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Metabolism
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"Cellular metabolism" redirects here. For the journal, see Cell Metabolism.

For the journal 'Metabolism', see Metabolism: Clinical and Experimental. For the architectural movement, see Metabolism (architecture).

Metabolism (/mǝˈtæbǝlɪzǝm/, from Greek: µεταβολή metabolē, "change") is the set of life-sustaining chemical reactions in
organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular
processes; the conversion of food to building blocks for proteins, lipids, nucleic acids, and some carbohydrates; and the elimination of
metabolic wastes. These enzyme-catalyzed reactions allow organisms to grow and reproduce, maintain their structures, and respond
to their environments. The word metabolism can also refer to the sum of all chemical reactions that occur in living organisms, including
digestion and the transportation of substances into and between different cells, in which case the above described set of reactions
within the cells is called intermediary (or intermediate) metabolism.

Metabolic reactions may be categorized as catabolic – the breaking down of compounds (for
example, of glucose to pyruvate by cellular respiration); or anabolic – the building up (synthesis) of
compounds (such as proteins, carbohydrates, lipids, and nucleic acids). Usually, catabolism releases
energy, and anabolism consumes energy.

The chemical reactions of metabolism are organized into metabolic pathways, in which one chemical
is transformed through a series of steps into another chemical, each step being facilitated by a
specific enzyme. Enzymes are crucial to metabolism because they allow organisms to drive desirable
reactions that require energy and will not occur by themselves, by coupling them to spontaneous Simplified view of the cellular
metabolism
reactions that release energy. Enzymes act as catalysts – they allow a reaction to proceed more
rapidly – and they also allow the regulation of the rate of a metabolic reaction, for example in
response to changes in the cell's environment or to signals from other cells.

The metabolic system of a particular organism determines which substances it will find nutritious
and which poisonous. For example, some prokaryotes use hydrogen sulfide as a nutrient, yet this gas
is poisonous to animals.[1] The basal metabolic rate of an organism is the measure of the amount of
energy consumed by all of these chemical reactions.
Structure of adenosine
A striking feature of metabolism is the similarity of the basic metabolic pathways among vastly triphosphate (ATP), a central
different species.[2] For example, the set of carboxylic acids that are best known as the intermediate in energy
metabolism
intermediates in the citric acid cycle are present in all known organisms, being found in species as
diverse as the unicellular bacterium Escherichia coli and huge multicellular organisms like elephants.[3] These similarities in metabolic
pathways are likely due to their early appearance in evolutionary history, and their retention is likely due to their efficacy.[4][5] In
various diseases, such as type II diabetes, metabolic syndrome, and cancer, normal metabolism is disrupted.[6] The metabolism of
cancer cells is also different from the metabolism of normal cells, and these differences can be used to find targets for therapeutic
intervention in cancer.[7]

Contents

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Key biochemicals

Further information: Biomolecule, Cell (biology), and Biochemistry

Most of the structures that make up animals, plants and microbes are made from four basic
classes of molecules: amino acids, carbohydrates, nucleic acid and lipids (often called fats). As
these molecules are vital for life, metabolic reactions either focus on making these molecules
during the construction of cells and tissues, or on breaking them down and using them to obtain
energy, by their digestion. These biochemicals can be joined to make polymers such as DNA and
proteins, essential macromolecules of life.[8]

Type of Name of monomer

Name of polymer forms Examples of polymer forms
molecule forms
Structure of a triacylglycerol lipid
Proteins (made of Fibrous proteins and globular
Amino acids Amino acids
polypeptides) proteins

Starch, glycogen and

Carbohydrates Monosaccharides Polysaccharides
cellulose

Nucleic acids Nucleotides Polynucleotides DNA and RNA

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Amino acids and proteins
Main article: Protein
This is a diagram depicting a large
set of human metabolic pathways.
Proteins are made of amino acids arranged in a linear chain joined by peptide bonds. Many proteins [image reference needed]

are enzymes that catalyze the chemical reactions in metabolism. Other proteins have structural or
mechanical functions, such as those that form the cytoskeleton, a system of scaffolding that maintains the cell shape.[9] Proteins are
also important in cell signaling, immune responses, cell adhesion, active transport across membranes, and the cell cycle.[10] Amino
acids also contribute to cellular energy metabolism by providing a carbon source for entry into the citric acid cycle (tricarboxylic acid
cycle),[11] especially when a primary source of energy, such as glucose, is scarce, or when cells undergo metabolic stress.[12]

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Lipids
Main article: Biolipid

Lipids are the most diverse group of biochemicals. Their main structural uses are as part of biological membranes both internal and
external, such as the cell membrane.[10] Their chemical energy can also be used. Lipids are the polymers of fatty acids[citation needed]
that contain a long, non-polar hydrocarbon chain with a small polar region containing oxygen. Lipids are usually defined as
hydrophobic or amphipathic biological molecules but will dissolve in organic solvents such as ethanol, benzene or chloroform.[13] The
fats are a large group of compounds that contain fatty acids and glycerol; a glycerol molecule attached to three fatty acids by ester
linkages is called a triacylglyceride.[14] Several variations on this basic structure exist, including backbones such as sphingosine in
sphingomyelin, and hydrophilic groups such as phosphate as in phospholipids. Steroids such as sterol are another major class of
lipids.[15]

