Photosynthesis

Steps of Photosynthesis

Photosynthesis uses carbon dioxide, water, and light energy to make sugar.
During photosynthesis, many reactions take place, but the overall process can be summarized by a chemical equation which states that the combination of carbon dioxide, water, and light energy produces a carbohydrate (glucose) and oxygen.
6CO2+12H2O+lightenergyC6H12O6+6O2+6H2O6\;{\mathrm{CO}}_2\;+\;12\;{\mathrm H}_2\mathrm O\;+\;\mathrm{light}\;\mathrm{energy}\;\rightarrow\;{\mathrm C}_6{\mathrm H}_{12}{\mathrm O}_6\;+\;6\;{\mathrm O}_2\;+\;6\;{\mathrm H}_2\mathrm O
The six-carbon sugar produced, C6H12O6, is glucose. The direct product of photosynthesis is actually a three-carbon molecule used to make sugar rather than glucose itself. Because the process that builds glucose from the three-carbon molecule immediately follows photosynthesis, the entire reaction is most easily summarized by the equation. However, the equation can be further simplified by showing only the net consumption of water:
6CO2+6H2O+lightenergyC6H12O6+6O26\;{\mathrm{CO}}_2\;+\;6\;{\mathrm H}_2\mathrm O\;+\;\mathrm{light}\;\mathrm{energy}\;\rightarrow\;{\mathrm C}_6{\mathrm H}_{12}{\mathrm O}_6\;+\;6\;{\mathrm O}_2
The sugar produced in this reaction is directly built not during photosynthesis but in the respiration process that follows photosynthesis. Photosynthesis consists of two stages: the light-dependent reactions and the light-independent reactions, which include the Calvin cycle (or Calvin-Benson cycle), the cycle that fixes carbon into glyceraldehyde 3-phosphate (G3P), the precursor to glucose.

Stages of Photosynthesis

In the first stage of photosynthesis, reactions in the thylakoid membrane convert light energy to chemical energy by splitting water molecules. In the second stage, chemical reactions in the stroma (the connective and supportive framework of a chloroplast) build sugar molecules from atmospheric carbon and use the chemical energy supplied by the first stage.

First Stage of Photosynthesis: The Light Reactions

The light reactions make up the first stage of photosynthesis. They use light energy to convert ADP into ATP, and to reduce NADP+ to NADPH.
Chlorophyll is able to harness light energy because of the way it interacts with light. Sunlight consists of light in a range of wavelengths, and chlorophyll absorbs almost all of the visible light. Thus, high-energy, short-wavelength light (toward the blue end of the spectrum) is absorbed and excites electrons inside the thylakoid membrane. The excitation of the electrons initiates the transformation of light energy into chemical energy. In order to ensure this process makes energy available as chemical energy, chloroplasts rely on structures known as photosystems. There are two photosystems, named in order of their discovery, photosystem I and photosystem II. Photosystem II is the first light-capturing complex, and photosystem I is the second light-capturing complex; both are found in the thylakoid membrane of a chloroplast that converts light energy into chemical energy. Photosystem I transfers the energy to the molecules of ATP and NADPH.

The light reactions are the first stage of photosynthesis. In these reactions, energy from light is converted into chemical energy. These light reactions take place in the thylakoid membrane. Water (a reactant in the photosynthesis equation) is split, providing electrons and hydrogen ions (H+). These electrons and hydrogen ions are accepted by nicotinamide adenine dinucleotide phosphate, NADP+, the oxidized form of NADPH, that serves as an electron carrier in the Calvin cycle. The H at the end of NADPH denotes that the molecule contains an extra hydrogen atom as compared to NADP+. A molecule of NADP+ contains an extra phosphate group compared to the electron carrier NAD+, which moves electrons through the process of cellular respiration. A separate reaction uses the energy from light to add an inorganic phosphate to adenosine diphosphate (ADP), which is the reduced form of the biological unit of energy, adenosine triphosphate, ATP, which consists of an adenosine (an adenine group and a ribose sugar) and three phosphate groups. ADP has one less phosphate group than ATP. Thus, at the end of the first stage of photosynthesis, chemical energy is available in the form of ATP and NADPH.

Photosystem II

A photosystem is a complex of proteins and pigments working together to absorb energy from light and transfer it to an electron acceptor, which is a molecule that accepts an electron and transfers it to another molecule. Each photosystem consists of a light-harvesting complex, which is a group of molecules that takes in light, surrounding a reaction-center complex, where the light is converted to chemical energy. Many pigments (including several types of chlorophyll, as well as others such as carotenoids) are distributed throughout the light-harvesting complex. They absorb energy from light as it enters the complex and pass the energy along to the reaction-center complex. When light strikes the chlorophyll, it excites an electron to a higher energy state. That electron then drops back to its initial state, which releases energy, exciting an electron in the next chlorophyll. Thus, the series of pigments creates a pathway to the reaction-center complex.

In the reaction-center complex of photosystem II, a pair of chlorophyll molecules known as P680 (so named because the molecules best absorb light at a wavelength of 680 nanometers) donate electrons to the primary electron acceptor. This leaves the P680 molecules positively charged (denoted P680+). An enzyme catalyzes the splitting of water into two electrons, two hydrogen ions (H+), and an oxygen atom. The electrons from this reaction are transferred to P680, returning P680 to its initial state. The oxygen atom combines with another oxygen atom to make O2. The hydrogen ions are released into the thylakoid lumen, where they create a proton gradient that will be used to form ATP. This process is chemiosmosis, the movement of ions across a semipermeable membrane down their electrochemical gradient, which results in a charge that can be used as a source of energy. The electrons from the primary electron acceptor are passed in a series of redox reactions from photosystem II to photosystem I via an electron carrier called plastoquinone (Pq), a cytochrome complex (an enzyme in the thylakoid membrane that forms part of the electron transport chain) and a protein called plastocyanin (Pc).

