Chapter 6 Where It Starts – Photosynthesis Albia Dugger • Miami Dade College 6.1 Biofuels • Coal, petroleum, and natural gas are fossil fuels – the remains of ancient forests, a limited resource • Biofuels such as oils, gases, or alcohols are made from organic matter that is not fossilized – a renewable resource • In the United States, biofuels are produced mainly from food crops such as corn, soybeans, and sugarcane • Researchers are looking for ways to use non-food-plant matter such as switchgrass and agricultural wastes Biofuels Research Autotrophs and Heterotrophs • Autotrophs harvest energy directly from the environment, and obtain carbon from inorganic molecules • Plants and most other autotrophs make their own food by photosynthesis, a process which uses the energy of sunlight to assemble carbohydrates from carbon dioxide and water • Animals and other heterotrophs get energy and carbon by breaking down organic molecules assembled by other organisms 6.2 Sunlight as an Energy Source • Energy flow through nearly all ecosystems on Earth begins when photosynthesizers intercept energy from the sun • Photosynthetic organisms use pigments to capture the energy of sunlight and convert it to chemical energy – the energy stored in chemical bonds Properties of Light • Visible light is part of an electromagnetic spectrum of energy radiating from the sun • Travels in waves • Organized into photons • Wavelength • The distance between the crests of two successive waves of light (nm) • Shorter wavelength have greater energy Electromagnetic Spectrum of Radiant Energy shortest wavelengths (highest energy) gamma rays x-rays range of heat range of most radiation escaping from reaching Earth’s surface Earth’s surface visible light near-infrared ultraviolet radiation radiation 400 nm 500 nm longest wavelengths (lowest energy) radiation microwaves radio waves infrared 600 nm 700 nm Wavelength and Energy Pigments: The Rainbow Catchers • Different wavelengths form colors of the rainbow • Photosynthesis uses wavelengths of 380-750 nm • Pigment • An organic molecule that selectively absorbs light of specific wavelengths • Chlorophyll a • The most common photosynthetic pigment • Absorbs violet and red light (appears green) Photosynthetic Pigments • Collectively, chlorophyll and accessory pigments absorb most wavelengths of visible light • Certain electrons in pigment molecules absorb photons of light energy, boosting electrons to a higher energy level • Energy is captured and used for photosynthesis Some Photosynthetic Pigments Take-Home Message: How do photosynthesizers absorb light? • Energy radiating from the sun travels through space in waves and is organized as packets called photons • The spectrum of radiant energy from the sun includes visible light; humans perceive different wavelengths of visible light as different colors; the shorter the wavelength, the greater the energy • Pigments absorb light at specific wavelengths; photosynthetic species use pigments such as chlorophyll a to harvest the energy of light for photosynthesis 6.3 Exploring the Rainbow • Photosynthetic pigments work together to harvest light of different wavelengths • Engelmann identified colors of light that drive photosynthesis (violet and red) by using a prism to divide light into colors – algae using these wavelengths gave off the most oxygen Photosynthesis and Wavelengths of Light bacteria alga 400 nm 500 nm 600 nm Wavelength 700 nm ANIMATED FIGURE: T. Englemann's experiment Absorption Spectra • Most photosynthetic organisms use a combination of pigments to drive photosynthesis • An absorption spectrum shows which wavelengths each pigment absorbs best • Organisms in different environments use different pigments Absorption Spectra chlorophyll b phycoerythrobilin phycocyanobilin Light absorption β-carotene 400 nm chlorophyll a 500 nm 600 nm Wavelength 700 nm Take-Home Message: Why do cells use more than one photosynthetic pigment? • A combination of pigments allows a photosynthetic organism to most efficiently capture the particular range of light wavelengths that reaches the habitat in which it evolved 6.4 Overview of Photosynthesis • In plants and other photosynthetic eukaryotes, photosynthesis occurs in chloroplasts • Photosynthesis occurs in two stages Two Stages of Photosynthesis • Light-dependent reactions (noncyclic pathway) • First stage of photosynthesis • Light energy is transferred to ATP and NADPH • Water molecules are split, releasing O2 • Light-independent reactions • Second stage of photosynthesis • Energy in ATP and NADPH drives synthesis of glucose and other carbohydrates from CO2 and water Summary: Photosynthesis 6CO2 + 6H2O → light energy → C6H12O6 + 6O2 The Chloroplast • Chloroplast • An organelle that specializes in photosynthesis in plants and many protists • Thylakoid membrane • Folded membrane that make up thylakoids • Contains clusters of light-harvesting pigments that absorb photons of different energies and convert light energy into chemical energy (first stage of photosynthesis) The Chloroplast • Stroma • A semifluid matrix surrounded by the two outer membranes of the chloroplast • Sugars are built in the stroma (second stage of photosynthesis) two outer membranes of chloroplast stroma part of thylakoid membrane system: thylakoid compartment, cutaway view Figure 6-5b p105 