Plans for the week  11/2 – Quiz  11/3 Photosynthesis overview – Quiz Review CH 8 outline due  11/4 Light Reactions  11/5 Genetic Update Conference – 8 th will be yours to catch up on work you will miss that day. See you teachers tomorrow to gather that work.  11/6 Calvin Cycle - 11/9 CAM and C4  Next week: Tuesday quiz on CH 8,  Thursday Unit Test on CH 6,7,8 (Energy, Respiration & Photosynthesis)

Photosynthesis 11/3  1.) Parts of a chloroplasts – Location Location Location  2.) What is used in Light independent and Calvin Cycle  3.) Some preassesment!

Overview: The Process That Feeds the Biosphere  Photosynthesis is the process that converts solar energy into chemical energy  Directly or indirectly, photosynthesis nourishes almost the entire living world © 2014 Pearson Education, Inc.

Figure 8.1

 Photosynthesis occurs in plants, algae, certain other protists, and some prokaryotes  These organisms feed not only themselves but also most of the living world © 2014 Pearson Education, Inc.

Figure 8.2 (a) Plants (d) Cyanobacteria (e) Purple sulfur bacteria (b) Multicellular alga (c) Unicellular eukaryotes 10  m 1  m 40  m

© 2014 Pearson Education, Inc. Figure 8.2a (a) Plants

© 2014 Pearson Education, Inc. Figure 8.2b (b) Multicellular alga

© 2014 Pearson Education, Inc. Figure 8.2c (c) Unicellular eukaryotes 10  m

© 2014 Pearson Education, Inc. Figure 8.2d (d) Cyanobacteria 40  m

© 2014 Pearson Education, Inc. Figure 8.2e (e) Purple sulfur bacteria 1  m

Concept 8.1: Photosynthesis converts light energy to the chemical energy of food  The structural organization of photosynthetic cells includes enzymes and other molecules grouped together in a membrane  This organization allows for the chemical reactions of photosynthesis to proceed efficiently  Chloroplasts are structurally similar to and likely evolved from photosynthetic bacteria © 2014 Pearson Education, Inc.

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  Chloroplasts are found mainly in cells of the mesophyll, the interior tissue of the leaf  Each mesophyll cell contains 30–40 chloroplasts © 2014 Pearson Education, Inc.

 CO 2 enters and O 2 exits the leaf through microscopic pores called stomata  The chlorophyll is in the membranes of thylakoids thylakoids may be stacked in columns called grana  Chloroplasts also contain stroma, a dense interior fluid © 2014 Pearson Education, Inc.

Figure 8.3 Leaf cross section 20  m Mesophyll Stomata Chloroplasts Vein CO 2 O2O2 Mesophyll cell Chloroplast Stroma Thylakoid space Outer membrane Intermembrane space Inner membrane Granum 1  m

© 2014 Pearson Education, Inc. Figure 8.3a Leaf cross section Mesophyll Stomata Chloroplasts Vein CO 2 O2O2

© 2014 Pearson Education, Inc. Figure 8.3b 20  m Mesophyll cell Chloroplast Stroma Thylakoid space Outer membrane Intermembrane space Inner membrane Granum 1  m

© 2014 Pearson Education, Inc. Figure 8.3c 20  m Mesophyll cell

© 2014 Pearson Education, Inc. Figure 8.3d Stroma Granum 1  m

Tracking Atoms Through Photosynthesis: Scientific Inquiry  Photosynthesis is a complex series of reactions that can be summarized as the following equation 6 CO 2  12 H 2 O  Light energy  C 6 H 12 O 6  6 O 2  6 H 2 O © 2014 Pearson Education, Inc.

The Splitting of Water  Chloroplasts split H 2 O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by-product © 2014 Pearson Education, Inc.

Figure 8.4 Products: Reactants: 6 CO 2 6 O 2 C 6 H 12 O 6 6 H 2 O 12 H 2 O

Photosynthesis as a Redox Process  Photosynthesis reverses the direction of electron flow compared to respiration  Photosynthesis is a redox process in which H 2 O is oxidized and CO 2 is reduced  Photosynthesis is an endergonic process; the energy boost is provided by light © 2014 Pearson Education, Inc.

Figure 8.UN01 becomes reduced becomes oxidized

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 the electron acceptor, NADP , to NADPH  Generate ATP from ADP by adding a phosphate group, photophosphorylation © 2014 Pearson Education, Inc.

