Tag: Carbon

  • DNA Structure

    What is DNA Structure?

    What do a human, a rose, and a bacterium have in common? Each of these things along with every other organism on Earth contains the molecular instructions for life, called deoxyribonucleic acid or DNA. Encoded within this DNA (Deoxyribonucleic Acid) are the directions for traits as diverse as the color of a person’s eyes, the scent of a rose, and the way in which bacteria infect a lung cell.

    DNA is found in nearly all living cells. However, its exact location within a cell depends on whether that cell possesses a special membrane-bound organelle called a nucleus. Organisms composed of cells that contain nuclei are classified as eukaryotes, whereas organisms composed of cells that lack nuclei are classified as prokaryotes. In eukaryotes, DNA is housed within the nucleus, but in prokaryotes, DNA is located directly within the cellular cytoplasm, as there is no nucleus available.

    But what, exactly, is DNA? In short, DNA is a complex molecule that consists of many components, a portion of which are passed from parent organisms to their offspring during the process of reproduction. Although each organism’s DNA is unique, all DNA is composed of the same nitrogen-based molecules. So how does DNA differ from organism to organism? It is simply the order in which these smaller molecules are arranged that differs among individuals. In turn, this pattern of arrangement ultimately determines each organism’s unique characteristics, thanks to another set of molecules that “read” the pattern and stimulate the chemical and physical processes it calls for.

    DNA is made up of molecules called nucleotides. Each nucleotide contains a phosphate group, a sugar group, and a nitrogen base. The four types of nitrogen bases are adenine (A), thymine (T), guanine (G) and cytosine (C). The order of these bases is what determines DNA’s instructions, or genetic code. Similar to the way the order of letters in the alphabet can be used to form a word, the order of nitrogen bases in a DNA sequence forms genes, which in the language of the cell, tells cells how to make proteins. Another type of nucleic acid, ribonucleic acid, or RNA, translates genetic information from DNA into proteins.

    The entire human genome contains about 3 billion bases and about 20,000 genes. Nucleotides are attached together to form two long strands that spiral to create a structure called a double helix. If you think of the double helix structure as a ladder, the phosphate and sugar molecules would be the sides, while the bases would be the rungs. The bases on one strand pair with the bases on another strand: adenine pairs with thymine, and guanine pairs with cytosine.

    DNA molecules are long so long, in fact, that they can’t fit into cells without the right packaging. To fit inside cells, DNA is coiled tightly to form structures we call chromosomes. Each chromosome contains a single DNA molecule. Humans have 23 pairs of chromosomes, which are found inside the cell’s nucleus.

    Why does a DNA molecule consist of two strands? The primary function of DNA is to store and transmit genetic information. To accomplish this function DNA must have two properties. It must be chemically stable so as to reduce the possibility of damage. DNA must also be capable of copying the information it contains. The two-stranded structure of DNA gives it both of these properties. The nucleotide sequence contains the information found in DNA. The nucleotides connect the two strands through hydrogen bonds. Because each nucleotide has a unique complementary nucleotide, each strand contains all the information required to synthesize a new DNA molecule. The double-stranded structure also makes the molecule more stable.

    DNA is the information molecule. It stores instructions for making other large molecules, called proteins. These instructions are stored inside each of your cells, distributed among 46 long structures called chromosomes. These chromosomes are made up of thousands of shorter segments of DNA, called genes. Each gene stores the directions for making protein fragments, whole proteins, or multiple specific proteins.

    DNA is well-suited to perform this biological function because of its molecular structure, and because of the development of a series of high-performance enzymes that are fine-tuned to interact with this molecular structure in specific ways. The match between DNA structure and the activities of these enzymes is so effective and well-refined that DNA has become, over evolutionary time, the universal information-storage molecule for all forms of life. Nature has yet to find a better solution than DNA for storing, expressing, and passing along instructions for making proteins.