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Carbohydrates
Main article: Carbohydrate

Carbohydrates are aldehydes or ketones, with many hydroxyl groups attached, that can exist as
straight chains or rings. Carbohydrates are the most abundant biological molecules, and fill
numerous roles, such as the storage and transport of energy (starch, glycogen) and structural
components (cellulose in plants, chitin in animals).[10] The basic carbohydrate units are called
monosaccharides and include galactose, fructose, and most importantly glucose.
Monosaccharides can be linked together to form polysaccharides in almost limitless ways.[16]
Glucose can exist in both a straight-
chain and ring form.
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Nucleotides
Main article: Nucleotide

The two nucleic acids, DNA and RNA, are polymers of nucleotides. Each nucleotide is composed of a phosphate attached to a ribose
or deoxyribose sugar group which is attached to a nitrogenous base. Nucleic acids are critical for the storage and use of genetic
information, and its interpretation through the processes of transcription and protein biosynthesis.[10] This information is protected by
DNA repair mechanisms and propagated through DNA replication. Many viruses have an RNA genome, such as HIV, which uses
reverse transcription to create a DNA template from its viral RNA genome.[17] RNA in ribozymes such as spliceosomes and ribosomes
is similar to enzymes as it can catalyze chemical reactions. Individual nucleosides are made by attaching a nucleobase to a ribose
sugar. These bases are heterocyclic rings containing nitrogen, classified as purines or pyrimidines. Nucleotides also act as coenzymes
in metabolic-group-transfer reactions.[18]

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Coenzymes
Main article: Coenzyme

Metabolism involves a vast array of chemical reactions, but most fall under a few basic
types of reactions that involve the transfer of functional groups of atoms and their bonds
within molecules.[19] This common chemistry allows cells to use a small set of metabolic
intermediates to carry chemical groups between different reactions.[18] These group- Structure of the coenzyme acetyl-CoA. The
transfer intermediates are called coenzymes. Each class of group-transfer reactions is transferable acetyl group is bonded to the
carried out by a particular coenzyme, which is the substrate for a set of enzymes that sulfur atom at the extreme left.

produce it, and a set of enzymes that consume it. These coenzymes are therefore
continuously made, consumed and then recycled.[20]

One central coenzyme is adenosine triphosphate (ATP), the energy currency of cells. This nucleotide is used to transfer chemical
energy between different chemical reactions. There is only a small amount of ATP in cells, but as it is continuously regenerated, the
human body can use about its own weight in ATP per day.[20] ATP acts as a bridge between catabolism and anabolism. Catabolism
breaks down molecules, and anabolism puts them together. Catabolic reactions generate ATP, and anabolic reactions consume it. It
also serves as a carrier of phosphate groups in phosphorylation reactions.[21]

A vitamin is an organic compound needed in small quantities that cannot be made in cells. In human nutrition, most vitamins function
as coenzymes after modification; for example, all water-soluble vitamins are phosphorylated or are coupled to nucleotides when they
are used in cells.[22] Nicotinamide adenine dinucleotide (NAD+), a derivative of vitamin B3 (niacin), is an important coenzyme that acts
as a hydrogen acceptor. Hundreds of separate types of dehydrogenases remove electrons from their substrates and reduce NAD+ into
NADH. This reduced form of the coenzyme is then a substrate for any of the reductases in the cell that need to transfer hydrogen
atoms to their substrates.[23] Nicotinamide adenine dinucleotide exists in two related forms in the cell, NADH and NADPH. The
NAD+/NADH form is more important in catabolic reactions, while NADP+/NADPH is used in anabolic reactions.[24]

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Mineral and cofactors
Further information: Bioinorganic chemistry

Inorganic elements play critical roles in metabolism; some are abundant (e.g. sodium and
potassium) while others function at minute concentrations. About 99% of a human's
body weight is made up of the elements carbon, nitrogen, calcium, sodium, chlorine,
potassium, hydrogen, phosphorus, oxygen and sulfur. Organic compounds (proteins,
lipids and carbohydrates) contain the majority of the carbon and nitrogen; most of the
oxygen and hydrogen is present as water.[25]

The abundant inorganic elements act as electrolytes. The most important ions are
sodium, potassium, calcium, magnesium, chloride, phosphate and the organic ion
bicarbonate. The maintenance of precise ion gradients across cell membranes maintains The structure of iron-containing hemoglobin.
osmotic pressure and pH.[26] Ions are also critical for nerve and muscle function, as The protein subunits are in red and blue, and
action potentials in these tissues are produced by the exchange of electrolytes between the iron-containing heme groups in green.
From PDB: 1GZX.
the extracellular fluid and the cell's fluid, the cytosol.[27] Electrolytes enter and leave cells
through proteins in the cell membrane called ion channels. For example, muscle contraction depends upon the movement of calcium,
sodium and potassium through ion channels in the cell membrane and T-tubules.[28]

Transition metals are usually present as trace elements in organisms, with zinc and iron being most abundant of those.[29] Metal
cofactors are bound tightly to specific sites in proteins; although enzyme cofactors can be modified during catalysis, they always
return to their original state by the end of the reaction catalyzed. Metal micronutrients are taken up into organisms by specific
transporters and bind to storage proteins such as ferritin or metallothionein when not in use.[30][31]

