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photosynthesis
(redirected from Photosyntheis)

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photosynthesis (fō'tōsĭn`thəsĭs), process in which green plants utilize the energy of sunlight to manufacture carbohydrates from carbon dioxide and water in the presence of chlorophyll. Some of the plants that lack chlorophyll, e.g., the Indian pipe Indian pipe, common name for the genus Monotropa and for the family Monotropaceae, low flowering plants of north temperate zones. They are chlorophylless saprophytes with a funguslike appearance.
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, secure their nutrients from organic material, as do animals, and a few bacteria manufacture their own carbohydrates with hydrogen and energy obtained from inorganic compounds (e.g., hydrogen sulfide) in a process called chemosynthesis chemosynthesis, process in which carbohydrates are manufactured from carbon dioxide and water using chemical nutrients as the energy source, rather than the sunlight used for energy in photosynthesis . Most life on earth is fueled directly or indirectly by sunlight.
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. However, the vast majority of plants contain chlorophyll—concentrated, in the higher land plants, in the leaves.

In these plants water is absorbed by the roots and carried to the leaves by the xylem, and carbon dioxide is obtained from air that enters the leaves through the stomata and diffuses to the cells containing chlorophyll. The green pigment chlorophyll chlorophyll (klôr`əfĭl')
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 is uniquely capable of converting the active energy of light into a latent form that can be stored (in food) and used when needed.

The Photosynthetic Process

The initial process in photosynthesis is the decomposition of water (H2O) into oxygen, which is released, and hydrogen; direct light is required for this process. The hydrogen and the carbon and oxygen of carbon dioxide (CO2) are then converted into a series of increasingly complex compounds that result finally in a stable organic compound, glucose (C6H12O6), and water. This phase of photosynthesis utilizes stored energy and therefore can proceed in the dark. The simplified equation used to represent this overall process is 6CO2+12H2O+energy=C6H12O6+6O2+6H2O. In general, the results of this process are the reverse of those in respiration, in which carbohydrates are oxidized to release energy, with the production of carbon dioxide and water.

The intermediary reactions before glucose is formed involve several enzymes, which react with the coenzyme ATP (see adenosine triphosphate adenosine triphosphate (ATP) (ədĕn`əsēn trī'fŏs`fāt)
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) to produce various molecules. Studies using radioactive carbon have indicated that among the intermediate products are three-carbon molecules from which acids and amino acids, as well as glucose, are derived. This suggests that fats and proteins are also products of photosynthesis. The main product, glucose, is the fundamental building block of carbohydrates (e.g., sugars, starches, and cellulose). The water-soluble sugars (e.g., sucrose and maltose) are used for immediate energy. The insoluble starches are stored as tiny granules in various parts of the plant—chiefly the leaves, roots (including tubers), and fruits—and can be broken down again when energy is needed. Cellulose is used to build the rigid cell walls that are the principal supporting structure of plants.

Importance of Photosynthesis

Animals and plants both synthesize fats and proteins from carbohydrates; thus glucose is a basic energy source for all living organisms. The oxygen released (with water vapor, in transpiration) as a photosynthetic byproduct, principally of phytoplankton, provides most of the atmospheric oxygen vital to respiration in plants and animals, and animals in turn produce carbon dioxide necessary to plants. Photosynthesis can therefore be considered the ultimate source of life for nearly all plants and animals by providing the source of energy that drives all their metabolic processes.

Bibliography

See I. Asimov, Photosynthesis (1969); R. M. Devlin and A. V. Barker, Photosynthesis (1972).


photosynthesis

Enlarge picture
The light reaction of photosynthesis. The light reaction occurs in two photosystems (units of …
(credit: © Merriam-Webster Inc.)
Process by which green plants and certain other organisms transform light into chemical energy. In green plants, light energy is captured by chlorophyll in the chloroplasts of the leaves and used to convert water, carbon dioxide, and minerals into oxygen and energy-rich organic compounds (simple and complex sugars) that are the basis of both plant and animal life. Photosynthesis consists of a number of photochemical and enzymatic reactions. It occurs in two stages. During the light-dependent stage (light reaction), chlorophyll absorbs light energy, which excites some electrons in the pigment molecules to higher energy levels; these leave the chlorophyll and pass along a series of molecules, generating formation of NADPH (an enzyme) and high-energy ATP molecules. Oxygen, released as a by-product, passes into the atmosphere through pores in the leaves. NADPH and ATP drive the second stage, the dark reaction (or Calvin cycle, discovered by Melvin Calvin), which does not require light. During this stage glucose is generated using atmospheric carbon dioxide. Photosynthesis is crucial for maintaining life on Earth; if it ceased, there would soon be little food or other organic matter on the planet, and most types of organisms would disappear.