    Alternative DNA structures

    DNA Structure A-DNA B-DNA and Z-DNA
    DNA Structure A-DNA B-DNA and Z-DNA

    DNA (Deoxyribonucleic Acid) exists in many possible conformations that include A-DNA, B-DNA, and Z-DNA forms, although, only B-DNA and Z-DNA have been directly observed in functional organisms. The conformation that DNA adopts depends on the hydration level, DNA sequence, the amount and direction of supercoiling, chemical modifications of the bases, the type and concentration of metal ions, and the presence of polyamines in solution.

    The first published reports of A-DNA X-ray diffraction patterns—and also B-DNA—used analyses based on Patterson transforms that provided only a limited amount of structural information for oriented fibers of DNA. An alternative analysis was then proposed by Wilkins et al., in 1953, for the in vivo B-DNA X-ray diffraction scattering patterns of highly hydrated DNA fibers in terms of squares of Bessel functions. In the same journal, James Watson and Francis Crick presented their molecular modeling analysis of the DNA X-ray diffraction patterns to suggest that the structure was a double-helix.

    Although the B-DNA form is most common under the conditions found in cells, it is not a well-defined conformation but a family of related DNA conformations that occur at the high hydration levels present in living cells. Their corresponding X-ray diffraction and scattering patterns are characteristic of molecular para crystals with a significant degree of disorder.

    Compared to B-DNA, the A-DNA form is a wider right-handed spiral, with a shallow, wide minor groove and a narrower, deeper major groove. The A form occurs under non-physiological conditions in partly dehydrated samples of DNA, while in the cell it may be produced in hybrid pairings of DNA and RNA strands, and in enzyme-DNA complexes. Segments of DNA where the bases have been chemically modified by methylation may undergo a larger change in conformation and adopt the Z form. Here, the strands turn about the helical axis in a left-handed spiral, the opposite of the more common B form. These unusual structures can be recognized by specific Z-DNA binding proteins and may be involved in the regulation of transcription.

  • LNA (Locked Nucleic Acid)

    LNA (Locked Nucleic Acid)

    What is an LNA? A locked nucleic acid (LNA), often referred to as inaccessible RNA, is a modified RNA nucleotide. The ribose moiety of an LNA nucleotide is modified with an extra bridge connecting the 2′ oxygen and 4′ carbon. The bridge “locks” the ribose in the 3′-endo (North) conformation, which is often found in the A-form duplexes. LNA nucleotides can be mixed with DNA or RNA residues in the oligonucleotide whenever desired and hybridize with DNA or RNA according to Watson-Crick base-pairing rules. Such oligomers are synthesized chemically and are commercially available. The locked ribose conformation enhances base stacking and backbone pre-organization. This significantly increases the hybridization properties (melting temperature) of oligonucleotides. LNA was independently synthesized by the group of Jesper Wengel in 1998, soon after the first synthesis by the group of Takeshi Imanishi in 1997. The exclusive rights to the LNA technology were secured in 1997 by Exiqon A/S, a Danish biotech company.

    LNA nucleotides are used to increase the sensitivity and specificity of expression in DNA microarrays, FISH probes, quantitative PCR probes and other molecular biology techniques based on oligonucleotides. For the in situ detection of miRNA, the use of LNA is currently (2005) the only efficient method. A triplet of LNA nucleotides surrounding a single-base mismatch site maximizes LNA probe specificity unless the probe contains the guanine base of G-T mismatch.

    Using LNA-based oligonucleotides therapeutically is an emerging field of biotechnology. The Danish pharmaceutical company Santaris Pharma a/s owns the sole rights to therapeutic uses of LNA technology and is now developing a new, LNA-based, hepatitis C drug called miravirsen, targeting miR-122, which is in Phase II clinical testing as of late 2010.

    Definition of an LNA?

    Locked nucleic acid (LNA) is a nucleic acid analogue containing one or more LNA nucleotide monomers with a bicyclic furanose unit locked in an RNA mimicking sugar conformation. LNA oligonucleotides display unprecedented hybridization affinity toward complementary single-stranded RNA and complementary single- or double-stranded DNA. Structural studies have shown that LNA oligonucleotides induce A-type (RNA-like) duplex conformations. The wide applicability of LNA oligonucleotides for gene silencing and their use for research and diagnostic purposes are documented in a number of recent reports, some of which are described herein.

