why is photosynthesis important essay

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Development of the idea

Overall reaction of photosynthesis.

• Basic products of photosynthesis
• Evolution of the process
• Light intensity and temperature
• Carbon dioxide
• Internal factors
• Energy efficiency of photosynthesis
• Structural features
• Light absorption and energy transfer
• The pathway of electrons
• Evidence of two light reactions
• Photosystems I and II
• Quantum requirements
• The process of photosynthesis: the conversion of light energy to ATP
• Elucidation of the carbon pathway
• Carboxylation
• Isomerization/condensation/dismutation
• Phosphorylation
• Regulation of the cycle
• Products of carbon reduction
• Photorespiration
• Carbon fixation in C 4 plants
• Carbon fixation via crassulacean acid metabolism (CAM)
• Differences in carbon fixation pathways
• The molecular biology of photosynthesis

Why is photosynthesis important?

What is the basic formula for photosynthesis, which organisms can photosynthesize.

photosynthesis

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• Khan Academy - Photosynthesis
• Biology LibreTexts - Photosynthesis
• University of Florida - Institute of Food and Agricultural Sciences - Photosynthesis
• Milne Library - Inanimate Life - Photosynthesis
• National Center for Biotechnology Information - Chloroplasts and Photosynthesis
• Roger Williams University Pressbooks - Introduction to Molecular and Cell Biology - Photosynthesis
• BCcampus Open Publishing - Concepts of Biology – 1st Canadian Edition - Overview of Photosynthesis
• photosynthesis - Children's Encyclopedia (Ages 8-11)
• photosynthesis - Student Encyclopedia (Ages 11 and up)
• Table Of Contents

Photosynthesis is critical for the existence of the vast majority of life on Earth. It is the way in which virtually all energy in the biosphere becomes available to living things. As primary producers, photosynthetic organisms form the base of Earth’s food webs and are consumed directly or indirectly by all higher life-forms. Additionally, almost all the oxygen in the atmosphere is due to the process of photosynthesis. If photosynthesis ceased, there would soon be little food or other organic matter on Earth, most organisms would disappear, and Earth’s atmosphere would eventually become nearly devoid of gaseous oxygen.

The process of photosynthesis is commonly written as: 6CO 2 + 6H 2 O → C 6 H 12 O 6 + 6O 2 . This means that the reactants, six carbon dioxide molecules and six water molecules, are converted by light energy captured by chlorophyll (implied by the arrow) into a sugar molecule and six oxygen molecules, the products. The sugar is used by the organism, and the oxygen is released as a by-product.

The ability to photosynthesize is found in both eukaryotic and prokaryotic organisms. The most well-known examples are plants, as all but a very few parasitic or mycoheterotrophic species contain chlorophyll and produce their own food. Algae are the other dominant group of eukaryotic photosynthetic organisms. All algae, which include massive kelps and microscopic diatoms , are important primary producers.  Cyanobacteria and certain sulfur bacteria are photosynthetic prokaryotes, in whom photosynthesis evolved. No animals are thought to be independently capable of photosynthesis, though the emerald green sea slug can temporarily incorporate algae chloroplasts in its body for food production.

photosynthesis , the process by which green plants and certain other organisms transform light energy into chemical energy . During photosynthesis in green plants, light energy is captured and used to convert water , carbon dioxide , and minerals into oxygen and energy-rich organic compounds .

It would be impossible to overestimate the importance of photosynthesis in the maintenance of life on Earth . If photosynthesis ceased, there would soon be little food or other organic matter on Earth. Most organisms would disappear, and in time Earth’s atmosphere would become nearly devoid of gaseous oxygen. The only organisms able to exist under such conditions would be the chemosynthetic bacteria , which can utilize the chemical energy of certain inorganic compounds and thus are not dependent on the conversion of light energy.

Energy produced by photosynthesis carried out by plants millions of years ago is responsible for the fossil fuels (i.e., coal , oil , and gas ) that power industrial society . In past ages, green plants and small organisms that fed on plants increased faster than they were consumed, and their remains were deposited in Earth’s crust by sedimentation and other geological processes. There, protected from oxidation , these organic remains were slowly converted to fossil fuels. These fuels not only provide much of the energy used in factories, homes, and transportation but also serve as the raw material for plastics and other synthetic products. Unfortunately, modern civilization is using up in a few centuries the excess of photosynthetic production accumulated over millions of years. Consequently, the carbon dioxide that has been removed from the air to make carbohydrates in photosynthesis over millions of years is being returned at an incredibly rapid rate. The carbon dioxide concentration in Earth’s atmosphere is rising the fastest it ever has in Earth’s history, and this phenomenon is expected to have major implications on Earth’s climate .

Requirements for food, materials, and energy in a world where human population is rapidly growing have created a need to increase both the amount of photosynthesis and the efficiency of converting photosynthetic output into products useful to people. One response to those needs—the so-called Green Revolution , begun in the mid-20th century—achieved enormous improvements in agricultural yield through the use of chemical fertilizers , pest and plant- disease control, plant breeding , and mechanized tilling, harvesting, and crop processing. This effort limited severe famines to a few areas of the world despite rapid population growth , but it did not eliminate widespread malnutrition . Moreover, beginning in the early 1990s, the rate at which yields of major crops increased began to decline. This was especially true for rice in Asia. Rising costs associated with sustaining high rates of agricultural production, which required ever-increasing inputs of fertilizers and pesticides and constant development of new plant varieties, also became problematic for farmers in many countries.

A second agricultural revolution , based on plant genetic engineering , was forecast to lead to increases in plant productivity and thereby partially alleviate malnutrition. Since the 1970s, molecular biologists have possessed the means to alter a plant’s genetic material (deoxyribonucleic acid, or DNA ) with the aim of achieving improvements in disease and drought resistance, product yield and quality, frost hardiness, and other desirable properties. However, such traits are inherently complex, and the process of making changes to crop plants through genetic engineering has turned out to be more complicated than anticipated. In the future such genetic engineering may result in improvements in the process of photosynthesis, but by the first decades of the 21st century, it had yet to demonstrate that it could dramatically increase crop yields.

Another intriguing area in the study of photosynthesis has been the discovery that certain animals are able to convert light energy into chemical energy. The emerald green sea slug ( Elysia chlorotica ), for example, acquires genes and chloroplasts from Vaucheria litorea , an alga it consumes, giving it a limited ability to produce chlorophyll . When enough chloroplasts are assimilated , the slug may forgo the ingestion of food. The pea aphid ( Acyrthosiphon pisum ) can harness light to manufacture the energy-rich compound adenosine triphosphate (ATP); this ability has been linked to the aphid’s manufacture of carotenoid pigments.

General characteristics

The study of photosynthesis began in 1771 with observations made by the English clergyman and scientist Joseph Priestley . Priestley had burned a candle in a closed container until the air within the container could no longer support combustion . He then placed a sprig of mint plant in the container and discovered that after several days the mint had produced some substance (later recognized as oxygen) that enabled the confined air to again support combustion. In 1779 the Dutch physician Jan Ingenhousz expanded upon Priestley’s work, showing that the plant had to be exposed to light if the combustible substance (i.e., oxygen) was to be restored. He also demonstrated that this process required the presence of the green tissues of the plant.

In 1782 it was demonstrated that the combustion-supporting gas (oxygen) was formed at the expense of another gas, or “fixed air,” which had been identified the year before as carbon dioxide. Gas-exchange experiments in 1804 showed that the gain in weight of a plant grown in a carefully weighed pot resulted from the uptake of carbon, which came entirely from absorbed carbon dioxide, and water taken up by plant roots; the balance is oxygen, released back to the atmosphere. Almost half a century passed before the concept of chemical energy had developed sufficiently to permit the discovery (in 1845) that light energy from the sun is stored as chemical energy in products formed during photosynthesis.

This equation is merely a summary statement, for the process of photosynthesis actually involves numerous reactions catalyzed by enzymes (organic catalysts ). These reactions occur in two stages: the “light” stage, consisting of photochemical (i.e., light-capturing) reactions; and the “dark” stage, comprising chemical reactions controlled by enzymes . During the first stage, the energy of light is absorbed and used to drive a series of electron transfers, resulting in the synthesis of ATP and the electron-donor-reduced nicotine adenine dinucleotide phosphate (NADPH). During the dark stage, the ATP and NADPH formed in the light-capturing reactions are used to reduce carbon dioxide to organic carbon compounds. This assimilation of inorganic carbon into organic compounds is called carbon fixation.

Van Niel’s proposal was important because the popular (but incorrect) theory had been that oxygen was removed from carbon dioxide (rather than hydrogen from water, releasing oxygen) and that carbon then combined with water to form carbohydrate (rather than the hydrogen from water combining with CO 2 to form CH 2 O).

By 1940 chemists were using heavy isotopes to follow the reactions of photosynthesis. Water marked with an isotope of oxygen ( 18 O) was used in early experiments. Plants that photosynthesized in the presence of water containing H 2 18 O produced oxygen gas containing 18 O; those that photosynthesized in the presence of normal water produced normal oxygen gas. These results provided definitive support for van Niel’s theory that the oxygen gas produced during photosynthesis is derived from water.

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Why Is Photosynthesis Important for All Organisms?

How Does a Plant Convert Light Energy to Chemical Energy?

Photosynthesis is important to living organisms because it is the number one source of oxygen in the atmosphere. Without photosynthesis, the carbon cycle could not occur, oxygen-requiring life would not survive and plants would die. Green plants and trees use photosynthesis to make food from sunlight, carbon dioxide and water in the atmosphere: It is their primary source of energy. The importance of photosynthesis in our life is the oxygen it produces. Without photosynthesis there would be little to no oxygen on the planet.

TL;DR (Too Long; Didn't Read)

Photosynthesis is important for all living organisms because it provides the oxygen needed by most living creatures for survival on the planet.

Reasons Why Photosynthesis Is Important

• It is the number one source of oxygen in the atmosphere.
• It contributes to the carbon cycle between the earth, the oceans, plants and animals.
• It contributes to the symbiotic relationship between plants, humans and animals.
• It directly or indirectly affects most life on Earth.
• It serves as the primary energy process for most trees and plants.

How Photosynthesis Works

Photosynthesis uses light energy from the sun and carbon dioxide and water in the atmosphere to make food for plants, trees, algae and even some bacteria. It releases oxygen as a byproduct. The chlorophyll in these living organisms, which also contributes to their green hues, absorbs the sunlight and combines it with carbon dioxide to convert these compounds into an organic chemical called adenosine triphosphate (ATP). ATP is crucial in the relationship between energy and living things, and is known as the "energy currency for all life."

Importance of Cellular Respiration to Photosynthesis

Cellular respiration allows all living cells to extract energy in the form of ATP from food and offer that energy for the vital processes of life. All living cells in plants, animals and humans take part in cellular respiration in one form or another. Cellular respiration is a three-step process. In step one, the cytoplasm of the cell breaks down glucose in a process called glycolysis, producing two pyruvate molecules from one glucose molecule and releasing a bit of ATP. In the second step, the cell transports the pyruvate molecules into the mitochondria, the energy center of the cells, without using oxygen, This is known as anaerobic respiration. The third step of cellular respiration involves oxygen and is called aerobic respiration, in which the food energy enters an electron transport chain where it produces ATP.

Cellular respiration in plants is essentially the opposite of photosynthesis. Living creatures breathe in oxygen and release carbon dioxide as a byproduct. A plant uses the carbon dioxide exhaled by animals and humans in combination with the sun's energy during cellular respiration to produce the food that it requires. Plants eventually release oxygen back into the atmosphere, resulting in a symbiotic relationship between plants, animals and humans.

Non-Photosynthetic Plants

While most plants use photosynthesis to produce energy, there are some that are non-photosynthetic. Plants that do not use photosynthesis to produce food are usually parasitic, which means they rely on a host for nutrient generation. Examples include Indian pipe ( Monotropa uniflora ) – also known as the ghost or corpse plant – and beechdrops ( Epifagus americana ), which steals nutrients found in beech tree roots. The Indian pipe plant is a ghostly white color because it contains no chlorophyll. Plants in the fungi kingdom – mushrooms, molds and yeasts – rely on their environment for food instead of photosynthesis.

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• University of California Santa Barbara: How Does Photosynthesis Affect Other Organisms?
• Columbia University: The Carbon Cycle and Earth's Climate
• State University of New York Cortland: Non-Photosynthetic Plants
• California State University, Sacramento: Kingdom Fungi

About the Author

As a journalist and editor for several years, Laurie Brenner has covered many topics in her writings, but science is one of her first loves. Her stint as Manager of the California State Mining and Mineral Museum in California's gold country served to deepen her interest in science which she now fulfills by writing for online science websites. Brenner is also a published sci-fi author. She graduated from San Diego's Coleman College in 1972.

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Essay on Photosynthesis

Students are often asked to write an essay on Photosynthesis in their schools and colleges. And if you’re also looking for the same, we have created 100-word, 250-word, and 500-word essays on the topic.

Let’s take a look…

100 Words Essay on Photosynthesis

What is photosynthesis.

Photosynthesis is how plants make their own food using sunlight. It happens in the leaves of plants. Tiny parts inside the leaves, called chloroplasts, use sunlight to turn water and carbon dioxide from the air into sugar and oxygen. The sugar is food for the plant.

The Ingredients

The main things needed for photosynthesis are sunlight, water, and carbon dioxide. Roots soak up water from the soil. Leaves take in carbon dioxide from the air. Then, using sunlight, plants create food and release oxygen.

The Process

In the chloroplasts, sunlight energy is changed into chemical energy. This energy turns water and carbon dioxide into glucose, a type of sugar. Oxygen is made too, which goes into the air for us to breathe.

Why It’s Important

Photosynthesis is vital for life on Earth. It gives us food and oxygen. Without it, there would be no plants, and without plants, animals and people would not survive. It also helps take in carbon dioxide, which is good for the Earth.

250 Words Essay on Photosynthesis

Why is photosynthesis important.

This process is very important because it is the main way plants make food for themselves and for us, too. Without photosynthesis, plants could not grow, and without plants, animals and humans would not have oxygen to breathe or food to eat.

How Photosynthesis Works

Photosynthesis happens in two main stages. In the first stage, the plant captures sunlight with its leaves. The sunlight gives the plant energy to split water inside its leaves into hydrogen and oxygen. The oxygen is released into the air, and the hydrogen is used in the next stage.

In the second stage, the plant mixes the hydrogen with carbon dioxide from the air to make glucose, which is a type of sugar that plants use for energy. This energy helps the plant to grow, make flowers, and produce seeds.

The Cycle of Life

Photosynthesis is a key part of the cycle of life on Earth. By making food and oxygen, plants support life for all creatures. When animals eat plants, they get the energy from the plants, and when animals breathe, they use the oxygen that plants release. It’s a beautiful cycle that keeps the planet alive.

500 Words Essay on Photosynthesis

Photosynthesis is a process used by plants, algae, and some bacteria to turn sunlight, water, and carbon dioxide into food and oxygen. This happens in the green parts of plants, mainly the leaves. The green color comes from chlorophyll, a special substance in the leaves that captures sunlight.

The Ingredients of Photosynthesis

The photosynthesis recipe.

When sunlight hits the leaves, the chlorophyll captures it and starts the food-making process. The energy from the sunlight turns water and carbon dioxide into glucose, a type of sugar that plants use for energy, and oxygen, which is released into the air. This process is like a recipe that plants follow to make their own food.

The Importance of Photosynthesis

Photosynthesis is very important for life on Earth. It gives us oxygen, which we need to breathe. Plants use the glucose they make for growth and to build other important substances like cellulose, which they use to make their cell walls. Without photosynthesis, there would be no food for animals or people, and no oxygen to breathe.

