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Respiration :
(Latin: re, repeatedly + spirare, to
breathe )
a chemical oxidation controlled and
catalyzed by enzymes that in living
protoplasm break down carbohydrates
and fats, thus releasing energy to be
used by the organism in doing work

Aerobic vs Anaerobic Respiration

There are many variations of cellular


respiration.
Some organisms require the presence of
oxygen for these processes, called
aerobic processes.
Other organisms carry out a form of
respiration that does not require oxygen;
these processes are called anaerobic.

Respiration

Respiration:
C12H22O11 + 12O2 12 CO2 + 11H2O
generation of energy;
generation of carbon precursors for biosynthesis
and cellular functions.
CO2, H2O, and ATP, NADPH, NADH, FADH
Four main processes:
Glycolysis;
The oxidative pentose phosphate pathway;
The citric acid cycle;
Oxidative phosphorylation
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Overview of respiration

Structures and reactions of the major electron-carrying nucleotides

Figure 11.3 Reactions of plant glycolysis and fermentation

Plant glycolysis and fermentation; (A) Main glycolytic pathway

(glyco = sugar; lysis = to split)


Carbohydrates are converted
into pyruvate in the cytosol,
and a small amount of ATP is
synthesized via the substratelevel phosphoralation. NADH
is also produced.
Plant glycolysis has alternative
enzymes for several steps. Start
from sucrose, end with
pyruvate and malate in plants.
Calvin-Benson
cycle

Sucrose is split into glucose


and fructose: invertase or
sucrose synthase pathway

Glycolysis

In the initial phase of glycolysis, each hexose


unit is phosphorylated twice and then split to
produce two triose phosphate molecules
eventually.
When O2 is not available, fermentation
regenerates NAD+ for glycolysis.
Plant glycolysis is regulated from the bottom up
by its products the primary regulation at the
level of PEP metabolism by pyruvate kinase and
PEP carboxylase.

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Gluconeogenesis

Reactions of the oxidative pentose phosphate pathway in plants

Irreversible
Reversible
Irreversible

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The oxidative pentose phosphate pathway


The reaction are carried out by soluble enzymes in the
cytosol and in plastids.
The first two reactions convert the 6C (glucose 6phosphate) into the 5C ribulose 5-phosphate, 1 CO2 and
2 NADPH.
Irreversible.
The remaining reactions convert ribulose 5-phosphate
into glyceraldehyde 3-phosphate and fructose 6phosphate.
The products (G3P) can be further metabolized by
glycolysis.
The pathway accounts for 10 -15% of glucose
breakdown in plants.
6 Glucose-6-P + 12 NADP+ + 7 H2O
5 glucose-P + 6 CO2 +Pi + 12 NADPH + 12 H+
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Roles of the oxidative pentose phosphate pathway plays


NADPH supply in the cotosol;
NADPH supply in the plastids;
Supply of substrates for biosynthetic processes - ribose-5phosphate is a precursor of ribose and deoxyribose
needed in the synthesis of nucleic acids.
This pathway is redox-regulated. The initial reaction
catalyzed by glucose-6-phosphate dedyhydrogenase is
inhibited by a high ratio of NADPH to NADP+.
In the light, there is little operation of this pathway in
chloroplast as the end products of the pathway being
synthesized by the Calvin-Benson cycle.

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Structure of plant mitochondria

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Figure 11.6 The plant citric acid cycle

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The Citric acid cycle

Pyruvate is oxidized to CO2 within the


mitochondrial matrix through the citric acid
cycle, generating a large number of
reducing equivalents in the form of NADH
and FADH2.
A unique feature of the plant citric acid
cycle is malic enzyme, which participates
in alternative pathways for the metabolism
of malate derived from glycolysis.

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Figure 11.7 Malic enzyme and PEP carboxylase provide


metabolic flexibility (A)

Malic enzyme converts malate into


pyruvate and thus makes it possible
for plant mitochondria to oxidize
both malate and citrate to CO2
without involving pyruvate
delivered by glycolysis

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Figure 11.7 Malic enzyme and PEP carboxylase provide metabolic flexibility (C)

With the added action of PEP carboxylase to the


standard pathway, glycolytic PEP can be
converted into 2-oxoglutarate, which is used for
nitrogen assimilation
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Organization of the electron transport chain and ATP synthesis

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Electron transport and ATP synthesis


Electron transport from NADH and FADH2 to oxygen is
coupled by enzyme complexes to proton transport across
the inner mitochondrial membrane. This generates an
electrochemical proton gradient used for powering
synthesis and export of ATP.

