C3 and C4 VS
CAM Plants
Topic Learning Outcome
To compare the photosynthesis process in C3,
C4 and CAM plants
C3
The majority of plants, e.g. rice
C4
CO2 temporarily stored as 4-C organic
acids resulting in more efficient C
exchange rate
Advantage in high light, high temperature,
low CO2
Many grasses and crops
e.g., corn, sorghum, millet, sugar cane
Crassulacean Acid Metabolism (CAM)
Stomata open during night
Advantage in arid climates
Many succulents
e.g., cacti, euphorbs, bromeliades, agaves
Types of Photosynthesis
CO2 fixation in CO2 fixation in CO2 fixation in
a C3 plant a C4 plant a CAM plant
C3 vs C4 Photosynthesis
In C3 plants the mesophyll cells have well-
formed chloroplasts and the cells are in parallel
layers
In C4 plants the bundle sheath cells surround
leaf veins also have chloroplasts, and mesophyll
cells are arranged concentrically
Leaf anatomy of a
C3 and C4 plant
C3 Plant C4 Plant
C4 plants fix CO2 by first forming a 4C molecule
prior to the Calvin cycle
4C molecule is then pumped into the bundle
sheath cells (BSCs) where it releases CO2 to the
Calvin cycle
In hot, dry climates oxygen accumulates in the
leaves of plants
C4 plants have an adaptive advantage in this
type of climate
In C3 plants, oxygen begins to compete with
CO2 for RuBP carboxylase and photosynthesis
stops
In C4 plants, CO2 is delivered to the Calvin cycle
in the BSCs which are sheltered from increased
oxygen levels
CAM Photosynthesis
CAM plants also produce a 4C molecule but this
occurs only at night
Stomata can open at night without much water
loss
This 4C molecule is stored in the mesophyll until
the next day
E.g. Succulents like cactuses, pineapples
CAM plants represent partitioning in time- allows
for conservation of water
C4 plants represent partitioning in space - light
rxns in mesophyll and Calvin cycle in bundle
sheath cells
Cross Section of a C3 Leaf
Photosynthesis occurs within the mesophyll cells
in C3 plants, which form a dense layer on the
upper surface of the leaf and a spongy layer on
the lower surface
Bundle-sheath cells
surrounding the
veins are not
photosynthetic
Stomata (singular stoma) are microscopic
openings on the undersurface of leaves that
allow gas exchange and water evaporation from
inside the leaf
Because dehydration
can be a serious
problem, the stomata
close when the plant
is under water stress.
When closed, CO2
needed for the Calvin
cycle cannot enter
Cross Section of a C4 Leaf
C4 Plants
Hot, moist environments
15% of plants (grasses, corn, sugarcane)
Divides photosynthesis spatially
Light rxn - mesophyll cells
Calvin cycle - bundle sheath cells
C4 Pathway
Some plants have developed a preliminary step
to the Calvin Cycle - This preamble step is
known as C4
While most C-fixation begins with RuBP, C4
begins with a new molecule,
phosphoenolpyruvate (PEP), a 3-C chemical
that is converted into oxaloacetic acid (OAA, a
4C chemical) when carbon dioxide is combined
with PEP
OAA is converted to Malic Acid, then transported
from the mesophyll cell into the bundle-sheath
cell, broken down into PEP & CO2
CO2 then enters the Calvin Cycle, with PEP
returning to the mesophyll cell
Produced sugars are now adjacent to the leaf
veins and can readily be transported throughout
the plant
The capture of CO2 by PEP is mediated by the
enzyme PEP carboxylase, which has a stronger
affinity for CO2 than does RuBP carboxylase
When CO2 levels decline below the threshold for