INTERACTION: Structure of a chloroplast To play movie you must be in Slide Show Mode PC Users: Please wait for content to load, then click to play Mac Users: CLICK HERE Take-Home Message: Where do the reactions of photosynthesis take place? • In the first stage of photosynthesis, light energy drives the formation of ATP and NADPH, and oxygen is released; in eukaryotic cells, these light-dependent reactions occur at the thylakoid membrane of chloroplasts • The second stage of photosynthesis, the light-independent reactions, occur in the stroma of chloroplasts; ATP and NADPH drive the synthesis of carbohydrates 3D ANIMATION: Photosynthesis Bio Experience 3D 6.5 Light-Dependent Reactions • Light-dependent reactions convert light energy to the energy of chemical bonds • Photons boost electrons in pigments to higher energy levels • Light-harvesting complexes absorb the energy • Electrons are released from special pairs of chlorophyll a molecules in photosystems • Electrons may be used in noncyclic or cyclic pathways of ATP formation Figure 6-7 p106 The Thylakoid Membrane photosystem light-harvesting complex The Noncyclic Pathway • Photosystems (type II and type I) contain “special pairs” of chlorophyll a molecules that eject electrons • Electrons lost from photosystem II are replaced by photolysis of water molecules – the process by which light energy breaks down a water molecule into hydrogen and oxygen • Electrons lost from a photosystem enter an electron transfer chain (ETC) in the thylakoid membrane The Noncyclic Pathway • In the ETC, electron energy is used to build up a H+ gradient across the membrane • H+ flows through ATP synthase, which attaches a phosphate group to ADP • ATP is formed in the stroma by chemiosmosis, or electron transfer phosphorylation The Noncyclic Pathway • Electrons from the first electron transfer chain (from photosystem II) are accepted by photosystem I • Electrons ejected from photosystem I enter a different electron transfer chain in which the coenzyme NADP+ accepts the electrons and H+, forming NADPH • ATP and NADPH are the energy products of light-dependent reactions in the noncyclic pathway Noncyclic Pathway of Photosynthesis H+ light energy electron transfer chain light energy to second stage of reactions ADP + Pi ATP synthase photosystem II photosystem I thylakoid compartment stroma The Cyclic Pathway • When NADPH accumulates in the stroma, the noncyclic pathway stalls • A cyclic pathway runs in type I photosystems to make ATP; electrons are cycled back to photosystem I and NADPH does not form What happens during the light-dependent reactions of photosynthesis? Take-Home Message: • In light-dependent reactions, chlorophylls and other pigments in thylakoid membrane transfer light energy to photosystems • Photosystems eject electrons that enter electron transfer chains in the membrane; electron flow through ETCs sets up hydrogen ion gradients that drive ATP formation • In the noncyclic pathway, oxygen is released and electrons end up in NADPH • A cyclic pathway involving only photosystem I allows the cell to continue making ATP when the noncyclic pathway is not running; NADPH does not form; O2 is not released 3D ANIMATION: Photophosphorylation ANIMATED FIGURE: Noncyclic pathway of electron flow To play movie you must be in Slide Show Mode PC Users: Please wait for content to load, then click to play Mac Users: CLICK HERE 6.6 Energy Flow in Photosynthesis • Energy flow in the light-dependent reactions is an example of how organisms harvest energy from their environment Photophosphorylation • Photophosphorylation is a light-driven reaction that attaches a phosphate group to a molecule • In noncyclic photophosphorylation, electrons move from water to photosystem II, to photosystem I, to NADPH • In cyclic photophosphorylation, electrons cycle within photosystem I Excited P700 energy Excited P680 P700 (photosystem I) P680 (photosystem II) light energy light energy Energy flow in the noncyclic reactions of photosynthesis Stepped Art Figure 6-9a p108 energy Excited P700 P700 (photosystem I) light energy Energy flow in the cyclic reactions of photosynthesis Stepped Art Figure 6-9b p108 How does energy flow during the reactions of photosynthesis? Take-Home Message: • Light provides energy inputs that keep electrons flowing through electron transfer chains • Energy lost by electrons as they flow through the chains sets up a hydrogen ion gradient that drives the synthesis of ATP alone, or ATP and NADPH 6.7 Light-Independent Reactions • The cyclic, light-independent reactions of the Calvin-Benson cycle are the “synthesis” part of photosynthesis • Calvin-Benson cycle • Enzyme-mediated reactions that build sugars in the stroma of chloroplasts Carbon Fixation • Carbon fixation • Extraction of carbon atoms from inorganic sources (atmosphere) and incorporating them into an organic molecule • Builds glucose from CO2 • Uses bond energy of molecules formed in light-dependent reactions (ATP, NADPH) The Calvin-Benson Cycle • The enzyme rubisco attaches CO2 to RuBP • Forms two 3-carbon PGA molecules • PGAL is formed • PGAs receive a phosphate group from ATP, and hydrogen and electrons from NADPH • Two PGAL combine to form a 6-carbon sugar • Rubisco is regenerated 1 4 Calvin– Benson Cycle 2 other molecules 3 glucose Stepped Art Figure 6-10 p109 