Write a summary equation for the reactions of photosynthesis. General and partial equations of photosynthesis

Photosynthesis is the process of transforming the energy of light absorbed by the body into the chemical energy of organic (and inorganic) compounds.

The process of photosynthesis is expressed by the overall equation:

6CO 2 + 6H 2 O ® C 6 H 12 O 6 + 6O 2.

In the light in a green plant, organic substances are formed from extremely oxidized substances - carbon dioxide and water, and molecular oxygen is released. In the process of photosynthesis, not only CO 2 is reduced, but also nitrates or sulfates, and energy can be directed to various endergonic processes, including the transport of substances.

The general equation for photosynthesis can be represented as:

12 H 2 O → 12 [H 2] + 6 O 2 (light reaction)

6 CO 2 + 12 [H 2] → C 6 H 12 O 6 + 6 H 2 O (dark reaction)

6 CO 2 + 12 H 2 O → C 6 H 12 O 6 + 6 H 2 O + 6 O 2

or in terms of 1 mol of CO 2:

CO 2 + H 2 O CH 2 O + O 2

All the oxygen released during photosynthesis comes from water. The water on the right side of the equation cannot be reduced because its oxygen comes from CO 2 . Using the methods of labeled atoms, it was obtained that H 2 O in chloroplasts is heterogeneous and consists of water coming from the external environment and water formed during photosynthesis. Both types of water are used in the process of photosynthesis. Evidence of the formation of O 2 in the process of photosynthesis is the work of the Dutch microbiologist Van Niel, who studied bacterial photosynthesis, and came to the conclusion that the primary photochemical reaction of photosynthesis is the dissociation of H 2 O, and not the decomposition of CO 2. Capable of photosynthetic assimilation of CO 2 bacteria (except cyanobacteria) are used as reducing agents H 2 S, H 2 , CH 3 and others, and do not emit O 2 . This type of photosynthesis is called photoreduction:

CO 2 + H 2 S → [CH 2 O] + H 2 O + S 2 or

CO 2 + H 2 A → [CH 2 O] + H 2 O + 2A,

where H 2 A - oxidizes the substrate, a hydrogen donor (in higher plants it is H 2 O), and 2A is O 2. Then the primary photochemical act in plant photosynthesis should be the decomposition of water into an oxidizing agent [OH] and a reducing agent [H]. [H] restores CO 2, and [OH] participates in the reactions of the release of O 2 and the formation of H 2 O.

Solar energy with the participation of green plants and photosynthetic bacteria is converted into free energy of organic compounds. To carry out this unique process, during evolution, a photosynthetic apparatus was created containing: I) a set of photoactive pigments capable of absorbing electromagnetic radiation of certain spectral regions and storing this energy in the form of electronic excitation energy, and 2) a special apparatus for converting electronic excitation energy into various forms chemical energy. First of all, this redox energy , associated with the formation of highly reduced compounds, electrochemical potential energy, due to the formation of electrical and proton gradients on the conjugating membrane (Δμ H +), phosphate bond energy of ATP and other macroergic compounds, which is then converted into free energy of organic molecules.

All these types of chemical energy can be used in the process of life for the absorption and transmembrane transport of ions and in most metabolic reactions, i.e. in a constructive exchange.

The ability to use solar energy and introduce it into biospheric processes determines the "cosmic" role of green plants, which was written about by the great Russian physiologist K.A. Timiryazev.

The process of photosynthesis is a very complex system of spatial and temporal organization. The use of high-speed methods of pulse analysis made it possible to establish that the process of photosynthesis includes reactions of different rates - from 10 -15 s (energy absorption and migration processes occur in the femtosecond time interval) to 10 4 s (formation of photosynthesis products). The photosynthetic apparatus includes structures with sizes from 10 -27 m 3 at the lowest molecular level to 10 5 m 3 at the crop level.

Concept of photosynthesis. The entire complex set of reactions that make up the process of photosynthesis can be represented by a schematic diagram, which displays the main stages of photosynthesis and their essence. In the modern scheme of photosynthesis, four stages can be distinguished, which differ in the nature and rate of reactions, as well as in the meaning and essence of the processes occurring at each stage:

* - SSC - light harvesting antenna complex of photosynthesis - a set of photosynthetic pigments - chlorophylls and carotenoids; RC - photosynthesis reaction center - chlorophyll dimer a; ETC - the electron transport chain of photosynthesis - is localized in the membranes of chloroplast thylakoids (conjugated membranes), includes quinones, cytochromes, iron-sulfur cluster proteins and other electron carriers.

Stage I - physical. It includes reactions of photophysical nature of the absorption of energy by pigments (P), its storage in the form of electronic excitation energy (P *) and migration to the reaction center (RC). All reactions are extremely fast and proceed at a rate of 10 -15 - 10 -9 s. Primary reactions of energy absorption are localized in light-harvesting antenna complexes (LSCs).

