§14. Autotrophic nutrition

§14. Autotrophic nutrition

The mechanism of photosynthesis. Light and dark stages. It has now been established that photosynthesis occurs in two stages: light and dark. The light stage is the process of using light to split water; At the same time, oxygen is released and energy-rich compounds are formed. The dark stage includes a group of reactions that use the high-energy products of the light stage to reduce CO2 to simple sugar, i.e. for carbon assimilation. Therefore, the dark stage is also called the synthesis stage. The term “dark stage” only means that light is not directly involved in it. Modern ideas about the mechanism of photosynthesis were formed on the basis of research conducted in the 1930-1950s. Previously, for many years, scientists were misled by a seemingly simple, but incorrect hypothesis, according to which O2 is formed from CO2, and the released carbon reacts with H2O, resulting in the formation of carbohydrates. In the 1930s, when it turned out that some sulfur bacteria do not produce oxygen during photosynthesis, biochemist K. van Niel suggested that the oxygen released during photosynthesis in green plants comes from water. In sulfur bacteria the reaction proceeds as follows:

Instead of O2, these organisms produce sulfur. Van Niel came to the conclusion that all types of photosynthesis can be described by the equation

where X is oxygen in photosynthesis, which occurs with the release of O2, and sulfur in the photosynthesis of sulfur bacteria. Van Niel also suggested that this process involves two stages: a light stage and a synthesis stage. This hypothesis was supported by the discovery of physiologist R. Hill. He discovered that destroyed or partially inactivated cells are capable of carrying out a reaction in the light in which oxygen is released, but CO2 is not reduced (it was called the Hill reaction). In order for this reaction to proceed, it was necessary to add some oxidizing agent capable of attaching electrons or hydrogen atoms given up by the oxygen of the water. One of Hill's reagents is quinone, which, by adding two hydrogen atoms, becomes dihydroquinone. Other Hill reagents contained ferric iron (Fe3+ ion), which, by adding one electron from the oxygen of water, was converted into divalent iron (Fe2+). Thus, it was shown that the transition of hydrogen atoms from oxygen in water to carbon can occur in the form of independent movement of electrons and hydrogen ions. It has now been established that for energy storage it is the transition of electrons from one atom to another that is important, while hydrogen ions can pass into an aqueous solution and, if necessary, be removed from it again. The Hill reaction, in which light energy is used to cause the transfer of electrons from oxygen to an oxidizing agent (electron acceptor), was the first demonstration of the conversion of light energy to chemical energy and a model for the light stage of photosynthesis. The hypothesis that oxygen is continuously supplied from water during photosynthesis was further confirmed in experiments using water labeled with a heavy isotope of oxygen (18O). Since the isotopes of oxygen (common 16O and heavy 18O) have the same chemical properties, plants use H218O in the same way as H216O. It turned out that the released oxygen contained 18O. In another experiment, plants carried out photosynthesis with H216O and C18O2. In this case, the oxygen released at the beginning of the experiment did not contain 18O. In the 1950s, plant physiologist D. Arnon and other researchers proved that photosynthesis includes light and dark stages. Preparations capable of carrying out the entire light stage were obtained from plant cells. Using them, it was possible to establish that in the light, electrons are transferred from water to the photosynthetic oxidizer, which as a result becomes an electron donor for the reduction of carbon dioxide at the next stage of photosynthesis. The electron carrier is nicotinamide adenine dinucleotide phosphate. Its oxidized form is designated NADP+, and its reduced form (formed after the addition of two electrons and a hydrogen ion) is designated NADPH. In NADP+ the nitrogen atom is pentavalent (four bonds and one positive charge), and in NADPHN it is trivalent (three bonds). NADP+ belongs to the so-called. coenzymes. Coenzymes, together with enzymes, carry out many chemical reactions in living systems, but unlike enzymes they change during the reaction. Most of the converted light energy stored in the light stage of photosynthesis is stored during the transfer of electrons from water to NADP+. The resulting NADPHN does not hold electrons as tightly as oxygen in water, and can give them away in the processes of synthesis of organic compounds, spending the accumulated energy on useful chemical work. A significant amount of energy is also stored in another way, namely in the form of ATP (adenosine triphosphate). It is formed by removing water from the inorganic phosphate ion (HPO42-) and the organic phosphate, adenosine diphosphate (ADP), according to the following equation:

ATP is an energy-rich compound, and its formation requires energy from some source. In the reverse reaction, i.e. When ATP is broken down into ADP and phosphate, energy is released. In many cases, ATP gives up its energy to other chemical compounds in a reaction in which hydrogen is replaced by phosphate. In the reaction below, sugar (ROH) is phosphorylated to form sugar phosphate:

Sugar phosphate contains more energy than non-phosphorylated sugar, so its reactivity is higher. ATP and NADPHN, formed (along with O2) in the light stage of photosynthesis, are then used at the stage of synthesis of carbohydrates and other organic compounds from carbon dioxide.

The structure of the photosynthetic apparatus. Light energy is absorbed by pigments (the so-called substances that absorb visible light). All plants that carry out photosynthesis have various forms of the green pigment chlorophyll, and all probably contain carotenoids, which are usually yellow in color. Higher plants contain chlorophyll a (C55H72O5N4Mg) and chlorophyll b (C55H70O6N4Mg), as well as four main carotenoids: b-carotene (C40H56), lutein (C40H55O2), violaxanthin and neoxanthin. This variety of pigments provides a wide spectrum of absorption of visible light, since each of them is “tuned” to its own region of the spectrum. Some algae have approximately the same set of pigments, but many of them have pigments that are somewhat different from those listed in their chemical nature. All these pigments, like the entire photosynthetic apparatus of the green cell, are enclosed in special organelles surrounded by a membrane, the so-called. chloroplasts. The green color of plant cells depends only on the chloroplasts; the remaining elements of the cells do not contain green pigments. The size and shape of chloroplasts vary quite widely. A typical chloroplast is shaped like a slightly curved cucumber measuring approx. 1 µm in diameter and length approx. 4 microns. Large cells of green plants, such as the leaf cells of most terrestrial species, contain many chloroplasts, but small unicellular algae, such as Chlorella pyrenoidosa, have only one chloroplast, occupying most of the cell.

