C10 photosynthesis

Photosynthesis converts light energy to the chemical energy of food

Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part)

light reactions

Calvin cycle

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The light reactions (in the thylakoids)

Split H2O

Release O2

Reduce the electron acceptor NADP+ to NADPH

Generate ATP from ADP by photophosphorylation

The Calvin cycle (in the stroma) forms sugar from CO2, using ATP and NADPH

The Calvin cycle begins with carbon fixation, incorporating CO2 into organic molecules

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Photosynthesis as a Redox Process

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Photosynthesis reverses the direction of electron flow compared to respiration

Photosynthesis is a redox process in which H2O is oxidized and CO2 is reduced

Photosynthesis is an endergonic process; the energy boost is provided by light

Chloroplasts: The Sites of Photosynthesis in Plants

Chloroplasts are found mainly in cells of the mesophyll, the interior tissue of the leaf

CO2 enters and O2 exits the leaf through microscopic pores called stomata

Thylakoids are connected sacs in the chloroplast that compose a third membrane system

Chlorophyll, the pigment that gives leaves their green color, resides in the thylakoid membranes

Photosynthesis is a complex series of reactions that can be summarized as the following equation:

6 CO2 + 12 H2O + Light energy → C6H12O6 + 6 O2 + 6 H2O

Chloroplasts split H2O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by-product

The light reactions convert solar energy to the chemical energy of ATP and NADPH

A Comparison of Chemiosmosis in Chloroplasts and Mitochondria

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Electron Flow

In summary, light reactions generate ATP and increase the potential energy of electrons by moving them from H2O to NADPH

Chloroplasts and mitochondria generate ATP by chemiosmosis, but use different sources of energy

Mitochondria transfer chemical energy from food to ATP; chloroplasts transform light energy into the chemical energy of ATP

Spatial organization of chemiosmosis differs between chloroplasts and mitochondria but also shows similarities

In mitochondria, protons are pumped to the intermembrane space and drive ATP synthesis as they diffuse back into the mitochondrial matrix
In chloroplasts, protons are pumped into the thylakoid space and drive ATP synthesis as they diffuse back into the stroma

ATP and NADPH are produced on the side facing the stroma, where the Calvin cycle takes place


cyclic

linear

the primary pathway, involves both photosystems and produces ATP and NADPH using light energy

8 steps

A photon hits a pigment in a light-harvesting complex of PS II, and its energy is passed among pigment molecules until it excites P680
An excited electron from P680 is transferred to the primary electron acceptor (we now call it P680+)

H2O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680+, thus reducing it to P680

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P680+ is the strongest known biological oxidizing agent

The H+ are released into the thylakoid space

O2 is released as a by-product of this reaction

Each electron “falls” down an electron transport chain from the primary electron acceptor of PS II to PS I. Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane
Potential energy stored in the proton gradient drives production of ATP by chemiosmosis

In PS I (like PS II), transferred light energy excites P700, which loses an electron to the primary electron acceptor

Each electron “falls” down an electron transport chain from the primary electron acceptor of PS I to the protein ferredoxin (Fd)
NADP+ reductase catalyzes the transfer of electrons to NADP+, reducing it to NADPH

electrons cycle back from Fd to the PS I reaction center via a plastocyanin molecule (Pc)

Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH
No oxygen is released

Some organisms such as purple sulfur bacteria have PS I but not PS II

Cyclic electron flow is thought to have evolved before linear electron flow
Cyclic electron flow may protect cells from light-induced damage

A Photosystem

A Reaction-Center Complex Associated with Light-Harvesting Complexes

reaction-center complex

an association of proteins holding a special pair of chlorophyll a molecules and a primary electron acceptor

consists of pigment molecules bound to proteins

transfer the energy of photons to the chlorophyll a molecules in the reaction-center complex

These chlorophyll a molecules are special because they can transfer an excited electron to a different molecule

primary electron acceptor

accepts excited electrons and is reduced as a result

Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions

Photosynthetic Pigments: The Light Receptors

Excitation of Chlorophyll by Light

Pigments are substances that absorb visible light

Different pigments absorb different wavelengths

Wavelengths that are not absorbed are reflected or transmitted

Leaves appear green because chlorophyll reflects and transmits green light

Chloroplasts are solar-powered chemical factories
Their thylakoids transform light energy into the chemical energy of ATP and NADPH

When a pigment absorbs light, it goes from a ground state to an excited state, which is unstable
When excited electrons fall back to the ground state, excess energy is released as heat

The Calvin cycle uses the chemical energy of ATP and NADPH to reduce CO2 to sugar

The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle

anabolic

The Calvin cycle is anabolic; it builds sugar from smaller molecules by using ATP and the reducing power of electrons carried by NADPH

three phases:

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Reduction

Regeneration of the CO2 acceptor (RuBP)

Carbon fixation (catalyzed by rubisco)

Alternative mechanisms of carbon fixation have evolved in hot, arid climates

Photorespiration

In photorespiration, rubisco adds O2 instead of CO2 in the Calvin cycle, producing a two-carbon compound

consumes O2 and organic fuel and releases CO2 without producing ATP or sugar

may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O2 and more CO2

limits damaging products of light reactions that build up in the absence of the Calvin cycle

C4 Plants

minimize the cost of photorespiration by incorporating CO2 into four-carbon compounds

Sugar production in C4 plants occurs in a three-step process:

1.The production of the four-carbon precursors is catalyzed by the enzyme PEP carboxylase in the mesophyll cells

2.These four-carbon compounds are exported to bundle-sheath cells
3.Within the bundle-sheath cells, they release CO2 that is then used in the Calvin cycle

climates

Suitable agricultural land is decreasing due to the effects of climate change, while the world demand for food continues to increase

Increasing levels of CO2 may affect C3 and C4 plants differently, perhaps changing the relative abundance of these species

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C4 photosynthesis uses less water and resources than C3 photosynthesis

Scientists have genetically modified rice, a C3 plant, to carry out C4 photosynthesis

They estimate 30–50% increase in yield compared to C3 rice

CAM Plants

crassulacean acid metabolism (CAM)

Some plants, including succulents, use crassulacean acid metabolism (CAM) to fix carbon

CAM plants open their stomata at night, incorporating CO2 into organic acids that are stored in the vacuoles

Stomata close during the day, and CO2 is released from organic acids and used in the Calvin cycle

Life depends on photosynthesis

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The energy entering chloroplasts as sunlight gets stored as chemical energy in organic compounds

Sugar made in the chloroplasts supplies chemical energy and carbon skeletons to synthesize the organic molecules of cells

Plants store excess sugar as starch in chloroplasts and other structures such as roots, tubers, seeds, and fruits