Stage II - photochemical. Reactions are localized in reaction centers and proceed at a rate of 10 -9 s. At this stage of photosynthesis, the energy of electronic excitation of the pigment of the reaction center (P (RC)) is used for charge separation. In this case, an electron with a high energy potential is transferred to the primary acceptor A, and the resulting system with separated charges (P (RC) - A) contains a certain amount of energy already in chemical form. The oxidized pigment P (RC) restores its structure due to the oxidation of the donor (D).

The transformation of one type of energy into another occurring in the reaction center is the central event of the photosynthesis process, which requires severe conditions for the structural organization of the system. At present, molecular models of reaction centers in plants and bacteria are generally known. Their similarity in structural organization was established, which indicates a high degree of conservatism of the primary processes of photosynthesis.

The primary products formed at the photochemical stage (P * , A -) are very labile, and the electron can return to the oxidized pigment P * (recombination process) with a useless loss of energy. Therefore, a rapid further stabilization of the formed reduced products with a high energy potential is necessary, which is carried out at the next, III stage of photosynthesis.

Stage III - electron transport reactions. A chain of carriers with different redox potential (E n ) forms the so-called electron transport chain (ETC). The redox components of ETC are organized in chloroplasts in the form of three main functional complexes - photosystem I (PSI), photosystem II (PSII), cytochrome b 6 f-complex, which provides a high speed of the electron flow and the possibility of its regulation. As a result of the work of the ETC, highly reduced products are formed: reduced ferredoxin (PD restore) and NADPH, as well as energy-rich ATP molecules, which are used in the dark reactions of CO 2 reduction that make up the IV stage of photosynthesis.

Stage IV - "dark" reactions of absorption and reduction of carbon dioxide. The reactions take place with the formation of carbohydrates, the end products of photosynthesis, in the form of which the solar energy is stored, absorbed and converted in the "light" reactions of photosynthesis. The speed of "dark" enzymatic reactions is 10 -2 - 10 4 s.

Thus, the entire course of photosynthesis is carried out with the interaction of three flows - the flow of energy, the flow of electrons and the flow of carbon. The conjugation of the three streams requires precise coordination and regulation of their constituent reactions.

The planetary role of photosynthesis

Photosynthesis, having arisen at the first stages of the evolution of life, remains the most important process of the biosphere. It is green plants through photosynthesis that provide the cosmic connection of life on Earth with the Universe and determine the ecological well-being of the biosphere up to the possibility of the existence of human civilization. Photosynthesis is not only a source of food resources and minerals, but also a factor in the balance of biospheric processes on Earth, including the constancy of the content of oxygen and carbon dioxide in the atmosphere, the state of the ozone screen, the content of humus in the soil, the greenhouse effect, etc.

The global net productivity of photosynthesis is 7–8·10 8 tons of carbon per year, of which 7% is directly used for food, fuel and building materials. Currently, the consumption of fossil fuels is approximately equal to the formation of biomass on the planet. Every year, in the course of photosynthesis, 70–120 billion tons of oxygen enters the atmosphere, which ensures the respiration of all organisms. One of the most important consequences of oxygen release is the formation of an ozone screen in the upper atmosphere at an altitude of 25 km. Ozone (O 3) is formed as a result of photodissociation of O 2 molecules under the action of solar radiation and traps most of the ultraviolet rays that have a detrimental effect on all living things.

Stabilization of CO 2 content in the atmosphere is also an essential factor in photosynthesis. At present, the content of CO 2 is 0.03–0.04% by volume of air, or 711 billion tons in terms of carbon. The respiration of organisms, the World Ocean, in the waters of which 60 times more CO 2 is dissolved than is in the atmosphere, the production activities of people, on the one hand, photosynthesis, on the other, maintain a relatively constant level of CO 2 in the atmosphere. Carbon dioxide in the atmosphere, as well as water, absorb infrared rays and retain a significant amount of heat on Earth, providing the necessary conditions for life.

However, over the past decades, due to increasing human burning of fossil fuels, deforestation and decomposition of humus, a situation has developed where technological progress has made the balance of atmospheric phenomena negative. The situation is aggravated by demographic problems: every day 200 thousand people are born on Earth, who need to be provided with vital resources. These circumstances put the study of photosynthesis in all its manifestations, from the molecular organization of the process to biospheric phenomena, into the rank of the leading problems of modern natural science. The most important tasks are to increase the photosynthetic productivity of agricultural crops and plantations, as well as to create effective biotechnologies for phototrophic syntheses.