    LNA Locked Nucleic Acid analogues

    What is an LNA?

    LNA (Locked Nucleic Acids) are synthetic modified nucleic acids where the carbohydrate part of the nucleic acid has been synthetically changed. The modification results in an increased bonding strength between the DNA-bases in a double-helix when one of the DNA-bases has been modified. The overall result is a higher melting point of a DNA double-helix containing LNA-modified nucleic acids and thereby an increased stability. By designing the complementary DNA-strand in a double helix so it consists more or less of LNA-units, it is possible to regulate the rate of transcription – even to block it completely. In this way, it is possible to control the expression of genes and thereby the synthesis of selected proteins. The LNA technology is, therefore, a promising tool in the treatment of diseases which originate from genetic defects.

  • DNA (Deoxyribonucleic Acid)

    DNA (Deoxyribonucleic Acid)

    Deoxyribonucleic Acid (DNA) is a molecule that carries the genetic instructions used in the growth, development, functioning and reproduction of all known living organisms and many viruses. DNA and RNA are nucleic acids; alongside proteins, lipids and complex carbohydrates (polysaccharides), they are one of the four major types of macromolecules that are essential for all known forms of life. Most DNA molecules consist of two biopolymer strands coiled around each other to form a double helix. DNA Structure, History of DNA Research.

    What is DNA?

    We all know that elephants only give birth to little elephants, giraffes to giraffes, dogs to dogs and so on for every type of living creature. But why is this so? The answer lies in a molecule called deoxyribonucleic acid (DNA), which contains the biological instructions that make each species unique. DNA, along with the instructions it contains, is passed from adult organisms to their offspring during reproduction. History of DNA Research.

    What is meaning of DNA?

    DNA stands for deoxyribonucleic acid, sometimes called “the molecule of life,” as almost all organisms have their genetic material codified as DNA. Since each person’s DNA is unique, “DNA typing” is a valuable tool in connecting suspects to crime scenes. You can also use the word less scientifically, as in “it’s just not in my DNA to sit through six hours of meetings.”

    You got your DNA from your parents, we call it ‘hereditary material’ (information that is passed on to the next generation). Nobody else in the world will have DNA the same as you, unless you have an identical twin. Deoxyribonucleic acid is a large molecule in the shape of a double helix. That’s a bit like a ladder that’s been twisted many times.

    The two DNA strands are termed polynucleotides since they are composed of simpler monomer units called nucleotides. Each nucleotide is composed of one of four nitrogen-containing nucleobases either cytosine (C), guanine (G), adenine (A), or thymine (T) and a sugar called deoxyribose and a phosphate group. The nucleotides are joined to one another in a chain by covalent bonds between the sugar of one nucleotide and the phosphate of the next, resulting in an alternating sugar-phosphate backbone. The nitrogenous bases of the two separate polynucleotide strands are bound together (according to base pairing rules (A with T, and C with G) with hydrogen bonds to make double-stranded DNA. The total amount of related DNA base pairs on Earth is estimated at 5.0 x 1037 and weighs 50 billion tonnes. In comparison, the total mass of the biosphere has been estimated to be as much as 4 trillion tons of carbon (TTC).

    DNA stores biological information. The DNA backbone is resistant to cleavage, and both strands of the double-stranded structure store the same biological information. This information is replicated as and when the two strands separate. A large part of DNA (more than 98% for humans) is non-coding, meaning that these sections do not serve as patterns for protein sequences.

    The two strands of DNA run in opposite directions to each other and are thus anti-parallel. Attached to each sugar is one of four types of nucleobases (informally, bases). It is the sequence of these four nucleobases along the backbone that encodes biological information. RNA strands are created using DNA strands as a template in a process called transcription. Under the genetic code, these RNA strands are translated to specify the sequence of amino acids within proteins in a process called translation.

