Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings PowerPoint ® Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp Chapter 10 Photosynthesis

Photosynthesis: The Energy Transfer Process That Feeds the Biosphere Photosynthesis is an energy transfer process that converts solar energy into chemical energy. Directly or indirectly, photosynthesis nourishes almost the entire living world. Photosynthesis is the key to Earth’s food and atmospheric oxygen. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Autotrophs “self feeders” make their own food. Autotrophs sustain themselves without eating anything derived from other organisms. Autotrophs are the producers of the biosphere, producing organic molecules (C-H- O) from CO 2 and other inorganic molecules. Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules (C-H-O) from H 2 O and CO 2 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Autotrophs = Producers (a) Plants (c) Unicellular protist 10 µm 1.5 µm 40 µm (d) Cyanobacteria (e) Purple sulfur bacteria (b) Multicellular alga

Heterotrophs obtain their organic material from other organisms. They eat … (-vores) Heterotrophs are the consumers of the biosphere. Almost all heterotrophs, including humans, depend on photoautotrophs for food and O 2 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Photosynthesis converts light energy to the chemical energy of food (C-H-O) Chloroplasts are structurally similar to and likely evolved from photosynthetic bacteria: cyanobacteria. The structural organization of these cells allows for the chemical reactions of photosynthesis. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chloroplasts: The Sites of Photosynthesis in Plants Leaves are the major locations of photosynthesis. Their green color is from chlorophyll, the green pigment within chloroplasts. Light energy absorbed by chlorophyll drives the synthesis of organic molecules in the chloroplast. CO 2 enters and O 2 exits the leaf through microscopic pores called stomata. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chloroplasts are found mainly in cells of the mesophyll, the interior tissue (layers) of the leaf. A typical mesophyll cell has 30–40 chloroplasts In the chloroplast, the chlorophyll is in the membranes of thylakoids (connected sacs); thylakoids may be stacked in columns called grana. Chloroplasts also contain stroma, a dense fluid region. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Chloroplasts

Zooming in on the location of photosynthesis in a plant 5 µm Mesophyll cell Stomata CO 2 O2O2 Chloroplast Mesophyll Vein Leaf cross section

Tracking Atoms Through Photosynthesis: Scientific Inquiry Photosynthesis can be summarized as the following equation: 6 CO H 2 O + Light energy  C 6 H 12 O O H 2 O or 6 CO H 2 O + Light energy  C 6 H 12 O O 2 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Splitting of Water = Photolysis In the light dependent reactions, chloroplasts split H 2 O into hydrogen and oxygen. In the light independent reactions, the chloroplasts incorporate the hydrogen from water into sugar molecules. Photosynthesis is a redox process in which H 2 O is oxidized to O 2 and C O 2 is reduced to C 6 H 12 O 6 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Two Stages of Photosynthesis: A Preview Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part) The light reactions (in the thylakoids): – Split H 2 O – Release O 2 – Reduce NADP + to NADPH – Generate ATP by photophosphorylation Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Calvin cycle (in the stroma) forms sugar C-H-O from CO 2, using ATP and NADPH. The Calvin cycle begins with carbon fixation, which incorporates CO 2 into organic molecules and then reduction produces sugar C-H-O. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Light Fig H2OH2O Chloroplast Light Reactions NADP + P ADP i +

Light Light Reactions H2OH2O Chloroplast Light Reactions NADP + P ADP i + ATP NADPH O2O2

Light H2OH2O Chloroplast Light Reactions NADP + P ADP i + ATP NADPH O2O2 Calvin Cycle CO 2

Light Photosynthesis H2OH2O Chloroplast Light Reactions NADP + P ADP i + ATP NADPH O2O2 Calvin Cycle CO 2 [CH 2 O] (sugar)

Light Reactions convert Solar Energy to the Chemical Energy of ATP and NADPH Chloroplasts are solar-powered chemical factories. Their thylakoids transform light energy into the chemical energy of ATP and NADPH. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Nature of Sunlight Light is a form of electromagnetic energy, also called electromagnetic radiation. Like other electromagnetic energy, light travels in rhythmic waves. Wavelength is the distance between crests of waves. Wavelength determines the type of electromagnetic energy. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The electromagnetic spectrum is the entire range of electromagnetic energy, or radiation. Visible light consists of wavelengths (including those that drive photosynthesis) that produce colors we can see. Light also behaves as though it consists of discrete energy particles, called photons. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