K.A. Timiryazev was the first to study space role green plants. Photosynthesis is the only process on Earth that takes place on a grandiose scale and is associated with the conversion of the energy of sunlight into the energy of chemical compounds. This cosmic energy, stored by green plants, forms the basis of the vital activity of all other heterotrophic organisms on Earth, from bacteria to humans. There are 5 main aspects of space and planetary activity of green plants.

1. Accumulation of organic matter. In the process of photosynthesis, land plants form 100-172 billion tons. biomass per year (in terms of dry matter), and plants of the seas and oceans - 60-70 billion tons. The total mass of plants on Earth is currently 2402.7 billion tons, and 90% of this mass is cellulose. About 2402.5 billion tons. accounted for by terrestrial plants and 0.2 billion tons. - on plants of the hydrosphere (lack of light!). The total mass of animals and microorganisms on Earth is 23 billion tons, that is, 1% of the mass of plants. Of this amount, ~ 20 billion tons. accounts for the inhabitants of the land and ~ 3 billion tons. - on the inhabitants of the hydrosphere. During the existence of life on Earth, the organic remains of plants and animals accumulated and modified (litter, humus, peat, and in the lithosphere - coal; in the seas and oceans - sedimentary rocks). When descending into deeper regions of the lithosphere, gas and oil were formed from these remains under the action of microorganisms, elevated temperatures and pressure. The mass of organic matter in the litter is ~ 194 billion tons; peat - 220 billion tons; humus ~ 2500 billion tons. Oil and gas - 10,000 - 12,000 billion tons. The content of organic matter in sedimentary rocks in terms of carbon is ~ 2 10 16 t. Especially intensive accumulation of organic matter occurred in Paleozoic(~ 300 million years ago). The stored organic matter is intensively used by man (wood, minerals).

2. Ensuring the constancy of the content of CO 2 in the atmosphere. The formation of humus, sedimentary rocks, combustible minerals removed significant amounts of CO 2 from the carbon cycle. In the Earth's atmosphere, CO 2 became less and less, and at present its content is ~ 0.03–0.04% by volume, or ~ 711 billion tons. in terms of carbon. In the Cenozoic era, the content of CO 2 in the atmosphere stabilized and experienced only daily, seasonal and geochemical fluctuations (plant stabilization at the modern level). Stabilization of the content of CO 2 in the atmosphere is achieved by balanced binding and release of CO 2 on a global scale. The binding of CO 2 in photosynthesis and the formation of carbonates (sedimentary rocks) is compensated by the release of CO 2 due to other processes: ~ 25 billion tons; breathing of humans and animals - ~ 1.6 billion tons. economic activities of people ~ 5 billion tons; geochemical processes ~ 0.05 billion tons. Total ~ 41.65 billion tons If CO 2 did not enter the atmosphere, its entire available supply would be bound in 6–7 years. The World Ocean is a powerful reserve of CO 2, 60 times more CO 2 is dissolved in its waters than it is in the atmosphere. So, photosynthesis, respiration and the carbonate system of the ocean maintain a relatively constant level of CO 2 in the atmosphere. Due to human economic activity (burning of combustible minerals, deforestation, decomposition of humus), the content of CO 2 in the atmosphere began to increase by ~ 0.23% per year. This circumstance may have global consequences, since the content of CO 2 in the atmosphere affects the thermal regime of the planet.

3. Greenhouse effect. The Earth's surface receives heat mainly from the Sun. Some of this heat is returned in the form of infrared rays. CO 2 and H 2 O contained in the atmosphere absorb infrared rays and thus retain a significant amount of heat on Earth (greenhouse effect). Microorganisms and plants in the process of respiration or fermentation supply ~ 85% of the total amount of CO 2 entering the atmosphere annually and, as a result, affect the thermal regime of the planet. The upward trend in CO 2 content in the atmosphere can lead to an increase in the average temperature on the Earth's surface, melting of glaciers (mountains and polar ice) and flooding of coastal zones. However, it is possible that an increase in the concentration of CO 2 in the atmosphere will enhance plant photosynthesis, which will lead to the fixation of excess amounts of CO 2 .

4. Accumulation of O 2 in the atmosphere. Initially, O 2 was present in the Earth's atmosphere in trace amounts. It currently accounts for ~21% by air volume. The appearance and accumulation of O 2 in the atmosphere is associated with the vital activity of green plants. Every year ~ 70–120 billion tons enter the atmosphere. O 2 formed in photosynthesis. Forests play a special role in this: 1 hectare of forest in 1 hour gives O 2, enough for 200 people to breathe.

5. Ozone shield formation at an altitude of ~ 25 km. O 3 is formed during the dissociation of O 2 under the action of solar radiation. The O 3 layer retains most of the UV (240-290 nm), which is detrimental to living things. The destruction of the planet's ozone screen is one of the global problems of our time.

Photosynthesis is the conversion of light energy into chemical bond energy. organic compounds.

