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PHOTOSYNTHESIS: ACQUIRING ENERGY FROM THE SUN CHAPTER 6 AN OVERVIEW OF PHOTOSYNTHESIS • Most of the energy used by almost all living cells ultimately comes from the sun. • Plants, algae, and some bacteria capture the sunlight energy by a process called photosynthesis. • Only about 1% of the available energy in sunlight is captured. Cross-section of leaf Cuticle Epidermis Mesophyll Vascular bundle Stoma Bundle Vacuole Nucleus sheath Cell wall Thylakoid Inner membrane Outer membrane Granum Stroma Chloroplast Chloroplasts Mesophyll cell AN OVERVIEW OF PHOTOSYNTHESIS • The leaf cells of plants contain chloroplasts. • The chloroplast contains internal membranes called thylakoids. • The thylakoids are stacked together in columns called grana. • The stroma is a semi liquid substance that surrounds the thylakoids. AN OVERVIEW OF PHOTOSYNTHESIS • The photosystem is the starting point of photosynthesis. • It is a network of pigments in the membrane of the thylakoid. • The primary pigment of a photosystem is chlorophyll. • The pigments act as an antenna to capture energy from sunlight. • Individual chlorophyll pigments pass the captured energy between them. AN OVERVIEW OF PHOTOSYNTHESIS • Photosynthesis takes place in three stages: 1. Capturing energy from sunlight. 2. Using the captured energy to produce ATP and NADPH. 3. Using the ATP and NADPH to make carbohydrates from CO2 in the atmosphere. AN OVERVIEW OF PHOTOSYNTHESIS • The process of photosynthesis is divided into two types of reactions. • Light-dependent reactions take place only in the presence of light and produce ATP and NADPH. • Light-independent reactions do not need light to occur and result in the formation of organic molecules. • More commonly known as the Calvin cycle. AN OVERVIEW OF PHOTOSYNTHESIS • The overall reaction for photosynthesis may be summarized by this simple equation: 6 CO2 + 6 H2O carbon dioxide water + Light energy C6H12O6 glucose + 6 O2 oxygen HOW PLANTS CAPTURE ENERGY FROM SUNLIGHT • Light is comprised of packets of energy called photons. • Sunlight has photons of varying energy levels. • The possible range of energy levels is represented by an electromagnetic spectrum. • Human eyes only perceive photons of intermediate energy levels. • This range of the spectrum is known as visible light. Increasing energy of photon Increasing wave length 0.001 nm 1 nm 1,000 nm 1 cm 10 nm 0.01 cm 1m 100 m Radioactive X-ray Light People Microwave FM radio Radar ovens elements machines bulbs AM radio X UV Gamma rays rays light Infrared Microwaves Radio waves Visible light 400 nm 430 nm 500 nm 560 nm 600 nm 650 nm 740 nm HOW PLANTS CAPTURE ENERGY FROM SUNLIGHT • Pigments are molecules that absorb light energy. • The main pigment in plants is chlorophyll. • Chlorophyll absorbs light at the ends of the visible spectrum, mainly blue and red light. HOW PLANTS CAPTURE ENERGY FROM SUNLIGHT • Plants also contain other pigments, called accessory pigments, that absorb light levels that chlorophyll does not. • These pigments give color to flowers, fruits, and vegetables. • They are present in leaves too, but are masked by chlorophyll until the fall when the chlorophyll is broken down. WHY ARE PLANTS GREEN? 