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admin | Natural Weight Loss Supplement | 14.07.2014
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ATP is often described as a ?high energy? compound, but hydrolysis of ATP to ADP or AMP yields only modest amounts of energy.
In general, living cells avoid big changes in free energy, because enzymes are only held together by weak hydrogen bonds and hydrophobic interactions, and it is difficult to build them strong enough to withstand the thump from a violent reaction. Allowing for a few leaks and overheads, complete oxidation of a molecule of glucose would yield about 25 - 30 molecules of ATP and the energy recovery would be about 50%.
About half the total energy available from food oxidation is ?captured? in the form of ATP. Under aerobic conditions 95% of the cellular ATP is produced within the mitochondria, which actively scavenge the cytosol for ADP, exporting ATP in exchange. Not only is the mitochondrial ATP factory much more efficient than the other parts of the cell, but all the major energy-yielding pathways are concentrated within the mitochondria. Mitochondria can really 'Hoover' the place clean, so that it is difficult to detect any free ADP in a eukaryotic cell. The energy available from splitting ATP depends on how far the hydrolysis reaction is displaced from equilibrium.
The mathematical relationship between DG and the distance from equilibrium was first described by the 19th century American chemist J. Standard DG0 values are measured with everything at 1 molar concentration, but the actual concentrations in living cells will differ from this, and the effective DG values may vary in different parts of the cell. Eukaryotic cells do not, however, get something for nothing by pumping ATP from the mitochondria into the cytosol. Mitochondria from actively respiring tissues like cardiac muscle have a larger internal membrane area that mitochondria from tissues (like liver) with a lower rate of oxygen consumption. One of the special phospholipids in the mitochondrial inner membrane is called cardiolipin, or disphosphatidyl glycerol. Acetyl-CoA produced from either fats or carbohydrates is completely oxidised to carbon dioxide and water by the Krebs (citric acid) cycle in the mitochondrial matrix (see Nelson & Cox pages 571-584). Isocitrate: Isocitrate is present at only about 5% of the citrate concentration because the aconitase equilibrium favours citrate. Oxoglutarate: This 2-keto-acid is chemically similar to pyruvate and it is metabolised in a similar way.
Succinate: This dibasic acid is a relatively poor reducing agent, which cannot convert NAD to NADH, so it requires a special flavoprotein dehydrogenase directly linked to the electron transport chain.
Malate: In addition to its role in the citric acid cycle, this dibasic acid is a major metabolic intermediate which is involved in gluconeogenesis, lipogenesis and mitochondrial redox shuttles.
Citrate synthase: Although this enzyme is the point where acetyl-CoA enters the Krebs cycle, it merely converts one cycle intermediate into another, and does not lead to any overall change in the total pool of cycle intermediates. Isocitrate dehydrogenase: Isocitrate is a good reducing agent, so its oxidation proceeds with a highly favourable delta G and is a major point for citric acid cycle regulation.
Oxoglutarate dehydrogenase: This huge multi-enzyme complex is structurally related to pyruvate dehydrogenase, although it is regulated in a different manner.
Succinate thiokinase: This enzyme catalyses the only substrate-level phosphorylation in the citric acid cycle.
Succinate dehydrogenase: This flavoprotein is an integral part of the mitochondrial inner memberane, where it forms part of the respiratory chain. Fumarase: This enzyme is freely reversible and probably operates very close equilibrium under physiological conditions. Malate dehydrogenase: This enzyme is freely reversible, and can operate in either direction in different circumstances. Acetyl-CoA: This high-energy compound can be formed from the oxidation of carbohydrates, amino acids and fats. Pyruvate dehydrogenase: This huge multi-enzyme complex catalyses the irreversible conversion of pyruvate into acetyl-CoA. Pyruvate carboxylase: This important regulatory enzyme is activated by acetyl-CoA and helps to control the total amount of citric acid cycle intermediates. Point (don't click) at this image with your mouse to view additional information about the Krebs cycle. The Krebs cycle and the respiratory chain form the final common pathway for substrate oxidation in eukaryotic cells, and the system generates about 95% of the cell's ATP supply. The inner and outer mitochondrial membranes differ enormously in their chemical composition and biological functions. Mitochondria cooperate with other organelles in order to achieve the complete breakdown of foodstuffs to CO2 and water.
