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The vast majority of energy found within any ecosystem on Earth originates from the Sun itself and is transferred between organisms through complex food webs. Photosynthetic organisms such as plants and algae form the basis of these webs by converting sunlight, water, and carbon dioxide into usable energy. That energy can then be utilized by the photosynthetic organism, which itself may be consumed by other organisms for their own energy requirements. This handout will focus on explaining the steps involved in the most common version of photosynthesis, known as C3 photosynthesis

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What Is Photosynthesis? | Biology | FuseSchool

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Photosynthetic reactions can be anoxygenic, thus they do not produce oxygen.

Learning Objectives

Discuss the characteristics that classify a specific type of chlorophototrophy as anoxygenic photosynthesis

Key Points

Anoxygenic photosynthesis produces cellular energy ( ATP ), without oxygen as a by-product.

As opposed to eukaryotic organisms, which rely on chlorophylls for photosynthesis, anoxygenic organisms rely on bacteriochlorophylls.

The electron transport chain of anoxygenic phototrophs is cyclic, meaning the electrons used during photosynthesis are fed back into the system, therefore no electrons are left over to oxidize water into oxygen.

Key Terms

electron transport chain: An electron transport chain (ETC) couples electron transfer between an electron donor (such as NADH) and an electron acceptor (such as O2) with the transfer of H+ ions (protons) across a membrane. The resulting electrochemical proton gradient is used to generate chemical energy in the form of adenosine triphosphate (ATP). Electron transport chains are the cellular mechanisms used for extracting energy from sunlight in photosynthesis and also from redox reactions, such as the oxidation of sugars (respiration).

electron donor: An electron donor is a chemical entity that donates electrons to another compound. It is a reducing agent that, by virtue of its donating electrons, is itself oxidized in the process.

anoxygenic: That does not involve the production of oxygen

Phototrophy is the process by which organisms trap light energy (photons) and store it as chemical energy in the form of ATP and/or reducing power in NADPH. There are two major types of phototrophy: chlorophyll-based chlorophototrophy and rhodopsin-based retinalophototrophy. Chlorophototrophy can further be divided into oxygenic photosynthesis and anoxygenic phototrophy. Oxygenic and anoxygenic photosynthesizing organisms undergo different reactions either in the presence of light or with no direct contribution of light to the chemical reaction (colloquially called "light reactions" and "dark reactions", respectively).

Figure: Green d winogradsky: A column containing green sulfur bacteria which uses anoxygenic photosynthesis.

Anoxygenic photosynthesis is the phototrophic process where light energy is captured and converted to ATP, without the production of oxygen. Water is therefore not used as an electron donor. There are several groups of bacteria that undergo anoxygenic photosynthesis: Green sulfur bacteria, green and red filamentous anoxygenic phototrophs (FAPs), phototrophic purple bacteria, phototrophic Acidobacteria, and phototrophic heliobacteria. Anoxygenic phototrophs have photosynthetic pigments called bacteriochlorophylls (similar to chlorophyll found in eukaryotes). Bacteriochlorophyll a and b have wavelengths of maximum absorption at 775 nm and 790 nm, respectively in ether. In vivo however, due to shared extended resonance structures, these pigments were found to maximally absorb wavelengths out further into the near-infrared. Bacteriochlorophylls c-g have the corresponding "peak" absorbance at more blue wavelengths when dissolved in an organic solvent, but are similarly red-shifted within their natural environment (with the exception of bacteriochlorophyll f, which has not been naturally observed).Unlike oxygenic phototrophs, anoxygenic photosynthesis only functions using (by phylum) either one of two possible types of photosystem. This restricts them to cyclic electron flow and are therefore unable to produce O ₂ from the oxidization of H ₂ O.

The cyclic nature of the electron flow is typified in purple non-sulfur bacteria. The electron transport chain of purple non-sulfur bacteria begins when the reaction centre bacteriochlorophyll pair, P870, becomes excited from the absorption of light. Excited P870 will then donate an electron to Bacteriopheophytin, which then passes it on to a series of electron carriers down the electron chain. In the process, it will generate a proton motor force (PMF) which can then be used to synthesize ATP by oxidative phosphorylation. The electron returns to P870 at the end of the chain so it can be used again once light excites the reaction-center. Therefore electrons are not left over to oxidize H ₂O into O ₂.