The Benefits to the Environment

Photosynthesis and the food chain.

All living things need energy to survive, and this energy usually comes from food. Plants are at the bottom of the food chain because they can make their own food using photosynthesis. Animals that eat plants get energy from the glucose in the plants. Then, animals that eat other animals get this energy too. So, photosynthesis is the start of the food chain that feeds almost every living thing on Earth.

Photosynthesis in Our Lives

Photosynthesis affects our lives in many ways. It gives us fruits, vegetables, and grains to eat. Trees and plants also give us wood, paper, and other materials. Plus, they provide shade and help make the air fresh and clean.

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Module 6: Metabolic Pathways

Photosynthesis, identify the basic components and steps of photosynthesis.

The processes in all organisms—from bacteria to humans—require energy. To get this energy, many organisms access stored energy by eating, that is, by ingesting other organisms. But where does the stored energy in food originate? All of this energy can be traced back to photosynthesis.

Photosynthesis is essential to all life on earth; both plants and animals depend on it. It is the only biological process that can capture energy that originates in outer space (sunlight) and convert it into chemical compounds (carbohydrates) that every organism uses to power its metabolism. In brief, the energy of sunlight is captured and used to energize electrons, which are then stored in the covalent bonds of sugar molecules. How long lasting and stable are those covalent bonds? The energy extracted today by the burning of coal and petroleum products represents sunlight energy captured and stored by photosynthesis around 300 million years ago.

Figure 1. Photoautotrophs including (a) plants, (b) algae, and (c) cyanobacteria synthesize their organic compounds via photosynthesis using sunlight as an energy source. Cyanobacteria and planktonic algae can grow over enormous areas in water, at times completely covering the surface. In a (d) deep sea vent, chemoautotrophs, such as these (e) thermophilic bacteria, capture energy from inorganic compounds to produce organic compounds. The ecosystem surrounding the vents has a diverse array of animals, such as tubeworms, crustaceans, and octopi that derive energy from the bacteria. (credit a: modification of work by Steve Hillebrand, U.S. Fish and Wildlife Service; credit b: modification of work by “eutrophication&hypoxia”/Flickr; credit c: modification of work by NASA; credit d: University of Washington, NOAA; credit e: modification of work by Mark Amend, West Coast and Polar Regions Undersea Research Center, UAF, NOAA)

Figure 2. The energy stored in carbohydrate molecules from photosynthesis passes through the food chain. The predator that eats these deer receives a portion of the energy that originated in the photosynthetic vegetation that the deer consumed. (credit: modification of work by Steve VanRiper, U.S. Fish and Wildlife Service)

Plants, algae, and a group of bacteria called cyanobacteria are the only organisms capable of performing photosynthesis (Figure 1). Because they use light to manufacture their own food, they are called photoautotrophs (literally, “self-feeders using light”). Other organisms, such as animals, fungi, and most other bacteria, are termed heterotrophs (“other feeders”), because they must rely on the sugars produced by photosynthetic organisms for their energy needs. A third very interesting group of bacteria synthesize sugars, not by using sunlight’s energy, but by extracting energy from inorganic chemical compounds; hence, they are referred to as chemoautotrophs .

The importance of photosynthesis is not just that it can capture sunlight’s energy. A lizard sunning itself on a cold day can use the sun’s energy to warm up. Photosynthesis is vital because it evolved as a way to store the energy in solar radiation (the “photo” part) as high-energy electrons in the carbon-carbon bonds of carbohydrate molecules (the “synthesis” part). Those carbohydrates are the energy source that heterotrophs use to power the synthesis of ATP via respiration. Therefore, photosynthesis powers 99 percent of Earth’s ecosystems. When a top predator, such as a wolf, preys on a deer (Figure 2), the wolf is at the end of an energy path that went from nuclear reactions on the surface of the sun, to light, to photosynthesis, to vegetation, to deer, and finally to wolf.

Learning Objectives

• Identify the reactants and products of photosynthesis
• Describe the visible and electromagnetic spectrums of light as they applies to photosynthesis
• Describe the light-dependent reactions that take place during photosynthesis
• Identify the light-independent reactions in photosynthesis

Photosynthesis is a multi-step process that requires sunlight, carbon dioxide (which is low in energy), and water as substrates (Figure 3). After the process is complete, it releases oxygen and produces glyceraldehyde-3-phosphate (GA3P), simple carbohydrate molecules (which are high in energy) that can subsequently be converted into glucose, sucrose, or any of dozens of other sugar molecules. These sugar molecules contain energy and the energized carbon that all living things need to survive.

Figure 3. Photosynthesis uses solar energy, carbon dioxide, and water to produce energy-storing carbohydrates. Oxygen is generated as a waste product of photosynthesis.

The following is the chemical equation for photosynthesis (Figure 4):

Figure 4. The basic equation for photosynthesis is deceptively simple. In reality, the process takes place in many steps involving intermediate reactants and products. Glucose, the primary energy source in cells, is made from two three-carbon GA3Ps.

Although the equation looks simple, the many steps that take place during photosynthesis are actually quite complex. Before learning the details of how photoautotrophs turn sunlight into food, it is important to become familiar with the structures involved.

In plants, photosynthesis generally takes place in leaves, which consist of several layers of cells. The process of photosynthesis occurs in a middle layer called the  mesophyll . The gas exchange of carbon dioxide and oxygen occurs through small, regulated openings called stomata (singular: stoma), which also play roles in the regulation of gas exchange and water balance. The stomata are typically located on the underside of the leaf, which helps to minimize water loss. Each stoma is flanked by guard cells that regulate the opening and closing of the stomata by swelling or shrinking in response to osmotic changes.

In all autotrophic eukaryotes, photosynthesis takes place inside an organelle called a  chloroplast . For plants, chloroplast-containing cells exist in the mesophyll. Chloroplasts have a double membrane envelope (composed of an outer membrane and an inner membrane). Within the chloroplast are stacked, disc-shaped structures called thylakoids . Embedded in the thylakoid membrane is chlorophyll, a pigment (molecule that absorbs light) responsible for the initial interaction between light and plant material, and numerous proteins that make up the electron transport chain. The thylakoid membrane encloses an internal space called the thylakoid lumen . As shown in Figure 5, a stack of thylakoids is called a granum , and the liquid-filled space surrounding the granum is called stroma or “bed” (not to be confused with stoma or “mouth,” an opening on the leaf epidermis).

Practice Question

Figure 5. Photosynthesis takes place in chloroplasts, which have an outer membrane and an inner membrane. Stacks of thylakoids called grana form a third membrane layer.

On a hot, dry day, plants close their stomata to conserve water. What impact will this have on photosynthesis?

The Two Parts of Photosynthesis

Photosynthesis takes place in two sequential stages: the light-dependent reactions and the light independent-reactions. In the  light-dependent reactions , energy from sunlight is absorbed by chlorophyll and that energy is converted into stored chemical energy. In the light-independent reactions , the chemical energy harvested during the light-dependent reactions drive the assembly of sugar molecules from carbon dioxide. Therefore, although the light-independent reactions do not use light as a reactant, they require the products of the light-dependent reactions to function. In addition, several enzymes of the light-independent reactions are activated by light. The light-dependent reactions utilize certain molecules to temporarily store the energy: These are referred to as energy carriers. The energy carriers that move energy from light-dependent reactions to light-independent reactions can be thought of as “full” because they are rich in energy. After the energy is released, the “empty” energy carriers return to the light-dependent reaction to obtain more energy. Figure 6 illustrates the components inside the chloroplast where the light-dependent and light-independent reactions take place.

Figure 6. Photosynthesis takes place in two stages: light dependent reactions and the Calvin cycle. Light-dependent reactions, which take place in the thylakoid membrane, use light energy to make ATP and NADPH. The Calvin cycle, which takes place in the stroma, uses energy derived from these compounds to make GA3P from CO 2 .

Photosynthesis at the Grocery Store

Figure 7. Foods that humans consume originate from photosynthesis. (credit: Associação Brasileira de Supermercados)

Major grocery stores in the United States are organized into departments, such as dairy, meats, produce, bread, cereals, and so forth. Each aisle (Figure 7) contains hundreds, if not thousands, of different products for customers to buy and consume.

Although there is a large variety, each item links back to photosynthesis. Meats and dairy link because the animals were fed plant-based foods. The breads, cereals, and pastas come largely from starchy grains, which are the seeds of photosynthesis-dependent plants. What about desserts and drinks? All of these products contain sugar—sucrose is a plant product, a disaccharide, a carbohydrate molecule, which is built directly from photosynthesis. Moreover, many items are less obviously derived from plants: for instance, paper goods are generally plant products, and many plastics (abundant as products and packaging) can be derived from algae or from oil, the fossilized remains of photosynthetic organisms. Virtually every spice and flavoring in the spice aisle was produced by a plant as a leaf, root, bark, flower, fruit, or stem. Ultimately, photosynthesis connects to every meal and every food a person consumes.

Spectrums of Light

How can light be used to make food? When a person turns on a lamp, electrical energy becomes light energy. Like all other forms of kinetic energy, light can travel, change form, and be harnessed to do work. In the case of photosynthesis, light energy is converted into chemical energy, which photoautotrophs use to build carbohydrate molecules. However, autotrophs only use a few specific components of sunlight.

What Is Light Energy?

The sun emits an enormous amount of electromagnetic radiation (solar energy). Humans can see only a fraction of this energy, which portion is therefore referred to as “visible light.” The manner in which solar energy travels is described as waves. Scientists can determine the amount of energy of a wave by measuring its wavelength, the distance between consecutive points of a wave. A single wave is measured from two consecutive points, such as from crest to crest or from trough to trough (Figure 8).

Figure 8. The wavelength of a single wave is the distance between two consecutive points of similar position (two crests or two troughs) along the wave.

Visible light constitutes only one of many types of electromagnetic radiation emitted from the sun and other stars. Scientists differentiate the various types of radiant energy from the sun within the electromagnetic spectrum. The electromagnetic spectrum is the range of all possible frequencies of radiation (Figure 9). The difference between wavelengths relates to the amount of energy carried by them.

Figure 9. The sun emits energy in the form of electromagnetic radiation. This radiation exists at different wavelengths, each of which has its own characteristic energy. All electromagnetic radiation, including visible light, is characterized by its wavelength.

Each type of electromagnetic radiation travels at a particular wavelength. The longer the wavelength (or the more stretched out it appears in the diagram), the less energy is carried. Short, tight waves carry the most energy. This may seem illogical, but think of it in terms of a piece of moving a heavy rope. It takes little effort by a person to move a rope in long, wide waves. To make a rope move in short, tight waves, a person would need to apply significantly more energy.

The electromagnetic spectrum (Figure 9) shows several types of electromagnetic radiation originating from the sun, including X-rays and ultraviolet (UV) rays. The higher-energy waves can penetrate tissues and damage cells and DNA, explaining why both X-rays and UV rays can be harmful to living organisms.

Absorption of Light

Light energy initiates the process of photosynthesis when pigments absorb the light. Organic pigments, whether in the human retina or the chloroplast thylakoid, have a narrow range of energy levels that they can absorb. Energy levels lower than those represented by red light are insufficient to raise an orbital electron to a populatable, excited (quantum) state. Energy levels higher than those in blue light will physically tear the molecules apart, called bleaching. So retinal pigments can only “see” (absorb) 700 nm to 400 nm light, which is therefore called visible light. For the same reasons, plants pigment molecules absorb only light in the wavelength range of 700 nm to 400 nm; plant physiologists refer to this range for plants as photosynthetically active radiation.

The visible light seen by humans as white light actually exists in a rainbow of colors. Certain objects, such as a prism or a drop of water, disperse white light to reveal the colors to the human eye. The visible light portion of the electromagnetic spectrum shows the rainbow of colors, with violet and blue having shorter wavelengths, and therefore higher energy. At the other end of the spectrum toward red, the wavelengths are longer and have lower energy (Figure 10).

Figure 10. The colors of visible light do not carry the same amount of energy. Violet has the shortest wavelength and therefore carries the most energy, whereas red has the longest wavelength and carries the least amount of energy. (credit: modification of work by NASA)

Understanding Pigments

Different kinds of pigments exist, and each has evolved to absorb only certain wavelengths (colors) of visible light. Pigments reflect or transmit the wavelengths they cannot absorb, making them appear in the corresponding color.

Chlorophylls and carotenoids are the two major classes of photosynthetic pigments found in plants and algae; each class has multiple types of pigment molecules. There are five major chlorophylls:  a , b , c and d and a related molecule found in prokaryotes called bacteriochlorophyll. Chlorophyll a and chlorophyll b are found in higher plant chloroplasts and will be the focus of the following discussion.

With dozens of different forms, carotenoids are a much larger group of pigments. The carotenoids found in fruit—such as the red of tomato (lycopene), the yellow of corn seeds (zeaxanthin), or the orange of an orange peel (β-carotene)—are used as advertisements to attract seed dispersers. In photosynthesis, carotenoids function as photosynthetic pigments that are very efficient molecules for the disposal of excess energy. When a leaf is exposed to full sun, the light-dependent reactions are required to process an enormous amount of energy; if that energy is not handled properly, it can do significant damage. Therefore, many carotenoids reside in the thylakoid membrane, absorb excess energy, and safely dissipate that energy as heat.

Each type of pigment can be identified by the specific pattern of wavelengths it absorbs from visible light, which is the  absorption spectrum . The graph in Figure 11 shows the absorption spectra for chlorophyll  a , chlorophyll b , and a type of carotenoid pigment called β-carotene (which absorbs blue and green light). Notice how each pigment has a distinct set of peaks and troughs, revealing a highly specific pattern of absorption. Chlorophyll a absorbs wavelengths from either end of the visible spectrum (blue and red), but not green. Because green is reflected or transmitted, chlorophyll appears green. Carotenoids absorb in the short-wavelength blue region, and reflect the longer yellow, red, and orange wavelengths.

Figure 11. (a) Chlorophyll a, (b) chlorophyll b, and (c) β-carotene are hydrophobic organic pigments found in the thylakoid membrane. Chlorophyll a and b, which are identical except for the part indicated in the red box, are responsible for the green color of leaves. β-carotene is responsible for the orange color in carrots. Each pigment has (d) a unique absorbance spectrum.

Figure 12. Plants that commonly grow in the shade have adapted to low levels of light by changing the relative concentrations of their chlorophyll pigments. (credit: Jason Hollinger)

Many photosynthetic organisms have a mixture of pigments; using them, the organism can absorb energy from a wider range of wavelengths. Not all photosynthetic organisms have full access to sunlight. Some organisms grow underwater where light intensity and quality decrease and change with depth. Other organisms grow in competition for light. Plants on the rainforest floor must be able to absorb any bit of light that comes through, because the taller trees absorb most of the sunlight and scatter the remaining solar radiation (Figure 12).

When studying a photosynthetic organism, scientists can determine the types of pigments present by generating absorption spectra. An instrument called a  spectrophotometer can differentiate which wavelengths of light a substance can absorb. Spectrophotometers measure transmitted light and compute from it the absorption. By extracting pigments from leaves and placing these samples into a spectrophotometer, scientists can identify which wavelengths of light an organism can absorb. Additional methods for the identification of plant pigments include various types of chromatography that separate the pigments by their relative affinities to solid and mobile phases.

Light-Dependent Reactions

The overall function of light-dependent reactions is to convert solar energy into chemical energy in the form of NADPH and ATP. This chemical energy supports the light-independent reactions and fuels the assembly of sugar molecules. The light-dependent reactions are depicted in Figure 13. Protein complexes and pigment molecules work together to produce NADPH and ATP.

