Hatch and Slack Cycle:

KV Calicut SRS
 C4 is the alternative pathway of Calvin
cycle (C3 cycle) taking place during the
dark phase of photosynthesis.
 In the C4 cycle, the first stable compound
is 4- Carbon compound, amely Oxaloacetic
acid (OAA).
Hence it is called C4 cycle. The primary
CO2 acceptor is Phosphoenole pyruvic acid
(PEP).
 It is a process of CO2 fixation.
CO2 fixation is fast and more efficient. This
pathway was worked out by Hatch and
Slack in 1966. Hence the C4 cycle is also
called as Hatch- Slack cycle.
•Fixation of one molecule of CO2 requires 5
ATP and 3 NADH.
• C4 cycle occurs in plants like maize, sorghum
etc.
• Chloroplasts are dimorphic in nature.
Chloroplasts of bundle sheath are larger and
contain starch grains
• In the leaves of these plants, the vascular
bundles are surrounded by bundle sheath of
larger parenchymatous
•The chloroplast in mesophyll cells are
smaller and always contain grana.
• This peculiar anatomy of leaves of C4
plants is called Kranz anatomy.
•The bundle sheath cells are bigger and
look like a ring or wreath.
• Kranz in German means Wreath and
hence it is called Kranz anatomy.
• The C4 cycle involves two carboxylation
reaction, one taking place in chloroplasts of
mesophyll cells and another in chloroplast
of bundle sheath cells.
There are four steps in Hatch and Slack cycle:
i. Carboxylation: It takes place in
the chloroplast of mesophyll cells.
Phosphoenol pyruvate (PEP), a 3 carbon
compound picks up CO2 and changes into 4 C-
Oxaloacetate in the presence of water. This
reaction is catalyzed by the enzyme, PEP
Carboxylase.
PEP + CO2+H2O -------- oxaloacetate (4C) +
H3PO4 (PEP carboxylase)
ii. Breakdown: Oxaloacetate breaks down
readily into 4 C- Malate and Aspartate in the
presence of the enzymes, transaminase and
Malate dehydrogenase.
Oxaloacetate (4C) ------------ Aspartate (4C)
+Malate (4C) (Transaminase, Malate
dehydrogenase)
These compounds diffuse from the mesophyll
cells into sheath cells.
 Splitting: In the sheath cells Malate and
Aspartate split enzymatically to yield free
CO2 and 3C- pyruvate. The CO2 is used in
Calvin cycle in the sheath cells.
Malate --------------- CO2 + Pyruvate (Decarboxylation)

.
The second carboxylation occurs in the
chloroplast of bundle sheath cells. The CO2
is accepted by 5C- compound, ribulose
bisphosphate (RuBP) in the presence of
enzyme; carboxydimutase and ultimalely
yield 3- phosphoglyceric acid. Some of the 3
Phosphoglyceric acids is utilized in the
formation of sugar and the rest to
regenerate ribulose biphosphate
 Phosphorylation: The pyruvate molecule
is transferred to chloroplasts of mesophyll
cells, where it is phosphorylated to
regenerate PEP in the presence of ATP. This
reaction is catalyzed by pyruvate
phosphokinase and the phosphoenol
pyruvate is regenerated.
Pyruvate + ATP+ Pi --------------- PEP + AMP
+ Pyrophosphate (Pyruvate
phosphokinase)
PHOTORESPIRATION:
In 1920s, German biochemist Otto
Warburg (1883-1970) discovered that plants
consumed oxygen at a higher rate when they
were illuminated. He also found that this
increased rate of O2 consumption inhibited
photosynthesis. Stimulation of O2
consumption by light is now referred as
photorespiration.
photorespiration consumes ATP and
NADPH, the high energy molecule made by
the light reaction.
Thus, photorespiration is a wasteful
process because it prevents plants from
using their ATP and NADPH to synthesize
carbohydrates.
Photorespiration is a biochemical process in
plants especially under conditions of water
stress and oxygen inhibits the Calvin cycle, the
carbon fixation process of photosynthesis.
 In Calvin cycle, first CO2 fixation step, this
is the reaction where RuBP combines with
CO2 to form two molecules of 3PGA i.e.
catalyzed by RuBisCo.
RuBP +CO2 – 2*3PGA
In C3 plants some O2 does bind to
RuBisCO and hence CO2 fixation is
decreased.
