Cellular respiration

Cellular respiration is a set of metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate (ATP), and then release waste products.[1] The reactions involved in respiration are catabolic reactions, which break large molecules into smaller ones, releasing energy in the process, as weak so-called "high-energy" bonds are replaced by stronger bonds in the products. Respiration is one of the key ways a cell releases chemical energy to fuel cellular activity. Cellular respiration is considered an exothermic redox reaction which releases heat. The overall reaction occurs in a series of biochemical steps, most of which are redox reactions themselves. Although cellular respiration is technically a combustion reaction, it clearly does not resemble one when it occurs in a living cell because of the slow release of energy from the series of reactions.

Nutrients that are commonly used by animal and plant cells in respiration include sugar, amino acids and fatty acids, and the most common oxidizing agent (electron acceptor) is molecular oxygen (O2). The chemical energy stored in ATP (its third phosphate group is weakly bonded to the rest of the molecule and is cheaply broken allowing stronger bonds to form, thereby transferring energy for use by the cell) can then be used to drive processes requiring energy, including biosynthesis, locomotion or transportation of molecules across cell membranes.

Aerobic respiration

Auto-and heterotrophs
Aerobic respiration (red arrows) is the main means by which both fungi and animals utilize chemical energy in the form of organic compounds that were previously created through photosynthesis (green arrow).

Aerobic respiration requires oxygen (O2) in order to create ATP. Although carbohydrates, fats, and proteins are consumed as reactants, it is the preferred method of pyruvate breakdown in glycolysis and requires that pyruvate enter the mitochondria in order to be fully oxidized by the Krebs cycle. The products of this process are carbon dioxide and water, but the energy transferred is used to break bonds in ADP as the third phosphate group is added to form ATP (adenosine triphosphate), by substrate-level phosphorylation, NADH and FADH2

Simplified reaction: C6H12O6 (s) + 6 O2 (g) → 6 CO2 (g) + 6 H2O (l) + heat
ΔG = −2880 kJ per mol of C6H12O6

The negative ΔG indicates that the reaction can occur spontaneously.

The potential of NADH and FADH2 is converted to more ATP through an electron transport chain with oxygen as the "terminal electron acceptor". Most of the ATP produced by aerobic cellular respiration is made by oxidative phosphorylation. This works by the energy released in the consumption of pyruvate being used to create a chemiosmotic potential by pumping protons across a membrane. This potential is then used to drive ATP synthase and produce ATP from ADP and a phosphate group. Biology textbooks often state that 38 ATP molecules can be made per oxidised glucose molecule during cellular respiration (2 from glycolysis, 2 from the Krebs cycle, and about 34 from the electron transport system).[2] However, this maximum yield is never quite reached because of losses due to leaky membranes as well as the cost of moving pyruvate and ADP into the mitochondrial matrix, and current estimates range around 29 to 30 ATP per glucose.[2]

Aerobic metabolism is up to 15 times more efficient than anaerobic metabolism (which yields 2 molecules ATP per 1 molecule glucose). However some anaerobic organisms, such as methanogens are able to continue with anaerobic respiration, yielding more ATP by using other inorganic molecules (not oxygen) as final electron acceptors in the electron transport chain. They share the initial pathway of glycolysis but aerobic metabolism continues with the Krebs cycle and oxidative phosphorylation. The post-glycolytic reactions take place in the mitochondria in eukaryotic cells, and in the cytoplasm in prokaryotic cells.


Respiration diagram
Out of the cytoplasm it goes into the Krebs cycle with the acetyl CoA. It then mixes with CO2 and makes 2 ATP, NADH, and FADH. From there the NADH and FADH go into the NADH reductase, which produces the enzyme. The NADH pulls the enzyme's electrons to send through the electron transport chain. The electron transport chain pulls H+ ions through the chain. From the electron transport chain, the released hydrogen ions make ADP for an end result of 32 ATP. O2 attracts itself to the left over electron to make water. Lastly, ATP leaves through the ATP channel and out of the mitochondria.