Figure 13. A photosystem consists of a light-harvesting complex and a reaction center. Pigments in the light-harvesting complex pass light energy to two special chlorophyll a molecules in the reaction center. The light excites an electron from the chlorophyll a pair, which passes to the primary electron acceptor. The excited electron must then be replaced. In (a) photosystem II, the electron comes from the splitting of water, which releases oxygen as a waste product. In (b) photosystem I, the electron comes from the chloroplast electron transport chain discussed below.

The actual step that converts light energy into chemical energy takes place in a multiprotein complex called a  photosystem , two types of which are found embedded in the thylakoid membrane, photosystem II (PSII) and photosystem I (PSI) (Figure 14). The two complexes differ on the basis of what they oxidize (that is, the source of the low-energy electron supply) and what they reduce (the place to which they deliver their energized electrons).

Both photosystems have the same basic structure; a number of antenna proteins to which the chlorophyll molecules are bound surround the reaction center where the photochemistry takes place. Each photosystem is serviced by the light-harvesting complex, which passes energy from sunlight to the reaction center; it consists of multiple antenna proteins that contain a mixture of 300–400 chlorophyll  a and b molecules as well as other pigments like carotenoids. The absorption of a single photon or distinct quantity or “packet” of light by any of the chlorophylls pushes that molecule into an excited state. In short, the light energy has now been captured by biological molecules but is not stored in any useful form yet. The energy is transferred from chlorophyll to chlorophyll until eventually (after about a millionth of a second), it is delivered to the reaction center. Up to this point, only energy has been transferred between molecules, not electrons.

Figure 14. The photosystem II (PSII) reaction center and the photosystem I (PSI).

In the photosystem II (PSII) reaction center, energy from sunlight is used to extract electrons from water. The electrons travel through the chloroplast electron transport chain to photosystem I (PSI), which reduces NADP + to NADPH. The electron transport chain moves protons across the thylakoid membrane into the lumen. At the same time, splitting of water adds protons to the lumen, and reduction of NADPH removes protons from the stroma. The net result is a low pH in the thylakoid lumen, and a high pH in the stroma. ATP synthase uses this electrochemical gradient to make ATP. What is the initial source of electrons for the chloroplast electron transport chain?

• carbon dioxide

The reaction center contains a pair of chlorophyll  a molecules with a special property. Those two chlorophylls can undergo oxidation upon excitation; they can actually give up an electron in a process called a photoact . It is at this step in the reaction center, that light energy is converted into an excited electron. All of the subsequent steps involve getting that electron onto the energy carrier NADPH for delivery to the Calvin cycle where the electron is deposited onto carbon for long-term storage in the form of a carbohydrate. PSII and PSI are two major components of the photosynthetic electron transport chain , which also includes the cytochrome complex . The cytochrome complex, an enzyme composed of two protein complexes, transfers the electrons from the carrier molecule plastoquinone (Pq) to the protein plastocyanin (Pc), thus enabling both the transfer of protons across the thylakoid membrane and the transfer of electrons from PSII to PSI.

The reaction center of PSII (called  P680 ) delivers its high-energy electrons, one at the time, to the primary electron acceptor , and through the electron transport chain (Pq to cytochrome complex to plastocyanine) to PSI. P680’s missing electron is replaced by extracting a low-energy electron from water; thus, water is split and PSII is re-reduced after every photoact. Splitting one H 2 O molecule releases two electrons, two hydrogen atoms, and one atom of oxygen. Splitting two molecules is required to form one molecule of diatomic O 2 gas. About 10 percent of the oxygen is used by mitochondria in the leaf to support oxidative phosphorylation. The remainder escapes to the atmosphere where it is used by aerobic organisms to support respiration.

As electrons move through the proteins that reside between PSII and PSI, they lose energy. That energy is used to move hydrogen atoms from the stromal side of the membrane to the thylakoid lumen. Those hydrogen atoms, plus the ones produced by splitting water, accumulate in the thylakoid lumen and will be used synthesize ATP in a later step. Because the electrons have lost energy prior to their arrival at PSI, they must be re-energized by PSI, hence, another photon is absorbed by the PSI antenna. That energy is relayed to the PSI reaction center (called  P700 ). P700 is oxidized and sends a high-energy electron to NADP + to form NADPH. Thus, PSII captures the energy to create proton gradients to make ATP, and PSI captures the energy to reduce NADP + into NADPH. The two photosystems work in concert, in part, to guarantee that the production of NADPH will roughly equal the production of ATP. Other mechanisms exist to fine tune that ratio to exactly match the chloroplast’s constantly changing energy needs.

Generating an Energy Carrier: ATP

As in the intermembrane space of the mitochondria during cellular respiration, the buildup of hydrogen ions inside the thylakoid lumen creates a concentration gradient. The passive diffusion of hydrogen ions from high concentration (in the thylakoid lumen) to low concentration (in the stroma) is harnessed to create ATP, just as in the electron transport chain of cellular respiration. The ions build up energy because of diffusion and because they all have the same electrical charge, repelling each other.

To release this energy, hydrogen ions will rush through any opening, similar to water jetting through a hole in a dam. In the thylakoid, that opening is a passage through a specialized protein channel called the ATP synthase. The energy released by the hydrogen ion stream allows ATP synthase to attach a third phosphate group to ADP, which forms a molecule of ATP (Figure 14). The flow of hydrogen ions through ATP synthase is called chemiosmosis because the ions move from an area of high to an area of low concentration through a semi-permeable structure.

Light-Independent Reactions

After the energy from the sun is converted into chemical energy and temporarily stored in ATP and NADPH molecules, the cell has the fuel needed to build carbohydrate molecules for long-term energy storage. The products of the light-dependent reactions, ATP and NADPH, have lifespans in the range of millionths of seconds, whereas the products of the light-independent reactions (carbohydrates and other forms of reduced carbon) can survive for hundreds of millions of years. The carbohydrate molecules made will have a backbone of carbon atoms. Where does the carbon come from? It comes from carbon dioxide, the gas that is a waste product of respiration in microbes, fungi, plants, and animals.

In plants, carbon dioxide (CO 2 ) enters the leaves through stomata, where it diffuses over short distances through intercellular spaces until it reaches the mesophyll cells. Once in the mesophyll cells, CO 2 diffuses into the stroma of the chloroplast—the site of light-independent reactions of photosynthesis. These reactions actually have several names associated with them. Another term, the Calvin cycle , is named for the man who discovered it, and because these reactions function as a cycle. Others call it the Calvin-Benson cycle to include the name of another scientist involved in its discovery. The most outdated name is dark reactions, because light is not directly required (Figure 15). However, the term dark reaction can be misleading because it implies incorrectly that the reaction only occurs at night or is independent of light, which is why most scientists and instructors no longer use it.

Figure 15. Light reactions harness energy from the sun to produce chemical bonds, ATP, and NADPH. These energy-carrying molecules are made in the stroma where carbon fixation takes place.

The light-independent reactions of the Calvin cycle can be organized into three basic stages: fixation, reduction, and regeneration.

Stage 1: Fixation

In the stroma, in addition to CO 2 , two other components are present to initiate the light-independent reactions: an enzyme called ribulose bisphosphate carboxylase (RuBisCO), and three molecules of ribulose bisphosphate (RuBP), as shown in Figure 16. RuBP has five atoms of carbon, flanked by two phosphates.

Figure 16. The Calvin cycle has three stages.

In stage 1, the enzyme RuBisCO incorporates carbon dioxide into an organic molecule, 3-PGA. In stage 2, the organic molecule is reduced using electrons supplied by NADPH. In stage 3, RuBP, the molecule that starts the cycle, is regenerated so that the cycle can continue. Only one carbon dioxide molecule is incorporated at a time, so the cycle must be completed three times to produce a single three-carbon GA3P molecule, and six times to produce a six-carbon glucose molecule.

Which of the following statements is true?

• In photosynthesis, oxygen, carbon dioxide, ATP, and NADPH are reactants. GA3P and water are products.
• In photosynthesis, chlorophyll, water, and carbon dioxide are reactants. GA3P and oxygen are products.
• In photosynthesis, water, carbon dioxide, ATP, and NADPH are reactants. RuBP and oxygen are products.
• In photosynthesis, water and carbon dioxide are reactants. GA3P and oxygen are products.

RuBisCO catalyzes a reaction between CO 2 and RuBP. For each CO 2 molecule that reacts with one RuBP, two molecules of another compound (3-PGA) form. PGA has three carbons and one phosphate. Each turn of the cycle involves only one RuBP and one carbon dioxide and forms two molecules of 3-PGA. The number of carbon atoms remains the same, as the atoms move to form new bonds during the reactions (3 atoms from 3CO 2 + 15 atoms from 3RuBP = 18 atoms in 3 atoms of 3-PGA). This process is called  carbon fixation , because CO 2 is “fixed” from an inorganic form into organic molecules.

Stage 2: Reduction

ATP and NADPH are used to convert the six molecules of 3-PGA into six molecules of a chemical called glyceraldehyde 3-phosphate (G3P). That is a reduction reaction because it involves the gain of electrons by 3-PGA. Recall that a  reduction is the gain of an electron by an atom or molecule. Six molecules of both ATP and NADPH are used. For ATP, energy is released with the loss of the terminal phosphate atom, converting it into ADP; for NADPH, both energy and a hydrogen atom are lost, converting it into NADP + . Both of these molecules return to the nearby light-dependent reactions to be reused and reenergized.

Stage 3: Regeneration

Interestingly, at this point, only one of the G3P molecules leaves the Calvin cycle and is sent to the cytoplasm to contribute to the formation of other compounds needed by the plant. Because the G3P exported from the chloroplast has three carbon atoms, it takes three “turns” of the Calvin cycle to fix enough net carbon to export one G3P. But each turn makes two G3Ps, thus three turns make six G3Ps. One is exported while the remaining five G3P molecules remain in the cycle and are used to regenerate RuBP, which enables the system to prepare for more CO 2 to be fixed. Three more molecules of ATP are used in these regeneration reactions.

Evolution of Photosynthesis

Figure 17. The harsh conditions of the desert have led plants like these cacti to evolve variations of the light-independent reactions of photosynthesis. These variations increase the efficiency of water usage, helping to conserve water and energy. (credit: Piotr Wojtkowski)

During the evolution of photosynthesis, a major shift occurred from the bacterial type of photosynthesis that involves only one photosystem and is typically anoxygenic (does not generate oxygen) into modern oxygenic (does generate oxygen) photosynthesis, employing two photosystems. This modern oxygenic photosynthesis is used by many organisms—from giant tropical leaves in the rainforest to tiny cyanobacterial cells—and the process and components of this photosynthesis remain largely the same. Photosystems absorb light and use electron transport chains to convert energy into the chemical energy of ATP and NADH. The subsequent light-independent reactions then assemble carbohydrate molecules with this energy.

Photosynthesis in desert plants has evolved adaptations that conserve water. In the harsh dry heat, every drop of water must be used to survive. Because stomata must open to allow for the uptake of CO 2 , water escapes from the leaf during active photosynthesis. Desert plants have evolved processes to conserve water and deal with harsh conditions. A more efficient use of CO 2 allows plants to adapt to living with less water. Some plants such as cacti (Figure 17) can prepare materials for photosynthesis during the night by a temporary carbon fixation/storage process, because opening the stomata at this time conserves water due to cooler temperatures. In addition, cacti have evolved the ability to carry out low levels of photosynthesis without opening stomata at all, a mechanism to face extremely dry periods.

Now that we’ve learned about the different pieces of photosynthesis, let’s put it all together. This video walks you through the process of photosynthesis as a whole:

In Summary: An Overview of Photosynthesis

The process of photosynthesis transformed life on Earth. By harnessing energy from the sun, photosynthesis evolved to allow living things access to enormous amounts of energy. Because of photosynthesis, living things gained access to sufficient energy that allowed them to build new structures and achieve the biodiversity evident today.

Only certain organisms, called photoautotrophs, can perform photosynthesis; they require the presence of chlorophyll, a specialized pigment that absorbs certain portions of the visible spectrum and can capture energy from sunlight. Photosynthesis uses carbon dioxide and water to assemble carbohydrate molecules and release oxygen as a waste product into the atmosphere. Eukaryotic autotrophs, such as plants and algae, have organelles called chloroplasts in which photosynthesis takes place, and starch accumulates. In prokaryotes, such as cyanobacteria, the process is less localized and occurs within folded membranes, extensions of the plasma membrane, and in the cytoplasm.

The pigments of the first part of photosynthesis, the light-dependent reactions, absorb energy from sunlight. A photon strikes the antenna pigments of photosystem II to initiate photosynthesis. The energy travels to the reaction center that contains chlorophyll  a to the electron transport chain, which pumps hydrogen ions into the thylakoid interior. This action builds up a high concentration of ions. The ions flow through ATP synthase via chemiosmosis to form molecules of ATP, which are used for the formation of sugar molecules in the second stage of photosynthesis. Photosystem I absorbs a second photon, which results in the formation of an NADPH molecule, another energy and reducing power carrier for the light-independent reactions.

Check Your Understanding

Answer the question(s) below to see how well you understand the topics covered in the previous section. This short quiz does  not  count toward your grade in the class, and you can retake it an unlimited number of times.

Use this quiz to check your understanding and decide whether to (1) study the previous section further or (2) move on to the next section.

• Authored by : Shelli Carter and Lumen Learning. Provided by : Lumen Learning. License : CC BY: Attribution
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• Photosynthesis: Crash Course Biology #8. Authored by : CrashCourse. Located at : https://youtu.be/sQK3Yr4Sc_k . License : All Rights Reserved . License Terms : Standard YouTube License

Photosynthesis

Plants are autotrophs, which means they produce their own food. They use the process of photosynthesis to transform water, sunlight, and carbon dioxide into oxygen, and simple sugars that the plant uses as fuel. These primary producers form the base of an ecosystem and fuel the next trophic levels. Without this process, life on Earth as we know it would not be possible. We depend on plants for oxygen production and food.

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Photosynthesis: Essay on Photosynthesis (2098 Words)

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Here is your essay on Photosynthesis!

[I] Photosynthesis:

Photosynthesis is one of the most fundamental biological reactions.

The chlorophyll bearing plants trap the free energy of sunlight as photons and transform and store it as chemical potential energy by combining CO 2 and water.

The end products of photosynthesis are carbohydrates with loss of oxygen. These directly or indirectly serve as the source of energy for all living beings, except chemosynthetic bacteria.

Image Courtesy : co2crc.com.au/Photosynthesis_media.jpg

[II] Food storage :

Some unpigmented plastids like leucoplasts store the essential food materials like protein, oil and starch. Later on these are used during germination of seeds and development.

[III] Hereditary carrier :

Recent studies show that these plastids, like chromosomes, are transmitted directly to the daughter cells during cell division. Cytoplasmic inheritance of plastids in Mirabilis is the well-known example. They produce phenotypic effects in Oenothera and other plants.

[IV] Chloroplasts as semiautonomous organoid :

The chloroplast matrix contains dissolved salts and enzymes of photo­synthesis. Besides these, like mitochondria, it contains RNA, DNA and ribosomes, and is capable of carrying on protein synthesis.

The chloroplast ribosomes are of the same size as ribosomes in prokaryotes. Chloroplasts are also semi-autonomous like mitochondria. They can grow and divide, and their DNA contains a portion of the genetic information needed for the synthesis of chloroplast proteins.

[V] Inheritance of chloroplasts :

Cells have the capacity to outgrow their chloroplasts and the rate of multiplication of chloroplasts is partly independent of the rate of multiplication of entire cells. Brawerman and Chargaff (1960) discovered it in Euglena gracilis after a temperature shock.