Most of the cellular lipids contain linear C16 and C18 fatty acids which are activated in the cytosol, and then transfered to the mitochondria as acyl-carnitine derivatives. Once inside the mitochondria, acyl-CoA is degraded stepwise using β-oxidation to acetyl CoA fragments, producing large amounts of NADH and FADH2. Fat metabolism nicely illustrates the metabolic specialisation and cooperation between compartments. Although the peroxisomal fat oxidation "spiral" looks similar to the mitochondrial pathway, it is not connected to the respiratory chain and it does not directly generate any ATP. Peroxisomal oxidation stops at 8 carbon atoms, so the octanoyl-CoA and acetyl-CoA products of peroxisomal lipid breakdown are converted to the corresponding carnitine derivatives and shuttled across to the mitochondria to complete their conversion to carbon dioxide and water. There has recently been great research interest in the various fat oxidation pathways, and in the control of gene expression by the peroxisome proliferator activated receptors (PPARs). Subcellular compartments increase the controllability and efficiency of eukaryotic cells by segregating key metabolites and keeping related processes together. There are SEVEN main compartments without counting the internal subdivisions within the organelles. Metabolic processes are precisely divided between the different compartments, and even when the same reaction is observed in two separate spaces, they are found to serve different functions. Electroneutral and electrical carriers differ hugely in their transport properties and we will return to this topic in subsequent lectures. The diagram below shows the malate - aspartate cycle, which normally accomplishes the re-oxidation of the NADH generated in the cytosol during aerobic glycolysis.
NAD and NADP both have the same standard redox potentials, which is calculated with equal amounts of the oxidised and reduced forms. Mitochondria have their own circular DNA, RNA and protein synthesis, which are all built on bacterial rather than eukaryotic lines. The mitochondrial chromosome is very small, but there are hundreds of copies in a typical cell.
Maternal inheritance arises because the sperm is so much smaller than the egg, so on fertilisation any mitochondria contributed by the sperm are completely swamped by the thousands of copies in the egg. Mitochondrial DNA mutations may affect only part of the body, a condition known as "heteroplasmy".
Almost all the ancestral bacterial genes were long since ?ripped? from the mitochondria and copied into nuclear DNA where they are easier to control.
The only protein coding genes remaining on the mitochondrial chromosome specify sticky hydrophobic proteins at the core of the mitochondrial inner membrane. In humans, the mitochondrial chromosome has only 13 protein-coding genes which specify a few of the core subunits from respiratory complexes I, III and IV, and part of ATP synthase. The mitochondrial organisation is destroyed in the membrane permeability transition, which is a key event during apoptosis, or programmed cell death. Apoptosis or programmed cell death is an ancient strategy for killing defective or unwanted cells which is probably present in all multicellular organisms, both animals and plants.

Intracellular apoptotic signalling exploits amplifying cascades of cysteine proteases called caspases that activate their protein substrates by cleaving them after aspartate residues. PARP stands for poly(ADP-ribose) polymerase, an enzyme which ADP-ribosylates a wide variety of nuclear proteins, using NAD as the ADP-ribosyl donor.
The life or death decision depends on a continuously shifting balance between pro- and anti-apoptotic factors.
There is cross-talk between the different pathways, so (for example) granzymes and caspase 8 can both activate the mitochondrial route, and are consequently subject to modulation by Bcl-2.
Oxygen is a powerful oxidant, which readily generates free radicals that damage many biological molecules. In reality, there is stacks of it, and with four negative charges it makes a significant contribution to the total anion content of each cell.
Although small, this additional energy tips the balance and is sufficient to drive many unfavourable processes in the required direction. The energy released from ATP hydrolysis is not much bigger than the peaks of thermal energy that are constantly rattling molecules around at body heat.
