8 Photosynthesis: Energy from Sunlight

8 Photosynthesis: Energy from Sunlight 8.1 What Is Photosynthesis? 8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? 8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? 8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? 8.5 How Is Photosynthesis Connected to Other Metabolic Pathways in Plants?

8.1 What Is Photosynthesis? Photosynthesis: “synthesis from light” The broad outline: Plants take in CO 2 and release water and O 2 Light is required

Figure 8.1 The Ingredients for Photosynthesis

8.1 What Is Photosynthesis? Ruben and Kamen determined the source of O 2 released during photosynthesis by using radioisotope tracers.

Figure 8.2 Water Is the Source of the Oxygen Produced by Photosynthesis

8.1 What Is Photosynthesis? Two pathways: Light reactions: light energy converted to chemical energy (in ATP and NADPH + H + ) Light-independent reactions: use the ATP and NADPH + H + plus CO 2 to produce sugars

Figure 8.3 An Overview of Photosynthesis

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Light is a form of electromagnetic radiation, which comes in discrete packets called photons and behave as particles. Light also behaves as if propagated as waves. Energy of a photon is inversely proportional to its wavelength.

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? When a photon meets a molecule it can be: Scattered or reflected Transmitted or pass through the molecule Absorbed—the molecule acquires the energy of the photon. The molecule goes from ground state to excited state.

Figure 8.4 Exciting a Molecule (A)

Figure 8.4 Exciting a Molecule (B) Energy from the photon boosts an electron into another shell

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Photons can have a wide range of wavelengths and energy levels. Molecules that absorb specific wavelengths in the visible range of the spectrum are called pigments.

Figure 8.5 The Electromagnetic Spectrum

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Absorption spectrum: plot of wavelengths absorbed by a pigment Action spectrum: plot of biological activity as a function of the wavelengths of light the organism is exposed to

Figure 8.6 Absorption and Action Spectra (Part 1)

Figure 8.6 Absorption and Action Spectra (Part 2)

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Several types of pigments absorb light energy used in photosynthesis: Chlorophylls: a and b Accessory pigments: absorb in red and blue regions, transfer the energy to chlorophylls—carotenoids, phycobilins

Figure 8.7 The Molecular Structure of Chlorophyll (Part 1)

Figure 8.7 The Molecular Structure of Chlorophyll (Part 2)

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? When a pigment returns to ground state, some of the energy may be given off as heat, some may be given off as fluorescence. Fluorescence has longer wavelengths and less energy than the absorbed light energy. No work is done.

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? If pigment can pass the energy to another molecule, there’s no fluorescence. The energy can be passed to a reaction center where it is converted to chemical energy.

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Pigments are arranged in antenna systems. Pigments are packed together on thylakoid membrane proteins. Excitation energy is passed from pigments that absorb short wavelengths to those that absorb longer wavelengths, and ends up in the reaction center pigment.

Figure 8.8 Energy Transfer and Electron Transport

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? The reaction center molecule is chlorophyll a. The excited chlorophyll (Chl*) is a reducing agent (electron donor).

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? A is the first in a chain of electron carriers on the thylakoid membrane—electron transport, a series of redox reactions. The final electron acceptor is NADP +

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Two systems of electron transport: Noncyclic electron transport—produces NADPH + H + and ATP Cyclic electron transport—produces ATP only

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Noncyclic electron transport: light energy is used to oxidize water → O 2, H +, and electrons Chl + is a strong oxidizing agent. It takes electrons from water, splitting the water molecule.

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Two photosystems required in noncyclic electron transport. Each photosystem consists of several chlorophyll and accessory pigment molecules. The photosystems complement each other, must be constantly absorbing light energy to power noncyclic electron transport.

Figure 8.9 Noncyclic Electron Transport Uses Two Photosystems

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Photosystem I Light energy reduces NADP + to NADPH + H + Reaction center is chlorophyll a molecule P 700 —absorbs in the 700nm range

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Photosystem II Light energy oxidizes water → O 2, H +, and electrons Reaction center is chlorophyll a molecule P 680 —absorbs at 680nm

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? The Z scheme models describes noncyclic electron transport. The reactions in the electron transport chain are coupled to a proton pump that results in the chemiosmotic formation of ATP.

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Cyclic electron transport: an electron from an excited chlorophyll molecule cycles back to the same chlorophyll molecule. A series of exergonic redox reactions, the released energy creates a proton gradient that is used to synthesize ATP.

Figure 8.10 Cyclic Electron Transport Traps Light Energy as ATP

8.2 How Does Photosynthesis Convert Light Energy into Chemical Energy? Photophosphorylation: light-driven production of ATP—a chemiosmotic mechanism. Electron transport is coupled to the transport of H + across the thylakoid membrane—from the stroma into the lumen.