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Catabolism

Main article: Catabolism

Catabolism is the set of metabolic processes that break down large molecules. These include breaking down and oxidizing food
molecules. The purpose of the catabolic reactions is to provide the energy and components needed by anabolic reactions which build
molecules.[32] The exact nature of these catabolic reactions differ from organism to organism, and organisms can be classified based
on their sources of energy, hydrogen, and carbon (their primary nutritional groups), as shown in the table below. Organic molecules
are used as a source of hydrogen atoms or electrons by organotrophs, while lithotrophs use inorganic substrates. Whereas
phototrophs convert sunlight to chemical energy,[33] chemotrophs depend on redox reactions that involve the transfer of electrons
from reduced donor molecules such as organic molecules, hydrogen, hydrogen sulfide or ferrous ions to oxygen, nitrate or sulfate. In
animals, these reactions involve complex organic molecules that are broken down to simpler molecules, such as carbon dioxide and
water. Photosynthetic organisms, such as plants and cyanobacteria, use similar electron-transfer reactions to store energy absorbed
from sunlight.[34]

Classification of organisms based on their metabolism [35]

sunlight photo-
Energy source
molecules chemo-

organic
organo-
Hydrogen or compound
electron donor inorganic -
litho-
compound troph

organic
hetero-
compound
Carbon source
inorganic
auto-
compound

The most common set of catabolic reactions in animals can be separated into three main stages. In the first stage, large organic
molecules, such as proteins, polysaccharides or lipids, are digested into their smaller components outside cells. Next, these smaller
molecules are taken up by cells and converted to smaller molecules, usually acetyl coenzyme A (acetyl-CoA), which releases some
energy. Finally, the acetyl group on acetyl-CoA is oxidized to water and carbon dioxide in the citric acid cycle and electron transport
chain, releasing more energy while reducing the coenzyme nicotinamide adenine dinucleotide (NAD+) into NADH.[32]

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Digestion
Further information: Digestion and Gastrointestinal tract

Macromolecules cannot be directly processed by cells. Macromolecules must be broken into smaller units before they can be used in
cell metabolism. Different classes of enzymes are used to digest these polymers. These digestive enzymes include proteases that
digest proteins into amino acids, as well as glycoside hydrolases that digest polysaccharides into simple sugars known as
monosaccharides.[36]

Microbes simply secrete digestive enzymes into their surroundings,[37][38] while animals only secrete these enzymes from specialized
cells in their guts, including the stomach and pancreas, and in salivary glands.[39] The amino acids or sugars released by these
extracellular enzymes are then pumped into cells by active transport proteins.[40][41]

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Energy from organic compounds
Further information: Cellular respiration, Fermentation (biochemistry), Carbohydrate catabolism,
Fat catabolism, and Protein catabolism

Carbohydrate catabolism is the breakdown of carbohydrates into smaller units.

Carbohydrates are usually taken into cells after they have been digested into
monosaccharides.[42] Once inside, the major route of breakdown is glycolysis, where
sugars such as glucose and fructose are converted into pyruvate and some ATP is
generated.[43] Pyruvate is an intermediate in several metabolic pathways, but the majority
is converted to acetyl-CoA through aerobic (with oxygen) glycolysis and fed into the citric
acid cycle. Although some more ATP is generated in the citric acid cycle, the most
A simplified outline of the catabolism of important product is NADH, which is made from NAD+ as the acetyl-CoA is oxidized. This
proteins, carbohydrates and oxidation releases carbon dioxide as a waste product. In anaerobic conditions, glycolysis
[image reference needed]
fats
produces lactate, through the enzyme lactate dehydrogenase re-oxidizing NADH to
NAD+ for re-use in glycolysis.[44] An alternative route for glucose breakdown is the pentose phosphate pathway, which reduces the
coenzyme NADPH and produces pentose sugars such as ribose, the sugar component of nucleic acids. [citation needed]

Fats are catabolized by hydrolysis to free fatty acids and glycerol. The glycerol enters glycolysis and the fatty acids are broken down
by beta oxidation to release acetyl-CoA, which then is fed into the citric acid cycle. Fatty acids release more energy upon oxidation
than carbohydrates. Steroids are also broken down by some bacteria in a process similar to beta oxidation, and this breakdown
process involves the release of significant amounts of acetyl-CoA, propionyl-CoA, and pyruvate, which can all be used by the cell for
energy. M. tuberculosis can also grow on the lipid cholesterol as a sole source of carbon, and genes involved in the cholesterol-use
pathway(s) have been validated as important during various stages of the infection lifecycle of M. tuberculosis.[45]

Amino acids are either used to synthesize proteins and other biomolecules, or oxidized to urea and carbon dioxide to produce energy.
[46] The oxidation pathway starts with the removal of the amino group by a transaminase. The amino group is fed into the urea cycle,
leaving a deaminated carbon skeleton in the form of a keto acid. Several of these keto acids are intermediates in the citric acid cycle,
for example α-ketoglutarate formed by deamination of glutamate.[47] The glucogenic amino acids can also be converted into glucose,
through gluconeogenesis (discussed below).[48]

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Energy transformations

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Oxidative phosphorylation
Further information: Oxidative phosphorylation, Chemiosmosis, and Mitochondrion