You may see slightly larger figures in some textbooks, but I think these are a bit optimistic. A reasonably active person (12MJ diet) turns over about 75kg of ATP every day, so a typical ATP molecule is broken down and resynthesised 1000 times each day. Of this ATP, about half the free energy can be converted into useful work, such as muscle contraction. In theory, a cell's energy pathways could achieve almost 100% efficiency, providing that they ran infinitely slowly. There is a fair amount of structural ADP permanently bound onto the cytoskeletal protein actin, but that is not available for use. The transport of ATP, ADP and phosphate across the inner mitochondrial membrane costs them 33% additional energy, over the bare minimum required for the synthesis of ATP within the mitochondrial compartment. It has a large fixed negative charge and four fatty acid side chains, instead of the usual two. It contains Krebs cycle enzymes, and the later stages of carbohydrate, fat and amino acid breakdown. These reactions generate 3 molecules of NADH and one of FADH all of which are re-cycled back to form NAD and FAD by the respiratory chain in the inner mitochondrial membrane. It is a substrate for the tricarboxylate antiporter present in liver and adipocyte mitochondria, so it can be exported to the cytosol and used for lipid synthesis.
Although isocitrate resembles citrate in many of its physical properties, chemically it is very different and is an excellent reducing agent. It is structurally related to glutamate, and is an important point for amino acid degradation products to enter the Krebs cycle. Succinate is a substrate for the dicarboxylate antiporter although it has no known cytosolic functions in most tissues.
Lack of oxaloacetate may limit Krebs cycle activity during periods of rapid fat oxidation, and this is an important factor promoting hepatic ketogenesis under these conditions. The NAD-linked isocitrate dehydrogenase is a large allosteric protein that is activated by calcium ions and ADP and inhibited by NAD(P)H. It contains three different types of protein subunit, which use thiamine pyrophosphate, FAD and lipoic acid as prosthetic groups. Under fasting conditions mitochondrial acetyl-CoA is either oxidised via the citric acid cycle, or it may be converted into acetoacetate by liver mitochondria and exported to the remainder of the body. It is structurally related to oxoglutarate dehydrogenase, although it is regulated in a different manner. We don't expect you to rote-learn this material, but we do expect you to recognise the main compounds, and appreciate their close relationship to key intermediates in amino-acid, carbohydrate and fat metabolism. In addition the Krebs cycle is the central clearing house for intermediary metabolism, and provides the essential interface between amino acids, carbohydrates and fats.
It contains numerous respiratory enzymes used for oxygen uptake, proteins involved in metabolite transport and proteins required for the manufacture and export of ATP. The initial stages of fatty acid activation are performed in the cytosol, and complex lipids are degraded in peroxisomes before the fragments are passed to the mitochondria for final processing. This can sometimes be treated by feeding short- or medium-chain length fatty acids, which can enter the mitochondria without using the carnitine shuttle. Fat and steroid biosynthesis takes place mainly in the cytosol, using acetyl-CoA derived from citrate cleavage, although subsequent steroid and bile acid processing occurs in the mitochondrial matrix space. The first enzyme, acyl-CoA oxidase, is a flavoprotein oxidase that converts oxygen to hydrogen peroxide.
This system is intimately connected with the development of obesity and type 2 diabetes, which are major medical problems in the modern world. If you were to include all these plus the various types of transport and storage vesicles there would be about twenty separate spaces. There is also great specialisation between the various tissues, which we will study in the later lectures. Complete the table below: "most" tissues means all tissues except red blood cells, "many" means all tissues except for red blood cells, adipocytes and liver.
Most of the membrane transport proteins catalyse electroneutral exchanges with other anions or hydroxide ions, so the resulting anion distribution is largely determined by the intracellular pH gradients.
Elaborate systems exist for transporting many hundreds of components that travel between the two compartments. Membrane carrier systems are needed whenever a metabolic pathway crosses from one compartment to another, and they are often expressed in a tissue-specific pattern. This allows cells to keep their cytosol more oxidising than their mitochondria, which suppresses lactate production under aerobic conditions.