Glycolysis is a metabolic pathway that takes place in the cytosol of cells in all living organisms. Glycolysis can be literally translated as "sugar splitting"[3], which functions with or without the presence of oxygne. In humans, aerobic conditions produce pyruvate and anaerobic conditions produce lactate. In aerobic conditions, the process converts one molecule of glucose into two molecules of pyruvate (pyruvic acid), generating energy in the form of two net molecules of ATP. Four molecules of ATP per glucose are actually produced, however, two are consumed as part of the preparatory phase. The initial phosphorylation of glucose is required to increase the reactivity (decrease its stability) in order for the molecule to be cleaved into two pyruvate molecules by the enzyme aldolase. During the pay-off phase of glycolysis, four phosphate groups are transferred to ADP by substrate-level phosphorylation to make four ATP, and two NADH are produced when the pyruvate are oxidized. The overall reaction can be expressed this way:

Glucose + 2 NAD+ + 2 Pi + 2 ADP → 2 pyruvate + 2 NADH + 2 ATP + 2 H+ + 2 H2O + heat

Starting with glucose, 1 ATP is used to donate a phosphate to glucose to produce glucose 6-phosphate. Glycogen can be converted into glucose 6-phosphate as well with the help of glycogen phosphorylase. During energy metabolism, glucose 6-phosphate becomes fructose 6-phosphate. An additional ATP is used to phosphorylate fructose 6-phosphate into fructose 1,6-bisphosphate by the help of phosphofructokinase. Fructose 1,6-biphosphate then splits into two phosphorylated molecules with three carbon chains which later degrades into pyruvate.

Oxidative decarboxylation of pyruvate

Pyruvate is oxidized to acetyl-CoA and CO2 by the pyruvate dehydrogenase complex (PDC). The PDC contains multiple copies of three enzymes and is located in the mitochondria of eukaryotic cells and in the cytosol of prokaryotes. In the conversion of pyruvate to acetyl-CoA, one molecule of NADH and one molecule of CO2 is formed.

Citric acid cycle

This is also called the Krebs cycle or the tricarboxylic acid cycle. When oxygen is present, acetyl-CoA is produced from the pyruvate molecules created from glycolysis. Once acetyl-CoA is formed, aerobic or anaerobic respiration can occur.[4] When oxygen is present, the mitochondria will undergo aerobic respiration which leads to the Krebs cycle. However, if oxygen is not present, fermentation of the pyruvate molecule will occur. In the presence of oxygen, when acetyl-CoA is produced, the molecule then enters the citric acid cycle (Krebs cycle) inside the mitochondrial matrix, and is oxidized to CO2 while at the same time reducing NAD to NADH. NADH can be used by the electron transport chain to create further ATP as part of oxidative phosphorylation. To fully oxidize the equivalent of one glucose molecule, two acetyl-CoA must be metabolized by the Krebs cycle. Two waste products, H2O and CO2, are created during this cycle.

The citric acid cycle is an 8-step process involving 18 different enzymes and co-enzymes.[4] During the cycle, acetyl-CoA (2 carbons) + oxaloacetate (4 carbons) yields citrate (6 carbons), which is rearranged to a more reactive form called isocitrate (6 carbons). Isocitrate is modified to become α-ketoglutarate (5 carbons), succinyl-CoA, succinate, fumarate, malate, and, finally, oxaloacetate.

The net gain of high-energy compounds from one cycle is 3 NADH, 1 FADH2, and 1 GTP; the GTP may subsequently be used to produce ATP. Thus, the total yield from 1 glucose molecule (2 pyruvate molecules) is 6 NADH, 2 FADH2, and 2 ATP.

Oxidative phosphorylation

In eukaryotes, oxidative phosphorylation occurs in the mitochondrial cristae. It comprises the electron transport chain that establishes a proton gradient (chemiosmotic potential) across the boundary of inner membrane by oxidizing the NADH produced from the Krebs cycle. ATP is synthesized by the ATP synthase enzyme when the chemiosmotic gradient is used to drive the phosphorylation of ADP. The electrons are finally transferred to exogenous oxygen and, with the addition of two protons, water is formed.

Efficiency of ATP production

The table below describes the reactions involved when one glucose molecule is fully oxidized into carbon dioxide. It is assumed that all the reduced coenzymes are oxidized by the electron transport chain and used for oxidative phosphorylation.

