2007-12-12T08:58:00.001-08:00

Photosynthesis

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Photosynthesis splits water to liberate O₂ and fixes CO₂ into sugar

The leaf is the primary site of photosynthesis in plants.

Photosynthesis is the conversion of light energy into chemical energy by living organisms. The raw materials are carbon dioxide and water, the energy source is sunlight, and the end-products include glucose and oxygen. It is arguably the most important biochemical pathway,^[1] since nearly all life depends on it. It is a complex process occurring in higher plants, phytoplankton, algae, as well as bacteria such as cyanobacteria. Photosynthetic organisms are also referred to as photoautotrophs.^[1]

The word comes from the Greek photo-, light, and synthesis, putting together.

Overview

Photosynthesis uses light energy and carbon dioxide to make triose phospates (G3P). G3P is generally considered the prime end-product of photosynthesis. It can be used as an immediate food nutrient, or combined and rearranged to form monosaccharide sugars, such as glucose, which can be transported to other cells, or packaged for storage as insoluble polysaccharides such as starch.

A general equation for photosynthesis is:

6 CO_2(gas) + 12 H₂O_(liquid) + photons → C₆H₁₂O_6(aqueous) + 6 O_2(gas) + 6 H₂O_(liquid)

carbon dioxide + water + light energy → glucose + oxygen + water

When written as a word equation the light energy appears above the arrow as it is required for photosynthesis but it is not actually a reactant.

The equation is often presented in introductory chemistry texts in simplified form as:^[2]

6 CO_2(gas) + 6 H₂O_(liquid) + photons → C₆H₁₂O_6(aqueous) + 6 O_2(gas)

Photosynthesis occurs in two stages. In the first phase light-dependent reactions or photosynthetic reactions (also called the Light reactions) capture the energy of light and use it to make high-energy molecules. During the second phase, the light-independent reactions (also called the Calvin-Benson Cycle, and formerly known as the Dark Reactions) use the high-energy molecules to capture carbon dioxide (CO₂) and make the precursors of carbohydrates.

In the light reactions one molecule of the pigment chlorophyll absorbs one photon and loses one electron. This electron is passed to a modified form of chlorophyll called pheophytin, which passes the electron to a quinone molecule, allowing the start of a flow of electrons down an electron transport chain that leads to the ultimate reduction of NADP into NADPH. In addition, it serves to create a proton gradient across the chloroplast membrane; its dissipation is used by ATP Synthase for the concomitant synthesis of ATP. The chlorophyll molecule regains the lost electron by taking one from a water molecule through a process called photolysis, that releases oxygen gas as a waste product.

In the Light-independent or dark reactions the enzyme RuBisCO captures CO₂ from the atmosphere and in a process that requires the newly formed NADPH, called the Calvin-Benson cycle releases three-carbon sugars which are later combined to form sucrose and starch.

Photosynthesis may simply be defined as the conversion of light energy into chemical energy by living organisms. It is affected by its surroundings and the rate of photosynthesis is affected by the concentration of carbon dioxide, the intensity of light, and the temperature.

In plants

Most plants are photoautotrophs, which means that they are able to synthesize food directly from inorganic compounds using light energy - for example from the sun, instead of eating other organisms or relying on nutrients derived from them. This is distinct from chemoautotrophs that do not depend on light energy, but use energy from inorganic compounds.

6 CO₂ + 12 H₂O → C₆H₁₂O₆ + 6 O₂ + 6 H₂O

The energy for photosynthesis ultimately comes from absorbed photons and involves a reducing agent, which is water in the case of plants, releasing oxygen as a waste product. The light energy is converted to chemical energy (known as light-dependent reactions), in the form of ATP and NADPH, which are used for synthetic reactions in photoautotrophs. The overall equation for the light-dependent reactions under the conditions of non-cyclic electron flow in green plants is:

2 H₂O + 2 NADP⁺ + 2 ADP + 2 P_i + light → 2 NADPH + 2 H⁺ + 2 ATP + O₂ ^[3]

Most notably, plants use the chemical energy to fix carbon dioxide into carbohydrates and other organic compounds through light-independent reactions. The overall equation for carbon fixation (sometimes referred to as carbon reduction) in green plants is:

3 CO₂ + 9 ATP + 6 NADPH + 6 H⁺ → C₃H₆O₃-phosphate + 9 ADP + 8 P_i + 6 NADP⁺ + 3 H₂O ^[3]

More specifically, carbon fixation produces an intermediate product, which is then converted to the final carbohydrate products. The carbon skeletons produced by photosynthesis are then variously used to form other organic compounds, such as the building material cellulose, as precursors for lipid and amino acid biosynthesis or as a fuel in cellular respiration. The latter not only occurs in plants, but also in animals when the energy from plants get passed through a food chain. Organisms dependent on photosynthetic and chemosynthetic organisms are called heterotrophs. In general outline, cellular respiration is the opposite of photosynthesis: glucose and other compounds are oxidised to produce carbon dioxide, water, and chemical energy. However, both processes take place through a different sequence of chemical reactions and in different cellular compartments.

Plants absorb light primarily using the pigment chlorophyll, which is the reason that most plants have a green color. The function of chlorophyll is often supported by other accessory pigments such as carotenes and xanthophylls. Both chlorophyll and accessory pigments are contained in organelles (compartments within the cell) called Chloroplasts. Although all cells in the green parts of a plant have chloroplasts, most of the energy is captured in the leaves. The cells in the interior tissues of a leaf, called the mesophyll, can contain between 450,000 and 800,000 chloroplasts for every square millimeter of leaf. The surface of the leaf is uniformly coated with a water-resistant waxy cuticle that protects the leaf from excessive evaporation of water and decreases the absorption of ultraviolet or blue light to reduce heating. The transparent epidermis layer allows light to pass through to the palisade mesophyll cells where most of the photosynthesis takes place.

Plants use up to 90% of the light that strikes them, whereas commercial solar panels use less than 30%. This is achieved by groups of chlorophyll molecules spending a long time in a superposition of states.^[4]

In algae and bacteria

Algae come in multiple forms from multicellular organisms like kelp, to microscopic, single-celled organisms. Although they are not as complex as land plants, photosynthesis takes place biochemically the same way. Very much like plants, algae have chloroplasts and chlorophyll, but various accessory pigments are present in some algae such as phycocyanin, carotenes and xanthophylls in green algae and phycoerythrin in red algae (rhodophytes), resulting in a wide variety of colors. All algae produce oxygen, and many are autotrophic. However, some are heterotrophic, relying on materials produced by other organisms. For example, in coral reefs, there is a mutualistic relationship between zooxanthellae and the coral polyps.^[5]

Photosynthetic bacteria do not have chloroplasts (or any membrane-bound organelles). Instead, photosynthesis takes place directly within the cell. Cyanobacteria contain thylakoid membranes very similar to those in chloroplasts and are the only prokaryotes that perform oxygen-generating photosynthesis. In fact chloroplasts are now considered to have evolved from an endosymbiotic bacterium, which was also an ancestor of and later gave rise to cyanobacterium. The other photosynthetic bacteria have a variety of different pigments, called bacteriochlorophylls, and do not produce oxygen. Some bacteria, such as Chromatium, oxidize hydrogen sulfide instead of water for photosynthesis, producing sulfur as waste.

Evolution

Plant cells with visible chloroplasts.

The ability to convert light energy to chemical energy confers a significant evolutionary advantage to living organisms. Early photosynthetic systems, such as those from green and purple sulfur and green and purple non-sulfur bacteria, are thought to have been anoxygenic, using various molecules as electron donors. Green and purple sulfur bacteria are thought to have used hydrogen and sulfur as an electron donor. Green nonsulfur bacteria used various amino and other organic acids. Purple nonsulfur bacteria used a variety of non-specific organic molecules. The use of these molecules is consistent with the geological evidence that the atmosphere was highly reduced at that time.^{[citation needed]}

Fossils of what are thought to be filamentous photosynthetic organisms have been dated at 3.4 billion years old.^[6]

Oxygen in the atmosphere exists due to the evolution of oxygenic photosynthesis, sometimes referred to as the oxygen catastrophe. Geological evidence suggests that oxygenic photosynthesis, such as that in cyanobacteria, became important during the Paleoproterozoic era around 2 billion years ago. Modern photosynthesis in plants and most photosynthetic prokaryotes is oxygenic. Oxygenic photosynthesis uses water as an electron donor which is oxidized into molecular oxygen by the absorption of a photon by the photosynthetic reaction centre.

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