As has been stated, carbohydrates are the most-important direct organic product of photosynthesis in the majority of green plants. The formation of a simple carbohydrate, glucose, is indicated by a chemical equation,Formation of a simple carbohydrate, glucose. basic products of photosynthesis

Little free glucose is produced in plants; instead, glucose units are linked to form starch or are joined with fructose, another sugar, to form sucrose (see carbohydrate).

Not only carbohydrates, as was once thought, but also amino acids, proteins, lipids (or fats), pigments, and other organic components of green tissues are synthesized during photosynthesis. Minerals supply the elements (e.g., nitrogen, N; phosphorus, P; sulfur, S) required to form these compounds. Chemical bonds are broken between oxygen (O) and carbon (C), hydrogen (H), nitrogen, and sulfur, and new bonds are formed in products that include gaseous oxygen (O2) and organic compounds. More energy is required to break the bonds between oxygen and other elements (e.g., in water, nitrate, and sulfate) than is released when new bonds form in the products. This difference in bond energy accounts for a large part of the light energy stored as chemical energy in the organic products formed during photosynthesis. Additional energy is stored in making complex molecules from simple ones.

Evolution of the process

Although life and the quality of the atmosphere today depend on photosynthesis, it is likely that green plants evolved long after the first living cells. When Earth was young, electrical storms and solar radiation probably provided the energy for the synthesis of complex molecules from abundant simpler ones, such as water, ammonia, and methane. The first living cells probably evolved from these complex molecules (see life: Production of polymers). For example, the accidental joining (condensation) of the amino acid glycine and the fatty acid acetate may have formed complex organic molecules known as porphyrins. These molecules, in turn, may have evolved further into colored molecules called pigments—e.g., chlorophylls of green plants, bacteriochlorophyll of photosynthetic bacteria, hemin (the red pigment of blood), and cytochromes, a group of pigment molecules essential in both photosynthesis and cellular respiration.

Primitive colored cells then had to evolve mechanisms for using the light energy absorbed by their pigments. At first, the energy may have been used immediately to initiate reactions useful to the cell. As the process for utilization of light energy continued to evolve, however, a larger part of the absorbed light energy probably was stored as chemical energy, to be used to maintain life. Green plants, with their ability to use light energy to convert carbon dioxide and water to carbohydrates and oxygen, are the culmination of this evolutionary process.

The first oxygenic (oxygen-producing) cells probably were the blue-green algae (cyanobacteria), which appeared about two billion to three billion years ago. These microscopic organisms are believed to have greatly increased the oxygen content of the atmosphere, making possible the development of aerobic (oxygen-using) organisms. Cyanophytes are prokaryotic cells; that is, they contain no distinct membrane-enclosed subcellular particles (organelles), such as nuclei and chloroplasts. Green plants, by contrast, are composed of eukaryotic cells, in which the photosynthetic apparatus is contained within membrane-bound chloroplasts. The complete genome sequences of cyanobacteria and higher plants provide evidence that the first photosynthetic eukaryotes were likely the red algae that developed when nonphotosynthetic eukaryotic cells engulfed cyanobacteria. Within the host cells, these cyanobacteria evolved into chloroplasts.

There are a number of photosynthetic bacteria that are not oxygenic (e.g., the sulfur bacteria previously discussed). The evolutionary pathway that led to these bacteria diverged from the one that resulted in oxygenic organisms. In addition to the absence of oxygen production, nonoxygenic photosynthesis differs from oxygenic photosynthesis in two other ways: light of longer wavelengths is absorbed and used by pigments called bacteriochlorophylls, and reduced compounds other than water (such as hydrogen sulfide or organic molecules) provide the electrons needed for the reduction of carbon dioxide.

Factors that influence the rate of photosynthesis

The rate of photosynthesis is defined in terms of the rate of oxygen production either per unit mass (or area) of green plant tissues or per unit weight of total chlorophyll. The amount of light, the carbon dioxide supply, temperature, water supply, and the availability of minerals are the most important environmental factors that affect the rate of photosynthesis in land plants. The rate of photosynthesis is also determined by the plant species and its physiological state—e.g., its health, its maturity, and whether it is in flower.