In oxidative phosphorylation, the electrons removed from organic molecules in areas such as the citric acid cycle are transferred to
oxygen and the energy released is used to make ATP. This is done in eukaryotes by a series of proteins in the membranes of
mitochondria called the electron transport chain. In prokaryotes, these proteins are found in the cell's inner membrane.[49] These
proteins use the energy from reduced molecules like NADH to pump protons across a membrane.[50]

Pumping protons out of the mitochondria creates a proton concentration difference across the
membrane and generates an electrochemical gradient.[51] This force drives protons back into the
mitochondrion through the base of an enzyme called ATP synthase. The flow of protons makes the
stalk subunit rotate, causing the active site of the synthase domain to change shape and
phosphorylate adenosine diphosphate – turning it into ATP.[20]

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Energy from inorganic compounds
Further information: Microbial metabolism and Nitrogen cycle Mechanism of ATP synthase. ATP
is shown in red, ADP and
Chemolithotrophy is a type of metabolism found in prokaryotes where energy is obtained from the phosphate in pink and the rotating
oxidation of inorganic compounds. These organisms can use hydrogen,[52] reduced sulfur stalk subunit in black.

compounds (such as sulfide, hydrogen sulfide and thiosulfate),[1] ferrous iron (Fe(II))[53] or
ammonia[54] as sources of reducing power and they gain energy from the oxidation of these compounds.[55] These microbial
processes are important in global biogeochemical cycles such as acetogenesis, nitrification and denitrification and are critical for soil
fertility.[56][57]

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Energy from light
Further information: Phototroph, Photophosphorylation, and Chloroplast

The energy in sunlight is captured by plants, cyanobacteria, purple bacteria, green sulfur bacteria and some protists. This process is
often coupled to the conversion of carbon dioxide into organic compounds, as part of photosynthesis, which is discussed below. The
energy capture and carbon fixation systems can, however, operate separately in prokaryotes, as purple bacteria and green sulfur
bacteria can use sunlight as a source of energy, while switching between carbon fixation and the fermentation of organic compounds.
[58][59]

In many organisms, the capture of solar energy is similar in principle to oxidative phosphorylation, as it involves the storage of energy
as a proton concentration gradient. This proton motive force then drives ATP synthesis.[60] The electrons needed to drive this electron
transport chain come from light-gathering proteins called photosynthetic reaction centres. Reaction centers are classified into two
types depending on the nature of photosynthetic pigment present, with most photosynthetic bacteria only having one type, while
plants and cyanobacteria have two.[61]

In plants, algae, and cyanobacteria, photosystem II uses light energy to remove electrons from water, releasing oxygen as a waste
product. The electrons then flow to the cytochrome b6f complex, which uses their energy to pump protons across the thylakoid
membrane in the chloroplast.[34] These protons move back through the membrane as they drive the ATP synthase, as before. The
[62]
electrons then flow through photosystem I and can then be used to reduce the coenzyme NADP+. This coenzyme can enter the
Calvin cycle, which is discussed below, or be recycled for further ATP generation. [citation needed]

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Anabolism

Further information: Anabolism

Anabolism is the set of constructive metabolic processes where the energy released by catabolism is used to synthesize complex
molecules. In general, the complex molecules that make up cellular structures are constructed step-by-step from smaller and simpler
precursors. Anabolism involves three basic stages. First, the production of precursors such as amino acids, monosaccharides,
isoprenoids and nucleotides, secondly, their activation into reactive forms using energy from ATP, and thirdly, the assembly of these
precursors into complex molecules such as proteins, polysaccharides, lipids and nucleic acids.[63]

Anabolism in organisms can be different according to the source of constructed molecules in their cells. Autotrophs such as plants
can construct the complex organic molecules in their cells such as polysaccharides and proteins from simple molecules like carbon
dioxide and water. Heterotrophs, on the other hand, require a source of more complex substances, such as monosaccharides and
amino acids, to produce these complex molecules. Organisms can be further classified by ultimate source of their energy:
photoautotrophs and photoheterotrophs obtain energy from light, whereas chemoautotrophs and chemoheterotrophs obtain energy
from oxidation reactions.[63]

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Carbon fixation
Further information: Photosynthesis, Carbon fixation, and Chemosynthesis

Photosynthesis is the synthesis of carbohydrates from sunlight and carbon dioxide (CO2). In plants,
cyanobacteria and algae, oxygenic photosynthesis splits water, with oxygen produced as a waste
product. This process uses the ATP and NADPH produced by the photosynthetic reaction centres,
as described above, to convert CO2 into glycerate 3-phosphate, which can then be converted into
glucose. This carbon-fixation reaction is carried out by the enzyme RuBisCO as part of the Calvin –
Benson cycle.[64] Three types of photosynthesis occur in plants, C3 carbon fixation, C4 carbon
fixation and CAM photosynthesis. These differ by the route that carbon dioxide takes to the Calvin
cycle, with C3 plants fixing CO2 directly, while C4 and CAM photosynthesis incorporate the CO2 into Plant cells (bounded by purple
walls) filled with chloroplasts
other compounds first, as adaptations to deal with intense sunlight and dry conditions.[65]
(green), which are the site of
photosynthesis
In photosynthetic prokaryotes the mechanisms of carbon fixation are more diverse. Here, carbon
dioxide can be fixed by the Calvin – Benson cycle, a reversed citric acid cycle,[66] or the carboxylation of acetyl-CoA.[67][68]
Prokaryotic chemoautotrophs also fix CO2 through the Calvin–Benson cycle, but use energy from inorganic compounds to drive the
reaction.[69]