Hence, the RuBP instead of being
converted to two molecule of PGA
binds with oxygen to form one
molecule of phosphoglycolate in a
pathway called photorespiration.
Photorespiration observed in all C3 plants
which have been examined but non-existent in
C4 plants.
 This is because C4 plants segregate their
RuBisCO enzyme in bundle sheath cells deep
within the leaf and the CO2 concentration in
these cells maintained at very high levels.
 C4 plants have higher growth rates than C3
plants because they do not waste their ATP and
NADPH in photorespiration.
 C3 plants face the problem of
photorespiration in hot dry days.
 These plants tend to close their stomata to
prevent excessive loss of water (from
transpiration).
 CO2 cannot enter the leaves (via the stomata)
resulting in the levels of CO2 within the leaves
to become low
. Since there are few CO2 molecule to fix the
O2 molecule used as a substrate to produce
G3P (Glyceraldehyde 3 Phosphate)
 ThreeFactors affecting
Photorespiration:
1) The rate of photorespiration
increases at any time when the
level of CO2 is low and O2 is
high. Such conditions occur
when stomata remain partially
closed or completely closed and
photosynthesis is underway.
3).
 2)The stomata of plants are open, resulting
in lowering down the rate of
photorespiration. But when plants become
water stressed, they close stomata to
prevent loss of water via transpiration. Thus,
on the other hand it restricts the exchange
of gases. The level of CO2 gradually rises
as water splits during light reaction.
 3) In desert and dry tropical areas,
photorespiration is reduced due to
water stress and on the other hand,
results in lowering down the potential
of plant growth. Some plants have
adapted to this problem by modifying
the way they carry out photosynthesis.
One of the common adaptations is
called CO2 metabolism in which plants
develop different leaf anatomy
called Kranz anatomy
Significance of C4 Cycle:
1. C4 plants have greater rate of carbon dioxide
assimilation than C3plants because PEPCO has
great affinity for CO2 and it shows no
photorespiration resulting in higher production
of dry matter.
2. C4 plants are better adapted to
environmental stress than C3 plants.
3. Carbon dioxide fixation by C4 plants requires
more ATP than C3plants for conversion of
pyruvic acid to PEPA.
4. Carbon dioxide acceptor in C4 plant is PEPA
and key enzyme is PEPCO.
5. They can very well grow in saline soils
because of presence of C4organic acid.
Crassulacean Acid Metabolism (CAM Pathway):
It is a mechanism of photosynthesis which occurs in succulents and
some other plants of dry habitats where the stomata remain closed
during the daytime and open only at night. The process of
photosynthesis is similar to that of C4 plants but instead of spatial
separation of initial PEPcase fixation and final Rubisco fixation of CO2,
the two steps occur in the same cells (in the stroma of mesophyll
chloroplasts) but at different times, night and day, e.g., Sedum,
Kalanchoe, Opuntia, Pineapple (Fig. 6.13). (CAM was for the first time
studied and reported by Ting (1971).
Factors Affecting Photosynthesis:
Photosynthesis is affected by both
environmental and genetic (internal) factors.
The environmental factors are light, CO2,
temperature, soil, water, nutrients etc. Internal
or genetic factors are all related with leaf and
include protoplasmic factors, chlorophyll
contents, structure of leaf, accumulation of
end product etc.
Principle of Limiting Factors:
Liebig (1843) proposed law of minimum which
states that the rate of a process is limited by
the pace (rapidity) of the slowest factor.
However, it was later on modified by Blackman
(1905) who formulated the “principle of
limiting factors”. It states that when a
metabolic process is conditioned as to its
rapidity by a number of separate factors, the
rate of the process is limited by the pace
(rapidity) of the slowest factor. This principle is
also known as “Blackman’s Law of Limiting
Factors.”
Blackman (1905) studied the effect of
CO2 concentration, light intensity and
temperature on rate of photosynthesis. All
other factors were maintained in optimum
concentration.
When sufficient light became available,
CO2 became limiting factor (Fig. 6.17). When
both are provided in sufficient quantity, the
rate of photosynthesis rose initially but again
reached a peak. It could not be increased
further. At this time, it was found that increase
in temperature could raise the rate of
photosynthesis up to 35°C. Further increase
was not possible. At this stage, some other
factor became limiting. Therefore, at one time
only one factor limits the rate of physiological
process.