LICENSES AND ATTRIBUTIONS

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Photosynthetic reactions can be anoxygenic, thus they do not produce oxygen. Learning Objectives Discuss the characteristics that classify a specific type of chlorophototrophy as anoxygenic photosynthesis Anoxygenic photosynthesis produces cellular energy ( ATP ), without oxygen as a by-product. As opposed to eukaryotic organisms, which rely on chlorophylls for photosynthesis, anoxygenic organisms rely on bacteriochlorophylls.

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Photosynthesis in organisms (article) | Khan Academy

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Learn for free about math, art, computer programming, economics, physics, chemistry, biology, medicine, finance, history, and more. Khan Academy is a nonprofit with the mission of providing a free, world-class education for anyone, anywhere.

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This video demonstrate the process by which green plants use sunlight to synthesize nutrients from carbon dioxide and water and generate oxygen as a by-product with the involvement of green pigment chlorophyll.

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Able to get an overall idea on Photosynthesis

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Introduction

Have you hugged a tree lately? If not, you might want to give it some thought. You, along with the rest of the human population, owe your existence to plants and other organisms that capture light. In fact, most life on Earth is possible because the sun provides a continuous supply of energy to ecosystems.

All organisms, including humans, need energy to fuel the metabolic reactions of growth, development, and reproduction. But organisms can't use light energy directly for their metabolic needs. Instead, it must first be converted into chemical energy through the process of photosynthesis.

What is photosynthesis?

Photosynthesis is the process in which light energy is converted to chemical energy in the form of sugars. In a process driven by light energy, glucose molecules (or other sugars) are constructed from water and carbon dioxide, and oxygen is released as a byproduct. The glucose molecules provide organisms with two crucial resources: energy and fixed-organic-carbon.

Energy. The glucose molecules serve as fuel for cells: their chemical energy can be harvested through processes like cellular respiration and fermentation, which generate adenosine triphosphate-, a small, energy-carrying molecule-for the cell's immediate energy needs.

Fixed carbon. Carbon from carbon dioxide-inorganic carbon-can be incorporated into organic molecules; this process is called carbon fixation, and the carbon in organic molecules is also known as fixed carbon. The carbon that's fixed and incorporated into sugars during photosynthesis can be used to build other types of organic molecules needed by cells.

The ecological importance of photosynthesis

Photosynthetic organisms, including plants, algae, and some bacteria, play a key ecological role. They introduce chemical energy and fixed carbon into ecosystems by using light to synthesize sugars. Since these organisms produce their own food-that is, fix their own carbon-using light energy, they are called photoautotrophs (literally, self-feeders that use light).

Humans, and other organisms that can't convert carbon dioxide to organic compounds themselves, are called heterotrophs, meaning different-feeders. Heterotrophs must get fixed carbon by eating other organisms or their by-products. Animals, fungi, and many prokaryotes and protists are heterotrophs.

Besides introducing fixed carbon and energy into ecosystems, photosynthesis also affects the makeup of Earth's atmosphere. Most photosynthetic organisms generate oxygen gas as a byproduct, and the advent of photosynthesis-over billion years ago, in bacteria resembling modern cyanobacteria-forever changed life on Earth. These bacteria gradually released oxygen into Earth's oxygen-poor atmosphere, and the increase in oxygen concentration is thought to have influenced the evolution of aerobic life forms-organisms that use oxygen for cellular respiration. If it hadn't been for those ancient photosynthesizers, we, like many other species, wouldn't be here today!

Photosynthetic organisms also remove large quantities of carbon dioxide from the atmosphere and use the carbon atoms to build organic molecules. Without Earth's abundance of plants and algae to continually suck up carbon dioxide, the gas would build up in the atmosphere. Although photosynthetic organisms remove some of the carbon dioxide produced by human activities, rising atmospheric levels are trapping heat and causing the climate to change. Many scientists believe that preserving forests and other expanses of vegetation is increasingly important to combat this rise in carbon dioxide levels.