    Within eukaryotic cells, DNA is organized into long structures called chromosomes. During cell division these chromosomes are duplicated in the process of DNA replication, providing each cell its own complete set of chromosomes. Eukaryotic organisms (animals, plants, fungi, and protists) store most of their DNA inside the cell nucleus and some of their DNA in organelles, such as mitochondria or chloroplasts. In contrast, prokaryotes (bacteria and archaea) store their DNA only in the cytoplasm. Within the eukaryotic chromosomes, chromatin proteins such as histones compact and organize DNA. These compact structures guide the interactions between DNA and other proteins, helping control which parts of the DNA are transcribed.

    DNA was first isolated by Friedrich Miescher in 1869. Its molecular structure was identified by James Watson and Francis Crick in 1953, whose model-building efforts were guided by X-ray diffraction data acquired by Raymond Gosling who was a post-graduate student of Rosalind Franklin. DNA is used by researchers as a molecular tool to explore physical laws and theories, such as the ergodic theorem and the theory of elasticity. The unique material properties of DNA have made it an attractive molecule for material scientists and engineers interested in micro- and nano-fabrication. Among notable advances in this field are DNA origami and DNA-based hybrid materials. And also read it DNA Structure, History of DNA Research.

  • Cellular Respiration

    What is Cellular Respiration?

    Some organisms, such as plants, can trap the energy in sunlight through photosynthesis (see Photosynthesis) and store it in the chemical bonds of carbohydrate molecules. The principal carbohydrate formed through photosynthesis is glucose. Other types of organisms, such as animals, fungi, many protozoa, and a large portion of bacteria, are unable to perform this process. Therefore, these organisms must rely on the carbohydrates formed in plants to obtain the energy necessary for their metabolic processes.

    Animals and other organisms obtain the energy available in carbohydrates through the process of cellular respiration. Cells take the carbohydrates into their cytoplasm, and through a complex series of metabolic processes, they break down the carbohydrates and release the energy. The energy is generally not needed immediately; rather, it is used to combine adenosine diphosphate (ADP) with phosphate ions to form adenosine triphosphate (ATP) molecules. The ATP can then be used for processes in the cells that require energy, much as a battery powers a mechanical device.

    During the process of cellular respiration, carbon dioxide is given off. This carbon dioxide can be used by plant cells during photosynthesis to form new carbohydrates. Also in the process of cellular respiration, oxygen gas is required to serve as an acceptor of electrons. This oxygen is identical to the oxygen gas given off during photosynthesis. Thus, there is an interrelationship between the processes of photosynthesis and cellular respiration, namely the entrapment of energy available in sunlight and the provision of the energy for cellular processes in the form of ATP.

    The overall mechanism of cellular respiration involves four processes: glycolysis, in which glucose molecules are broken down to form pyruvic acid molecules; the Krebs cycle, in which pyruvic acid is further broken down and the energy in its molecule is used to form high-energy compounds, such as nicotinamide adenine dinucleotide (NADH); the electron transport system, in which electrons are transported along a series of coenzymes and cytochromes and the energy in the electrons is released; and chemiosmosis, in which the energy given off by electrons pumps protons across a membrane and provides the energy for ATP synthesis. The general chemical equation for cellular respiration is:

    C6H12O6 + 6 O2 → 6 H2O + 6CO2 + energy

    Figure 1. provides an overview of cellular respiration. Glucose is converted to pyruvic acid in the cytoplasm, which is then used to produce acetyl CoA in the mitochondrion. Finally, the Krebs cycle proceeds in the mitochondrion. Electron transport and chemiosmosis result in energy release; ATP synthesis also occurs in the mitochondrion.

    Glycolysis

    Glycolysis is the process in which one glucose molecule is broken down to form two molecules of pyruvic acid (also called pyruvate). The glycolysis process is a multi-step metabolic pathway that occurs in the cytoplasm of animal cells, plant cells, and the cells of microorganisms. At least six enzymes operate in the metabolic pathway.

    In the first and third steps of the pathway, ATP energizes the molecules. Thus, two ATP molecules must be expended in the process. Further along in the process, the six-carbon glucose molecule converts into intermediary compounds and is then split into two three-carbon compounds. The latter undergo additional conversions and eventually form pyruvic acid at the conclusion of the process.