Step coenzyme yield ATP yield Source of ATP
Glycolysis preparatory phase −2 Phosphorylation of glucose and fructose 6-phosphate uses two ATP from the cytoplasm.
Glycolysis pay-off phase 4 Substrate-level phosphorylation
2 NADH 3 or 5 Oxidative phosphorylation : Each NADH produces net 1.5 ATP (instead of usual 2.5) due to NADH transport over the mitochondrial membrane
Oxidative decarboxylation of pyruvate 2 NADH 5 Oxidative phosphorylation
Krebs cycle 2 Substrate-level phosphorylation
6 NADH 15 Oxidative phosphorylation
2 FADH2 3 Oxidative phosphorylation
Total yield 30 or 32 ATP From the complete oxidation of one glucose molecule to carbon dioxide and oxidation of all the reduced coenzymes.

Although there is a theoretical yield of 38 ATP molecules per glucose during cellular respiration, such conditions are generally not realized because of losses such as the cost of moving pyruvate (from glycolysis), phosphate, and ADP (substrates for ATP synthesis) into the mitochondria. All are actively transported using carriers that utilize the stored energy in the proton electrochemical gradient.

  • Pyruvate is taken up by a specific, low Km transporter to bring it into the mitochondrial matrix for oxidation by the pyruvate dehydrogenase complex.
  • The phosphate carrier (PiC) mediates the electroneutral exchange (antiport) of phosphate (H2PO4; Pi) for OH or symport of phosphate and protons (H+) across the inner membrane, and the driving force for moving phosphate ions into the mitochondria is the proton motive force.
  • The ATP-ADP translocase (also called adenine nucleotide translocase, ANT) is an antiporter and exchanges ADP and ATP across the inner membrane. The driving force is due to the ATP (−4) having a more negative charge than the ADP (−3), and thus it dissipates some of the electrical component of the proton electrochemical gradient.

The outcome of these transport processes using the proton electrochemical gradient is that more than 3 H+ are needed to make 1 ATP. Obviously this reduces the theoretical efficiency of the whole process and the likely maximum is closer to 28–30 ATP molecules.[2] In practice the efficiency may be even lower because the inner membrane of the mitochondria is slightly leaky to protons.[5] Other factors may also dissipate the proton gradient creating an apparently leaky mitochondria. An uncoupling protein known as thermogenin is expressed in some cell types and is a channel that can transport protons. When this protein is active in the inner membrane it short circuits the coupling between the electron transport chain and ATP synthesis. The potential energy from the proton gradient is not used to make ATP but generates heat. This is particularly important in brown fat thermogenesis of newborn and hibernating mammals.

Cellular respiration
Stoichiometry of aerobic respiration and most known fermentation types in eucaryotic cell. [6] Numbers in circles indicate counts of carbon atoms in molecules, C6 is glucose C6H12O6, C1 carbon dioxide CO2. Mitochondrial outer membrane is omitted.

According to some of newer sources the ATP yield during aerobic respiration is not 36–38, but only about 30–32 ATP molecules / 1 molecule of glucose [6], because:

  • ATP : NADH+H+ and ATP : FADH2 ratios during the oxidative phosphorylation appear to be not 3 and 2, but 2.5 and 1.5 respectively. Unlike in the substrate-level phosphorylation, the stoichiometry here is difficult to establish.
    • ATP synthase produces 1 ATP / 3 H+. However the exchange of matrix ATP for cytosolic ADP and Pi (antiport with OH or symport with H+) mediated by ATP–ADP translocase and phosphate carrier consumes 1 H+ / 1 ATP as a result of regeneration of the transmembrane potential changed during this transfer, so the net ratio is 1 ATP : 4 H+.
    • The mitochondrial electron transport chain proton pump transfers across the inner membrane 10 H+ / 1 NADH+H+ (4 + 2 + 4) or 6 H+ / 1 FADH2 (2 + 4).
So the final stoichiometry is
1 NADH+H+ : 10 H+ : 10/4 ATP = 1 NADH+H+ : 2.5 ATP
1 FADH2 : 6 H+ : 6/4 ATP = 1 FADH2 : 1.5 ATP
  • ATP : NADH+H+ coming from glycolysis ratio during the oxidative phosphorylation is
    • 1.5, as for FADH2, if hydrogen atoms (2H++2e) are transferred from cytosolic NADH+H+ to mitochondrial FAD by the glycerol phosphate shuttle located in the inner mitochondrial membrane.
    • 2.5 in case of malate-aspartate shuttle transferring hydrogen atoms from cytosolic NADH+H+ to mitochondrial NAD+