 The Calvin cycle (in the stroma) forms sugar from CO 2, using ATP and NADPH  The Calvin cycle begins with carbon fixation, incorporating CO 2 into organic molecules © 2014 Pearson Education, Inc. Animation: Photosynthesis

11/4 The Light Dependant Reactions  1.) Light and photons  2.) excited electrons and pigments  3.) Electron Transport Chain and Chemiosmosis  4.) Linear Electron Flow  But first most importantly

 of the light reactions is to create ATP and NADPH for the Calvin Cycle  The biproduct is Oxygen The “GOAL”

What is needed for light dependant reaction  Water  Light

Figure 8.6 Gamma rays 10 −5 nm10 −3 nm1 nm 10 3 nm 10 6 nm 1 m (10 9 nm) 10 3 m Radio waves Micro- waves X-rays Infrared UV Visible light Shorter wavelength Longer wavelength Lower energy Higher energy nm

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 and transmits green light © 2014 Pearson Education, Inc. Animation: Light and Pigments

Figure 8.7 Reflected light Light Absorbed light Chloroplast Granum Transmitted light

Figure 8.8 Refracting prism White light Green light Blue light Chlorophyll solution Photoelectric tube Galvanometer Slit moves to pass light of selected wavelength. 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. Technique 1243

Figure 8.9 Chloro- phyll a Rate of photosynthesis (measured by O 2 release) Results Absorption of light by chloroplast pigments Chlorophyll b Carotenoids Filament of alga Aerobic bacteria (a) Absorption spectra (b) Action spectrum (c) Engelmann’s experiment Wavelength of light (nm)

Figure 8.9a Chloro- phyll a Absorption of light by chloroplast pigments Chlorophyll b Carotenoids (a) Absorption spectra Wavelength of light (nm)

Figure 8.9b (b) Action spectrum Rate of photosynthesis (measured by O 2 release)

Figure 8.9c Filament of alga Aerobic bacteria (c) Engelmann’s experiment

© 2014 Pearson Education, Inc. Figure 8.5 Light CO 2 H2OH2O P i Chloroplast Light Reactions Calvin Cycle [CH 2 O] (sugar) O2O2 ADP ATP NADP   NADPH

© 2014 Pearson Education, Inc. Figure Light H2OH2O Chloroplast Light Reactions P i ADP NADP  

© 2014 Pearson Education, Inc. Figure Light H2OH2O P i Chloroplast Light Reactions O2O2 ADP ATP NADP   NADPH

© 2014 Pearson Education, Inc. Figure Light CO 2 H2OH2O P i Chloroplast Light Reactions Calvin Cycle O2O2 ADP ATP NADP   NADPH

© 2014 Pearson Education, Inc. Figure Light CO 2 H2OH2O P i Chloroplast Light Reactions Calvin Cycle [CH 2 O] (sugar) O2O2 ADP ATP NADP   NADPH

Location!  Chloroplasts are solar-powered chemical factories  Their thylakoids transform light energy into the chemical energy of ATP and NADPH © 2014 Pearson Education, Inc.

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 and transmits green light © 2014 Pearson Education, Inc. Animation: Light and Pigments

© 2014 Pearson Education, Inc. Figure 8.7 Reflected light Light Absorbed light Chloroplast Granum Transmitted light

 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 © 2014 Pearson Education, Inc.

Figure 8.8 Refracting prism White light Green light Blue light Chlorophyll solution Photoelectric tube Galvanometer Slit moves to pass light of selected wavelength. 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. Technique 1243

© 2014 Pearson Education, Inc. Figure 8.9 Chloro- phyll a Rate of photosynthesis (measured by O 2 release) Results Absorption of light by chloroplast pigments Chlorophyll b Carotenoids Filament of alga Aerobic bacteria (a) Absorption spectra (b) Action spectrum (c) Engelmann’s experiment Wavelength of light (nm)

© 2014 Pearson Education, Inc. Figure 8.9a Chloro- phyll a Absorption of light by chloroplast pigments Chlorophyll b Carotenoids (a) Absorption spectra Wavelength of light (nm)

© 2014 Pearson Education, Inc. Figure 8.9b (b) Action spectrum Rate of photosynthesis (measured by O 2 release)