An electron microscope allows you to get acquainted with the very complex structure of chloroplasts. It makes it possible to identify much smaller structures than those visible in a conventional light microscope. In a light microscope, particles smaller than 0.5 microns cannot be distinguished. By 1961, the resolution of electron microscopes made it possible to observe particles that were a thousand times smaller (about 0.5 nm). Using an electron microscope, very thin membrane structures, the so-called, were identified in chloroplasts. thylakoids. These are flat sacs, closed at the edges and collected in stacks called grana; In the photographs, the grains look like stacks of very thin pancakes. Inside the sacs there is a space - the thylakoid cavity, and the thylakoids themselves, collected in grana, are immersed in a gel-like mass of soluble proteins that fills the internal space of the chloroplast and is called the stroma. The stroma also contains smaller and thinner thylakoids that connect individual grana to each other. All thylakoid membranes are composed of approximately equal amounts of proteins and lipids. Regardless of whether they are collected in grana or not, it is in them that the pigments are concentrated and the light stage occurs. The dark stage, as is commonly believed, occurs in the stroma.

Photosystems. Chlorophyll and carotenoids, embedded in the thylakoid membranes of chloroplasts, are assembled into functional units - photosystems, each of which contains approximately 250 pigment molecules. The structure of the photosystem is such that of all these molecules capable of absorbing light, only one specially located chlorophyll a molecule can use its energy in photochemical reactions - it is the reaction center of the photosystem. The remaining pigment molecules, absorbing light, transfer its energy to the reaction center; these light-harvesting molecules are called antenna molecules. There are two types of photosystems. In photosystem I, the specific chlorophyll a molecule, which makes up the reaction center, has an absorption optimum at a light wavelength of 700 nm (designated P700; P - pigment), and in photosystem II - at 680 nm (P680). Typically, both photosystems operate synchronously and (in light) continuously, although photosystem I can operate separately.

Biology [Complete reference book for preparing for the Unified State Exam] Lerner Georgy Isaakovich

2.5.3. Photosynthesis and chemosynthesis

All living things need food and nutrients. When feeding, they use energy stored primarily in organic compounds - proteins, fats, carbohydrates. Heterotrophic organisms, as already mentioned, use food of plant and animal origin, already containing organic compounds. Plants create organic matter through the process of photosynthesis. Research into photosynthesis began in 1630 with the experiments of the Dutchman van Helmont. He proved that plants do not obtain organic matter from the soil, but create it themselves. Joseph Priestley in 1771 proved the “correction” of air with plants. Placed under a glass cover, they absorbed carbon dioxide released by the smoldering splinter. Research has continued and it has now been established that photosynthesis is the process of formation of organic compounds from carbon dioxide (CO 2) and water using light energy and takes place in the chloroplasts of green plants and the green pigments of some photosynthetic bacteria.

Chloroplasts and folds of the cytoplasmic membrane of prokaryotes contain a green pigment - chlorophyll. The chlorophyll molecule is capable of being excited by sunlight and donating its electrons and moving them to higher energy levels. This process can be compared to throwing a ball up. As the ball rises, it stores potential energy; falling, he loses her. The electrons do not fall back, but are picked up by electron carriers (NADP + - nicotinamide diphosphate). In this case, the energy they previously accumulated is partially spent on the formation of ATP. Continuing the comparison with a thrown ball, we can say that the ball, as it falls, heats the surrounding space, and part of the energy of the falling electrons is stored in the form of ATP. The process of photosynthesis is divided into reactions caused by light and reactions associated with carbon fixation. They are called light And dark phases.

"Light phase"- This is the stage in which the light energy absorbed by chlorophyll is converted into electrochemical energy in the electron transport chain. It is carried out in the light, in gran membranes with the participation of transporter proteins and ATP synthetase.

Reactions caused by light occur on the photosynthetic membranes of grana chloroplasts:

1) excitation of chlorophyll electrons by light quanta and their transition to a higher energy level;

2) reduction of electron acceptors – NADP + to NADP H

2Н + + 4е - + NADP + ? NADP N;

3) photolysis of water, occurring with the participation of light quanta: 2H 2 O? 4H + + 4e - + O 2.

This process takes place inside thylakoids– folds of the inner membrane of chloroplasts. Thylakoids form grana - stacks of membranes.

Since exam papers ask not about the mechanisms of photosynthesis, but about the results of this process, we will move on to them.

The results of light reactions are: photolysis of water with the formation of free oxygen, ATP synthesis, reduction of NADP+ to NADP H. Thus, light is needed only for the synthesis of ATP and NADP-H.

"Dark phase"- the process of converting CO 2 into glucose in the stroma (the space between grana) of chloroplasts using the energy of ATP and NADP H.

The result of dark reactions is the conversion of carbon dioxide into glucose and then into starch. In addition to glucose molecules, the formation of amino acids, nucleotides, and alcohols occurs in the stroma.

The overall equation for photosynthesis is -

The meaning of photosynthesis. During the process of photosynthesis, free oxygen is formed, which is necessary for the respiration of organisms:

oxygen forms a protective ozone screen that protects organisms from the harmful effects of ultraviolet radiation;

photosynthesis provides the production of raw organic substances, and therefore food for all living beings;

photosynthesis helps reduce the concentration of carbon dioxide in the atmosphere.

Chemosynthesis – the formation of organic compounds from inorganic ones due to the energy of redox reactions of nitrogen, iron, and sulfur compounds. There are several types of chemosynthetic reactions:

1) oxidation of ammonia to nitrous and nitric acid by nitrifying bacteria:

NH3? HNQ 2? HNO 3 + Q;

2) conversion of ferrous iron into ferric iron by iron bacteria:

Fe 2+ ? Fe 3+ + Q;

3) oxidation of hydrogen sulfide to sulfur or sulfuric acid by sulfur bacteria

H 2 S + O 2 = 2H 2 O + 2S + Q,

H 2 S + O 2 = 2H 2 SO 4 + Q.

The released energy is used for the synthesis of organic substances.

The role of chemosynthesis. Bacteria are chemosynthetics, destroy rocks, purify wastewater, and participate in the formation of minerals.