1. All wave lengths except 500- to600-nanometer wave lengths (green) absorbed by leaf 4. Brain perceives “green” 2. Green reflected by leaf 3. Green absorbed by eye pigment ABSORPTION SPECTRA OF CHLOROPHYLLS AND CAROTENOIDS Relative light absorption CarotenoidsChlorophyll bChlorophyll a 400 450 500 550 600 650 Wavelength (nm) 700 ORGANIZING PIGMENTS INTO PHOTOSYSTEMS • In plants, the light-dependent reactions occur within a complex of proteins and pigments called a photosystem. ORGANIZING PIGMENTS INTO PHOTOSYSTEMS Electron • Light energy is first captured acceptor Reaction by any one of the chlorophyll Photon center e– pigments. chlorophyll e– • The energy is passed along to other pigments until it reaches the reaction center chlorophyll molecule. Chlorophyll • The reaction center then molecules releases an excited electron, Photosystem which is then transferred to an electron acceptor. • The excited electron that is lost is then replaced by an electron donor. Electro donor ORGANIZING PIGMENTS INTO PHOTOSYSTEMS • The light-dependent reactions in plants and algae use two photosystems. • Photosystem II • Captures a photon of light and releases an excited electron to the electron transport system (ETS). • The ETS then produces ATP. ORGANIZING PIGMENTS INTO PHOTOSYSTEMS Energy of electrons • Photosystem I • Absorbs another photon of light and releases an excited electron to another ETS. • The ETS produces NADPH. • The electron from photosystem II replaces the electron from the reaction center. Excited reaction center Excited reaction center 4 e– e– Photon Reaction center P680 1 Photosystem II Reaction center 3 H+ Proton gradient Water-splitting formed for ATP synthesis enzyme – e 2H2O 4H+ + O2 Electron transport system 5 NADP+ + H+ e– ATP 2 e– NADPH Photon P700 Photosystem I Electron transport system HOW PHOTOSYSTEMS CONVERT LIGHT TO CHEMICAL ENERGY • Plants produce both ATP and NADPH by noncyclic photophosphorylation. • The excited electrons flow through both photosystems and end up in NADPH. • High energy electrons generated by photosystem II are used to make ATP and are then passed along to photosystem I to drive the production of NADPH. THE PHOTOSYNTHETIC ELECTRON TRANSPORT SYSTEM Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Stroma Photon Calvin cycle Antenna complex Thylakoid membrane ATP Photon H+ + NADP+ NADP H+ NADP e– e– e– e– Light-dependent reactions 2H2O Thylakoid s pace Proton gradient H+ Water-splitting enzyme O2 H+ 4 H+ Photosystem II H+ Electron transport system H+ Photosystem I H+ Thylakoid space Electron transport system HOW PHOTOSYSTEMS CONVERT LIGHT TO CHEMICAL ENERGY • As a result of the proton pump of the ETS, a large concentration of protons builds up in the thylakoid space. • The thylakoid membrane is impermeable to protons. • Protons can only re-enter the stroma by traveling through a protein channel called ATP synthase. • As the protons follow their concentration gradient and pass through ATP synthase channels, ADP is phosphorylated into ATP. • This process is called chemiosmosis. CHEMIOSMOSIS IN A CHLOROPLAST Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Stroma Calvin cycle H+ Photon ADP ATP H+ H+ H+ ATP NADPH Light-dependent reactions Thylakoid space e– e– 2H2O H+ Thylakoid O2 space 4 H+ Photosystem II H+ H+ Electron transport system ATP synthase http://highered.mcgraw-hill.com/olcweb/cgi/pluginpop.cgi?it=swf::535::535::/sites/dl/free/0072437316/120072/bio13.swf::Photosynthetic%20Electron%20Transport%20and%20ATP%20Synthesis BUILDING NEW MOLECULES • Cells use the products of the light-dependent reactions to build organic molecules. • ATP is needed to drive endergonic reactions. • NADPH is needed to provide reducing power in the form of hydrogens. BUILDING NEW MOLECULES • The synthesis of new molecules employs the light-independent, or Calvin cycle, reactions. • These reactions are also known as C3 photosynthesis. BUILDING NEW MOLECULES • The Calvin cycle reactions occur in 3 stages • 1. Carbon fixation • Carbon from CO2 in the air is attached to an organic molecule, RuBP. • 2. Making sugars • The carbons are shuffled about through a series of reactions to make sugars. • 3. Reforming RuBP • The remaining molecules are used to reform RuBP. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 1 2 3 CO2 3 P 3 RuBP (Starting material) P 6 3-phosphoglycerate 6 3-phosphoglycerate 6 3 RuBP (Starting material) ATP 6 NADPH 3 P P Glyceraldehyde 3-phosphate 6 P 1 ATP 5 Glyceraldehyde 3-phosphate Glyceraldehyde 3-phosphate Glucose The Calvin cycle begins when a carbon atom from a CO2 molecule is added to a five-carbon molecule (the starting material). The resulting six-carbon molecule is unstable and immediately splits into three-carbon molecules. (Three “turns” of the cycle are indicated here with three molecules of CO2 entering the cycle.) Then, through a series of reactions, energy from ATP and hydrogens from NADPH (the products of the lightdependent reactions) are added to the three-carbon molecules. The nowreduced three-carbon molecules either combine to make glucose or are used to make other molecules. Most of the reduced three-carbon molecules are used to regenerate the five-carbon starting material, thus completing the cycle. http://highered.mcgraw-hill.com/sites/0070960526/student_view0/chapter5/animation_quiz_1.htmla BUILDING NEW MOLECULES • The Calvin cycle must “turn” 6 times in order to form a new glucose molecule. • Only one carbon is added from CO2 per turn. • The Calvin cycle also recycles reactants needed for the light-dependent reactions. • It returns ADP so that it is available for chemiosmosis in photosystem II. • It returns NADP+ back to the ETS of photosystem I. REACTIONS OF THE CALVIN CYCLE Stroma of chloroplast 3 CO2 Calvin cycle P AT P NADPH Light-dependent reactions Thylakoid space • Rubisco is thought to be the most abundant protein on earth 6 3-phosphoglycerate Rubisco 3 RuBP 6 ATP Carbon fixation 6 ADP 3 ADP P 3 ATP Reforming RuBP P 6 1,3-bisphosphoglycera 2 Pi 6 NADPH Making sugars 6 NADP 6 Pi P P 5 Glyceraldehyde 3-phosphate 6 Glyceraldehyde 3-phosphate P 1 Glyceraldehyde 3-phosphate Glucose and other sugars PHOTORESPIRATION: PUTTING THE BRAKES ON PHOTOSYNTHESIS • Many plants have trouble carrying out C3 photosynthesis when it is hot. • Plants close openings in their leaves, called stomata (singular, stoma), in order to prevent water loss. • The closed stoma also prevent gas exchange. • O2 levels build up inside the leaves while the concentrations of CO2 fall. • The enzyme rubisco fixes oxygen instead of carbon. • This process is called photorespiration and shortcircuits the Calvin cycle and photosynthesis. PLANT RESPONSE IN HOT, ARID WEATHER Leaf epidermis Heat Under hot, arid conditions, leaves lose water by evaporation through openings in the leaves called stomata. H2O H2O Stoma The stomata close to conserve water but as a result, O2 builds up inside the leaves, and CO2 cannot enter the leaves. This leads to photorespiration. O2 CO2 O2 CO2 CARBON FIXATION IN C4 PLANTS CO2 • Some plants have adapted to hot climates by performing C4 photosynthesis. • These C4 plants include sugarcane, corn, and many grasses. • They fix carbon using different types of cells and reactions than C3 plants and do not run out of CO2 even in hot weather. • CO2 becomes trapped in cells called bundle-sheath cells. Phosphoenol- Oxalopyruvate (PEP) acetate PPi + AMP Mesophyll cell Pi + ATP Pyruvate Malate Pyruvate Malate CO2 Calvin cycle Glucose Bundlesheath cell PHOTORESPIRATION: PUTTING THE BRAKES ON PHOTOSYNTHESIS • Another strategy to avoid a reduction in photosynthesis in hot weather occurs in many succulent (water-storing) plants, such as cacti and pineapples. • These plants undergo crassulacean acid metabolism (CAM). • Photosynthesis occurs via the C4 pathway at night and the C3 pathway during the day. COMPARING CARBON FIXATION IN C4 AND CAM PLANTS CO2 CO2 Mesophyll cell C4 pathway CO2 Bundlesheath cell C4 pathway Night COMesophyll 2 cell Calvin cycle Calvin cycle Glucose Glucose C4 plants CAM plants Day