Leaves are sites of photosynthesis

Plants are the most common autotrophs in terrestrial-land-ecosystems. All green plant tissues can photosynthesize, but in most plants, but the majority of photosynthesis usually takes place in the leaves. The cells in a middle layer of leaf tissue called the mesophyll are the primary site of photosynthesis.

Small pores called stomata -singular, stoma-are found on the surface of leaves in most plants, and they let carbon dioxide diffuse into the mesophyll layer and oxygen diffuse out.

Each mesophyll cell contains organelles called chloroplasts, which are specialized to carry out the reactions of photosynthesis. Within each chloroplast, disc-like structures called thylakoids are arranged in piles like stacks of pancakes that are known as grana-singular, granum. The membrane of each thylakoid contains green-colored pigments called chlorophylls that absorb light. The fluid-filled space around the grana is called the stroma, and the space inside the thylakoid discs is known as the thylakoid space. Different chemical reactions occur in the different parts of the chloroplast.

The light-dependent reactions and the Calvin cycle

The light-dependent reactions take place in the thylakoid membrane and require a continuous supply of light energy. Chlorophylls absorb this light energy, which is converted into chemical energy through the formation of two compounds, -an energy storage molecule-and -a reduced (electron-bearing) electron carrier. In this process, water molecules are also converted to oxygen gas-the oxygen we breathe!

The Calvin cycle, also called the light-independent reactions, takes place in the stroma and does not directly require light. Instead, the Calvin cycle uses and from the light-dependent reactions to fix carbon dioxide and produce three-carbon sugars-glyceraldehyde-3-phosphate, or G3P, molecules-which join up to form glucose.

Overall, the light-dependent reactions capture light energy and store it temporarily in the chemical forms of and . There, is broken down to release energy, and donates its electrons to convert carbon dioxide molecules into sugars. In the end, the energy that started out as light winds up trapped in the bonds of the sugars.

Photosynthesis vs. cellular respiration

At the level of the overall reactions, photosynthesis and cellular respiration are near-opposite processes. They differ only in the form of energy absorbed or released, as shown in the diagram below.

At the level of individual steps, photosynthesis isn't just cellular respiration run in reverse. Instead, as we'll see the rest of this section, photosynthesis takes place in its own unique series of steps. However, there are some notable similarities between photosynthesis and cellular respiration.

For instance, photosynthesis and cellular respiration both involve a series of redox reactions (reactions involving electron transfers). In cellular respiration, electrons flow from glucose to oxygen, forming water and releasing energy. In photosynthesis, they go in the opposite direction, starting in water and winding up in glucose-an energy-requiring process powered by light. Like cellular respiration, photosynthesis also uses an electron transport chain to make a concentration gradient, which drives synthesis by chemiosmosis.

If those things don't sound familiar, though, don't worry! You don't need to know cellular respiration to understand photosynthesis. Just keep reading and watching, and you'll learn all the ins and outs of this life-sustaining process.

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Intro to photosynthesis (article) | Khan Academy

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How photosynthesis takes place in plants

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How photosynthesis takes place in plants

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how photosynthesis takes place in plants & Process Of Photosynthesis (animated)

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Explains concept of photosynthesis from basic to advance

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Page ID

35693

1. What you'll learn to do: Identify the basic components and steps of photosynthesis
Learning Objectives

2. An Overview of Photosynthesis

1. Solar Dependence and Food Production
Practice Question

2. The Two Parts of Photosynthesis

1. Try It

2. Learning Objectives

3. Light Energy

1. What Is Light Energy?

2. Absorption of Light

3. Understanding Pigments

4. The Light-Dependent Reactions of Photosynthesis

1. Generating an Energy Carrier: ATP

2. Generating Another Energy Carrier: NADPH

1. Learning Objectives

2. Practice Question

5. The Calvin Cycle

1. The Innerworkings of the Calvin Cycle
Try It

2. Photosynthesis in Prokaryotes

1. Learning Objectives

2. Practice Question

6. Summary

7. Check Your Understanding

8. Contributors and Attributions

What you'll learn to do: Identify the basic components and steps of photosynthesis

No matter how complex or advanced a machine, such as the latest cellular phone, the device cannot function without energy. Living things, similar to machines, have many complex components; they too cannot do anything without energy, which is why humans and all other organisms must "eat" in some form or another. That may be common knowledge, but how many people realize that every bite of every meal ingested depends on the process of photosynthesis?