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Carbohydrates and glycans
Further information: Gluconeogenesis, Glyoxylate cycle, Glycogenesis, and Glycosylation

In carbohydrate anabolism, simple organic acids can be converted into monosaccharides such as glucose and then used to assemble
polysaccharides such as starch. The generation of glucose from compounds like pyruvate, lactate, glycerol, glycerate 3-phosphate
and amino acids is called gluconeogenesis. Gluconeogenesis converts pyruvate to glucose-6-phosphate through a series of
intermediates, many of which are shared with glycolysis.[43] However, this pathway is not simply glycolysis run in reverse, as several
steps are catalyzed by non-glycolytic enzymes. This is important as it allows the formation and breakdown of glucose to be regulated
separately, and prevents both pathways from running simultaneously in a futile cycle.[70][71]

Although fat is a common way of storing energy, in vertebrates such as humans the fatty acids in these stores cannot be converted to
glucose through gluconeogenesis as these organisms cannot convert acetyl-CoA into pyruvate; plants do, but animals do not, have
the necessary enzymatic machinery.[72] As a result, after long-term starvation, vertebrates need to produce ketone bodies from fatty
acids to replace glucose in tissues such as the brain that cannot metabolize fatty acids.[73] In other organisms such as plants and
bacteria, this metabolic problem is solved using the glyoxylate cycle, which bypasses the decarboxylation step in the citric acid cycle
and allows the transformation of acetyl-CoA to oxaloacetate, where it can be used for the production of glucose.[72][74] Other than fat,
glucose is stored in most tissues, as an energy resource available within the tissue through glycogenesis which was usually being
used to maintained glucose level in blood.[75]

Polysaccharides and glycans are made by the sequential addition of monosaccharides by glycosyltransferase from a reactive sugar-
phosphate donor such as uridine diphosphate glucose (UDP-Glc) to an acceptor hydroxyl group on the growing polysaccharide. As
any of the hydroxyl groups on the ring of the substrate can be acceptors, the polysaccharides produced can have straight or branched
structures.[76] The polysaccharides produced can have structural or metabolic functions themselves, or be transferred to lipids and
proteins by enzymes called oligosaccharyltransferases.[77][78]

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Fatty acids, isoprenoids and sterol
Further information: Fatty acid synthesis and Steroid metabolism

Fatty acids are made by fatty acid synthases that polymerize and then reduce
acetyl-CoA units. The acyl chains in the fatty acids are extended by a cycle of
reactions that add the acyl group, reduce it to an alcohol, dehydrate it to an alkene
group and then reduce it again to an alkane group. The enzymes of fatty acid
biosynthesis are divided into two groups: in animals and fungi, all these fatty acid
synthase reactions are carried out by a single multifunctional type I protein,[79]
while in plant plastids and bacteria separate type II enzymes perform each step in
the pathway.[80][81]

Terpenes and isoprenoids are a large class of lipids that include the carotenoids
and form the largest class of plant natural products.[82] These compounds are
made by the assembly and modification of isoprene units donated from the
reactive precursors isopentenyl pyrophosphate and dimethylallyl pyrophosphate.
[83] These precursors can be made in different ways. In animals and archaea, the
mevalonate pathway produces these compounds from acetyl-CoA,[84] while in
plants and bacteria the non-mevalonate pathway uses pyruvate and
glyceraldehyde 3-phosphate as substrates.[83][85] One important reaction that Simplified version of the steroid synthesis pathway
with the intermediates isopentenyl pyrophosphate
uses these activated isoprene donors is sterol biosynthesis. Here, the isoprene
(IPP), dimethylallyl pyrophosphate (DMAPP), geranyl
units are joined to make squalene and then folded up and formed into a set of rings
pyrophosphate (GPP) and squalene shown. Some
to make lanosterol.[86] Lanosterol can then be converted into other sterols such as intermediates are omitted for clarity.
cholesterol and ergosterol.[86][87]

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Proteins
Further information: Protein biosynthesis and Amino acid synthesis

Organisms vary in their ability to synthesize the 20 common amino acids. Most bacteria and plants can synthesize all twenty, but
mammals can only synthesize eleven nonessential amino acids, so nine essential amino acids must be obtained from food.[10] Some
simple parasites, such as the bacteria Mycoplasma pneumoniae, lack all amino acid synthesis and take their amino acids directly from
their hosts.[88] All amino acids are synthesized from intermediates in glycolysis, the citric acid cycle, or the pentose phosphate
pathway. Nitrogen is provided by glutamate and glutamine. Nonessensial amino acid synthesis depends on the formation of the
appropriate alpha-keto acid, which is then transaminated to form an amino acid.[89]

Amino acids are made into proteins by being joined in a chain of peptide bonds. Each different protein has a unique sequence of amino
acid residues: this is its primary structure. Just as the letters of the alphabet can be combined to form an almost endless variety of
words, amino acids can be linked in varying sequences to form a huge variety of proteins. Proteins are made from amino acids that
have been activated by attachment to a transfer RNA molecule through an ester bond. This aminoacyl-tRNA precursor is produced in
an ATP-dependent reaction carried out by an aminoacyl tRNA synthetase.[90] This aminoacyl-tRNA is then a substrate for the
ribosome, which joins the amino acid onto the elongating protein chain, using the sequence information in a messenger RNA.[91]