UV Visible light Infrared Micro- waves Radio waves X-rays Gamma rays 10 3 m 1 m (10 9 nm) 10 6 nm 10 3 nm 1 nm 10 –3 nm 10 –5 nm nm Longer wavelength Lower energyHigher energy Shorter wavelength Electromagnetic Spectrum

Photosynthetic Pigments: The Light Receptors 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 green light. We see reflected light. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Why leaves are green: interaction of light with chloroplasts Reflected light Absorbed light Light Chloroplast Transmitted light Granum

A spectrophotometer measures a pigment’s ability to absorb various wavelengths. This machine sends light through pigments and measures the fraction of light transmitted at each wavelength. Transmitted light is light that is not absorbed. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Determining an absorption spectrum Galvanometer Slit moves to pass light of selected wavelength White light Green light Blue light The low transmittance (high absorption) reading indicates that chlorophyll absorbs most blue light. The high transmittance (low absorption) reading indicates that chlorophyll absorbs very little green light. Refracting prism Photoelectric tube Chlorophyll solution TECHNIQUE

An absorption spectrum is a graph plotting a pigment’s light absorption vs. wavelength. The absorption spectrum of chlorophyll a suggests that violet-blue and red light work best for photosynthesis. An action spectrum profiles the relative effectiveness of different wavelengths of radiation in driving a process such as photosynthesis. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Wavelength of light (nm) Action spectrum Absorption spectra Engelmann’s experiment Aerobic bacteria RESULTS Rate of photosynthesis (measured by O 2 release) Absorption of light by chloroplast pigments Filament of alga Chloro- phyll a Chlorophyll b Carotenoids

The action spectrum of photosynthesis was first demonstrated in 1883 by Theodor W. Engelmann. In his experiment, he exposed different segments of a filamentous alga to different wavelengths. Areas receiving wavelengths favorable to photosynthesis produced excess O 2 He used the growth of aerobic bacteria clustered along the alga as a measure of O 2 production. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chlorophyll a is the main photosynthetic pigment. Accessory pigments, such as chlorophyll b, broaden the spectrum used for photosynthesis. Accessory pigments called carotenoids absorb excessive light that would damage chlorophyll. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Chlorophyll Porphyrin ring: light-absorbing “head” of molecule; note magnesium atom at center in chlorophyll a CH 3 Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts; H atoms not shown CHO in chlorophyll b

Excitation of Chlorophyll by Light 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, photons are given off, an afterglow called fluorescence. If illuminated, an isolated solution of chlorophyll will fluoresce, giving off light and heat. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Excitation of isolated chlorophyll by light (a) Excitation of isolated chlorophyll molecule Heat Excited state (b) Fluorescence Photon Ground state Photon (fluorescence) Energy of electron e–e– Chlorophyll molecule

A Photosystem: A Reaction-Center Complex Associated with Light-Harvesting Complexes A photosystem consists of a reaction-center complex (a type of protein complex) surrounded by light-harvesting complexes. The light-harvesting complexes (pigment molecules bound to proteins) funnel the energy of photons to the reaction center. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

A primary electron acceptor in the reaction center accepts an excited electron from chlorophyll a. Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

How a photosystem harvests light THYLAKOID SPACE (INTERIOR OF THYLAKOID) STROMA e–e– Pigment molecules Photon Transfer of energy Special pair of chlorophyll a molecules Thylakoid membrane Photosystem Primary electron acceptor Reaction-center complex Light-harvesting complexes

There are two types of photosystems in the thylakoid membranes. Photosystem II (PS II) functions first (the numbers reflect order of discovery) and is best at absorbing a wavelength of 680 nm. The reaction-center chlorophyll a of PS II is called P680. Photosystem I (PS I) is best at absorbing a wavelength of 700 nm. The reaction-center chlorophyll a of PS I is called P700. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Linear Electron Flow: Non-Cyclic Flow is Normal. During the light reactions, there are two possible routes for electron flow: cyclic and linear (non-cyclic). Linear electron flow, non-cyclic electron flow, is the primary pathway. Non-cyclic flow involves both photosystems II & I, and produces ATP and NADPH using light energy. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

In Photosystem II electron flow, P680 + (P680 that is missing an electron) is a very strong oxidizing agent. H 2 O is split (photolysis) by enzymes, and the electrons are transferred from the hydrogen atoms to P680 +, thus reducing it to P680. O 2 is released as a by-product of this reaction. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Pigment molecules Light P680 e–e– Primary acceptor 2 1 e–e– e–e– 2 H + O2O2 + 3 H2OH2O 1/21/2 4 Pq Pc Cytochrome complex Electron transport chain 5 ATP Photosystem II (PS II) Diffusion of H + (protons) across the thylakoid membrane drives ATP synthesis Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane.