Figure 1. This sage thrasher's diet, like that of almost all organisms, depends on photosynthesis. (credit: modification of work by Dave Menke, U.S. Fish and Wildlife Service)

Learning Objectives

Summarize the process of photosynthesis

Describe how the wavelength of light affects its energy and color

Describe the light-dependent reactions that take place during photosynthesis

Describe the steps and processes in the Calvin Cycle

An Overview of Photosynthesis

All living organisms on earth consist of one or more cells. Each cell runs on the chemical energy found mainly in carbohydrate molecules (food), and the majority of these molecules are produced by one process: photosynthesis. Through photosynthesis, certain organisms convert solar energy (sunlight) into chemical energy, which is then used to build carbohydrate molecules. The energy used to hold these molecules together is released when an organism breaks down food. Cells then use this energy to perform work, such as cellular respiration.

The energy that is harnessed from photosynthesis enters the ecosystems of our planet continuously and is transferred from one organism to another. Therefore, directly or indirectly, the process of photosynthesis provides most of the energy required by living things on earth.

Photosynthesis also results in the release of oxygen into the atmosphere. In short, to eat and breathe, humans depend almost entirely on the organisms that carry out photosynthesis.
Learn more about photosynthesis

Solar Dependence and Food Production

Some organisms can carry out photosynthesis, whereas others cannot. An autotroph is an organism that can produce its own food. The Greek roots of the word autotroph mean "self" ( auto) "feeder" ( troph). Plants are the best-known autotrophs, but others exist, including certain types of bacteria and algae (Figure 2). Oceanic algae contribute enormous quantities of food and oxygen to global food chains. Plants are also photoautotrophs, a type of autotroph that uses sunlight and carbon from carbon dioxide to synthesize chemical energy in the form of carbohydrates. All organisms carrying out photosynthesis require sunlight.

Figure 2. (a) Plants, (b) algae, and (c) certain bacteria, called cyanobacteria, are photoautotrophs that can carry out photosynthesis. Algae can grow over enormous areas in water, at times completely covering the surface. (credit a: Steve Hillebrand, U.S. Fish and Wildlife Service; credit b: "eutrophication&hypoxia"/Flickr; credit c: NASA; scale-bar data from Matt Russell)

Figure 3. The energy stored in carbohydrate molecules from photosynthesis passes through the food chain. The predator that eats these deer is getting energy that originated in the photosynthetic vegetation that the deer consumed. (credit: Steve VanRiper, U.S. Fish and Wildlife Service)

Heterotrophs are organisms incapable of photosynthesis that must therefore obtain energy and carbon from food by consuming other organisms. The Greek roots of the word heterotroph mean "other" ( hetero) "feeder" ( troph), meaning that their food comes from other organisms. Even if the food organism is another animal, this food traces its origins back to autotrophs and the process of photosynthesis. Humans are heterotrophs, as are all animals. Heterotrophs depend on autotrophs, either directly or indirectly. Deer and wolves are heterotrophs. A deer obtains energy by eating plants. A wolf eating a deer obtains energy that originally came from the plants eaten by that deer. The energy in the plant came from photosynthesis, and therefore it is the only autotroph in this example (Figure 3). Using this reasoning, all food eaten by humans also links back to autotrophs that carry out photosynthesis.

Photosynthesis is a multi-step process that requires sunlight, carbon dioxide (which is low in energy), and water as substrates (Figure 4). 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 4. 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 5):

Figure 5. 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 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).

Practice Question

Figure 6. 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?

Show Answer

Levels of carbon dioxide (a necessary photosynthetic substrate) will immediately fall. As a result, the rate of photosynthesis will be inhibited.

