Chapter 7 Photosynthesis

7.1 Photosynthetic Organisms Photosynthesis converts solar energy into chemical energy

Autotrophs produce their own organic molecules from inorganic molecules; also called producers photoautotrophs: use sunlight as source of energy chemoautotrophs: use inorganic chemicals as source of energy this chapter is about the former

7.1 Photosynthetic Organisms Photosynthesis converts solar energy into chemical energy
Heterotrophs must consume organic molecules; also called consumers both autotrophs and heterotrophs use organic molecules produced by photosynthesis for growth, repair, and energy

Fig. 7.1 Photosynthetic organisms

7.1 Photosynthetic Organisms

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Flowering plant anatomy

raw materials for photosynthesis: water and carbon dioxide water is absorbed by roots and transported to leaves by stem carbon dioxide enters leaf through openings called stomata water and carbon dioxide diffuse into chloroplasts, organelles specialized for photosynthesis

7.1 Photosynthetic Organisms

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Flowering plant anatomy, cont.

chloroplasts stroma: fluid-filled interior thylakoids: flattened, membranous sacs

inside called thylakoid space grouped into stacks called grana

chlorophyll: pigment embedded in thylakoid membrane

chlorophyll a chlorophyll b

Fig. 7.2 Leaves and photosynthesis

cuticle

Leaf cross section

upper epidermis mesophyll lower epidermis

CO2 leaf vein inner membrane outer membrane stroma granum

O2 stoma

stroma

Chloroplast
thylakoid space thylakoid membrane

37,000x

Grana

independent thylakoid overlapping thylakoid in a granum in a granum

Chlorophyll in thylakoid membrane

7.2 Solar Energy Conversion Photosynthesis takes place in two sets of reactions q Photosynthetic Pigments
pigments are molecules that absorb light; they reflect or transmit light they dont absorb absorption spectrum: range of light wavelengths that a molecule absorbs measured by spectrophotometer

Absorption, reflection, transmission

One way to measure absorption

Fig. 7.3a Absorption spectra

7.2 Solar Energy Conversion Photosynthesis takes place in two sets of reactions q Photosynthetic Pigments, cont.
not all absorbed light is necessarily used for photosynthesis the range of light wavelengths that is used in photosynthesis is called an action spectrum can be measured by the rate of oxygen production

Fig. 7.3b Action spectrum

Engelmanns experiment

7.2 Solar Energy Conversion

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Photosynthetic Reaction

Niel (1930) discovered that O2 given off comes from water, not CO2 discovered by tagging CO and 2 H2O with 18O.

7.2 Solar Energy Conversion

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Photosynthetic Reaction, cont.

Usual equation:

Net equation:

7.2 Solar Energy Conversion

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Photosynthetic Reaction, cont.

Two Sets of Reactions Blackman (1905) suggested two sets of reactions involved in photosynthesis
light reactions Calvin cycle reactions

7.2 Solar Energy Conversion

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Photosynthetic Reaction, cont.

Two Sets of Reactions, cont. Light reactions: solar energy chemical energy (ATP, NADPH)
chlorophyll absorbs sunlight and energizes electrons H O is split, H+ ions used for ATP 2 production ADP + P ATP NADP+ NADPH

7.2 Solar Energy Conversion

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Photosynthetic Reaction, cont.

Two Sets of Reactions, cont. Calvin cycle: chemical energy (ATP, NADPH) chemical energy (carbohydrate)
CO2 absorbed and reduced to a carbohydrate ATP and NADPH drive the reactions

Fig. 7.4 Overview of Photosynthesis

Fate of reactant atoms

7.3 Light Reactions The light reactions store energy in ATP and NADPH q Noncyclic Electron Pathway
Point: produce ATP and NADPH uses photosystems: pigment complexes with electron acceptors; act like solar energy antennas pigments include chlorophyll a, chlorophyll b, and carotenoids photosystem I and photosystem II

Schematic of a photosystem

Chlorophyll excitation

A mechanical analogy

Fig. 7.5 Noncyclic electron pathway

H2O solar energy
ADP+P NADP+

CO2

sun

Light reactions NADPH ATP thylakoid membrane

Calvin cycle

sun

energy level

electron acceptor

O2

CH2O

electron acceptor

ele ctr on

e
tra nsp ort ch

e NADP+

e
ain

ATP e reaction center

H+ NADPH reaction center pigment complex

pigment complex e

Photosystem I Photosystem II
CO2 CH2O Calvin cycle reactions H2O
1 2 O2

2H+

7.3 Light Reactions

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Noncyclic Electron Pathway, cont.