So finally we have, per molecule of glucose

Altogether this gives 4 + 3 (or 5) + 20 + 3 = 30 (or 32) ATP per molecule of glucose

These figures may still require further tweaking as new structural details become available. The above value of 3 H+/ATP for the synthase assumes that the synthase translocates 9 protons, and produces 3 ATP, per rotation. The number of protons depends on the number of c subunits in the Fo c-ring, and it is now known that this is 10 in yeast Fo[7] and 8 for vertebrates.[8] Including one H+ for the transport reactions, this means that synthesis of one ATP requires 1+10/3=4.33 protons in yeast and 1+8/3 = 3.67 in vertebrates. This would imply that in human mitochondria the 10 protons from oxidizing NADH would produce 2.72 ATP (instead of 2.5) and the 6 protons from oxidizing succinate or ubiquinol would produce 1.64 ATP (instead of 1.5). This is consistent with experimental results within the margin of error described in a recent review.[9]

The total ATP yield in ethanol or lactic acid fermentation is only 2 molecules coming from glycolysis, because pyruvate is not transferred to the mitochondrion and finally oxidized to the carbon dioxide (CO2), but reduced to ethanol or lactic acid in the cytoplasm.[6]


Without oxygen, pyruvate (pyruvic acid) is not metabolized by cellular respiration but undergoes a process of fermentation. The pyruvate is not transported into the mitochondrion, but remains in the cytoplasm, where it is converted to waste products that may be removed from the cell. This serves the purpose of oxidizing the electron carriers so that they can perform glycolysis again and removing the excess pyruvate. Fermentation oxidizes NADH to NAD+ so it can be re-used in glycolysis. In the absence of oxygen, fermentation prevents the buildup of NADH in the cytoplasm and provides NAD+ for glycolysis. This waste product varies depending on the organism. In skeletal muscles, the waste product is lactic acid. This type of fermentation is called lactic acid fermentation. In strenuous exercise, when energy demands exceed energy supply, the respiratory chain cannot process all of the hydrogen atoms joined by NADH. During anaerobic glycolysis, NAD+ regenerates when pairs of hydrogen combine with pyruvate to form lactate. Lactate formation is catalyzed by lactate dehydrogenase in a reversible reaction. Lactate can also be used as an indirect precursor for liver glycogen. During recovery, when oxygen becomes available, NAD+ attaches to hydrogen from lactate to form ATP. In yeast, the waste products are ethanol and carbon dioxide. This type of fermentation is known as alcoholic or ethanol fermentation. The ATP generated in this process is made by substrate-level phosphorylation, which does not require oxygen.

Fermentation is less efficient at using the energy from glucose: only 2 ATP are produced per glucose, compared to the 38 ATP per glucose nominally produced by aerobic respiration. This is because the waste products of fermentation still contain chemical potential energy that can be released by oxidation. Ethanol, for example, can be burned in an internal combustion engine like gasoline. Glycolytic ATP, however, is created more quickly. For prokaryotes to continue a rapid growth rate when they are shifted from an aerobic environment to an anaerobic environment, they must increase the rate of the glycolytic reactions. For multicellular organisms, during short bursts of strenuous activity, muscle cells use fermentation to supplement the ATP production from the slower aerobic respiration, so fermentation may be used by a cell even before the oxygen levels are depleted, as is the case in sports that do not require athletes to pace themselves, such as sprinting.

Anaerobic respiration

Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor (such as oxygen) to produce large amounts of energy, to drive the bulk production of ATP.

Anaerobic respiration is used by some microorganisms in which neither oxygen (aerobic respiration) nor pyruvate derivatives (fermentation) is the final electron acceptor. Rather, an inorganic acceptor such as sulfate or nitrate is used. Such organisms are typically found in unusual places such as underwater caves or near hydrothermal vents at the bottom of the ocean.