Photosynthesis is characteristic of plants, including all algae, a number of prokaryotes, including cyanobacteria, and some unicellular eukaryotes.

In most cases, photosynthesis produces oxygen (O2) as a by-product. However, this is not always the case as there are several different pathways for photosynthesis. In the case of oxygen release, its source is water, from which hydrogen atoms are split off for the needs of photosynthesis.

Photosynthesis consists of many reactions in which various pigments, enzymes, coenzymes, etc. participate. The main pigments are chlorophylls, in addition to them, carotenoids and phycobilins.

In nature, two ways of plant photosynthesis are common: C 3 and C 4. Other organisms have their own specific reactions. What unites these different processes under the term “photosynthesis” is that in all of them, in total, the conversion of photon energy into a chemical bond occurs. For comparison: during chemosynthesis, the energy of the chemical bond of some compounds (inorganic) is converted into others - organic.

There are two phases of photosynthesis - light and dark. The first depends on the light radiation (hν), which is necessary for the reactions to proceed. The dark phase is light independent.

In plants, photosynthesis takes place in chloroplasts. As a result of all reactions, primary organic substances are formed, from which carbohydrates, amino acids, fatty acids, etc. are then synthesized. Usually, the total reaction of photosynthesis is written in relation to glucose - the most common product of photosynthesis:

6CO 2 + 6H 2 O → C 6 H 12 O 6 + 6O 2

The oxygen atoms that make up the O 2 molecule are not taken from carbon dioxide, but from water. Carbon dioxide is a source of carbon which is more important. Due to its binding, plants have the opportunity to synthesize organic matter.

The chemical reaction presented above is a generalized and total. It is far from the essence of the process. So glucose is not formed from six individual molecules of carbon dioxide. The binding of CO 2 occurs in one molecule, which first attaches to an already existing five-carbon sugar.

Prokaryotes have their own characteristics of photosynthesis. So in bacteria, the main pigment is bacteriochlorophyll, and oxygen is not released, since hydrogen is not taken from water, but often from hydrogen sulfide or other substances. In blue-green algae, the main pigment is chlorophyll, and oxygen is released during photosynthesis.

Light phase of photosynthesis

In the light phase of photosynthesis, ATP and NADP·H 2 are synthesized due to radiant energy. It happens on the thylakoids of chloroplasts, where pigments and enzymes form complex complexes for the functioning of electrochemical circuits, through which electrons and partly hydrogen protons are transferred.

The electrons end up at the coenzyme NADP, which, being negatively charged, attracts some of the protons and turns into NADP H 2 . Also, the accumulation of protons on one side of the thylakoid membrane and electrons on the other creates an electrochemical gradient, the potential of which is used by the ATP synthetase enzyme to synthesize ATP from ADP and phosphoric acid.

The main pigments of photosynthesis are various chlorophylls. Their molecules capture the radiation of certain, partly different spectra of light. In this case, some electrons of chlorophyll molecules move to a higher energy level. This is an unstable state, and, in theory, electrons, by means of the same radiation, should give the energy received from outside into space and return to the previous level. However, in photosynthetic cells, excited electrons are captured by acceptors and, with a gradual decrease in their energy, are transferred along the chain of carriers.

On thylakoid membranes, there are two types of photosystems that emit electrons when exposed to light. Photosystems are a complex complex of mostly chlorophyll pigments with a reaction center from which electrons are torn off. In a photosystem, sunlight catches a lot of molecules, but all the energy is collected in the reaction center.

The electrons of photosystem I, having passed through the chain of carriers, restore NADP.

The energy of the electrons detached from photosystem II is used to synthesize ATP. And the electrons of photosystem II fill the electron holes of photosystem I.

The holes of the second photosystem are filled with electrons formed as a result of water photolysis. Photolysis also occurs with the participation of light and consists in the decomposition of H 2 O into protons, electrons and oxygen. It is as a result of the photolysis of water that free oxygen is formed. Protons are involved in the creation of an electrochemical gradient and the reduction of NADP. Electrons are received by the chlorophyll of photosystem II.

Approximate summary equation of the light phase of photosynthesis:

H 2 O + NADP + 2ADP + 2P → ½O 2 + NADP H 2 + 2ATP

Cyclic electron transport

The so-called non-cyclic light phase of photosynthesis. Is there some more cyclic electron transport when NADP reduction does not occur. In this case, electrons from photosystem I go to the carrier chain, where ATP is synthesized. That is, this electron transport chain receives electrons from photosystem I, not II. The first photosystem, as it were, implements a cycle: the emitted electrons return to it. On the way, they spend part of their energy on the synthesis of ATP.