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Nucleotide synthesis and salvage
Further information: Nucleotide salvage, Pyrimidine biosynthesis, and Purine § Metabolism

Nucleotides are made from amino acids, carbon dioxide and formic acid in pathways that require large amounts of metabolic energy.
[92] Consequently, most organisms have efficient systems to salvage preformed nucleotides.[92][93] Purines are synthesized as
nucleosides (bases attached to ribose).[94] Both adenine and guanine are made from the precursor nucleoside inosine
monophosphate, which is synthesized using atoms from the amino acids glycine, glutamine, and aspartic acid, as well as formate
transferred from the coenzyme tetrahydrofolate. Pyrimidines, on the other hand, are synthesized from the base orotate, which is
formed from glutamine and aspartate.[95]

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Xenobiotics and redox metabolism

Further information: Xenobiotic metabolism, Drug metabolism, Alcohol metabolism, and Antioxidant

All organisms are constantly exposed to compounds that they cannot use as foods and that would be harmful if they accumulated in
cells, as they have no metabolic function. These potentially damaging compounds are called xenobiotics.[96] Xenobiotics such as
synthetic drugs, natural poisons and antibiotics are detoxified by a set of xenobiotic-metabolizing enzymes. In humans, these include
cytochrome P450 oxidases,[97] UDP-glucuronosyltransferases,[98] and glutathione S-transferases.[99] This system of enzymes acts in
three stages to firstly oxidize the xenobiotic (phase I) and then conjugate water-soluble groups onto the molecule (phase II). The
modified water-soluble xenobiotic can then be pumped out of cells and in multicellular organisms may be further metabolized before
being excreted (phase III). In ecology, these reactions are particularly important in microbial biodegradation of pollutants and the
bioremediation of contaminated land and oil spills.[100] Many of these microbial reactions are shared with multicellular organisms, but
due to the incredible diversity of types of microbes these organisms are able to deal with a far wider range of xenobiotics than
multicellular organisms, and can degrade even persistent organic pollutants such as organochloride compounds.[101]

A related problem for aerobic organisms is oxidative stress.[102] Here, processes including oxidative phosphorylation and the formation
of disulfide bonds during protein folding produce reactive oxygen species such as hydrogen peroxide.[103] These damaging oxidants
are removed by antioxidant metabolites such as glutathione and enzymes such as catalases and peroxidases.[104][105]

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Thermodynamics of living organisms

Further information: Biological thermodynamics

Living organisms must obey the laws of thermodynamics, which describe the transfer of heat and work. The second law of
thermodynamics states that in any isolated system, the amount of entropy (disorder) cannot decrease. Although living organisms'
amazing complexity appears to contradict this law, life is possible as all organisms are open systems that exchange matter and energy
with their surroundings. Living systems are not in equilibrium, but instead are dissipative systems that maintain their state of high
complexity by causing a larger increase in the entropy of their environments.[106] The metabolism of a cell achieves this by coupling
the spontaneous processes of catabolism to the non-spontaneous processes of anabolism. In thermodynamic terms, metabolism
maintains order by creating disorder.[107]

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Regulation and control

Further information: Metabolic pathway, Metabolic control analysis, Hormone, Regulatory enzymes, and Cell signaling

As the environments of most organisms are constantly changing, the reactions of metabolism must be finely regulated to maintain a
constant set of conditions within cells, a condition called homeostasis.[108][109] Metabolic regulation also allows organisms to respond
to signals and interact actively with their environments.[110] Two closely linked concepts are important for understanding how
metabolic pathways are controlled. Firstly, the regulation of an enzyme in a pathway is how its activity is increased and decreased in
response to signals. Secondly, the control exerted by this enzyme is the effect that these changes in its activity have on the overall
rate of the pathway (the flux through the pathway).[111] For example, an enzyme may show large changes in activity (i.e. it is highly
regulated) but if these changes have little effect on the flux of a metabolic pathway, then this enzyme is not involved in the control of
the pathway.[112]

There are multiple levels of metabolic regulation. In intrinsic regulation, the metabolic
pathway self-regulates to respond to changes in the levels of substrates or products; for
example, a decrease in the amount of product can increase the flux through the pathway
to compensate.[111] This type of regulation often involves allosteric regulation of the
activities of multiple enzymes in the pathway.[113] Extrinsic control involves a cell in a
multicellular organism changing its metabolism in response to signals from other cells.
These signals are usually in the form of water-soluble messengers such as hormones and
growth factors and are detected by specific receptors on the cell surface.[114] These Effect of insulin on glucose uptake and
signals are then transmitted inside the cell by second messenger systems that often metabolism. Insulin binds to its receptor (1),
which in turn starts many protein activation
involved the phosphorylation of proteins.[115]
cascades (2). These include: translocation of
Glut-4 transporter to the plasma membrane
A very well understood example of extrinsic control is the regulation of glucose
and influx of glucose (3), glycogen synthesis
metabolism by the hormone insulin.[116] Insulin is produced in response to rises in blood (4), glycolysis (5) and fatty acid synthesis (6).
[image reference needed]
glucose levels. Binding of the hormone to insulin receptors on cells then activates a
cascade of protein kinases that cause the cells to take up glucose and convert it into
storage molecules such as fatty acids and glycogen.[117] The metabolism of glycogen is controlled by activity of phosphorylase, the
enzyme that breaks down glycogen, and glycogen synthase, the enzyme that makes it. These enzymes are regulated in a reciprocal
fashion, with phosphorylation inhibiting glycogen synthase, but activating phosphorylase. Insulin causes glycogen synthesis by
activating protein phosphatases and producing a decrease in the phosphorylation of these enzymes.[118]