photosynthesis
(in plants) the synthesis of organic compounds from carbon dioxide and water (with the release of oxygen) using light energy absorbed by chlorophyll

photosynthesis [¦fōd·ō′sin·thə·səs]
(biochemistry)
Synthesis of chemical compounds in light, especially the manufacture of organic compounds (primarily carbohydrates) from carbon dioxide and a hydrogen source (such as water), with simultaneous liberation of oxygen, by chlorophyll-containing plant cells.

Photosynthesis

The manufacture in light of organic compounds (primarily certain carbohydrates) from inorganic materials by chlorophyll- or bacteriochlorophyll-containing cells. This process requires a supply of energy in the form of light. In chlorophyll-containing plant cells and in cyanobacteria, photosynthesis involves oxidation of water (H2O) to oxygen molecules, which are released into the environment. In contrast, bacterial photosynthesis does not involve O2 evolution—instead of H2O, other electron donors, such as H2S, are used. This article will focus on photosynthesis in plants. See Bacterial physiology and metabolism, Chlorophyll, Plant respiration

The light energy absorbed by the pigments of photosynthesizing cells, especially by the pigment chlorophyll or bacteriochlorophyll, is efficiently converted into stored chemical energy. Together, the two aspects of photosynthesis—the conversion of inorganic into organic matter, and the conversion of light energy into chemical energy—make it the fundamental process of life on Earth: it is the ultimate source of all living matter and of all life energy.

The net overall chemical reaction of plant photosynthesis is shown in the equation below,

where {CH2O} stands for a carbohydrate (sugar).

The photochemical reaction in photosynthesis belongs to the type known as oxidation-reduction, with CO2 acting as the oxidant (hydrogen or electron acceptor) and water as the reductant (hydrogen or electron donor). The unique characteristic of this particular oxidation-reduction is that it goes “in the wrong direction” energetically; that is, it converts chemically stable materials into chemically unstable products. Light energy is used to make this “uphill” reaction possible.

Photosynthesis is a complex, multistage process. Its main parts are (1) the primary photochemical process in which light energy absorbed by chlorophyll is converted into chemical energy, in the form of some energy-rich intermediate products; and (2) the enzyme-catalyzed “dark” (that is, not photochemical) reactions by which these intermediates are converted into the final products—carbohydrates and free oxygen.

Experiments suggest that plants contain two pigment systems. One (called photosystem I, or PS I, sensitizing reaction I) contains the major part of chlorophyll a; the other (called photosystem II, or PS II, sensitizing reaction II) contains some chlorophyll a and the major part of chlorophyll b or other auxiliary pigments (for example, the red and blue pigments, called phycobilins, in red and blue-green algae, and the brown pigment fucoxanthol in brown algae and diatoms). It appears that efficient photosynthesis requires the absorption of an equal number of light quanta in PS I and in PS II; and that within both systems excitation energy undergoes resonance migration from one pigment to another until it ends in special molecules of chlorophyll a called the reaction centers. The latter molecules then enter into a series of chemical reactions that result in the oxidation of water to produce O2 and the reduction of nicotinamide adenine dinucleotide phosphate (NADP+). Chromatophores from photosynthetic bacteria and chloroplasts from green plants, when illuminated in the presence of adenosine diphosphate (ADP) and inorganic phosphate, also use light energy to synthesize adenosine triphosphate (ATP); this photophosphorylation could be associated with some energy-releasing step in photosynthesis.

Schematic outline of the Calvin (C 3 ) carbon dioxide assimilation cycleenlarge picture
Schematic outline of the Calvin (C3) carbon dioxide assimilation cycle

The light-dependent conversion of radiant energy into chemical energy as ATP and reduced nicotinamide adenine dinucleotide phosphate (NADPH) serves as a prelude to the utilization of these compounds for the reductive fixation of CO2 into organic molecules. Such molecules, broadly designated as photosynthates, are usually but not invariably in the form of carbohydrates such as glucose polymers or sucrose, and form the base for the nutrition of all living things. Collectively, the biochemical processes by which CO2 is assimilated into organic molecules are known as the photosynthetic dark reactions, not because they must occur in darkness, but because light—in contrast to the photosynthetic light reactions—is not required.