    During the latter stages of glycolysis, four ATP molecules are synthesized using the energy given off during the chemical reactions. Thus, four ATP molecules are synthesized and two ATP molecules are used during glycolysis, for a net gain of two ATP molecules.

    An overview of cellular respiration

    Figure 1. An overview of cellular respiration.

    Another reaction during glycolysis yields enough energy to convert NAD to NADH (plus a hydrogen ion). The reduced coenzyme (NADH) will later be used in the electron transport system, and its energy will be released. During glycolysis, two NADH molecules are produced.

    Because glycolysis does not require oxygen, the process is considered to be anaerobic. For certain anaerobic organisms, such as some bacteria and fermentation yeasts, glycolysis is the sole source of energy.

    Glycolysis is a somewhat inefficient process because much of the cellular energy remains in the two molecules of pyruvic acid that are created. Interestingly, this process is somewhat similar to a reversal of photosynthesis (see Photosynthesis).

    Krebs Cycle

    Following glycolysis, the mechanism of cellular respiration involves another multi-step process—the Krebs cycle, which is also called the citric acid cycle or the tricarboxylic acid cycle. The Krebs cycle uses the two molecules of pyruvic acid formed in glycolysis and yields high-energy molecules of NADH and Flavin adenine dinucleotide (FADH2), as well as some ATP.

    Krebs Cycle
    Krebs Cycle

    The Krebs cycle occurs in the mitochondrion of a cell (see Figure 6). This sausage-shaped organelle possesses inner and outer membranes and, therefore, inner and outer compartments. The inner membrane is folded over itself many times; the folds are called cristae. They are somewhat similar to the thylakoid membranes in chloroplasts (see Photosynthesis). Located along the cristae are the important enzymes necessary for the proton pump and for ATP production.

    Prior to entering the Krebs cycle, the pyruvic acid molecules are altered. Each three-carbon pyruvic acid molecule undergoes conversion to a substance called acetyl-coenzyme A, or Acetyl-CoA. During the process, the pyruvic acid molecule is broken down by an enzyme, one carbon atom is released in the form of carbon dioxide, and the remaining two carbon atoms are combined with a coenzyme called coenzyme A. This combination forms Acetyl-CoA. In the process, electrons and a hydrogen ion are transferred to NAD to form high-energy NADH.

    Acetyl-CoA enters the Krebs cycle by combining with a four-carbon acid called oxaloacetic acid. The combination forms the six-carbon acid called citric acid. Citric acid undergoes a series of enzyme-catalyzed conversions. The conversions, which involve up to ten chemical reactions, are all brought about by enzymes. In many of the steps, high-energy electrons are released to NAD. The NAD molecule also acquires a hydrogen ion and becomes NADH. In one of the steps, FAD serves as the electron acceptor, and it acquires two hydrogen ions to become FADH2. Also, in one of the reactions, enough energy is released to synthesize a molecule of ATP. Because for each glucose molecule there are two pyruvic acid molecules entering the system, two ATP molecules are formed.

    Also during the Krebs cycle, the two carbon atoms of Acetyl-CoA are released, and each forms a carbon dioxide molecule. Thus, for each Acetyl-CoA entering the cycle, two carbon dioxide molecules are formed. Two Acetyl-CoA molecules enter the cycle, and each has two carbon atoms, so four carbon dioxide molecules will form. Add these four molecules to the two carbon dioxide molecules formed in the conversion of pyruvic acid to Acetyl-CoA, and it adds up to six carbon dioxide molecules. These six CO2 molecules are given off as waste gas in the Krebs cycle. They represent the six carbons of glucose that originally entered the process of glycolysis.

    At the end of the Krebs cycle, the final product is oxaloacetic acid. This is identical to the oxaloacetic acid that begins the cycle. Now the molecule is ready to accept another Acetyl-CoA molecule to begin another turn of the cycle. All told, the Krebs cycle forms (per two molecules of pyruvic acid) two ATP molecules, ten NADH molecules, and two FADH2 molecules. The NADH and the FADH2 will be used in the electron transport system.