Cells which were permitted to multiply rapidly became irreversibly bleached, whereas cells prevented from dividing regained their normal ability to produce chloroplasts. They concluded that Euglena contains an autonomously replicating factor which is necessary for chloroplast formation.

[VI] DNA in chloroplasts :

Chloroplasts contain both DNA and the necessary mechanism for synthesizing specific RNA’s and proteins from a DNA template. DNA is found in chloroplasts (Stocking and Gifford, 1959). Ris and Plaut (1962) have also found DNA in the chloroplasts of alga Chlamydomonas. It has now been generally accepted that characteristic chloroplast DNA’s or chloroplast chromosomes occur in the photosynthetic organelles of algae and higher plants.

According to Brawerman (1966) this DNA differs from nuclear DNA in GC (guanosine and cytosine) content. Chloroplasts also contain a DNA-dependent RNA polymerase; it appears that specific RNA’s are synthesized from chloroplast DNA as a template (Kirk 1966). Chloroplasts DNA are capable of self-duplication.

[VII] Chloroplast ribosomes :

Lyttleton (1962) isolated chloroplast ribosomes, which are estimated to make up 3 to 7% of the chloroplast dry mass. Chloroplast ribosomes are smaller than cytoplasmic ribosomes. These are 60-66S. Chloroplast ribosomes also dissociate reversibly into 50S and 35S subunits, in a way that is found in E. coli ribosomes (Boardman et al., 1966). Chloroplasts have three types of RNA required for protein synthesis: ribosomal, transfer and messenger. Chloroplast ribosomes associate to form polysomes for synthesis of proteins (Gunning and Steer, 1975).

[VIII] Protein synthesis :

Protein synthesis in mitochondria and chloroplasts is similar to that of prokaryotes. For example, the size of chloroplast ribosomes is the same as ribosomes of blue-green algae, and ribosomes of chloroplasts and mitochondria more closely resemble prokaryotic ribosomes in antibiotic sensitivity than they do eukaryotic ribosomes.

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Why is Photosynthesis Important?

How it works

“Photosynthesis is an important process which allows organisms to produce food without the intervention of any other organisms. This specialized reaction, occurring only in autotrophs, has several key features that are fundamental for understanding the process as a whole. Factors of where the process takes place, the reactants and products of the reactions, and what happens in the process are all important elements in the success of photosynthesis.

Photosynthesis occurs in the chloroplast, an organelle found only in plant cells. Contained in the chloroplast are multiple structures where different element of photosynthesis take place.

One such structure are thylakoids; flattened membrane sacs that store pigments like chlorophyll. This is where the light-dependent reactions occur. Thylakoids arrange in stacks called grana. Grana are surrounded by stroma, a fluid filling the chloroplast similar to the cytoplasm in cells. In the stroma is where light-independent reactions take place. Through these individual structures, photosynthesis is able to take place.

Photosynthesis is a biochemical reaction, so there are specific reactants and products to enter into and exit from the process each time the cycle takes place. For the reaction to occur, there must be carbon dioxide, CO2, and water, H2O. Produced from this are oxygen, O2, and high energy sugars, C6H12O6. However, for the equation to be balanced with both reactants and products, there must be 6 carbon dioxide reactant molecules and 6 water reactant molecules, or, 6CO2 + 6H2O. There will also be 6 oxygen molecules produced, or 6O2. This equation written in full looks like carbon dioxide + water + light †’ high energy sugars + oxygen, or in numerical form, 6CO2 + 6H2O + light †’ 62O + C6H12O6+602. Overall, the equation of photosynthesis shows the what enters the reaction and what is produced.

While photosynthesis is often viewed as having a single step, the process actually has two parts; the light-dependent reactions and the light-independent reactions. Each process has its own role in making all of the products of this reaction. The light-dependent reactions, occuring in the thylakoids, need sunlight to take place. They transform water into oxygen and use the energy of sunlight to convert ADP and NADP+ into ATP and NADPH which carry energy. On the other hand, light-independent reactions, also known as the Calvin Cycle, occur in the stroma and do not need sunlight. They use the energy in ATP and NADPH made during the light-dependent reactions to convert carbon dioxide from the environment into high energy sugars. The energy is then turned into ADP and NADP+ which then cycle back to the thylakoids for the next set of light-dependent reactions. It is through both of these processes working simultaneously that plants can continuously make food.

In conclusion, each of the individual elements of photosynthesis, including the location of the process, the ingredients of the reaction, and the parts of the reaction have a key role in this function. Through these many pieces, autotrophs are able to thrive. With an awareness of these elements, one has a better understanding of the process of photosynthesis.

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Understanding Global Change

Discover why the climate and environment changes, your place in the Earth system, and paths to a resilient future.

Photosynthesis

Photosynthesis is the processes of using sunlight to convert chemical compounds (specifically carbon dioxide and water ) into food . Photosynthesizing organisms (plants, algae, and bacteria) provide most of the chemical energy that flows through the biosphere.  They also produced most of the biomass that led to the fossil fuels that power much of our modern world. Photosynthesis takes place on land, in the ocean, and in freshwater environments. The first photosynthesizing single-celled bacteria evolved over 3.5 billion years ago. The subsequent rise in atmospheric oxygen (a byproduct of photosynthesis) about a billion years later played a major role in shaping the evolution of life on Earth over the last 2.5 billion years. Today the vast majority of land, freshwater, and oceanic organisms require oxygen for respiration , the biochemical process that generates energy from food.

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Global Change Infographic

Photosynthesus is an essential part of How the Earth System Works.  Click the image on the left to open the Understanding Global Change Infographic . Locate the photosynthesis icon and identify other Earth system processes and phenomena that cause changes to, or are affected by, photosynthesis.

Photosynthesis is the chemical process by which plants, algae, and some bacteria use the energy from sunlight to transform carbon dioxide (a greenhouse gas ) from the atmosphere, and water , into organic compounds such as sugars. These sugars are then used to make complex carbohydrates, lipids, and proteins, as well as the wood, leaves, and roots of plants.  The amount of organic matter made by photosynthesizing organisms in an ecosystem is defined as the productivity of that ecosystem.  Energy flows through the biosphere as organisms (including some animals) eat photosynthesizing organisms (called herbivores), and as organisms then eat those herbivores (carnivores) , etc., to get their energy for growth, reproduction, and other functions.  This energy is acquired through the process of cellular respiration , which usually requires oxygen.   Oxygen is a byproduct of photosynthesis. About 70% of the oxygen in the atmosphere that we breathe comes from algae in the ocean. Atmospheric oxygen from photosynthesis also forms the ozone layer , which protects organisms from harmful high-energy ultraviolet (UV) radiation from the Sun . Because photosynthesis also requires water , the availability of water affects the productivity and biomass of the ecosystem, which in turn affects how much and how rapidly water cycles through the ecosystem.

Fossil fuels are derived from the burial of photosynthetic organisms, including plants on land (which primarily form coal) and plankton in the oceans (which primarily form oil and natural gas). While buried, the carbon in the organic material is removed from the carbon cycle for thousands of years to hundreds of millions of years. The burning of fossil fuels has dramatically increased the exchange of carbon from the ground back into the atmosphere and oceans. This return of carbon back into atmosphere as carbon dioxide is occurring at a rate that is hundreds to thousands of times faster than it took to bury it, and much faster than it can be removed by photosynthesis or weathering . Thus, the carbon dioxide released from the burning of fossil fuels is accumulating in the atmosphere, increasing average temperatures and causing ocean acidification .

A simplified diagram showing the overall inputs – carbon dioxide, water, and sunlight, and products – oxygen and sugar (glucose), of photosynthesis.

The rate of photosynthesis in ecosystems is affected by various environmental conditions, including:

• Climatic conditions, such as the amount of sunlight available at different latitudes , temperature , and precipitation For example, ecosystems at low latitudes, such as tropical rainforests, have higher productivity and biomass than ecosystems near the poles because of they receive more sunlight and rainfall than regions at higher latitudes.
• Nutrients , especially nitrogen and phosphorus , which when limited can decrease productivity, but when abundant can increase productivity and biomass. Photosynthesizing organisms extract nutrients from the environment, and return them to the soil when they die and decay.
• Numerous other abiotic environmental factors, including soil quality (often related to nutrient levels), wildfires , water acidity , and oxygen levels .
• Species interactions , including the resources species provide for each other, and how they compete for resources such as water, light, and/or space. Species that reduce or increase the success of other species alter population sizes , thus affecting productivity and biomass .
• Evolutionary processes that can change the growth and reproduction rates of photosynthesizing organisms over time, as well as the growth and reproduction of rates of the organisms that eat them.

Humans have altered the rate of photosynthesis, and in turn productivity , in ecosystems through a variety of activities, including:

• Deforestation , habitat destruction , and urbanization , which remove plants and trees from the environment and disrupt ecosystems.
• Agricultural activities that increase the amount of crops available to feed the growing global human population .
• The use of fertilizers for agricultural activities that increase the amount of nutrients , especially nitrogen and phosphorous , in soil or water. These nutrients increase plant and algae growth, including growth of species that are toxic to other organisms. Increased nutrients is not always a good thing. For example, in aquatic environments, nutrient-rich runoff can cause large amounts of algae to grow – when these algae die, they are consumed by bacteria which can reduce oxygen levels in the water, killing fish and other species. This process is known as eutrophication.
• Human freshwater use , which can limit the amount of water available for plants and trees in an ecosystem.
• The release of pollutants and waste , which can reduce growth and reproduction or kill plants.
• Activities that release carbon dioxide and other greenhouse gases that cause global warming, such as the burning of fossil fuels , agricultural activities , and deforestation . Increasing carbon dioxide levels may increase photosynthesis rates in some plants, but this can also make plants less nutritious . Increasing average global land and ocean temperatures and changes in precipitation patterns also affect plant and algae growth, and can make certain species more susceptible to disease .
• Activities such as the burning of fossil fuels , agricultural activities , and deforestation that release carbon dioxide into the atmosphere, which is absorbed by the ocean causing acidification . The decreasing pH of ocean waters (along with ocean warming) causes physiological stress for many plant and algae species, which can decrease growth, reproduction, species population sizes, and biomass .
• The increase in carbon dioxide to the atmosphere is also thought to impact global photosynthesis rates. During photosynthesis, plants convert carbon dioxide to biomass such as sugars and wood. However, the same enzyme (rubisco) that fixes carbon dioxide can also use oxygen. When oxygen is used, plants undergo a process known as photorespiration where biomass is not produced and instead carbon dioxide is emitted to the atmosphere (see this teaching resource for more information). Photorespiration is often considered a negative process for plants. It has been proposed that as carbon dioxide levels rise in the atmosphere rates of photorespiration will decrease and rates of photosynthesis will increase. This change is termed carbon dioxide fertilization and demonstrates the complex interactions between life and climate change.
• Introducing invasive species that compete with native plant or algae species for nutrients, water, light, or other resources, reducing native species populations.

The Earth system model below includes some of the processes and phenomena related to photosynthesis.  These processes operate at various rates and on different spatial and temporal scales. For example, carbon dioxide is transferred among plants and animals over relatively short time periods (hours-weeks), but the deforestation alters ecosystems over decades to centuries, or longer.  Can you think of additional cause and effect relationships between photosynthesis and other processes in the Earth system?

Click the bolded terms (e.g. respiration , productivity and biomass , and burning of fossil fuels ) on this page to learn more about these process and phenomena. Alternatively, explore the Understanding Global Change Infographic and find new topics that are of interest and/or locally relevant to you.

Learn more in these real-world examples, and challenge yourself to  construct a model  that explains the Earth system relationships.

• The bacteria that changed the world
• New York Times: Antarctic Ice Reveals Earth’s Accelerating Plant Growth
• USGCRP: Climate and Health Assessment, Food Safety, Nutrition, and Distribution
• HHMI BioInteractive: Photosynthesis
• Evolution Connection: More on photorespiration

What is photosynthesis?

Photosynthesis is arguably the most important biological process on earth. By liberating oxygen and consuming carbon dioxide, it has transformed the world into the hospitable environment we know today. Directly or indirectly, photosynthesis fills all of our food requirements and many of our needs for fiber and building materials. The energy stored in petroleum, natural gas and coal all came from the sun via photosynthesis, as does the energy in firewood, which is a major fuel in many parts of the world. This being the case, scientific research into photosynthesis is vitally important. If we can understand and control the intricacies of the photosynthetic process, we can learn how to increase crop yields of food, fiber, wood, and fuel, and how to better use our lands. The energy-harvesting secrets of plants can be adapted to man-made systems which provide new, efficient ways to collect and use solar energy. These same natural "technologies" can help point the way to the design of new, faster, and more compact computers, and even to new medical breakthroughs. Because photosynthesis helps control the makeup of our atmosphere, understanding photosynthesis is crucial to understanding how carbon dioxide and other "greenhouse gases" affect the global climate. In this document, we will briefly explore each of the areas mentioned above, and illustrate how photosynthesis research is critical to maintaining and improving our quality of life.

Photosynthesis and food. All of our biological energy needs are met by the plant kingdom, either directly or through herbivorous animals. Plants in turn obtain the energy to synthesize foodstuffs via photosynthesis. Although plants draw necessary materials from the soil and water and carbon dioxide from the air, the energy needs of the plant are filled by sunlight. Sunlight is pure energy. However, sunlight itself is not a very useful form of energy; it cannot be eaten, it cannot turn dynamos, and it cannot be stored. To be beneficial, the energy in sunlight must be converted to other forms. This is what photosynthesis is all about. It is the process by which plants change the energy in sunlight to kinds of energy that can be stored for later use. Plants carry out this process in photosynthetic reaction centers. These tiny units are found in leaves, and convert light energy to chemical energy, which is the form used by all living organisms. One of the major energy-harvesting processes in plants involves using the energy of sunlight to convert carbon dioxide from the air into sugars, starches, and other high-energy carbohydrates. Oxygen is released in the process. Later, when the plant needs food, it draws upon the energy stored in these carbohydrates. We do the same. When we eat a plate of spaghetti, our bodies oxidize or "burn" the starch by allowing it to combine with oxygen from the air. This produces carbon dioxide, which we exhale, and the energy we need to survive. Thus, if there is no photosynthesis, there is no food. Indeed, one widely accepted theory explaining the extinction of the dinosaurs suggests that a comet, meteor, or volcano ejected so much material into the atmosphere that the amount of sunlight reaching the earth was severely reduced. This in turn caused the death of many plants and the creatures that depended upon them for energy.

Photosynthesis and energy. One of the carbohydrates resulting from photosynthesis is cellulose, which makes up the bulk of dry wood and other plant material. When we burn wood, we convert the cellulose back to carbon dioxide and release the stored energy as heat. Burning fuel is basically the same oxidation process that occurs in our bodies; it liberates the energy of "stored sunlight" in a useful form, and returns carbon dioxide to the atmosphere. Energy from burning "biomass" is important in many parts of the world. In developing countries, firewood continues to be critical to survival. Ethanol (grain alcohol) produced from sugars and starches by fermentation is a major automobile fuel in Brazil, and is added to gasoline in some parts of the United States to help reduce emissions of harmful pollutants. Ethanol is also readily converted to ethylene, which serves as a feedstock to a large part of the petrochemical industry. It is possible to convert cellulose to sugar, and then into ethanol; various microorganisms carry out this process. It could be commercially important one day.

Our major sources of energy, of course, are coal, oil and natural gas. These materials are all derived from ancient plants and animals, and the energy stored within them is chemical energy that originally came from sunlight through photosynthesis. Thus, most of the energy we use today was originally solar energy!

