LECTURE PRESENTATIONS
                                    For CAMPBELL BIOLOGY, NINTH EDITION
                Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson



Chapter 10

Photosynthesis




                                                                                                                    Lectures by
                                                                                                                    Erin Barley
                                                                                                            Kathleen Fitzpatrick

© 2011 Pearson Education, Inc.
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




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• Autotrophs sustain themselves without eating
       anything derived from other organisms
     • Autotrophs are the producers of the biosphere,
       producing organic molecules from CO2 and other
       inorganic molecules
     • Almost all plants are photoautotrophs, using the
       energy of sunlight to make organic molecules




© 2011 Pearson Education, Inc.
Figure 10.1
• Photosynthesis occurs in plants, algae, certain
       other protists, and some prokaryotes
     • These organisms feed not only themselves but
       also most of the living world




© 2011 Pearson Education, Inc.
Figure 10.2




                                                   (b) Multicellular
        (a) Plants                                     alga




                                                                  (d) Cyanobacteria
                                                                                      40 µm
               (c) Unicellular
                                 10 µm
                   protists

                                         (e) Purple sulfur
                                                           1 µm
                                             bacteria
• Heterotrophs obtain their organic material from
       other organisms
     • Heterotrophs are the consumers of the
       biosphere
     • Almost all heterotrophs, including humans,
       depend on photoautotrophs for food and O2




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Photosynthesis converts light energy to the
 chemical energy of food
     • Chloroplasts are structurally similar to and
       likely evolved from photosynthetic bacteria
     • The structural organization of these cells
       allows for the chemical reactions of
       photosynthesis




© 2011 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


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• CO2 enters and O2 exits the leaf through
       microscopic pores called stomata
     • The chlorophyll is in the membranes of
       thylakoids (connected sacs in the chloroplast);
       thylakoids may be stacked in columns called
       grana
     • Chloroplasts also contain stroma, a dense
       interior fluid




© 2011 Pearson Education, Inc.
Figure 10.4
                                                Leaf cross section
                                                Chloroplasts Vein


                                  Mesophyll




                                               Stomata
                                                         CO2   O2

                                        Chloroplast   Mesophyll
                                                        cell




                                             Outer
                                             membrane
                        Thylakoid        Intermembrane
              Stroma Granum Thylakoid    space                 20 µm
                             space      Inner
                                        membrane




                                 1 µm
Tracking Atoms Through Photosynthesis:
Scientific Inquiry
     • Photosynthesis is a complex series of reactions
       that can be summarized as the following
       equation:
              6 CO2 + 12 H2O + Light energy → C6H12O6 + 6 O2 + 6 H2O




© 2011 Pearson Education, Inc.
The Splitting of Water
     • Chloroplasts split H2O into hydrogen and oxygen,
       incorporating the electrons of hydrogen into
       sugar molecules and releasing oxygen as a by-
       product




© 2011 Pearson Education, Inc.
Figure 10.5




              Reactants:         6 CO2       12 H2O




              Products:    C6H12O6       6 H2O        6 O2
Photosynthesis as a Redox Process
     • Photosynthesis reverses the direction of electron
       flow compared to respiration
     • Photosynthesis is a redox process in which H2O
       is oxidized and CO2 is reduced
     • Photosynthesis is an endergonic process; the
       enery boost is provided by light




© 2011 Pearson Education, Inc.
Figure 10.UN01




                      becomes reduced

      Energy + 6 CO2 + 6 H2O            C6 H12 O6 + 6 O2
                                  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 H2O
                – Release O2
                – Reduce NADP+ to NADPH
                – Generate ATP from ADP by
                  photophosphorylation

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• The Calvin cycle (in the stroma) forms sugar
       from CO2, using ATP and NADPH
     • The Calvin cycle begins with carbon fixation,
       incorporating CO2 into organic molecules




© 2011 Pearson Education, Inc.
Figure 10.6-4

                   H2O                CO2



      Light

                             NADP+
                              ADP
                              +Pi
                                     Calvin
                   Light             Cycle
                 Reactions

                              ATP

                             NADPH



   Chloroplast

                    O2               [CH2O]
                                     (sugar)
The 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




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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



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• 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 particles, called photons




© 2011 Pearson Education, Inc.
Figure 10.7


                                                                1m
  10−5 nm 10−3 nm       1 nm        10 nm
                                      3
                                                  106 nm      (109 nm)    103 m

          Gamma                                        Micro-        Radio
                X-rays         UV         Infrared     waves         waves
           rays




                                Visible light



       380       450    500     550         600      650       700       750 nm
       Shorter wavelength                                  Longer wavelength
              Higher energy                                Lower energy
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




© 2011 Pearson Education, Inc.
Figure 10.8




                            Light
                                             Reflected
                                             light

              Chloroplast




                    Absorbed                   Granum
                    light


                               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




© 2011 Pearson Education, Inc.
Figure 10.9
              TECHNIQUE
                     Refracting Chlorophyll Photoelectric
               White prism      solution    tube
               light                             Galvanometer




                                             High transmittance
                  Slit moves to   Green      (low absorption):
                  pass light      light      Chlorophyll absorbs
                  of selected                very little green light.
                  wavelength.




                                             Low transmittance
                                  Blue       (high absorption):
                                  light      Chlorophyll absorbs
                                             most blue light.
• An absorption spectrum is a graph plotting a
       pigment’s light absorption versus 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




© 2011 Pearson Education, Inc.
Figure 10.10
               RESULTS




                                     Absorption of light by
                                     chloroplast pigments
                                                                Chloro-
                                                                phyll a      Chlorophyll b


                                                                                    Carotenoids


               (a) Absorption
                   spectra                                      400          500          600             700
                                                                             Wavelength of light (nm)




                                     (measured by O2 release)
                                     Rate of photosynthesis




               (b) Action spectrum                              400           500            600        700


                                                                          Aerobic bacteria
                                                                                Filament
                                                                                 of alga


               (c) Engelmann’s
                   experiment                                   400           500            600         700
• 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 O2
     • He used the growth of aerobic bacteria clustered
       along the alga as a measure of O2 production



© 2011 Pearson Education, Inc.
• 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




© 2011 Pearson Education, Inc.
Figure 10.11
                        CH3 in chlorophyll a
               CH3
                        CHO in chlorophyll b


                        Porphyrin ring




                     Hydrocarbon tail
                     (H atoms not shown)
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




© 2011 Pearson Education, Inc.
Figure 10.12




                                             Excited
                                   e−        state
 Energy of electron




                                                       Heat




                                               Photon
                                               (fluorescence)
                      Photon
                                             Ground
                               Chlorophyll   state
                                molecule


   (a) Excitation of isolated chlorophyll molecule              (b) Fluorescence
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



© 2011 Pearson Education, Inc.
Figure 10.13




                                       Photosystem
                                                              STROMA
                      Photon
                                 Light-           Reaction-   Primary
                                 harvesting       center      electron
                                 complexes        complex     acceptor
 Thylakoid membrane




                                                                                                   Chlorophyll    STROMA




                                                                         Thylakoid membrane
                                              e   −




                          Transfer    Special pair of     Pigment
                          of energy   chlorophyll a       molecules
                                      molecules
                                                                                              Protein
                                               THYLAKOID SPACE                                subunits           THYLAKOID
                                           (INTERIOR OF THYLAKOID)                                                 SPACE
    (a) How a photosystem harvests light                                    (b) Structure of photosystem II
• A primary electron acceptor in the reaction
       center accepts an excited electron and is
       reduced as a result
     • Solar-powered transfer of an electron from a
       chlorophyll a molecule to the primary electron
       acceptor is the first step of the light reactions




© 2011 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




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• Photosystem I (PS I) is best at absorbing a
       wavelength of 700 nm
     • The reaction-center chlorophyll a of PS I is
       called P700




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Linear Electron Flow
     • During the light reactions, there are two possible
       routes for electron flow: cyclic and linear
     • Linear electron flow, the primary pathway,
       involves both photosystems and produces ATP
       and NADPH using light energy




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• A photon hits a pigment and its energy is passed
       among pigment molecules until it excites P680
     • An excited electron from P680 is transferred to
       the primary electron acceptor (we now call it
       P680+)




© 2011 Pearson Education, Inc.
Figure 10.14-1




                   Primary
                   acceptor
                             2
                      e−




                           P680

  1 Light




                                  Pigment
                                  molecules

                 Photosystem II
                     (PS II)
• P680+ is a very strong oxidizing agent
     • H2O is split by enzymes, and the electrons are
       transferred from the hydrogen atoms to P680 +,
       thus reducing it to P680
     • O2 is released as a by-product of this reaction




© 2011 Pearson Education, Inc.
Figure 10.14-2




                      Primary
                      acceptor
                               2
            H2O         e−
   2 H+
     +
1
  /2 O2     3
                e−
                 e−
                             P680

  1 Light




                                    Pigment
                                    molecules

                 Photosystem II
                     (PS II)
• Each electron “falls” down an electron transport
       chain from the primary electron acceptor of PS II
       to PS I
     • 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




© 2011 Pearson Education, Inc.
Figure 10.14-3




                                         Ele
                      Primary               c   tron       4
                      acceptor                         tran
                                                           s   por
                                    Pq                             t ch
                               2                                        ai   n
            H2O         e−
   2 H+                                         Cytochrome
     +                                           complex
1
  /2 O2     3
                e−                                                       Pc
                 e−
                             P680                      5

  1 Light

                                                  ATP



                                    Pigment
                                    molecules

                 Photosystem II
                     (PS II)
• 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




© 2011 Pearson Education, Inc.
Figure 10.14-4




                                                                                   Primary
                                         Ele                                       acceptor
                      Primary               c   tron       4
                      acceptor                         tran
                                                           s   por
                                    Pq                             t ch               e−
                               2                                        ai   n
            H2O         e−
   2 H+                                         Cytochrome
     +                                           complex
1
  /2 O2     3
                e−                                                       Pc
                 e−                                                                    P700
                             P680                      5                                             Light

  1 Light                                                                                        6

                                                  ATP



                                    Pigment
                                    molecules
                                                                                 Photosystem I
                                                                                     (PS I)
                 Photosystem II
                     (PS II)
• Each electron “falls” down an electron transport
       chain 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
     • This process also removes an H+ from the
       stroma



© 2011 Pearson Education, Inc.
Figure 10.14-5




                                                                                                     E
                                                                                                   tra lect
                                                                                                  ch ns ron
                                                                                                    ai po
                                                                                   Primary            n rt
                                         Ele                                       acceptor
                      Primary               c   tron       4                                                 7
                      acceptor                         tran
                                                           s   por                               Fd
                                    Pq                             t ch               e−
                               2                                        ai   n                     e−            8
                        e−                                                                              e−               NADP+
            H2O                                 Cytochrome
   2H +
                                                                                                               NADP+      + H+
     +                                           complex
            3                                                                                                reductase
1
  /2 O2                                                                                                                  NADPH
                e−                                                       Pc
                 e−                                                                    P700
                             P680                      5                                                     Light

  1 Light                                                                                               6

                                                  ATP



                                    Pigment
                                    molecules
                                                                                 Photosystem I
                                                                                     (PS I)
                 Photosystem II
                     (PS II)
Figure 10.15



                                                               e−



                                    e−                                       e−
                                                   Mill
                                                  makes
                                                   ATP                       NADPH
                                          e   −
                                                          e−
                                                     e−




                                                                         n
                                                                    Photo
                        e−
                                                   ATP
                    n
               Photo




                             Photosystem II               Photosystem I
Cyclic Electron Flow
     • Cyclic electron flow uses only photosystem I
       and produces ATP, but not NADPH
     • No oxygen is released
     • Cyclic electron flow generates surplus ATP,
       satisfying the higher demand in the Calvin cycle




© 2011 Pearson Education, Inc.
Figure 10.16




                                       Primary
      Primary                          acceptor
                        Fd
      acceptor                                       Fd
                  Pq                                             NADP+
                                                       NADP+      + H+
                                                     reductase
                   Cytochrome                                    NADPH
                    complex

                                Pc



                                     Photosystem I
 Photosystem II        ATP
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; chloroplasts transform light energy
       into the chemical energy of ATP
     • Spatial organization of chemiosmosis differs
       between chloroplasts and mitochondria but also
       shows similarities

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• 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




© 2011 Pearson Education, Inc.
Figure 10.17
                     Mitochondrion                         Chloroplast




               MITOCHONDRION                                    CHLOROPLAST
                   STRUCTURE                                    STRUCTURE
                                             H+   Diffusion
                 Intermembrane                                  Thylakoid
                         space                                  space
                                  Electron
                       Inner                                    Thylakoid
                                 transport
                    membrane                                    membrane
                                   chain

                                       ATP
                                     synthase
                        Matrix                                  Stroma
                                      ADP + P i
                                                          ATP
        Key       Higher [H ]
                           +
                                                  H   +

                  Lower [H+ ]
• 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 H2O to NADPH




© 2011 Pearson Education, Inc.
Figure 10.18




  STROMA
  (low H+ concentration)                  Cytochrome                                    NADP+
                                           complex       Photosystem I                reductase
                 Photosystem II
                                                   Light
               Light               4 H+                                                        3
                                                                                                   NADP+ + H+
                                                                                Fd


                                          Pq
                                                                                              NADPH
                                                             Pc
                                               2
                   H2O
                           1   /2 O2
                               1

  THYLAKOID SPACE                +2 H+             4 H+
  (high H+ concentration)
                                                                                                          To
                                                                                                         Calvin
                                                                                                         Cycle

                               Thylakoid
                               membrane                     ATP
                                                          synthase
                                                                     ADP
                                                                      +         ATP
  STROMA
                                                                      Pi
  (low H+ concentration)                                                   H+
The Calvin cycle uses the chemical energy
of ATP and NADPH to reduce CO2 to sugar

     • The Calvin cycle, like the citric acid cycle,
       regenerates its starting material after molecules
       enter and leave the cycle
     • The cycle builds sugar from smaller molecules
       by using ATP and the reducing power of
       electrons carried by NADPH



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• Carbon enters the cycle as CO2 and leaves as
       a sugar named glyceraldehyde 3-phospate
       (G3P)
     • For net synthesis of 1 G3P, the cycle must take
       place three times, fixing 3 molecules of CO2
     • The Calvin cycle has three phases
                – Carbon fixation (catalyzed by rubisco)
                – Reduction
                – Regeneration of the CO2 acceptor (RuBP)


© 2011 Pearson Education, Inc.
Figure 10.19-3
                                                              Input
                                                              3  (Entering one
                                                              CO2 at a time)



                                                                                        Phase 1: Carbon fixation

                                                          Rubisco
                                                                      3 P                    P
                                                                       Short-lived
                                                                      intermediate
                                 3P                   P                                6         P
                                Ribulose bisphosphate                                3-Phosphoglycerate
                                       (RuBP)                                                                  6    ATP
                                                                                                            6 ADP


                        3 ADP                                   Calvin
                                                                Cycle
                 3                                                                       6 P           P
                     ATP
                                                                                     1,3-Bisphosphoglycerate
                                                                                                               6 NADPH
                      Phase 3:
                      Regeneration of                                                                      6 NADP+
                      the CO2 acceptor                                                                    6 Pi
                      (RuBP)                       P
                                         5
                                             G3P
                                                                               6           P
                                                                         Glyceraldehyde 3-phosphate        Phase 2:
                                                                                    (G3P)                  Reduction




                                                          1                P
                                                                  G3P                Glucose and
                                                               (a sugar)             other organic
                                                                Output               compounds
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 H2O but also limits photosynthesis
     • The closing of stomata reduces access to CO2
       and causes O2 to build up
     • These conditions favor an apparently wasteful
       process called photorespiration
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Photorespiration: An Evolutionary Relic?
     • In most plants (C3 plants), initial fixation of CO2,
       via rubisco, forms a three-carbon compound (3-
       phosphoglycerate)
     • In photorespiration, rubisco adds O2 instead of
       CO2 in the Calvin cycle, producing a two-carbon
       compound
     • Photorespiration consumes O2 and organic fuel
       and releases CO2 without producing ATP or
       sugar



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• Photorespiration may be an evolutionary relic because
        rubisco first evolved at a time when the atmosphere had
        far less O2 and more CO2
      • Photorespiration limits damaging products of light
        reactions that build up in the absence of the Calvin cycle
      • In many plants, photorespiration can drain as much as
        50% of the carbon fixed by the Calvin cycle on a hot, dry
        day
      • However, studies have shown that photorespiration is
        essential for plant growth as demonstrated by
        photorespiration mutants that are inviable in normal air
        and only grow in elevated CO2 conditions


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C4 Plants
     • C4 plants minimize the cost of photorespiration
       by incorporating CO2 into four-carbon
       compounds in mesophyll cells
     • This step requires the enzyme PEP
       carboxylase
     • PEP carboxylase has a higher affinity for CO2
       than rubisco does; it can fix CO2 even when CO2
       concentrations are low
     • These four-carbon compounds are exported to
       bundle-sheath cells, where they release CO2
       that is then used in the Calvin cycle
© 2011 Pearson Education, Inc.
Figure 10.20


                                    C4 leaf anatomy             The C4 pathway

                                                       Mesophyll
                   Mesophyll cell                      cell                                CO2
 Photosynthetic                                              PEP carboxylase
 cells of C4    Bundle-
 plant leaf     sheath
                cell
                                                        Oxaloacetate (4C)     PEP (3C)
       Vein                                                                         ADP
       (vascular tissue)
                                                          Malate (4C)                ATP


                                                                        Pyruvate (3C)
                                                      Bundle-
                                     Stoma            sheath         CO2
                                                      cell
                                                                            Calvin
                                                                            Cycle


                                                                        Sugar



                                                                   Vascular
                                                                   tissue
CAM Plants
     • Some plants, including succulents, use
       crassulacean acid metabolism (CAM) to fix
       carbon
     • CAM plants open their stomata at night,
       incorporating CO2 into organic acids
     • Stomata close during the day, and CO2 is
       released from organic acids and used in the
       Calvin cycle



© 2011 Pearson Education, Inc.
Figure 10.21




           Sugarcane                                   Pineapple
                                C4                                   CAM           CO2
                                         CO2
                                               1 CO2 incorporated
               Mesophyll Organic acid                              Organic acid          Night
               cell                              (carbon fixation)


                                  CO2                                       CO2
           Bundle-                             2 CO2 released                            Day
           sheath               Calvin                                   Calvin
                                Cycle            to the Calvin           Cycle
           cell
                                                 cycle

                                 Sugar                                     Sugar
                (a) Spatial separation of steps            (b) Temporal separation of steps
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 O2 in our atmosphere


© 2011 Pearson Education, Inc.

10 photosynthesis

  • 1.
    LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 10 Photosynthesis Lectures by Erin Barley Kathleen Fitzpatrick © 2011 Pearson Education, Inc.
  • 2.
    Overview: The ProcessThat Feeds the Biosphere • Photosynthesis is the process that converts solar energy into chemical energy • Directly or indirectly, photosynthesis nourishes almost the entire living world © 2011 Pearson Education, Inc.
  • 3.
    • Autotrophs sustainthemselves without eating anything derived from other organisms • Autotrophs are the producers of the biosphere, producing organic molecules from CO2 and other inorganic molecules • Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules © 2011 Pearson Education, Inc.
  • 4.
  • 5.
    • Photosynthesis occursin plants, algae, certain other protists, and some prokaryotes • These organisms feed not only themselves but also most of the living world © 2011 Pearson Education, Inc.
  • 6.
    Figure 10.2 (b) Multicellular (a) Plants alga (d) Cyanobacteria 40 µm (c) Unicellular 10 µm protists (e) Purple sulfur 1 µm bacteria
  • 7.
    • Heterotrophs obtaintheir organic material from other organisms • Heterotrophs are the consumers of the biosphere • Almost all heterotrophs, including humans, depend on photoautotrophs for food and O2 © 2011 Pearson Education, Inc.
  • 8.
    Photosynthesis converts lightenergy to the chemical energy of food • Chloroplasts are structurally similar to and likely evolved from photosynthetic bacteria • The structural organization of these cells allows for the chemical reactions of photosynthesis © 2011 Pearson Education, Inc.
  • 9.
    Chloroplasts: The Sitesof 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 © 2011 Pearson Education, Inc.
  • 10.
    • CO2 entersand O2 exits the leaf through microscopic pores called stomata • The chlorophyll is in the membranes of thylakoids (connected sacs in the chloroplast); thylakoids may be stacked in columns called grana • Chloroplasts also contain stroma, a dense interior fluid © 2011 Pearson Education, Inc.
  • 11.
    Figure 10.4 Leaf cross section Chloroplasts Vein Mesophyll Stomata CO2 O2 Chloroplast Mesophyll cell Outer membrane Thylakoid Intermembrane Stroma Granum Thylakoid space 20 µm space Inner membrane 1 µm
  • 12.
    Tracking Atoms ThroughPhotosynthesis: Scientific Inquiry • Photosynthesis is a complex series of reactions that can be summarized as the following equation: 6 CO2 + 12 H2O + Light energy → C6H12O6 + 6 O2 + 6 H2O © 2011 Pearson Education, Inc.
  • 13.
    The Splitting ofWater • Chloroplasts split H2O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by- product © 2011 Pearson Education, Inc.
  • 14.
    Figure 10.5 Reactants: 6 CO2 12 H2O Products: C6H12O6 6 H2O 6 O2
  • 15.
    Photosynthesis as aRedox Process • Photosynthesis reverses the direction of electron flow compared to respiration • Photosynthesis is a redox process in which H2O is oxidized and CO2 is reduced • Photosynthesis is an endergonic process; the enery boost is provided by light © 2011 Pearson Education, Inc.
  • 16.
    Figure 10.UN01 becomes reduced Energy + 6 CO2 + 6 H2O C6 H12 O6 + 6 O2 becomes oxidized
  • 17.
    The Two Stagesof 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 H2O – Release O2 – Reduce NADP+ to NADPH – Generate ATP from ADP by photophosphorylation © 2011 Pearson Education, Inc.
  • 18.
    • The Calvincycle (in the stroma) forms sugar from CO2, using ATP and NADPH • The Calvin cycle begins with carbon fixation, incorporating CO2 into organic molecules © 2011 Pearson Education, Inc.
  • 19.
    Figure 10.6-4 H2O CO2 Light NADP+ ADP +Pi Calvin Light Cycle Reactions ATP NADPH Chloroplast O2 [CH2O] (sugar)
  • 20.
    The light reactionsconvert 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 © 2011 Pearson Education, Inc.
  • 21.
    The Nature ofSunlight • 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 © 2011 Pearson Education, Inc.
  • 22.
    • The electromagneticspectrum 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 particles, called photons © 2011 Pearson Education, Inc.
  • 23.
    Figure 10.7 1m 10−5 nm 10−3 nm 1 nm 10 nm 3 106 nm (109 nm) 103 m Gamma Micro- Radio X-rays UV Infrared waves waves rays Visible light 380 450 500 550 600 650 700 750 nm Shorter wavelength Longer wavelength Higher energy Lower energy
  • 24.
    Photosynthetic Pigments: TheLight 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 © 2011 Pearson Education, Inc.
  • 25.
    Figure 10.8 Light Reflected light Chloroplast Absorbed Granum light Transmitted light
  • 26.
    • A spectrophotometermeasures a pigment’s ability to absorb various wavelengths • This machine sends light through pigments and measures the fraction of light transmitted at each wavelength © 2011 Pearson Education, Inc.
  • 27.
    Figure 10.9 TECHNIQUE Refracting Chlorophyll Photoelectric White prism solution tube light Galvanometer High transmittance Slit moves to Green (low absorption): pass light light Chlorophyll absorbs of selected very little green light. wavelength. Low transmittance Blue (high absorption): light Chlorophyll absorbs most blue light.