EXAMPLES OF TASKS

A1. Photosynthesis is a process that occurs in green plants. It is related to:

1) the breakdown of organic substances into inorganic ones

2) the creation of organic substances from inorganic

3) chemical conversion of glucose into starch

4) formation of cellulose

A2. The starting material for photosynthesis is

1) proteins and carbohydrates 3) oxygen and ATP

2) carbon dioxide and water 4) glucose and oxygen

A3. The light phase of photosynthesis occurs

1) in the grana of chloroplasts 3) in the stroma of chloroplasts

2) in leukoplasts 4) in mitochondria

A4. The energy of excited electrons in the light stage is used for:

1) ATP synthesis 3) protein synthesis

2) glucose synthesis 4) carbohydrate breakdown

A5. As a result of photosynthesis, chloroplasts produce:

1) carbon dioxide and oxygen

2) glucose, ATP and oxygen

3) proteins, fats, carbohydrates

4) carbon dioxide, ATP and water

A6. Chemotrophic organisms include

1) pathogens of tuberculosis

2) lactic acid bacteria

3) sulfur bacteria

Part B

IN 1. Select the processes occurring in the light phase of photosynthesis

1) photolysis of water

2) glucose formation

3) synthesis of ATP and NADP H

4) use of CO 2

5) formation of free oxygen

6) use of ATP energy

AT 2. Select the substances involved in the process of photosynthesis

cellulose 4) carbon dioxide

glycogen 5) water

chlorophyll 6) nucleic acids

Part WITH

C1. What conditions are necessary for the process of photosynthesis to begin?

C2. How does the structure of a leaf ensure its photosynthetic functions?

From the book 100 Great Scientific Discoveries author Samin Dmitry

PHOTOSYNTHESIS For several years, the French chemists Peltier (1788–1842) and Cavantou (1795–1877) worked together. This fruitful collaboration led to the discovery of strychnine and brucine. Their greatest fame was brought to them by the discovery of quinine, a surefire remedy against malaria. In 1817 scientists

From the book Great Soviet Encyclopedia (FO) by the author TSB

From the book Great Soviet Encyclopedia (XE) by the author TSB

From the book The Newest Book of Facts. Volume 1 [Astronomy and astrophysics. Geography and other earth sciences. Biology and Medicine] author

What is photosynthesis and what does it mean for life on Earth? Photosynthesis is the formation by higher plants, algae, and photosynthetic bacteria of complex organic substances necessary for the life of both the plants themselves and all other plants.

From the book Biology [Complete reference book for preparing for the Unified State Exam] author Lerner Georgy Isaakovich

2.5. Metabolism: energy and plastic metabolism, their relationship. Enzymes, their chemical nature, role in metabolism. Stages of energy metabolism. Fermentation and respiration. Photosynthesis, its significance, cosmic role. Phases of photosynthesis. Light and dark reactions

From the book The Newest Book of Facts. Volume 1. Astronomy and astrophysics. Geography and other earth sciences. Biology and medicine author Kondrashov Anatoly Pavlovich

Checking work on the topic “Photosynthesis”

Option 1.

Exercise 1.

An object

Function

Photosynthesis

Cell center

Cell division

1) EPS 2) chloroplast 3) ribosome 4) nucleus

Task 2. In the table below, there is a relationship between the positions in the first and second columns.

An object

Function

Glucose

DNA

Nucleotide

What concept should be entered in the blank in this table?

1) amino acid 2) chitin 3) cellulose 4) RNA

Task 3. Insert into the text “Light phase of photosynthesis” the missing terms from the proposed list, using numerical notations. Write down the resulting sequence of numbers.

LIGHT PHASE OF PHOTOSYNTHESIS

It has now been established that photosynthesis occurs in two phases: light and __________ (A). In the light phase, due to solar energy, excitation of molecules __________ (B) and synthesis of molecules __________ (C) occurs. Simultaneously with this reaction, water decomposes under the influence of light, releasing free __________ (G). This process is called photolysis.

LIST OF TERMS:

1) DNA 2) dark 3) oxygen 4) ATP 5) dark 6) hemoglobin

7) chlorophyll 8) carbon dioxide

X the concentration of carbon dioxide is plotted (in %), and along the axis at

Which of the proposed descriptions most accurately reflects this dependence of carbon dioxide concentrations in the range of 0.03 0.16%? Rate of photosynthesis in this interval

    grows smoothly throughout the entire graph

    grows sharply throughout the graph

    increases smoothly at the beginning, and then does not change

Task 5. Study the graph of the dependence of the relative rate of photosynthesis on light intensity (the x-axis shows the relative light intensity in candelas, and the y-axis relative rate of photosynthesis (in arbitrary units)).

Determine at what light intensity, from those listed, the relative rate of photosynthesis will be maximum.


in the chloroplast. Which of the following substances corresponds to number 3 on this diagram?

Test work on the topic “Photosynthesis”

Option 2.

Exercise 1. In the table below, there is a relationship between the positions in the first and second columns.

An object

Function

Mitochondria

Breath

Photosynthesis

What concept should be entered in the blank in this table?

1) Golgi complex 2) chloroplast 3) ribosome 4) nucleus

Task 2. In the table below, there is a relationship between the positions in the first and second columns.

An object

Function

stroma

glucose synthesis

grains

What concept should be entered in the blank in this table?

1) protein synthesis 2) water photolysis 3) lipid synthesis 4) glycolysis

Task 3. Insert into the text “Dark phase of photosynthesis” the missing terms from the proposed list, using numerical notations. Write down the resulting sequence of numbers.

DARK PHASE OF PHOTOSYNTHESIS

It has now been established that photosynthesis occurs in two phases: __________ (A) and dark. For dark phase reactions to occur, the presence of light __________ (B). At this time, the assimilation of __________ (B) from the air occurs, its reduction by hydrogen ions and the formation of organic matter __________ (D) due to the energy accumulated in the light phase.

LIST OF TERMS

1) light 2) carbon dioxide 3) oxygen 4) protein 5) twilight 6) optional

7) glucose 8) required

Task 4. Study the graph of the dependence of the relative rate of photosynthesis on the concentration of carbon dioxide (along the axisat the relative rate of photosynthesis is plotted (in arbitrary units), and along the axis X – concentration of carbon dioxide (in %)).

Determine the concentration of carbon dioxide at which the relative rate of photosynthesis will be 24 conventional units.

    0,08 % 2) 0,05 % 3) 0,03 % 4) 0,01 %

Task 5. Study the graph of photosynthesis rate versus carbon dioxide concentration (x-axisthe concentration of carbon dioxide is plotted (in %), and along the y-axisrelative rate of photosynthesis (in arbitrary units)).