In PS I (like PS II), transferred light energy excites P700, which loses an electron to an electron acceptor. P700 + (P700 that is missing an electron) accepts an electron passed down from PS II via the electron transport chain, ETC. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Each electron “falls” down an ETC from the primary electron acceptor of PS I to the protein ferredoxin (Fd). The electrons are then transferred to NADP + and reduce it to NADPH. The electrons of NADPH are available for the reactions of the Calvin cycle in the stroma. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Pigment molecules Light P680 e–e– Primary acceptor 2 1 e–e– e–e– 2 H + O2O2 + 3 H2OH2O 1/21/2 4 Pq Pc Cytochrome complex Electron transport chain 5 ATP Photosystem I (PS I) Light Primary acceptor e–e– P700 6 Fd Electron transport chain NADP + reductase NADP + + H + NADPH 8 7 e–e– e–e– 6 Photosystems II and I: Non-Cyclic Electron Flow: Produces: O 2 NADPH and ATP Photosystem II (PS II)

A mechanical analogy for the light reactions Mill makes ATP e–e– NADPH Photon e–e– e–e– e–e– e–e– e–e– ATP Photosystem IIPhotosystem I e–e–

Cyclic Electron Flow: not the normal flow Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH. Cyclic electron flow generates surplus ATP, satisfying the higher demand in the Calvin cycle. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Cyclic Electron Flow makes only ATP ATP Photosystem II Photosystem I Primary acceptor Pq Cytochrome complex Fd Pc Primary acceptor Fd NADP + reductase NADPH NADP + + H +

A Comparison of Chemiosmosis in Chloroplasts and Mitochondria Chloroplasts and mitochondria generate ATP by chemiosmosis, but use different sources of energy. Mitochondria transfer chemical energy from food to ATP : oxidative phosphosphorlation. Chloroplasts transform light energy into the chemical energy of ATP: photophosphorylation. Spatial organization of chemiosmosis differs between chloroplasts and mitochondria but also shows similarities. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Proton Gradient drives ATP Synthesis

chemiosmosis in mitochondria and chloroplasts Key Mitochondrion Chloroplast CHLOROPLAST STRUCTURE MITOCHONDRION STRUCTURE Intermembrane space Inner membrane Electron transport chain H+H+ Diffusion Matrix Higher [H + ] Lower [H + ] Stroma ATP synthase ADP + P i H+H+ ATP Thylakoid space Thylakoid membrane

ATP and NADPH are produced on the side facing the stroma where the Calvin cycle takes place. In summary, light reactions generate ATP and increase the potential energy of electrons by moving them from H 2 O to NADPH. O 2, ATP, and NADPH are produced in the Light Reactions. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Light Fd Cytochrome complex ADP + i H+H+ ATP P synthase To Calvin Cycle STROMA (low H + concentration) Thylakoid membrane THYLAKOID SPACE (high H + concentration) STROMA (low H + concentration) Photosystem II Photosystem I 4 H + Pq Pc Light NADP + reductase NADP + + H + NADPH +2 H + H2OH2O O2O2 e–e– e–e– 1/21/ Photochemical = Light Reactions

The Calvin Cycle uses ATP and NADPH to convert CO 2 to sugar The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle. The Calvin Cycle builds sugar from smaller molecules using: CO 2, ATP, and NADPH. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Carbon enters the cycle as CO 2 and leaves as a sugar named glyceraldehyde-3-phospate (G3P) a triose sugar. For net synthesis of 1 G3P, the cycle must take place three times, fixing 3 molecules of CO 2 The Calvin cycle has three phases: – Carbon fixation (CO 2 attaches to RuBP catalyzed by rubisco) – Reduction “sugar making” – Regeneration of the CO 2 acceptor (RuBP). Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Calvin Cycle Ribulose bisphosphate ( RuBP) 3-Phosphoglycerate Short-lived intermediate Phase 1: Carbon fixation (Entering one at a time) Rubisco Input CO 2 P P P P P ATP 6 6 ADP P P 6 1,3-Bisphosphoglycerate 6 P P NADP + NADPH i Phase 2: Reduction: “sugar making ” Glyceraldehyde-3-phosphate (G3P) 1 P Output G3P ( a triose sugar ) Glucose and other organic compounds Calvin Cycle 3 3 ADP ATP 5 P Phase 3: Regeneration of the CO 2 acceptor RuBP G3P