Edit
Evolution

Further information: Molecular evolution and Phylogenetics

The central pathways of metabolism described above, such as glycolysis

and the citric acid cycle, are present in all three domains of living things and
were present in the last universal common ancestor.[3][119] This universal
ancestral cell was prokaryotic and probably a methanogen that had
extensive amino acid, nucleotide, carbohydrate and lipid metabolism.[120]
[121] The retention of these ancient pathways during later evolution may be
the result of these reactions having been an optimal solution to their
particular metabolic problems, with pathways such as glycolysis and the
citric acid cycle producing their end products highly efficiently and in a
minimal number of steps.[4][5] The first pathways of enzyme-based
metabolism may have been parts of purine nucleotide metabolism, while
previous metabolic pathways were a part of the ancient RNA world.[122] Evolutionary tree showing the common ancestry of organisms
from all three domains of life. Bacteria are colored blue,
Many models have been proposed to describe the mechanisms by which eukaryotes red, and archaea green. Relative positions of some
novel metabolic pathways evolve. These include the sequential addition of of the phyla included are shown around the tree.
novel enzymes to a short ancestral pathway, the duplication and then
divergence of entire pathways as well as the recruitment of pre-existing enzymes and their assembly into a novel reaction pathway.
[123] The relative importance of these mechanisms is unclear, but genomic studies have shown that enzymes in a pathway are likely to
have a shared ancestry, suggesting that many pathways have evolved in a step-by-step fashion with novel functions created from pre-
existing steps in the pathway.[124] An alternative model comes from studies that trace the evolution of proteins' structures in metabolic
networks, this has suggested that enzymes are pervasively recruited, borrowing enzymes to perform similar functions in different
metabolic pathways (evident in the MANET database)[125] These recruitment processes result in an evolutionary enzymatic mosaic.
[126] A third possibility is that some parts of metabolism might exist as "modules" that can be reused in different pathways and
perform similar functions on different molecules.[127]

As well as the evolution of new metabolic pathways, evolution can also cause the loss of metabolic functions. For example, in some
parasites metabolic processes that are not essential for survival are lost and preformed amino acids, nucleotides and carbohydrates
may instead be scavenged from the host.[128] Similar reduced metabolic capabilities are seen in endosymbiotic organisms.[129]

Edit
Investigation and manipulation

Further information: Protein methods, Proteomics, Metabolomics, and Metabolic network modelling

Classically, metabolism is studied by a reductionist approach that focuses on a single

metabolic pathway. Particularly valuable is the use of radioactive tracers at the whole-
organism, tissue and cellular levels, which define the paths from precursors to final
products by identifying radioactively labelled intermediates and products.[130] The
enzymes that catalyze these chemical reactions can then be purified and their kinetics
and responses to inhibitors investigated. A parallel approach is to identify the small
molecules in a cell or tissue; the complete set of these molecules is called the
metabolome. Overall, these studies give a good view of the structure and function of
simple metabolic pathways, but are inadequate when applied to more complex systems
such as the metabolism of a complete cell.[131]

An idea of the complexity of the metabolic networks in cells that contain thousands of
different enzymes is given by the figure showing the interactions between just 43
proteins and 40 metabolites to the right: the sequences of genomes provide lists Metabolic network of the Arabidopsis thaliana
containing anything up to 26.500 genes.[132] However, it is now possible to use this citric acid cycle. Enzymes and metabolites are
shown as red squares and the interactions
genomic data to reconstruct complete networks of biochemical reactions and produce
between them as black lines.
more holistic mathematical models that may explain and predict their behavior.[133] These
models are especially powerful when used to integrate the pathway and metabolite data obtained through classical methods with data
on gene expression from proteomic and DNA microarray studies.[134] Using these techniques, a model of human metabolism has now
been produced, which will guide future drug discovery and biochemical research.[135] These models are now used in network analysis,
to classify human diseases into groups that share common proteins or metabolites.[136][137]

Bacterial metabolic networks are a striking example of bow-tie[138][139][140] organization, an architecture able to input a wide range of
nutrients and produce a large variety of products and complex macromolecules using a relatively few intermediate common
currencies.

