in Entrevaux. Similar to the Swiss Bindenfleisch, it is typically eaten as a starter. Skerpikjøt – a type of wind-dried mutton, is a delicacy of the Faroe Islands which is traditionally eaten at Christmas but also at other times of the year. Slinzega – a type of air-dried meat produced in Valtellina, in the Italian Alps. It is made in a similar manner to Bresaola, with smaller pieces of meat, which therefore bear a stronger taste. Smithfield ham – a specific form of the country ham, A 1926 Statute of Virginia (passed by the Virginia General Assembly) first regulated the usage of the term "Smithfield Ham".
Certain countries adopt immigration policies designed to be favourable towards members of diaspora communities with a connection to the country. For example, the Indian government confers Overseas Citizenship of India (OCI) status on foreign citizens of Indian origin to enable them to live and work indefinitely in India. OCI status was introduced in response to demands for dual citizenship by the Indian diaspora, particularly in countries with large populations of Indian origin. It was introduced by The Citizenship (Amendment) Act, 2005 in August 2005. Similar to OCI status, the UK Ancestry visa exempts members of the British diaspora from usual immigration controls. Poland issued the Karta Polaka to citizens of certain northeast European countries with Polish ancestry, but later expanded it to the worldwide Polonia. Some nations recognise a right of return for people with ancestry in that country or a connection to a particular ethnic group. A notable example of this is the right of Sephardi Jews to acquire Spanish nationality by virtue of their community's Spanish origins. Similar exemptions to immigration controls exist for people of Armenian origin seeking to acquire Armenian citizenship. Ghana similarly grants an indefinite right to stay in Ghana to members of the African diaspora, regardless of citizenship. Similarly, Israel maintains a policy permitting members of the Jewish diaspora to immigrate to Israel regardless of prior nationality.
==== Volatile organic compounds ==== Volatile organic compounds (VOCs) include a variety of chemicals, some of which may have short- and long-term adverse health effects. There are numerous sources of VOCs indoors, which means that their concentrations are consistently higher indoors (up to ten times higher) than outdoors. Some VOCs are emitted directly indoors, and some are formed through the subsequent chemical reactions that can occur in the gas-phase, or on surfaces. VOCs presenting health hazards include benzene, formaldehyde, tetrachloroethylene and trichloroethylene. VOCs are emitted by thousands of indoor products. Examples include: paints, varnishes, waxes and lacquers, paint strippers, cleaning and personal care products, pesticides, building materials and furnishings, office equipment such as copiers and printers, correction fluids and carbonless copy paper, graphics and craft materials including glues and adhesives, permanent markers, and photographic solutions. Chlorinated drinking water releases chloroform when hot water is used in the home. Benzene is emitted from fuel stored in attached garages. Human activities such as cooking and cleaning can also emit VOCs. Cooking can release long-chain aldehydes and alkanes when oil is heated and terpenes can be released when spices are prepared and/or cooked. Leaks of natural gas from cooking appliances have been linked to elevated levels of VOCs including benzene in homes in the USA. Cleaning products contain a range of VOCs, including monoterpenes, sesquiterpenes, alcohols and esters.
The DIT and other forms of digital mass analyzers scan ions by scanning the frequency of the drive waveform. The AC voltage is typically fixed during the scan. Digital devices use a duty cycle which allows them to operate completely independent of a DC voltage and without resonant excitation. When the DC voltage is zero the parameter a is also zero. Consequently, ion stability will depend on q. With these considerations it was possible to design a new type of stability diagram that is more suitable for planning and performing experiment. In 2014 Brabeck and Reilly created a stability diagram that maps the range of stable mass-to-charge ratios, m/z to the corresponding range of drive frequencies based on several user inputs. For a particular duty cycle, the operator can quickly reference the range of stable masses at each frequency of a scan. Fig 4 (a) and (b) shows the frequency-m/z stability diagram for a linear DIT with a duty cycle of d = 0.50 and d= 0.60 respectively.
Sources: en.wikipedia.org
Secondary radionuclides in a decay chain will occur in proportion to their half-lives, so short-lived ones will be very rare. For example, polonium can be found in uranium ores at a concentration about 1 part 1010 of uranium (0.1 mg per metric ton) by calculating the ratio of half-lives of polonium-210 to uranium-238, its ultimate parent.