Photosynthesis, fiber, and materials. Wood, of course, is not only burned, but is an important material for building and many other purposes. Paper, for example, is nearly pure photosynthetically produced cellulose, as is cotton and many other natural fibers. Even wool production depends on photosynthetically-derived energy. In fact, all plant and animal products including many medicines and drugs require energy to produce, and that energy comes ultimately from sunlight via photosynthesis. Many of our other materials needs are filled by plastics and synthetic fibers which are produced from petroleum, and are thus also photosynthetic in origin. Even much of our metal refining depends ultimately on coal or other photosynthetic products. Indeed, it is difficult to name an economically important material or substance whose existence and usefulness is not in some way tied to photosynthesis.

Photosynthesis and the environment. Currently, there is a lot of discussion concerning the possible effects of carbon dioxide and other "greenhouse gases" on the environment. As mentioned above, photosynthesis converts carbon dioxide from the air to carbohydrates and other kinds of "fixed" carbon and releases oxygen to the atmosphere. When we burn firewood, ethanol, or coal, oil and other fossil fuels, oxygen is consumed, and carbon dioxide is released back to the atmosphere. Thus, carbon dioxide which was removed from the atmosphere over millions of years is being replaced very quickly through our consumption of these fuels. The increase in carbon dioxide and related gases is bound to affect our atmosphere. Will this change be large or small, and will it be harmful or beneficial? These questions are being actively studied by many scientists today. The answers will depend strongly on the effect of photosynthesis carried out by land and sea organisms. As photosynthesis consumes carbon dioxide and releases oxygen, it helps counteract the effect of combustion of fossil fuels. The burning of fossil fuels releases not only carbon dioxide, but also hydrocarbons, nitrogen oxides, and other trace materials that pollute the atmosphere and contribute to long-term health and environmental problems. These problems are a consequence of the fact that nature has chosen to implement photosynthesis through conversion of carbon dioxide to energy-rich materials such as carbohydrates. Can the principles of photosynthetic solar energy harvesting be used in some way to produce non-polluting fuels or energy sources? The answer, as we shall see, is yes.

Why study photosynthesis?

Because our quality of life, and indeed our very existence, depends on photosynthesis, it is essential that we understand it. Through understanding, we can avoid adversely affecting the process and precipitating environmental or ecological disasters. Through understanding, we can also learn to control photosynthesis, and thus enhance production of food, fiber and energy. Understanding the natural process, which has been developed by plants over several billion years, will also allow us to use the basic chemistry and physics of photosynthesis for other purposes, such as solar energy conversion, the design of electronic circuits, and the development of medicines and drugs. Some examples follow.

Photosynthesis and agriculture. Although photosynthesis has interested mankind for eons, rapid progress in understanding the process has come in the last few years. One of the things we have learned is that overall, photosynthesis is relatively inefficient. For example, based on the amount of carbon fixed by a field of corn during a typical growing season, only about 1 - 2% of the solar energy falling on the field is recovered as new photosynthetic products. The efficiency of uncultivated plant life is only about 0.2%. In sugar cane, which is one of the most efficient plants, about 8% of the light absorbed by the plant is preserved as chemical energy. Many plants, especially those that originate in the temperate zones such as most of the United States, undergo a process called photorespiration. This is a kind of "short circuit" of photosynthesis that wastes much of the plants' photosynthetic energy. The phenomenon of photorespiration including its function, if any, is only one of many riddles facing the photosynthesis researcher.

If we can fully understand processes like photorespiration, we will have the ability to alter them. Thus, more efficient plants can be designed. Although new varieties of plants have been developed for centuries through selective breeding, the techniques of modern molecular biology have speeded up the process tremendously. Photosynthesis research can show us how to produce new crop strains that will make much better use of the sunlight they absorb. Research along these lines is critical, as recent studies show that agricultural production is leveling off at a time when demand for food and other agricultural products is increasing rapidly.

Because plants depend upon photosynthesis for their survival, interfering with photosynthesis can kill the plant. This is the basis of several important herbicides, which act by preventing certain important steps of photosynthesis. Understanding the details of photosynthesis can lead to the design of new, extremely selective herbicides and plant growth regulators that have the potential of being environmentally safe (especially to animal life, which does not carry out photosynthesis). Indeed, it is possible to develop new crop plants that are immune to specific herbicides, and to thus achieve weed control specific to one crop species.

Photosynthesis and energy production. As described above, most of our current energy needs are met by photosynthesis, ancient or modern. Increasing the efficiency of natural photosynthesis can also increase production of ethanol and other fuels derived from agriculture. However, knowledge gained from photosynthesis research can also be used to enhance energy production in a much more direct way. Although the overall photosynthesis process is relatively wasteful, the early steps in the conversion of sunlight to chemical energy are quite efficient. Why not learn to understand the basic chemistry and physics of photosynthesis, and use these same principles to build man-made solar energy harvesting devices? This has been a dream of chemists for years, but is now close to becoming a reality. In the laboratory, scientists can now synthesize artificial photosynthetic reaction centers which rival the natural ones in terms of the amount of sunlight stored as chemical or electrical energy. More research will lead to the development of new, efficient solar energy harvesting technologies based on the natural process.

The role of photosynthesis in control of the environment. How does photosynthesis in temperate and tropical forests and in the sea affect the quantity of greenhouse gases in the atmosphere? This is an important and controversial issue today. As mentioned above, photosynthesis by plants removes carbon dioxide from the atmosphere and replaces it with oxygen. Thus, it would tend to ameliorate the effects of carbon dioxide released by the burning of fossil fuels. However, the question is complicated by the fact that plants themselves react to the amount of carbon dioxide in the atmosphere. Some plants, appear to grow more rapidly in an atmosphere rich in carbon dioxide, but this may not be true of all species. Understanding the effect of greenhouse gases requires a much better knowledge of the interaction of the plant kingdom with carbon dioxide than we have today. Burning plants and plant products such as petroleum releases carbon dioxide and other byproducts such as hydrocarbons and nitrogen oxides. However, the pollution caused by such materials is not a necessary product of solar energy utilization. The artificial photosynthetic reaction centers discussed above produce energy without releasing any byproducts other than heat. They hold the promise of producing clean energy in the form of electricity or hydrogen fuel without pollution. Implementation of such solar energy harvesting devices would prevent pollution at the source, which is certainly the most efficient approach to control.

Photosynthesis and electronics. At first glance, photosynthesis would seem to have no association with the design of computers and other electronic devices. However, there is potentially a very strong connection. A goal of modern electronics research is to make transistors and other circuit components as small as possible. Small devices and short connections between them make computers faster and more compact. The smallest possible unit of a material is a molecule (made up of atoms of various types). Thus, the smallest conceivable transistor is a single molecule (or atom). Many researchers today are investigating the intriguing possibility of making electronic components from single molecules or small groups of molecules. Another very active area of research is computers that use light, rather than electrons, as the medium for carrying information. In principle, light-based computers have several advantages over traditional designs, and indeed many of our telephone transmission and switching networks already operate through fiber optics. What does this have to do with photosynthesis? It turns out that photosynthetic reaction centers are natural photochemical switches of molecular dimensions. Learning how plants absorb light, control the movement of the resulting energy to reaction centers, and convert the light energy to electrical, and finally chemical energy can help us understand how to make molecular-scale computers. In fact, several molecular electronic logic elements based on artificial photosynthetic reaction centers have already been reported in the scientific literature.

Photosynthesis and medicine. Light has a very high energy content, and when it is absorbed by a substance this energy is converted to other forms. When the energy ends up in the wrong place, it can cause serious damage to living organisms. Aging of the skin and skin cancer are only two of many deleterious effects of light on humans and animals. Because plants and other photosynthetic species have been dealing with light for eons, they have had to develop photoprotective mechanisms to limit light damage. Learning about the causes of light- induced tissue damage and the details of the natural photoprotective mechanisms can help us can find ways to adapt these processes for the benefit of humanity in areas far removed from photosynthesis itself. For example, the mechanism by which sunlight absorbed by photosynthetic chlorophyll causes tissue damage in plants has been harnessed for medical purposes. Substances related to chlorophyll localize naturally in cancerous tumor tissue. Illumination of the tumors with light then leads to photochemical damage which can kill the tumor while leaving surrounding tissue unharmed. Another medical application involves using similar chlorophyll relatives to localize in tumor tissue, and thus act as dyes which clearly delineate the boundary between cancerous and healthy tissue. This diagnostic aid does not cause photochemical damage to normal tissue because the principles of photosynthesis have been used to endow it with protective agents that harmlessly convert the absorbed light to heat.

Conclusions

The above examples illustrate the importance of photosynthesis as a natural process and the impact that it has on all of our lives. Research into the nature of photosynthesis is crucial because only by understanding photosynthesis can we control it, and harness its principles for the betterment of mankind. Science has only recently developed the basic tools and techniques needed to investigate the intricate details of photosynthesis. It is now time to apply these tools and techniques to the problem, and to begin to reap the benefits of this research.

Written by and Copyright ©1996 Devens Gust Professor of Chemistry and Biochemistry, Arizona State University

A  translation  of this article into Belorussian by Martha Ruszkowski is available

Learn more about Devens Gust

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Photosynthesis

Matthew p. johnson.

Department of Molecular Biology and Biotechnology, University of Sheffield, Firth Court, Western Bank, Sheffield S10 2TN, U.K.

Photosynthesis sustains virtually all life on planet Earth providing the oxygen we breathe and the food we eat; it forms the basis of global food chains and meets the majority of humankind's current energy needs through fossilized photosynthetic fuels. The process of photosynthesis in plants is based on two reactions that are carried out by separate parts of the chloroplast. The light reactions occur in the chloroplast thylakoid membrane and involve the splitting of water into oxygen, protons and electrons. The protons and electrons are then transferred through the thylakoid membrane to create the energy storage molecules adenosine triphosphate (ATP) and nicotinomide–adenine dinucleotide phosphate (NADPH). The ATP and NADPH are then utilized by the enzymes of the Calvin–Benson cycle (the dark reactions), which converts CO 2 into carbohydrate in the chloroplast stroma. The basic principles of solar energy capture, energy, electron and proton transfer and the biochemical basis of carbon fixation are explained and their significance is discussed.

An overview of photosynthesis

Introduction.

Photosynthesis is the ultimate source of all of humankind's food and oxygen, whereas fossilized photosynthetic fuels provide ∼87% of the world's energy. It is the biochemical process that sustains the biosphere as the basis for the food chain. The oxygen produced as a by-product of photosynthesis allowed the formation of the ozone layer, the evolution of aerobic respiration and thus complex multicellular life.

Oxygenic photosynthesis involves the conversion of water and CO 2 into complex organic molecules such as carbohydrates and oxygen. Photosynthesis may be split into the ‘light’ and ‘dark’ reactions. In the light reactions, water is split using light into oxygen, protons and electrons, and in the dark reactions, the protons and electrons are used to reduce CO 2 to carbohydrate (given here by the general formula CH 2 O). The two processes can be summarized thus:

Light reactions:

Dark reactions:

The positive sign of the standard free energy change of the reaction (Δ G °) given above means that the reaction requires energy ( an endergonic reaction ). The energy required is provided by absorbed solar energy, which is converted into the chemical bond energy of the products ( Box 1 ).

Standard free energy change

Photosynthesis converts ∼200 billion tonnes of CO 2 into complex organic compounds annually and produces ∼140 billion tonnes of oxygen into the atmosphere. By facilitating conversion of solar energy into chemical energy, photosynthesis acts as the primary energy input into the global food chain. Nearly all living organisms use the complex organic compounds derived from photosynthesis as a source of energy. The breakdown of these organic compounds occurs via the process of aerobic respiration, which of course also requires the oxygen produced by photosynthesis.

Unlike photosynthesis, aerobic respiration is an exergonic process (negative Δ G °) with the energy released being used by the organism to power biosynthetic processes that allow growth and renewal, mechanical work (such as muscle contraction or flagella rotation) and facilitating changes in chemical concentrations within the cell (e.g. accumulation of nutrients and expulsion of waste). The use of exergonic reactions to power endergonic ones associated with biosynthesis and housekeeping in biological organisms such that the overall free energy change is negative is known as ‘ coupling’.

Photosynthesis and respiration are thus seemingly the reverse of one another, with the important caveat that both oxygen formation during photosynthesis and its utilization during respiration result in its liberation or incorporation respectively into water rather than CO 2 . In addition, glucose is one of several possible products of photosynthesis with amino acids and lipids also being synthesized rapidly from the primary photosynthetic products.

The consideration of photosynthesis and respiration as opposing processes helps us to appreciate their role in shaping our environment. The fixation of CO 2 by photosynthesis and its release during breakdown of organic molecules during respiration, decay and combustion of organic matter and fossil fuels can be visualized as the global carbon cycle ( Figure 1 ).

The relationship between respiration, photosynthesis and global CO 2 and O 2 levels.

At present, this cycle may be considered to be in a state of imbalance due to the burning of fossil fuels (fossilized photosynthesis), which is increasing the proportion of CO 2 entering the Earth's atmosphere, leading to the so-called ‘greenhouse effect’ and human-made climate change.

Oxygenic photosynthesis is thought to have evolved only once during Earth's history in the cyanobacteria. All other organisms, such as plants, algae and diatoms, which perform oxygenic photosynthesis actually do so via cyanobacterial endosymbionts or ‘chloroplasts’. An endosymbiotoic event between an ancestral eukaryotic cell and a cyanobacterium that gave rise to plants is estimated to have occurred ∼1.5 billion years ago. Free-living cyanobacteria still exist today and are responsible for ∼50% of the world's photosynthesis. Cyanobacteria themselves are thought to have evolved from simpler photosynthetic bacteria that use either organic or inorganic compounds such a hydrogen sulfide as a source of electrons rather than water and thus do not produce oxygen.

The site of photosynthesis in plants

In land plants, the principal organs of photosynthesis are the leaves ( Figure 2 A). Leaves have evolved to expose the largest possible area of green tissue to light and entry of CO 2 to the leaf is controlled by small holes in the lower epidermis called stomata ( Figure 2 B). The size of the stomatal openings is variable and regulated by a pair of guard cells, which respond to the turgor pressure (water content) of the leaf, thus when the leaf is hydrated, the stomata can open to allow CO 2 in. In contrast, when water is scarce, the guard cells lose turgor pressure and close, preventing the escape of water from the leaf via transpiration.

( A ) The model plant Arabidopsis thaliana . ( B ) Basic structure of a leaf shown in cross-section. Chloroplasts are shown as green dots within the cells. ( C ) An electron micrograph of an Arabidopsis chloroplast within the leaf. ( D ) Close-up region of the chloroplast showing the stacked structure of the thylakoid membrane.

Within the green tissue of the leaf (mainly the mesophyll) each cell (∼100 μm in length) contains ∼100 chloroplasts (2–3 μm in length), the tiny organelles where photosynthesis takes place. The chloroplast has a complex structure ( Figure 2 C, D) with two outer membranes (the envelope), which are colourless and do not participate in photosynthesis, enclosing an aqueous space (the stroma) wherein sits a third membrane known as the thylakoid, which in turn encloses a single continuous aqueous space called the lumen.

The light reactions of photosynthesis involve light-driven electron and proton transfers, which occur in the thylakoid membrane, whereas the dark reactions involve the fixation of CO 2 into carbohydrate, via the Calvin–Benson cycle, which occurs in the stroma ( Figure 3 ). The light reactions involve electron transfer from water to NADP + to form NADPH and these reactions are coupled to proton transfers that lead to the phosphorylation of adenosine diphosphate (ADP) into ATP. The Calvin–Benson cycle uses ATP and NADPH to convert CO 2 into carbohydrates ( Figure 3 ), regenerating ADP and NADP + . The light and dark reactions are therefore mutually dependent on one another.