© 2014 Pearson Education, Inc. Figure 8.9c Filament of alga Aerobic bacteria (c) Engelmann’s experiment

 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 © 2014 Pearson Education, Inc. Video: Chlorophyll Model

© 2014 Pearson Education, Inc. Figure 8.10 Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts; H atoms not shown Porphyrin ring: light-absorbing “head” of molecule; note magnesium atom at center CH 3 in chlorophyll a CHO in chlorophyll b CH 3

© 2014 Pearson Education, Inc. Figure 8.11 Photon (fluorescence) Ground state (b) Fluorescence Excited state Chlorophyll molecule Photon Heat e−e− (a) Excitation of isolated chlorophyll molecule Energy of electron

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) transfer the energy of photons to the reaction center © 2014 Pearson Education, Inc.

Figure 8.12 (b) Structure of a photosystem (a) How a photosystem harvests light Chlorophyll STROMA THYLAKOID SPACE Protein subunits STROMA THYLAKOID SPACE (INTERIOR OF THYLAKOID) Photosystem Photon Light- harvesting complexes Reaction- center complex Primary electron acceptor Special pair of chlorophyll a molecules Transfer of energy Pigment molecules Thylakoid membrane ee

© 2014 Pearson Education, Inc. Figure 8.12a (a) How a photosystem harvests light STROMA THYLAKOID SPACE (INTERIOR OF THYLAKOID) Photosystem Photon Light- harvesting complexes Reaction- center complex Primary electron acceptor Special pair of chlorophyll a molecules Transfer of energy Pigment molecules Thylakoid membrane e−e−

© 2014 Pearson Education, Inc. Figure 8.12b (b) Structure of a photosystem Chlorophyll STROMA THYLAKOID SPACE Protein subunits Thylakoid membrane

 A primary electron acceptor in the reaction center accepts excited electrons and is reduced as a result © 2014 Pearson Education, Inc.

 There are two types of photosystems in the thylakoid membrane  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 © 2014 Pearson Education, Inc.

 Photosystem I (PS I) is best at absorbing a wavelength of 700 nm  The reaction-center chlorophyll a of PS I is called P700 © 2014 Pearson Education, Inc.

Linear Electron Flow  Linear electron flow involves the flow of electrons through both photosystems to produce ATP and NADPH using light energy © 2014 Pearson Education, Inc.

 Linear electron flow can be broken down into a series of steps 1.A photon hits a pigment and its energy is passed among pigment molecules until it excites P680 2.An excited electron from P680 is transferred to the primary electron acceptor (we now call it P680  ) 3.H 2 O is split 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 © 2014 Pearson Education, Inc.

Figure 8.UN02 Calvin Cycle NADPH NADP  ATP ADP Light CO 2 [CH 2 O] (sugar) Light Reactions O2O2 H2OH2O

© 2014 Pearson Education, Inc. Figure Primary acceptor Photosystem II (PS II ) Light P680 Pigment molecules 1 2 e−e−

© 2014 Pearson Education, Inc. Figure Primary acceptor 2 H  O2O2  Photosystem II (PS II ) H2OH2O Light  2 1 P680 Pigment molecules 1 23 e−e− e−e− e−e−

© 2014 Pearson Education, Inc. Figure Primary acceptor 2 H  O2O2  ATP Photosystem II (PS II ) H2OH2O Light  2 1 P680 Pq Electron transport chain Cytochrome complex Pc Pigment molecules e−e− e−e− e−e−

© 2014 Pearson Education, Inc. Figure Primary acceptor 2 H  O2O2  ATP Photosystem II (PS II ) H2OH2O Light  2 1 P680 Pq Electron transport chain Cytochrome complex Pc Pigment molecules Primary acceptor Photosystem I (PS I ) P700 Light e−e− e−e− e−e− e−e−

© 2014 Pearson Education, Inc. Figure Primary acceptor 2 H  O2O2  ATP NADPH Photosystem II (PS II ) H2OH2O e−e− e−e− e−e− Light  2 1 P680 Pq Electron transport chain Cytochrome complex Pc Pigment molecules Primary acceptor Photosystem I (PS I ) e−e− P700 e−e− e−e− Fd Light Electron transport chain HH  NADP  reductase

4.Each electron “falls” down an electron transport chain from the primary electron acceptor of PS II to PS I 5.Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane; diffusion of H  (protons) across the membrane drives ATP synthesis © 2014 Pearson Education, Inc.