Light intensity and temperature

As has been mentioned, the complex mechanism of photosynthesis includes a photochemical, or light-harvesting, stage and an enzymatic, or carbon-assimilating, stage that involves chemical reactions. These stages can be distinguished by studying the rates of photosynthesis at various degrees of light saturation (i.e., intensity) and at different temperatures. Over a range of moderate temperatures and at low to medium light intensities (relative to the normal range of the plant species), the rate of photosynthesis increases as the intensity increases and is relatively independent of temperature. As the light intensity increases to higher levels, however, the rate becomes saturated; light “saturation” is achieved at a specific light intensity, dependent on species and growing conditions. In the light-dependent range before saturation, therefore, the rate of photosynthesis is determined by the rates of photochemical steps. At high light intensities, some of the chemical reactions of the dark stage become rate-limiting. In many land plants, a process called photorespiration occurs, and its influence upon photosynthesis increases with rising temperatures. More specifically, photorespiration competes with photosynthesis and limits further increases in the rate of photosynthesis, especially if the supply of water is limited (see below Photorespiration).

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Carbon dioxide

Included among the rate-limiting steps of the dark stage of photosynthesis are the chemical reactions by which organic compounds are formed by using carbon dioxide as a carbon source. The rates of these reactions can be increased somewhat by increasing the carbon dioxide concentration. Since the middle of the 19th century, the level of carbon dioxide in the atmosphere has been rising because of the extensive combustion of fossil fuels, cement production, and land-use changes associated with deforestation. The atmospheric level of carbon dioxide climbed from about 0.028 percent in 1860 to 0.032 percent by 1958 (when improved measurements began) and to 0.041 percent by 2020. This increase in carbon dioxide directly increases plant photosynthesis up to a point, but the size of the increase depends on the species and physiological condition of the plant. Furthermore, most scientists maintain that increasing levels of atmospheric carbon dioxide affect climate, increasing global temperatures and changing rainfall patterns. Such changes will also affect photosynthesis rates.

Water

For land plants, water availability can function as a limiting factor in photosynthesis and plant growth. Besides the requirement for a small amount of water in the photosynthetic reaction itself, large amounts of water are transpired from the leaves; that is, water evaporates from the leaves to the atmosphere via the stomata. Stomata are small openings through the leaf epidermis, or outer skin; they permit the entry of carbon dioxide but inevitably also allow the exit of water vapor. The stomata open and close according to the physiological needs of the leaf. In hot and arid climates the stomata may close to conserve water, but this closure limits the entry of carbon dioxide and hence the rate of photosynthesis. The decreased transpiration means there is less cooling of the leaves and hence leaf temperatures rise. The decreased carbon dioxide concentration inside the leaves and the increased leaf temperatures favor the wasteful process of photorespiration. If the level of carbon dioxide in the atmosphere increases, more carbon dioxide could enter through a smaller opening of the stomata, so more photosynthesis could occur with a given supply of water.

Minerals

Several minerals are required for healthy plant growth and for maximum rates of photosynthesis. Nitrogen, sulfate, phosphate, iron, magnesium, calcium, and potassium are required in substantial amounts for the synthesis of amino acids, proteins, coenzymes, deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), chlorophyll and other pigments, and other essential plant constituents. Smaller amounts of such elements as manganese, copper, and chloride are required in photosynthesis. Some other trace elements are needed for various nonphotosynthetic functions in plants.

Internal factors

Each plant species is adapted to a range of environmental factors. Within this normal range of conditions, complex regulatory mechanisms in the plant’s cells adjust the activities of enzymes (i.e., organic catalysts). These adjustments maintain a balance in the overall photosynthetic process and control it in accordance with the needs of the whole plant. With a given plant species, for example, doubling the carbon dioxide level might cause a temporary increase of nearly twofold in the rate of photosynthesis; a few hours or days later, however, the rate might fall to the original level because photosynthesis produced more sucrose than the rest of the plant could use. By contrast, another plant species provided with such carbon dioxide enrichment might be able to use more sucrose, because it had more carbon-demanding organs, and would continue to photosynthesize and to grow faster throughout most of its life cycle.