C3 photosynthesis

The essential details of C3 photosynthesis can be seen in Fig. 1. Three molecules of CO2 combine with three molecules of the five-carbon compound ribulose bisphosphate (RuBP) in a reaction catalyzed by RuBP carboxylase to form three molecules of an enzyme-bound six-carbon compound. These are hydrolyzed into six molecules of the three-carbon compound phosphoglyceric acid (PGA), which are phosphorylated by the conversion of six molecules of ATP (releasing ADP for photophosphorylation via the light reactions). The resulting compounds are reduced by the NADPH formed in photosynthetic light reactions to form six molecules of the three-carbon compound phosphoglyceraldehyde (PGAL). One molecule of PGAL is made available for combination with another three-carbon compound, dihydroxyacetone phosphate, which is isomerized from a second PGAL (requiring a second “turn” of the Calvin-cycle wheel) to form a six-carbon sugar. The other five PGAL molecules, through a complex series of enzymatic reactions, are rearranged into three molecules of RuBP, which can again be carboxylated with CO2 to start the cycle turning again. The net product of two “turns” of the cycle, a six-carbon sugar (glucose-6-phosphate) is formed either within the chloroplast in a pathway leading to starch (a polymer of many glucose molecules), or externally in the cytoplasm in a pathway leading to sucrose (condensed from two six-carbon sugars, glucose and fructose).

Schematic outline of the Hatch-Slack (C 4 ) carbon dioxide assimilation route in two cell types of a NADP-ME-type plantenlarge picture
Schematic outline of the Hatch-Slack (C4) carbon dioxide assimilation route in two cell types of a NADP-ME-type plant

C4 photosynthesis

Initially, the C3 cycle was thought to be the only route for CO2 assimilation, although it was recognized by plant anatomists that some rapidly growing plants (such as maize, sugarcane, and sorghum) possessed an unusual organization of the photosynthetic tissues in their leaves (Kranz morphology). It was then demonstrated that plants having the Kranz anatomy utilized an additional CO2 assimilation route now known as the C4-dicarboxylic acid pathway (Fig. 2). Carbon dioxide enters a mesophyll cell, where it combines with the three-carbon compound phosphoenolpyruvate (PEP) to form a four-carbon acid, oxaloacetic acid, which is reduced to malic acid or transaminated to aspartic acid. The four-carbon acid moves into bundle sheath cells, where the acid is decarboxylated, the CO2 assimilated via the C3 cycle, and the resulting three-carbon compound, pyruvic acid, moves back into the mesophyll cell and is transformed into PEP, which can be carboxylated again. The two cell types, mesophyll and bundle sheath, are not necessarily adjacent, but in all documented cases of C4 photosynthesis the organism had two distinct types of green cells. C4 metabolism is classified into three types, depending on the decarboxylation reaction used with the four-carbon acid in the bundle sheath cells:

1. NADP-ME type (sorghum):

2. NAD-ME type (Atriplex species):

3. PCK type (Panicum species):

CAM photosynthesis

Under arid and desert conditions, where soil water is in short supply, transpiration during the day when temperatures are high and humidity is low may rapidly deplete the plant of water, leading to desiccation and death. By keeping stomata closed during the day, water can be conserved, but the uptake of CO2, which occurs entirely through the stomata, is prevented. Desert plants in the Crassulaceae, Cactaceae, Euphorbiaceae, and 15 other families evolved, apparently independently of C4 plants, an almost identical strategy of assimilating CO2 by which the CO2 is taken in at night when the stomata open; water loss is low because of the reduced temperatures and correspondingly higher humidities. First studied in plants of the Crassulaceae, the process has been called crassulacean acid metabolism (CAM).

In contrast to C4, where two cell types cooperate, the entire process occurs within an individual cell; the separation of C4 and C3 is thus temporal rather than spatial. At night, CO2 combines with PEP through the action of PEP carboxylase, resulting in the formation of oxaloacetic acid and its conversion into malic acid. The PEP is formed from starch or sugar via the glycolytic route of respiration. Thus, there is a daily reciprocal relationship between starch (a storage product of C3 photosynthesis) and the accumulation of malic acid (the terminal product of nighttime CO2 assimilation).



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