It is necessary because neither NAD nor NADH can directly cross the mitochondrial inner membrane. The actual redox potentials inside living cells are very different, because cells actively keep these ratios a long way from 1:1 There is a redox pump inside the mitochondria called the energy linked transhydrogenase which actively transfers hydride ions from NADH to NADP+ using energy derived from mitochondrial respiration. It is claimed that all the mitochondria in modern humans derive from an African lady "mitochondrial Eve" who lived about 200,000 years ago.
One possible explanation is that the mutation occured during cleavage of the embryo, with the result that only part of the body, or a restricted group of tissues received the mutated stock. Most mitochondrial proteins are synthesised on cytosolic ribosomes, and laboriously imported across the outer and the inner mitochondrial membranes.
Proteins must be correctly targetted for the matrix space, inner membrane, inter-membrane space or outer membrane. These integral membrane proteins spontaneously insert into the first phospholipid bilayer they encounter, so the only safe place to express them is in the interior of the mitochondrial matrix space, where there is limited opportunity to get it wrong. In addition, it codes for ribosomal RNA and tRNAs, but the DNA and RNA polymerases, and all the ribosomal proteins are imported from the cytosol! Various membrane components are reorganised to form a large pore which permits the escape of cytochrome c. A major event in some of these signalling pathways is the mitochondrial permeability transition, which involves a bizarre change in mitochondrial compartmentation. Radiation and cytotoxic drugs are effective against susceptible tumours because these cells are already teetering on the brink of self-destruction, and the slightest push will tilt the balance. Once triggered, apoptosis is auto-catalytic because activated caspase 3 can itself activate caspase 8, which can in turn initiate the mitochondrial permeability transition and the release of cytochrome c.
It can be a fiercely toxic gas, and pre-capillary sphincters normally regulate oxygen delivery to the tissues.
ATP was quickly discovered by the early biochemists because there is plenty of it around and it is difficult to miss. But for reactions that are already close to equilibrium, the extra input from ATP is still enough to bias the process in the desired direction.

As we will see in the next lecture, mitochondrial respiration is normally restricted by shortage of ADP. Eukaryotic cells increase the energy yield from each molecule of ATP by transporting ATP and ADP between cell compartments. It applies to all chemical reactions (in fact, to all physical processes) and in essence is little more than common sense. For different chemical reactions each graph will be displaced along the horizontal axis, but it will always have the same general shape.
This extra energy must be paid for by oxidising additional food, and supplied by the mitochondrial respiratory chain. It contains porin, an integral membrane protein that self-assembles into ?grommets? with a central hole. About a dozen metabolites can cross the inner membrane using highly specific protein carriers, but all other movements are blocked. These provide a very stable anchor, and prevent the membrane pulling to pieces under the huge electrical stress. In liver (but not in other tissues) it also contains part of the urea cycle, and the ancillary enzymes needed to transfer material between the major divisions of metabolism.
The respiratory chain is composed of flavoproteins, ubiquinone, non-haem iron proteins and cytochromes. Oxaloacetate is structurally related to aspartate and plays a major role in gluconeogenesis, lipogenesis and mitochondrial redox shuttles. Compartmentation increases metabolic efficiency by bringing substrates and enzymes closer together.
Movement of small molecules is catalysed and regulated by membrane transport proteins located in the inner mitochondrial membrane. Examine the transport stoichiometry and functions of the mitochondrial carriers tabulated above and try to decide which category they belong to. There are very few coenzyme transporters, and elaborate metabolite shuttle networks are used instead to move material from one compartment to another. This cycle must revolve twice for every molecule of glucose that completes the glycolytic pathway. The difference in effective redox potential between the two coenzymes is roughly equivalent to the energy stored in ATP.
Nucleic acid processing is less reliable in mitochondria and it is a mystery how these copies are normally kept in synchrony.