Steps: 1. photosystem II (PS II) absorbs solar energy (electrons are excited) 2. electrons are passed to the reaction center (a particular pair of chlorophyll a molecules) 3. electrons move to electron acceptor molecules

7.3 Light Reactions

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Noncyclic Electron Pathway, cont.

Steps: 4. electrons are replaced by removing one from water, leaving 2H+ and O2 (water is oxidized); H+ accumulates in thylakoid space 5. meanwhile, electrons move down an electron transport chain

7.3 Light Reactions

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Noncyclic Electron Pathway, cont.

Steps: 6. electron transport chain pumps more H+ ions into the thylakoid space, building a gradient 7. PS I absorbs solar energy and transfers electrons to electron acceptors 8. PS I gets replacement electrons from adjacent transport chain

7.3 Light Reactions

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Noncyclic Electron Pathway, cont.

Steps: 9. PS I electron acceptors pass electrons to NADP+; NADP+ accepts two electrons and H+, making NADPH 10. H+ ions flow down electrochemical gradient (through ATP synthase) to form ATP; also known as photophosphorylation

ATP production due to gradient

Fig. 7.5 Noncyclic electron pathway

H2O solar energy
ADP+P NADP+

CO2

sun

Light reactions NADPH ATP thylakoid membrane

Calvin cycle

sun

energy level

electron acceptor

O2

CH2O

electron acceptor

ele ctr on

e
tra nsp ort ch

e NADP+

e
ain

ATP e reaction center

H+ NADPH reaction center pigment complex

pigment complex e

Photosystem I Photosystem II
CO2 CH2O Calvin cycle reactions H2O
1 2 O2

2H+

7.3 Light Reactions

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Cyclic Electron Pathway

Point: Calvin cycle needs more ATP than NADPH, so cyclic pathway provides the extra ATP; ATP also used in other reactions (NADPH not so much) does not involve PS II

7.3 Light Reactions

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Cyclic Electron Pathway, cont.

Steps: 1. PS I absorbs solar energy and passes electrons to reaction center 2. energized electrons are sent down electron transport chain (instead of going to NADP+) 3. electron transport chain builds H+ gradient

7.3 Light Reactions

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Cyclic Electron Pathway, cont.

Steps: 4. H+ ions flow through ATP synthase to make ATP (also called chemiosmosis because it involves a gradient) 5. electrons return to PS 1, hence the term cyclic

Fig. 7.6 Cyclic electron pathway

H2 O solar energy ADP+P NADP+ Light reactions NADPH ATP CO2 Calvin cycle

thylakoid membrane

O2

CH2O

sun

energy level

electron acceptor e

el ec tro n

tra ns po rt ch ai n

ATP

reaction center

CO2

CH2O

Photosystem I

pigment complex

Calvin cycle reactions and other enzymatic reactions

Fig. 7.7 Organization of a thylakoid

H2O solar energy ADP+P NADP+ Light reactions NADPH ATP thylakoid membrane CO2 Calvin cycle

stroma O2 CH2O

thylakoid membrane granum thylakoid space

thylakoid

photosystem II electron transport H+ chain stroma photosystem I NADP H+ reductase Pq e NADP+ e e e H+ H2O 2 H
+

NADPH

e + O2 H+

1 2

H+ Thylakoid space

H+

ATP synthase ATP

H+ chemiosmosis P + ADP

Stroma

7.4 Calvin Cycle Reactions The Calvin cycle uses CO2 and energy from ATP and NADPH to make carbohydrates

Three major steps: 1. carbon dioxide fixation 2. carbon dioxide reduction 3. regeneration of RuBP

RuBP (ribulose-1,5-biphosphate) is a C5 (5-carbon) molecule

Fig. 7.8 Calvin cycle reactions

H2 O solar energy P ADP+ NADP+ CO2 Light reactions NADPH ATP Calvin cycle

Metabolites of the Calvin Cycle

stroma O2 CH2O

3 CO2 intermediate 3 C6

RuBPribulose-1,5-bisphosphate 3PG 3-phosphoglycerate BPG 1,3-bisphosphoglycerate G3P glyceraldehyde-3-phosphate

3 RuBP C5

CO2 fixation

6 3PG C3

6 ATP These ATP and NADPH molecules were produced by the light reactions

3 ADP + 3 P

Calvin cycle regeneration of RuBP

CO2 reduction

6ADP + 6 P

These ATP molecules were produced by the light reactions.

