CHAPTER 8:

Photosynthesis

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Photosynthesis
 Energy within light is captured and used to
synthesize carbohydrates

CO2 + H2O + light energy → C6H12O6 + O2 + H2O

 CO2 is reduced
 H2O is oxidized
 Energy from light drives this endergonic reaction

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Photosynthesis powers the biosphere
 Regions on the surface of the Earth and in
the atmosphere where living organisms
exist
 Life is largely driven by the photosynthetic
power of green plants
 Cycle where cells use organic molecules
for energy and plants replenish those
molecules using photosynthesis
 Plants also produce oxygen
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Trophic levels
 Heterotroph
 Must eat food, organic molecules from their
environment, to sustain life
 Autotroph
 Make organic molecules from inorganic
sources like CO2 and H2O
 Photoautotroph
 Use light as a source of energy
 Green plants, algae, cyanobacteria

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Green tissue is Photosynthetic
 Photosynthesis occurs in the green parts
of plants, leaves, stems etc.
 Inside cells, inside the chloroplasts, in the
thylakoid membranes
Chloroplast
 Organelles in plants and algae that carry
out photosynthesis
 Chlorophyll- green pigment
 Majority of photosynthesis occurs in
internal part of leaves called mesophyll
 Stomata- pores where carbon dioxide
enters and oxygen exits leaf
 H2O comes from roots

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Chloroplast anatomy
 Outer and inner membrane
 Between is intermembrane space
 3rd membrane - thylakoid membrane
contains chlorophyll
 Forms thylakoids (think of stacked disks/pies)
 Enclose thylakoid lumen (pie filling)
 Granum- stack of thylakoids
 Stroma- fluid filled region between
thylakoid membrane and inner membrane

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2 stages of photosynthesis
 Light reactions
 Uses light energy
 Take place in thylakoid membranes
 Produce ATP, NADPH and O2

 Calvin cycle
 Occurs in stroma
 Uses ATP and NADPH to incorporate CO2 into
organic molecules
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Light energy
 Type of electromagnetic radiation
 Energy derived from both electric and
magnetic fields
 Travels as waves
 Short to long wavelengths
 Also behaves as particles- photons
 Shorter wavelengths have more energy

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Light as a wave
 Travels in waves
 Similar to what you see if you drop a rock into
a pond
 The distance between the tops of the
waves = wavelength
 The wavelengths can vary enormously
1 nanometer - 1 kilometer
 Electromagnetic spectrum
 Visible light has wavelengths from 350nm-
750nm!
Light as a particle
 But light also behaves as a particle
called photons (A. Einstein)
 Photons act like particles, but aren’t
 No mass
 Each has a FIXED quantity of energy
 The amount of energy in a photon is
inversely proportional to its wavelength.
 Theshorter the wavelength, the more
energy
Electromagnetic spectrum

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Light receptors
 When light hits matter it can be
 Reflected
 Absorbed
 Transmitted (pass through)
 Pigments absorb light
Photosynthetic pigments
 Photosynthetic pigments absorb some light
energy and reflect others
 Leaves are green because they reflect green
wavelengths
 Absorption boosts electrons to higher
energy levels
 To orbitals of higher energy
 Excited state

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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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Photosynthetic pigments
 After an electron absorbs light energy, it is
in an excited state and usually unstable
 Electron drops back down to lower orbital
 Most pigments release energy as
 Heat
 Light
 Excited electrons in photosynthetic
pigments are transferred to another
molecule or “captured”
 Able to hold energy
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 Pigments in plants that absorb light energy
 Chlorophyll a
 Chlorophyll b
 Carotenoids (color)
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 Absorption vs. action spectrum
 Absorption spectrum
 Wavelengths that are absorbed by different pigments in the plant
 Action spectrum
 Rate of photosynthesis by whole plant at specific wavelengths

Corresponds to
highest light energy
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absorption
Photosystems
 Captured light energy can be transferred to
other molecules to ultimately produce
energy intermediates for cellular work
 Happens in photosystems
 Thylakoid membranes of chloroplast
contain…
 Photosystem I (PSI)
 Photosystem II (PSII)
 Both are complexes of proteins and pigments

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Photosystems
 Light excites pigment molecules (ex,
chlorophylls) in PSII and PSI
 PSII - excited electrons travel to PSI
 Water is oxidized- generates O2 and H+
 Releases energy in electron transport chain
 Energy used to make H + electrochemical gradient

(using pumps)
 PSI – primary role to make NADPH
 Addition of H+ to NADP contributes to H+
electrochemical gradient
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ATP synthesis
 ATP synthesis
 Chemiosmotic mechanism
 Driven by flow of H+ from thylakoid lumen into
stroma via ATP synthase
 H+ gradient generated by
 ↑H+ in thylakoid lumen by splitting of water
 ↑H+ by ETC pumping H+ into lumen
 ↓H + from formation of NADPH in stroma

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 Electrons boosted to
higher energy level Reduce H+
outside