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 7 illustrates the components inside the chloroplast where the light-dependent and light-independent reactions take place.

Figure 7. 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 CO₂.

Click the link to learn more about photosynthesis.

Try It

Figure 8. 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 8) 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.

Learning Objectives

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.

Light Energy

Figure 9. Autotrophs can capture light energy from the sun, converting it into chemical energy used to build food molecules. (credit: modification of work by Gerry Atwell, U.S. Fish and Wildlife Service)

How can light be used to make food? It is easy to think of light as something that exists and allows living organisms, such as humans, to see, but light is a form of energy. Like all energy, light can travel, change form, and be harnessed to do work. In the case of photosynthesis, light energy is transformed into chemical energy, which autotrophs use to build carbohydrate molecules. However, autotrophs only use a specific component of sunlight (Figure 9).

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 is referred to as "visible light." The manner in which solar energy travels can be described and measured as waves. Scientists can determine the amount of energy of a wave by measuring its wavelength, the distance between two consecutive, similar points in a series of waves, such as from crest to crest or trough to trough (Figure 10).

Figure 10. The wavelength of a single wave is the distance between two consecutive points along the wave.

Visible light constitutes only one of many types of electromagnetic radiation emitted from the sun. The electromagnetic spectrum is the range of all possible wavelengths of radiation (Figure 11). Each wavelength corresponds to a different amount of energy carried.

Figure 11. The sun emits energy in the form of electromagnetic radiation. This radiation exists in different wavelengths, each of which has its own characteristic energy. Visible light is one type of energy emitted from the sun.

Each type of electromagnetic radiation has a characteristic range of wavelengths. The longer the wavelength (or the more stretched out it appears), 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 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 sun emits (Figure 11) a broad range of electromagnetic radiation, including X-rays and ultraviolet (UV) rays. The higher-energy waves are dangerous to living things; for example, X-rays and UV rays can be harmful to humans.

Absorption of Light

Light energy enters the process of photosynthesis when pigments absorb the light. In plants, pigment molecules absorb only visible light for photosynthesis. 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 these colors to the human eye. The visible light portion of the electromagnetic spectrum is perceived by the human eye as a 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.

Understanding Pigments

Figure 12. Plants that commonly grow in the shade benefit from having a variety of light-absorbing pigments. Each pigment can absorb different wavelengths of light, which allows the plant to absorb any light that passes through the taller trees. (credit: Jason Hollinger)

Different kinds of pigments exist, and each absorbs only certain wavelengths (colors) of visible light. Pigments reflect the color of the wavelengths that they cannot absorb.

All photosynthetic organisms contain a pigment called chlorophyll a, which humans see as the common green color associated with plants. Chlorophyll a absorbs wavelengths from either end of the visible spectrum (blue and red), but not from green. Because green is reflected, chlorophyll appears green.

Other pigment types include chlorophyll b (which absorbs blue and red-orange light) and the carotenoids. Each type of pigment can be identified by the specific pattern of wavelengths it absorbs from visible light, which is its absorption spectrum.

Many photosynthetic organisms have a mixture of pigments; between them, the organism can absorb energy from a wider range of visible-light wavelengths. Not all photosynthetic organisms have full access to sunlight. Some organisms grow underwater where light intensity decreases with depth, and certain wavelengths are absorbed by the water. 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 block most of the sunlight (Figure 12).

The Light-Dependent Reactions of Photosynthesis

The overall purpose of the light-dependent reactions is to convert light energy into chemical energy. This chemical energy will be used by the Calvin cycle to fuel the assembly of sugar molecules.

The light-dependent reactions begin in a grouping of pigment molecules and proteins called a photosystem. Photosystems exist in the membranes of thylakoids. A pigment molecule in the photosystem absorbs one photon, a quantity or "packet" of light energy, at a time.

A photon of light energy travels until it reaches a pigment molecule, such as chlorophyll. The photon causes an electron in the chlorophyll to become "excited." The energy given to the electron then travels from one pigment molecule to another until it reaches a pair of chlorophyll a molecules called the reaction center. This energy then excites an electron in the reaction center causing it to break free and be passed to the primary electron acceptor. The reaction center is therefore said to "donate" an electron to the primary electron acceptor (Figure 13).