  • 28.
    • An absorptionspectrum is a graph plotting a pigment’s light absorption versus 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 © 2011 Pearson Education, Inc.
  • 29.
    Figure 10.10 RESULTS Absorption of light by chloroplast pigments Chloro- phyll a Chlorophyll b Carotenoids (a) Absorption spectra 400 500 600 700 Wavelength of light (nm) (measured by O2 release) Rate of photosynthesis (b) Action spectrum 400 500 600 700 Aerobic bacteria Filament of alga (c) Engelmann’s experiment 400 500 600 700
  • 30.
    • The actionspectrum 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 O2 • He used the growth of aerobic bacteria clustered along the alga as a measure of O2 production © 2011 Pearson Education, Inc.
  • 31.
    • Chlorophyll ais 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 © 2011 Pearson Education, Inc.
  • 32.
    Figure 10.11 CH3 in chlorophyll a CH3 CHO in chlorophyll b Porphyrin ring Hydrocarbon tail (H atoms not shown)
  • 33.
    Excitation of Chlorophyllby 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 © 2011 Pearson Education, Inc.
  • 34.
    Figure 10.12 Excited e− state Energy of electron Heat Photon (fluorescence) Photon Ground Chlorophyll state molecule (a) Excitation of isolated chlorophyll molecule (b) Fluorescence
  • 35.
    A Photosystem: AReaction-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 © 2011 Pearson Education, Inc.
  • 36.
    Figure 10.13 Photosystem STROMA Photon Light- Reaction- Primary harvesting center electron complexes complex acceptor Thylakoid membrane Chlorophyll STROMA Thylakoid membrane e − Transfer Special pair of Pigment of energy chlorophyll a molecules molecules Protein THYLAKOID SPACE subunits THYLAKOID (INTERIOR OF THYLAKOID) SPACE (a) How a photosystem harvests light (b) Structure of photosystem II
  • 37.
    • A primaryelectron acceptor in the reaction center accepts an excited electron and is reduced as a result • Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions © 2011 Pearson Education, Inc.
  • 38.
    • There aretwo 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 © 2011 Pearson Education, Inc.
  • 39.
    • Photosystem I(PS I) is best at absorbing a wavelength of 700 nm • The reaction-center chlorophyll a of PS I is called P700 © 2011 Pearson Education, Inc.
  • 40.
    Linear Electron Flow • During the light reactions, there are two possible routes for electron flow: cyclic and linear • Linear electron flow, the primary pathway, involves both photosystems and produces ATP and NADPH using light energy © 2011 Pearson Education, Inc.
  • 41.
    • A photonhits a pigment and its energy is passed among pigment molecules until it excites P680 • An excited electron from P680 is transferred to the primary electron acceptor (we now call it P680+) © 2011 Pearson Education, Inc.
  • 42.
    Figure 10.14-1 Primary acceptor 2 e− P680 1 Light Pigment molecules Photosystem II (PS II)
  • 43.
    • P680+ isa very strong oxidizing agent • H2O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680 +, thus reducing it to P680 • O2 is released as a by-product of this reaction © 2011 Pearson Education, Inc.
  • 44.
    Figure 10.14-2 Primary acceptor 2 H2O e− 2 H+ + 1 /2 O2 3 e− e− P680 1 Light Pigment molecules Photosystem II (PS II)
  • 45.
    • Each electron“falls” down an electron transport chain from the primary electron acceptor of PS II to PS I • 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 © 2011 Pearson Education, Inc.
  • 46.
    Figure 10.14-3 Ele Primary c tron 4 acceptor tran s por Pq t ch 2 ai n H2O e− 2 H+ Cytochrome + complex 1 /2 O2 3 e− Pc e− P680 5 1 Light ATP Pigment molecules Photosystem II (PS II)
  • 47.
    • In PSI (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 © 2011 Pearson Education, Inc.
  • 48.
    Figure 10.14-4 Primary Ele acceptor Primary c tron 4 acceptor tran s por Pq t ch e− 2 ai n H2O e− 2 H+ Cytochrome + complex 1 /2 O2 3 e− Pc e− P700 P680 5 Light 1 Light 6 ATP Pigment molecules Photosystem I (PS I) Photosystem II (PS II)
  • 49.
    • Each electron“falls” down an electron transport chain 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 • This process also removes an H+ from the stroma © 2011 Pearson Education, Inc.
  • 50.
    Figure 10.14-5 E tra lect ch ns ron ai po Primary n rt Ele acceptor Primary c tron 4 7 acceptor tran s por Fd Pq t ch e− 2 ai n e− 8 e− e− NADP+ H2O Cytochrome 2H + NADP+ + H+ + complex 3 reductase 1 /2 O2 NADPH e− Pc e− P700 P680 5 Light 1 Light 6 ATP Pigment molecules Photosystem I (PS I) Photosystem II (PS II)
  • 51.
    Figure 10.15 e− e− e− Mill makes ATP NADPH e − e− e− n Photo e− ATP n Photo Photosystem II Photosystem I
  • 52.
    Cyclic Electron Flow • Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH • No oxygen is released • Cyclic electron flow generates surplus ATP, satisfying the higher demand in the Calvin cycle © 2011 Pearson Education, Inc.
  • 53.
    Figure 10.16 Primary Primary acceptor Fd acceptor Fd Pq NADP+ NADP+ + H+ reductase Cytochrome NADPH complex Pc Photosystem I Photosystem II ATP