In July 2019, a scientific study of Kidd Mine in Canada discovered sulfur-breathing organisms which live 7900 feet below the surface, and which breathe sulfur in order to survive. these organisms are also remarkable due to eating rocks such as pyrite as their regular food source. [10][11][12]

See also


  1. ^ Bailey, Regina. "Cellular Respiration". Archived from the original on 2012-05-05.
  2. ^ a b c Rich, P. R. (2003). "The molecular machinery of Keilin's respiratory chain". Biochemical Society Transactions. 31 (Pt 6): 1095–1105. doi:10.1042/BST0311095. PMID 14641005.
  3. ^ Reece1 Urry2 Cain3 Wasserman4 Minorsky5 Jackson6, Jane1 Lisa2 Michael3 Steven4 Peter5 Robert6 (2010). Campbell Biology Ninth Edition. Pearson Education, Inc. p. 168.
  4. ^ a b "Cellular Respiration" (PDF). Archived (PDF) from the original on 2017-05-10.
  5. ^ Porter, R.; Brand, M. (1 September 1995). "Mitochondrial proton conductance and H+/O ratio are independent of electron transport rate in isolated hepatocytes". The Biochemical Journal (Free full text). 310 (Pt 2): 379–382. doi:10.1042/bj3100379. ISSN 0264-6021. PMC 1135905. PMID 7654171.
  6. ^ a b c Stryer, Lubert (1995). Biochemistry (fourth ed.). New York – Basingstoke: W. H. Freeman and Company. ISBN 978-0716720096.
  7. ^ Stock D, Leslie AG, Walker JE (1999). "Molecular architecture of the rotary motor in ATP synthase". Science. 286: 1700–5. doi:10.1126/science.286.5445.1700. PMID 10576729.CS1 maint: uses authors parameter (link)
  8. ^ Watt, I.N., Montgomery, M.G., Runswick, M.J., Leslie, A.G.W., Walker, J.E. (2010). "Bioenergetic Cost of Making an Adenosine Triphosphate Molecule in Animal Mitochondria". Proc. Natl. Acad. Sci. USA. 107: 16823. doi:10.1073/pnas.1011099107. PMC 2947889. PMID 20847295.CS1 maint: uses authors parameter (link)
  9. ^ P.Hinkle (2005). "P/O ratios of mitochondrial oxidative phosphorylation". Biochimica et Biophysica Acta. 1706: 1–11. doi:10.1016/j.bbabio.2004.09.004. PMID 15620362.
  10. ^ ‘Follow the Water’: Hydrogeochemical Constraints on Microbial Investigations 2.4 km Below Surface at the Kidd Creek Deep Fluid and Deep Life Observatory, Garnet S. Lollar, Oliver Warr, Jon Telling, Magdalena R. Osburn & Barbara Sherwood Lollar, Received 15 Jan 2019, Accepted 01 Jul 2019, Published online: 18 Jul 2019.
  11. ^ World’s Oldest Groundwater Supports Life Through Water-Rock Chemistry, July 29, 2019, deepcarbon.net.
  12. ^ Strange life-forms found deep in a mine point to vast 'underground Galapagos', By Corey S. Powell, Sept. 7, 2019, nbcnews.com.

External links

Adenosine diphosphate

Adenosine diphosphate (ADP), also known as adenosine pyrophosphate (APP), is an important organic compound in metabolism and is essential to the flow of energy in living cells. ADP consists of three important structural components: a sugar backbone attached to adenine and two phosphate groups bonded to the 5 carbon atom of ribose. The diphosphate group of ADP is attached to the 5’ carbon of the sugar backbone, while the adenosine attaches to the 1’ carbon.ADP can be interconverted to adenosine triphosphate (ATP) and adenosine monophosphate (AMP). ATP contains one more phosphate group than does ADP. AMP contains one fewer phosphate group. Energy transfer used by all living things is a result of dephosphorylation of ATP by enzymes known as ATPases. The cleavage of a phosphate group from ATP results in the coupling of energy to metabolic reactions and a by-product of ADP. Being the "molecular unit of currency", ATP is continually being reformed from lower-energy species ADP and AMP. The biosynthesis of ATP is achieved throughout processes such as substrate-level phosphorylation, oxidative phosphorylation, and photophosphorylation, all of which facilitating the addition of a phosphate group to ADP.


Aerobic means "requiring air," in which "air" usually means oxygen.

Aerobic may also refer to

Aerobic exercise, prolonged exercise of moderate intensity

Aerobics, a form of aerobic exercise

Aerobic respiration, the aerobic process of cellular respiration

Aerobic organism, a living thing with an oxygen-based metabolism

Aerobic organism

An aerobic organism or aerobe is an organism that can survive and grow in an oxygenated environment. In contrast, an anaerobic organism (anaerobe) is any organism that does not require oxygen for growth. Some anaerobes react negatively or even die if oxygen is present.

Anaerobic respiration

Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain.In aerobic organisms undergoing respiration, electrons are shuttled to an electron transport chain, and the final electron acceptor is oxygen. Molecular oxygen is a highly oxidizing agent and, therefore, is an excellent electron acceptor. In anaerobes, other less-oxidizing substances such as sulphate (SO42−), nitrate (NO3−), sulphur (S), or fumarate are used. These terminal electron acceptors have smaller [[reduction potential]s than O2, meaning that less energy is released per oxidized molecule. Therefore, generally speaking, anaerobic respiration is less efficient than aerobic.

Anaplerotic reactions

Anaplerotic reactions (from the Greek ἀνά= 'up' and πληρόω= 'to fill') are chemical reactions that form intermediates of a metabolic pathway. Examples of such are found in the citric acid cycle (TCA cycle). In normal function of this cycle for respiration, concentrations of TCA intermediates remain constant; however, many biosynthetic reactions also use these molecules as a substrate. Anaplerosis is the act of replenishing TCA cycle intermediates that have been extracted for biosynthesis (in what are called anaplerotic reactions).

The TCA cycle is a hub of metabolism, with central importance in both energy production and biosynthesis. Therefore, it is crucial for the cell to regulate concentrations of TCA cycle metabolites in the mitochondria. Anaplerotic flux must balance cataplerotic flux in order to retain homeostasis of cellular metabolism.

Carbohydrate metabolism

Carbohydrate metabolism is the whole of biochemical processes responsible for the metabolic formation, breakdown, and interconversion of carbohydrates in living organisms.

Carbohydrates are central to many essential metabolic pathways. Plants synthesize carbohydrates from carbon dioxide and water through photosynthesis, allowing them to store energy absorbed from sunlight internally. When animals and fungi consume plants, they use cellular respiration to break down these stored carbohydrates to make energy available to cells. Both animals and plants temporarily store the released energy in the form of high energy molecules, such as ATP, for use in various cellular processes.Although humans consume a variety of carbohydrates, digestion breaks down complex carbohydrates into a few simple monomers (monosaccharides) for metabolism: glucose, fructose, and galactose. Glucose constitutes about 80% of the products, and is the primary structure that is distributed to cells in the tissues, where it is broken down or stored as glycogen. In aerobic respiration, the main form of cellular respiration used by humans, glucose and oxygen are metabolized to release energy, with carbon dioxide and water as byproducts. Most of the fructose and galactose travel to the liver, where they can be converted to glucose.Some simple carbohydrates have their own enzymatic oxidation pathways, as do only a few of the more complex carbohydrates. The disaccharide lactose, for instance, requires the enzyme lactase to be broken into its monosaccharide components, glucose and galactose.


Chemiosmosis is the movement of ions across a semipermeable membrane, down their electrochemical gradient. An example of this would be the generation of adenosine triphosphate (ATP) by the movement of hydrogen ions (H+) across a membrane during cellular respiration or photosynthesis.

Hydrogen ions, or protons, will diffuse from an area of high proton concentration to an area of lower proton concentration, and an electrochemical concentration gradient of protons across a membrane can be harnessed to make ATP. This process is related to osmosis, the diffusion of water across a membrane, which is why it is called "chemiosmosis".

ATP synthase is the enzyme that makes ATP by chemiosmosis. It allows protons to pass through the membrane and uses the free energy difference to phosphorylate adenosine diphosphate (ADP), making ATP. The generation of ATP by chemiosmosis occurs in mitochondria and chloroplasts, as well as in most bacteria and archaea, an electron transport chain pumps H+ ions in the thylakoid spaces through thylakoid membranes to stroma (fluid). The energy from the electron movement through electron transport chains cross through ATP synthase which allows the proton to pass through them and use this free energy difference to photophosphorylate ADP making ATP.


A crista (; plural cristae) is a fold in the inner membrane of a mitochondrion. The name is from the Latin for crest or plume, and it gives the inner membrane its characteristic wrinkled shape, providing a large amount of surface area for chemical reactions to occur on. This aids aerobic cellular respiration, because the mitochondrion requires oxygen. Cristae are studded with proteins, including ATP synthase and a variety of cytochromes.

