essay for photosynthesis

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.

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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

Photosynthesis is a process used by plants, algae, and some bacteria to turn sunlight, water, and carbon dioxide into food and oxygen. Think of it like a recipe that plants use to make their own food. This happens in the leaves of plants, which have a green substance called chlorophyll.

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

To make their food, plants need three main things: sunlight, water, and carbon dioxide. Sunlight is the energy plants use to create their food. They get water from the ground through their roots. Carbon dioxide, a gas found in the air, is taken in through tiny holes in the leaves called stomata.

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 also helps the environment. Plants take in carbon dioxide, which is a gas that can make the Earth warmer when there is too much of it in the air. By using carbon dioxide to make food, plants help keep the air clean and the Earth’s temperature just right.

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.

In conclusion, photosynthesis is a vital process that allows plants to make food and oxygen using sunlight, water, and carbon dioxide. It is the foundation of the food chain and has a big impact on the environment and our lives. Understanding photosynthesis helps us appreciate how important plants are and why we need to take care of them and the environment they live in.

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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.

Biology, Earth Science, Chemistry, Ecology

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.

On this page:

What is photosynthesis, earth system model about photosynthesis, explore the earth system, investigate, links to learn more.

For the classroom:

• Teaching Resources

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

84 Photosynthesis Essay Topic Ideas & Examples

🏆 best photosynthesis topic ideas & essay examples, 👍 good essay topics on photosynthesis, 📌 simple & easy photosynthesis essay titles, ❓ photosynthesis essay questions.