Figure 13. Light energy is absorbed by a chlorophyll molecule and the photon is passed along a pathway to other chlorophyll molecules. The energy culminates in a molecule of chlorophyll found in the reaction center. The energy "excites" one of its electrons enough to leave the molecule and be transferred to a nearby primary electron acceptor. A molecule of water splits to release an electron, which is needed to replace the one donated. Oxygen and hydrogen ions are also formed from the splitting of water.

To replace the electron in the reaction center, a molecule of water is split. This splitting releases an electron and results in the formation of oxygen (O ₂) and hydrogen ions (H⁺) in the thylakoid space. Technically, each breaking of a water molecule releases a pair of electrons, and therefore can replace two donated electrons.

The replacing of the electron enables the reaction center to respond to another photon. The oxygen molecules produced as byproducts find their way to the surrounding environment. The hydrogen ions play critical roles in the remainder of the light-dependent reactions.

Keep in mind that the purpose of the light-dependent reactions is to convert solar energy into chemical carriers that will be used in the Calvin cycle. In eukaryotes, two photosystems exist, the first is called photosystem II, which is named for the order of its discovery rather than for the order of function.

After the photon hits, photosystem II transfers the free electron to the first in a series of proteins inside the thylakoid membrane called the electron transport chain. As the electron passes along these proteins, energy from the electron fuels membrane pumps that actively move hydrogen ions against their concentration gradient from the stroma into the thylakoid space. This is quite analogous to the process that occurs in the mitochondrion in which an electron transport chain pumps hydrogen ions from the mitochondrial stroma across the inner membrane and into the intermembrane space, creating an electrochemical gradient. After the energy is used, the electron is accepted by a pigment molecule in the next photosystem, which is called photosystem I (Figure 14).

Figure 14. From photosystem II, the excited electron travels along a series of proteins. This electron transport system uses the energy from the electron to pump hydrogen ions into the interior of the thylakoid. A pigment molecule in photosystem I accepts the electron.

Generating an Energy Carrier: ATP

In the light-dependent reactions, energy absorbed by sunlight is stored by two types of energy-carrier molecules: ATP and NADPH. The energy that these molecules carry is stored in a bond that holds a single atom to the molecule. For ATP, it is a phosphate atom, and for NADPH, it is a hydrogen atom. NADH will be discussed further in relation to cellular respiration, which occurs in the mitochondrion, where it carries energy from the citric acid cycle to the electron transport chain. When these molecules release energy into the Calvin cycle, they each lose atoms to become the lower-energy molecules ADP and NADP⁺.

The buildup of hydrogen ions in the thylakoid space forms an electrochemical gradient because of the difference in the concentration of protons (H⁺) and the difference in the charge across the membrane that they create. This potential energy is harvested and stored as chemical energy in ATP through chemiosmosis, the movement of hydrogen ions down their electrochemical gradient through the transmembrane enzyme ATP synthase, just as in the mitochondrion.

The hydrogen ions are allowed to pass through the thylakoid membrane through an embedded protein complex called ATP synthase. This same protein generated ATP from ADP in the mitochondrion. The energy generated by the hydrogen ion stream allows ATP synthase to attach a third phosphate to ADP, which forms a molecule of ATP in a process called photophosphorylation. The flow of hydrogen ions through ATP synthase is called chemiosmosis, because the ions move from an area of high to low concentration through a semi-permeable structure.

Generating Another Energy Carrier: NADPH

The remaining function of the light-dependent reaction is to generate the other energy-carrier molecule, NADPH. As the electron from the electron transport chain arrives at photosystem I, it is re-energized with another photon captured by chlorophyll. The energy from this electron drives the formation of NADPH from NADP⁺ and a hydrogen ion (H⁺). Now that the solar energy is stored in energy carriers, it can be used to make a sugar molecule.