Photosystem I

In photosystem I, a process similar to the process in photosystem II captures light energy and transfers it to a primary electron acceptor via P700 (which best absorbs light at 700 nm). This creates a positive charge, making the resulting ion (P700+) able to accept the electrons from photosystem II. The excited electrons from the primary electron acceptor of photosystem I are then passed in a series of redox reactions through the protein ferrodoxin (Fd). Redox reactions are reactions in which substances gain or lose electrons. The enzyme NADP reductase catalyzes the transfer of these electrons to NADP+. It also transfers a hydrogen ion (H+) from the stroma to NADP+. This makes the molecule NADPH, which is used as an electron carrier in the Calvin cycle.
Energy from light is captured in photosystem II and transferred to the primary electron acceptor (a molecule that receives or accepts electrons). Water is split into electrons, H+, and an oxygen atom. Electrons are moved out of photosystem II to photosystem I via redox reactions (transfer of electrons) along Pq (plastoquinone), the cytochrome complex, and Pc (plastocyanin). Energy is first captured by photosystem II, which absorbs light at a wavelength of 680 nm, and then during photosystem I, when light is absorbed at a wavelength of 700 nanometers. The proton gradient created by H+ supplies the energy needed to make ATP through chemiosmosis. In photosystem I, energy is captured and transferred to the primary electron acceptor. It is transported by Fd (ferrodoxin) to NADP reductase, which reduces NADP+ to NADPH.

Chemiosmosis

Chemiosmosis is the movement of charged particles across a semipermeable membrane, down their electrochemical gradient. This means they are moved from an area with more particles of that charge to an area with fewer particles of that charge. After water is split in photosystem II, H+ are released out of the photosystem into the thylakoid lumen. This creates a proton gradient that makes the interior of the thylakoid positively charged. This chemical potential is used by the enzyme ATP synthase to generate ATP from ADP. ATP synthase resides in the thylakoid membrane. H+ release energy in the form of hydrogen bonds as ATP synthase moves them from inside the thylakoid to the stroma. This provides the energy needed to add an inorganic phosphate to ADP, forming ATP (inorganic phosphates and ADP reside within the stroma). H+ are not consumed in this reaction and are released into the stroma. This makes H+ available to NADP reductase, which uses H+ to reduce NADP+ to NADPH, following the reactions in photosystem I. Thus, at the end of the first stage of photosynthesis, ATP and NADPH have been produced.
ATP synthase, an enzyme that synthesizes ATP (adenosine triphosphate), uses the proton gradient supplied by photosystem II and the splitting of water to generate ATP from ADP in chemiosmosis. Chemiosmosis is the movement of charged particles across a semipermeable membrane, down their electrochemical gradient. This makes ATP available as an energy source for other reactions.

The Second Stage of Photosynthesis: The Calvin Cycle

The second stage of photosynthesis is the Calvin cycle, also known as the light-independent reactions.
The second stage of photosynthesis is the Calvin cycle, also called the Calvin-Benson cycle, is named for its discoverers, Melvin Calvin and Andrew Benson. The Calvin cycle is an anabolic process (a chemical reaction that synthesizes molecules in metabolism) that builds the molecules that make up glucose, the six-carbon sugar that is the product of reactions following the Calvin cycle. The cycle uses energy to build a large molecule from smaller ones. The cycle takes place in the stroma (the fluid inside chloroplasts) in three main steps.

The first step is known as carbon fixation because it takes in CO2 from the atmosphere and "fixes" it into organic molecules that can be used by living things. In this step, the enzyme ribulose bisphosphate carboxylase-oxygenase (abbreviated rubisco) adds one carbon to a five-carbon sugar called ribulose bisphosphate (RuBP) during carbon fixation. This forms a six-carbon sugar that is energetically unstable, meaning it cannot hold its form. It immediately splits into two 3-carbon molecules called 3-phosphoglycerate (3-PGA).

In the second step, each molecule of 3-phosphoglycerate receives a phosphate group from ATP, forming 1,3-bisphosphoglycerate. Because this process changes ATP to ADP, it is endergonic (uses energy). Then NADPH donates a pair of electrons to 1,3-bisphosphoglycerate, which reduces the molecule. This causes it to lose a phosphate group. The product of this reaction is glyceraldehyde 3-phosphate (G3P), a three-carbon sugar formed in the Calvin cycle that is a precursor to glucose. Importantly, for every three molecules of CO2 that enter the cycle, six molecules of G3P are formed. However, five of these G3P molecules will be recycled into RuBP in the next step. Only one molecule leaves the cycle, resulting in a net production of one G3P for every three CO2 consumed.

The final step is the regeneration of RuBP from the five molecules of G3P in the previous step. This final step involves a complex series of reactions that rearrange the carbon atoms into the five-carbon sugar, consuming an additional three ATP in the process. At the end of this step, RuBP is again ready to receive CO2, and the cycle begins again. Because only one G3P is produced from a single turn of the cycle, it takes six turns of the cycle to produce a single molecule of glucose.
In the first step of the Calvin cycle, CO2 is fixed with RuBP to form 3-phosphoglycerate. In the second step, 3-phosphoglycerate is reduced and becomes G3P. One molecule leaves the cycle, while the others go on to the third step. In the third step, the remaining G3P regenerate RuBP, and the cycle begins again.