The light reactions of photosynthesis take place in the thylakoid membrane, whereas the dark reactions are located in the chloroplast stroma.

Photosynthetic electron and proton transfer chain

The light-driven electron transfer reactions of photosynthesis begin with the splitting of water by Photosystem II (PSII). PSII is a chlorophyll–protein complex embedded in the thylakoid membrane that uses light to oxidize water to oxygen and reduce the electron acceptor plastoquinone to plastoquinol. Plastoquinol in turn carries the electrons derived from water to another thylakoid-embedded protein complex called cytochrome b 6 f (cyt b 6 f ). cyt b 6 f oxidizes plastoquinol to plastoquinone and reduces a small water-soluble electron carrier protein plastocyanin, which resides in the lumen. A second light-driven reaction is then carried out by another chlorophyll protein complex called Photosystem I (PSI). PSI oxidizes plastocyanin and reduces another soluble electron carrier protein ferredoxin that resides in the stroma. Ferredoxin can then be used by the ferredoxin–NADP + reductase (FNR) enzyme to reduce NADP + to NADPH. This scheme is known as the linear electron transfer pathway or Z-scheme ( Figure 4 ).

The linear electron transfer pathway from water to NADP + to form NADPH results in the formation of a proton gradient across the thylakoid membrane that is used by the ATP synthase enzyme to make ATP.

The Z-scheme, so-called since it resembles the letter ‘Z’ when turned on its side ( Figure 5 ), thus shows how the electrons move from the water–oxygen couple (+820 mV) via a chain of redox carriers to NADP + /NADPH (−320 mV) during photosynthetic electron transfer. Generally, electrons are transferred from redox couples with low potentials (good reductants) to those with higher potentials (good oxidants) (e.g. during respiratory electron transfer in mitochondria) since this process is exergonic (see Box 2 ). However, photosynthetic electron transfer also involves two endergonic steps, which occur at PSII and at PSI and require an energy input in the form of light. The light energy is used to excite an electron within a chlorophyll molecule residing in PSII or PSI to a higher energy level; this excited chlorophyll is then able to reduce the subsequent acceptors in the chain. The oxidized chlorophyll is then reduced by water in the case of PSII and plastocyanin in the case of PSI.

The main components of the linear electron transfer pathway are shown on a scale of redox potential to illustrate how two separate inputs of light energy at PSI and PSII result in the endergonic transfer of electrons from water to NADP + .

Relationship between redox potentials and standard free energy changes

The water-splitting reaction at PSII and plastoquinol oxidation at cyt b 6 f result in the release of protons into the lumen, resulting in a build-up of protons in this compartment relative to the stroma. The difference in the proton concentration between the two sides of the membrane is called a proton gradient. The proton gradient is a store of free energy (similar to a gradient of ions in a battery) that is utilized by a molecular mechanical motor ATP synthase, which resides in the thylakoid membrane ( Figure 4 ). The ATP synthase allows the protons to move down their concentration gradient from the lumen (high H + concentration) to the stroma (low H + concentration). This exergonic reaction is used to power the endergonic synthesis of ATP from ADP and inorganic phosphate (P i ). This process of photophosphorylation is thus essentially similar to oxidative phosphorylation, which occurs in the inner mitochondrial membrane during respiration.

An alternative electron transfer pathway exists in plants and algae, known as cyclic electron flow. Cyclic electron flow involves the recycling of electrons from ferredoxin to plastoquinone, with the result that there is no net production of NADPH; however, since protons are still transferred into the lumen by oxidation of plastoquinol by cyt b 6 f , ATP can still be formed. Thus photosynthetic organisms can control the ratio of NADPH/ATP to meet metabolic need by controlling the relative amounts of cyclic and linear electron transfer.

How the photosystems work

Light absorption by pigments.

Photosynthesis begins with the absorption of light by pigments molecules located in the thylakoid membrane. The most well-known of these is chlorophyll, but there are also carotenoids and, in cyanobacteria and some algae, bilins. These pigments all have in common within their chemical structures an alternating series of carbon single and double bonds, which form a conjugated system π–electron system ( Figure 6 ).

The chemical structures of the chlorophyll and carotenoid pigments present in the thylakoid membrane. Note the presence in each of a conjugated system of carbon–carbon double bonds that is responsible for light absorption.

The variety of pigments present within each type of photosynthetic organism reflects the light environment in which it lives; plants on land contain chlorophylls a and b and carotenoids such as β-carotene, lutein, zeaxanthin, violaxanthin, antheraxanthin and neoxanthin ( Figure 6 ). The chlorophylls absorb blue and red light and so appear green in colour, whereas carotenoids absorb light only in the blue and so appear yellow/red ( Figure 7 ), colours more obvious in the autumn as chlorophyll is the first pigment to be broken down in decaying leaves.

Chlorophylls absorb light energy in the red and blue part of the visible spectrum, whereas carotenoids only absorb light in the blue/green.

Light, or electromagnetic radiation, has the properties of both a wave and a stream of particles (light quanta). Each quantum of light contains a discrete amount of energy that can be calculated by multiplying Planck's constant, h (6.626×10 −34 J·s) by ν, the frequency of the radiation in cycles per second (s −1 ):

The frequency (ν) of the light and so its energy varies with its colour, thus blue photons (∼450 nm) are more energetic than red photons (∼650 nm). The frequency (ν) and wavelength (λ) of light are related by:

where c is the velocity of light (3.0×10 8 m·s −1 ), and the energy of a particular wavelength (λ) of light is given by:

Thus 1 mol of 680 nm photons of red light has an energy of 176 kJ·mol −1 .

The electrons within the delocalized π system of the pigment have the ability to jump up from the lowest occupied molecular orbital (ground state) to higher unoccupied molecular electron orbitals (excited states) via the absorption of specific wavelengths of light in the visible range (400–725 nm). Chlorophyll has two excited states known as S 1 and S 2 and, upon interaction of the molecule with a photon of light, one of its π electrons is promoted from the ground state (S 0 ) to an excited state, a process taking just 10 −15 s ( Figure 8 ). The energy gap between the S 0 and S 1 states is spanned by the energy provided by a red photon (∼600–700 nm), whereas the energy gap between the S 0 and S 2 states is larger and therefore requires a more energetic (shorter wavelength, higher frequency) blue photon (∼400–500 nm) to span the energy gap.

Photons with slightly different energies (colours) excite each of the vibrational substates of each excited state (as shown by variation in the size and colour of the arrows).

Upon excitation, the electron in the S 2 state quickly undergoes losses of energy as heat through molecular vibration and undergoes conversion into the energy of the S 1 state by a process called internal conversion. Internal conversion occurs on a timescale of 10 −12 s. The energy of a blue photon is thus rapidly degraded to that of a red photon. Excitation of the molecule with a red photon would lead to promotion of an electron to the S 1 state directly. Once the electron resides in the S 1 state, it is lower in energy and thus stable on a somewhat longer timescale (10 −9 s). The energy of the excited electron in the S 1 state can have one of several fates: it could return to the ground state (S 0 ) by emission of the energy as a photon of light (fluorescence), or it could be lost as heat due to internal conversion between S 1 and S 0 . Alternatively, if another chlorophyll is nearby, a process known as excitation energy transfer (EET) can result in the non-radiative exchange of energy between the two molecules ( Figure 9 ). For this to occur, the two chlorophylls must be close by (<7 nm), have a specific orientation with respect to one another, and excited state energies that overlap (are resonant) with one another. If these conditions are met, the energy is exchanged, resulting in a mirror S 0 →S 1 transition in the acceptor molecule and a S 1 →S 0 transition in the other.

Two chlorophyll molecules with resonant S 1 states undergo a mirror transition resulting in the non-radiative transfer of excitation energy between them.

Light-harvesting complexes

In photosynthetic systems, chlorophylls and carotenoids are found attached to membrane-embedded proteins known as light-harvesting complexes (LHCs). Through careful binding and orientation of the pigment molecules, absorbed energy can be transferred among them by EET. Each pigment is bound to the protein by a series of non-covalent bonding interactions (such as, hydrogen bonds, van der Waals interactions, hydrophobic interaction and co-ordination bonds between lone pair electrons of residues such as histidine in the protein and the Mg 2+ ion in chlorophyll); the protein structure is such that each bound pigment experiences a slightly different environment in terms of the surrounding amino acid side chains, lipids, etc., meaning that the S 1 and S 2 energy levels are shifted in energy with respect to that of other neighbouring pigment molecules. The effect is to create a range of pigment energies that act to ‘funnel’ the energy on to the lowest-energy pigments in the LHC by EET.

Reaction centres

A photosystem consists of numerous LHCs that form an antenna of hundreds of pigment molecules. The antenna pigments act to collect and concentrate excitation energy and transfer it towards a ‘special pair’ of chlorophyll molecules that reside in the reaction centre (RC) ( Figure 10 ). Unlike the antenna pigments, the special pair of chlorophylls are ‘redox-active’ in the sense that they can return to the ground state (S 0 ) by the transfer of the electron residing in the S 1 excited state (Chl*) to another species. This process is known as charge separation and result in formation of an oxidized special pair (Chl + ) and a reduced acceptor (A − ). The acceptor in PSII is plastoquinone and in PSI it is ferredoxin. If the RC is to go on functioning, the electron deficiency on the special pair must be made good, in PSII the electron donor is water and in PSI it is plastocyanin.

Light energy is captured by the antenna pigments and transferred to the special pair of RC chlorophylls which undergo a redox reaction leading to reduction of an acceptor molecule. The oxidized special pair is regenerated by an electron donor.

It is worth asking why photosynthetic organisms bother to have a large antenna of pigments serving an RC rather than more numerous RCs. The answer lies in the fact that the special pair of chlorophylls alone have a rather small spatial and spectral cross-section, meaning that there is a limit to the amount of light they can efficiently absorb. The amount of light they can practically absorb is around two orders of magnitude smaller than their maximum possible turnover rate, Thus LHCs act to increase the spatial (hundreds of pigments) and spectral (several types of pigments with different light absorption characteristics) cross-section of the RC special pair ensuring that its turnover rate runs much closer to capacity.

Photosystem II

PSII is a light-driven water–plastoquinone oxidoreductase and is the only enzyme in Nature that is capable of performing the difficult chemistry of splitting water into protons, electrons and oxygen ( Figure 11 ). In principle, water is an extremely poor electron donor since the redox potential of the water–oxygen couple is +820 mV. PSII uses light energy to excite a special pair of chlorophylls, known as P680 due to their 680 nm absorption peak in the red part of the spectrum. P680* undergoes charge separation that results in the formation of an extremely oxidizing species P680 + which has a redox potential of +1200 mV, sufficient to oxidize water. Nonetheless, since water splitting involves four electron chemistry and charge separation only involves transfer of one electron, four separate charge separations (turnovers of PSII) are required to drive formation of one molecule of O 2 from two molecules of water. The initial electron donation to generate the P680 from P680 + is therefore provided by a cluster of manganese ions within the oxygen-evolving complex (OEC), which is attached to the lumen side of PSII ( Figure 12 ). Manganese is a transition metal that can exist in a range of oxidation states from +1 to +5 and thus accumulates the positive charges derived from each light-driven turnover of P680. Progressive extraction of electrons from the manganese cluster is driven by the oxidation of P680 within PSII by light and is known as the S-state cycle ( Figure 12 ). After the fourth turnover of P680, sufficient positive charge is built up in the manganese cluster to permit the splitting of water into electrons, which regenerate the original state of the manganese cluster, protons, which are released into the lumen and contribute to the proton gradient used for ATP synthesis, and the by-product O 2 . Thus charge separation at P680 provides the thermodynamic driving force, whereas the manganese cluster acts as a catalyst for the water-splitting reaction.

The organization of PSII and its light-harvesting antenna. Protein is shown in grey, with chlorophylls in green and carotenoids in orange. Drawn from PDB code 3JCU

Progressive extraction of electrons from the manganese cluster is driven by the oxidation of P680 within PSII by light. Each of the electrons given up by the cluster is eventually repaid at the S 4 to S 0 transition when molecular oxygen (O 2 ) is formed. The protons extracted from water during the process are deposited into the lumen and contribute to the protonmotive force.

The electrons yielded by P680* following charge separation are not passed directly to plastoquinone, but rather via another acceptor called pheophytin, a porphyrin molecule lacking the central magnesium ion as in chlorophyll. Plastoquinone reduction to plastoquinol requires two electrons and thus two molecules of plastoquinol are formed per O 2 molecule evolved by PSII. Two protons are also taken up upon formation of plastoquinol and these are derived from the stroma. PSII is found within the thylakoid membrane of plants as a dimeric RC complex surrounded by a peripheral antenna of six minor monomeric antenna LHC complexes and two to eight trimeric LHC complexes, which together form a PSII–LHCII supercomplex ( Figure 11 ).

Photosystem I

PSI is a light-driven plastocyanin–ferredoxin oxidoreductase ( Figure 13 ). In PSI, the special pair of chlorophylls are known as P700 due to their 700 nm absorption peak in the red part of the spectrum. P700* is an extremely strong reductant that is able to reduce ferredoxin which has a redox potential of −450 mV (and is thus is, in principle, a poor electron acceptor). Reduced ferredoxin is then used to generate NADPH for the Calvin–Benson cycle at a separate complex known as FNR. The electron from P700* is donated via another chlorophyll molecule and a bound quinone to a series of iron–sulfur clusters at the stromal side of the complex, whereupon the electron is donated to ferredoxin. The P700 species is regenerated form P700 + via donation of an electron from the soluble electron carrier protein plastocyanin.

The organization of PSI and its light-harvesting antenna. Protein is shown in grey, with chlorophylls in green and carotenoids in orange. Drawn from PDB code 4XK8.

PSI is found within the thylakoid membrane as a monomeric RC surrounded on one side by four LHC complexes known as LHCI. The PSI–LHCI supercomplex is found mainly in the unstacked regions of the thylakoid membrane ( Figure 13 ).

Other electron transfer chain components

Plastoquinone/plastoquinol.

Plastoquinone is a small lipophilic electron carrier molecule that resides within the thylakoid membrane and carries two electrons and two protons from PSII to the cyt b 6 f complex. It has a very similar structure to that of the molecule ubiquinone (coenzyme Q 10 ) in the mitochondrial inner membrane.

Cytochrome b 6 f complex

The cyt b 6 f complex is a plastoquinol–plastocyanin oxidoreductase and possess a similar structure to that of the cytochrome bc 1 complex (complex III) in mitochondria ( Figure 14 A). As with Complex III, cyt b 6 f exists as a dimer in the membrane and carries out both the oxidation and reduction of quinones via the so-called Q-cycle. The Q-cycle ( Figure 14 B) involves oxidation of one plastoquinol molecule at the Qp site of the complex, both protons from this molecule are deposited in the lumen and contribute to the proton gradient for ATP synthesis. The two electrons, however, have different fates. The first is transferred via an iron–sulfur cluster and a haem cofactor to the soluble electron carrier plastocyanin (see below). The second electron derived from plastoquinol is passed via two separate haem cofactors to another molecule of plastoquinone bound to a separate site (Qn) on the complex, thus reducing it to a semiquinone. When a second plastoquinol molecule is oxidized at Qp, a second molecule of plastocyanin is reduced and two further protons are deposited in the lumen. The second electron reduces the semiquinone at the Qn site which, concomitant with uptake of two protons from the stroma, causes its reduction to plastoquinol. Thus for each pair of plastoquinol molecules oxidized by the complex, one is regenerated, yet all four protons are deposited into the lumen. The Q-cycle thus doubles the number of protons transferred from the stroma to the lumen per plastoquinol molecule oxidized.

