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Bios 305 Topic 8.

Photosynthesis page 1
Prof. J. Stafstrom

8. C3 and C4 photosynthesis
________________________________________________________________________
Reading: Ch. 7
Review: overview of photosynthesis; light reactions: Calvin cycle (115-128)
Photorespiration: C3, C4 and CAM pathways (127-138)
Grass leaves: 566-567
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Outline
1. Over view of the light reactions
2. Review of the Calvin cycle (C3 pathway)
3. Photorespiration
4. C4 photosynthesis and Kranz anatomy
5. CAM

1. Overview
31-1. Photosynthesis is important for all life. Plants and other autotrophs produce
reduced carbon (sugars), which is the source of virtually all energy used by all
organisms. (Note: Chapters 31 and 32 are available on the Raven website)
7-4. You do not need to know too much about the physics of light (photons, wavelength,
pigments).
7-11. You do not need to know details about electron transport and how ATP and
NADPH are formed during the light reactions.
7-12. You do not need to understand how ATP and NADPH are made by chemiosmosis.
7-18. You DO need to know about RuBISCO and the Calvin cycle.
7-1, 7-7. Review of chloroplast structure: 2 envelope membranes; thylakoid membranes
(light reactions, electron transport); stroma (soluble enzymes, DNA, ribosomes, etc.)

7-9. Photosynthesis occurs in chloroplasts.


1) Light reactions: pigments, light-harvesting proteins, electron transport chain, etc.
occurs in association with thylakoid membranes; products are ATP (energy) and
NADPH (high energy electrons or reducing power), which are used to drive carbon
fixation in the Calvin cycle; electrons needed to reduce NADP+ to NADPH come
from “splitting” of water; O2 from water is a waste product of photosynthesis.

2) Calvin cycle [7-18]: soluble enzymes are in the stroma of the chloroplast; use
energy from the light reactions (ATP and NADPH) to "fix" gaseous CO2 into sugars;
carbon becomes reduced.
7-15. Stomata open and close to allow gas exchange (CO2 in; H2O and O2 out).
7-3. Oxygen (O2) is a waste product of photosynthesis.
Bios 305 Topic 8. Photosynthesis page 2
Prof. J. Stafstrom

2. Review of the Calvin cycle (C3 pathway)


7-18. RuBISCO and the Calvin cycle.
CO2 + ATP + NADPH Æ sugars, reduced carbon compounds

- "dark" or "light-independent" reactions


- named for Melvin Calvin, who worked out many of the reactions (1950s); Nobel
prize
- need ATP (energy) and NADPH (reducing power) for carbon fixation (gas to
soluble sugar)

Calvin cycle (C3) reactions


Rxn 1 (most important):
3 RuBP [total 15C] + 3 CO2 [total 3C] Æ [3 6C] Æ 6 PGA [3C each, 18C total]
7-16. RuBISCO catalyzes fixes CO2 fixation. This reaction is catalyzed by
RuBISCO (ribulose bisphosphate carboxylase/oxygenase)
- PGA, the first detectable product of the Calvin cycle, contains 3 carbons (the 6C
cpd. Plants in which O2 is initially fixed by the Calvin cycle are called C3 plants.

Rxn. 2: 6 PGA + 6 ATP Æ 6 bPG [18C]


Rxn. 3: 6 bPG + 6 NADPH Æ 6 GP [18C]
Rxn. 4a: 5 GP (15C) + 3 ATP Æ Æ Æ 3 RuBP [15C]
Rxn. 4b: 1 GP = net yield of Calvin cycle

NOTE: The complete Calvin cycle is considerably more complicated, especially with
regard to reactions used to regenerate RuBP (indicated as "4a").