    Electron Transport System

    The electron transport system occurs in the cristae of the mitochondria, where a series of cytochromes (enzymes) and coenzymes exist. These cytochromes and coenzymes act as carrier molecules and transfer molecules. They accept high-energy electrons and pass the electrons to the next molecule in the system. At key proton-pumping sites, the energy of the electrons transports protons across the membrane into the outer compartment of the mitochondrion.

    Each NADH molecule is highly energetic, which accounts for the transfer of six protons into the outer compartment of the mitochondrion. Each FADH2 molecule accounts for the transfer of four protons. The flow of electrons is similar to that taking place in photosynthesis. Electrons pass from NAD to FAD, to other cytochromes and coenzymes, and eventually they lose much of their energy. In cellular respiration, the final electron acceptor is an oxygen atom. In their energy-depleted condition, the electrons unite with an oxygen atom. The electron-oxygen combination then reacts with two hydrogen ions (protons) to form a water molecule (H2O).

    The role of oxygen in cellular respiration is substantial. As a final electron acceptor, it is responsible for removing electrons from the electron transport system. If oxygen were not available, electrons could not be passed among the coenzymes, the energy in electrons could not be released, the proton pump could not be established, and ATP could not be produced. In humans, breathing is the essential process that brings oxygen into the body for delivery to the cells to participate in cellular respiration.

    Chemiosmosis

    The actual production of ATP in cellular respiration takes place through the process of chemiosmosis (see Cells and Energy). Chemiosmosis involves the pumping of protons through special channels in the membranes of mitochondria from the inner to the outer compartment. The pumping establishes a proton (H+) gradient. After the gradient is established, protons diffuse down the gradient through a transport protein called ATP synthase. The flow of hydrogens catalyzes the pairing of a phosphate with ADP, forming ATP.

    The energy production of cellular respiration is substantial. Most biochemists agree that 36 molecules of ATP can be produced for each glucose molecule during cellular respiration as a result of the Krebs cycle reactions, the electron transport system, and chemiosmosis. Also, two ATP molecules are produced through glycolysis, so the net yield is 38 molecules of ATP. These ATP molecules may then be used in the cell for its needs. However, the ATP molecules cannot be stored for long periods of time, so cellular respiration must constantly continue in order to regenerate the ATP molecules as they are used. Each ATP molecule is capable of releasing 7.3 kilocalories of energy per mole.

    Fermentation

    Fermentation is an anaerobic process in which energy can be released from glucose even though oxygen is not available. Fermentation occurs in yeast cells, and a form of fermentation takes place in bacteria and in the muscle cells of animals.

    In yeast cells (the yeast used for baking bread and producing alcoholic beverages), glucose can be metabolized through cellular respiration as in other cells. When oxygen is lacking, however, glucose is still metabolized to pyruvic acid via glycolysis. The pyruvic acid is converted first to acetaldehyde and then to ethyl alcohol. The net gain of ATP to the yeast cell is two molecules—the two molecules of ATP normally produced in glycolysis.

    Yeasts are able to participate in fermentation because they have the necessary enzyme to convert pyruvic acid to ethyl alcohol. This process is essential because it removes electrons and hydrogen ions from NADH during glycolysis. The effect is to free the NAD so it can participate in future reactions of glycolysis. The net gain to the yeast cell of two ATP molecules permits it to remain alive for some time. However, when the percentage of ethyl alcohol reaches approximately 15 percent, the alcohol kills the yeast cells.

    Yeast is used in both bread and alcohol production. Alcohol fermentation is the process that yields beer, wine, and other spirits. The carbon dioxide given off during fermentation supplements the carbon dioxide given off during the Krebs cycle and causes bread to rise.

    In muscle cells, another form of fermentation takes place. When muscle cells contract too frequently (as in strenuous exercise), they rapidly use up their oxygen supply. As a result, the electron transport system and Krebs cycle slow considerably, and ATP production is slowed. However, muscle cells have the ability to produce a small amount of ATP through glycolysis in the absence of oxygen. The muscle cells convert glucose to pyruvic acid. An enzyme in the muscle cells then converts the pyruvic acid to lactic acid. As in the yeast, this reaction frees up the NAD while providing the cells with two ATP molecules from glycolysis. Eventually, however, the lactic acid buildup causes intense fatigue, and the muscle stops contracting.