( A ) Structure drawn from PDB code 1Q90. ( B ) The protonmotive Q-cycle showing how electrons from plastoquinol are passed to both plastocyanin and plastoquinone, doubling the protons deposited in the lumen for every plastoquinol molecule oxidized by the complex.

Plastocyanin

Plastocyanin is a small soluble electron carrier protein that resides in the thylakoid lumen. The active site of the plastocyanin protein binds a copper ion, which cycles between the Cu 2+ and Cu + oxidation states following its oxidation by PSI and reduction by cyt b 6 f respectively.

Ferredoxin is a small soluble electron carrier protein that resides in the chloroplast stroma. The active site of the ferredoxin protein binds an iron–sulfur cluster, which cycles between the Fe 2+ and Fe 3+ oxidation states following its reduction by PSI and oxidation by the FNR complex respectively.

Ferredoxin–NADP + reductase

The FNR complex is found in both soluble and thylakoid membrane-bound forms. The complex binds a flavin–adenine dinucleotide (FAD) cofactor at its active site, which accepts two electrons from two molecules of ferredoxin before using them reduce NADP + to NADPH.

ATP synthase

The ATP synthase enzyme is responsible for making ATP from ADP and P i ; this endergonic reaction is powered by the energy contained within the protonmotive force. According to the structure, 4.67 H + are required for every ATP molecule synthesized by the chloroplast ATP synthase. The enzyme is a rotary motor which contains two domains: the membrane-spanning F O portion which conducts protons from the lumen to the stroma, and the F 1 catalytic domain that couples this exergonic proton movement to ATP synthesis.

Membrane stacking and the regulation of photosynthesis

Within the thylakoid membrane, PSII–LHCII supercomplexes are packed together into domains known as the grana, which associate with one another to form grana stacks. PSI and ATP synthase are excluded from these stacked PSII–LHCII regions by steric constraints and thus PSII and PSI are segregated in the thylakoid membrane between the stacked and unstacked regions ( Figure 15 ). The cyt b 6 f complex, in contrast, is evenly distributed throughout the grana and stromal lamellae. The evolutionary advantage of membrane stacking is believed to be a higher efficiency of electron transport by preventing the fast energy trap PSI from ‘stealing’ excitation energy from the slower trap PSII, a phenomenon known as spillover. Another possible advantage of membrane stacking in thylakoids may be the segregation of the linear and cyclic electron transfer pathways, which might otherwise compete to reduce plastoquinone. In this view, PSII, cyt b 6 f and a sub-fraction of PSI closest to the grana is involved in linear flow, whereas PSI and cyt b 6 f in the stromal lamellae participates in cyclic flow. The cyclic electron transfer pathway recycles electrons from ferredoxin back to plastoquinone and thus allows protonmotive force generation (and ATP synthesis) without net NADPH production. Cyclic electron transfer thereby provides the additional ATP required for the Calvin–Benson cycle (see below).

( A ) Electron micrograph of the thylakoid membrane showing stacked grana and unstacked stromal lamellae regions. ( B ) Model showing the distribution of the major complexes of photosynthetic electron and proton transfer between the stacked grana and unstacked stromal lamellae regions.

‘Dark’ reactions: the Calvin–Benson cycle

CO 2 is fixed into carbohydrate via the Calvin–Benson cycle in plants, which consumes the ATP and NADPH produced during the light reactions and thus in turn regenerates ADP, P i and NADP + . In the first step of the Calvin–Benson cycle ( Figure 16 ), CO 2 is combined with a 5-carbon (5C) sugar, ribulose 1,5-bisphosphate in a reaction catalysed by the enzyme ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). The reaction forms an unstable 6C intermediate that immediately splits into two molecules of 3-phosphoglycerate. 3-Phosphoglycerate is first phosphorylated by 3-phosphoglycerate kinase using ATP to form 1,3-bisphosphoglycerate. 1,3-Bisphosphoglycerate is then reduced by glyceraldehyde 3-phosphate dehydrogenase using NADPH to form glyceraldehyde 3-phosphate (GAP, a triose or 3C sugar) in reactions, which are the reverse of glycolysis. For every three CO 2 molecules initially combined with ribulose 1,5-bisphopshate, six molecules of GAP are produced by the subsequent steps. However only one of these six molecules can be considered as a product of the Calvin–Benson cycle since the remaining five are required to regenerate ribulose 1,5-bisphosphate in a complex series of reactions that also require ATP. The one molecule of GAP that is produced for each turn of the cycle can be quickly converted by a range of metabolic pathways into amino acids, lipids or sugars such as glucose. Glucose in turn may be stored as the polymer starch as large granules within chloroplasts.

Overview of the biochemical pathway for the fixation of CO 2 into carbohydrate in plants.

A complex biochemical ‘dance’ ( Figure 16 ) is then involved in the regeneration of three ribulose 1,5-bisphosphate (5C) from the remaining five GAP (3C) molecules. The regeneration begins with the conversion of two molecules of GAP into dihydroxyacetone phosphate (DHAP) by triose phosphate isomerase; one of the DHAP molecules is the combined with another GAP molecule to make fructose 1,6-bisphosphate (6C) by aldolase. The fructose 1,6-bisphosphate is then dephosphorylated by fructose-1,6-bisphosphatase to yield fructose 6-phosphate (6C) and releasing P i . Two carbons are then removed from fructose 6-phosphate by transketolase, generating erythrose 4-phosphate (4C); the two carbons are transferred to another molecule of GAP generating xylulose 5-phosphate (5C). Another DHAP molecule, formed from GAP by triose phosphate isomerase is then combined with the erythrose 4-phosphate by aldolase to form sedoheptulose 1,7-bisphosphate (7C). Sedoheptulose 1,7-bisphosphate is then dephosphorylated to sedoheptulose 7-phosphate (7C) by sedoheptulose-1,7-bisphosphatase releasing P i . Sedoheptulose 7-phosphate has two carbons removed by transketolase to produce ribose 5-phosphate (5C) and the two carbons are transferred to another GAP molecule producing another xylulose 5-phosphate (5C). Ribose 5-phosphate and the two molecules of xylulose 5-phosphate (5C) are then converted by phosphopentose isomerase to three molecules of ribulose 5-phosphate (5C). The three ribulose 5-phosphate molecules are then phosphorylated using three ATP by phosphoribulokinase to regenerate three ribulose 1,5-bisphosphate (5C).

Overall the synthesis of 1 mol of GAP requires 9 mol of ATP and 6 mol of NADPH, a required ratio of 1.5 ATP/NADPH. Linear electron transfer is generally thought to supply ATP/NADPH in a ratio of 1.28 (assuming an H + /ATP ratio of 4.67) with the shortfall of ATP believed to be provided by cyclic electron transfer reactions. Since the product of the Calvin cycle is GAP (a 3C sugar) the pathway is often referred to as C 3 photosynthesis and plants that utilize it are called C 3 plants and include many of the world's major crops such as rice, wheat and potato.

Many of the enzymes involved in the Calvin–Benson cycle (e.g. transketolase, glyceraldehyde-3-phosphate dehydrogenase and aldolase) are also involved in the glycolysis pathway of carbohydrate degradation and their activity must therefore be carefully regulated to avoid futile cycling when light is present, i.e. the unwanted degradation of carbohydrate. The regulation of the Calvin–Benson cycle enzymes is achieved by the activity of the light reactions, which modify the environment of the dark reactions (i.e. the stroma). Proton gradient formation across the thylakoid membrane during the light reactions increases the pH and also increases the Mg 2+ concentration in the stroma (as Mg 2+ flows out of the lumen as H + flows in to compensate for the influx of positive charges). In addition, by reducing ferredoxin and NADP + , PSI changes the redox state of the stroma, which is sensed by the regulatory protein thioredoxin. Thioredoxin, pH and Mg 2+ concentration play a key role in regulating the activity of the Calvin–Benson cycle enzymes, ensuring the activity of the light and dark reactions is closely co-ordinated.

It is noteworthy that, despite the complexity of the dark reactions outlined above, the carbon fixation step itself (i.e. the incorporation of CO 2 into carbohydrate) is carried out by a single enzyme, Rubisco. Rubisco is a large multisubunit soluble protein complex found in the chloroplast stroma. The complex consists of eight large (56 kDa) subunits, which contain both catalytic and regulatory domains, and eight small subunits (14 kDa), which enhance the catalytic function of the L subunits ( Figure 17 A). The carboxylation reaction carried out by Rubisco is highly exergonic (Δ G °=−51.9 kJ·mol- 1 ), yet kinetically very slow (just 3 s −1 ) and begins with the protonation of ribulose 1,5-bisphosphate to form an enediolate intermediate which can be combined with CO 2 to form an unstable 6C intermediate that is quickly hydrolysed to yield two 3C 3-phosphoglycerate molecules. The active site in the Rubisco enzyme contains a key lysine residue, which reacts with another (non-substrate) molecule of CO 2 to form a carbamate anion that is then able to bind Mg 2+ . The Mg 2+ in the active site is essential for the catalytic function of Rubisco, playing a key role in binding ribulose 1,5-bisphosphate and activating it such that it readily reacts with CO 2.. Rubisco activity is co-ordinated with that of the light reactions since carbamate formation requires both high Mg 2+ concentration and alkaline conditions, which are provided by the light-driven changes in the stromal environment discussed above ( Figure 17 B).

( A ) Structure of the Rubisco enzyme (the large subunits are shown in blue and the small subunits in green); four of each type of subunit are visible in the image. Drawn from PDB code 1RXO. ( B ) Activation of the lysine residue within the active site of Rubisco occurs via elevated stromal pH and Mg 2+ concentration as a result of the activity of the light reactions.

In addition to carboxylation, Rubisco also catalyses a competitive oxygenation reaction, known as photorespiration, that results in the combination of ribulose 1,5-bisphosphate with O 2 rather than CO 2 . In the oxygenation reaction, one rather than two molecules of 3-phosphoglycerate and one molecule of a 2C sugar known as phosphoglycolate are produced by Rubisco. The phosphoglycolate must be converted in a series of reactions that regenerate one molecule of 3-phosphoglycerate and one molecule of CO 2 . These reactions consume additional ATP and thus result in an energy loss to the plant. Although the oxygenation reaction of Rubisco is much less favourable than the carboxylation reaction, the relatively high concentration of O 2 in the leaf (250 μM) compared with CO 2 (10 μM) means that a significant amount of photorespiration is always occurring. Under normal conditions, the ratio of carboxylation to oxygenation is between 3:1 and 4:1. However, this ratio can be decreased with increasing temperature due to decreased CO 2 concentration in the leaf, a decrease in the affinity of Rubisco for CO 2 compared with O 2 and an increase in the maximum rate of the oxygenation reaction compared with the carboxylation reaction. The inefficiencies of the Rubisco enzyme mean that plants must produce it in very large amounts (∼30–50% of total soluble protein in a spinach leaf) to achieve the maximal photosynthetic rate.

CO 2 -concentrating mechanisms

To counter photorespiration, plants, algae and cyanobacteria have evolved different CO 2 -concentrating mechanisms CCMs that aim to increase the concentration of CO 2 relative to O 2 in the vicinity of Rubisco. One such CCM is C 4 photosynthesis that is found in plants such as maize, sugar cane and savanna grasses. C 4 plants show a specialized leaf anatomy: Kranz anatomy ( Figure 18 ). Kranz, German for wreath, refers to a bundle sheath of cells that surrounds the central vein within the leaf, which in turn are surrounded by the mesophyll cells. The mesophyll cells in such leaves are rich in the enzyme phosphoenolpyruvate (PEP) carboxylase, which fixes CO 2 into a 4C carboxylic acid: oxaloaceatate. The oxaloacetate formed by the mesophyll cells is reduced using NADPH to malate, another 4C acid: malate. The malate is then exported from the mesophyll cells to the bundle sheath cells, where it is decarboxylated to pyruvate thus regenerating NADPH and CO 2 . The CO 2 is then utilized by Rubisco in the Calvin cycle. The pyruvate is in turn returned to the mesophyll cells where it is phosphorylated using ATP to reform PEP ( Figure 19 ). The advantage of C 4 photosynthesis is that CO 2 accumulates at a very high concentration in the bundle sheath cells that is then sufficient to allow Rubisco to operate efficiently.

Plants growing in hot, bright and dry conditions inevitably have to have their stomata closed for large parts of the day to avoid excessive water loss and wilting. The net result is that the internal CO 2 concentration in the leaf is very low, meaning that C 3 photosynthesis is not possible. To counter this limitation, another CCM is found in succulent plants such as cacti. The Crassulaceae fix CO 2 into malate during the day via PEP carboxylase, store it within the vacuole of the plant cell at night and then release it within their tissues by day to be fixed via normal C 3 photosynthesis. This is termed crassulacean acid metabolism (CAM).

Acknowledgments

I thank Professor Colin Osborne (University of Sheffield, Sheffield, U.K.) for useful discussions on the article, Dr Dan Canniffe (Penn State University, Pennsylvania, PA, U.S.A.) for providing pure pigment spectra and Dr P.J. Weaire (Kingston University, Kingston-upon-Thames, U.K.) for his original Photosynthesis BASC article (1994) on which this essay is partly based.

Abbreviations

This article is a reviewed, revised and updated version of the following ‘Biochemistry Across the School Curriculum’ (BASC) booklet: Weaire, P.J. (1994) Photosynthesis . For further information and to provide feedback on this or any other Biochemical Society education resource, please contact [email protected]. For further information on other Biochemical Society publications, please visit www.biochemistry.org/publications .

Competing Interests

The Author declares that there are no competing interests associated with this article.

Recommended reading and key publications

• Blankenship R.E. Early evolution of photosynthesis. Plant Physiol. 2010; 154 :434–438. doi: 10.1104/pp.110.161687. [ PMC free article ] [ PubMed ] [ CrossRef ] [ Google Scholar ]
• Blankenship R.E. Molecular Mechanisms of Photosynthesis. Chichester: Wiley–Blackwell Publishing; 2014. [ Google Scholar ]
• Nelson N., Ben Shem A. The complex architecture of oxygenic photosynthesis. Nat. Rev. 2004; 5 :1–12. doi: 10.1038/nrm1525. [ PubMed ] [ CrossRef ] [ Google Scholar ]
• Raines C. The Calvin cycle revisited. Photosynth. Res. 2003; 75 :1–10. doi: 10.1023/A:1022421515027. [ PubMed ] [ CrossRef ] [ Google Scholar ]
• Ruban A.V. Evolution under the sun: optimizing light harvesting in photosynthesis. J. Exp. Bot. 2015; 66 :7–23. doi: 10.1093/jxb/eru400. [ PubMed ] [ CrossRef ] [ Google Scholar ]
• Sage R.F. The evolution of C 4 photosynthesis. New Phytol. 2004; 161 :341–370. doi: 10.1111/j.1469-8137.2004.00974.x. [ PubMed ] [ CrossRef ] [ Google Scholar ]

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Article Contents

Photosynthetic research in plant science.

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Ayumi Tanaka, Amane Makino, Photosynthetic Research in Plant Science, Plant and Cell Physiology , Volume 50, Issue 4, April 2009, Pages 681–683, https://doi.org/10.1093/pcp/pcp040

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Photosynthesis is a highly regulated, multistep process. It encompasses the harvest of solar energy, transfer of excitation energy, energy conversion, electron transfer from water to NADP + , ATP generation and a series of enzymatic reactions that assimilate carbon dioxide and synthesize carbohydrate.