With the discovery of the dual-membrane nature of mitochondria, the pioneers of mitochondrial ultrastructural research proposed different models for the organization of the mitochondrial inner membrane. Three models proposed were:

Baffle model – According to Palade, the mitochondrial inner membrane is convoluted in a baffle-like manner with broad openings towards the intra-cristal space. This model entered most textbooks and was widely believed for a long time.

Septa model – Sjöstrand suggested that sheets of inner membrane are spanned like septa (plural of septum) through the matrix, separating it into several distinct compartments.

Crista junction model – Daems and Wisse proposed that cristae are connected to the inner boundary membrane via tubular structures characterized by rather small diameters, termed crista junctions (CJs). These structures were rediscovered recently by EM tomography, leading to the establishment of this currently widely accepted model.

D-amino acid dehydrogenase

D-amino-acid dehydrogenase (EC is a bacterial enzyme that catalyses the oxidation of D-amino acids into their corresponding oxoacids. It contains both flavin and nonheme iron as cofactors. The enzyme has a very broad specificity and can act on most D-amino acids.D-amino acid + H2O + acceptor <=> a 2-oxo acid + NH3 + reduced acceptor

This reaction is distinct from the oxidation reaction catalysed by D-amino acid oxidase that uses oxygen as a second substrate, as the dehydrogenase can use many different compounds as electron acceptors, with the physiological substrate being coenzyme Q.

Facultative anaerobic organism

A facultative anaerobe is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent.

Some examples of facultatively anaerobic bacteria are Staphylococcus spp., Streptococcus spp., Escherichia coli, Salmonella, Listeria spp., Shewanella oneidensis and Yersinia pestis. Certain eukaryotes are also facultative anaerobes, including fungi such as Saccharomyces cerevisiae and many aquatic invertebrates such as Nereid (worm) polychaetes.

Flavin group

Flavin (from Latin flavus, "yellow") is the common name for a group of organic compounds based on pteridine, formed by the tricyclic heterocycle isoalloxazine. The biochemical source is the vitamin riboflavin. The flavin moiety is often attached with an adenosine diphosphate to form flavin adenine dinucleotide (FAD), and, in other circumstances, is found as flavin mononucleotide (or FMN), a phosphorylated form of riboflavin. It is in one or the other of these forms that flavin is present as a prosthetic group in flavoproteins.

The flavin group is capable of undergoing oxidation-reduction reactions, and can accept either one electron in a two-step process or two electrons at once. Reduction is made with the addition of hydrogen atoms to specific nitrogen atoms on the isoalloxazine ring system:

In aqueous solution, flavins are yellow-coloured when oxidized, taking a red colour in the semi-reduced anionic state or blue in the neutral (semiquinone) state, and colourless when totally reduced. The oxidized and reduced forms are in fast equilibrium with the semiquinone (radical) form, shifted against the formation of the radical:

Flox + FlredH2 ⇌ FlH•where Flox is the oxidized flavin, FlredH2 the reduced flavin (upon addition of two hydrogen atoms) and FlH• the semiquinone form (addition of one hydrogen atom).

In the form of FADH2, it is one of the cofactors that can transfer electrons to the electron transfer chain.

Food energy

Food energy is chemical energy that animals (including humans) derive from food through the process of cellular respiration. Cellular respiration may either involve the chemical reaction of food molecules with molecular oxygen (aerobic respiration) or the process of reorganizing the food molecules without additional oxygen (anaerobic respiration).

Glycerol phosphate shuttle

The glycerol-3-phosphate shuttle is a mechanism that regenerates NAD+ from NADH, a by-product of glycolysis. Its importance in transporting reducing equivalents is secondary to the malate-aspartate shuttle.

Malate-aspartate shuttle

The malate-aspartate shuttle (sometimes simply the malate shuttle) is a biochemical system for translocating electrons produced during glycolysis across the semipermeable inner membrane of the mitochondrion for oxidative phosphorylation in eukaryotes. These electrons enter the electron transport chain of the mitochondria via reduction equivalents to generate ATP. The shuttle system is required because the mitochondrial inner membrane is impermeable to NADH, the primary reducing equivalent of the electron transport chain. To circumvent this, malate carries the reducing equivalents across the membrane.