  • Photosynthesis

    What is Photosynthesis?

    A great variety of living things on Earth, including all green plants, synthesize their foods from simple molecules, such as carbon dioxide and water. For this process, the organisms require energy, and that energy is derived from sunlight.

    Figure 1. shows the energy relationships in living cells. Light energy is captured in the chloroplast of plant cells and used to synthesize glucose molecules, shown as C6H12O6. In the process, oxygen (O2) is released as a waste product. The glucose and oxygen are then used in the mitochondrion of the plant cell, and the energy is released and used to fuel the synthesis of ATP from ADP and P. In the reaction, CO2 and water are released in the mitochondrion to be reused in photosynthesis in the chloroplast.

    Energy relationships in living cells Cycles
    Energy relationships in living cells Cycles

    Energy relationships in living cells

    Figure 1. Energy relationships in living cells.

    The process of utilizing energy to synthesize carbohydrate molecules is called photosynthesis. Photosynthesis is actually two separate processes. in the first process, energy-rich electrons flow through a series of coenzymes and other molecules. This electron energy is trapped. During the trapping process, adenosine triphosphate (ATP) molecules and molecules of nicotinamide adenine dinucleotide phosphate hydrogen (NADPH) are formed. Both ATP and NADPH are rich in energy. These molecules are used in the second process, where carbon dioxide molecules are bound into carbohydrates too form organic substances such as glucose.

    Chloroplast

    The organelle in which photosynthesis occurs (in the leaves and green stems of plants, for example) is called the chloroplast. Chloroplasts are relatively large organelles, containing a watery, protein-rich fluid called stroma. The stroma contains many small structures composed of membranes that resemble stacks of coins. Each stack is a granum (the plural form is grana). Each membrane in the stack is a thylakoid. Within the thylakoid membranes of the granum, many of the reactions of photosynthesis take place. The thylakoids are somewhat similar to the cristae of mitochondria (see Cellular Respiration).

    Photosystems

    Pigment molecules organized into photosystems capture sunlight in the chloroplast. Photosystems are clusters of light-absorbing pigments with some associated molecules—proton (hydrogen ion) pumps, enzymes, coenzymes, and cytochromes (see Cells and Energy). Each photosystem contains about 200 molecules of a green pigment called chlorophyll and about 50 molecules of another family of pigments called carotenoids. In the reaction center of the photosystem, the energy of sunlight is converted to chemical energy. The center is sometimes called a light-harvesting antenna.

    There are two photosystems within the thylakoid membranes, designated photosystem I and photosystem II. The reaction centers of these photosystems are P700 and P680, respectively. The energy captured in these reaction centers drives chemiosmosis, and the energy of chemiosmosis stimulates ATP production in the chloroplasts.

    Process of Photosynthesis

    The process of photosynthesis is conveniently divided into two parts: the energy-fixing reaction (also called the light reaction) and the carbon-fixing reaction (also called the light-independent reaction or the dark reaction).

    Energy-fixing reaction

    The energy-fixing reaction of photosynthesis begins when light is absorbed in photosystem II in the thylakoid membranes. The energy of the sunlight, captured in the P680 reaction center, causes the electrons from P680’s chlorophyll to move to a higher, unstable energy level. These electrons pass through a series of cytochromes in the nearby electron-transport system.

    After passing through the electron transport system, the energy-rich electrons eventually enter Photosystem-I. Some of the energy of the electron is used to pump protons across the thylakoid membrane, and this pumping sets up the potential for chemiosmosis.

    The spent electrons from P680 enter the P700 reaction center in photosystem I. Sunlight activates the electrons, which receive a second boost out of the chlorophyll molecules. There they reach a high energy level. The electrons progress through a second electron transport system, but this time there is no proton pumping. Rather, the energy reduces NADP. This reduction occurs as two electrons join NADP and energize the molecule. Because NADP acquires two negatively charged electrons, it attracts two positively charged protons to balance the charges. Consequently, the NADP molecule is reduced to NADPH, a molecule that contains much energy.

