Photosynthetic electron transport

Light and photosynthetic electron transport convert DPEs into free radicals of undetermined stmcture. The radicals produced in the presence of the bipyridinium and DPE herbicides decrease leaf chlorophyll and carotenoid content and initiate general destmction of chloroplasts with concomitant formation of short-chain hydrocarbons from polyunsaturated fatty acids (37,97). 

A. Trebst and M. Avron, eds.. Photosynthesis P. Photosynthetic Electron Transport andPhotophosphorylation, Tnyclopedia of Plant Physiolog i, NS., Springer-Vedag, Berlin, 1977. 

The quantum yield of photosynthesis, the amount of product formed per equivalent of light input, has traditionally been expressed as the ratio of COg fixed or Og evolved per quantum absorbed. At each reaction center, one photon or quantum yields one electron. Interestingly, an overall stoichiometry of one translocated into the thylakoid vesicle for each photon has also been observed. Two photons per center would allow a pair of electrons to flow from HgO to NADP (Figure 22.12), resulting in the formation of 1 NADPH and Og. If one ATP were formed for every 3 H translocated during photosynthetic electron transport, 1 ATP would be synthesized. More appropriately, 4 hv per center (8 quanta total) would drive the evolution of 1 Og, the reduction of 2 NADP, and the phosphorylation of 2 ATP. 

The thylakoid membrane is asymmetrically organized, or sided, like the mitochondrial membrane. It also shares the property of being a barrier to the passive diffusion of H ions. Photosynthetic electron transport thus establishes an electrochemical gradient, or proton-motive force, across the thylakoid membrane with the interior, or lumen, side accumulating H ions relative to the stroma of the chloroplast. Like oxidative phosphorylation, the mechanism of photophosphorylation is chemiosmotic. 

Photosynthetic electron transport, which pumps into the thylakoid lumen, can occur in two modes, both of which lead to the establishment of a transmembrane proton-motive force. Thus, both modes are coupled to ATP synthesis and are considered alternative mechanisms of photophosphorylation even though they are distinguished by differences in their electron transfer pathways. The two modes are cyclic and noncyclic photophosphorylation. 

FIGURE 22.21 The mechanism of photophosphorylation. Photosynthetic electron transport establishes a proton gradient that is tapped by the CFiCFo ATP synthase to drive ATP synthesis. Critical to this mechanism is the fact that the membrane-bound components of light-induced electron transport and ATP synthesis are asymmetrical with respect to the thylakoid membrane so that vectorial discharge and uptake of ensue, generating the proton-motive force. 

Assuming that the concentrations of ATP, ADP, and P in chloroplasts are 3 mM, 0.1 mM, and 10 mM, respectively, what is the AG for ATP synthesis under these conditions Photosynthetic electron transport establishes the proton-motive force driving photophosphorylation. What redox potential difference is necessary to achieve ATP synthesis under the foregoing conditions, assuming an electron pair is transferred per molecule of ATP generated ... 

If noncyclic photosynthetic electron transport leads to the translocation of 3 H /e and cyclic photosynthetic electron transport leads to the translocation of 2 H /A, what is the relative photosynthetic efficiency of ATP synthesis (expressed as the number of photons absorbed per ATP synthesized) for noncyclic versus cyclic photophosphorylation (Assume that the CFiCEq ATP synthase yields 1 ATP/3 H. )... 

In contrast to common usage, the distinction between photosynthetic and respiratory Rieske proteins does not seem to make sense. The mitochondrial Rieske protein is closely related to that of photosynthetic purple bacteria, which represent the endosymbiotic ancestors of mitochondria (for a review, see also (99)). Moreover, during its evolution Rieske s protein appears to have existed prior to photosynthesis (100, 101), and the photosynthetic chain was probably built around a preexisting cytochrome be complex (99). The evolution of Rieske proteins from photosynthetic electron transport chains is therefore intricately intertwined with that of respiration, and a discussion of the photosynthetic representatives necessarily has to include excursions into nonphotosynthetic systems. 

Sharkey, T.D. Badger, M.R. (1982). Effects of water stress on photosynthetic electron transport, photophosphorylation and metabolite levels of Xanthium strumarium mesophyll cells. Planta, 156, 199-206. 

While phototactic action spectra measured in some Phormidium species indicate that chlorophyll a is not involved in the absorption of phototactically active light (see below), the phototactic action spectrum of Anabaena variabilis106) shows slight activity around 440 nm and a distinct peak at around 670 nm, both indicating chlorophyll a. Since blockers of the photosynthetic electron transport, such as DCMU andDBMIB, (see below) do not affect phototactic orientation, the active light seems not to be utilized via the photosynthetic electron transport chain (for further information see below). 

These experiments show that, as in the case of chlorophyll a and the carotenoids, the energy absorbed by the phycobiliproteins is utilized via the photosynthetic apparatus furthermore, they provide evidence that photophobic responses in blue-green algae are caused by sudden changes in the steady state of the photosynthetic electron transport, especially the non-cyclic one. 

Phototactic action spectra of Phormidium autumnale and Phormidium uncinatum, measured by Nultsch86>89), show prominent maxima in the absorption range of C-phycoerythrin and smaller, but distinct, peaks in the absorption range of C-phyco-cyanin. Red light absorbed by chlorophyll a is not active, while in the blue range absorbedby the Soret band, the action spectrum shows aminimum(Fig. 6). Nultsch87) concluded that biliproteins are photoreceptors of phototaxis, but independently of the photosynthetic electron transport and phosphorylation. 

These results were interpreted using the electron pool hypothesis There is an electron pool situated in the linear photosynthetic electron transport chain between photosystems II and I (Fig. 9). A phobic response is triggered by a decrease in the flow rate through the pool. This can be accomplished in two ways ... 

Fig. 10. Redox systems of the photosynthetic electron transport chain incorporated in the thylakoid membrane. Irradiation causes the generation of a proton gradient (after Trebst and Hauska135))...

L. Florin, A. Tsokoglou, T. Happe (2001) A novel type of iron hydrogenase in the green alga Scenedesmus obliquus is linked to the photosynthetic electron transport chain. J. Biol. Chem., 276 6125-6132... 

The key enzyme hydrogenase catalyses the reversible reduction of protons to molecular hydrogen. Inhibitor experiments indicate that the ferredoxin PetF functions as natural electron donor linking the hydrogenase to the photosynthetic electron transport chain [Florin et al., 2001],... 

The water acts as a large reserve of electrons for the photosynthetic electron transport process in the Z scheme, with the oxygen being produced as a waste product. 

The first cytochrome to be recognised as a component of the photosynthetic electron transport chain was cytochrome f [142]. The properties of cytochrome f have been reviewed [143,144], and amino-acid sequence information is available for pea, spinach, wheat and tobacco [145]. The axial ligand to the heme-Fe... 

Cu Laccase, oxidases Plastocyanin photosynthetic electron transport Cytochrome c oxidase mitochondrial electron transport... 

Fe Cytochrome oxidase reduction of oxygen to water Cytochrome P-450 0-insertion from O2, and detoxification Cytochromes b and c electron transport in respiration and photosynthesis Cytochrome f photosynthetic electron transport Ferredoxin electron transport in photosynthesis and nitrogen fixation Iron-sulfur proteins electron transport in respiration and photosynthesis Nitrate and nitrite reductases reduction to ammonium... 

Ferredoxins of the 2Fe-2S type play a role in the photosynthetic electron transport as an essential electron acceptor of photosystem I. The solution... 

Because photosystem 11 and the cytochrome b/f complex release protons from reduced plastoquinone into the lumen (via a Q. cycle), photosynthetic electron transport establishes an electrochemical gradient across the thylakoid membrane (see p. 126), which is used for ATP synthesis by an ATP synthase. ATP and NADPH+H", which are both needed for the dark reactions, are formed in the stroma. 

The photosynthetic electron transport chain in plants starts in photosystem II (PS 11 see p. 128). PS 11 consists of numerous protein subunits (brown) that contain bound pigments—i.e., dye molecules that are involved in the absorption and transfer of light energy. 

It can be seen from the normal potentials E° (see p. 18) of the most important redox systems involved in the light reactions why two excitation processes are needed in order to transfer electrons from H2O to NADP"". After excitation in PS II, E° rises from around -IV back to positive values in plastocyanin (PC)—i. e., the energy of the electrons has to be increased again in PS I. If there is no NADP" available, photosynthetic electron transport can still be used for ATP synthesis. During cyclic photophosphorylation, electrons return from ferredoxin (Fd) via the plastoquinone pool to the b/f complex. This type of electron transport does not produce any NADPH, but does lead to the formation of an gradient and thus to ATP synthesis. 

Howell JM, Vieth WR (1982) Biophotolytic membranes simplified kinetic model of photosynthetic electron transport, JMol Catal 16 245-298... 

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