Learning Objectives

In the first part of photosynthesis, the light-dependent reaction, pigment molecules absorb energy from sunlight. The most common and abundant pigment is chlorophyll a. A photon strikes photosystem II to initiate photosynthesis. Energy travels through the electron transport chain, which pumps hydrogen ions into the thylakoid space. This forms an electrochemical gradient. The ions flow through ATP synthase from the thylakoid space into the stroma in a process called 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 carrier for the Calvin cycle reactions.

Practice Question

Describe the pathway of energy in light-dependent reactions.

Show Answer

The energy is present initially as light. A photon of light hits chlorophyll, causing an electron to be energized. The free electron travels through the electron transport chain, and the energy of the electron is used to pump hydrogen ions into the thylakoid space, transferring the energy into the electrochemical gradient. The energy of the electrochemical gradient is used to power ATP synthase, and the energy is transferred into a bond in the ATP molecule. In addition, energy from another photon can be used to create a high-energy bond in the molecule NADPH.

The Calvin Cycle

After the energy from the sun is converted and packaged into ATP and NADPH, the cell has the fuel needed to build food in the form of carbohydrate molecules. The carbohydrate molecules made will have a backbone of carbon atoms. Where does the carbon come from? The carbon atoms used to build carbohydrate molecules comes from carbon dioxide, the gas that animals exhale with each breath. The Calvin cycle is the term used for the reactions of photosynthesis that use the energy stored by the light-dependent reactions to form glucose and other carbohydrate molecules. This process may also be called the light-independent reaction, as it does not directly require sunlight (but it does require the products produced from the light-dependent reactions).

The Innerworkings of the Calvin Cycle

Figure 15. Light-dependent reactions harness energy from the sun to produce ATP and NADPH. These energy-carrying molecules travel into the stroma where the Calvin cycle reactions take place.

In plants, carbon dioxide (CO ₂) enters the chloroplast through the stomata and diffuses into the stroma of the chloroplast-the site of the Calvin cycle reactions where sugar is synthesized. The reactions are named after the scientist who discovered them, and reference the fact that the reactions function as a cycle. Others call it the Calvin-Benson cycle to include the name of another scientist involved in its discovery (Figure 15).

The Calvin cycle reactions (Figure 16) can be organized into three basic stages: fixation, reduction, and regeneration. In the stroma, in addition to CO ₂, two other chemicals are present to initiate the Calvin cycle: an enzyme abbreviated RuBisCO, and the molecule ribulose bisphosphate (RuBP). RuBP has five atoms of carbon and a phosphate group on each end.

RuBisCO catalyzes a reaction between CO ₂ and RuBP, which forms a six-carbon compound that is immediately converted into two three-carbon compounds. This process is called carbon fixation, because CO ₂ is "fixed" from its inorganic form into organic molecules.

ATP and NADPH use their stored energy to convert the three-carbon compound, 3-PGA, into another three-carbon compound called G3P. This type of reaction is called a reduction reaction, because it involves the gain of electrons. A reduction is the gain of an electron by an atom or molecule. The molecules of ADP and NAD⁺, resulting from the reduction reaction, return to the light-dependent reactions to be re-energized.

One of the G3P molecules leaves the Calvin cycle to contribute to the formation of the carbohydrate molecule, which is commonly glucose (C ₆H ₁₂O ₆). Because the carbohydrate molecule has six carbon atoms, it takes six turns of the Calvin cycle to make one carbohydrate molecule (one for each carbon dioxide molecule fixed). The remaining G3P molecules regenerate RuBP, which enables the system to prepare for the carbon-fixation step. ATP is also used in the regeneration of RuBP.

Figure 16. The Calvin cycle has three stages. In stage 1, the enzyme RuBisCO incorporates carbon dioxide into an organic molecule. In stage 2, the organic molecule is reduced. In stage 3, RuBP, the molecule that starts the cycle, is regenerated so that the cycle can continue.

In summary, it takes six turns of the Calvin cycle to fix six carbon atoms from CO ₂. These six turns require energy input from 12 ATP molecules and 12 NADPH molecules in the reduction step and 6 ATP molecules in the regeneration step.