• Similarities and Differences of Photosynthesis and Cellular Respiration The starting reactants are the light of the sun, carbon dioxide, and water. Both processes are required to support the being of plants and animals.
• Photosynthetic Pigments in the Chlorophyll Extract Write a concise figure legend with a minimal number of words that allows the reader to understand what he sees. Note: Do not repeat all details of the methods in the legend!
• Photosynthesis as a Biological Process During the process water from the soil is taken up by the roots all the way to the leaves via the xylem.
• Planting Bamboo: The Role of Photosynthesis Lastly, I think bamboo’s photosynthetic exceptionalism is manifested in the fact that light is one of the most important factors of photosynthesis, as it is a source of the necessary radiant energy for biochemical processes.
• Photosynthetic Organism: Aesculus Hippocastanum Aesculus hippocastanum is not picky about food and can grow for a long time in the shade and on sub-hourly soils.
• Research and Experiments: Molecules in Food, Photosynthesis The saliva of humans and some mammals contains amylase: the enzyme that begins the chemical process of food digestion. I took the pollution of the environment with heavy metals and the effect on photosynthesis.
• Runaway Photosynthesis and Its Effects on Life in the Universe According to extant literature, runaway photosynthesis describes a scenario in which the machinery of photosynthesis could be adversely affected by shifts in the environment, leading to the crumbling of all ecosystems that depend on it […]
• The Photosynthetic Rate in Terrestrial Plants In this regard, one is expected to establish the necessities of photosynthesis and discover the requirement of carbon dioxide and light for oxygen evolution.
• Recent Advances in Artificial Photosynthesis Specifically, scientists strive to use the known and change them into “functional, efficient, synthetic systems that will tap the endless supply of energy coming from the sun”.[1] Researchers believe that artificial photosynthesis can work on […]
• The Complexity of Photosynthesis and Respiration The dark reaction which happens in the absence of light but inside the stroma utilizes CO2 and ATP to form sugars in what is referred to as the Calvin Cycle.
• Photosynthesis & Fermentation: Plant Cellular Metabolism The process of photosynthesis and fermentation was the main focus of the practical exercise. The role of the fermentation process to provide energy for the yeast cells is shown in this experiment.
• “Why Study Photosynthesis?” by Devens Gust By revealing the information about the details of photosynthesis, Gust encourages people to investigate and to develop the innovated techniques thus applying them to the problem solution in the different spheres of life.
• Photosynthesis Rate Determination From the Oxygen Gas Evolution The results showed that the rate of release of oxygen determined the rate of rising of the leaf segment subject to utilization of carbon dioxide proving that the evolution of oxygen could be used to […]
• Photosynthesis and Cellular Respiration The reactants of the respiration are glucose circulating in the blood and oxygen obtained from breathing, while the product is ATP.
• The Photosynthesis Rate of Waterweed In this experiment, it was hypothesized that the rate of photosynthesis rises with an increase in the concentration of carbon dioxide and went on to test the hypothesis.
• Environmental Studies: Photosynthesis Concept Similarly, the effect of carbon dioxide and light intensity on the rate of photosynthesis was also measured by varying the concentration and distance of carbon dioxide respectively.
• Comparing Photosynthesis and Cellular Respiration
• How Light Intensity Affects the Rate of Photosynthesis
• The Connection Between Carbon Dioxide and the Rate of Photosynthesis
• Could Earth Survive With Artificial Photosynthesis
• Light-Dependent and Light-Independent Reactions of Photosynthesis
• Differences and Similarities Between Respiration and Photosynthesis
• How Different Coloured Rays of Light Affect Photosynthesis
• Factors Affecting the Rate of Photosynthesis in Leafs
• General Information About Hydrilla and Photosynthesis
• How Are the Reactants and Products of Photosynthesis
• How Can the Light Reactions of Photosynthesis Be Improved in Plants
• The Evolution of Oxygen-Releasing Photosynthesis
• How Does Photosynthesis Affect Our Own
• Interactions Between Circadian Clocks and Photosynthesis
• The Limiting Factors of Photosynthesis
• Investigating the Chemical Equation for Photosynthesis
• Photosynthesis: Adenosine Triphosphate and Energy
• Photosynthesis and Its Effect on the Environment
• Components of Natural Photosynthetic Apparatus in Solar Cells
• Photosynthesis: Process That Converts Light Energy Organisms Into Chemical Energy
• The Basic Needs for Photosynthesis
• The Environmental Effects and Limitations of Drought on Photosynthesis
• The Experiment That Led to the Discovery of Photosynthesis
• The Floating Leaf Disk Assay for Investigating Photosynthesis
• The Stages and Special Conditions Needed for Photosynthesis
• What Are the Chemical Products of Photosynthesis
• The Investigation of the Color of Light and Its Effect on Photosynthesis
• What Role Does Molecular Oxygen Play in Photosynthesis
• Investigation of How Light Affects Photosynthesis
• Finding the Factor That Can Increase the Rate of Photosynthesis
• An In-Depth Overview of the Process of Photosynthesis in Plants
• Experimenting on the Rate of Photosynthesis in Chloroplasts
• The Features of the Process of Photosynthesis
• The Positive Charges of Electron and Intercellular Fluid in the Photosynthesis
• The Determination of Rates of Photosynthesis Through Chloroplasts
• The Impact of Different Types of Environments on Photosynthesis
• The Process of Marine Photosynthesis and Factors Affecting It
• Peculiarities of Photosynthesis of Sugar Plants
• Photosynthesis Concept, Equation & Steps
• How Does Photosynthesis Yield Sugar
• Why Is Photosynthesis Important?
• Who Discovered Photosynthesis?
• Why Is Photosynthesis Good for Plants?
• What Factors Affect Photosynthesis?
• Why Is It Impossible to Create Photosynthesis in Artificial Conditions?
• In Which Plant Organs Does Photosynthesis Take Place?
• What Are the Different Stages of Photosynthesis?
• Can Photosynthesis Happen Without Light?
• Why Is It Not Possible for Humans to Photosynthesize?
• What Is the Primary Purpose of Photosynthesis?
• What Are the Products of Photosynthesis?
• What Is Rate of Photosynthesis?
• Where Does the Energy for Photosynthesis Come From?
• Which Organelle Is Responsible for Photosynthesis?
• What Are the Main Steps of Photosynthesis?
• How Does Photosynthesis Yield Sugar?
• What Are the Developmental Stages of Photosynthesis?
• How Carbon Dioxide Can Affect the Rate of Photosynthesis?
• Why Can Photosynthesis Not Occur at Very High Temperatures?
• What Would Happen if Photosynthesis Stopped Happening on Earth?
• What Is the Maximum Temperature for Photosynthesis?
• Which Element Increases Photosynthesis in Plant?
• Which Plants Do Photosynthesis at Night?
• What Gas Is Released During Photosynthesis?
• How Are the Reactants and Products of Photosynthesis?
• How Can the Light Reactions of Photosynthesis Be Improved in Plants?
• How Did the Evolution of Oxygen-Releasing Photosynthesis?
• Sugar Paper Topics
• Climate Change Titles
• Deforestation Research Ideas
• Genetic Engineering Topics
• Energy Essay Ideas
• Renewable Energy Essay Titles
• Solar Energy Essay Ideas
• Environment Research Topics
• Chicago (A-D)
• Chicago (N-B)

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8.1 Overview of Photosynthesis

Learning objectives.

By the end of this section, you will be able to do the following:

• Explain the significance of photosynthesis to other living organisms
• Describe the main structures involved in photosynthesis
• Identify the substrates and products of 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 from sunlight and convert it into chemical compounds (carbohydrates) that every organism uses to power its metabolism. It is also a source of oxygen necessary for many living organisms. In brief, the energy of sunlight is “captured” to energize electrons, whose energy is 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 350 to 200 million years ago during the Carboniferous Period.

Plants, algae, and a group of bacteria called cyanobacteria are the only organisms capable of performing photosynthesis ( Figure 8.2 ). 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. For this reason, they are referred to as chemoautotrophs .

The importance of photosynthesis is not just that it can capture sunlight’s energy. After all, a lizard sunning itself on a cold day can use the sun’s energy to warm up in a process called behavioral thermoregulation . In contrast, photosynthesis is vital because it evolved as a way to store the energy from solar radiation (the “photo-” part) to energy 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 8.3 ), the wolf is at the end of an energy path that went from nuclear reactions on the surface of the sun, to visible light, to photosynthesis, to vegetation, to deer, and finally to the wolf.

Main Structures and Summary of Photosynthesis

Photosynthesis is a multi-step process that requires specific wavelengths of visible sunlight, carbon dioxide (which is low in energy), and water as substrates ( Figure 8.4 ). After the process is complete, it releases oxygen and produces glyceraldehyde-3-phosphate (G3P), as well as simple carbohydrate molecules (high in energy) that can then 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.

The following is the chemical equation for photosynthesis ( Figure 8.5 ):

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.

Basic Photosynthetic Structures

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 due to high temperatures on the upper surface of the leaf. 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 mostly in the mesophyll. Chloroplasts have a double membrane envelope (composed of an outer membrane and an inner membrane), and are ancestrally derived from ancient free-living cyanobacteria. 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 8.6 , 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).

Visual Connection

On a hot, dry day, the guard cells of 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 drives 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, however, 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 8.7 illustrates the components inside the chloroplast where the light-dependent and light-independent reactions take place.

Link to Learning

Click the link to learn more about photosynthesis.

Everyday Connection

Photosynthesis at the grocery store.

Major grocery stores in the United States are organized into departments, such as dairy, meats, produce, bread, cereals, and so forth. Each aisle ( Figure 8.8 ) contains hundreds, if not thousands, of different products for customers to buy and consume.

Although there is a large variety, each item ultimately can be linked 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) are derived from “algae” (unicellular plant-like organisms, and cyanobacteria). 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.

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Home — Essay Samples — Science — Light — Photosynthesis Process