  • 54.
    A Comparison ofChemiosmosis 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; chloroplasts transform light energy into the chemical energy of ATP • Spatial organization of chemiosmosis differs between chloroplasts and mitochondria but also shows similarities © 2011 Pearson Education, Inc.
  • 55.
    • 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 © 2011 Pearson Education, Inc.
  • 56.
    Figure 10.17 Mitochondrion Chloroplast MITOCHONDRION CHLOROPLAST STRUCTURE STRUCTURE H+ Diffusion Intermembrane Thylakoid space space Electron Inner Thylakoid transport membrane membrane chain ATP synthase Matrix Stroma ADP + P i ATP Key Higher [H ] + H + Lower [H+ ]
  • 57.
    • ATP andNADPH 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 H2O to NADPH © 2011 Pearson Education, Inc.
  • 58.
    Figure 10.18 STROMA (low H+ concentration) Cytochrome NADP+ complex Photosystem I reductase Photosystem II Light Light 4 H+ 3 NADP+ + H+ Fd Pq NADPH Pc 2 H2O 1 /2 O2 1 THYLAKOID SPACE +2 H+ 4 H+ (high H+ concentration) To Calvin Cycle Thylakoid membrane ATP synthase ADP + ATP STROMA Pi (low H+ concentration) H+
  • 59.
    The Calvin cycleuses the chemical energy of ATP and NADPH to reduce CO2 to sugar • The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle • The cycle builds sugar from smaller molecules by using ATP and the reducing power of electrons carried by NADPH © 2011 Pearson Education, Inc.
  • 60.
    • Carbon entersthe cycle as CO2 and leaves as a sugar named glyceraldehyde 3-phospate (G3P) • For net synthesis of 1 G3P, the cycle must take place three times, fixing 3 molecules of CO2 • The Calvin cycle has three phases – Carbon fixation (catalyzed by rubisco) – Reduction – Regeneration of the CO2 acceptor (RuBP) © 2011 Pearson Education, Inc.
  • 61.
    Figure 10.19-3 Input 3 (Entering one CO2 at a time) Phase 1: Carbon fixation Rubisco 3 P P Short-lived intermediate 3P P 6 P Ribulose bisphosphate 3-Phosphoglycerate (RuBP) 6 ATP 6 ADP 3 ADP Calvin Cycle 3 6 P P ATP 1,3-Bisphosphoglycerate 6 NADPH Phase 3: Regeneration of 6 NADP+ the CO2 acceptor 6 Pi (RuBP) P 5 G3P 6 P Glyceraldehyde 3-phosphate Phase 2: (G3P) Reduction 1 P G3P Glucose and (a sugar) other organic Output compounds
  • 62.
    Alternative mechanisms ofcarbon 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 H2O but also limits photosynthesis • The closing of stomata reduces access to CO2 and causes O2 to build up • These conditions favor an apparently wasteful process called photorespiration © 2011 Pearson Education, Inc.
  • 63.
    Photorespiration: An EvolutionaryRelic? • In most plants (C3 plants), initial fixation of CO2, via rubisco, forms a three-carbon compound (3- phosphoglycerate) • In photorespiration, rubisco adds O2 instead of CO2 in the Calvin cycle, producing a two-carbon compound • Photorespiration consumes O2 and organic fuel and releases CO2 without producing ATP or sugar © 2011 Pearson Education, Inc.
  • 64.
    • Photorespiration maybe an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O2 and more CO2 • Photorespiration limits damaging products of light reactions that build up in the absence of the Calvin cycle • In many plants, photorespiration can drain as much as 50% of the carbon fixed by the Calvin cycle on a hot, dry day • However, studies have shown that photorespiration is essential for plant growth as demonstrated by photorespiration mutants that are inviable in normal air and only grow in elevated CO2 conditions © 2011 Pearson Education, Inc.
  • 65.
    C4 Plants • C4 plants minimize the cost of photorespiration by incorporating CO2 into four-carbon compounds in mesophyll cells • This step requires the enzyme PEP carboxylase • PEP carboxylase has a higher affinity for CO2 than rubisco does; it can fix CO2 even when CO2 concentrations are low • These four-carbon compounds are exported to bundle-sheath cells, where they release CO2 that is then used in the Calvin cycle © 2011 Pearson Education, Inc.
  • 66.
    Figure 10.20 C4 leaf anatomy The C4 pathway Mesophyll Mesophyll cell cell CO2 Photosynthetic PEP carboxylase cells of C4 Bundle- plant leaf sheath cell Oxaloacetate (4C) PEP (3C) Vein ADP (vascular tissue) Malate (4C) ATP Pyruvate (3C) Bundle- Stoma sheath CO2 cell Calvin Cycle Sugar Vascular tissue
  • 67.
    CAM Plants • Some plants, including succulents, use crassulacean acid metabolism (CAM) to fix carbon • CAM plants open their stomata at night, incorporating CO2 into organic acids • Stomata close during the day, and CO2 is released from organic acids and used in the Calvin cycle © 2011 Pearson Education, Inc.
  • 68.
    Figure 10.21 Sugarcane Pineapple C4 CAM CO2 CO2 1 CO2 incorporated Mesophyll Organic acid Organic acid Night cell (carbon fixation) CO2 CO2 Bundle- 2 CO2 released Day sheath Calvin Calvin Cycle to the Calvin Cycle cell cycle Sugar Sugar (a) Spatial separation of steps (b) Temporal separation of steps
  • 69.
    The Importance ofPhotosynthesis: 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 O2 in our atmosphere © 2011 Pearson Education, Inc.