What will be the relative rate of photosynthesis if the concentration of carbon dioxide in the greenhouse air is 0.03%?

Task 6. Study the diagram showing the process of photosynthesis taking place
in the chloroplast. Which of the following substances corresponds to number 2 in this diagram?

3)

oxygen

4)

glucose

PHOTOSYNTHESIS

the formation by living plant cells of organic substances, such as sugars and starch, from inorganic ones - from CO2 and water - using the energy of light absorbed by plant pigments. It is the process of food production on which all living things - plants, animals and humans - depend. All terrestrial plants and most aquatic plants release oxygen during photosynthesis. Some organisms, however, are characterized by other types of photosynthesis that occur without the release of oxygen. The main reaction of photosynthesis, which occurs with the release of oxygen, can be written in the following form:

Organic substances include all carbon compounds with the exception of its oxides and nitrides. The largest quantities of organic substances produced during photosynthesis are carbohydrates (primarily sugars and starch), amino acids (from which proteins are built) and, finally, fatty acids (which, in combination with glycerophosphate, serve as material for the synthesis of fats). Of inorganic substances, the synthesis of all these compounds requires water (H2O) and carbon dioxide (CO2). Amino acids also require nitrogen and sulfur. Plants can absorb these elements in the form of their oxides, nitrate (NO3-) and sulfate (SO42-), or in other, more reduced forms, such as ammonia (NH3) or hydrogen sulfide (hydrogen sulfide H3S). The composition of organic compounds can also include phosphorus during photosynthesis (plants absorb it in the form of phosphate) and metal ions - iron and magnesium. Manganese and some other elements are also necessary for photosynthesis, but only in trace amounts. In terrestrial plants, all these inorganic compounds, with the exception of CO2, enter through the roots. Plants obtain CO2 from atmospheric air, in which its average concentration is 0.03%. CO2 enters the leaves and O2 is released from them through small openings in the epidermis called stomata. The opening and closing of stomata is regulated by special cells - they are called guard cells - also green and capable of carrying out photosynthesis. When light falls on the guard cells, photosynthesis begins in them. The accumulation of its products forces these cells to stretch. In this case, the stomatal opening opens wider, and CO2 penetrates to the underlying layers of the leaf, the cells of which can now continue photosynthesis. Stomata also regulate the evaporation of water by leaves, the so-called. transpiration, since most of the water vapor passes through these openings. Aquatic plants obtain all the nutrients they need from the water in which they live. CO2 and bicarbonate ion (HCO3-) are also found in both sea and fresh water. Algae and other aquatic plants obtain them directly from water. Light in photosynthesis plays the role of not only a catalyst, but also one of the reactants. A significant part of the light energy used by plants during photosynthesis is stored in the form of chemical potential energy in the products of photosynthesis. For photosynthesis, which occurs with the release of oxygen, any visible light from violet (wavelength 400 nm) to medium red (700 nm) is more or less suitable. Some types of bacterial photosynthesis that are not accompanied by the release of O2 can effectively use light with a longer wavelength, up to the far red (900 nm). Clarification of the nature of photosynthesis began at the time of the birth of modern chemistry. The works of J. Priestley (1772), J. Ingenhaus (1780), J. Senebier (1782), as well as the chemical studies of A. Lavoisier (1775, 1781) led to the conclusion that plants convert carbon dioxide into oxygen and for this process it is necessary light. The role of water remained unknown until it was pointed out in 1808 by N. Saussure. In his very precise experiments, he measured the increase in dry weight of a plant growing in a pot of soil, and also determined the amount of carbon dioxide absorbed and oxygen released. Saussure confirmed that all the carbon incorporated into organic matter by a plant comes from carbon dioxide. At the same time, he discovered that the increase in plant dry matter was greater than the difference between the weight of carbon dioxide absorbed and the weight of oxygen released. Since the weight of the soil in the pot did not change significantly, the only possible source of weight gain was water. Thus, it was shown that one of the reactants in photosynthesis is water. The importance of photosynthesis as one of the energy conversion processes could not be appreciated until the very idea of ​​chemical energy arose. In 1845, R. Mayer came to the conclusion that during photosynthesis, light energy is converted into chemical potential energy stored in its products.

PHOTOSYNTHESIS is a process on which all life on Earth depends. It occurs only in plants. During photosynthesis, a plant produces organic substances necessary for all living things from inorganic substances. Carbon dioxide contained in the air enters the leaf through special openings in the epidermis of the leaf, which are called stomata; water and minerals come from the soil to the roots and from there are transported to the leaves through the plant's conducting system. The energy necessary for the synthesis of organic substances from inorganic ones is supplied by the Sun; this energy is absorbed by plant pigments, mainly chlorophyll. In the cell, the synthesis of organic substances occurs in chloroplasts, which contain chlorophyll. Free oxygen, also produced during photosynthesis, is released into the atmosphere.

PHOTOSYNTHESIS SCHEME

The role of photosynthesis. The total result of the chemical reactions of photosynthesis can be described for each of its products by a separate chemical equation. For the simple sugar glucose, the equation is:

The equation shows that in a green plant, due to light energy, one molecule of glucose and six molecules of oxygen are formed from six molecules of water and six molecules of carbon dioxide. Glucose is just one of many carbohydrates synthesized in plants. Below is the general equation for the formation of a carbohydrate with n carbon atoms per molecule:

The equations describing the formation of other organic compounds are not so simple. Amino acid synthesis requires additional inorganic compounds, such as the formation of cysteine:

The role of light as a reactant in the process of photosynthesis is easier to demonstrate if we turn to another chemical reaction, namely combustion. Glucose is one of the subunits of cellulose, the main component of wood. The combustion of glucose is described by the following equation:

This equation is a reversal of the equation for glucose photosynthesis, except that instead of light energy, it produces mostly heat. According to the law of conservation of energy, if energy is released during combustion, then during the reverse reaction, i.e. During photosynthesis, it must be absorbed. The biological analogue of combustion is respiration, so respiration is described by the same equation as non-biological combustion. For all living cells, with the exception of green plant cells in the light, biochemical reactions serve as a source of energy. Respiration is the main biochemical process that releases energy stored during photosynthesis, although long food chains may lie between these two processes. A constant supply of energy is necessary for any manifestation of life, and light energy, which photosynthesis converts into chemical potential energy of organic substances and uses to release free oxygen, is the only important primary source of energy for all living things. Living cells then oxidize ("burn") these organic substances with oxygen, and some of the energy released when oxygen combines with carbon, hydrogen, nitrogen and sulfur is stored for use in various life processes, such as movement or growth. Combining with the listed elements, oxygen forms their oxides - carbon dioxide, water, nitrate and sulfate. Thus the cycle ends. Why is free oxygen, the only source of which on Earth is photosynthesis, so necessary for all living things? The reason is its high reactivity. The electron cloud of a neutral oxygen atom has two fewer electrons than required for the most stable electron configuration. Therefore, oxygen atoms have a strong tendency to acquire two additional electrons, which is achieved by combining (forming two bonds) with other atoms. An oxygen atom can form two bonds with two different atoms or form a double bond with one atom. In each of these bonds, one electron is supplied by an oxygen atom, and the second electron is supplied by another atom participating in the formation of the bond. In a water molecule (H2O), for example, each of the two hydrogen atoms supplies its only electron to form a bond with oxygen, thereby satisfying the inherent desire of oxygen to acquire two additional electrons. In a CO2 molecule, each of the two oxygen atoms forms a double bond with the same carbon atom, which has four bonding electrons. Thus, in both H2O and CO2, the oxygen atom has as many electrons as necessary for a stable configuration. If, however, two oxygen atoms bond to each other, then the electron orbitals of these atoms allow only one bond to form. The need for electrons is thus only half satisfied. Therefore, the O2 molecule, compared to the CO2 and H2O molecules, is less stable and more reactive. Organic products of photosynthesis, such as carbohydrates, (CH2O)n, are quite stable, since each of the carbon, hydrogen and oxygen atoms in them receives as many electrons as necessary to form the most stable configuration. The process of photosynthesis, which produces carbohydrates, therefore converts two very stable substances, CO2 and H2O, into one completely stable substance, (CH2O)n, and one less stable substance, O2. The accumulation of huge amounts of O2 in the atmosphere as a result of photosynthesis and its high reactivity determine its role as a universal oxidizing agent. When an element gives up electrons or hydrogen atoms, we say that the element is oxidized. The addition of electrons or the formation of bonds with hydrogen, as with carbon atoms in photosynthesis, is called reduction. Using these concepts, photosynthesis can be defined as the oxidation of water coupled with the reduction of carbon dioxide or other inorganic oxides.

The mechanism of photosynthesis. Light and dark stages. It has now been established that photosynthesis occurs in two stages: light and dark. The light stage is the process of using light to split water; At the same time, oxygen is released and energy-rich compounds are formed. The dark stage includes a group of reactions that use the high-energy products of the light stage to reduce CO2 to simple sugar, i.e. for carbon assimilation. Therefore, the dark stage is also called the synthesis stage. The term “dark stage” only means that light is not directly involved in it. Modern ideas about the mechanism of photosynthesis were formed on the basis of research conducted in the 1930-1950s. Previously, for many years, scientists were misled by a seemingly simple, but incorrect hypothesis, according to which O2 is formed from CO2, and the released carbon reacts with H2O, resulting in the formation of carbohydrates. In the 1930s, when it turned out that some sulfur bacteria do not produce oxygen during photosynthesis, biochemist K. van Niel suggested that the oxygen released during photosynthesis in green plants comes from water. In sulfur bacteria the reaction proceeds as follows:

Instead of O2, these organisms produce sulfur. Van Niel came to the conclusion that all types of photosynthesis can be described by the equation

where X is oxygen in photosynthesis, which occurs with the release of O2, and sulfur in the photosynthesis of sulfur bacteria. Van Niel also suggested that this process involves two stages: a light stage and a synthesis stage. This hypothesis was supported by the discovery of physiologist R. Hill. He discovered that destroyed or partially inactivated cells are capable of carrying out a reaction in the light in which oxygen is released, but CO2 is not reduced (it was called the Hill reaction). In order for this reaction to proceed, it was necessary to add some oxidizing agent capable of attaching electrons or hydrogen atoms given up by the oxygen of the water. One of Hill's reagents is quinone, which, by adding two hydrogen atoms, becomes dihydroquinone. Other Hill reagents contained ferric iron (Fe3+ ion), which, by adding one electron from the oxygen of water, was converted into divalent iron (Fe2+). Thus, it was shown that the transition of hydrogen atoms from oxygen in water to carbon can occur in the form of independent movement of electrons and hydrogen ions. It has now been established that for energy storage it is the transition of electrons from one atom to another that is important, while hydrogen ions can pass into an aqueous solution and, if necessary, be removed from it again. The Hill reaction, in which light energy is used to cause the transfer of electrons from oxygen to an oxidizing agent (electron acceptor), was the first demonstration of the conversion of light energy to chemical energy and a model for the light stage of photosynthesis. The hypothesis that oxygen is continuously supplied from water during photosynthesis was further confirmed in experiments using water labeled with a heavy isotope of oxygen (18O). Since the isotopes of oxygen (common 16O and heavy 18O) have the same chemical properties, plants use H218O in the same way as H216O. It turned out that the released oxygen contained 18O. In another experiment, plants carried out photosynthesis with H216O and C18O2. In this case, the oxygen released at the beginning of the experiment did not contain 18O. In the 1950s, plant physiologist D. Arnon and other researchers proved that photosynthesis includes light and dark stages. Preparations capable of carrying out the entire light stage were obtained from plant cells. Using them, it was possible to establish that in the light, electrons are transferred from water to the photosynthetic oxidizer, which as a result becomes an electron donor for the reduction of carbon dioxide at the next stage of photosynthesis. The electron carrier is nicotinamide adenine dinucleotide phosphate. Its oxidized form is designated NADP+, and its reduced form (formed after the addition of two electrons and a hydrogen ion) is designated NADPH. In NADP+ the nitrogen atom is pentavalent (four bonds and one positive charge), and in NADPHN it is trivalent (three bonds). NADP+ belongs to the so-called. coenzymes. Coenzymes, together with enzymes, carry out many chemical reactions in living systems, but unlike enzymes they change during the reaction. Most of the converted light energy stored in the light stage of photosynthesis is stored during the transfer of electrons from water to NADP+. The resulting NADPHN does not hold electrons as tightly as oxygen in water, and can give them away in the processes of synthesis of organic compounds, spending the accumulated energy on useful chemical work. A significant amount of energy is also stored in another way, namely in the form of ATP (adenosine triphosphate). It is formed by removing water from the inorganic phosphate ion (HPO42-) and the organic phosphate, adenosine diphosphate (ADP), according to the following equation:

ATP is an energy-rich compound, and its formation requires energy from some source. In the reverse reaction, i.e. When ATP is broken down into ADP and phosphate, energy is released. In many cases, ATP gives up its energy to other chemical compounds in a reaction in which hydrogen is replaced by phosphate. In the reaction below, sugar (ROH) is phosphorylated to form sugar phosphate:

Sugar phosphate contains more energy than non-phosphorylated sugar, so its reactivity is higher. ATP and NADPHN, formed (along with O2) in the light stage of photosynthesis, are then used at the stage of synthesis of carbohydrates and other organic compounds from carbon dioxide.

The structure of the photosynthetic apparatus. Light energy is absorbed by pigments (the so-called substances that absorb visible light). All plants that carry out photosynthesis have various forms of the green pigment chlorophyll, and all probably contain carotenoids, which are usually yellow in color. Higher plants contain chlorophyll a (C55H72O5N4Mg) and chlorophyll b (C55H70O6N4Mg), as well as four main carotenoids: b-carotene (C40H56), lutein (C40H55O2), violaxanthin and neoxanthin. This variety of pigments provides a wide spectrum of absorption of visible light, since each of them is “tuned” to its own region of the spectrum. Some algae have approximately the same set of pigments, but many of them have pigments that are somewhat different from those listed in their chemical nature. All these pigments, like the entire photosynthetic apparatus of the green cell, are enclosed in special organelles surrounded by a membrane, the so-called. chloroplasts. The green color of plant cells depends only on the chloroplasts; the remaining elements of the cells do not contain green pigments. The size and shape of chloroplasts vary quite widely. A typical chloroplast is shaped like a slightly curved cucumber measuring approx. 1 µm in diameter and length approx. 4 microns. Large cells of green plants, such as the leaf cells of most terrestrial species, contain many chloroplasts, but small unicellular algae, such as Chlorella pyrenoidosa, have only one chloroplast, occupying most of the cell.

An electron microscope allows you to get acquainted with the very complex structure of chloroplasts. It makes it possible to identify much smaller structures than those visible in a conventional light microscope. In a light microscope, particles smaller than 0.5 microns cannot be distinguished. By 1961, the resolution of electron microscopes made it possible to observe particles that were a thousand times smaller (about 0.5 nm). Using an electron microscope, very thin membrane structures, the so-called, were identified in chloroplasts. thylakoids. These are flat sacs, closed at the edges and collected in stacks called grana; In the photographs, the grains look like stacks of very thin pancakes. Inside the sacs there is a space - the thylakoid cavity, and the thylakoids themselves, collected in grana, are immersed in a gel-like mass of soluble proteins that fills the internal space of the chloroplast and is called the stroma. The stroma also contains smaller and thinner thylakoids that connect individual grana to each other. All thylakoid membranes are composed of approximately equal amounts of proteins and lipids. Regardless of whether they are collected in grana or not, it is in them that the pigments are concentrated and the light stage occurs. The dark stage, as is commonly believed, occurs in the stroma.

Photosystems. Chlorophyll and carotenoids, embedded in the thylakoid membranes of chloroplasts, are assembled into functional units - photosystems, each of which contains approximately 250 pigment molecules. The structure of the photosystem is such that of all these molecules capable of absorbing light, only one specially located chlorophyll a molecule can use its energy in photochemical reactions - it is the reaction center of the photosystem. The remaining pigment molecules, absorbing light, transfer its energy to the reaction center; these light-harvesting molecules are called antenna molecules. There are two types of photosystems. In photosystem I, the specific chlorophyll a molecule, which makes up the reaction center, has an absorption optimum at a light wavelength of 700 nm (designated P700; P - pigment), and in photosystem II - at 680 nm (P680). Typically, both photosystems operate synchronously and (in light) continuously, although photosystem I can operate separately.

Transformations of light energy. Consideration of this issue should begin with photosystem II, where light energy is utilized by the reaction center P680. When light enters this photosystem, its energy excites the P680 molecule, and a pair of excited, energized electrons belonging to this molecule are detached and transferred to an acceptor molecule (probably quinone), denoted by the letter Q. The situation can be imagined in such a way that the electrons as would jump from the received light “push” and the acceptor catches them in some upper position. If it were not for the acceptor, the electrons would return to their original position (to the reaction center), and the energy released during the downward movement would turn into light, i.e. would be spent on fluorescence. From this point of view, the electron acceptor can be considered as a fluorescence quencher (hence its designation Q, from the English quench - to quench).

The P680 molecule, having lost two electrons, has oxidized, and in order for the process not to stop there, it must be restored, i.e. gain two electrons from some source. Water serves as such a source: it splits into 2H+ and 1/2O2, donating two electrons to oxidized P680. This light-dependent splitting of water is called photolysis. Enzymes that carry out photolysis are located on the inner side of the thylakoid membrane, as a result of which all hydrogen ions accumulate in the thylakoid cavity. The most important cofactor for photolysis enzymes are manganese atoms. The transition of two electrons from the reaction center of the photosystem to the acceptor is an “uphill” climb, i.e. to a higher energy level, and this rise is provided by light energy. Next, in photosystem II, a pair of electrons begins a gradual “descent” from acceptor Q to photosystem I. The descent occurs along an electron transport chain, very similar in organization to the similar chain in mitochondria (see also METABOLISM). It consists of cytochromes, proteins containing iron and sulfur, copper-containing protein and other components. The gradual descent of electrons from a more energized state to a less energized one is associated with the synthesis of ATP from ADP and inorganic phosphate. As a result, light energy is not lost, but is stored in the phosphate bonds of ATP, which can be used in metabolic processes. The production of ATP during photosynthesis is called photophosphorylation. Simultaneously with the described process, light is absorbed in photosystem I. Here, its energy is also used to separate two electrons from the reaction center (P700) and transfer them to an acceptor - an iron-containing protein. From this acceptor, through an intermediate carrier (also a protein containing iron), both electrons go to NADP+, which as a result becomes capable of attaching hydrogen ions (formed during photolysis of water and preserved in thylakoids) - and turns into NADPH. As for the reaction center P700, which was oxidized at the beginning of the process, it accepts two (“descended”) electrons from photosystem II, which returns it to its original state. The total reaction of the light stage occurring during photoactivation of photosystems I and II can be represented as follows:

The total energy output of the electron flow in this case is 1 ATP molecule and 1 NADPH molecule per 2 electrons. By comparing the energy of these compounds with the energy of light that provides their synthesis, it was calculated that approximately 1/3 of the energy of absorbed light is stored in the process of photosynthesis. In some photosynthetic bacteria, photosystem I operates independently. In this case, the flow of electrons moves cyclically from the reaction center to the acceptor and - along a roundabout path - back to the reaction center. In this case, photolysis of water and the release of oxygen do not occur, NADPH is not formed, but ATP is synthesized. This mechanism of light reaction can also occur in higher plants under conditions when an excess of NADPH occurs in the cells.

Dark reactions (synthesis stage). The synthesis of organic compounds by reduction of CO2 (as well as nitrate and sulfate) also occurs in chloroplasts. ATP and NADPH, supplied by the light reaction occurring in thylakoid membranes, serve as a source of energy and electrons for synthesis reactions. The reduction of CO2 is the result of the transfer of electrons to CO2. During this transfer, some of the C-O bonds are replaced by C-H, C-C, and O-H bonds. The process consists of a number of stages, some of which (15 or more) form a cycle. This cycle was discovered in 1953 by the chemist M. Calvin and his colleagues. Using a radioactive isotope of carbon instead of the usual (stable) isotope in their experiments, these researchers were able to trace the path of carbon in the reactions being studied. In 1961, Calvin was awarded the Nobel Prize in Chemistry for this work. The Calvin cycle involves compounds with the number of carbon atoms in molecules from three to seven. All components of the cycle, with the exception of one, are sugar phosphates, i.e. sugars in which one or two OH groups are replaced by a phosphate group (-OPO3H-). An exception is 3-phosphoglyceric acid (PGA; 3-phosphoglycerate), which is a sugar acid phosphate. It is similar to phosphorylated three-carbon sugar (glycerophosphate), but differs from it in that it has a carboxyl group O=C-O-, i.e. one of its carbon atoms is connected to oxygen atoms by three bonds. It is convenient to begin the description of the cycle with ribulose monophosphate, which contains five carbon atoms (C5). ATP formed in the light stage reacts with ribulose monophosphate, converting it into ribulose diphosphate. The second phosphate group gives ribulose diphosphate additional energy, since it carries part of the energy stored in the ATP molecule. Therefore, the tendency to react with other compounds and form new bonds is more pronounced in ribulose diphosphate. It is this C5 sugar that adds CO2 to form a six-carbon compound. The latter is very unstable and under the influence of water breaks down into two fragments - two FHA molecules. If we keep in mind only the change in the number of carbon atoms in sugar molecules, then this main stage of the cycle in which the fixation (assimilation) of CO2 occurs can be represented as follows:

The enzyme that catalyzes CO2 fixation (specific carboxylase) is present in chloroplasts in very large quantities (over 16% of their total protein content); Given the enormous mass of green plants, it is probably the most abundant protein in the biosphere. The next step is that the two molecules of PGA formed in the carboxylation reaction are each reduced by one molecule of NADPH to a three-carbon sugar phosphate (triose phosphate). This reduction occurs as a result of the transfer of two electrons to the carbon of the carboxyl group of FHA. However, in this case, ATP is also needed to provide the molecule with additional chemical energy and increase its reactivity. This task is performed by an enzyme system that transfers the terminal phosphate group of ATP to one of the oxygen atoms of the carboxyl group (a group is formed), i.e. PGA is converted to diphosphoglyceric acid. Once NADPHN donates one hydrogen atom plus an electron to the carbon of the carboxyl group of this compound (equivalent to two electrons plus a hydrogen ion, H+), the C-O single bond is broken and the oxygen bound to the phosphorus is transferred to the inorganic phosphate, HPO42-, and the carboxyl group O =C-O- turns into aldehyde O=C-H. The latter is characteristic of a certain class of sugars. As a result, PGA, with the participation of ATP and NADPH, is reduced to sugar phosphate (triose phosphate). The entire process described above can be represented by the following equations: 1) Ribulose monophosphate + ATP -> Ribulose diphosphate + ADP 2) Ribulose diphosphate + CO2 -> Unstable C6 compound 3) Unstable C6 compound + H2O -> 2 PGA 4) PGA + ATP + NADPH -> ADP + H3PO42- + Triose phosphate (C3). The end result of reactions 1-4 is the formation of two molecules of triose phosphate (C3) from ribulose monophosphate and CO2 with the consumption of two molecules of NADPH and three molecules of ATP. It is in this series of reactions that the entire contribution of the light stage - in the form of ATP and NADPH - to the carbon reduction cycle is represented. Of course, the light stage must additionally supply these cofactors for the reduction of nitrate and sulfate and for the conversion of PGA and triose phosphate formed in the cycle into other organic substances - carbohydrates, proteins and fats. The significance of subsequent stages of the cycle is that they lead to the regeneration of the five-carbon compound, ribulose monophosphate, necessary to restart the cycle. This part of the loop can be written as follows:

which gives a total of 5C3 -> 3C5. Three molecules of ribulose monophosphate, formed from five molecules of triose phosphate, are converted - after the addition of CO2 (carboxylation) and reduction - into six molecules of triose phosphate. Thus, as a result of one revolution of the cycle, one molecule of carbon dioxide is included in the three-carbon organic compound; three revolutions of the cycle in total give a new molecule of the latter, and for the synthesis of a molecule of six-carbon sugar (glucose or fructose), two three-carbon molecules and, accordingly, 6 revolutions of the cycle are required. The cycle gives the increase in organic matter to reactions in which various sugars, fatty acids and amino acids are formed, i.e. "building blocks" of starch, fats and proteins. The fact that the direct products of photosynthesis are not only carbohydrates, but also amino acids, and possibly fatty acids, was also established using an isotope label - a radioactive isotope of carbon. A chloroplast is not just a particle adapted for the synthesis of starch and sugars. This is a very complex, well-organized “factory”, capable of not only producing all the materials from which it itself is built, but also supplying with reduced carbon compounds those parts of the cell and those plant organs that do not carry out photosynthesis themselves.