This is a list of Nobel laureates who are Poles (ethnic) or Polish (citizenship). The Nobel Prize is a set of annual international awards bestowed on "those who conferred the greatest benefit on humankind", first instituted in 1901. Since 1903, there have been nineteen Poles who were awarded twenty Nobel Prizes. Poles have been the recipients of all Nobel prize categories: Physics, Chemistry, Physiology or Medicine, Literature, Peace and Economics.
=== Suppression of the pro-Alid groups === Mu'awiya died in 680 and was succeeded by his son Yazid I. Mu'awiya's designation of his son was an unprecedented act and shocked many in the Muslim community, particularly the Arab nobility of Kufa. They long sympathized with Caliph Ali, Mu'awiya's former rival, and Ali's family. One of Ali's sons, Husayn dispatched his cousin Muslim ibn Aqil to Kufa to set the stage for Husayn's accession to the caliphate. Ibn Aqil garnered significant support and was hosted by a prominent pro-Alid nobleman. Ubayd Allah became aware of Ibn Aqil's activities, prompting the latter to launch a premature assault against the governor. Ubayd Allah was holed up in his palace, but thirty men from his shurta (security forces) fended off Ibn Aqil's partisans, while he persuaded many Kufan noblemen to back him against Ibn Aqil, who was abandoned by his supporters and slain on 10 September 680. Husayn had already been en route to Kufa from Medina when he received news of Ibn Aqil's execution. Ubayd Allah was prepared for Husayn's arrival and sent troops to intercept him. They prevented Husayn and his small retinue from reaching the watered areas of the province. The two sides negotiated for weeks, but Ubayd Allah refused Husayn entry into Kufa or return to Arabia while Husayn refused to recognize Yazid's caliphate. In the end, a short battle was fought at Karbala on 10 October 680, in which Husayn and nearly all of his partisans were slain.
=== Conductive ink === In 2012 Vorbeck Materials started shipping the Siren anti-theft packaging device, which uses their graphene-based Vor-Ink circuitry to replace the metal antenna and external wiring to an RFID chip. This was the world's first commercially available product based on graphene.
The affinity, given as the dissociation constant (Kd), between a TCR and a pMHC was determined by surface plasmon resonance (SPR) to be in the range of 1–100 μM, with an association rate (kon) of 1000 -10000 M−1 s−1 and a dissociation rate (koff) of 0.01 -0.1 s−1. In comparison, cytokines have an affinity of KD = 10–600 pM to their receptor. It has been shown that even a single amino acid change in the presented peptide that affects the affinity of the pMHC to the TCR reduces the T-cell response and cannot be compensated by a higher pMHC concentration. A negative correlation between the dissociation rate of the pMHC-TCR complex and the strength of the T-cell response has been observed. That means, pMHC that bind the TCR for a longer time initiate a stronger activation of the T cell. Furthermore, T cells are highly sensitive; interaction with a single pMHC is enough to trigger activation. T cells move on quickly from antigens that do not trigger responses, rapidly scanning pMHC on an antigen-presenting cell (APC) to increase the chance of finding a specific pMHC. On average, a T cell encounters 20 APCs per hour. Different models for the molecular mechanisms that underlie this highly specific and highly sensitive process of antigen discrimination have been proposed. The occupational model simply suggests that the TCR response is proportional to the number of pMHC bound to the receptor. Given this model, a shorter lifetime of a peptide can be compensated by higher concentration such that the maximum response of the T cell stays the same.
Sources: en.wikipedia.org
GSH is the reduced form with a free thiol group, while GSSG is the oxidized disulfide-linked dimer. Most assays distinguish the two because their balance reflects redox conditions. The names are not interchangeable.
It is synthesized in the body from amino acids, so it is not classified as an essential dietary nutrient for most people. Dietary and supplemental forms are studied for their effects on tissue levels and health markers. Evidence varies by population and outcome.
The phrase highlights its high intracellular concentration and its role in several antioxidant and detoxification reactions. It is not the only antioxidant, and the term can oversimplify its functions. Scientific descriptions usually specify the pathway or enzyme involved.
GSH is the reduced, thiol-containing form of glutathione, while GSSG is the oxidized disulfide dimer formed when two GSH molecules react. Cells maintain a high GSH-to-GSSG ratio under normal conditions. A shift toward GSSG is often interpreted as oxidative stress, though sample handling can affect the measured ratio.