Photophosphorylation and oxidative phosphorylation

The light phase of photosynthesis can be compared with the stage of cellular respiration - oxidative phosphorylation, which occurs on the mitochondrial cristae. There, too, ATP synthesis occurs due to the transfer of electrons and protons along the carrier chain. However, in the case of photosynthesis, energy is stored in ATP not for the needs of the cell, but mainly for the needs of the dark phase of photosynthesis. And if during respiration organic substances serve as the initial source of energy, then during photosynthesis it is sunlight. The synthesis of ATP during photosynthesis is called photophosphorylation rather than oxidative phosphorylation.

Dark phase of photosynthesis

For the first time the dark phase of photosynthesis was studied in detail by Calvin, Benson, Bassem. The cycle of reactions discovered by them was later called the Calvin cycle, or C 3 -photosynthesis. In certain groups of plants, a modified photosynthesis pathway is observed - C 4, also called the Hatch-Slack cycle.

In the dark reactions of photosynthesis, CO 2 is fixed. The dark phase takes place in the stroma of the chloroplast.

Recovery of CO 2 occurs due to the energy of ATP and the reducing power of NADP·H 2 formed in light reactions. Without them, carbon fixation does not occur. Therefore, although the dark phase does not directly depend on light, it usually also proceeds in light.

Calvin cycle

The first reaction of the dark phase is the addition of CO 2 ( carboxylatione) to 1,5-ribulose biphosphate ( ribulose 1,5-diphosphate) – RiBF. The latter is a doubly phosphorylated ribose. This reaction is catalyzed by the enzyme ribulose-1,5-diphosphate carboxylase, also called rubisco.

As a result of carboxylation, an unstable six-carbon compound is formed, which, as a result of hydrolysis, decomposes into two three-carbon molecules phosphoglyceric acid (PGA) is the first product of photosynthesis. FHA is also called phosphoglycerate.

RiBP + CO 2 + H 2 O → 2FGK

FHA contains three carbon atoms, one of which is part of the acidic carboxyl group (-COOH):

FHA is converted into a three-carbon sugar (glyceraldehyde phosphate) triose phosphate (TF), which already includes an aldehyde group (-CHO):

FHA (3-acid) → TF (3-sugar)

This reaction consumes the energy of ATP and the reducing power of NADP · H 2 . TF is the first carbohydrate of photosynthesis.

After that, most of the triose phosphate is spent on the regeneration of ribulose bisphosphate (RiBP), which is again used to bind CO 2 . Regeneration involves a series of ATP-consuming reactions involving sugar phosphates with 3 to 7 carbon atoms.

It is in this cycle of RiBF that the Calvin cycle is concluded.

A smaller part of the TF formed in it leaves the Calvin cycle. In terms of 6 bound molecules of carbon dioxide, the yield is 2 molecules of triose phosphate. The total reaction of the cycle with input and output products:

6CO 2 + 6H 2 O → 2TF

At the same time, 6 RiBP molecules participate in the binding and 12 FHA molecules are formed, which are converted into 12 TF, of which 10 molecules remain in the cycle and are converted into 6 RiBP molecules. Since TF is a three-carbon sugar, and RiBP is a five-carbon one, then in relation to carbon atoms we have: 10 * 3 = 6 * 5. The number of carbon atoms that provide the cycle does not change, all the necessary RiBP is regenerated. And six molecules of carbon dioxide included in the cycle are spent on the formation of two molecules of triose phosphate leaving the cycle.

The Calvin cycle, based on 6 bound CO 2 molecules, consumes 18 ATP molecules and 12 NADP · H 2 molecules, which were synthesized in the reactions of the light phase of photosynthesis.

The calculation is carried out for two triose phosphate molecules leaving the cycle, since the glucose molecule formed later includes 6 carbon atoms.

Triose phosphate (TP) is the end product of the Calvin cycle, but it can hardly be called the end product of photosynthesis, since it almost does not accumulate, but, reacting with other substances, turns into glucose, sucrose, starch, fats, fatty acids, amino acids. In addition to TF, FHA plays an important role. However, such reactions occur not only in photosynthetic organisms. In this sense, the dark phase of photosynthesis is the same as the Calvin cycle.

PHA is converted into a six-carbon sugar by stepwise enzymatic catalysis. fructose-6-phosphate, which turns into glucose. In plants, glucose can be polymerized into starch and cellulose. The synthesis of carbohydrates is similar to the reverse process of glycolysis.

photorespiration

Oxygen inhibits photosynthesis. The more O 2 in the environment, the less efficient the CO 2 sequestration process. The fact is that the enzyme ribulose bisphosphate carboxylase (rubisco) can react not only with carbon dioxide, but also with oxygen. In this case, the dark reactions are somewhat different.

Phosphoglycolate is phosphoglycolic acid. The phosphate group is immediately cleaved from it, and it turns into glycolic acid (glycolate). For its "utilization" oxygen is needed again. Therefore, the more oxygen in the atmosphere, the more it will stimulate photorespiration and the more oxygen the plant will need to get rid of the reaction products.