6.In PS I (like PS II), transferred light energy excites P700, causing it to lose an electron to an electron acceptor (we now call it P700  )  P700  accepts an electron passed down from PS II via the electron transport chain © 2014 Pearson Education, Inc.

7.Excited electrons “fall” down an electron transport chain from the primary electron acceptor of PS I to the protein ferredoxin (Fd) 8.The electrons are transferred to NADP , reducing it to NADPH, and become available for the reactions of the Calvin cycle  This process also removes an H  from the stroma © 2014 Pearson Education, Inc.

 The energy changes of electrons during linear flow can be represented in a mechanical analogy © 2014 Pearson Education, Inc.

Figure 8.14 Photosystem II Photosystem I NADPH Mill makes ATP Photon

A Comparison of Chemiosmosis in Chloroplasts and Mitochondria  chemiosmosis but use different sources of energy  Chem to chem, radiant to chem  Where is H+ pumped? © 2014 Pearson Education, Inc.

 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 © 2014 Pearson Education, Inc.

Figure 8.15 Electron transport chain Higher [H  ] P i HH CHLOROPLAST STRUCTURE Inter- membrane space MITOCHONDRION STRUCTURE Thylakoid space Inner membrane Matrix Key Lower [H  ] Thylakoid membrane Stroma ATP synthase ADP  HH Diffusion

© 2014 Pearson Education, Inc. Figure 8.15a Electron transport chain Higher [H  ] HH CHLOROPLAST STRUCTURE Inter- membrane space MITOCHONDRION STRUCTURE Thylakoid space Inner membrane Matrix Key Lower [H  ] Thylakoid membrane Stroma ATP synthase ADP  HH Diffusion P i

 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 © 2014 Pearson Education, Inc.

Figure 8.UN02 Calvin Cycle NADPH NADP  ATP ADP Light CO 2 [CH 2 O] (sugar) Light Reactions O2O2 H2OH2O

© 2014 Pearson Education, Inc. Figure 8.16 Photosystem II Photosystem I To Calvin Cycle HH THYLAKOID SPACE (high H  concentration) Thylakoid membrane STROMA (low H  concentration) ATP synthase NADPH e−e− Light NADP  ATP ADP  NADP  reductase Fd HH  Pq Pc Cytochrome complex 4 H  Light  2 H  O2O2 H2OH2O  H  e−e− 1 23 P i

© 2014 Pearson Education, Inc. Figure 8.16a Photosystem II Photosystem I HH THYLAKOID SPACE (high H  concentration) Thylakoid membrane STROMA (low H  concentration) ATP synthase e−e− Light ATP ADP  Fd Pq Pc Cytochrome complex 4 H  Light  2 H  O2O2 H2OH2O  H  e−e− 12 P i

© 2014 Pearson Education, Inc. Figure 8.16b 23 Photosystem I To Calvin Cycle HH ATP synthase NADPH Light NADP  ATP ADP  NADP  reductase Fd HH  Pc Cytochrome complex 4 H  THYLAKOID SPACE (high H  concentration) STROMA (low H  concentration) P i

Concept 8.3: The Calvin cycle uses the chemical energy of ATP and NADPH to reduce CO 2 to sugar  The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle  Unlike the citric acid cycle, the Calvin cycle is anabolic  It builds sugar from smaller molecules by using ATP and the reducing power of electrons carried by NADPH © 2014 Pearson Education, Inc.

 Carbon enters the cycle as CO 2 and leaves as a sugar named glyceraldehyde 3-phospate (G3P)  For net synthesis of one G3P, the cycle must take place three times, fixing three molecules of CO 2  The Calvin cycle has three phases  Carbon fixation  Reduction  Regeneration of the CO 2 acceptor © 2014 Pearson Education, Inc.

 Phase 1, carbon fixation, involves the incorporation of the CO 2 molecules into ribulose bisphosphate (RuBP) using the enzyme rubisco © 2014 Pearson Education, Inc.

Figure 8.UN03 Calvin Cycle NADPH NADP  ATP ADP Light CO 2 [CH 2 O] (sugar) Light Reactions O2O2 H2OH2O

© 2014 Pearson Education, Inc. Figure Input 3 Calvin Cycle as 3 CO 2 Rubisco Phase 1: Carbon fixation RuBP 3-Phosphoglycerate P P PP P

© 2014 Pearson Education, Inc. Figure P i NADPH Input 3 ATP Calvin Cycle as 3 CO 2 Rubisco Phase 1: Carbon fixation Phase 2: Reduction G3P Output Glucose and other organic compounds G3P RuBP 3-Phosphoglycerate 1,3-Bisphosphoglycerate 6 ADP NADP  P P P P P PP P P

© 2014 Pearson Education, Inc. Figure P i NADPH Input 3 ATP Calvin Cycle as 3 CO 2 Rubisco Phase 1: Carbon fixation Phase 2: Reduction Phase 3: Regeneration of RuBP G3P Output Glucose and other organic compounds G3P RuBP 3-Phosphoglycerate 1,3-Bisphosphoglycerate 6 ADP P 3 P P P 6 NADP  6 P 5 P G3P ATP 3 ADP 3 3 PP 1 P P

 Phase 2, reduction, involves the reduction and phosphorylation of 3-phosphoglycerate to G3P © 2014 Pearson Education, Inc.

 Phase 3, regeneration, involves the rearrangement of G3P to regenerate the initial CO 2 receptor, RuBP © 2014 Pearson Education, Inc.

Evolution of Alternative Mechanisms of Carbon Fixation in Hot, Arid Climates  Adaptation to dehydration is a problem for land 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 an apparently wasteful process called photorespiration © 2014 Pearson Education, Inc.

 In most plants (C 3 plants), initial fixation of CO 2, via rubisco, forms a three-carbon compound (3- phosphoglycerate)  In photorespiration, rubisco adds O 2 instead of CO 2 in the Calvin cycle, producing a two-carbon compound  Photorespiration decreases photosynthetic output by consuming ATP, O 2, and organic fuel and releasing CO 2 without producing any ATP or sugar © 2014 Pearson Education, Inc.

 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 © 2014 Pearson Education, Inc.

 C 4 plants minimize the cost of photorespiration by incorporating CO 2 into a four-carbon compound  An enzyme in the mesophyll cells has a high affinity for CO 2 and can fix carbon 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 C 4 Plants © 2014 Pearson Education, Inc.

Figure 8.18 Bundle- sheath cell Sugarcane CO 2 Pineapple CO 2 (a) Spatial separation of steps C4C4 CO 2 CAM Day Night Sugar Calvin Cycle Calvin Cycle Sugar Organic acid Organic acid Mesophyll cell (b) Temporal separation of steps

© 2014 Pearson Education, Inc. Figure 8.18a Sugarcane

© 2014 Pearson Education, Inc. Figure 8.18b Pineapple

© 2014 Pearson Education, Inc. Figure 8.18c Bundle- sheath cell CO 2 (a) Spatial separation of steps C4C4 CO 2 CAM Day Night Sugar Calvin Cycle Calvin Cycle Sugar Organic acid Organic acid Mesophyll cell (b) Temporal separation of steps 1212

CAM Plants  Some plants, including succulents, use crassulacean acid metabolism (CAM) to fix carbon  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 © 2014 Pearson Education, Inc.

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 the chloroplasts and in structures such as roots, tubers, seeds, and fruits  In addition to food production, photosynthesis produces the O 2 in our atmosphere © 2014 Pearson Education, Inc.

Figure 8.19 Photosystem II Electron transport chain Calvin Cycle NADPH Light NADP  ATP CO 2 H2OH2O ADP  3-Phosphpglycerate G3P RuBP Sucrose (export) Starch (storage) Chloroplast O2O2 Light Reactions: Photosystem I Electron transport chain P i

© 2014 Pearson Education, Inc. Figure 8.UN04

© 2014 Pearson Education, Inc. Figure 8.UN05 Photosystem II Photosystem I NADP  ATP Fd HH  Pq Cytochrome complex O2O2 H2OH2O Pc NADP  reductase NADPH Primary acceptor Electron transport chain Primary acceptor Electron transport chain

© 2014 Pearson Education, Inc. Figure 8.UN06 Calvin Cycle Regeneration of CO 2 acceptor Carbon fixation Reduction 1 G3P (3C) 3 CO 2 3  5C 6  3C 5  3C

© 2014 Pearson Education, Inc. Figure 8.UN07 pH 7 pH 4pH 8 pH 4