Chapt08 Lecture Photosynthesis 4 1

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    Copyright (c) The McGraw-Hill

    Companies, Inc. Permission requiredfor reproduction or display. 1

    CHAPTER 8

    PHOTOSYNTHESIS

    Prepared by

    Brenda Leady, Universi ty of Toledo

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    PHOTOSYNTHESISSWBAT:

    Describe overall equation of photosynthesis

    Distinguish: autotrophs, heterotrophs, photoautotrophs andchemoautotrophs

    Describe stages of photosynthesis (light-dependent reactionsand Calvin Cycle)

    Outline chemiosmosis in photophosphorilation

    Describe alternative ways of Carbon fixation (C4 and CAM

    plants)

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    PHOTOSYNTHESISSWBAT:

    Describe overall equation of photosynthesis

    Distinguish: autotrophs, heterotrophs, photoautotrophs andchemoautotrophs

    Describe stages of photosynthesis (light-dependent reactionsand Calvin Cycle)

    Outline chemiosmosis in photophosphorilation

    Describe alternative ways of Carbon fixation (C4 and CAM

    plants)

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    Trophic organization

    Heterotroph

    Must eat food, organic molecules from their

    environment, to sustain lifeAutotroph

    Make organic molecules from inorganicsources Photoautotroph

    Use light as a source of energy

    Green plants, algae, cyanobacteria

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    Photosynthesis

    Energy within light is captured and used to

    synthesize carbohydrates

    CO2+ H2O + light energy C6H12O6+ O2+ H2O

    CO2is reduced H2O is oxidized

    Energy from light drives this endergonic

    reaction

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    Link between Photosynthesis and Respiration:

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    Who does it?

    Plants

    Algae

    Cyanobacteria

    Photosynthetic bacteria

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    Biosphere

    Regions on the surface of the Earth and in

    the atmosphere where living organisms

    exist Largely driven by the photosynthetic

    power of green plants

    Cycle where cells use organic moleculesfor energy and plants replenish those

    molecules using photosynthesis

    Plants also produce oxygen

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    Chloroplast

    Organelles in plants and algae that carry

    out photosynthesis

    Chlorophyll- green pigment

    Majority of photosynthesis occurs in

    leaves in central mesophyll

    Stomata- carbon dioxide enters and

    oxygen exits leaf

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    Chloroplast anatomy

    Outer and inner membrane

    Intermembrane space

    Thylakoid membrane contains pigmentmolecules

    Thylakoid membrane forms thylakoids

    Enclose thylakoid lumen

    Granum- stack of thylakoids

    Stroma- fluid filled region betweenthylakoid membrane and inner membrane

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    http://localhost/var/www/apps/conversion/tmp/scratch_7/chapt04_lecture-2.ppt
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    2 stages of photosynthesis

    Light reactions

    Take place in thylakoid membranes

    Produce ATP, NADPH and O2

    Calvin cycle

    Occurs in stroma

    Uses ATP and NADPH to incorporate CO2into organic molecules

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    Light energy

    Type of electromagnetic radiation

    Travels as waves

    Short to long wavelengths

    Also behaves as particles- photons

    Shorter wavelengths have more energy

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    Photosynthetic pigments absorb some lightenergy and reflect others

    Leaves are green because they reflect green

    wavelengths

    Absorption boosts electrons to higher energylevels

    Wavelength of light that a pigment absorbs

    depends on the amount of energy needed to

    boost an electron to a higher orbital

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    After an electron absorbs energy, it is anexcited state and usually unstable

    Releases energy asHeatLight

    Excited electrons in pigments can be

    transferred to another molecule orcaptured

    Captured light energy can be transferredto other molecules to ultimately produce

    energy intermediates for cellular work

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    Pigments

    Chlorophyll a

    Chlorophyll b

    Carotenoids

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    Spectrophotometers are used tomeasure light absorption

    White

    light

    Refracting

    prism

    Chlorophyll

    solution Photoelectric

    tube

    The low transmittance

    (high absorption)reading indicates that

    chlorophyll absorbs

    most blue light.