Complex I NADH dehydrogenase


Complex II succinate dehydrogenase
Complex III cytochrome bc1 complex
Complex IV cytochrome c oxidase
Complex V ATP synthase

During aerobic respiration, up to 60 ATP molecules are


produced per sucrose molecule.
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ATP synthesis

Typical for plant respiration is the presence of


several proteins (alternative oxidase, NAD(P)H
dehydrogenases, and uncoupling proteins) that
lower the energy recovery.
The main products of the respiratory process are
ATP and metabolic intermediates used in
biosynthesis. The cellular demand for these
compounds regulates respiration via control
points in the electron transport chain, the citric
acid cycle, and glycolysis.

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Mechanisms lower the ATP yield

Alternative oxidase
Plants display a capacity of cyanide-resistant respiration
due to alternative oxidase
It plays a role to lower ATP production and generate heat
in certain tissues or stressed conditions

The uncoupling protein


Increase the proton permeability of the membrane thus
as an uncoupler. Less ATP and more heat are
generated.

The rotenone-insensitive NADH dehydrogenases


work as a non-proton-pumping bypass when
complex I is overloaded.
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Figure 11.8 Organization of the electron transport chain and ATP


synthesis

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Figure 11.9 Transmembrane transport in plant mitochondria

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Figure 11.10 Metabolic interactions between mitochondria and cytosol

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Figure 11.11 Metabolic regulation of pyruvate dehydrogenase (PDH) activity

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Figure 11.12 Model of bottom-up regulation of plant respiration

Green arrows: stimulation (ADP,


Pi)
Red lines and squares: inhibition
ATP, NADH, citrate, PEP

Respiration can be up- or downregulated in response to the


changing demands for its products:
ATP and organic acids

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Glycolysis, the oxidative pentose phosphate pathway, and the


citric acid cycle

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Respiration in intact plants and tissues

More than 50% of the daily photosynthetic yield


may be respired by a plant;
Many factors can affect the respiration rate
observed at the whole-plant level. These factors
include the nature and age of the plant tissue
and environmental factors such as light,
temperature, nutrient and water supply, and O2
and CO2 concentrations.
Maintenance respiration
Growth respiration
Respiratory quotient (RQ): RA =
CO2 release/O2 consumption
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Figure 11.14 Structural features of triacylglycerols and polar


glycerolipids in higher plants

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Figure 11.15 Major polar glycerolipid classes in plant membranes

Glyceroglycolipids: sugar
forms the head group

Glycerophospholipids:
head contains phosphate

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Lipid metabolism

Triacylglycerol (fats and oils) are an


efficient form for storage of reduced
carbon, particularly in seeds. Polar
glycerolipids are the primary structural
components of membranes.
Triacylglycerols are synthesized in the ER
and accumulate within the phospholipid
bilayer, forming oil bodies.

Figure 11.16 Cycle of fatty acid synthesis in plastids of plant cells


ACP: acyl carrier protein

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Figure 11.17 The two pathways for glycerolipid synthesis

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Lipid metabolism

Fatty acids are synthesized in plastids using


acetyl-CoA, in cycles of two-carbon addition.
Fatty acids from the plastids can be transported
to the ER, where they are further modified.
The function of a membrane may be influenced
by its lipid composition. The degree of
unsaturation of the fatty acids influences the
sensitivity of plants to cold, but does not seem to
be involved in normal chiling injury.
Certain membrane lipid breakdown products,
such as jasmonic acid, can act as signaling
agents in plant cells
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Figure 11.18 Conversion of fats into sugars; (A) Carbon flow during fatty acid
breakdown

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Figure 11.18 Conversion of fats into sugars; (B) Electron micrograph

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Germination

During germination in oil-storing seeds,


the stored lipids are metabolized to
carbohydrates in a series of reactions that
include the glyoxylate cycle. The
glyoxylate cycle takes place in
glyoxysomes, and subsequent steps occur
in mitochondria.
The reduced carbon generated during lipid
breakdown in the glyoxysomes is
ultimately converted into carbohydrates in
the cytosol by gluconeogenesis.
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