Photosynthesis Process

• Categories: Light Photosynthesis

About this sample

Words: 423 |

Published: Feb 12, 2019

Words: 423 | Page: 1 | 3 min read

Works Cited

• Campbell, N. A., & Reece, J. B. (2008). Photosynthesis and cellular respiration. In Biology (8th ed., pp. 190-220). Benjamin-Cummings Publishing Company.
• Taiz, L., & Zeiger, E. (2010). Photosynthesis: Carbon reactions. In Plant physiology (5th ed., pp. 174-207). Sinauer Associates.
• Raven, P. H., Evert, R. F., & Eichhorn, S. E. (2016). Photosynthesis and respiration. In Biology of Plants (8th ed., pp. 186-229). W. H. Freeman and Company.
• Niyogi, K. K. (1999). Photoprotection revisited: Genetic and molecular approaches. Annual Review of Plant Physiology and Plant Molecular Biology, 50, 333-359. doi:10.1146/annurev.arplant.50.1.333
• Siedow, J. N., & Day, D. A. (2000). Respiration and photorespiration. In Plant physiology (3rd ed., pp. 500-548). Academic Press.
• Allen, J. F. (2002). Photosynthesis and cellular respiration considered as coupled redox cycles: A chemiosmotic bridge linking two epochs. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 357(1426), 707-717. doi:10.1098/rstb.2001.0993
• Geigenberger, P. (2003). Response of plant metabolism to too little oxygen. Current Opinion in Plant Biology, 6(3), 247-256. doi:10.1016/S1369-5266(03)00038-8
• Foyer, C. H., & Noctor, G. (2005). Redox homeostasis and antioxidant signaling: A metabolic interface between stress perception and physiological responses. The Plant Cell, 17(7), 1866-1875. doi:10.1105/tpc.105.033589
• Sharkey, T. D. (2005). Effects of moderate heat stress on photosynthesis: Importance of thylakoid reactions, rubisco deactivation, reactive oxygen species, and thermotolerance provided by isoprene. Plant, Cell & Environment, 28(3), 269-277. doi:10.1111/j.1365-3040.2005.01324.x
• Sweetlove, L. J., & Fernie, A. R. (2018). The impact of oxidative stress on metabolism: A compartmental analysis. Frontiers in Plant Science, 9, 1647. doi:10.3389/fpls.2018.01647

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Photosynthesis

Matthew p johnson.

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Correspondence: Matthew P. Johnson (email [email protected] ).

Received 2016 Jan 14; Revised 2016 Jul 22; Accepted 2016 Jul 26; Issue date 2016 Oct 31.

This is an open access article published by Portland Press Limited on behalf of the Biochemical Society and distributed under the Creative Commons Attribution Licence 4.0 (CC BY) .

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.

Keywords: membrane, photosynthesis, thylakoid

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 ).

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 ).

Figure 1. The global carbon cycle.

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.

Figure 2. Location of the photosynthetic machinery.

( 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.

Figure 3. Division of labour within the chloroplast.

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 ).

Figure 4. The photosynthetic electron and proton transfer chain.

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.

Figure 5. Z-scheme of photosynthetic electron transfer.

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 + .

Box 2. 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 ).

Figure 6. Major photosynthetic pigments in plants.

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.

Figure 7. Basic absorption spectra of the major chlorophyll and carotenoid pigments found in plants.

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.

Figure 8. Jablonski diagram of chlorophyll showing the possible fates of the S 1 and S 2 excited states and timescales of the transitions involved.

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.

Figure 9. Basic mechanism of excitation energy transfer between chlorophyll molecules.

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.

Figure 10. Basic structure of a photosystem.

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.

Figure 11. Basic structure of the PSII–LHCII supercomplex from spinach.

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

Figure 12. S-state cycle of water oxidation by the manganese cluster (shown as circles with roman numerals representing the manganese ion oxidation states) within the PSII oxygen-evolving complex.

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.

Figure 13. Basic structure of the PSI–LHCI supercomplex from pea.

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.

Figure 14. Cytochrome b 6 f complex.

( 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).

Figure 15. Lateral heterogeneity in thylakoid membrane organization.

( 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.

Figure 16. The Calvin–Benson cycle.

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).

Figure 17. Rubisco.

( 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.

Figure 18. Diagram of a C 4 plant leaf showing Kranz anatomy.

Figure 19. The C 4 pathway (NADP + –malic enzyme type) for fixation of CO 2 .

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

adenosine diphosphate

adenosine triphosphate

carbohydrate

cytochrome b 6 f

dihydroxyacetone phosphate

excitation energy transfer

ferredoxin–NADP + reductase

glyceraldehyde 3-phosphate

light-harvesting complex

nicotinomide–adenine dinucleotide phosphate

phosphoenolpyruvate

inorganic phosphate

reaction centre

ribulose-1,5-bisphosphate carboxylase/oxygenase

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. [ DOI ] [ PMC free article ] [ PubMed ] [ 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. [ DOI ] [ PubMed ] [ Google Scholar ]
• Raines C. The Calvin cycle revisited. Photosynth. Res. 2003;75:1–10. doi: 10.1023/A:1022421515027. [ DOI ] [ PubMed ] [ 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. [ DOI ] [ PubMed ] [ Google Scholar ]
• Sage R.F. The evolution of C4 photosynthesis. New Phytol. 2004;161:341–370. doi: 10.1111/j.1469-8137.2004.00974.x. [ DOI ] [ PubMed ] [ Google Scholar ]
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