ANIMATED FIGURE: Photosynthesis overview To play movie you must be in Slide Show Mode PC Users: Please wait for content to load, then click to play Mac Users: CLICK HERE Take-Home Message: What happens in light- independent reactions of photosynthesis? • Light-independent reactions of photosynthesis run on the bond energy of ATP and energy of electrons donated by NADPH; both formed in the light-dependent reactions • Collectively called the Calvin–Benson cycle, these carbonfixing reaction use hydrogen (from NADPH), and carbon and oxygen (from CO2) to build sugars 6.8 Adaptations: Different Carbon-Fixing Pathways • Environments differ, and so do details of photosynthesis: • C3 plants • C4 plants • CAM plants Stomata • Stomata • Small openings through the waxy cuticle covering epidermal surfaces of leaves and green stems • Allow CO2 in and O2 out • Close on dry days to minimize water loss C3 Plants • C3 plants • Plants that use only the Calvin–Benson cycle to fix carbon • Forms 3-carbon PGA in mesophyll cells • Used by most plants, but inefficient in dry weather when stomata are closed • Example: barley Photorespiration • When stomata are closed, CO2 needed for light-independent reactions can’t enter, O2 produced by light-dependent reactions can’t leave • Photorespiration • At high O2 levels, rubisco attaches to oxygen instead of carbon • CO2 is produced rather than fixed A C3 Plant: Barley palisade mesophyll cell spongy mesophyll cell mesophyll cell CO2 O2 glycolate RuBP Calvin– Benson PGA Cycle ATP NADPH B On dry days, stomata close and oxygen accumulates inside leaves. The excess causes rubisco to attach oxygen instead of carbon to RuBP. This is photorespiration, and it makes sugar production inefficient in C3 plants. sugars Figure 6-11b p110 C4 Plants • C4 plants • Plants that have an additional set of reactions for sugar production on dry days when stomata are closed; compensates for inefficiency of rubisco • Forms 4-carbon oxaloacetate in mesophyll cells, then bundle-sheath cells make sugar • Examples: Corn, switchgrass, bamboo A C4 Plant: Millet mesophyll cell bundle-sheath cell B C4 plants. Oxygen also builds up inside leaves when stomata close during photosynthesis. An additional pathway in these plants keeps the CO2 concentration high enough in bundle-sheath cells to prevent photorespiration. CO2 from inside plant mesophyll cell oxaloacetate bundle-sheath cell C4 Cycle CO2 PGA RuBP Calvin– Benson Cycle sugars Figure 6-12b p110 CAM Plants • CAM plants (Crassulacean Acid Metabolism) • Plants with an alternative carbon-fixing pathway that allows them to conserve water in climates where days are hot • Forms 4-carbon oxaloacetate at night, which is later broken down to CO2 for sugar production • Example: succulents, cactuses A CAM Plant: Jade Plant mesophyll cell CO2 from outside plant oxaloacetate C4 Cycle night day CO2 PGA RuBP Calvin– Benson Cycle sugars Figure 6-13a p111 ANIMATED FIGURE: Carbon-fixing adaptations To play movie you must be in Slide Show Mode PC Users: Please wait for content to load, then click to play Mac Users: CLICK HERE Take-Home Message: How do carbon-fixing reactions vary? • When stomata are closed, oxygen builds up inside leaves of C3 plants; rubisco then can attach oxygen (instead of carbon dioxide) to RuBP; photorespiration reduces the efficiency of sugar production, so it can limit the plant’s growth • Plants adapted to dry conditions limit photorespiration by fixing carbon twice: C4 plants separate the two sets of reactions in space; CAM plants separate them in time Biofuels Revisited • The first cells on Earth were chemoautotrophs that extracted energy and carbon from inorganic molecules in the environment, such as hydrogen sulfide and methane • The evolution of photosynthesis dramatically and permanently changed Earth’s atmosphere • Photoautotrophs use photosynthesis to make food from CO2 and water, releasing O2 into the atmosphere Earth’s Early Atmosphere Earth With an Oxygen Atmosphere Effects of Atmospheric Oxygen • Selection pressure on evolution of life • Oxygen radicals • Development of ATP-forming reactions • Aerobic respiration • Formation of ozone (O3) layer • Protection from UV radiation The Atmospheric Carbon Cycle • Photosynthesis removes carbon dioxide from the atmosphere, and locks carbon atoms in organic compounds • Aerobic organisms break down organic compounds for energy, and release CO2 into the atmosphere • Since photosynthesis evolved, these two processes have constituted a more or less balanced cycle of the biosphere • Today, Earth’s atmosphere is out of balance – the level of CO2 is increasing, mainly as a result of human activity Fossil Fuels • When we burn fossil fuels, carbon that has been locked for hundreds of millions of years is released back into the atmosphere, mainly as carbon dioxide • Today, we release about 28 billion tons of carbon dioxide into the atmosphere each year, more than ten times the amount we released in the year 1900 • Increased atmospheric CO2 contributes to global warming and disrupts natural biological systems Fossil Fuel Emissions Renewable Energy Sources • Biofuels are a renewable source of energy • The carbon in plant matter comes from atmospheric CO2, fixed by photosynthesis • Making biofuel production economically feasible is a high priority for today’s energy researchers