Light to chemical energy

The light energy is converted to chemical energy using the light-dependent reactions. The products of the light dependent reactions are ATP from photophosphorylation and NADPH from photoreduction. Both are then utilized as an energy source for the light-independent reactions.

In plants, the light-dependent reactions occur in the thylakoid membranes of the chloroplasts and use light energy to synthesize ATP and NADPH. The photons are captured in the antenna complexes of photosystem I and II by chlorophyll and accessory pigments (see diagram at right). When a chorophyll a molecule at a photosystem's reaction center absorbs energy, an electron is excited and transferred to an electron-acceptor molecule through a process called Photoinduced charge separation. These electrons are shuttled through an electron transport chain that initially functions to generate a chemiosmotic potential across the membrane, the so called Z-scheme shown in the diagram. An ATP synthase enzyme uses the chemiosmotic potential to make ATP during photophosphorylation while NADPH is a product of the terminal redox reaction in the Z-scheme.

A photosystem: a light-harvesting cluster of photosynthetic pigments in a chloroplast thylakoid membrane.

 

Water photolysis

The NADPH is the main reducing agent in chloroplasts, providing a source of energetic electrons to other reactions. Its production leaves chlorophyll with a deficit of electrons (oxidized), which must be obtained from some other reducing agent. The excited electrons lost from chlorophyll in photosystem I are replaced from the electron transport chain by plastocyanin. However, since photosystem II includes the first steps of the Z-scheme, an external source of electrons is required to reduce its oxidized chlorophyll a molecules. This role is played by water during a reaction known as photolysis and results in water being split to give electrons, oxygen and hydrogen ions. Photosystem II is the only known biological enzyme that carries out this oxidation of water. Initially, the hydrogen ions from photolysis contribute to the chemiosmotic potential but eventually they combine with the hydrogen carrier molecule NADP⁺ to form NADPH. Oxygen is a waste product of light-independent reactions, but the majority of organisms on Earth use oxygen for cellular respiration, including photosynthetic organisms.

Oxygen and photosynthesis

With respect to oxygen and photosynthesis, there are two important concepts.

• Plant and algal cells also use oxygen for cellular respiration, although they have a net output of oxygen since much more is produced during photosynthesis.

• Oxygen is a product of the photolysis reaction not the fixation of carbon dioxide, during the light-independent reactions. Consequently, the source of oxygen during photosynthesis is water, not carbon dioxide.

Bacterial variations

The concept that oxygen production is not directly associated with the fixation of carbon dioxide was first proposed by Cornelis Bernadus van Neil in the 1930s, who studied photosynthetic bacteria. Aside from the cyanobacteria, bacteria only have one photosystem and use reducing agents other than water. They get electrons from a variety of different inorganic chemicals including sulfide or hydrogen, so for most of these bacteria oxygen is not produced.

The 'Z-scheme' of electron flow in light-dependent reactions.

Others, such as the halophiles (an Archeae) produced so called purple membranes where the bacteriorhodopsin could harvest light and produce energy. The purple membranes was one of the first to be used to demonstrate the chemiosmotic theory: light hit the membranes and the pH of the solution that contained the purple membranes dropped as protons were pumping out of the membrane.

Carbon fixation

The fixation of carbon dioxide is a light-independent process in which carbon dioxide combines with a five-carbon sugar, ribulose bisphosphate (RuBP), to give two molecules of a three-carbon compound, glycerate 3-phosphate (GP). This compound is also sometimes known as 3-phosphoglycerate (PGA). GP, in the presence of ATP and NADPH from the light-dependent stages, is reduced to glyceraldehyde 3-phosphate (G3P). This product is also referred to as 3-phosphoglyceraldehyde (PGAL) or even as triose phosphate (a three-carbon sugar). This is the point at which carbohydrates are produced during photosynthesis. Some of the triose phosphates condense to form hexose phosphates, sucrose, starch and cellulose or are converted to acetylcoenzyme A to make amino acids and lipids. Others go on to regenerate RuBP so the process can continue (see Calvin cycle).

Cite: Wikipedia