    Slit moves to

    pass light

    of selectedwavelength

    Blue

    light

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    Absorption vs. action spectrum Absorption spectrum

    Wavelengths that are absorbed bydifferent pigments in the plant

    Action spectrumRate of photosynthesis by wholeplant at specific wavelengths

    The color of the pigment comes from the

    wavelengths of light reflected (in other words, those

    not absorbed). Chlorophyll, the green pigment

    common to all photosynthetic cells, absorbs all

    wavelengths of visible light except green

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    The color of the pigment comes

    from the wavelengths of light

    reflected (in other words, those

    not absorbed). Chlorophyll, the

    green pigment common to all

    photosynthetic cells, absorbs all

    wavelengths of visible light

    except green

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    Photosystems

    Thylakoid membrane:

    o Photosystem I (PSI)

    o Photosystem II (PSII)

    Each of them has a light harvesting complex

    (antenna complex) and a reaction center

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    Photosytem II (PSII)

    2 main components Light-harvesting complex or antenna complex

    Directly absorbs photons

    Energy transferred via resonance energy transfer to P680in the reaction center

    Reaction center P680 P680* (Relatively unstable)

    P680* actually releases its high-energy electronto theprimary electron acceptor and becomes P680+(more stable).

    P680 has to be regenerated, by replacing the electron so

    P680 can work again: : This missing electron of P680+isreplaced with a low-energy electronfrom water which yieldsoxygen gas

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    Photosystem II (PSII)

    Redox machine

    Recent research in biochemical

    composition of protein complex and role ofcomponents

    3 dimensional structure determined in

    2004 using x-ray crystallography

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    D1 and D2, contain the reaction center that carries out the redox reactions

    CP43 and CP47, bind the pigment molecules that form the light-harvesting complex wraparound D1 and D2 so that pigments in CP43 and CP47 can transfer energy to P680 byresonance energy transfer.Pp: primary electron acceptor: a chlorophyll molecule lacking Mg2+, called pheophytin (Pp).QA:plastoquinone molecule, designated QAQBanother plastoquinone molecule which can accept two high-energy electrons and bind

    two H+

    . QBcan diffuse away from the reaction center.

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    The oxidation of water occurs in a region called the manganese cluster. This site is

    located on the side of D1 that faces the thylakoid lumen.The manganese cluster has four Mn2+, one Ca2+, and one Cl.Two water molecules bind to this site.D1 catalyzes the removal of four low-energy electrons from the two water moleculesto create four H+and O2.Each low-energy electronis transferred, one at a time, to an amino acid in D1 (a

    tyrosine, Tyr) and then to P680+to produce P680.

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    Electrons accepted by primary electron acceptorin PSII are transferred to a pigment molecule in

    the reaction center of PSI

    Electron releases some of its energy along the

    way

    Establishes H+electrochemical gradient

    ATP synthesis uses chemiosmotic mechanism similar

    to mitochondria

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    Photosystem I (PSI)

    Key role to make NADPH

    Light striking light-harvesting complex ofPSI transfers energy to a reaction center

    High energy electron removed from P700and transferred to a primary electronacceptor

    NADP+reductaseNADP++ 2 electrons + H + NADPH

    P700+replaces its electrons fromplastocyanin

    No splitting water, no oxygen gas formed

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    Summary1. O2produced in thylakoid lumenby oxidation of

