MadSci Network: Biochemistry |
Philipp: That is a good question. Photosynthesis is an extremely complicated series of biochemical reactions utilizing light that we do not completely yet understand. Only recently, were scientists able to visualize (Through x-ray crystallography) part of the ligh-harvesting complex of photosynthesis. First of all, I will break your question up into two parts... the pH/charge gradient created by the proton pumping... In this case, the pH change is created simply because you have more protons on one side of a membrane than the other... And the potential generated here, is utilized by the ATPase to synthesize ATP from AMP/ADP and ppi (inorganic phosphate). This reaction is comparable to that performed in respiration, as observed in the mitochondria, so I will not delve further into it now. Essentially electrons are converted into chemical energy through a multitude of reactions, but how are these electrons transduced? In simplest terms, there are a series of chemicals in plants that are labile for accepting electrons and are able to 'release' them as well with different energetic properties. These molecules, absorb a photon (Light) which in turn excites and removes an electron from an orbital. This excitation is passed through the photosystem and in photosynthetic bacteria (The best understood photosynthetic system) the energy of the excited electron is traded off with the exchange of a proton across the membrane... A special protein called cytochrome b1 facilitates the pumping, and another component recycles the electrons, cytochrome b2. So, essentially there is no net loss in electrons, as you move an electron one way (-1 negative charge) it is reduced in it's excited energy state by the oxidation and pushing of 4 protons across the membrane... The actual chemistry of the electron is mediated by iron in the cytochrome, which as photosynthesis progresses continually is reduced and oxideized from Fe3+ to Fe2+ and back again as cytochrome b2 takes the electron (Fe2+ >> Fe3_) back to the PSI/II. The source of the hydrogen is from the breaking of water molecules, and the energy of the electron is used to reduce NADP+ to NADPH, which, comparable to NADH+ H+ in respirative organisms, is what is actually used to generate the ATP. If you have had chemistry, the half reactions for this are: O2 + 4e- + 4H+ <==> 2H2O Eo'= +.815V NADP+ + H+ +2e- <==> NADPH Eo'= -.320V 2NADP+ + 2H2O <===> 2NADPH + O2 + 2H+ Eo' = -1.135V As for the transfer of electrons between photosystems, again, an iron containing molecule plays an important part, cytochrome f. This cytochrome allows electron transport from PSII to PSI. There are several herbicides that block this reaction, killing the plant, such as DCMU. The sum of photosynthesis is mediated by three main membrane bound proteins with these mobile electron carrying proteins around them, they are: PSII complex,--->cytochrome F complex--->PSI, and then the NADPH is generated through oxidation/reduction of FAD, and ferredoxin, and as you mentioned, the ATP is generated by the F1 Fo ATPase complex, which through all of that pH potential you mentioned, lets protons back through the membrane and makes ATP from ADP + Pi... There are dozens of other reactions, (oxidations/reductions) occuring here, but those are primarily the major players. Remember, the take home message is that everything is ultimately recycled, there are limited quantities of most of the chemicals involved, (Particularly NADP+) and the electrons need to be 'taken back' to the beginning to regenerate the system. Any good biochemistry text, such as Voet&Voet, Suzuki, or Strier, has lots of really good color drawings and explanations of the multitude of electron hand-off's that are occuring... Good luck and have fun with your photons and protons... -Matt-
Try the links in the MadSci Library for more information on Biochemistry.