Photorespiration is the light-dependent consumption of oxygen and the release of carbon dioxide. That is, the exchange of gases occurs as during respiration, but takes place in chloroplasts and depends on light radiation. Photorespiration depends on light only because ribulose biphosphate is formed only during photosynthesis.

During photorespiration, carbon atoms are returned from glycolate to the Calvin cycle in the form of phosphoglyceric acid (phosphoglycerate).

2 Glycolate (C 2) → 2 Glyoxylate (C 2) → 2 Glycine (C 2) - CO 2 → Serine (C 3) → Hydroxypyruvate (C 3) → Glycerate (C 3) → FGK (C 3)

As you can see, the return is not complete, since one carbon atom is lost when two molecules of glycine are converted into one molecule of the amino acid serine, while carbon dioxide is released.

Oxygen is needed at the stages of conversion of glycolate to glyoxylate and glycine to serine.

The conversion of glycolate to glyoxylate and then to glycine occurs in peroxisomes, and serine is synthesized in mitochondria. Serine again enters the peroxisomes, where it first produces hydroxypyruvate, and then glycerate. Glycerate already enters the chloroplasts, where FHA is synthesized from it.

Photorespiration is typical mainly for plants with C3-type photosynthesis. It can be considered harmful, since energy is wasted on the conversion of glycolate into FHA. Apparently, photorespiration arose due to the fact that ancient plants were not ready for a large amount of oxygen in the atmosphere. Initially, their evolution took place in an atmosphere rich in carbon dioxide, and it was he who mainly captured the reaction center of the rubisco enzyme.

C 4 -photosynthesis, or the Hatch-Slack cycle

If in C 3 photosynthesis the first product of the dark phase is phosphoglyceric acid, which includes three carbon atoms, then in the C 4 pathway, the first products are acids containing four carbon atoms: malic, oxaloacetic, aspartic.

C 4 -photosynthesis is observed in many tropical plants, for example, sugar cane, corn.

C 4 -plants absorb carbon monoxide more efficiently, they have almost no photorespiration.

Plants in which the dark phase of photosynthesis proceeds along the C 4 pathway have a special leaf structure. In it, the conducting bundles are surrounded by a double layer of cells. The inner layer is the lining of the conducting beam. The outer layer is mesophyll cells. Chloroplast cell layers differ from each other.

Mesophilic chloroplasts are characterized by large grains, high activity of photosystems, absence of the enzyme RiBP carboxylase (rubisco) and starch. That is, the chloroplasts of these cells are adapted mainly for the light phase of photosynthesis.

In the chloroplasts of the cells of the conducting bundle, the grana are almost not developed, but the concentration of RiBP carboxylase is high. These chloroplasts are adapted for the dark phase of photosynthesis.

Carbon dioxide first enters the mesophyll cells, binds with organic acids, is transported in this form to the sheath cells, is released, and then binds in the same way as in C3 plants. That is, the C 4 -path complements rather than replaces C 3 .

In the mesophyll, CO 2 is added to phosphoenolpyruvate (PEP) to form oxaloacetate (acid), which includes four carbon atoms:

The reaction takes place with the participation of the PEP-carboxylase enzyme, which has a higher affinity for CO 2 than rubisco. In addition, PEP-carboxylase does not interact with oxygen, and therefore is not spent on photorespiration. Thus, the advantage of C4 photosynthesis lies in more efficient fixation of carbon dioxide, an increase in its concentration in the sheath cells, and, consequently, more efficient operation of RiBP carboxylase, which is almost not consumed for photorespiration.

Oxaloacetate is converted into a 4-carbon dicarboxylic acid (malate or aspartate), which is transported to the chloroplasts of the cells lining the vascular bundles. Here, the acid is decarboxylated (removal of CO2), oxidized (removal of hydrogen) and converted to pyruvate. Hydrogen restores NADP. Pyruvate returns to the mesophyll, where PEP is regenerated from it with the consumption of ATP.

The torn off CO 2 in the chloroplasts of the lining cells goes to the usual C 3 path of the dark phase of photosynthesis, i.e., to the Calvin cycle.

Photosynthesis along the Hatch-Slack pathway requires more energy.

It is believed that the C 4 pathway evolved later than the C 3 pathway and is in many ways an adaptation against photorespiration.

1. Give definitions of concepts.
Photosynthesis- the process of formation of organic substances from carbon dioxide and water in the light with the participation of photosynthetic pigments.
Autotrophs organisms that synthesize organic substances from inorganic substances.
Heterotrophs are organisms that are unable to synthesize organic substances from inorganic substances by photosynthesis or chemosynthesis.
Mixotrophs- organisms that can use various sources of carbon and energy.