A major technological application of this information is metabolic engineering. Here, organisms such as yeast, plants or bacteria are
genetically modified to make them more useful in biotechnology and aid the production of drugs such as antibiotics or industrial
chemicals such as 1,3-propanediol and shikimic acid.[141][142][143] These genetic modifications usually aim to reduce the amount of
energy used to produce the product, increase yields and reduce the production of wastes.[144]

Edit
History

Further information: History of biochemistry and History of molecular biology

The term metabolism is derived from the Ancient Greek word µεταβολή – "Metabole" for "a change" which derived from µεταβάλλ
–"Metaballein" means "To change"[145]

Edit
Greek philosophy

Aristotle's The Parts of Animals sets out enough details of his views on metabolism for
an open flow model to be made. He believed that at each stage of the process, materials
from food were transformed, with heat being released as the classical element of fire,
and residual materials being excreted as urine, bile, or faeces.[146]

Ibn al-Nafis described metabolism in his 1260 AD work titled Al-Risalah al-Kamiliyyah fil
Siera al-Nabawiyyah (The Treatise of Kamil on the Prophet's Biography) which included
the following phrase "Both the body and its parts are in a continuous state of dissolution
and nourishment, so they are inevitably undergoing permanent change."[147] Aristotle's metabolism as an open flow model

Edit
Application of the scientific method and Modern metabolic theories

The history of the scientific study of metabolism spans several centuries and has moved from examining whole animals in early
studies, to examining individual metabolic reactions in modern biochemistry. The first controlled experiments in human metabolism
were published by Santorio Santorio in 1614 in his book Ars de statica medicina.[148] He described how he weighed himself before and
after eating, sleep, working, sex, fasting, drinking, and excreting. He found that most of the food he took in was lost through what he
called "insensible perspiration".

In these early studies, the mechanisms of these metabolic processes had not been identified and a vital force
was thought to animate living tissue.[149] In the 19th century, when studying the fermentation of sugar to
alcohol by yeast, Louis Pasteur concluded that fermentation was catalyzed by substances within the yeast
cells he called "ferments". He wrote that "alcoholic fermentation is an act correlated with the life and
organization of the yeast cells, not with the death or putrefaction of the cells."[150] This discovery, along with
the publication by Friedrich Wöhler in 1828 of a paper on the chemical synthesis of urea,[151] and is notable for
being the first organic compound prepared from wholly inorganic precursors. This proved that the organic
compounds and chemical reactions found in cells were no different in principle than any other part of
chemistry.

Santorio Santorio in his

It was the discovery of enzymes at the beginning of the 20th century by Eduard Buchner that separated the
steelyard balance, from
study of the chemical reactions of metabolism from the biological study of cells, and marked the beginnings of Ars de statica
biochemistry.[152] The mass of biochemical knowledge grew rapidly throughout the early 20th century. One of medicina, first
the most prolific of these modern biochemists was Hans Krebs who made huge contributions to the study of published 1614

metabolism.[153] He discovered the urea cycle and later, working with Hans Kornberg, the citric acid cycle and
the glyoxylate cycle.[154][155][74] Modern biochemical research has been greatly aided by the development of new techniques such as
chromatography, X-ray diffraction, NMR spectroscopy, radioisotopic labelling, electron microscopy and molecular dynamics
simulations. These techniques have allowed the discovery and detailed analysis of the many molecules and metabolic pathways in
cells. [citation needed]

Edit
See also

Anthropogenic metabolism

Antimetabolite – Chemical that inhibits the use of a metabolite

Calorimetry – Determining heat transfer in a system by measuring its other properties

Isothermal microcalorimetry – Measuring versus elapsed time the net rate of heat flow

Inborn errors of metabolism – Class of genetic diseases

Iron–sulfur world hypothesis – Hypothetical scenario for the origin of life, a "metabolism first" theory of the origin of life

Metabolic disorder – disease that involving errors in metabolic processes of building or degradation of molecules

Microphysiometry

Primary nutritional groups – Group of organisms

Respirometry – Estimation of metabolic rates by measuring heat production

Stream metabolism

Sulfur metabolism – Set of chemical reactions involving sulfur in living organisms

Thermic effect of food – effect of food

Urban metabolism

Water metabolism – Aspect of homeostasis concerning control of the amount of water in an organism

Overflow metabolism – Cellular phenomena

Oncometabolism

Reactome – Database of biological pathways

KEGG – Collection of bioinformatics databases

Edit
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Further reading

Introductory

Rose S, Mileusnic R (1999). The Chemistry of Life. Penguin Press Science. ISBN 0-14-027273-9.

Schneider EC, Sagan D (2005). Into the Cool: Energy Flow, Thermodynamics, and Life. University of Chicago Press. ISBN 0-226-73936-8.

Lane N (2004). Oxygen: The Molecule that Made the World. USA: Oxford University Press. ISBN 0-19-860783-0.

Advanced

Price N, Stevens L (1999). Fundamentals of Enzymology: Cell and Molecular Biology of Catalytic Proteins. Oxford University Press. ISBN 0-19-
850229-X.

Berg J, Tymoczko J, Stryer L (2002). Biochemistry. W. H. Freeman and Company. ISBN 0-7167-4955-6.

Cox M, Nelson DL (2004). Lehninger Principles of Biochemistry. Palgrave Macmillan. ISBN 0-7167-4339-6.

Brock TD, Madigan MR, Martinko J, Parker J (2002). Brock's Biology of Microorganisms. Benjamin Cummings. ISBN 0-13-066271-2.

Da Silva JJ, Williams RJ (1991). The Biological Chemistry of the Elements: The Inorganic Chemistry of Life. Clarendon Press. ISBN 0-19-855598-9.

Nicholls DG, Ferguson SJ (2002). Bioenergetics. Academic Press Inc. ISBN 0-12-518121-3.

Wood HG (February 1991). "Life with CO or CO2 and H2 as a source of carbon and energy". FASEB Journal. 5 (2): 156–63.
doi:10.1096/fasebj.5.2.1900793 . PMID 1900793 . S2CID 45967404 .

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