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Introduction/training [organization of the study] [polarography] [calibrating] [research paper] Mitochondria theory: [overview] [structure] [Krebs reactions] [electron transport] [the gradient] [oxidative phosphorylation] Mitochondria in vitro: [preparation] [fate of substrates] [state IV] [state III] [metabolic poisons] [mitotraces] [rationale] [experiments] Additional topics: [glossary of terms ] [Hans Krebs] [origin of mitochondria] [other functions]
The standard reduction potential of NADH is -0.315V, while that of coenzyme Q is 0.045V (difference of 0.345 V). Therefore there is a strong 'pull' by Coenzyme Q on electrons through the components of Complex I. Just for the sake of understanding the principles, let Complex I (NADH dehydrogenase complex), embedded in an intact inner membrane, be the only component of an experimental electron transport system. We'll simply take the electrons from Coenzyme Q when they reach it, so the system can keep going. The removal of protons from the matrix and deposition of protons in the intermembrane space creates a concentration difference of protons across the inner membrane. This is called the chemiosmotic gradient. As the gradient builds up, more and more energy is required to push protons across. When the amount of energy required to push protons reaches 69.5 kJ/mole, electron transport has to stop. In fact, the second law of thermodynamics requires that electron transport stop before the gradient builds up to that point. www.ruf.rice.edu/~bioslabs/studies/mitochondria/mitogradient.html
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point.
If there was no way of draining energy from the system, electron transport could not continue despite the presence of adequate substrate. However, a mitochondrion is always in a steady state of respiration, in which the energy lost by processes that dissipate the gradient is constantly replaced by electron transport.
Respiratory Control
The limitation placed on electron transport by the chemisosmotic gradient is termed respiratory control. Mitochondria are said to exercise respiratory control as long as they can restrict electron transport by means of the gradient. If the gradient is destroyed by damaging the membranes, respiratory control is abolished and electron transport can run freely.
www.ruf.rice.edu/~bioslabs/studies/mitochondria/mitogradient.html
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balloon. Blowing with all your might you succeed in filling the balloon so that it is two feet in diameter. You cannot force any more air into it so all you are doing now is holding pressure. Now some joker comes along and pokes a couple of holes in your balloon, so that air leaks out at a controlled rate. You have maintained constant pressure, so now you find yourself moving air into the balloon as it leaks out. The balloon diameter, proportional to internal pressure, remains constant. If you plug one of the holes you move less air. Open up more holes and you move more air. The electron transport system applies constant "pressure," holding the gradient at a constant level. The rate of electron tranport (analagous to air flow in the balloon example) varies as energy is drained from the system at different rates. Adding ADP in vitro, for example, opens up avenues for protons to be forced into the matrix, draining energy from the gradient. Its addition is the equivalent of poking additional holes in your balloon. Electron transport spontaneously increases.
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Mitochondria function: the chemiosmotic gradient Endotherms like ourselves (formerly called homeotherms) maintain a constant body temperature. The setpoint for that temperature is determined by a region the hypothalamus (part of the brain, for you newcomers). But just how is the temperature regulated? That is, how does the body maintain a balance between heat production and heat loss?
The answer to that one is very complex. There are many mechanisms for heat dissipation and retention (sweating, changes in distribution of the circulation, shivering, piloerection - no, it's not a dirty word, etc.). Heat production results from the loss of some of the free energy of every chemical reaction as heat - that is, every chemical reaction results in the loss of some of the total free energy in the universe. That's one of the laws of thermodynamics - I forgot which. A major source of heat production is electron transport in mitochondria. How do mitochondria produce heat? Much of the energy released by electron transport does no useful work at all (that law of thermodynamics is at work again). That is, each exchange of electrons results in a loss of some energy as heat. Furthermore, dissipation of the chemiosmotic gradient without doing work results in the transformation of a great deal more energy to heat. The mechanism of respiratory control can be exploited to increase heat production by dissipating the gradient at a faster rate, or to slow heat production by making the ETS more efficient. One example of that type of regulation is the hormone thyroxine (thyroid hormone). Thyroid hormone has many functions, mostly associated with promotion of anabolic activity synthesis of compounds and cellular growth. However thyroxine is also a mildly effective uncoupling agent. Uncoupling agents dissipate the chemical gradient, usually by creating a mechanism by which protons can escape the intermembrane space (or otherwise 'short out' the gradient), allowing an increase in electron transport. In colder regions of the world a noticible change in thyroid hormone levels takes place circannually, that is, levels are higher in winter and lower in summer.
Copyright and I nte nde d Use Visitors: to e nsure that your me ssage is not mistake n for S PAM, ple ase include the acronym "Bios211" in the subje ct line of e -mail communications Cre ate d by David R. Capre tte (capre tte @rice .e du), Rice Unive rsity 12 De c 96 Update d 31 May 05
www.ruf.rice.edu/~bioslabs/studies/mitochondria/mitogradient.html
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