    Because electrons have flowed out of the P680 reaction center, the chlorophyll molecules are left without a certain number of electrons. Electrons secured from water molecules replace these electrons. Each split water molecule releases two electrons that enter the chlorophyll molecules to replace those lost. The split water molecules also release two protons that enter the cytoplasm near the thylakoid and are available to increase the chemiosmotic gradient.

    The third product of the split water molecules is oxygen. Two oxygen atoms combine with one another to form molecular oxygen (O2), which is given off as the by-product of photosynthesis; it fills the atmosphere and is used by all oxygen-requiring organisms, including plant and animal cells.

    Described above are the noncyclic energy-fixing reactions (see Figure 2). Certain plants and autotrophic prokaryotes are also known to participate in cyclic energy-fixing reactions. These reactions involve only photosystem I and the P700 reaction center. Excited electrons leave the reaction center, pass through coenzymes of the electron transport system, and follow a special pathway back to P700. Each electron powers the proton pump and encourages the transport of a proton across the thylakoid membrane. This process enriches the proton gradient and eventually leads to the generation of ATP.

    The energy-fixing reactions of photosynthesis

    Figure 2. The energy-fixing reactions of photosynthesis.

    ATP production in the energy-fixing reactions of photosynthesis occurs by the process of chemiosmosis (explained in Cells and Energy). Essentially, this process consists of a rush of protons across a membrane (the thylakoid membrane, in this case), accompanied by the synthesis of ATP molecules. Biochemists have calculated that the proton concentration on one side of the thylakoid is 10,000 times that of the opposite side of the membrane.

    In photosynthesis, the protons pass back across the membranes through channels lying alongside sites where enzymes are located. As the protons pass through the channels, the energy of the protons is released to form high-energy ATP bonds. ATP is formed in the energy-fixing reactions along with the NADPH formed in the main reactions. Both ATP and NADPH provide the energy necessary for the synthesis of carbohydrates that occurs in the second major set of events in photosynthesis.

    Carbon-fixing reaction

    Glucose and other carbohydrates are synthesized in the carbon-fixing reaction of photosynthesis, often called the Calvin cycle after Melvin Calvin, who performed much of the biochemical research (see Figure 3). This phase of photosynthesis occurs in the stroma of the plant cell.

    A carbon-fixing reaction or the Calvin cycle
    A carbon-fixing reaction or the Calvin cycle

    Figure 3. A carbon-fixing reaction, also called the Calvin cycle.

    In the carbon-fixing reaction, an essential material is carbon dioxide, which is obtained from the atmosphere. The carbon dioxide is attached to a five-carbon compound called ribulose bisphosphate. Ribulose bisphosphate carboxylase catalyzes this reaction.

    After carbon dioxide has been joined to ribulose bisphosphate, a six-carbon product forms, which immediately breaks into two three-carbon molecules called phosphoglycerate. Each phosphoglycerate molecule converts to another organic compound, but only in the presence of ATP. The ATP used is the ATP synthesized in the energy-fixing reaction. The organic compound formed converts to still another organic compound using the energy present in NADPH. Again, the energy-fixing reaction provides the essential energy. Each of the organic compounds that results consists of three carbon atoms. Eventually, the compounds interact with one another and join to form a single molecule of six-carbon glucose. This process also generates additional molecules of ribulose bisphosphate to participate in further carbon-fixing reactions.

    Glucose can be stored in plants in several ways. In some plants, the glucose molecules are joined to one another to form starch molecules. Potato plants, for example, store starch in tubers (underground stems). In some plants, glucose converts to fructose (fruit sugar), and the energy is stored in this form. In still other plants, fructose combines with glucose to form sucrose, commonly known as table sugar. The energy is stored in carbohydrates in this form. Plant cells obtain energy for their activities from these molecules. Animals use the same forms of glucose by consuming plants and delivering the molecules to their cells.

    All living things on Earth depend in some way on photosynthesis. It is the main mechanism for bringing the energy of sunlight into living systems and making that energy available for the chemical reactions taking place in cells.