Photosynthesis has a unique place in the history of plant science, as its central concepts were established by the middle of the last century, and the detailed mechanisms have since been elucidated. For example, measurements of photosynthetic efficiency (quantum yield) at different wavelengths of light (Emerson and Lews 1943 ) led to the insight that two distinct forms of Chl must be excited in oxygenic photosynthesis. These results suggested the concept of two photochemical systems. The reaction center pigments of PSII and PSI (P680 and P700, respectively) were found by studying changes in light absorbance in the red region (Kok 1959 , Döring et al. 1969 ). Chls with absorbance maxima corresponding to these specific wavelengths were proposed as the final light sink. These Chls were shown to drive electron transfer by charge separation. The linkage of electron transfer and CO 2 assimilation was suggested by studies on Hill oxidant (Hill 1937 ). A linear electron transport system with two light-driven reactions (Z scheme) was proposed based upon observations of the redox state of cytochromes (Hill and Bendall 1960 , Duysens et al. 1961 ), and photophosphorylation was found to be associated with thylakoid fragments (Arnon et al. 1954 ). The metabolic pathway that assimilates carbon by fixation of CO 2 was discovered by Calvin's group who used 14 CO 2 radioactive tracers in the 1950s (Bassham and Calvin 1957 ). This was the first significant discovery in biochemistry made using radioactive tracers. The primary reaction of CO 2 fixation is catalyzed by Rubisco (Weissbach et al. 1956 ), initially called Fraction 1 protein (Wildman and Bonner 1947 ). Rubisco is the most abundant protein in the world, largely because it is also the most inefficient with the lowest catalytic turnover rate (1–3 s –1 ). Another CO 2 fixation pathway was then found in sugarcane (Kortschak et al. 1964, Hatch and Slack 1965) and named C 4 photosynthesis.

Although photosynthesis plays the central role in the energy metabolism of plants, historically there have not been strong interactions between photosynthesis research and other fields of plant science. Many techniques and tools developed for photosynthesis research have not been widely used in other fields because they were developed to examine phenomena unique to photosynthesis. For example, excitation energy transfer and charge separation are fundamental but unique processes of photosynthesis. Another reason for the historic isolation of photosynthesis research within plant science is that it was long believed that CO 2 fixation and carbohydrate production are the sole function of photosynthesis, with carbohydrates representing the only link between photosynthesis and other biological phenomena.

However, this situation has begun to change. Recent research has revealed that photosynthesis is closely related to a variety of other physiological processes. It is a major system for controlling the redox state of cells, playing an important role in regulating enzyme activity and many other cellular processes (Buchanan and Balmer 2005 , Hisabori et al. 2007 ). Photosynthesis also generates reactive oxygen species, which are now appreciated as being regulatory factors for many biological processes rather than inevitable by-products of photosynthesis (Wagner et al. 2004 , Beck 2005 ). Precursor molecules of Chl, which are a major component of photosynthesis, act as a chloroplast-derived signal, and are involved in regulating the cell cycle (Kobayashi et al. 2009 ). In light of this new information, it seems important to re-evaluate the function(s), both potential and demonstrated, of photosynthesis from a variety of view points. Photosynthesis research now employs the methods and tools of molecular biology and genetics, which are central methods for plant science in general. Meanwhile, Chl fluorescence and gas exchange measurements, developed especially for photosynthesis research, are now widely used in stress biology and ecology.

Photosynthesis research also contributes to our understanding of ecological phenomena and even the global environments (Farquhar et al. 1980 , de Pury and Farquhar 1997 , Monsi and Saeki 2005 ). Indeed, photosynthesis is now an integral component of simulation models used to predict the future of our planet. Improving the efficiency of photosynthesis by artificial modification of photosynthetic proteins and pathways has long been considered impossible or at best problematic, because, over evolutionary time, photosynthesis has become complex and tightly regulated. However, recent advances have made it possible to manipulate photosynthesis using molecular genetic technology (Andrews and Whiney 2003 , Raines 2006 ). These advances may have positive influences on crop productivity (Parry et al. 2007 ) as photosynthetic rates have frequently been correlated with biomass accretion (Kruger and Volin 2006 ). Thus, we can expect many more novel concepts to be added to this history of photosynthetic research.

As photosynthesis research tackles new challenges, we should also continue to re-evaluate past research. Oxygen evolution, energy dissipation and cyclic electron transport are crucial processes during photosynthesis, yet their mechanisms still remain to be clarified. We have very limited knowledge of the formation and degradation of photosynthetic apparatus. Also, although photosynthesis plays a central role in C and N metabolism in plants, we do not yet understand how potential photosynthesis is related to crop productivity.

Plant and Cell Physiology would like to contribute to the development of novel concepts, pioneering new fields and solving the unanswered questions of photosynthesis. This special issue covers a wide range of topics in photosynthesis research. Terashima et al. (pp. 684–697) readdress the enigmatic question of why leaves are green. They show that the light profile through a leaf is steeper than that of photosynthesis, and that the green wavelengths in white light are more effective in driving photosynthesis than red light. Evans (pp. 698–706) proposes a new model using Chl fluorescence to explore modifications in quantum yield with leaf depth. This new multilayered model can be applied to study variations in light absorption profiles, photosynthetic capacity and calculation of chloroplastic CO 2 concentration at different depths through the leaf.

Singlet oxygen, 1 O 2 , is produced by the photosystem and Chl pigments. 1 O 2 not only causes physiological damage but also activates stress response programs. The flu mutant of Arabidopsis thaliana overaccumulates protochlorophyllide that upon illumination generates singlet oxygen, causing growth cessation and cell death. Coll et al. (pp. 707–718) have isolated suppressor mutants, dubbed ‘singlet oxygen-linked death activator’ (soldat), that specifically abrogate 1 O 2 -mediated stress responses in young flu seedlings, and they discuss the processes of acclimation to stresses. Phephorbide a is a degradation product of Chl and one of the most powerful photosensitzing molecules. Mutants defective in pheophorbide a oxygenase, which converts phephorbide a to open tetrapyrrole, accumulate pheophorbide a and display cell death in a light-dependent manner. Hirashima et al. (pp. 719–729) report that pheophorbide a is involved in this light-independent cell death.

Plants regulate the redox level of the plastoquinone pool in response to the light environment. In acclimation to high-light conditions, the redox level is kept in an oxidized state by the plastoquinone oxidation system (POS). Miyake et al. (pp. 730–743) investigated the mechanism of POS using the Chl fluorescence parameter, qL.

Nagai and Makino (pp. 744–755) examine in detail the differences between rice and wheat, the two most commercially important crops, in the temperature responses of CO 2 assimilation and plant growth. They find that the difference in biomass production between the two species at the level of the whole plant depends on the difference in N-use efficiency in leaf photosynthesis and growth rate. Sage and Sage (pp. 756–772) examine chlorenchyma structure in rice and related Oryza species in relation to photosynthetic function. They find that rice chlorenchyma architecture includes adaptations to maximize the scavenging of photorespired CO 2 and to enhance the diffusive conductance of CO 2 . In addition, they consider that the introduction of Kranz anatomy does not require radical anatomical alterations in engineering C 4 rice.

Bioinformatics has become a powerful tool, especially in photosynthetic research, because photosynthetic organisms have a wide taxonomic distribution among prokaryotes and eukaryotes. Ishikawa et al. (pp. 773–788) present the results of a pilot study of functional orthogenomics, combining bioinformatic and experimental analyses to identify nuclear-encoded chloroplast proteins of endosymbiontic origin (CRENDOs). They conclude that phylogenetic profiling is useful in finding CPRENDOs, although the physiological functions of orthologous genes may be different in chloroplasts and cyanobacteria.

We hope you enjoy this special issue, and would like to invite you to submit more excellent papers to Plant and Cell Physiology in the field of photosynthesis.

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The important role of photosynthesis.

Bill Cook, Michigan State University Extension - April 09, 2013

Photosynthesis is not just about oxygen production it is also about energy production.

Most people would agree that photosynthesis is a great thing. I’ve never heard anyone argue against it. However, some folks have missed the purpose of photosynthesis. It’s not oxygen production.

The primary function of photosynthesis is to convert solar energy into chemical energy and then store that chemical energy for future use. For the most part, the planet’s living systems are powered by this process. It’s not particularly efficient by human engineering standards, but it does the job. Photosynthesis happens in regions of a cell called chloroplasts. The chemistry and physics are complex.

It’s a bit humbling to consider that the energy in our bodies travels 93 million miles in a little more than eight minutes, and that life has tapped into that energy stream. For a short time that energy is tied up in biological systems before it continues on its merry way into the dark of space.

In essence, green plants take carbon, hydrogen and oxygen from the molecules of carbon dioxide and water, and then recombine them into a new molecule called glucose. This happens in the presence of sunlight, of course. Energy is stored in the bonds of the glucose molecule. Glucose is a fairly simple sugar, easy to break down. Ever wonder why kids bounce off the walls and ceilings soon after a good dose of sugar?

Chemically speaking, the inputs to photosynthesis are six carbon atoms, 12 hydrogen atoms and 18 oxygen atoms. Glucose uses six carbon, 12 hydrogen, and six oxygen molecules. Simple math shows 12 leftover oxygen atoms, or six oxygen molecules. Oxygen atoms prefer mates.

Interestingly, and not coincidentally, the process of respiration breaks apart the glucose molecule. Respiration occurs in the cells of nearly all living things. The released energy is then used for all sorts of metabolic activity, including the energy that you are using to read this article. Respiration happens in regions of a cell called mitochondria. The chemical reactions are the reverse of photosynthesis, using a glucose molecule and six oxygen molecules (12 atoms) as inputs. Energy is released along with some carbon dioxide and water.

But this is enough chemistry.

Trees and other green plants practice respiration, too, just like animals, but they also practice photosynthesis. This is why ecologists categorize green plants as “producers” and most every other life form as a “consumer.” It’s about the energy. OK, there are decomposers, too, but that’s another story and they’re still dependent upon the energy captured by the producers.

Oxygen is a byproduct of photosynthesis and, correspondingly, carbon dioxide the byproduct of respiration. Trees are often credited as the major oxygen generator for the planet, but that would be false. Most of the planet is covered with water and the collective photosynthesis of lowly algae is the true oxygen machine.

Nevertheless, trees and forests are, indeed, significant oxygen producers. However, if oxygen was the only benefit of trees and forests, we could easily live without them. And some forests actually produce more carbon dioxide than oxygen. Fortunately, the benefits of both trees and forests extend far beyond something as narrow as oxygen production.

Much of the basic structural material of plants and wood is cellulose, which is an especially complex sugar. The constituent molecules of carbon, hydrogen and oxygen can be recombined to form lots of useful chemicals such as ethanol, perfumes, bioplastics, clothing fabrics and a range of industrial ingredients. It’s generally agreed that sources from within renewable living ecosystems have distinct advantages over using the ancient materials that make up fossil fuels.

Plants and photosynthesis are the basis of fossil fuels, too, but from millions and millions of years ago. Bringing huge volumes of those molecules back into living ecosystems has a few drawbacks that science has gotten pretty good at measuring and describing.

Trees, forests, forest soils and forest products are mighty important in the cycling of carbon and the relative size of various carbon pools. There are other elements that also cycle through forests. Science has a pretty good handle on these relationships, too. Michigan residents might do well to place a bit more weight on these service benefits of trees, forests, and forest management.

As for photosynthesis itself, maybe it’s better if we think more about the energy capture and less about the oxygen production.

This article was published by Michigan State University Extension . For more information, visit https://extension.msu.edu . To have a digest of information delivered straight to your email inbox, visit https://extension.msu.edu/newsletters . To contact an expert in your area, visit https://extension.msu.edu/experts , or call 888-MSUE4MI (888-678-3464).

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Why Photosynthesis is Essential to Life on Earth

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Life is powered by sunlight. The energy used by most living cells comes ultimately from the sun. Plants, algae, and some bacteria use energy from sunlight, particularly blue and red wavelengths, to build molecules which later can be split through cellular respiration to retrieve some of that energy. Storing energy in molecules and then oxidizing those molecules to retrieve the stored energy maintains all life on Earth. Plants are often called ‘producers’ because they produce energy-storing molecules used by almost all other organisms on Earth. By eating plants, herbivores ‘steal’ these energy-storing molecules to maintain their own life processes. By eating animals, carnivores ‘plunder’ the molecules that store the energy oringinally captured by plants. By feeding on dead tissue, decomposers exploit whatever molecules remain in the dead the plants, herbivores, and carnivores. Ultimately, the process of photosynthesis is the most important chemical reaction on Earth. As biologists are well aware, “Roses are red, violets are blue. If the green plants go, then so do you!” Pardon Our Interruption As you were browsing something about your browser made us think you were a bot. There are a few reasons this might happen: You've disabled JavaScript in your web browser. You're a power user moving through this website with super-human speed. You've disabled cookies in your web browser. A third-party browser plugin, such as Ghostery or NoScript, is preventing JavaScript from running. Additional information is available in this support article . To regain access, please make sure that cookies and JavaScript are enabled before reloading the page. 282 IMAGES Photosynthesis Explained Why Photosynthesis Is Essential To Life On Earth: [Essay Example], 326 15 Reasons Why Photosynthesis Is Important? What Is Photosynthesis? 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Oxygenic photosynthesis is a very important process, not only because it is the source of our food, fibre and many useful substances, but also because almost all life on the Earth depends on it, either directly or indirectly. Recent advances in understanding and improving photosynthesis Introduction. Photosynthesis is the chemical reaction that sustains most life on Earth. Since the description of the Hill reaction and the Calvin-Benson cycle 1-3, knowledge about their components, regulation, and limitations experienced a vertiginous increase.It is widely known that plants have important handicaps related to photosynthesis. Photosynthesis Unlike photosynthesis, aerobic respiration is an exergonic process (negative ΔG°) with the energy released being used by the organism to power biosynthetic processes that allow growth and renewal, mechanical work (such as muscle contraction or flagella rotation) and facilitating changes in chemical concentrations within the cell (e.g. accumulation of nutrients and expulsion of waste). Photosynthetic Research in Plant Science Photosynthesis is a highly regulated, multistep process. It encompasses the harvest of solar energy, transfer of excitation energy, energy conversion, electron transfer from water to NADP +, ATP generation and a series of enzymatic reactions that assimilate carbon dioxide and synthesize carbohydrate. Photosynthesis has a unique place in the ... Photosynthesis Overview, Benefits & Importance Photosynthesis takes place in specialized structures called chloroplasts, which are small organelles within plant cells. These chloroplasts contain a number of important chemicals and structures ... The important role of photosynthesis The primary function of photosynthesis is to convert solar energy into chemical energy and then store that chemical energy for future use. For the most part, the planet's living systems are powered by this process. It's not particularly efficient by human engineering standards, but it does the job. Why Photosynthesis is Essential to Life on Earth The plants requires glucose for their growth and energy. All the living people and animals on the earth depend on fats, proteins, and carbohydrates to obtain their energy, that is what makes this is one of the reasons photosynthesis is essential to life on earth. Another product of photosynthesis is oxygen which we need to breathe. Biology Laboratory Manual By feeding on dead tissue, decomposers exploit whatever molecules remain in the dead the plants, herbivores, and carnivores. Ultimately, the process of photosynthesis is the most important chemical reaction on Earth. As biologists are well aware, "Roses are red, violets are blue. If the green plants go, then so do you!". How is photosynthesis essential to life on earth? Photosynthesis is a chemical process whereby plants and algae that contain chlorophyll capture radiant energy from the sun, and use carbon dioxide and water from the environment to then convert the sunlight to food (glucose), while at the same time creating oxygen and water as byproducts. Of course, we humans get the benefit of all that food ... assignment essay homework paper phd report resume review speech thesis