Other eukaryotic organelles may have been acquired in a similar fashion, from other free-living precursors that joined the eukaryotic federation. Protein import is expensive energetically, but it is nevertheless worthwhile because of the better opportunities for regulation that exist in the nuclear genome. This is a key signaling event in the apoptotic cascade, which is used to destroy tumour cells and invading viruses, and to reshape the body during embryogenesis and growth. The adenine nucleotide carrier and other proteins are incorporated into new transmembrane pores. The proto-oncogene Bcl-2 is an anti-apoptotic gene whose protein product inhibits the mitochondrial permeability transition. The cross-talk and positive feedback normally ensures a clear decision on whether each cell will live or die. Mitochondria have a great avidity for oxygen, and the free oxygen concentration inside most cells is usually kept very low. BioTechUSA представляет непосредственно сам креатин фосфат в виде добавки Creatine Phosphate 5000. Rather than one big heave at infrequent intervals, living cells use ATP to apply an endless series of little nudges at every point where the flow needs a boost.
Diffusion rate is proportional to concentration, so the real bottleneck is returning the "empties" for recycling. On the other hand, wasteful animals will get very hot and require excesssive amounts of food, so 50% efficiency may be the optimal compromise. If you begin with large amounts of reactants and no products then the reaction will proceed to equilibrium with formation of products, but conversely, starting with all products and no reactants the reaction will proceed in the opposite direction to the same final equilibrium position, where DG is zero. Barth syndrome is rare genetic problem, affecting only male children, where there is a defect in cardiolipin synthesis. Acetyl-CoA from all three sources is finally oxidised via the Krebs cycle within the mitochondrial matrix space. Xenobiotics, drugs and really awkward molecules get a preliminary free radical assault from the cytochrome p450 system anchored in the endoplasmic reticulum, which introduces additional hydroxyl groups and opens up new lines of chemical attack. Hydration and dehydrogenation of the enoyl-CoA are catalysed by a single enzyme, which produces NADH. These porters are highly specific for particular charged species, and often catalyse an exchange between two molecules such that their charges exactly balance. This creates holes in both mitochondrial membranes, collapsing the inner membrane potential, preventing ATP synthesis and releasing cytochrome c from the inter-membrane space into the cytosol. Inactivation of PARP implies that the cell has "given up" on DNA repair, and the cellular machinery now regards death as inevitable. It is named after B cell lymphomas, which are tumours of the MALT tissue in the small intestine where redundant B cells fail to die at the appropriate time.
The minuscule amount of ADP in working muscle has only milliseconds to get back from the contractile proteins to the mitochondria for reconversion into ATP.
There are very large ionic and electrical gradients across the inner membrane, which are exploited for the synthesis and export of ATP.
It conveys electrons stepwise from good reducing agents at the NADH end to good oxidising agents at the oxygen end. The dicarboxylic products of these reactions, together with branched chain fatty acids and very long chain linear fatty acids (>C20) transfer to the peroxisomes, where they are converted to coenzyme A derivatives which undergo a modified type of β-oxidation to reduce them to a manageable size. Peroxisomes lack a respiratory chain, and they are impermeable to pyridine nucleotides so the only available method to recycle the NADH back to NAD is through lactate dehydrogenase, as shown in the diagram below.
The cytosol is normally the largest compartment, except for adipocytes which are dominated by a huge globule of fat.
This suggests that mitochondria are descended from captured bacteria that were enslaved by our eukaryotic ancestors ? the endosymbiont hypothesis. You don't need all this detail for ICU3!) Apotosis is a major anti-cancer and anti-viral mechanism in humans, and it is also involved in growth and differentiation, heart attacks and stroke.
Once released, the cytochrome c associates with another protein called Apaf-1 and initiates a proteolytic cascade.
Despite elaborate measures to protect themselves from oxidation, they become irreversibly damaged and have to be discarded after about 4 months. This is a simple example of a shuttle system (see below) which are widely used to balance up metabolic pathways in different compartments.
In most tissues mitochondria are the second largest space (up to 40% of cell volume in cardiac muscle) and they play a central role in metabolism.
ATP дает энергию в процессе выпуска одной из своих фосфатных молекул, после чего превращается в ADP (аденозин трифосфат). Creatine Phosphate 5000 позволяет превратить ADP в ATP и таким образом обеспечивает мышцы дополнительной энергией.

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