RuBP carboxylase, RuBP is catalyzed with O2
instead of CO2, forms glycolic acid, can be broken
down by photorespiration, producing neither NADH
nor ATP, in effect dismantling the Calvin Cycle
C4 plants evolved in the tropics and are adapted to
higher temperatures than are the C3 plants found at
higher latitudes
CO2 Fixation in C4 Plants
CO2 C3 C3
C4
CO2
Calvin
cycle
The Calvin cycle occurs in
bundle sheath cells
C4 Plants
Advantages of C4 metabolism
Avoid the photorespiratory loss of carbon
Improves the water use efficiency of the plants
Results in higher rates of photosynthesis at high
temperature
Improves the efficiency of N utilization (because
C3 require lots of Rubisco)
Crassulacean Acid Metabolism -
CAM Plants
Hot, dry environments
5% of plants (cactus and ice plants)
Stomata closed during day; open during the
night
Light rxn - occurs during the day
Calvin Cycle - occurs when CO2 is present
Mesophlly cell
CAM Plants
Mesophlly cell
CAM: Sequence of Events
NIGHT
Stomata open
Nocturnal transpiration & carbon fixation by
PEPcase
OAA produced
Reduced with NADPH to malate
Shuttled into vacuole
Acid content of vacuole increases
Starch depleted to provide PEP for carboxylation
DAY
Stomata close
Transpiration decreased
Acid content decreases
Malate decarboxylated to provide CO2 for Calvin
cycle
Starch content increases
Comparison of C4 and CAM Plants
Sugarcane Pineapple
1)CO2 incorporated
into four carbon
organic acids
(carbon fixation)
2)Organic acids
released CO2 into
Calvin cycle
a) Spatial separation of steps b) Temporal separation of steps
Environmental Change &
Photosynthesis:
C3 vs. C4 vs. CAM
Increasing CO2
Increasing N
Changes in water
CO2 Effects On Photosynthesis
C4 > C3 at low CO2
But, C3 > C4 at high CO2
If CO2 is low
6 CO2
CO2
6 C-C-C-C-C-C
O2
RUBISCO
RuBP 6 C-C-C-C-C
When the concentration of CO2 is low, oxygen will bind to
the active site of RUBISCO. When oxygen is bound to
RUBISCO, RuBP is broken down and CO2 is
released. This wastes energy and is no use to the plant.
It is called photorespiration because oxygen is taken up
and CO2 is released.
Photosynthetic N-use efficiency
C4 plants need (have) less leaf N than C3
Photosynthesis higher per unit N in C4
Humans are increasing global N, which benefits
C3 more than C4
Increasing CO2 decreases leaf N content, more
in C3 than C4
Photosynthetic Water-use
Efficiency
C4 plants use less water than C3 (because
stomata open less)
Water availability may increase or decrease the
rate of photosynthesis
C3 vs C4 vs CAM
Feature C3 C4 CAM
Leaf anatomy no distinct Kranz Usually no
bundle (wreath) palisade cells,
sheath anatomy large
vacuoles
Initial carboxylating rubisco PEPcase PEPcase
enzyme
Product of CO2 fixation PGA (3C) OAA (4C) OAA (4C)
Theoretical energy 1: 3: 2 1: 5 : 2 1: 6.5: 2
requirements (CO2:
ATP: NADPH)
Transpiration ratio (g 450-950 250-350 18-125
H2O/g dry wt)
Photosynthesis rate (mg 15 - 30 40 - 80 (low)
CO2 fixed dm-2 h-1)
chl a/b ratio 2.8 3.9 2.5 - 3.0
Carbon dioxide 50 - 150 (Hi) 0-10 (low) 0-5 (in dark)
compensation point (ppm)
Response to light Light No light -
saturation saturation
easily
achieved
Photosynthesis inhibited Yes No Yes
by oxygen?
Photorespiration Yes Only in bundle Late afternoon
detectable? sheath
Temperature optimum for 15-25 30-47 35
photosynthesis
Dry matter production Low High low, variable
(bushels/acre) (26 – soybean; (87 – maize;
30 – wheat) 50 –
sorghum)
Chloroplasts one type dimorphic one type