Check out this animation of the Calvin cycle. Click Stage 1, Stage 2, and then Stage 3 to see G3P and ATP regenerate to form RuBP.

Try It

Figure 17. Living in the harsh conditions of the desert has led plants like this cactus to evolve variations in reactions outside the Calvin cycle. These variations increase efficiency and help conserve water and energy. (credit: Piotr Wojtkowski)

The shared evolutionary history of all photosynthetic organisms is conspicuous, as the basic process has changed little over eras of time. Even between the giant tropical leaves in the rainforest and tiny cyanobacteria, the process and components of photosynthesis that use water as an electron donor remain largely the same. Photosystems function to absorb light and use electron transport chains to convert energy. The Calvin cycle reactions assemble carbohydrate molecules with this energy.

However, as with all biochemical pathways, a variety of conditions leads to varied adaptations that affect the basic pattern. Photosynthesis in dry-climate plants (Figure 17) has evolved with adaptations that conserve water. In the harsh dry heat, every drop of water and precious energy must be used to survive. Two adaptations have evolved in such plants. In one form, a more efficient use of CO ₂ allows plants to photosynthesize even when CO ₂ is in short supply, as when the stomata are closed on hot days. The other adaptation performs preliminary reactions of the Calvin cycle at night, because opening the stomata at this time conserves water due to cooler temperatures. In addition, this adaptation has allowed plants to carry out low levels of photosynthesis without opening stomata at all, an extreme mechanism to face extremely dry periods.

Photosynthesis in Prokaryotes

The two parts of photosynthesis-the light-dependent reactions and the Calvin cycle-have been described, as they take place in chloroplasts. However, prokaryotes, such as cyanobacteria, lack membrane-bound organelles. Prokaryotic photosynthetic autotrophic organisms have infoldings of the plasma membrane for chlorophyll attachment and photosynthesis (Figure 18). It is here that organisms like cyanobacteria can carry out photosynthesis.

Figure 18. A photosynthetic prokaryote has infolded regions of the plasma membrane that function like thylakoids. Although these are not contained in an organelle, such as a chloroplast, all of the necessary components are present to carry out photosynthesis. (credit: scale-bar data from Matt Russell)

Learning Objectives

Using the energy carriers formed in the first stage of photosynthesis, the Calvin cycle reactions fix CO ₂ from the environment to build carbohydrate molecules. An enzyme, RuBisCO, catalyzes the fixation reaction, by combining CO ₂ with RuBP. The resulting six-carbon compound is broken down into two three-carbon compounds, and the energy in ATP and NADPH is used to convert these molecules into G3P. One of the three-carbon molecules of G3P leaves the cycle to become a part of a carbohydrate molecule. The remaining G3P molecules stay in the cycle to be formed back into RuBP, which is ready to react with more CO ₂. Photosynthesis forms a balanced energy cycle with the process of cellular respiration. Plants are capable of both photosynthesis and cellular respiration, since they contain both chloroplasts and mitochondria.

Practice Question

Which part of the Calvin cycle would be affected if a cell could not produce the enzyme RuBisCO?

Show Answer

None of the cycle could take place, because RuBisCO is essential in fixing carbon dioxide. Specifically, RuBisCO catalyzes the reaction between carbon dioxide and RuBP at the start of the cycle.

Summary

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:

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.
https://assessments.lumenlearning.co...sessments/6880

Contributors and Attributions

Introduction to Photosynthesis. Authored by: Shelli Carter and Lumen Learning. Provided by: Lumen Learning. License: CC BY: Attribution

Photosystem figure. Authored by: Adapted by Stephen Snyder. License: CC BY: Attribution. License Terms: Derivative of OpenStax original work

Photosynthesis: Crash Course Biology #8. Authored by: CrashCourse. Located at: https://youtu.be/sQK3Yr4Sc_k. License: All Rights Reserved. License Terms: Standard YouTube License

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Page ID 35693 What you'll learn to do: Identify the basic components and steps of photosynthesis Learning Objectives An Overview of Photosynthesis Solar Dependence and Food Production Practice Question The Two Parts of Photosynthesis Try It Learning Objectives Light Energy What Is Light Energy?