Editor's Notes

  • #5 Figure 10.1 How can sunlight, seen here as a spectrum of colors in a rainbow, power the synthesis of organic substances?
  • #7 Figure 10.2 Photoautotrophs.
  • #12 Figure 10.4 Zooming in on the location of photosynthesis in a plant.
  • #15 Figure 10.5 Tracking atoms through photosynthesis.
  • #17 Figure 10.UN01 In-text figure, p. 188
  • #20 Figure 10.6 An overview of photosynthesis: cooperation of the light reactions and the Calvin cycle.
  • #24 Figure 10.7 The electromagnetic spectrum.
  • #26 Figure 10.8 Why leaves are green: interaction of light with chloroplasts.
  • #28 Figure 10.9 Research Method: Determining an Absorption Spectrum
  • #29 For the Cell Biology Video Space-Filling Model of Chlorophyll a, go to Animation and Video Files.
  • #30 Figure 10.10 Inquiry: Which wavelengths of light are most effective in driving photosynthesis?
  • #33 Figure 10.11 Structure of chlorophyll molecules in chloroplasts of plants.
  • #35 Figure 10.12 Excitation of isolated chlorophyll by light.
  • #37 Figure 10.13 The structure and function of a photosystem.
  • #43 Figure 10.14 How linear electron flow during the light reactions generates ATP and NADPH.
  • #45 Figure 10.14 How linear electron flow during the light reactions generates ATP and NADPH.
  • #47 Figure 10.14 How linear electron flow during the light reactions generates ATP and NADPH.
  • #49 Figure 10.14 How linear electron flow during the light reactions generates ATP and NADPH.
  • #51 Figure 10.14 How linear electron flow during the light reactions generates ATP and NADPH.
  • #52 Figure 10.15 A mechanical analogy for linear electron flow during the light reactions.
  • #54 Figure 10.16 Cyclic electron flow.
  • #57 Figure 10.17 Comparison of chemiosmosis in mitochondria and chloroplasts.
  • #59 Figure 10.18 The light reactions and chemiosmosis: the organization of the thylakoid membrane.
  • #62 Figure 10.19 The Calvin cycle.
  • #67 Figure 10.20 C 4 leaf anatomy and the C 4 pathway.
  • #69 Figure 10.21 C 4 and CAM photosynthesis compared.