1/20/21

Photosynthesis 1 – light reactions

What are ’light’ and ‘dark’ reactions?

How is light energy captured for biochemistry?

What light is important?

Where does photosynthesis take place?

What is the ‘Z’ scheme?

Biology, Campbell&Reese 7th ed., Chap. 10

What is photosynthesis?

Photosynthesis: a process that converts atmospheric CO2 and H2O to

carbohydrates

Solar energy is captured in chemical form as ATP and NADPH

ATP and NADPH are used to convert CO2 to hexose phosphates

Phototrophs: photosynthetic organisms (some bacteria, algae, higher plants)

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Photosynthesis divides between

‘light’ and ‘dark’ reactions

Light and Dark reactions

Light (-dependent) reactions ...

Are membrane-bound

Release O2 from splitting 2H2O molecules, with

H+ from H2O used in the Chemiosmotic synthesis of ATP, and

Hydride ion (H:-) from H2O reduces NADP+ to NADPH

Dark (light-independent reactions ...

Occur in solution

Reduce gaseous CO2 to carbohydrate

Require energy of NADPH and ATP

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Chloroplast structure

Chloroplasts
4-8 mm diameter organelle
two surrounding membranes
internal membrane stacking

Chloroplast structure

Chloroplasts
4-8 mm diameter organelle
two surrounding membranes
internal membrane stacking

EM of chloroplast
Freeze-fracture EM of grana

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Chlorophyll and other pigments

capture light
Chlorophylls are most abundant and most important in light harvesting ...
contain tetrapyrrole ring (chlorin) similar to heme, but contains Mg2+
chlorophylls a (Chl a) and b (Chl b) in plants
bacteriochlorophylls a (BChl a) and b (BChl b) are similar and major
pigments in anaerobic photosynthetic bacteria

Chlorophyll and other pigments

capture light
‘Antenna’ pigments include carotenoids (carotene pigments), phycocyanin
and phycoerythrin

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Chlorophyll and other pigments

capture light

Antenna pigments extend the range of light capture,

especially in the green (where chlorophyll absorbs
poorly)

Antenna pigments and chlorophylls exchange

light energy (excited electrons) until
captured

Light absorption is organised between

two Photosystems

Photosystems I (PSI) and II (PSII):

contain many proteins and pigments embedded in the thylakoid membrane
PSI and PSII operate in series, connected by cytochrome bf complex
Electrons are conducted from H2O to NADP+

Z-scheme

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Light absorption is organised between

two Photosystems

PSI and PSII each contain a reaction centre (site of the photochemical reaction)
chlorophylls in each reaction center are paired to capture light
PSI paired for P700 (absorb light maximally at 700nm)
PSII paired for P680 (absorb light maximally at 680nm)

Z-scheme

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Physical layout of light reaction steps

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The Z-scheme

Z-scheme: path of electron flow and reduction potentials of the components in

photosynthesis

Absorption of light energy converts P680 and P700 (poor reducing agents) to
excited molecules (good reducing agents)

Light energy drives the electron flow uphill

NADP+ is ultimately reduced to NADPH

Light is captured by antenna complexes

For 2 H2O oxidized to O2, 2 NADP+ are reduced to 2 NADPH

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Chemiososmosis and energy conversion

Mitchell’s Chemiosmosis Theory – conversion of energy from electron

transport via formation of a transmembrane electrochemical gradient

Requires e- (H+) transfer across a ‘coupling’ membrane to generate DµH

DµH = DY + DpH (DµH also sometime called the ‘proton motive force’)

DµH used to drive protons through the ATP synthase (H+-ATPase) to make
ATP

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Photophosphorylation - H+-ATP synthase

Photophosphorylation: synthesis of ATP which is dependant upon light energy

Chloroplast ATP synthase consists of two major particles: CFo and CF1

CFo spans the membrane, forms a pore for H+

CF1 protrudes into the stroma and catalyzes ATP synthesis from ADP and Pi

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Photophosphorylation - H+-ATP synthase

Mitchell’s Chemiosmosis Theory – conversion of energy from electron
transport via formation of a transmembrane electrochemical gradient

Requires e- (H+) transfer across a ‘coupling’ membrane to generate DµH

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Photophosphorylation - H+-ATP synthase

Mitchell’s Chemiosmosis Theory – conversion of energy from electron
transport via formation of a transmembrane electrochemical gradient

Requires e- (H+) transfer across a ‘coupling’ membrane to generate DµH

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Cyclic electron flow and phosphorylation

For 4e- transferred to 2 NADPH, 2 ATP are produced from the proton gradient
However, for each CO2 reduced to (CH2O) in carbohydrate synthesis, 2 NADPH and
3 ATP are required
Cyclic electron transport yields ATP but not NADPH, thus balancing the need for 3
ATP for every 2 NADPH

Ferridoxin donates e- not to NADP+, but back to the PQ pool via a specialized
cytochrome
Cyclic flow increases the protonmotive force and increases ATP production, but
no NADP+ is produced

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Summary

Photosynthesis uses light energy to fix carbon into sugar

Photosynthesis takes place in the chloroplast

Light reactions capture and use light energy to reduce NADP
and phosphorylate ADP
Light drives photosynthetic electron transport to generate µH

The µH is used by H+-ATP synthase to phosphorylate ADP

ATP and NADPH energy is used for cellular reactions

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