3. Photorespitation
7-19. RuBISCO has two activities: 1) carboxylase (Good); 2 oxygenase (Bad)
1) carobxylase reaction (useful): RuBP [5C] + CO2 [1C] Æ 2 PGA
- as described above, this reaction stores energy in reduced carbon cpds.; anabolic;
carbon is fixed
2) oxygenase reaction (wastes energy): RuBP [5C] + O2 Æ 1 PGA [3C] + 1
phosphoglycolate [2C]
- only 1 PGA is made instead of 2
- the carbon in phosphoglycolate must be broken down or salvaged through a
complex set of reactions referred to collectively as “photorespitation”

7-20. Photorespiration salvage pathway:


- Involves enzymes in 3 organelles (chloroplasts, peroxisomes and mitochondria).
- Consumes O2 and releases CO2 (hence “respiration”)
- Consumes a lot of energy (1 ATP, 2 NADH)
Bios 305 Topic 8. Photosynthesis page 3
Prof. J. Stafstrom

The "Problem": Air contains ca. 21% O2 and 0.036% CO2 (a ratio of about 600:1).
RuBISCO works fine under these conditions. Some physiological or environmental
conditions tend to skew this ratio even higher, which will promote the oxygenase reaction
or photorespiration. When it is hot and dry, plants keep their stomata closed to prevent
water loss. At the same time, O2 (a waste product of the light reactions) builds up in the
leaf, and more CO2 cannot be acquired from the atmosphere.

The "Solution": Plants can avoid photorespiration by separating the initial fixation of CO2
from the final fixation of CO2 in the Calvin cycle, thus protecting RuBISCO from high
levels of O2.

4. C4 photosynthesis and Kranz anatomy: separation in SPACE


7-22. The C4 pathway
1) Initial CO2 fixation occurs in outer mesophyll cells. CO2 + PEP yield
oxaloacetate (OAA). OAA is converted to malate.
2) Malate is transported to inner bundle sheath cells. CO2 is released and used by the
Calvin cycle. Pyruvate is returned to mesophyll cells.
3) Conversion of pyruvate to PEP uses energy (ATP).
4) Initial and final CO2 fixation are separated in space.

7-24. Chloroplasts of mesophyll cells (M cells) and bundle sheath cells (BS cells) are
distinctive. Transport is through plasmodesmata. BS cells are surrounded by thick,
impermeable walls, which limits access of O2 to RuBISCO. The C4 shuttle leads to
relatively high concentrations of CO2 in BS cells. These processes allow the Calvin
cycle to function efficiently within BS cells.

7-23. C4 plants such as corn have Kranz anatomy. Comparison of the leaves of two
grasses, sugarcane (C4 plant; 25-25) and wheat (C3 plant; 25-25). (C3 leaves also
contain BS and M cells; however, their structure and arrangement is different, and
each carries out different biochemical reactions).

C4 photosynthesis occurs in many monocot and dicot families.

C4 pathway requires expenditure of some additional energy (ATP). If temperatures are mild
and water is abundant, C3 photosynthesis is more efficient. Under hot and dry conditions,
however, C4 plants are more efficient. The ability of C4 plants to avoid photorespiration
more than offsets the additional energetic costs of this pathway.
Bios 305 Topic 8. Photosynthesis page 4
Prof. J. Stafstrom

5. CAM pathway: separation in TIME


7-25b. CAM = crassulacean acid metabolism: Initial fixation of CO2 (into a C4
compound) occurs at night. Initial and final fixation are separated in time.

7-26. Details of CAM C4 pathway.


Initial fixation of CO2 (into OAA and then malate) occurs when stomata are open at
night. Malate is stored in large vaculoes within mesophyll cells.
Stomata are closed during the day; malate is transferred from the vacuole to the
cytoplasm and then to chloroplasts; Calvin cycle/C3 photosynthesis then occurs; light is
used to drive the light reactions, which produce ATP and NADPH needed for the Calvin
cycle

In addition to Crassulaceae, CAM occurs in many monocot and dicot families (and some
gymnosperms and ferns). Evolved independently many times.

CAM photosynthesis conserves water better than even C4 photosynthesis, but plants fix
little carbon and grow slowly. Most efficient in arid regions; cacti are CAM plants.

7-25. Summary: Initial and final fixation of CO2 are separated spatially in C4 plants and
temporally in CAM plants. Remember: both C4 and CAM pathways also include the
Calvin cycle or C3 pathway; the distinction is where CO2 is fixed initially.

Relative efficiencies of different types of photosynthesis with regard to water loss


(see text).

Essay: The carbon cycle, greenhouse gases, and global warming (pp 136-137).
We won’t discuss this is class, but you ought to read and digest this information.

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