    H2O by PSII

    2 electrons transferred to P680+

    2. ATP produced in stroma by H+electrochemicalgradient(chemiosmosis) due to:

    1. Splitting of water places H+in the lumen

    2. High-energy electrons move from PSII to PSI,pumping H+into the lumen

    3. Formation of NADPH consumes H+in the stroma

    3. NADPH produced in the stromafrom high-energyelectrons that start in PSII and boosted in PSI

    NADP++ 2 electrons + H + NADPH

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    QBcan diffuse away from the reaction center PSII carrying a pair of electronsEach electron enters an electron transport chain: a series of electron carriers located in the thylakoidmembrane. The electron transport chain functions similarly to the one found in mitochondria.The electrons go from QB, to a cytochrome complex, then to plastocyanin (Pc), a small protein; and then toPSIAlong its journey from PSII to PSI, the electron releases some of its energy at particular steps and is transferred tothe next component that has a higher electronegativity. In the cyochrome complex the energy released is harnessedto pump H+into the thylakoid lumen. One result of the electron movement is to establish a H+electrochemical gradient

    From FSII to FSI

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    When light strikes the light-harvesting complex of PSI, this energy is also transferred to a reactioncenter, where a high-energy electron is removed from a pigment molecule, designated P700, andtransferred to a primary electron acceptor.

    A protein called ferredoxin (Fd) can accept two high-energy electrons, one at a time, from the primaryelectron acceptor.

    Fd then transfers the two electrons to the enzyme NADP+reductase. This enzyme transfers the twoelectrons to NADP+and together with a H+creates NADPH. The formation of NADPH results in fewerH+in the stroma and thereby contributes to the formation of a H+electrochemical gradient across thethylakoid membrane.

    FSI

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    Synthesis of ATP in chloroplasts is achieved by a chemiosmotic mechanism similar to that used tomake ATP in mitochondria.

    ATP synthesis is driven by the flow of H+from the thylakoid lumen into the stroma via ATPsynthase

    A H+gradient is generated in three ways:(1) the splitting of water, which places H+in the thylakoid lumen;(2) the movement of high-energy electrons from photosystem II to photosystem I, which pumps H+into

    the thylakoid lumen(3) the formation of NADPH, which consumes H+in the stroma.

    Chemiosmosis

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    Cyclic and noncyclic electron flow

    Noncyclic

    Electrons begin at PSII and eventually

    transfer to NADPHLinear process produces ATP and NADPH in

    equal amounts

    Cyclic photophosphorylation

    Electron cycling releases energy to transportH+into lumen driving synthesis of ATP

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    Cyclic photophosphorylation

    Electron cycling releases energy to transport H+into lumen driving synthesis of ATP.

    NADPH IS NOT produced

    H2O is not splitter and O2 is not produced

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    Photosystems II and I work together to produce

    ATP and NADPH

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    Z scheme (energy curve) Robin Hill and Fay Bendall

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    Photosynthesis animations

    Photosynthetic Electron Transport and ATP Synthesis

    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%20Synthesishttp://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
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    The cytochrome complexes of mitochondria and

    chloroplasts have evolutionarily related proteins in

    common

    Homologous genes are similar because theyare derived from a common ancestor

    Comparing the electron transport chains ofmitochondria and chloroplasts revealshomologous genes

    Family of cytochrome b-type proteins plays

    similar but specialized roles

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    Calvin cycle

    ATP and NADPH used to make

    carbohydrates

    Somewhat similar to citric acid cycle

    CO2incorporated into carbohydrates

    Precursors to all organic molecules

    Energy storage

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    Overview of Calvin Cycle

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    CO2incorporation

    Also called Calvin-Benson cycle

    Requires massive input of energy: For every 6CO

    2

    incorporated, 18 ATP and 12 NADPH used

    Glucose is not directly made. Instead, moleculesof glyceraldehyde-3-phosphate, which areproducts of the Calvin cycle, are used as startingmaterials for the synthesis of glucose and othermolecules, including sucrose. After glucose molecules aremade, they may be linked together to form a polymer of glucose called starch, whichis stored in the chloroplast for later use. Alternatively, the disaccharide sucrose maybe made and transported out of the leaf to other parts of the plant.