6 BPG C3 6 NADPH 6 G3P C3 6 NADP+

3 ATP

5 G3P C3

net gain of one G3P x 2 Other organic molecules Glucose

7.4 Calvin Cycle Reactions

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Fixation of Carbon Dioxide

1. CO2 attaches to RuBP, forming a C6 molecule that immediately splits into two C3 molecules called 3PG (3phosphoglycerate) RuBP carboxylase is the enzyme that speeds the reaction

20-50% of the protein content of chloroplasts

7.4 Calvin Cycle Reactions

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Reduction of Carbon Dioxide

2. 3PG is reduced to G3P (glyceraldehyde-3-phosphate) in two steps ATP is oxidized to ADP and P in first step NADPH is oxidized to NADP+ in second step CO has been reduced because 2 RCO2 has become RCH2O

Reduction of carbon dioxide

7.4 Calvin Cycle Reactions

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Regeneration of RuBP

3. G3P is used to regenerate RuBP using energy from ATP 5 G3P 3 RuBP

because 5 C3 = 3 C5 15 carbons = 15 carbons

therefore, 3 turns of the calvin cycle allow one G3P to exit (3 carbons in, 3 carbons out)

Regeneration of RuBP from G3P

Fig. 7.8 Calvin cycle reactions

H2 O solar energy P ADP+ NADP+ CO2 Light reactions NADPH ATP Calvin cycle

Metabolites of the Calvin Cycle

stroma O2 CH2O

3 CO2 intermediate 3 C6

RuBPribulose-1,5-bisphosphate 3PG 3-phosphoglycerate BPG 1,3-bisphosphoglycerate G3P glyceraldehyde-3-phosphate

3 RuBP C5

CO2 fixation

6 3PG C3

6 ATP These ATP and NADPH molecules were produced by the light reactions

3 ADP + 3 P

Calvin cycle regeneration of RuBP

CO2 reduction

6ADP + 6 P

These ATP molecules were produced by the light reactions.

6 BPG C3 6 NADPH 6 G3P C3 6 NADP+

3 ATP

5 G3P C3

net gain of one G3P x 2 Other organic molecules Glucose

7.4 Calvin Cycle Reactions

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Importance of the Calvin Cycle

Glyceraldehyde-3-phosphate can be used to make glucose, other sugars, starch, cellulose, amino acids, fatty acids, and glycerol G3P is also commonly known as PGAL (phosphoglyceraldehyde)

Fig. 7.9 Fate of G3P

7.5 Other Types of Photosynthesis C3, C4, and CAM photosynthesis are three types of photosynthesis

plants that do photosynthesis as described are called C3 plants when they close stomata to conserve water, [CO2] in leaf decreases and [O2] increases RuBP combines with O ; CO 2 2 eventually released; called photorespiration (wasteful)

Fig. 7.10a C3 carbon fixation

7.5 Other Types of Photosynthesis

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C4 Photosynthesis

C4 photosynthesis avoids photorespiration by separating CO2 fixation and the Calvin cycle spatially

7.5 Other Types of Photosynthesis

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C4 Photosynthesis, cont.

Steps: 1. mesophyll cells fix CO 2 2. PEP carboxylase fixes CO to 2 PEP, forming a C4 molecule

PEPCase PEP + CO2 oxaloacetate

7.5 Other Types of Photosynthesis

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C4 Photosynthesis, cont.

Steps: 3. malate pumped into bundle sheath cells, where CO2 enters Calvin cycle
requires energy, but still more efficient than C 3 photosynthesis in hot, dry climates because no photorespiration PEP does not bind O 2

Fig. 7.10b C4 carbon fixation

7.5 Other Types of Photosynthesis

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CAM Photosynthesis

CAM photosynthesis avoids photorespiration by separating CO2 fixation and the Calvin cycle temporally CAM stands for crassulaceanacid metabolism after Crassulaceae family of plants also found in cacti and other plants in warm, dry climates

7.5 Other Types of Photosynthesis

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CAM Photosynthesis, cont.

Steps: 1. PEPCase is used to fix CO at 2 night (when stomata are open) 2. C molecules are stored in large 4 vacuoles in mesophyll cells 3. C molecules (malate) release 4 CO2 to the Calvin cycle when NADPH and ATP are available from light reactions (during day)

Fig. 7.11 CAM carbon fixation

Fig. 7.10

Fig. 7.11 C3, C4, CAM carbon fixation