Alternative mechanisms of carbon fixation have evolved in hot, arid climates Dehydration is a problem for plants, sometimes requiring trade-offs with other metabolic processes, especially photosynthesis. On hot, dry days, plants close stomata, which conserves H 2 O but also limits photosynthesis The closing of stomata reduces access to CO 2 and causes O 2 to build up. These conditions favor a seemingly wasteful process called photorespiration. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Photorespiration: An Evolutionary Relic? In most plants, C 3 plants, initial fixation of CO 2 by rubisco forms a three-carbon compound. In photorespiration, rubisco bonds with and adds O 2 instead of CO 2 in the Calvin cycle. Photorespiration consumes O 2 and organic fuel and releases CO 2 without producing ATP or sugar. Wasteful. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Photorespiration may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O 2 and more CO 2 Photorespiration limits damaging products of light reactions that build up in the absence of the Calvin cycle In many plants, photorespiration is a problem because on a hot, dry day it can drain as much as 50% of the carbon fixed by the Calvin cycle. This slows / harms plant growth significantly. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

C 4 Plants = Most Efficient at Carbon Fixation C 4 plants minimize the cost of photorespiration by incorporating CO 2 into four-carbon compounds in mesophyll cells. This step requires the enzyme PEP carboxylase PEP carboxylase has a higher affinity for CO 2 than rubisco does; it can fix CO 2 even when CO 2 concentrations are low. These four-carbon compounds are exported to bundle-sheath cells, where they release CO 2 that is then used in the Calvin cycle. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

C 4 leaf anatomy Mesophyll cell Photosynthetic cells of C 4 plant leaf Bundle- sheath cell Vein (vascular tissue) Stoma The C 4 pathway Mesophyll cell CO 2 PEP carboxylase Oxaloacetate (4C) Malate (4C) PEP (3C) ADP ATP Pyruvate (3C) CO 2 Bundle- sheath cell Calvin Cycle Sugar Vascular tissue Carbon fixation occurs in Bundle Sheath, a region with low O 2

CAM Plants: Stomates Open at Night Some plants, including succulents, use crassulacean acid metabolism (CAM) for carbon fixation. CAM plants open their stomata at night, incorporating CO 2 into organic acids. Stomata close during the day, and CO 2 is released from organic acids and used in the Calvin Cycle. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

CO 2 Sugarcane Mesophyll cell CO 2 C4C4 Bundle- sheath cell Organic acids release CO 2 to Calvin cycle CO 2 incorporated into four-carbon organic acids (carbon fixation) Pineapple Night Day CAM Sugar Calvin Cycle Calvin Cycle Organic acid (a) Spatial separation of steps (b) Temporal separation of steps CO 2 1 2

The Importance of Photosynthesis: A Review 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 structures such as roots, tubers, seeds, and fruits. In addition to food production, photosynthesis produces the O 2 in our atmosphere. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Photosynthesis Light Reactions: Photosystem II Electron transport chain Photosystem I Electron transport chain CO 2 NADP + ADP P i + RuBP 3-Phosphoglycerate Calvin Cycle G3P ATP NADPH Starch (storage) Sucrose (export) Chloroplast Light H2OH2O O2O2

Photochemical = Light Reactions CO 2 NADP + reductase Photosystem II H2OH2O O2O2 ATP Pc Cytochrome complex Primary acceptor Primary acceptor Photosystem I NADP + + H + Fd NADPH Electron transport chain Electron transport chain O2O2 H2OH2O Pq

Calvin Cycle = Light Independent Reactions Regeneration of CO 2 acceptor 1 G3P (3C) Reduction Carbon fixation 3 CO 2 Calvin Cycle 6  3C 5  3C 3  5C

You should now be able to : 1.Describe the structure of a chloroplast. 2.Describe the relationship between an action spectrum and an absorption spectrum. 3.Trace the movement of electrons in linear, noncyclic electron flow. 4.Trace the movement of electrons in cyclic electron flow. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

5.Describe the similarities and differences between oxidative phosphorylation in mitochondria and photophosphorylation in chloroplasts..; 6.Describe the role of ATP and NADPH in the Calvin cycle. 7.Describe the major consequences of photorespiration. 8.Describe two important photosynthetic adaptations that minimize photorespiration. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings