Biochemistry Online: An Approach Based on Chemical Logic

Biochemistry Online

CHAPTER 8 - OXIDATION/PHOSPHORYLATION

D:  THE LIGHT REACTION OF PHOTOSYNTHESIS

BIOCHEMISTRY - DR. JAKUBOWSKI

04/15/16

Learning Goals/Objectives for Chapter 8D: 

After class and this reading, students will be able to

  • described the mechanistic similarities between mitochondrial oxidative/phosophorylation in which NADH and FADH2 are regenerated on reduction of O2 and the light reaction of photosynthesis in which O2 and a reducing agent, NADPH are produced;
  • describe similarities in fluorescence resonance energy transfer and exciton transfer;
  • describe the difference in properties between chlorophylls acting as antennae and chlorophylls at the reaction center;
  • describe how sunlight driven excitation of chlorophyll molecules at the reaction center produces an oxidzing agent strong enough to oxide water and form O2, itself a powerful oxidizing agent;
  • explain the general flow of electrons from dioxgen to NADP+ through a series of mobile and membrane protein bound electron carriers in the Z scheme of electron transport in the chloroplast thylacoid membranes;
  • explain with picture diagrams how oxidation of H2O and phosphorylation reactions (to produce ATP) are coupled in in the Z scheme;

D3.  The Light Reactions of Photosynthesis

Photoexcitation of the non-reaction center chlorophyll turns that molecule into a good reducing agent, which transfers its electron to the nearest excited state level of the reaction center chlorophyll. If you count both step together, the non-reaction center chlorophyll gets "photooxidized", in the process producing the "strong" oxidizing agent which is the positively charged chlorophyll derivative. The extra electron passed onto the second molecule will eventually be passed on to NADP+ to produce NADPH.   The light reaction of photosynthesis in green plants is shown below.  In this process, in a scheme that is reminiscent of electron transport in mitochondria, water is oxidized by photosystem II. Electrons from water are moved through PSII to a mobile, hydrophobic molecule, plastaquinone (PQ) to form its reduced form, PQH2.   PSII is a complicated structure with many polypeptide chains, lots of chlorophylls, and Mn, Ca, and Fe ions.  A Mn cluster, called the oxygen evolving complex, OEC, is directly involved in the oxidation of wate.  Two key homologous 32 KD protein subunits, D1 and D2, in PSII are transmembrane proteins and are at the heart of the PSII complex.  Another photosystem, PS1, is also found further "downstream" in the electron transport pathway.  It takes electrons from another reduced mobile carrier of electrons, plastocyanin (PCred) to ferredoxin, which becomes a strong reducing agent. Ferrodoxin is a protein with an Fe-S cluster (Fe-S-Fe-S in a 4 membered ring, with 2 additions Cys residues coordinating each Fe).  It  ultimately passes its electrons along to NADP+ to form NADPH. A summary of the light reaction in plants and standard reduction potentials of the participants, are shown below.  Note that many of the complexes produce a transmembrane proton gradient.  In contrast to mitochondria, the lumen (as compared to the mitochondrial matrix) becomes more acidic that the other stroma.  Protons then can move down a concentration gradient through the C0C1ATPase to produce ATP required for reductive biosythesis of glucose.

Figure:   THE LIGHT REACTIONS OF PHOTOSYNTHESIS

 Figure:  Detailed View of Light Reaction of Photosynthesis (reprinted with permission from Kanehisa Laboratories and the KEGG project: www.kegg.org )

Boxed number represent Enzyme Commission Number.  Original KEGG Map with imbedded links.

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