PHOTOSYNTHESIS :

Regulation of Calvin Cycle

Photorespiration
Hatch and Slack Pathway
Crassulacean Acid Metabolism

Regulation of Calvin Cycle

 Regulation prevents the Calvin Cycle from being

active in the dark, when it might function in a futile
cycle with Glycolysis & Pentose Phosphate Pathway,
wasting ATP & NADPH.

Light activates, or dark inhibits, the Calvin Cycle

(previously called the dark reaction) in several
ways.
Light-activated e- transfer is linked to pumping of H+
into thylakoid disks. pH in the stroma increases to
about 8.
Alkaline pH activates stromal Calvin Cycle enzymes
RuBP Carboxylase, Fructose-1,6-Bisphosphatase &
Sedoheptulose Bisphosphatase.

H2O OH + H

Chloroplast

stroma
(alkaline)

(acid inside
thylakoid disks)

The light-activated H+ shift is countered by Mg++ release

from thylakoids to stroma. RuBP Carboxylase (in stroma)
requires Mg++ binding to carbamate at the active site.
Some plants synthesize a transition-state inhibitor,
carboxyarabinitol-1-phosphate (CA1P), in the dark.
RuBP Carboxylase Activase facilitates release of CA1P
from RuBP Carboxylase, when it is activated under
conditions of light by thioredoxin.

 Thioredoxin is a small protein with a disulfide that is

reduced in chloroplasts via light-activated electron transfer.
During illumination, the thioredoxin disulfide is reduced
to a dithiol by ferredoxin, a constituent of the
photosynthetic light reaction pathway, via an enzyme
Ferredoxin-Thioredoxin Reductase.
Reduced thioredoxin activates several Calvin Cycle
enzymes, including Fructose-1,6-bisphosphatase,
Sedoheptulose-1,7-bisphosphatase, and RuBP Carboxylase
Activase, by reducing disulfides in those enzymes to thiols.
PDB 1FAA

disulfide

thioredoxin

ferredoxinRed

ferredoxinOx

-S
|

-S

FerredoxinThioredoxin
Reductase

thioredoxin

Thioredoxin f

-SH
-SH

PHOTORESPIRATION

 Photorespiration occurs when the CO2 levels inside a leaf

become low. This happens on hot dry days. On hot dry days the
plant is forced to close its stomata to prevent excess water loss.

The plant continues fix CO2 when its stomata are closed, the
CO2 will get used up and the O2 ratio in the leaf will increase
relative to CO2 concentrations. When the CO2 levels inside the
leaf drop to around 50 ppm, RuBisCO starts to combine O2 with
RuBP instead of CO2.

 The net result of this is that instead of producing 2 x 3C PGA

molecules, only one molecule of PGA is produced and a toxic 2C
molecule called phosphoglycolate is produced

 The plant must get rid of the phosphoglycolate since it

is highly toxic. It converts the molecule to glycolic acid.
The glycolic acid is then transported to the peroxisome
and there converted to glycine.
Glycine is transported to mitochondria and converted
to serine
The serine is then used to make other
organic
molecules.
All these conversions cost the plant energy and results
in the net loss of CO2 from the plant
To prevent this process, two specialized biochemical
additions have been evolved in the plant world: C4 and
CAM metabolism.

Hatch and Slack (C4)Pathway

C4 Leaf Anatomy
The C4 plants vascular bundles
are surrounded by two rings of
cells
- the inner ring, called bundle
sheath cells, contain starchrich chloroplasts lacking grana
- mesophyll cells present as
the outer ring.
This peculiar anatomy is called
kranz anatomy.
kranz anatomy is to provide a
site in which CO2 can be
concentrated around RuBisCO,
thereby avoiding
photorespiration

 The C4 pathway is
designed to efficiently fix
CO2 at low concentrations
and plants that use this
pathway are known as C4
plants.
These plants fix CO2 into a
four carbon compound
(C4) called oxaloacetate.
This occurs in cells called
mesophyll cells.

STEPS IN C4 CYCLE
1. CO2 is fixed to a three-carbon
compound called
phosphoenolpyruvate to produce
the four-carbon compound
oxaloacetate. The enzyme
catalyzing this reaction, PEP
carboxylase, fixes CO2 very
efficiently so the C4 plants don't
need to have their stomata open as
much. The oxaloacetate is then
converted to another four-carbon
compound called malate in a step
requiring the reducing power of
NADPH

STEPS IN C4 CYCLE
2. The malate then exits the mesophyll
cells and enters the chloroplasts of
specialized cells called bundle sheath
cells. Here the four-carbon malate is
decarboxylated to produce CO2, a
three-carbon compound called
pyruvate, and NADPH. The CO2
combines with ribulose bisphosphate
and goes through the Calvin cycle.
3. The pyruvate re-enters the mesophyll
cells, reacts with ATP, and is
converted back to
phosphoenolpyruvate, the starting
compound of the C4 cycle.

Differences between Calvin (C3) and C4 cycles

C4

C3

Temp 15-250 C
Absence of malate
One carboxylation reaction
CO2 is the substrate
Usual leaf structures

Temp 30-350 C
Presence of malate
2 carboxylation reactions
HCO3 is the substrate
Closed stomata to reduce
water loss and
concentrating CO2 in the
bundle sheet cells
Additional ATP is required

Crassulacean Acid Metabolism

 CAM plants live in very dry condition and,

unlike other plants, open their stomata to fix
CO2 only at night.
Like C4 plants, the use PEP carboxylase to fix
CO2, forming oxaloacetate.
The oxaloacetate is converted to malate which
is stored in cell vacuoles. During the day when
the stomata are closed, CO2 is removed from
the stored malate and enters the Calvin cycle

Comparison
C3 Plants

C4 Plants

CAM Plants

Stomata

Open at Night
Closed during Day

Partially closed
during day and
partially closed at
night

Closed during day

and Open at night

Carbon fixation

Carbon dioxide is
fixed into a three
carbon compound
that is stable

Carbon dioxide is
Carbon dioxide is
temporarily stored temporarily stored
as a 4 carbon stable as an organic acid.
compound

Water Loss

Has trouble with

water loss due to
transpiration.
Trouble with
photorespiration

Less water loss than

C3 plants.
Photorespiration is
not a problem

Grow very slowly

and no problems
with
photorespiration