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    3 phases

    1. Carbon fixation CO2incorporated in ribulose bisphosphate (RuBP)

    using RuBP carboxylase/oxygenase (rubisco)

    6 carbon intermediate splits into 2 3PG

    2. Reduction and carbohydrate production ATP is used to convert 3 phosphoglycerate (3PG)

    into 1,3-bisphosphoglycerate (1,3 BPG)

    NADPH electrons reduce it to glyceraldehyde 3 P(G3P)

    6 CO2 12 G3P 2 for carbohydrates

    10 for regeneration

    3. Regeneration of RuBP

    10 G3P converted into 6 RuBP using 6 ATP

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    Photosynthesis animations

    calvinCycle.swf

    http://highered.mcgraw-hill.com/olcweb/cgi/pluginpop.cgi?it=swf::525::530::/sites/dl/free/0072464631/291136/calvinCycle.swf::calvinCycle.swfhttp://highered.mcgraw-hill.com/olcweb/cgi/pluginpop.cgi?it=swf::525::530::/sites/dl/free/0072464631/291136/calvinCycle.swf::calvinCycle.swf
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    The Calvin cycle was determined by isotope

    labeling methods

    14C-labeled CO2injected into cultures of greenalgae

    Allowed to incubate different lengths of time

    Separated newly made radiolabeled moleculesusing two-dimensional paper chromatography

    Autoradiography- radiation from 14C-labeled

    molecules makes dark spots on the film Identified 14C-labeled spots and the order they

    appeared

    Calvin awarded Nobel Prize in 1961

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    Variations in photosynthesis

    Certain environmental conditions can

    influence both the efficiency and way the

    Calvin cycle worksLight intensity

    Temperature

    Water availability

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    Photorespiration

    RuBP + CO22 3PGRubisco functions as a carboxylase

    C3plants make 3PG

    Rubisco can also be an oxygenaseAdds O2to RuBP eventually releasing CO2

    Photorespiration

    Using O2

    and liberating CO2

    is wasteful

    More likely in hot and dry environment

    Favored when CO2low and O2high

    Rubisco: RuBP carboxylase/oxygenase

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    54Wheat plants Oak leaves

    C3 plants

    90% of plants are C3

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    C4plants: To avoid photorespiration

    C4plants make a 4 carbon compound inthe first step of carbon fixation

    Hatch-Slack pathway

    Leaves have 2-cell layer organizationMesophyll cells

    CO2enters via stomata and 4 carbon compoundformed (PEP carboxylase does not promote

    photorespiration)Bundle-sheath cells

    4 carbon compound transferred that releasessteady supply CO2

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    C4 plants

    In warm and dry climates, C4plants have an advantage. During the day, they cankeep their stomata partially closed to conserve water. Furthermore, they can avoidphotorespiration. C4plants are well adapted to habitats with high daytimetemperatures and intense sunlight.

    Examples of C4plants are sugarcane, crabgrass, and corn.

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    CAM plants

    Some C4plants separate processes using time

    Crassulacean Acid Metabolism

    CAM plants open their stomata at night CO2enters and is converted to malate

    Stomata close during the day to conserve water

    Malate broken down into CO2to drive Calvincycle

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    Process in separated cells Process at different times

    C4 CAM

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    Adaptations for hot weather: C4 and CAM

    plants

    C4: corn, sugarcane andsorghum

    CAM: succulents (aloe, jade),pineapple, cactiCAM = crassulasean acid

    metabolism

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    C4 and CAM compared

    Both fix CO2into a C4 compound

    Then CO2is transferred to the Calvin cycle

    In C4 plants there is a spatial separation (2 celltypes)

    In CAM plants there is a temporal separation (C4accumulates at night, Calvin cycle during theday)

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    What is better? C3 or C4 strategy?

    In warm dry climates C4plants have the

    advantage in conserving water and preventing

    photorespiration

    In cooler climates, C3plants use less energy to

    fix CO2

    90% of plants are C3