Mitochondrial shuttle

The mitochondrial shuttles are systems used to transport reducing agents across the inner mitochondrial membrane. NADH cannot cross the membrane, but it can reduce another molecule that can cross the membrane, so that its electrons can reach the electron transport chain.

The two main systems in humans are:

In humans, the glycerol phosphate shuttle is primarily found in brown adipose tissue, as the conversion is less efficient, thus generating heat, which is one of the main purposes of brown fat. It is primarily found in babies, though it is present in small amounts in adults around the kidneys and on the back of our necks. The malate-aspartate shuttle is found in much of the rest of the body.


Nanaerobes are organisms that cannot grow in the presence of micromolar concentrations of oxygen, but can grow with and benefit from the presence of nanomolar concentrations of oxygen (e.g. Bacteroides fragilis). Like other anaerobes, these organisms do not require oxygen for growth. This growth benefit requires the expression of an oxygen respiratory chain that is typically associated with microaerophilic respiration. Recent studies suggest that respiration in low concentrations of oxygen is an ancient process which predates the emergence of oxygenic photosynthesis.

Pyruvate dehydrogenase complex

Pyruvate dehydrogenase complex (PDC) is a complex of three enzymes that converts pyruvate into acetyl-CoA by a process called pyruvate decarboxylation. Acetyl-CoA may then be used in the citric acid cycle to carry out cellular respiration, and this complex links the glycolysis metabolic pathway to the citric acid cycle. Pyruvate decarboxylation is also known as the "pyruvate dehydrogenase reaction" because it also involves the oxidation of pyruvate.This multi-enzyme complex is related structurally and functionally to the oxoglutarate dehydrogenase and branched-chain oxo-acid dehydrogenase multi-enzyme complexes.

Respiration (physiology)

In physiology, respiration is the movement of oxygen from the outside environment to the cells within tissues, and the transport of carbon dioxide in the opposite direction.

The physiological definition of respiration differs from the biochemical definition, which refers to a metabolic process by which an organism obtains energy (in the form of ATP and NADPH) by oxidising nutrients and releasing waste products. Although physiologic respiration is necessary to sustain cellular respiration and thus life in animals, the processes are distinct: cellular respiration takes place in individual cells of the organism, while physiologic respiration concerns the diffusion and transport of metabolites between the organism and the external environment.

In animals with lungs, physiological respiration involves respiratory cycles of inhaled and exhaled breaths. Inhalation (breathing in) is usually an active movement. The contraction of the diaphragm muscle cause a pressure variation, which is equal to the pressures caused by elastic, resistive and inertial components of the respiratory system. In contrast, exhalation (breathing out) is usually a passive process. Breathing in brings air into the lungs where the process of gas exchange takes place between the air in the alveoli and the blood in the pulmonary capillaries

The process of breathing does not fill the alveoli with atmospheric air during each inhalation (about 350 ml per breath), but the inhaled air is carefully diluted and thoroughly mixed with a large volume of gas (about 2.5 liters in adult humans) known as the functional residual capacity which remains in the lungs after each exhalation, and whose gaseous composition differs markedly from that of the ambient air. Physiological respiration involves the mechanisms that ensure that the composition of the functional residual capacity is kept constant, and equilibrates with the gases dissolved in the pulmonary capillary blood, and thus throughout the body. Thus, in precise usage, the words breathing and ventilation are hyponyms, not synonyms, of respiration; but this prescription is not consistently followed, even by most health care providers, because the term respiratory rate (RR) is a well-established term in health care, even though it would need to be consistently replaced with ventilation rate if the precise usage were to be followed.


A triose is a monosaccharide, or simple sugar, containing three carbon atoms. There are only three possible trioses (including dihydroxyacetone): L-glyceraldehyde and D-glyceraldehyde, the two enantiomers of glyceraldehyde, which are aldotrioses because the carbonyl group is at the end of the chain, and dihydroxyacetone, the only ketotriose, which is symmetrical and therefore has no enantiomers.Trioses are important in cellular respiration. During glycolysis, fructose-1,6-bisphosphate is broken down into glyceraldehyde-3-phosphate and dihydroxyacetone phosphate. Lactic acid and pyruvic acid are later derived from these molecules.



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