Increases H+
inside 25
Summary of light reactions
1. O2 produced in thylakoid lumen by
oxidation of H2O by PSII
 2 electrons transferred to P680+
2. NADPH produced in the stroma from high-
energy electrons that start in PSII and
boosted in PSI
 NADP reductase
 NADP+ + 2 electrons + H + → NADPH
3. ATP produced in stroma by H+
electrochemical gradient
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Cyclic and noncyclic electron flow
 Noncyclic
 Electrons begin at PSII and eventually
transfer to NADPH
 Linear process produces ATP and NADPH in
equal amounts
 Cyclic
 PSI electrons excited, release energy and
eventually return to PSI
 Electron cycling releases energy to transport
H+ into lumen driving synthesis of ATP
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Further increases H+ gradient
to produce more ATP
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The cytochrome complexes of mitochondria and
chloroplasts contain evolutionarily related proteins
 Recurring theme is descent with modification
 Homologous genes are similar because they
are derived from a common ancestor
 Comparing the electron transport chains of
mitochondria and chloroplasts reveals
homologous genes
 Family of cytochrome b-type proteins plays
similar but specialized roles
Photosytem II (PSII)
 2 main components
 Light-harvesting complex or antenna complex
 Directly absorbs photons
 Energy transferred via resonance energy transfer
to P680 pigment
 Reaction center
 P680 →P680* (higher energy state)
 Relatively unstable

 Releases electron to primary electron acceptor

 Replaces lost electrons from water

 Oxidation of water yields oxygen gas

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To ETC

Oxidation of
H2O replaces
lost electron 32
Redox machine
 PSII removes high energy electrons from a
pigment molecule and transfers them to a
primary electron acceptor
 Recent research in biochemical composition
of protein complex and role of components
 3 dimensional structure determined in 2004
using x-ray crystallography

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Z scheme
 Zigzag shape of energy curve
 Based on electron energy changes
 Events involve increases and decreases in
the energy of an electron as it moves from
PSII through PSI to NADPH
 Electron of nonexcited pigment molecule
has lowest energy in PSII
 Highest energy level from being boosted by
PSI

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Calvin cycle
 Also called Calvin-Benson cycle
 ATP and NADPH used to make
carbohydrates
 CO2 incorporated into carbohydrates
 Precursors to all organic molecules
 Energy storage

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CO2 incorporation
 Requires massive input of energy
 For every 6 CO2 incorporated, 18 ATP and
12 NADPH used
 Product is glyceraldehyde-3-phosphate
(G3P)
 Starting material to make glucose and other
carbs
 Glucose is not directly made

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3 phases of Calvin cycle
1. Carbon fixation
 CO2 incorporated in ribulose bisphosphate (RuBP) – 5
C sugar using rubisco
 Results in a 6 carbon intermediate that splits into 2 3-
phosphoglycerate molecules (3PG)
2. Reduction and carbohydrate production
 ATP is used to convert 3PG into 1,3-
bisphosphoglycerate (1,3-BPG)
 NADPH electrons reduce it to G3P
 6 CO2 → 12 G3P
 2 for carbohydrates
 10 for regeneration of RuBP
3. Regeneration of RuBP
 10 G3P converted into 6 RuBP using 6 ATP 39
 Phase 1

reduction

Phase 3 Phase 2
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The Calvin cycle was determined by isotope labeling
methods

 14
C-labeled CO2 injected into cultures of green algae
 Allowed to incubate different lengths of time
 Separated newly made radiolabeled molecules using
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
Variations in photosynthesis
 Certain environmental conditions can
influence both the efficiency and way the
Calvin cycle works
 Light
intensity
 Temperature
 Water availability

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Photorespiration
 RuBP (5 C’s) + CO2 → 2 3PG

 Rubisco functions as a carboxylase

 C plants make 3PG (3PG has 3 C’s)
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 Rubisco can also be an oxygenase
 Adds O2 to RuBP in place of CO2
 Photorespiration
 More likely in hot and dry environments
 Deserts
 Favored when CO2 low and O2 high
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So What?
 The result is a 3 carbon and a 2 carbon
molecule instead of 2 3-Carbon molecules
 The 2 carbon molecule is exported from the
chloroplast
 To the peroxisome and/or mitochondria
where it is degraded down to CO2!
 Called Photorespiration because it occurs in
the light AND consumes Oxygen
 BUT NO ATP is produced!!!!
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C4 plants – minimize photorespiration
 C4 plants make oxaloacetate (4 carbon
compound) in the first step of carbon fixation
NOT 3PG
 Hatch-Slack pathway
 Leaves have 2-cell layer organization
 Mesophyll cells
 CO2 enters via stomata and 4 carbon compound
formed (PEP carboxylase does not promote
photorespiration)
 Bundle-sheath cells
 4 carbon compound transferred that releases steady
supply CO2
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Different carbon
fixation process

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 Which is better – C3 or C4?
 In warm dry climates C4 plants have
the advantage in conserving water
and preventing photorespiration
 In cooler climates, C3 plants use less
energy to fix CO2
90% of plants are C3

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Crassulacean Acid Metabolism
(CAM) plants
 Some C4 plants separate processes using time
 Cacti
 CAM plants open their stomata at night
 CO2 enters and is converted to oxaloacetate then to
malate
 Stomata close during the day to conserve water
 Malate broken down into CO2 to drive Calvin
cycle during the day

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