2. Fill in the table.

3. Fill in the table.

4. Explain the essence of the statement of the great Russian scientist K. A. Timiryazev: "A log is a canned solar energy."
A log is a part of a tree, its tissues consist of accumulated organic compounds (cellulose, sugar, etc.), which were formed during photosynthesis.

5. Write the overall photosynthesis equation. Do not forget to specify the required conditions for the reactions to take place.

12. Choose a term and explain how its modern meaning corresponds to the original meaning of its roots.
The chosen term is mixotrophs.
Conformity. The term is specified, as organisms with a mixed type of nutrition are called, which are able to use various sources of carbon and energy.

13. Formulate and write down the main ideas of § 3.3.
According to the type of nutrition, all living organisms are divided into:
Autotrophs that synthesize organic substances from inorganic substances.
Heterotrophs that feed on ready-made organic matter.
Mixotrophs with mixed nutrition.
Photosynthesis is the process of formation of organic substances from carbon dioxide and water in the light with the participation of photosynthetic pigments by phototrophs.
It is divided into a light phase (water and H+ molecules are formed, which are necessary for the dark phase, and oxygen is also released) and dark (glucose is formed). The total photosynthesis equation: 6CO2 + 6H2O → C6H12O6 + 6O2. It flows in the light in the presence of chlorophyll. Thus, light energy is converted into
the energy of chemical bonds, and plants form for themselves glucose and sugars.

Organic (and inorganic) compounds.

The process of photosynthesis is expressed by the overall equation:

6CO 2 + 6H 2 O ® C 6 H 12 O 6 + 6O 2.

In the light, in a green plant, organic substances are formed from extremely oxidized substances - carbon dioxide and water, and molecular oxygen is released. In the process of photosynthesis, not only CO 2 is reduced, but also nitrates or sulfates, and energy can be directed to various endergonic processes, including the transport of substances.

The general equation for photosynthesis can be represented as:

12 H 2 O → 12 [H 2] + 6 O 2 (light reaction)

6 CO 2 + 12 [H 2] → C 6 H 12 O 6 + 6 H 2 O (dark reaction)

6 CO 2 + 12 H 2 O → C 6 H 12 O 6 + 6 H 2 O + 6 O 2

or in terms of 1 mol of CO 2:

CO 2 + H 2 O CH 2 O + O 2

All the oxygen released during photosynthesis comes from water. The water on the right side of the equation cannot be reduced because its oxygen comes from CO 2 . Using the methods of labeled atoms, it was obtained that H 2 O in chloroplasts is heterogeneous and consists of water coming from the external environment and water formed during photosynthesis. Both types of water are used in the process of photosynthesis.

Evidence of the formation of O 2 in the process of photosynthesis is the work of the Dutch microbiologist Van Niel, who studied bacterial photosynthesis, and came to the conclusion that the primary photochemical reaction of photosynthesis is the dissociation of H 2 O, and not the decomposition of CO 2. Capable of photosynthetic assimilation of CO 2 bacteria (except cyanobacteria) are used as reducing agents H 2 S, H 2 , CH 3 and others, and do not emit O 2 .

This type of photosynthesis is called photoreduction:

CO 2 + H 2 S → [CH 2 O] + H 2 O + S 2 or

CO 2 + H 2 A → [CH 2 O] + H 2 O + 2A,

where H 2 A - oxidizes the substrate, a hydrogen donor (in higher plants it is H 2 O), and 2A is O 2. Then the primary photochemical act in plant photosynthesis should be the decomposition of water into an oxidizing agent [OH] and a reducing agent [H]. [H] restores CO 2, and [OH] participates in the reactions of the release of O 2 and the formation of H 2 O.

Solar energy with the participation of green plants and photosynthetic bacteria is converted into free energy of organic compounds.

To implement this unique process, a photosynthetic apparatus was created in the course of evolution, containing:

I) a set of photoactive pigments capable of absorbing electromagnetic radiation of certain spectral regions and storing this energy in the form of electronic excitation energy, and

2) a special apparatus for converting the energy of electronic excitation into various forms of chemical energy.

First of all, this redox energy , associated with the formation of highly reduced compounds, electrochemical potential energy, due to the formation of electrical and proton gradients on the conjugating membrane (Δμ H +), energy of ATP phosphate bonds and other macroergic compounds, which is then converted into free energy of organic molecules.

All these types of chemical energy can be used in the process of life for the absorption and transmembrane transport of ions and in most metabolic reactions, i.e. in a constructive exchange.

The ability to use solar energy and introduce it into biospheric processes determines the “cosmic” role of green plants, which was written about by the great Russian physiologist K.A. Timiryazev.