In chloroplasts. Significance photosynthesis to invigorate the biosphere... chlorophyll is called photosynthesis. Process photosynthesis appears so gloomy... vikorist in the dark phase photosynthesis. Dark phase photosynthesis or the Calvin cycle...

Remember from the textbook “Plants. Bacteria. Fungi and lichens,” what is the essence of photosynthesis. In which cell organelles does it occur? What substances are involved and which are synthesized during photosynthesis? What conditions are necessary for photosynthesis?

Life on Earth depends on autotrophic organisms. Almost all organic substances necessary for living cells are produced through the process of photosynthesis.

Photosynthesis (from the Greek photos - light and synthesis - connection, combination) is the transformation by green plants and photosynthetic microorganisms of inorganic substances (water and carbon dioxide) into organic ones due to solar energy, which is converted into the energy of chemical bonds in the molecules of organic substances.

Rice. 55. J. Priestley (1783-1804) and his experience

History of the discovery and study of photosynthesis. For several centuries, biologists have tried to unravel the mystery of the green leaf. It has long been believed that plants create nutrients from water and minerals.

The discovery of the role of the green leaf belongs not to a biologist, but to a chemist - the English scientist Joseph Priestley (Fig. 55). In 1771, while studying the importance of air for the combustion of substances and respiration, he performed the following experiment. He placed the mouse in a sealed glass vessel and after some time became convinced that it had consumed all the oxygen in the air and died. But if a living plant was placed next to it, the mouse continued to live. Consequently, the air in the vessel remained good. Priestley made an important conclusion: plants improve the air, saturating it with oxygen - making it suitable for breathing. This was the first time the role of green plants was established. Priestley was the first to suggest the role of light in the life of plants.

A great contribution to the study of photosynthesis was made by the Russian scientist K.A. Timiryazev (Fig. 56). He studied the influence of different parts of the sunlight spectrum on the process of photosynthesis and found that photosynthesis is most effective in red rays. Timiryazev proved that by assimilating carbon in the presence of sunlight, the plant converts its energy into the energy of organic substances.

In his work “The Sun, Life and Chlorophyll,” K. A. Timiryazev described in detail and scientifically substantiated his experiments. His laboratory research methods were used by other scientists for subsequent work on photosynthesis. An act of authoritative recognition of the scientist’s scientific merits was the invitation of Kliment Arkadyevich Timiryazev to the Royal Society of London in 1903 to give the famous lecture “The Cosmic Role of Plants.” For his work on photosynthesis, he was elected honorary doctor of a number of Western European universities.

Phases of photosynthesis. During the process of photosynthesis, energy-poor water and carbon dioxide are converted into energy-intensive organic matter - glucose. In this case, solar energy is accumulated in the chemical bonds of this substance. In addition, during the process of photosynthesis, oxygen is released into the atmosphere, which is used by organisms for respiration.

Rice. 56. Kliment Arkadyevich Timiryazev (1843 - 1920)

It has now been established that photosynthesis occurs in two phases - light and dark (Fig. 57).

Rice. 57. General scheme of photosynthesis

Rice. 58. The intensity of photosynthesis in different light spectra

During the light phase, due to solar energy, chlorophyll molecules are excited and ATP is synthesized. Simultaneously with this reaction, water (H20) decomposes under the influence of light, releasing free oxygen (02). This process was called photolysis (from the Greek photos - light and lysis - dissolution). The resulting hydrogen ions bind to a special substance - the hydrogen ion transporter (NADP) and are used in the next phase.

The presence of light is not necessary for tempo phase reactions to occur. The source of energy here is ATP molecules synthesized in the light phase. In the tempo phase, carbon dioxide is absorbed from the air, its reduction with hydrogen ions and the formation of glucose due to the use of ATP energy.

The influence of environmental conditions on photosynthesis. Photosynthesis uses only 1% of the solar energy falling on the leaf. Photosynthesis depends on a number of environmental conditions. Firstly, this process occurs most intensively under the influence of red rays of the solar spectrum (Fig. 58). The intensity of photosynthesis is determined by the amount of oxygen released, which displaces water from the cylinder. The rate of photosynthesis also depends on the degree of illumination of the plant. An increase in daylight hours leads to an increase in the productivity of photosynthesis, i.e., the amount of organic substances produced by the plant.

The meaning of photosynthesis. Photosynthesis products are used:

  • organisms as nutrients, a source of energy and oxygen for life processes;
  • in human food production;
  • as a building material for housing construction, in the production of furniture, etc.

Humanity owes its existence to photosynthesis. All fuel reserves on Earth are products formed as a result of photosynthesis. Using coal and wood, we obtain energy that was stored in organic matter during photosynthesis. At the same time, oxygen is released into the atmosphere. Scientists estimate that without photosynthesis, the entire supply of oxygen would be used up in 3,000 years.

Chemosynthesis. In addition to photosynthesis, there is another known method for obtaining energy and synthesizing organic substances from inorganic ones. Some bacteria are capable of extracting energy by oxidizing various inorganic substances. They do not need light to create organic substances. The process of synthesis of organic substances from inorganic ones, which takes place thanks to the energy of oxidation of inorganic substances, is called chemosynthesis (from the Latin chemistry - chemistry and the Greek synthesis - connection, combination).

Chemosynthesizing bacteria were discovered by Russian scientist S.N. Vinogradsky. Depending on the oxidation of which substance releases energy, chemosynthesizing iron bacteria, sulfur bacteria and azotobacteria are distinguished.

Exercises based on the material covered

  1. Define photosynthesis. What is the significance of this process for life on Earth?
  2. What substances are formed during the light phase of photosynthesis?
  3. Name the main reactions of the tempo phase. What energy is used to synthesize glucose?
  4. What is the main difference between chemosynthesis and photosynthesis?
  5. Explain why, in the process of historical development of the organic world, photosynthetic organisms took a dominant position compared to chemosynthetic ones.


© 2024 globusks.ru - Car repair and maintenance for beginners