Historically, the role of light in photosvnthesis has been ascribed either to a photolysis of carbon dioxide or to a photolysis of water and a resultant rearrangement of constituent atoms into molecules of oxygen and glucose (or formaldehyde). The discovery of photophosphorylation demonstrated that photosynthesis includes a light-induced phosphorus metabolism that precedes, and is independent from, a photolysis of water or CO2. ATP formation could best be accounted for not by a photolytic disruption of the covalent bonds in C02 or water but by the operation of a light-induced electron flow that results in a release of free energy which is trapped in the pyrophosphate bonds of ATP.Photophosphorylation is now divided into (a) a noncyclic type, in which the formation of ATP is coupled with a light-induced electron transport from water to ferredoxin and a concomitant evolution of oxygen and (b) a cyclic type which yields only ATP and produces no net change in the oxidation-reduction state of any electron donor or acceptor. Reduced ferredoxin formed in (a) serves as an electron donor for the reduction of NADP by an enzymic reaction that is independent of light. ATP, from both cyclic and noncyclic photophosphorylation, and reduced NADP jointly constitute the assimilatory power for the conversion of C02 to carbohydrates (3 moles of ATP and 2 moles of reduced NADP are required per mole of C02).Investigations, mainly with whole cells, have shown that photosynthesis in green plants involves two photosystems, one (System II) that best uses light of "short" wavelength (X < 685 nm) and another (System I) that best uses light of "long" wavelength (X > 685 nm). Cyclic photophosphorylation in chloroplasts involves a System I photoreaction. Noncyclic photophosphorylation is widely held to involve a collaboration of two photoreactions: a short-wavelength photoreaction belonging to System II and a long-wavelength photoreaction belonging to System I. Recent findings, however, indicate that noncyclic photophosphorylation may include two short-wavelength, System II, photoreactions that operate in series and are joined by a "dark" electron-transport chain to which is coupled a phosphorylation site.
Early conceptsThe first hypothesis about the role of light in photosynthesis came very appropriately from Jan Ingenhousz, who some years earlier had made the epochal discovery that it is "the influence of the light of the sun upon the plant" (1) that is responsible for the "restorative" effect of vegetation on "bad" air-an observation first made in 1771 by Joseph Priestley without reference to light (2). In 1796, Ingenhousz wrote that the green plant absorbs from "carbonic acid in the sunshine, the carbon, throwing out at that time the oxygen alone, and keeping the carbon to itself as nourishment" (3).The idea that light liberates oxygen by photodecomposing C02 had, with some modifications, persisted for well over a century. It seemed to have had a special attraction for some of the most illustrious chemists in their day, e.g., von ...