The process of photosynthesis is a very complex system of spatial and temporal organization. The use of high-speed methods of pulsed analysis made it possible to establish that the process of photosynthesis includes reactions of different rates - from 10 -15 s (energy absorption and migration processes occur in the femtosecond time interval) to 10 4 s (formation of photosynthesis products). The photosynthetic apparatus includes structures with sizes from 10 -27 m 3 at the lowest molecular level to 10 5 m 3 at the level of crops.

Concept of photosynthesis.

The whole complex set of reactions that make up the process of photosynthesis can be represented by a schematic diagram, which displays the main stages of photosynthesis and their essence. In the modern scheme of photosynthesis, four stages can be distinguished, which differ in the nature and rate of reactions, as well as in the meaning and essence of the processes occurring at each stage:

I stage - physical. It includes reactions of photophysical nature of the absorption of energy by pigments (P), its storage in the form of electronic excitation energy (P *) and migration to the reaction center (RC). All reactions are extremely fast and proceed at a rate of 10 -15 - 10 -9 s. Primary reactions of energy absorption are localized in light-harvesting antenna complexes (SSCs).

Stage II - photochemical. Reactions are localized in reaction centers and proceed at a rate of 10 -9 s. At this stage of photosynthesis, the energy of the electronic excitation of the pigment (P (RC)) of the reaction center is used to separate charges. In this case, an electron with a high energy potential is transferred to the primary acceptor A, and the resulting system with separated charges (P (RC) - A) contains a certain amount of energy already in chemical form. The oxidized pigment P (RC) restores its structure due to the oxidation of the donor (D).

The transformation of one type of energy into another occurring in the reaction center is the central event of the photosynthesis process, requiring strict conditions for the structural organization of the system. At present, molecular models of reaction centers in plants and bacteria are generally known. Their similarity in structural organization was established, which indicates a high degree of conservatism of the primary processes of photosynthesis.

The primary products formed at the photochemical stage (P * , A -) are very labile, and the electron can return to the oxidized pigment P * (recombination process) with a useless loss of energy. Therefore, fast further stabilization of the formed reduced products with a high energy potential is necessary, which is carried out at the next, III stage of photosynthesis.

Stage III - electron transport reactions. A chain of carriers with a different redox potential (E n ) forms the so-called electron transport chain (ETC). The redox components of ETC are organized in chloroplasts in the form of three main functional complexes - photosystem I (PSI), photosystem II (PSII), cytochrome b 6 f-complex, which provides a high speed of the electron flow and the possibility of its regulation. As a result of the work of the ETC, highly reduced products are formed: reduced ferredoxin (PD restored) and NADPH, as well as energy-rich ATP molecules, which are used in the dark reactions of CO 2 reduction that make up the IV stage of photosynthesis.

Stage IV - "dark" reactions of absorption and reduction of carbon dioxide. The reactions take place with the formation of carbohydrates, the end products of photosynthesis, in the form of which solar energy is stored, absorbed and converted in the "light" reactions of photosynthesis. The speed of "dark" enzymatic reactions - 10 -2 - 10 4 s.

Thus, the entire course of photosynthesis is carried out with the interaction of three flows - the energy flow, the electron flow and the carbon flow. The conjugation of the three streams requires precise coordination and regulation of their constituent reactions.

Development of the doctrine of photosynthesis

General photosynthesis equation

Chloroplast structure

10. Origin of chloroplasts

11. Ontogeny of chloroplasts

12.

13. Pigments of photosynthetic plants

14. Chlorophyll

15.

16. Phycobilins

17. Carotenoids

18.

19. Groups of carotenoids:

20. Flavonoid pigments

21. Flavonoid pigments:

22. Stages of photosynthesis

23.

24.

25. Light stage of photosynthesis

26.

27. Chlorophyll arousal levels

28. Pigment systems

29. Localization of electron and proton transport reactions in the thylakoid membrane

30. Non-cyclic photosynthetic phosphorylation (Z - scheme, or Govindzhi scheme)

31. Photosynthetic phosphorylation

32. Cyclic photosynthetic phosphorylation

33. Cyclic and non-cyclic transport of electrons in chloroplasts

34.

35. Dark stage of photosynthesis

36. C3 photosynthesis, Calvin cycle

37. General equation of the Calvin cycle

38. C4 photosynthesis (Hatch-Slack-Karpilov path)

39.

40. Crassula type photosynthesis

41. CAM photosynthesis

42.

43.

44. Influence of internal and external factors on photosynthesis

45. Factors affecting photosynthesis

46. ​​Factors affecting photosynthesis

47. Factors affecting photosynthesis

48. Factors affecting photosynthesis

49. Factors affecting photosynthesis

50. Factors affecting photosynthesis

51. Factors affecting photosynthesis

52. Factors affecting photosynthesis

53. Daily course of photosynthesis

54. Conclusion