Figure 8.11 Chloroplasts Form ATP Chemiosmotically (Part 1)

Figure 8.11 Chloroplasts Form ATP Chemiosmotically (Part 2)

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? CO 2 fixation—CO 2 is reduced to carbohydrates. Enzymes in the stroma use the energy in ATP and NADPH to reduce CO 2. Because the ATP and NADPH are not “stockpiled” these light-independent reactions must also take place in the light.

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? Calvin and Benson used the 14 C radioisotope to determine the sequence of reactions in CO 2 fixation. They exposed Chlorella to 14 CO 2, then extracted the organic compounds and separated them by paper chromatography.

Figure 8.12 Tracing the Pathway of CO 2 (Part 1)

Figure 8.12 Tracing the Pathway of CO 2 (Part 2)

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? 3-second exposure of the Chlorella to 14 CO 2 revealed that the first compound to be formed is 3PG, a 3-carbon sugar phosphate.

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? The pathway of CO 2 fixation is the Calvin cycle. CO 2 is first added to a 5-C RuBP; the 6-C compound immediately breaks down into two molecules of 3PG. The enzyme is rubisco—the most abundant protein in the world.

Figure 8.13 The Calvin Cycle (Part 1)

Figure 8.13 The Calvin Cycle (Part 2)

Figure 8.14 RuBP Is the Carbon Dioxide Acceptor

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? The Calvin cycle consists of 3 processes: Fixation of CO 2 Reduction of 3PG to G3P Regeneration of RuBP

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? G3P: glyceraldehyde 3-phosphate Most is recycled into RuBP The rest is converted to starch and sucrose

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? Covalent bonds in carbohydrates produced in the Calvin cycle represent the total energy yield of photosynthesis. This energy is used by the autotrophs themselves, and by heterotrophs—other organisms that cannot photosynthesize.

8.3 How Is Chemical Energy Used to Synthesize Carbohydrates? The Calvin cycle is stimulated by light: Proton pumping from stroma into thylakoids increases the pH which favors the activation of rubisco. Electron flow from photosystem I reduces disulfide bonds to activate Calvin cycle enzymes.

Figure 8.15 The Photochemical Reactions Stimulate the Calvin Cycle

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? Rubisco is an oxygenase as well as a carboxylase. It can add O 2 to RuBP instead of CO 2, reduces amount of CO 2 fixed, and limits plant growth. Products: 3PG and phosphoglycolate

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? Photorespiration: The phosphoglycolate forms glycolate— moves into peroxisomes—converted to glycine. Glycine diffuses into mitochondria, 2 glycines are converted into glycerate + CO 2

Figure 8.16 Organelles of Photorespiration

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? Rubisco has 10 times more affinity for CO 2. In the leaf, if O 2 concentration is high, photorespiration occurs. If CO 2 concentration is high, CO 2 is fixed. Photorespiration is more likely at high temperatures, such as hot days when stomata are closed.

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? C 3 plants: first product of CO 2 fixation is 3PG. Palisade cells in the mesophyll have abundant rubisco. C 4 plants: first product of CO 2 fixation is oxaloacetate, a 4-C compound. Mesophyll cells contain PEP carboxylase.

Figure 8.17 Leaf Anatomy of C 3 and C 4 Plants

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? C 4 plants—corn, sugar cane, tropical grasses—can keep stomata closed on hot days, but photorespiration does not occur. In mesophyll cells, CO 2 is accepted by PEP (phosphoenolpyruvate) to form oxaloacetate. PEP carboxylase has no affinity for O 2.

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? Oxaloacetate diffuses to bundle sheath cells which have abundant rubisco. The oxaloacetate is decarboxylated (PEP returns to mesophyll cells), CO 2 enters the Calvin cycle.

Figure 8.18 The Anatomy and Biochemistry of C 4 Carbon Fixation (A)

Figure 8.18 The Anatomy and Biochemistry of C 4 Carbon Fixation (B)

8.4 How Do Plants Adapt to the Inefficiencies of Photosynthesis? CAM plants—crassulacean acid metabolism Fix CO 2 with PEP carboxylase—at night—stomata can open with less water loss. Oxaloacetate is converted to malic acid. Day—malic acid goes to chloroplasts and is decarboxylated—CO 2 enters the Calvin cycle.

8.5 How Is Photosynthesis Connected to Other Metabolic Pathways in Plants? Photosynthesis and respiration are closely linked by the Calvin cycle. Glycolysis in the cytosol, respiration in the mitochondria, and photosynthesis in the chloroplasts can occur simultaneously.

Figure 8.19 Metabolic Interactions in a Plant Cell (Part 1)

Figure 8.19 Metabolic Interactions in a Plant Cell (Part 2)

8.5 How Is Photosynthesis Connected to Other Metabolic Pathways in Plants? Photosynthesis results in only 5 percent of total sunlight energy being transformed to the energy of chemical bonds. Understanding the inefficiencies of photosynthesis may be important as climate change drives changes in photosynthetic activity of plants.

Figure 8.20 Energy Losses During Photosynthesis